The Potato Crop

THE POTATO CROP
World Crop Series

Available
The Grass Crop
The physiological basis of production
M.B. Jones and A. Lazenby
The Tomato Crop
A scientific basis for improvement
J.G. Atherton and J. Rudich
Wheat Breeding
Its scientific basis
F.G.H. Lupton
Forthcoming titles
Sugar Beet
D.A. Cooke and R.K. Scott
Bananas and Plantains
S. Gowen
Oats
R.W. Welch
The Groundnut Crop
J. Smartt
THEPOTATO
CROP
The scientific basis for improvement
Edited by
Paul M. Harris
Department of Agriculture
University of Reading
fiji Springer-Science+Business Media, B.V.

First edition 1978
Second edition 1992
© 1992 Springer Science+Business Media Dordrecht
Originally published by Chapman & HalI 1992
Softcover reprint of the hardcover 2nd edition 1992
Chapter 13 © 1992 B.D. Witney and D.C. McRae

Typeset in 10112pt Times by J&L Composition Ltd, Filey, North Yorkshire
Suffolk
ISBN 978-94-010-5034-0 ISBN 978-94-011-2340-2 (eBook)
DOI 10.1007/978-94-011-2340-2
Apart from any fair dealing for the purposes of research or private study, or
criticism or review, as permitted under the UK Copyright Designs and
Patents Act, 1988, this publication may not be reproduced, stored, or
transmitted, in any form or by any means, without the prior permission in
writing of the publishers, or in the case of reprographic reproduction only in
accordance with the terms of the licences issued by the Copyright Licensing
Agency in the UK, or in accordance with the terms of licences issued by the
appropriate Reproduction Rights Organization outside the UK. Enquiries
concerning reproduction outside the terms stated here should be sent to the
publishers at the London address printed on this page.
The publisher makes no representation, express or implied, with regard to
the accuracy of the information contained in this book and cannot accept
any legal responsibility orliability for any errors or omissions that may be
made. .
A catalogue record for this book is available from the British Library
Library of Congress Cataloging-in-Publication Data
The Potato crop: the scientific basis for improvementledited by
Paul M. Harris.
p. cm.
Includes bibliographical reference.s and index.
ISBN 4-010-5034-0
1. Potatoes. I. Harris, Pald M., 1931-
SB211.P8P775 1991
635' .21--dc20 91-23381
CIP
Contents
Contributors xi
Preface to the second edition xv
Preface to the first edition xvi
Foreword xviii
1 History of the potato 1
J.G. Hawkes
1.1 Introduction 1
1.2 The status of the potato in South America 1
1.3 Evidence for domestication - archaeological data 2
1.4 The potato at the time of the Spanish conquest 3
1.5 Historical and linguistic evidence 4
1.6 The introduction of the potato into Europe 5
1.7 The nature and exact source of the early European potato 6
1.8 The spread of the potato into other parts of the world 9
References 11
2 Biosyst,ematics of the potato 13
J.G. Hawkes
2.1 Introduction 13
2.2 Brief outline of potato classification 13
2.3 Distribution and ecology of potatoes 41
2.4 Species concepts in potatoes 46
2.5 Crossability, sterility and breeding behaviour 48
2.6 Endosperm balance numbers 51
2.7 Cytology of potato species 52
2.8 Chemotaxonomic relationships of potatoes 53
2.9 Evolutionary divergence and genome differentiation in wild
potato species 55
2.10 Potato evolution 56
2.11 Evolutionary relationships of cultivated potato species 57
References 60
vi Contents
3 Structure and development of the potato plant 65

Elizabeth G. Cutter
3.1 Introduction 65
3.2 Development of the seedling 66
3.3 The root system 67
3.4 The leafy shoot 70
3.5 The stolon 86
3.6 The tuber 96
3.7 Dormancy and its control 128
3.8 Development and structure of the flower 133
3.9 Embryogenesis 137
3.10 Tissue culture 138
3.11 Genetic engineering 144
3.12 Conclusions 145
Acknowledgements 146
References 146
4 Mineral nutrition 162

P.M. Harris
4.1 Introduction 162
4.2 Nutrient accumulation and use 163
4.3 Relations between mineral nutrients, growth and yield 168
4.4 Fertilizer requirements 177
4.5 Some factors modifying the yield response to fertilizers 194
4.6 Fertilizer use and potato quality 205
4.7 Conclusions 207
References 209
5 Water relations and growth of potatoes 214

P.J. Gregory and L. P. Simmonds
5.2 Growth and water use 214
5.3 Y.ield and water supply 219
5.4 Root growth 223
5.5 The physiological basis for the drought-sensitivity of
potato growth 228
5.6 Water supply and tuber quality 236
References 241
6 Seed tuber production and management 247

E.J. Allen, P.l. O'Brien and D. Firman
6.2 Growth and development of the seed tuber 247
6.3 Seed multiplication and certification 271
6.4 Seed production and utilization 278
Contents vii
6.5 True seed 286

References 287
7 Plant density 292
E.l. Allen and D.C.E. Wurr
7.2 Units of density 293
7.3 Spatial arrangement 306
7.4 Effects of plant density 312
References 330
8 Breeding new varieties 334
P.D.S. Caligari
8.2 Breeding methods 337
8.3 Breeding objectives 353
8.4 The potential role of other breeding techniques 362
References 364
9 Weed control 373
P.l. W. Lutman
9.1 Weeds in potatoes 373
9.2 Weed biology 376
9.3 Weed control 379
9.4 Recommendations for the use of herbicides 382
9.5 Potato haulm destruction 394
9.6 Potatoes as weeds 396
References 399
10 Disease aspects of potato production 403
G.A. Hide and D.H. Lapwood
10.2 Effects of diseases 405
10.3 Survival and spread of pathogens 419
10.4 Control of diseases 421
10.5 Future prospects 431
References 432
11 Pest aspects of potato production
Part 1 The nematode pests of potatoes 438
K. Evans and D.L. Trudgill
11.1 Introduction to their biology 438
11.2 Effects of root-feeding nematodes on growth and yield 447
11.3 Interactions with other organisms 452
11.4 Methods of decreasing nematode damage 453
11.5 Future developments in nematode research 462
References 466
Vlll Contents
Part 2 Insect pests 476

K. V. Raman and E. B. Radcliffe
11.7 Pest biology, damage and distribution 480
11.8 Pest control measures 490
11.9 Integrated pest management (IPM) 495
References 497
12 Tuber quality 507

R. M.J. Storey and H. V. Davies
12.2 Quality associated with the fresh tuber 512
12.3 Quality associated with internal factors 521
12.4 Aspects affecting quality of the cooked potato 533
12.5 Processing quality 540
12.6 Irradiation 548
12.7 Concluding comments 551
References 552
13 Mechanization of crop production and handling operations 570

B.D. Witney and D.C. McRae
13.2 Seedbed preparation 570
13.3 Fertilizer application 573
13.4 Potato planting 574
13.5 Harvesting 578
13.6 Handling and grading 593
13.7 Conclusions and future prospects 604
References 604
14 The physics and physiology of storage 608

W. G. Burton, A. van Es and K.J. Hartmans
14.2 Physics 612
14.3 Wound healing and curing 651
14.4 Dormancy, sprout growth and ageing 657
14.5 Changes in composition during storage 677
References 709
15 Potato production in the tre,pics 728

D.J. Midmore
15.2 Production statistics for the tropics and sub tropics 729
15.3 Tropical highland potato production (> 1500 m) 731
15.4 Tropical lowland potato production « 1500 m) -
constraints and solutions 742
Contents ix
15.5 Tropical lowland potato production - two practical

examples 776
15.6 General comments and future 779
References 780
16 Potato production in the context of the world and farm economy 794
D.E. Horton and 1.L. Anderson
16.1 Global patterns and trends 794
16.2 Potatoes in the food systems of developing countries 804
16.3 Potatoes in developed countries 806
References 815
17 Principles of agronomy and their application in the potato industry 816

E.J. Allen and R. K. Scott
17.1 Crop growth and development 816
17.2 The use of agronomic principles in understanding the
effects of agronomic treatments 845
17.3 Relevance of agronomic understanding to improvements
in the potato industry 873
References 878
Index 882
Contributors
E.J. Allen
Cambridge University Farm
Huntingdon Road
Cambridge CB30LH, UK
J.L. Anderson
The Scottish Agricultural College
Rural Resource Management Division
42 South Oswald Road
Edinburgh EHO 2HH, UK
Dr W.G. Burton
deceased
P.D.S. Caligari
Department of Agricultural Botany
Plant Science Laboratories
Whiteknights
PO Box 221
Reading RG6 2AS, UK
E.G. Cutter
School of Biological Sciences
Department of Cell and Structural Biology
University of Manchester
Williamson Building, Oxford Road
Manchester M13 9PL, UK
H.V. Davies
Scottish Crop Research 1nstitute
1nvergowrie
Dundee DD2 5DA, UK
XlI Contributors
A. van Es
Pootakkerweg 28
6706 BX
Wageningen, The Netherlands
K. Evans
Crop and Environment Protection Division
Entomology and Nematology Department
Rothamsted Experimental Station
Harpenden
Herts AL5 2JQ, UK
D. Firman
Huntingdon Road
Cambridge CB3 OLH, UK
P .J. Gregory
CSIRO
Dryland Crops and Soils Research Unit
Private Bag
PO Wembley
WA6014
Western Australia
P.M. Harris
Earley Gate
PO Box 236
Reading RG6 2AT, UK
K.J. Hartmans
IBVL
PO Box)8
6700 Wageningen, The Netherlands
J.G. Hawkes
c/o School of Continuing Studies
University of Birmingham
PO Box 363
Birmingham BI52TT, UK
G.A. Hide
Plant Pathology Department
Harpenden
Herts AL5 2JQ, UK
Contributors Xlll
D.E. Horton
ISNAR
PO Box 93375
The Hague, 2509 Al
The Netherlands
D.H. Lapwood
Plant Pathology Department
Harpenden
Herts AL5 2IQ, UK
P.I.W. Lutman
Crop Management Department
AFRC-IACR Rothamstead Experimental Station
Harpenden, Herts, AL5 2IQ
D.C. McRae
Scottish Centre of Agricultural Engineering
Bush Estate
Penicuik
Midlothian, EH260PH, Scotland, UK
D.I. Midmore
The Asian Vegetable Research and Development Center
PO Box 42, Shanhua
Tainan, Taiwan, 74199
Republic of China
P.I. O'Brien
Huntingdon Road
Cambridge CB3 OLH, UK
E.B. Radcliffe
Department of Entomology
College of Agriculture
University of Minnesota
219 Hodson Hall, 1980 Folwell Avenue
St Paul, Minnesota 55108-1385, USA
K.V. Raman
Nematology and Entomology Department
International Potato Center
PO Box 5969
Lima, Peru
XIV Contributors
R.K. Scott
School of Agriculture
Nottingham University
Sutton Bonington
Loughborough
Leics LEl2 5RD, UK
L.P. Simmonds
Department of Soil Science
London Road
Reading RGI 5AQ, UK
R.M.J. Storey
Potato Marketing Board
Broadfield House
4 Between Towns Road
Cowley
Oxford OX43NA, UK
D.L. Trudgill
Scottish Crop Research Institute
Invergowrie
Dundee DD2 5DA, UK
B.D. Witney
Scottish Centre of Agricultural Engineering
Bush Estate
Penicuik
Midlothian, EH260PH, Scotland, UK
D.C.E. Wurr
Horticulture Research International
Wellesbourne
Warwick CV35 9EF, UK
Preface to the second edition
Research and publications on the potato crop have burgeoned since the
first edition of this book was published in 1978. However, the warm
reception of the first edition suggested that it had a useful part to play in
promoting the scientific basis for understanding and improving the yield
and quality of the crop. Since the first edition was out of print and a second
reprint would not have taken into account the contributions made by
research over the intervening years, it became obvious that a complete
revision was necessary. There was, in particular, a need to take account of
the rapid extension of interest in the crop into climates and farming
systems with which it has not been traditionally associa,ted.
Those involved with the crop will be sadly aware that a number of
contributors to the first edition are no longer with us. Their contribution to
our knowledge of the crop will however be a permanent legacy of their
achievement. I would like to thank all those who have contributed to the
book for their willingness to cooperate in the difficult task of bringing their
particular subject up to date. This is even more noteworthy for the
pressure of time appears to be almost an order of magnitude higher than it
was when the first edition was tackled. Under the circumstances it is
perhaps not surprising that not all the initial objectives have been
achieved; that most of them have is a tribute to the contributors, to whom
I would like to dedicate this edition of the book.
Paul M. Harris
November 1991
Preface to the first edition
One of the major objectives of this book on the potato has been to review
the knowledge which is fundamental to successful potato production
and upon which further improvement must be based. Underlying this
objective is the belief that high yields, efficiently obtained are of prime
importance to producers and consumers and to research workers who serve
the interests of both. Despite the paradox that the greatest financial
returns are often achieved when yields are lower than normal, profitability
through scarcity is not in the long-term interest of either producers or
consumers.
This attempt to review the knowledge on which the scientific production
of potatoes is based has necessitated seeking contributions from a fairly
large number of specialists, which is a well-recognized method of dealing
with subjects which are too large to be considered in the necessary depth
by anyone author. However this tendency to specialize within fairly
narrow limits carries with it the attendant dangers that few research
workers see the complete picture; on the other hand the producer is
concerned with the whole system of producing a crop but may not have
sufficient knowledge of particular components of the system which may
significan~ly affect the efficiency with which his enterprise is operated. It is
the hope that this book, read in its entirety, will provide the basic
biological principles on which more productive and economic systems of
potato production can be based.
Inevitably the views presented in this book will have been coloured by
the particular experiences and enthusiasm of the contributors, and no
doubt much has been omitted that might have been included and possibly
vice versa; however the overall balance is the responsibility of the editor
and any shortcomings in this respect should be attributed to him. It is my
hope that anyone reading the book who h,as constructive suggestions to
make will have no hesitation in contacting the editor or contributors.
I am indebted to Professor Rober,ts for asking me to edit this book and to
the contributors who have so willingly cooperated in its production.
Thanks are also due to my wife and family for their patient forebearance
Preface to first edition xvii
during the book's preparation. Finally I would like to dedicate this book to
all those who have contributed to our knowledge of this fascinating, useful
and enjoyable vegetable.
University of Reading P.M.H.
June, 1977
Foreword
The first edition of this book was published twelve years ago. It established
the first of a continuing series of monographs on important crops. This
series arose from the perception that, while the disciplines which contri-
bute to improving crop production are becoming more specialized, so it is
becoming increasingly necessary to integrate this knowledge in order to
improve production systems. Farmers, growers and agronomists have
always had to try and do this; but it is also important for the scientific
specialists who serve them to have some grasp of cognate disciplines, so
that an understanding of the wider context can help them to target their
contribution more effectively.
It is no longer really possible for one author to write an up-to-date book
which adequately and critically deals with all facets of a crop. An account
which satisfactorily covers the field requires the efforts of a number of
specialists. And if this account is to be coherent and well-balanced it needs
careful planning and editing. The first edition of this book was based on
this simple principle and it became the model for the series.
The reception and demand for the first edition suggests that many people
concerned with the potato crop found this approach helpful. In the
meanwhile, considerable progress has been made in some areas; some of
the original authors have moved on to other interests, and sadly, others
have died. A new edition was clearly called for and the opportunity has
been taken for a complete revision.
The underlying philosophy which concentrated on unde.rlying principles,
which consequently are not limited by geographical boundaries, remains
the same. But the contributions of several new authors, and the recogni-
tion of an increasing interest in the crop in the tropics, has given this new
edition an even greater international flavour. I believe that anyone who is
seriously interested in this important crop will wish to have a copy near to
hand.
Eric H. Roberts
World Crop Series Editor
November 1991
CHAPTER 1
History of the potato

J.G. Hawkes
1.1. INTRODUCTION
The time and place of origin of cultivated plants and their subsequent
evolution under domestication have caught the imagination of botanists
and agricultural scientists from at least the early days of the last century.
The studies of de Candolle, Vavilov and others have shown the need for a
synthesis of information from such diverse fields as cytogenetics, history,
linguistics, botany and archaeology to help trace the complex pattern of
evolution of our ancient crops, many of which were already being
cultivated some nine or ten thousand years ago.
The potato is undoubtedly of ancient origin, although our knowledge of
its early stages of domestication is not so precise as that of some other
crops, such as wheat and barley. We know, however, that it was domesti-
cated in South America and that it had been dispersed by man over a con-
siderable area by the time the Spaniards arrived in the sixteenth century ..
1.2 THE STATUS OF THE POTATO IN SOUTH AMERICA
At one time, on the evidence of Gerard (1597), the potato was thought to
have come from the colony of Virginia (now North Carolina), in the North
American continent. Nevertheless, all other evidence points away from
this hypothesis and places the source of the potato in South America. Data
from early Spanish post-conquest chronicles as well as archaeological
remains show clearly that the potato was an ancient domesticated plant of
South America by the time the Spaniards arrived. Furthermore, many wild
species occur in South America, and more specifically in the Andes of Peru
and Bolivia, from one or more of which the cultivated potato must have
been derived.
Many wild species occur also in Mexico and Central America, as we shall
see in more detail in the next chapter. However, these never seem to have
been taken into cultivation, and the apparently indigenous forms of
The Potato Crop Edited by Paul Harris
Published in 1992 by Chapman & Hall, London. ISBN 0 412 29640 3
2 History of the potato
Figure 1.1 Ancient Peruvian vessel (Moche culture) representing a potato tuber.
Solanum tuberosum in Mexico and Central America all seem to have come
from South America, probably in early post-conquest days.
1.3 EVIDENCE FOR DOMESTICATION - ARCHAEOLOGICAL DATA
The most striking archaeological evidence for the antiquity of potato

cultivation is afforded by the truly amazing ceramics from the northern
coast of Peru belonging to the Moche, Chimu and Inca cultures (about the
fourth century AD onwards) (see Fig. 1.1). These ceramics, made in the
form of potatoes, have been described and analysed in some detail by
Salaman (1937, 1939, 1949). The Moche people, especially, were remark-
ably inventive and portrayed with accuracy and artistic feeling all kinds of
birds, fish, mammals and plants, as well as scenes of everyday life, portrait
heads and natural objects. Amongst these the vessels depicting potatoes
are rather rare, which is not so surprising when one realizes that these were
coastal peoples who probably obtained potatoes by trade or barter from
the mountains but who did not themselves grow this crop in their warm
coastal valley river oases.
Unfortunately, the pottery of the Highland peoples (Tiahuanaco,
Chavin, etc.) although very interestingly decorated with abstract patterns
The potato at the time of the Spanish conquest 3
and stylized representations of deities, reptiles, birds and felines, did not,
in general, depict plants. There is, however, one remarkable exception.
This is a series of ceremonial vessels or urns of great size (> 1 m tall) from
South Central Peru, of Tiahuanaco-Huari style dating from about AD 900.
These urns, known as 'Pacheco', apparently contained offerings of food
and were ceremonially destroyed and then buried. After restoration they
were found to depict Andean crop plants, such as potatoes, maize, oca
(Oxalis tuberosa) , ulluco (Ullucus tuberosus) , mashua (Tropaeolum
tuberosum) , and other plants, possibly quinoa (Chenopodium quinoa) ,
peppers (Capsicum spp.), and lupins.
All ceramics representing potatoes are restricted to Peru, and none have
been recovered from Colombia, Ecuador, Bolivia, Argentina or Chile,
even though the potato is certain to have been an ancient crop in those
countries also.
Actual remains of the potatoes themselves are found in tombs, store-
houses or dwellings and rubbish heaps but do not occur with much
frequency. A kind of dried potato (chuno or tunta), which is still produced
by Indians today by freezing and drying, after trampling and sometimes
washing, is known from some archaeological sites (see Hawkes, 1967).
Although certain materials were found from as far back as about 400 Be,
no really ancient remains had been identified with certainty until recently.
The situation at present is now more hopeful. Detailed studies of starch
and cell structures using light and scanning electron microscope techniques
have enabled us to identify potatoes from the Chilca valley near Lima,
radiocarbon dated at an age of c. 7000 years before present (b.p.)
(Martins, 1976). These were provided by Frederic Engel (1970) and co"
workers, whilst other remains of early food plants excavated by Moseley
(1975) have also been examined in which potatoes were dated to 4500 b.p.
More recent finds of potatoes at Casma in Peru were reported by Ugent et
al. (1982) dated to 4000 b.p. Ugent and his co-workers also described wild
potatoes from Chile that were apparently eaten though not cultivated, and
dated at 13 000 years b.p. (Ugent et al., 1987).
The Chilca canyon evidence takes the origin of potato cultivation ba,ck to
an age when maize first became cultivated in Mexico and equates it with
the approximate time of agricultural origins in the New World, based on
excavations in Mexico and elsewhere. From studies of the distribution of
existing primitive cultivated potatoes and the wild species most similar to
them it seems highly probable that the potato was first taken into
cultivation in the northern Bolivian region of Lake Titicaca/Lake Poopo.
Its ancestor could have been the wild species Solanum leptophyes.
1.4 THE POTATO AT THE TIME OF THE SPANISH CONQUEST

It seems quite clear from what has just been said that the potato was an
ancient cultivated plant at the time of the discovery of South America. We
do not know the geographical extent of its early cultivation since the oldest
archaeological finds are wholly from Peru. Because botanical evidence
points to an origin for the cultivated potato in the Peru-Bolivian region, it
is only to be expected that, once domesticated, it would have diffused
rapidly throughout the high Andes of those two countries. Indeed Weston
La Barre (1947) has asserted that the Highland cultures owed their
existence to the potato since there are few other plants that can be grown
at altitudes of 3500 m or more. Maize does poorly at such height,
and although there are certain other tuber crops, as has already been
mentioned when describing the 'pacheco' urns, these are grown on a small
scale. The small grained Chenopodium quinoa would hardly rank as a main
staple food, and the cultivated lupin is a difficult food plant to handle.
Unfortunately the archaeological evidence is limited to certain areas only.
1.5 HISTORICAL AND LINGUISTIC EVIDENCE
At the time of the conquest in the early sixteenth century we have accounts
by chroniclers (Fig. 1.2) of potato cultivation in what are now Colombia,
Ecuador, Peru, Bolivia and Chile (for detailed literature sources, see
Hawkes, 1967). Even at that early time it seems to have been an intensively
cultivated crop, and we can assume with some degree of certainty that
Figure 1.2 Planting and harvesting potatoes in Inca times; from a sixteenth-
seventeenth century manuscript by Guaman Poma de Ayala (reproduced by kind
permission of the Institut d'Ethnologie, Paris).
The introduction of the potato into Europe 5
many hundreds of varieties were grown over a distribution area not very
dissimilar from that in which it is cultivated today.
The native names of the potato also indicate an ancient and widespread
cultivation (see Hawkes, 1947), since they differ completely in the main
Indian languages that were spoken in the areas where the potato was
grown. Thus in the Chibcha language of Central Columbia the names
iomza, iomuy, etc., were used; in Quechua, the language of the Inca
empire, the usual name was papa, but this replaced earlier names (ajsu,
impari, etc.) in the languages of the vanquished tribes. In Bolivia, the
Aymani Indians used the words amka and choque, whilst in Chile, the
Araucanians gave it the name pOfU.
The Spaniards adopted the name papa for the potato throughout their
western American colonies, and other names as well as the languages
themselves have largely died out (although not in Aymara). Even so, the
old names for the potato in various regions still persist here and there in
varietal combinations or in dialect phrases.
In Europe the word papa was never adopted. On the contrary, our word
is derived from batata, the Caribbean Arawak Indian word for Ipomoea,
the sweet potato. This is because the Spaniards encountered the sweet
potato first, and not having a name for a tuber of that sort they used the
Indian word batata. Subsequently, other tuberous plants which they found
in their American colonies were given the same name. Patata and potato
are obviously cognate forms of batata; so the word papa, which is still
widely used in the whole of Spanish Latin America, never spread outside
this area, even though the plant itself is now grown in most other parts of
the world.
To sum up this section we can say with some certainty that historical and
linguistic evidence clearly corroborate archaeological evidence as to the
origin of the cultivated potato in the Andes of South America.
1.6 THE INTRODUCTION OF THE POTATO INTO EUROPE
The potato appeared in Europe during the last quarter of the sixteenth
century. Unfortunately, we know of no account describing its introduction
and we must perforce rely on circumstantial evidence only.
Legend has associated Sir Walter Raleigh and Sir Francis Drake with the
introduction of the potato into England. There is no factual evidence to
substantiate these claims, and much evidence which tends to negate them.
Thus Drake saw potatoes in Chile during his world voyage in 1578, but
since he did not return to England until 1580 he could not have kept
potatoes with him in a living condition for two years (see Drake, 1628).
Furthermore, Clusius (1582) who described Drake's plants collected
during this voyage from a visit he made to England in 1581 makes no
mention of the potato either.
Raleigh is reported to have seen and collected potatoes on one of his
voyages, but from a knowledge of the locality we can be certain that these
were sweet potatoes and not S. tuberosum. Indeed, Raleigh evidently
never went near any area where he might have found potatoes; however,
there may be some truth in the statement that Raleigh planted potatoes
from England on his estates at Youghal in southern Ireland.
Salaman (1937) quotes evidence that potatoes were bought in 1573 at
Seville and concludes that they could have arrived in southern Spain about
1570 and have been cultivated somewhere in the Seville region for some
years so as to build up stocks. This date of 1570 is generally quoted as the
approximate time of first arrival of the potato in continental Europe. We
assume that it came merely as ships' stores and was not at that time
regarded as a plant of special significance. From Spain it seems to have
spread to Italy, whence it was sent to th,e famous herbalist Clusius at
Vienna by Philippe de Sivry, Prefect of Mons in Belgium. From this
Spanish source the potato spread through Europe as a botanical curiosity
and no more.
A separate introduction was recorded in England by the herbalist John
Gerard (1596, 1597), who claimed erroneously, as we have already said, to
have received it from Virginia in North America. The reason for this is by
no means certain but it has been suggested that he confused plants brought
by Hariot from Virginia in 1586 (see Hariot, 1588) with the potato which
was received on another occasion, between 1588 and 1593 (see Hawkes,
1967, pp. 255-62), probably from a 'prize' Spanish ship captured by
English seamen.
1.7 THE NATURE AND EXACT SOURCE OF THE EARLY EUROPEAN

POTATO
Although there is no doubt that the European potato belongs to the species
S. tuberosum there has been considerable controversy as to what region of
South America it came from and thus whether subspecies tuberosum from
Chile or subspecies andigena from the Andes was first brought to Europe.
The Russian botanists, Juzepczuk and Bukasov (1929) regarded the
early European potato as having come from Chile and thus having been
adapted to the long day conditions of Europe right from the time of
its arrival. Salaman (1946, 1954), however, provided historical and geo-
graphical evidence which made an introduction by 1570 from Chile highly
unlikely. Since the Andean subspecies andigena is adapted to form tubers
under the 12 h day of tropical latitudes and does not tuberize in Europe
until very late in the season when the natural day length has shortened to
more or less 12 h in late September and early October, the Russian
botanists believed it could not for this reason have been the subspecies first
introduced into Europe.
The nature and exact source of the early European potato 7
It was nevertheless suggested by Salaman and Hawkes (see Salaman,
1946, 1949, 1954; Salaman and Hawkes, 1949; Hawkes, 1967) that the
early European potato came from the Andes, and perhaps from the
northern Colombian part. This would have meant that through strong
artificial selection in Europe from 1570 to about 1750 when Linnaeus
described the European S. tuberosum the potato would have changed from
one subspecies to the other, just as it had presumably done over the
previous millennia when it was taken by Indian tribes from the Andes to
Chile. This is not so far-fetched as it may seem at first glance, since
Simmonds (1966) succeeded in carrying out the same process for a third
time during the space of a very few generations of selection for earliness in
Britain.
If this hypothesis is correct we might expect to see evidence of subspecies
andigena in Europe in the late sixteenth and early seventeenth centuries
before it became changed into subspecies tuberosum. In fact, these
changes can be seen very clearly, in herbarium specimens, woodcuts and
descriptions (Salaman and Hawkes, 1949).
Andean potatoes when grown in the European long summer days
tuberize late and the stems continue to grow, producing tier after tier of
flowers and long branched stolons which often shoot up to form new
plantlets. The stems are thinner and more frequent than tuberosum stems
and the leaf is more highly dissected. Other differences (see Hawkes, 1956)
can also be noted. The tuber yield is poor since the night temperatures
become too low in September and October for efficient tuberization, and
the plants are often killed by frost.
The early descriptions and woodcuts agree very well with the features·
described above. Thus, Clusius (1601) describes stems up to 7.5 ft (2.286 m)
and 10.5 ft (3.2 m) long, whilst several authors (Clusius, 1601; Bauhin,
1620) mention that the plants are harvested in November. The many
stolons and small tubers of andigena when grown in Europe are featured in
the early illustrations (see Fig. 1.3; Bauhin, 1620), whilst a water-colour
painting of 1598 sent to Clusius by Philippe de Sivry also shows the
dissected leaves and small tubers. Finally, perhaps the most interesting
piece of evidence is the herbarium specimen of Caspar Bauhin made in
about 1620 which is still very similar to andigena.
A final piece of evidence is that the early potato in Europe was regarded
as a botanical curiosity and was not generally grown as a field crop until
about the mid-eighteenth century, despite the fact that it was a completely
accepted cultivated plant in South America. Only in very favoured places
in southern France and southern Ireland and elsewhere, with mild winter
climates, could it be grown as a crop. The reason for this is obvious. The
yields were too late and too small, due to the adverse photoperiodic
response in respect of tuberization, until selection for earliness had
resulted in varieties adapted to the long summer days of western and
northern Europe.
LIB E R Q V I N T V S.
LIBER QVINTVS
'DE SOLANO.
CAl'VT I.
Solanum tubcro[ul'Ilefc:ulcacum.
I'SOLANvMcuberofumcfculentum:caulem ha- admodlll1l figura nonn lk Matthiolo & huicli-

bet ad duomm criumvc cubicorum, rarius in bro addira eleganter, cum omnibus plantz.pani-
virialticudinem exurgentem,cralfum,angulofum, bus,exprimit) ex atro purpurafccntia, '.lbhlr(uta.
firiarum, leuicct hicfutum, in multos ramos infir- cJ:teris pallide vitentibus. abfque pedlculo. pal-
mos,in cecram,nili adminiculo Culbneantur, pro- mum 101l<'is. in Cex,odo, plurcs&pauciores par-
cumbentcs, brachi;ltum, cx quorum Ollis pediculi tcs,ex vna"'coIb depcndcntes.diuilis,ex rotundica-
crall, angulo{j, flores ClIllinenccs,prodeunt.Folia tc: oblongis,intcgris,quibus {jngulis,duo Cextuplo
qu:r primona[cunrur,Barbarc:r fimilia 1unt (que- minora intctijciuncur, IX [empc:rpars vna, qUJ: {e-
M ptima
Figure 1.3 The potato in Europe in the early seventeenth century (C. Bauhin,
1620). Note the long branched stolons, small irregular tubers, small daughter plants
formed from stolons, many inflorescences and abundant fruits. These features are
all typical of short-day adapted andigena potatoes when grown under the long
summer days of Europe. (Note: The bottom right-hand tuber may be a Jerusalem
articho ke. )
The spread of the potato into other parts of the world 9
We know that plants were raised from true seed at a very early date, thus
providing segregants on which selection could act. Clusius (1601) remarks
on this and describes the results in some detail; and there are other
accounts of selection for earliness in the literature. We thus have in the
potato a good example of selection for physiological characters long before
the development of the science of plant breeding.
1.8 THE SPREAD OF THE POTATO INTO OTHER PARTS OF THE

WORLD
We have already mentioned that the potato was introduced into Europe on
two occasions, firstly into Spain in about 1570 and secondly into England a
little later. This latter introduction took place between 1588 and 1593, with
a strong suggestion that 1590 was the actual year of arrival (Hawkes, 1967,
p.261).
From these two introductions it seems likely that the potato spread into
nearly every part of the world. From the Spanish introduction it diffused
through continental Europe and parts of Asia. From the English one, it
spread to Ireland, Scotland, and Wales and parts of northern Europe, and
from one or more of the British sources to most of the British overseas
colonies, including what was later to become the USA.
Let us look first at the diffusion of the potato through continental
Europe. From Spain it was said to have been taken to Italy by the
Carmelite Friars, and Clusius (1601) mentions that it was grown in Italy
before 1587 (see also Magazzini de Vallombrosa, 1623). In Portugal
potatoes appeared even earlier.
Clusius received potatoes in Vienna, from Italy via Mons in the low
countries and sent samples in the succeeding years to botanists in many
parts of Germany and Austria. They gradually spread by the end of the
seventeenth century to most German-speaking parts of Europe although
they only began to be grown on a large scale after about 1780 and did not
become really popular until the nineteenth century.
The Swiss herbalist C. Bauhin and his brother J. Bauhin evidently
obtained tubers at second hand from Clusius in the late sixteenth century
and had sent them to France by about 1600. Potatoes were extensively
grown in southern France by the mid-seventeenth century and probably
were more successful as a field crop earlier than in Germany because of the
milder autumns. However, there was considerable resistance to their
cultivation by all but the very poorest people until Parmentier made them
popular in about 1773.
The Slavic nations seem to have obtained their potatoes chiefly from
Germany since their words for this plant are derived from German ones,
such as Kartoffel, Grundbirne, etc. There are also words in Czechoslovakia,
Bulgaria and southern Russia cognate with Brandenburg, indicating the
German source for those regions also. Evidently the general adoption in
eastern Europe was late eighteenth to early ninteenth century (see Laufer,
1938).
The potato was first said to have been brought to Russia by Peter the
Great at the end of the seventeenth century from Holland, where of course
it had become well-known in gardens since Clusius' time. Only small
amounts were produced as delicacies for royal banquets up to the mid-
eighteenth century but by 1764 it began to be grown around St Petersburg
to a reasonable extent (Lechnovicz, 1970). Its adoption in all parts was
greatly speeded up by a royal decree in 1765 and potatoes were then sent to
all parts of the country for cultivation.
Now let us turn to potatoes derived from the introduction into England
in about 1590. For reasons already stated above the potato began in
England, as in most other parts of Europe, by assuming the role of
botanical curiosity and was esteemed as a delicacy in the court of James I.
It was not until the mid-eighteenth century that potatoes are recorded as
having been grown on a field scale, and this fits in rather well with accounts
from other parts of northern Europe. The same time scale is recorded also
for Scotland and Wales (Salaman, 1949). On the other hand there is much
evidence that the potato was grown on a field scale in Ireland, at least in
the south-west, by the early seventeenth century, probably due to the very
mild climate which allowed it to survive and set its tubers from October to
December. Southwell (1693) for instance, states that his grandfather
brought potatoes to Ireland 'who had them from Sir Walter Raleigh after
his return from Virginia'. Even though the latter part of this quotation
must be false, the fact that Raleigh died in 1618 would certainly give a last
date to the introduction (see also Salaman, 1949).
From Scotland the potato was said to have been taken to Norway in the
mid-eighteenth century and possibly thence to Sweden and Denmark,
although it seems probable that these last-named countries might have
already received samples earlier from German or other sources.
In North America the potato was completely unknown until the early
seventeenth century, and there seems to be not a shred of evidence to
support Gerard's account of its being native to Virginia or anywhere else on
the North American continent. One should perhaps say that wild potatoes are
known in the south-western states but this is quite another matter.
In fact the North American continent first received potatoes from
England via Bermuda in 1621, where they had been grown since an initial
importation from the mother country in 1613. Strangely enough, these first
North American potatoes were grown in Virginia, where Gerard some 35
years ago had erroneously assumed them to be native. Many other
introductions from England and Ireland were made later in the century,
but no records of an introduction from South America were made before
Goodrich (1863) obtained some varieties in a Panama market which were
said to have come from Chile.
References 11
Potatoes were said to have been taken to India by British missionaries in
the late seventeenth century and to China a little earlier. They were grown
in Japan by the late seventeenth century and in parts of Africa and the
West Indies by about the same period. Nowhere, even in tropical regions,
did they become widely grown before the mid-eighteenth century (Laufer,
1938). According to Yen (1961/2) the potato was first introduced into New
Zealand in 1769 by the French explorer, de Surville, followed by several
small introductions in the nineteenth century. They were adopted widely
by the Maoris in the early nineteenth century and were well established by
1840.
Thus a plant confined to South America until the late sixteenth century,
has in the course of four hundred years become a crop of world importance
ranking fourth in production after wheat, rice and maize.
REFERENCES
Bauhin, C. (1620) Prodromus Theatri Botanici, Frankfurt-on-Main.

Clusius, C. (1582) Aliquot Notae in Garciae Aromatum Historiam, Antwerp.
Clusius, C. (1601) Rariorum Plantarum Historia, Antwerp.
Drake, Sir F. (1628) The World Encompassed, (ed. W.S.W. Vaux), Hakluyt
Society, 16, 1854, London.
Engel, F. (1970) Exploration of the Chilca Canyon, Peru. Curro Anthrop., 11,
55-8.
Gerard, J. (1596) Catalogus Arborum, Fruticum ac Plantarum, etc. 1st edn,
London.
Gerard, J. (1597) The Herball or General Historie of Plantes, 1st edn, London.
Goodrich, C.E. (1863) The potato: its diseases, with incidental remarks on its soils
and culture. Trans. NY State Agric. Soc., 23, 103-39.
Hariot, T. (1588) A Briefe and True Report of the New Found Land of Virginia,
London.
Hawkes, J.G. (1947) On the origin and meaning of South American Indian potato
names. J. Linn. Soc. (Bot.), 53, 205-50.
Hawkes, J.G. (1956) Taxonomic studies on the tuber-bearing solanums. I.
Solanum tuberosum and the tetraploid species complex. Proc. Linn. Soc., 166,
97-144.
Hawkes, J.G. (1967) The history of the potato. J. Roy. Hort. Soc., 92, 207-24,
249-62, 288-302, 364-5.
Juzepczuk, S.W. and Bukasov, S.M. (1929) A contribution to the question of the
origin of the potato. Proc. USSR Congr. Genet. Pl. and Animal Breed., 3,
593-611.
La Barre, Weston (1947) Potato taxonomy among the Ayman'i Indians of Bolivia.
Acta Americana, 5, 83-103.
Laufer, B. (1938) The American plant migration. Part I. The potato. Field Mus.
Publ. 418, Anthrop. Ser., 28, 132pp.
Lechnovicz, V.S. (1970) The Potato, Leningrad, pp. 7-10.
Magazzini De Vallombrosa, Father (1623) Dell' Agricoltura Toscana,
Vallombrosa, Italy.
Martins, R.(1976) New Archaeological Techniques for the Study of Ancient Root
Crops in Peru, Ph.D. Thesis, University of Birmingham.
Moseley, M.E. (1975) The Maritime Foundations of Andean Civilization,
California.
Salaman, R.N. (1937) The potato in its early home and its introduction into
Europe. J. Roy. Hart. Soc., 62, 61-77, 112-23, 156--62, 253-66.
Salaman, R.N. (1939) Deformities and mutilations of the face as depicted on the
Chimu pottery of Peru. J. Roy. Anthrop. Inst., 69, 109-22.
Salaman, R.N. (1946) The early European potato; its character and place of origin.
J. Linn. Soc. (Bot.) 53, 1-27.
Salaman, R.N. (1949) The History and Social Influence of the Potato, Cambridge.
New edn (1985), ed. J.G. Hawkes.
Salaman, R.N. (1954) The origin of the early European potato. J. Linn. Soc.
(Bot.), 55, 185-90.
Salaman, R.N. and Hawkes, J.G. (1949) The character of the early European
potato. Proc. Linn. Soc., 161, 71-84.
Simmonds, N.W. (1966) Studies of the tetraploid potatoes. III. Progress in the
experimental re-creation of the Tuberosum Group. J. Linn. Soc. (BOlt..), 59,
279-88.
Southwell, Sir Robert (1693) Journal Book of the Royal Society of London, 13
Dec. 1693.
Ugent, D., Pozorski, S. and Pozorski, T. (1982) Archaeological potato tuber
remains from the Casma valley of Peru. Econ. Bot., 36, 182-92.
Ugent, D., Dillehay, T. and Ramirez, C. (1987) Potato remains from a late
Pleistocene settlement in southcentral Chile. Econ. Bot., 41,17-27.
Yen, D.E. (1961/62) The potato in early New Zealand. Potato 1., 1961/62, 2-5.
CHAPTER 2
Biosystematics of the potato

J.G. Hawkes
2.1 INTRODUCTION
The potato of commerce belongs to a single species, Solanum tuberosum

L., apart from certain cultivated forms in South America which will be
mentioned later and which are generally regarded as distinct. Other well-
known cultivated plants in the genus Solanum are the egg-plant or
aubergine (S. melongena) , the pepino (S. muricatum) and the naranjillo
(S. quitoense). The chilli peppers (Capsicum spp.) and the tomato
(Lycopersicon esculentum) are better known and more widely grown than
the last two Solanum species mentioned above. Less well-known fruits
such as Cyphomandra and Physalis as well as a number of ornamentals
such as Petunia, Schizanthus, Salpiglossis and Lycium also belong to the
Solanaceae. Several genera, such as Nicotiana, Datura, Atropa and
Mandragora, have acquired a more sinister reputation because of their
poisonous alkaloids. Even so, probably all members of the family possess
some alkaloids, though in small amounts in many cases.
2.2 BRIEF OUTLINE OF POTATO CLASSIFICATION
The genus Solanum, to which the cultivated potato belongs, is an extremely

large one, containing about 1000 species. It extends all over the world
except for the far north and south, with a strong concentration of species
diversity in South and Central America on the one hand and Australia on
the other. In this it mirrors to a large extent the distribution pattern of the
family itself.
In addition to S. tuberosum, some six other cultivated species and over
230 wild species of potato are generally recognized. The rest of the genus
consists of non-tuberiferous species, such as S. nigrum (black nightshade)
and many spiny herbs and shrubs. The tuber-bearing species are com-
pletely confined to the American continent and many are of considerable
14 Biosystematics of the potato
interest to potato breeders because of their resistance to pests and
pathogens and their adaptation to climatic extremes.
There have been many attempts at grouping the potatoes in a satis-
factory way within the genus, and some authors have suggested that they
should be separated from it altogether. On the whole, such a course of
action has not received much support since in floral structure, which is
generally considered of prime value in generic distinctions, no species of
Solanum differs essentially from any other.
The tuber-bearing species, once included within the section Tuberarium,
are now grouped under the older name of Petota whiCh seems indeed most
appropriate. They are distinguished by actinomorphic flowers, a jointed or
articulated pedicel or flower stalk, unbranched glandular or eglandular
hairs, no thorns or spines and generally (although not invariably) com-
pound leaves. The typical potato leaf is described botanically as inter-
ruptedly imparipinnate, since the major sequence of paired leaflets and
terminal unpaired one, is broken or interrupted by smaller folioles or
interjected leaflets. The number of pairs of lateral leaflets is often used to
characterize a leaf, which can be spoken of as, for instance, 2-3-jugate,
indicating 2-3 pairs of main laterals.
Although most species in Petota possess underground stolons, which are
of course stems botanically, bearing at their tips the characteristic potato
tubers, some related species that must be grouped with them on the basis
of other morphological characters, lack stolons and tubers altogether
(series luglandifolia and Etuberosa).
Section Petola has been divided for the sake of convenience into two
subsections, namely, ESTOLONIFERA (containing the two non-tuber-
bearing series Etuberosa and luglandifolia); and POTATOE (including 19
other series) as shown below.
The series are here set out in taxonomic sequence and the spedes in each
of them are arranged alphabetically. Species known only from herbarium
specimens or not available from seed collections or gene banks are
omitted. Those included therefore are limited to actual listed and pub-
lished inventories. (For complete species lists reference can be made to
Hawkes (1990) in which details are given also of the seed banks holding
living materials of the species in question.)
In the present work a short description of each species is followed by a
note on its distribution, habitat, chromosome number and endosperm
balance number (EBN), where known. (See Table 2.1.)
2.2.1 Classification of potato species

Section· Petota Dumort
Subsection ESTOLONIFERA Hawkes
Series I ETUBEROSA Juz.
Herbs with very low almost basal pedical articulation. The plants bear no
stolons or tubers, although they are sometimes rhizomatous. Corolla
Table 2.1 Classification and chromosome number of the more important wild
potato species in Subsection Potatoe*
Species arranged according to chromosome number (x = 12)
Series 2x 3x 4x 5x 6x
MorelliformiaS. morelli[orme
II Bulbocastana S. bulbocastanum S. bulbocastanum
S. darum
III Pinnatisecta S. brachistotrichum
S. cardiophyllum S. cardiophyllum
S. jamesii S. jamesii
S. pinnatisectum
S. tarnii
S. trilidum
IV Polyadenia S. polyadenium
S. lesteri
V Commersoniana S. commersonii S. commersonii
VI Circaeifolia S. capsicibaccatum
S. circaei[olium
VII Lignicaulia S. lignicaule
VIII Olmosiana S. olmosense
IX Yungasensa S. arnezii
S. chacoense
S. tarijense
S. yungflsense
X Megistacroloba S. astleyi
S. boliviense
S. megistacrolobum
S. sanctae-rosae
S. toralapanum
• Excluding Subsection Estolonifera, series Etuberosa and Juglandifolia.
Table 2.1 Continued
XI Cuneoalata S. infundibuliforme
XII Conicibaccata S. chomatophilum S. agrimonifolium S. moscopanum
S. santolallae S. colombianum
S. violaceimarmoratum S. oxycarpum
XIII Piurana S. piurae S. tuquerrense
XIV Ingifolia S. ingifolium
XV Maglia S. maglia S. maglia
XVI Tuberosa S. alandiae
(wild) S. berthaultii
S. brevicaule
S. bukasovii
S. canasense
S. gourlayi S. gourlayi
S. hondelmannii
S. kurtzianum
S. leptophyes
S. medians
S. microdontum s. microdontum
S. multidissectum
S. multiinterruptum
S. neocardenasii
S.okadae
S. oplocense S. oplocense S. oplocense
S. sparsipilum
S. spegazzinii S. sucrense
S. vernei
S. verrucosum
Table 2.1 Continued
XVI Tuberosa S. x ajanhuiri S. x chaucha S. x curtilobum
(cultivated) S. phureja S. x juzepczukii
S. stenotomum S. tuberosum
subsp. tuberosum
S. tuberosum
subsp. andigena
XVII Acaulia S. acaule S. albicans
XVIII Longipedicellata S. x vallis-mexici S. fendleri
S. hjertingii
S. papita
S. polytrichon
S. stoloniferum
XIX Demissa S. x semidemissum S. brachycarpum
S. demissum
S. guerreroense
S. hougasii
S. iopetalum
S. schenckii
rotate, purplish or bluish. Central Chile as far south as the island of Chiloe;
Nahuel Huapi region of South Argentina; Islands of Juan Fernandez.
1 S. brevidens Phil. A highly polymorphic species with pubescence
varying from dense to almost absent; pedicel articulation 2-4 mm above
the base; calyx teeth very short (0-0.5 mm); corolla sky-blue, paling to
white at the edges. S. Central Chile, island of Chiloe, Nahuel Huapi region
of S. Argentina. Wet forests from sea level to about 1000 m. 2n=24;
EBN=1.
2 S. etuberosum Lind!. Distinguished by yellowish-green stems and
leaves, with short velvety pubescence and often crisped leaflet margins;
pedicel articulation 4-5 mm above the base; calyx with well marked teeth
(1-1.5 mm); corolla rich purple, very showy. N. Central Chile. Dry
mountain forests, from 1250-2500 m. 2n=24; EBN=1.
3 S. femandizianum Phil. Plant glabrous or glabrescent; pedicel articu-
lation right at the base. Islands of Juan Fernandez. Wet forests, from
100--600 m. 2n=24; EBN=1.
Series II JUGLANDIFOLIA (Rydb.) Hawkes

Plants with herbaceous to woody stems, bright yellow flowers and no
stolons or tubers. They are included with the true potatoes because of the
position of the pedicel articulation at some distance above the base. They
bear obvious resemblances to the genus Lycopersicon, differing from it in
the absence of floral bracts and sterile anther tips. Mountains of Central
America and north-western South America as far south as the coastal
ranges of Peru and north Chile.
1 S. juglandifolium Dun. A perennial woody climber (liane); leaf
pinnate, rugose and hispid above; interjected leaflets few or absent; fruit
up to 4.5 cm diam. Costa Rica, Venezuela, Colombia, Ecuador. Forests,
scrub woodland and hedges at 1000-3000 m. 2n=24.
2 S. lycopersicoides Dun. A bush or shrub, to 2.5 m tall. Leaf bi-
pinnatisect, very similar to that of the tomato. Peru. Open ground, from
2800-3150 m. 2n=24.
3 S. ochranthum Dun. Habit as for S. juglandifolium; leaf pinnate,
smooth and velvety; interjected leaflets very numerous. Very large berries
to 6 cm diam., resembling a green tomato, although much harder.
Colombia, Ecuador, Peru. Forests, scrub woodland and hedges at 1800-
3500 m. 2n=24.
4 S. sitiens Johnston. A herb up to 50 cm tall, glabrous, with bi-
pinnatisect leaves and stellate flowers; pericarp rapidly becoming dry and
papery at maturity. North Chile. Rocky dry ravines at 3000 m. 2n=24.
Subsection POTATOE G. Don.

This subsection contains the true potatoes whose tubers are borne on
underground stolons, which are true stems, not roots.
Brief outline of potato classification 19
Series I MORELLIFORMIA Hawkes
Small tuber-bearing herbs with a strong resemblance to S. nigrum, with
predominantly epiphytic habit, simple leaves, very small white stellate
flowers (less than 1.5 cm diam.), long slender anthers which are slightly
coherent laterally, and small few-seeded berries. Mountain forests of
Mexico and Guatemala.
1 S. morelliforme Bitt. et Muench. A small epiphyte growing on trees
and also on moss-covered walls and rocks in dense shade, never on the
ground. Distinguished by the simple leaves, small stellate flowers, slender
anthers and small few-seeded berries. Central to S. Mexico; Guatemala. In
wet mountain forests from about 2000-3000 m. 2n=24.
Series II BULBOCASTANA (Rydb.) Hawkes

Small tuber-bearing terrestrial herbs with simple leaves, stellate flowers
(1.5-3 cm diam.), short thick anthers and round berries larger than those
of Morelliformia. Central and S. Mexico; Guatemala.
1 S. bulbocastanum Dun. Leaf with rounded to cuneate base, generally
densely pubescent, varying from ovate to linear-lanceolate according to
subspecies. Flowers white to deep cream. Phytophthora resistant. Both
diploid and (apparently) autotriploid forms are known. Central to S.
Mexico; Guatemala. Woods, grassland, rocks and field borders. Often
grows under quite dry conditions. Altitude range 1500-2300 m.
la Subspecies bulbocastanum. Leaf and stem densely pubescent; leaf
ovate in shape, less than 2.5 times as long as broad; corolla white. Central
to S. Mexico. 2n=24 (36); EBN=1.
1b Subspecies dolichophyUum (Bitt.) Hawkes. Leaf linear-Ianceolate,
more than 2.5 times as long as broad, attenuate at base and apex; style
varying in length from 6 to 10 mm. Corolla white. Central Mexico, States
of Morelos and Guerrero. 2n=24; EBN=1.
lc Subspecies partitum (Corr.) Hawkes. Leaf ovate to ovate-lanceolate,
less than 2.5 times as long as broad; leaf and stem less densely pubescent
than in subsp. bulbocastanum; pedical above articulation and calyx
completely glabrous; calyx pale yellow-green; corolla deep cream, the
lobes very deeply separated from each other. Guatemala and the State of
Chiapas in S. Mexico. 2n=24; EBN=1.
2 S. darum Corr. Leaves typically cordate; corolla lobes purple above,
white below; base of anther 3-5 lobed. Guatemala, Mexico. High
mountain forests. 2n=24.
Series III PINNATISECTA (Rydb.) Hawkes

Herbs with stolons and tubers, imparipinnate to imparipinnatisect leaves
and stellate corolla which is white or deep cream coloured, sometimes
tinged with purple; berries globular or conical. United States southwards
to central Mexico.
1 S. brachistotrichum (Bitt.) Rydb. Distinguished from most other
species in this series by the characteristic minute 2-3-celled triangular hairs
on leaves and stem. Leaflets sub-sessile to petiolulate, always slightly
decurrent on to rachis, 2-3(-4)-jugate. Corolla slightly mauve-tinted
towards acumens, very reflexed. N.W. Mexico. Dry pinon and juniper
scrub vegetation at 1750--2500 m. 2n=24; EBN=1.
2 S. cardiophyllum Lindl. Leaflets ovate--cordate to lanceolate, 2-4-
jugate, petiolulate, not decurrent (or only occasionally so). Calyx acumens
0.5-1.5 mm long, according to subspecies. Blight-resistant. N.W. to S.
Mexico, in dry scrub vegetation, field borders, old lava fields, and
especially as weeds of cultivation; the tubers of subsp. ehrenbergii are
edible. Altitude: 1600--2500 m.
2a Subspecies cardiophyllum. Leaflets glabrous, shining, dark green
above; calyx acumens not more than 0.5 mm long; corolla deep cream to
buff-coloured. Anthers short, 5 mm, not attenuate above, often tinged
with violet inside. Both diploid and (apparently) autotriploid forms are
known, the latter being distinguished generally by very much wider
(broadly ovate--cordate) leaflets. Central Mexico. 2n=24 (36); EBN=1.
2b Subspecies ehrenbergii Bitt. Differs from the type subspecies by the
paler green, non-shining, oblong-Ianceolate to lanceolate leaflets and the
frequent presence of hairs of varying lengths on leaves and stems. Calyx
acumens 1-1.5 mm long; anthers narrow, generally attenuate. Petals often
tinged with mauve towards the tips. Mostly diploid forms are known,
although a triploid (presumably autotriploid) has been recorded in San
Luis Potosi State. Central to N.W. and W. Mexico. 2n=24; EBN=1.
2c Subspecies lanceolatum (Berth.) Bitt. This does not differ suf-
ficiently from the diploid forms of subsp. cardiophyllum to be ranked as a
separate species. Distinguished from subsp. cardiophyllum by the
narrower leaflets (more than 2.5 times as long as broad), the longer
peduncle (more than 3.5 cm long) and the 0.5-1.0 mm long calyx acumens.
Corolla white or cream-coloured. Central to S. Mexico. 2n=24; EBN=1.
3 S. hintonii Corr. Leaf 2-3-jugate, with no interjected leaflets. Peduncle
branches rather long. Fruit conical. Mexico, Mexico State, by stone walls,
at 1700 m altitude.
4 S. jamesii Torr. Plants without typical semilunate pseudostipular
leaflets; if present they are pinnatisect and do not clasp the stem. Leaves
slightly decurrent, generally pubescent, occasionally glabrous, with (2)-3-
4--(5) pairs of leaflets and generally no interjected leaflets. Corolla lobes
narrow. N.W. Mexico; S.W. United States; dry scrub vegetation from
1500--2300 m. 2n=24; EBN=1.
5 S. X michoacanum (Bitt.) Rydb. Leaf (2)-3-jugate; corolla white,
stellate; fruit unknown. A natural hybrid of S. pinnatisectum x S.
bulbocastanum. W. Central Mexico amongst rocks and along field borders,
at altitudes of 2000--2100 m. 2n=24.
6 S. nayaritense (Bitt.) Rydb. Distinguished from all other species in this
series by the long weak spreading hairs, pointing downwards on the stem
and inflorescence branches. W. Mexico, States of Nayarit and Zacatecas,
probably native to mountain pine forests.
7 S. pinnatisectum Dun. Plant without semilunate pseudostipular
kaflets, as in S. jamesii. Distinguished from S. jamesii by the glabrous
leaves, ~8 leaflet pairs, generally with a few interjected leaflets and large
showy corolla with broad triangular lobes. Berry round to conical. Some
blight-resistance. Central Mexico. Cultivated fields, waste places and field
borders, from 1800-2100 m. 2n=24; EBN=l.
8 S. x sambucinum Rydb. A natural hybrid between S. pinnatisectum
and S. cardiophyllum subsp. ehrenbergii, which shows segregation to the
parental types when grown from seed. Leaf fairly dark green, with 4-5-
paired leaflets which are narrowly lanceolate and both petiolulate and
decurrent (thus intermediate between the two parents). Corolla with lobes
broader than those of S. cardiophyllum, narrower than those of S.
pinnatisectum. Some blight-resistance. Central Mexico, States of
Queretaro, Guanajuato and Michoacan; a weed of fields and field borders,
found in the distribution areas of the two parents, at about 1800-2000 m.
2n=24; EBN=l.
9 S. stenophyllidium Bitt. Leaflets or leaf lobes sessile, at least six times
as long as broad, sub-linear, attenuate, strongly decurrent on to the rachis.
Hairs as for S. brachistotrichum. Corolla white. W. Mexico, State of
Jalisco, and perhaps elsewhere. In dry grassy places and scrub at about
2000 m. 2n=24.
10 S. tarnii Hawkes et Hjerting. Leaflets narrow lanceolate, petiolulate.
Corolla white; berries globular to ovoid. Central Mexico, States of
Hidalgo, Queretaro and Veracruz at 2360-2650 m in open vegetation.
2n=24.
11 S. trifidum Corr. Leaf 1-3-jugate; stem softly pubescent, provided
with hairs and glands; corolla creamy white, with fairly broad lobes.
Berries long oval, pointed. W. Mexico (States of Michoacan and Jalisco),
in open pine forests, maize fields, roadsides, etc., at altitudes of 2200-2400
m. 2n=24; EBN=l.
Series IV POLYADENIA Buk.

Tuber-bearing herbs with a very dense indumentum of glandular hairs of
an objectionable odour; most of these glands possess 2-celled stalks - a
feature not known in any other tuber-bearing Solanum species. Corolla
white, pentagonal to sub-stellate. Berries ovate, cordate or conical,
somewhat flattened, with black streaks, especially on the edges. Central to
S. Mexico on dry stony hillsides or in damp rain forests, according to the
species.
1 S. lesteri Hawkes et Hjerting. Differs from S. polyadenium in the more
robust habit, thick purple-pigmented stem, dense indumentum of long
spreading multicellular hairs on all green parts, in addition to the glandular
ones, white sub-stellate-pentagonal corolla, and long-conical pointed
2-grooved and flattened berries. A distinct species, obviously related to S.
polyadenium, but occurring under quite different ecological conditions.
Endemic to S. Mexico (Oaxaca State), in damp mountain forests at 2300
m.2n=24.
2 S. polyadenium Greenm. Whole plant covered with very frequent,
stalked glands; stalks generally 2-celled. Very sparse multicellular non-
glandular hairs also present but not easily visible without a lens. Whole
plant more slender and less robust than S. lesteri. Corolla white,sub-
stellate to pentagonal. Insect-resistant and with some blight-resistance.
Central Mexico, from Veracruz to Jalisco; on dry stony hillsides, by old
walls, on old lava and amongst trees and shrubs, from 1900-2000 m.
2n=24.
Series V COMMERSONIANA Buk.

Tuber-bearing herbs with imparipinnate leaves and stellate corolla with
rather broad lobes, generally less than twice as long as broad. Flowers
white or mauve-tinted. South America: Argentina, Paraguay, Uruguay,
Brazil.
1 S. calvescens Bitt. Probably related to S. commersonii, and especially
to subsp. malmeamum, from which it may be distinguished by the larger
size, upright growth, curved leaves, more acute lateral leaflets, large
flowers, and especially by the curious stamens in which the anthers and
filaments are not clearly delimited from each other, either in form or
colour. Brazil, State of Minas Gerais, and perhaps elsewhere; on shady
river banks and in cultivated fields, at altitudes of about 1200 m. 2n=36,
but diploids probably occur also.
2 S. commersonii Dun. Differs from S. chacoense in the sessile obtusely
rounded leaflets at the apex; terminal leaflet typically much larger than the
laterals and the leaf often sub-Iyrate in shape. Peduncle not or once-
branched; branches often very short. Corolla white or often tinted purple
on the external surface; berry cordate or conical. Occurs in both diploid
and (probably) auto triploid forms.
2a Subspecies commersonii. Lateral leaflets decreasing rapidly to base
of leaf, often markedly decurrent, normally sessile; peduncle once-forked,
the branches somewhat contracted; corolla generally purple, the lobes
about 1.5 times as long as broad or even longer. Most plant breeding and
cytological work in the past has utilized this subspecies, often in its triploid
forms. Moderately frost-resistant. Coastal belt of Argentina and Uruguay,
and coastal regions of S. Brazil. Grows in a wide variety of habitats but
very frequently in marshy places, fields, river banks, woods and sandy
shores (dune slacks), from sea level up to about 400 m. 2n=24 (36);
EBN=l.
2b Subspecies malmeanum (Bitt.) Hawkes et Hjerting. Differs from
subsp. commersonii in the lateral leaflets gradually decreasing to base of
leaf, narrowly decurrent and slightly petiolulate; peduncles unbranched or
the branches not markedly contracted; corolla always white, the lobes
about as long as broad. This subspecies is known in diploid and triploid
forms, the latter having been found chiefly in Argentina, provo Misiones.
Like subsp. commersonii, it possesses conical berries and is resistant to
frost. Has a more inland distribution and is known in N.B. Argentina,
Brazil, N. Uruguay and S. Paraguay. Inhabits similar places to those in
which subsp. commersonii is found but seems to prefer shady thickets and
woodlands; it has about the same altitude range as subsp. commersonii.
2n=24 (36); EBN=1.
Series VI CIRCAEIFOLIA Hawkes

Tuber-bearing herbs with small leaves, the terminal leaflet enlarged and
laterals reduced in size. Corolla white, stellate, less than 1.5 cm diam.;
berries narrow-conical. N. Bolivia; hedges and bushy places at high
altitudes.
1 S. capsicibaccatum Card. Distinguished from S. circaeifolium by the
softly hairy leaves, 1-2 pairs of lateral leaflets which are more than half the
length of the terminal, rather pubescent inflorescence and grooved stigma.
Central Bolivia, depts Cochabamba, Santa Cruz, La Paz and Potosi, in
cloud forest and among bushes and scrub vegetation at 2000--4000 m
altitude. 2n=24; EBN=1.
2 S. circaeifolium Bitt. Leaf glabrescent or sparsely pubescent, simple,
or sometimes 1-2(-4)-jugate, the laterals much less than half the length of
the terminal leaflet; stigma simple. Bolivia, depts La Paz and Cochabamba,
in cloud forest and scrub vegetation at altitudes of 250~3900 m.
2a Subspecies circaeifolium. Leaf generally simple, entirely glabrous.
Bolivia, dept La Paz. 2n=24; EBN=1.
2b Subspecies quimense Hawkes et Hjerting. Leaf generally 2-3-
jugate and with long silky hairs. Bolivia, depts La Paz, Cochabamba and
Santa Cruz. 2n=24; EBN=1.
Series VII LIGNICAULIA Hawkes

Plants with cylindrical rather woody stems, covered with a velvety
pubescence. Corolla white, stellate to sub-stellate. Berries sharp-pointed,
ovate-conical. S. Peru.
1 S. lignicaule Vargas. Stem terete, somewhat woody, with velvety
pubescence; leaf softly pubescent, yellow-green. Corolla creamy-white.
Berry ovate-conical. S. Peru, dept Cuzco, on dry bushy slopes and
amongst stones at 300~3500 m. 2n=24; EBN=1.
Series VIII OLMOSIANA Ochoa

Tuber pubescent; leaf 2-jugate with broad irregularly swollen rachis
wing hardly differentiated into leaflets or even lobes; corolla white,
stellate, deeply divided into narrow lobes, hooded at apex; berry ovoid to
pyriform.
1 S. olmosense Ochoa. Characters as for series. Peru, dept Lambayeque
above Olmos, in rain forest at 1800 m. 2n=24.
Series IX YUNGASENSA Corr.

Corolla stellate, white to creamy yellow. Berries sphericaL S. America:
Bolivia, Argentina, Uruguay, Paraguay and Brazil (one species in Peru).
All species basically diploid.
1 S. arnezii Card. Stem mottled purple with conspicuous wings. Corolla
creamy yellow, deeply stellate. Bolivia, depts Chuquisaca and Santa Cruz.
2n=24(?).
2 S. chacoense Bitt. Leaflets petiolulate, acute to acuminate, the
terminal leaflet hardly larger than the laterals; corolla uniformly white;
berries globular. An extremely polymorphic species, spreading from
central Bolivia, southwards into Argentina, Paraguay, Uruguay and S.
Brazil, generally as a field weed in lowland pastures. The great range of
variation has induced many authors to divide it into a large number of
microspecies which cannot now be maintained s,ince they are all apparently
fertile with each other and intergrade considerably. Some forms with insect
and virus Y resistance.
2a Subspecies chacoense. Lateral leaflets 2-3 times as long as broad;
plains forms are almost glabrous, forming a semi-rosette in the early stages
of growth, and also possess very short (not more than 0.75 mm) calyx
acumens. Forms of subsp. chacoense occurring in the mountain valleys of
N. W. Argentina and to central Bolivia are more pubescent, with shorter
petiolules, longer calyx acumens and more developed stem wings. They
rarely form the semi-rosette of basal leaves that is seen in S. chacoense in
the plains. Some of these forms seem to be due to introgression with the
species S. microdontum and possibly certain other mountain species.
Bolivia, Argentina, Paraguay, Uruguay. Waysides, pastures, arable land,
scrub and woodland margins, from sea level to 2350 m. 2n=24; EBN=2.
2b Subspecies muelleri (Bitt.) Hawkes et Hjerting. Differs from subsp.
chacoense by the long narrow leaflets, about 3.3-4.3 times as long as broad,
the long petiolules, up to 25 mm on the acroscopic side, the very oblique
leaflet bases and rather low pedical articulation. Plant generally glabrous,
very occasionally pubescent. Argentina (prov. Misiones), S. Brazil (States
of Rio Grande, Parana, Santa Catarina). Grasslands, river banks, way-
sides, fields, forest margins and clearings, at fairly low altitudes (up to
800m?).
3 S. huancabambense Ochoa. Leaflets oblong-elliptic, obtuse, petiolulate;
calyx with coarse hairs; corolla white, sub-stellate, the lobes about 9 mm
long X 14 mm wide. N. Peru, dept Piura, at 1800--3000 m. Bushy places.
2n=24; EBN =2.
4 S. tarijense Hawkes. Plant with a dense pubescence of simple and
short-stalked glandular hairs, pleasantly aromatic; calyx acumens well-
marked, linear; corolla white, stellate; berry globular, with white raised
spots. Some insect-resistance. Natural hybrids with S. berthaultii in Bolivia
are frequent. Central Bolivia to N. W. Argentina. Scrub and cactus
vegetation in dry inter-an dine valleys at altitudes of about 2000-2800 m.
2n=24; EBN=2.
5 S. yungasense Hawkes. Plant with small (2 cm diam.) creamy yellow
corolla with narrow lobes more than twice as long as broad. N. Bolivia.,
dept La Paz, sub-tropical forests (Yungas region) at 1100-1900 m. 2n=24.
6 S. x trigalense Card. A natural hybrid of S. chacoense x S. tarijense.
Bolivia, dept Santa Cruz, at 2100 m. 2n=24.
7 S. x zudaniense Card. A natural hybrid of S. tarijense X S. berthaultii.
Bolivia, depts Chuquisaca, Cochabamba and Potosi, at about 2000 m.
2n=24.
Series X MEGISTACROLOBA Card. et Hawkes

Rather short-stemmed or straggling tuber-bearing herbs whose leaves bear
a very enlarged terminal leaflet, with the lateral leaflets or leaf lobes much
smaller than the terminal or sometimes completely absent. The laterals
when present are generally broadly decurrent on to the rachis at the
basiscopic side; peduncle often very short; pedicel long, with very high
articulation; corolla sub-stellate to rotate, purple. N. Peru to N.W.
Argentina, growing in waste places, open mountain pastures, etc.
1 S. astleyi Hawkes et Hjerting. Stem slender, erect; leaf narrow-
lanceolate to ovate-lanceolate, generally simple; no interjected leaflets.
Corolla blue-violet, sub-stellate to pentagonal. Bolivia, dept Potosi at
3000-3300 m in fields and waste places. 2n=24; EBN=2.
2 S. boliviense Dun. A rosette or sometimes caulescent species, with
generally completely simple leaves; lateral leaflets when present much
shorter than terminal, shortly petiolulate; corolla rotate, dark purple.
Central Bolivia, depts Chuquisaca and Potosi; dry bushy places and
cultivated fields, from 2600-3750 m. 2n=24.
3 S. chavinense Corr. A species with 1-2 pairs of lateral leaflets, densely
pubescent pedicels and large showy stellate deep purple corolla. N. Peru,
amongst rocks, trees and shrubs at 3500-4200 m.
4 S. hastiforme Corr. Lateral leaflets reduced to minute decurrent lobes
at base of terminal; leaf somewhat hastate; corolla rich purple, rotate-
pentagonal, with well-marked lobes and distinct acumens. N. Peru, along a
bushy rocky stream, at 3200 m. 2n=24.
5 S. megistacrolobum Bitt. Rosette-forming or with straggling stem;
terminal leaflet long ovate to obvate, long-ellipsoid or rhomboid, rounded
or obtuse at apex (or with a minute mucron only); laterals lobed, small or
absent. Leaf smelling of parsley. Globodera and frost-resistant. S. Peru to
N.W. Argentina, high altitude plateaux and mountain slopes, from 2600-
4300 m. 2n=24; EBN =2.
6 S. raphanifolium Card. et Hawkes. Distinguished from S. megistacro-
lobum chiefly by the broadly ovate to orbicular terminal leaflet and the
paler sub-stellate to rotate corolla. S. Peru, waste places, hillsides,
cultivated fields, etc., from 2800-3800 m. 2n=24; EBN=2.
7 S. sanctae-rosae Hawkes. Distinguished from S. megistacrolobum and
S. raphanifolium by the acuminate lateral and terminal leaflets, the small
rosette habit, deep blue-purple corolla, with poorly marked acumens and
large globose stigma. Globodera-resistant. N.W. Argentina, in high
mountain pastures, sandy and rocky places, etc., from 2500-3800 m.
2n=24; EBN =2.
8 S. sogarandinum Ochoa. Very closely related to S. megistacrolobum,
of which it may perhaps represent a northern subspecies; differs chiefly in
the very robust inflorescence and rotate (not sub-stellate) corolla. Some
frost-resistance. N. Peru, paramos (high Andean grassland), at 3550 m.
2n=24; EBN =2.
9 S. toralapanum Card. et Hawkes. Distinguished from S. megistacro-
lobum by the very broadly triangular decurrent rachis wing, or in the
simple-leaved forms by the very long spathulate or long obovate leaf blade,
long common peduncle (6-8 cm), spreading to recurved calyx acumens and
lack of parsley-like leaf odour. Bolivia to N. Argentina, field margins,
rocky and grassy slopes, etc., from 3000-4500 m. 2n=24; EBN=2.
Series XI CUNEOALAT A Hawkes

Small straggling tuberiferous herbs with pinnatifid leaf, the rachis with
narrow wedge-shaped decurrent wings between each pair of leaflets;
corolla purplish, sub-stellate to rotate with well delimited petal acumens.
N. Peru to N.W. Argentina and N. Chile, in dry cactus deserts and scrub.
Possibly drought-resistant.
1 S. infundibuliforme Phil. Leaflets roughly lanceolate, linear-lanceolate
to linear. Interjected leaflets entirely absent. Corolla pale mauve to
purplish, occasionally white, in shape varying from sub-stellate to
completely rotate (circular with well-marked acumens, as in series
Longipedicellata). Possibly drought resistant. Central Bolivia to N. W.
Argentina and N. Chile; dry cactus and scrub deserts at 2450-4100 m.
2n=24; EBN =2.
Series XII CONICIBACCAT A Bitt.

Tuber-bearing herbs with generally well-dissected leaves and acuminate
leaflets, rotate to sub-stellate whitish to generally purple flowers and
ovoid-conical to long conical berries. Mexico, southwards to Bolivia. All
species grow in humid mountain forests and in other regions of high rainfall.
Conicibaccata is one of the three taxonomic series of wild potatoes to be
found both north and south of the Panama isthmus (together with
Juglandifolia and Tuberosa). Many of the species are not very well known
and are difficult to cultivate. The polyploid series of species in Conicibaccata,
with diploid, tetraploid and hexaploid species, is of great theoretical
interest and indicates a complex evolutionary history.
1 S. agrimonifolium Rydb. Berries narrow-conical, grooved, 3-4.5 cm
long. Corolla large, rotate. Leaf with (6)-7-9 pairs of leaflets and
numerous interjected leaflets. S. Mexico to Guatemala. Cloud forests at
about 2000-3300 m. 2n=48; EBN=2.
2 S. chomatophilum Bitt. Leaf 3-4-jugate with 0-4 pairs of small
interjected leaflets; leaflets elliptic or broad lanceolate-elliptic, entirely
glabrous; corolla blue, to 4 cm diam. with short lobes; filaments pubescent
externally; berry ovoid. N. to Central Peru in woods and grassy places,
from 2000-4000 m. 2n=24; EBN =2.
3 S. colombianum Dun. Corolla rotate-pentagonal; berries broadly
ovoid-conical with generally blunt apex, less than 3 cm long; leaves barely
acuminate. A very polymorphic species occurring in Venezuela, Colombia
and Ecuador. Cloud forests at 2200-3500 m. 2n=48; EBN =2.
4 S. flahaultii Bitt. All green parts with long white thick spreading hairs;
leaf with large oval terminal leaflet. Corolla with very flat lobes and small
acumens. Colombia, Ecuador and Venezuela. Bushy places in paramo
vegetation, 2700-3700 m. 2n=48.
5 S. laxissimum Bitt. Leaf 4-6-jugate; corolla pale blue, sub-stellate,
3-3.5 cm diam.; filaments pubescent; berries long conical, 2.5-3 cm long.
S. to central Peru, growing at the edges of cloud forests, from 1900-3600 m.
2n=24; EBN=2.
6 S. longiconicum Bitt. Leaf glabrous, dark green, with acuminate
leaflets; interjected leaflets sparse; corolla rotate; berries narrow conical.
Costa Rica and Panama, at 1000-3150 m in mountain forest and paramo
vegetation. 2n=48.
7 S. moscopanum Hawkes. Corolla with flattened or even slightly
concave lobes, appearing somewhat lO-pointed. Berries and leaves similar
to S. colombianum from which this species was probably in part derived. S.
Colombia. High altitude forests and clearings at 2900-3400 m. 2n=72;
EBN=4.
8 S. neovalenzuelae Lopez. Leaf 2(-3)-jugate, with few or no interjected
leaflets; pedicel articulation very low. N.E. Colombia, 3650 m. 2n=48.
9 S. oxycarpum Schiede. Berries narrow-conical, 3-4 cm long, sharp-
pointed; leaflets pubescent, acuminate; interjected leaflets generally
absent; corolla rotate. East central to southern Mexico. Cloud forests at
about 2000-2200 m. 2n=48; EBN=2.
10 S. paucijugum Bitt. Terminal leaflet much longer and broader than
the 2-3-jugate laterals which decrease rapidly towards base of leaf; corolla
rotate, purple, with rounded lobes and large acumens; berries blunt ovoid.
Ecuador; mountain forests and pastures at altitudes of about 3200-4100 m
and perhaps lower also. 2n=48.
11 S. santolallae Vargas. Leaf sub-glabrous, with apically attenuate
leaflets, glabrous inflorescence branches and calyx, and pale corolla. S. to
central Peru at 2500-3600 m in cloud forests and thickets. 2n=24.
12 S. subpanduratum Ochoa. Terminal leaflet somewhat violin-shaped
to obovate; pubescence delicately silky. Venezuela, in woods at 3400-3600
m.2n=48.
13 S. urubambae Juz. Leaf 2(-3)-jugate, without interjected leaflets,
densely pubescent. Corolla dark purple. S. Peru among stones at 2000-
2700 m. 2n=24.
14 S. violaceimarmoratum Bitt. Terminal leaflet wider, although barely
longer than lateral leaflets; leaflets 3-4-jugate; stem often violet-marbled;
pedicel swelling into calyx base; corolla rotate-pentagonal to sub-stellate.
Berries to 3 cm long. N. Bolivia, in cloud forests and clearings, pathsides,
etc., at 3000-3600 m. 2n=24; EBN=2.
Series XIII PIURANA Hawkes

Tuber-bearing herbs with shining glabrous or glabrescent leaves which
become leathery and with revolute margin when dry; corolla large,
rotate, showy, blue, purple or white; berries ovate, with flattened apex.
Colombia, Ecuador, N. and central Peru, in a wide range of habitats.
1 S. acroglossum Juz. Leaf simple to 1(-2)-jugate; lateral leaflets with
long tongue-like acumens, obtuse at apex. Corolla rotate, dark blue-violet.
Central Peru, amongst bushes at 2700-3000 m. 2n=24.
2 S. albornozii Corr. Leaf surface dark green above, paler below.
Corolla white, with lilac streak on the back of each petal, rotate. Berry
ovoid. S. Ecuador, amongst clearings at 3100 m. 2n=24.
3 S. hypacrarthrum Bitt. Leaf 0-2-jugate with very large terminal and
minute laterals, if present; terminal apically cuspidate; articulation very
high. Corolla white. Berry more or less globose. Central Peru amongst
grasses and bushes at 1800-2800 m. 2n=24.
4 S. jalcae Ochoa. Leaf 2-3-jugate with winged rachis and larger
terminal leaflet; leaf pigmented purple below; calyx dark purple; corolla
blue, with short lobes and very small acumens. N. Peru, growing on
paramos and high alpine pastures from 3100-3500 m. Frost-resistant.
5 S. paucissectum Ochoa. Terminal leaflet much larger than the 0-2-
paired laterals; corolla white with lilac streaks or lilac acumens, 2.5-3 cm
diam. N. Peru, dept Piura, in open places and amongst trees, at 2900-3200
m. 2n=24; EBN =2.
6 S. piurae Bitt. Leaf 2-3(-4)-jugate; 1-4 pairs of interjected leaflets;
leaflets lanceolate; pedicels 3-5.5 cm long; corolla purple, 2-4 cm diam.;
filaments glabrous. Berry ovate to cordate. N. Peru, rocky slopes amongst
shrubs and grasses at 2000-2900 m. 2n=24.
7 S. solisii Hawkes. Leaf 1-2-jugate, with no interjected leaflets. Corolla
rotate, with very short lobes, to 3.5 cm diam., purple. S. Ecuador, at 3400-
3600 m in damp grassy and bushy places.
8 S. tuquerrense Hawkes. Leaf 4-{i (-lO)-jugate, glabrous or with some long
spreading hairs; corolla blue, 2-3.5 cm diam.; filaments glabrous. Unique
amongst tetraploids in being self-incompatible; some frost-resistance. Colombia,
Ecuador, on paramos and amongst shrubs at 2600-3300 m. 2n=48; EBN=2.
Series XIV INGIFOLIA Ochoa
Tuber-bearing herbs with erect or decumbent habit, thick winged stems,
imparipinnate leaves with broadly winged rachis, the wing continuing right
to the leaf base, and rotate corolla. Andes of N. Peru.
1 S. ingifolium Ochoa. A very curious and distinct species with leaves
similar to those of the Leguminous genus Inga. Leaf rachis with broad
wings which swell out from the point of insertion of each leaflet pair and
become slightly narrowed towards the next pair below but continuing to
leaf base; no interjected leaflets; corolla lilac, rotate. Endemic to the
humid mountain forests of N. Peru (Piura) at altitudes of 2800-3000 m.
(Note: the other species in this series, S. raquialatum Ochoa, is not

included in any of the published seed lists.)
Series XV MAGLIA Bitt.

Distinguished from all other series by the loose barrel-shaped anther
column, and the anthers and filaments not well demarcated from each
other, either in colour or form. Distribution: Central Chile and W.
Argentina.
1 S. maglia Schlechtd. A large bushy plant with (1-)2-3-paired broadly
ovate leaflets, and broader terminal leaflet. Corolla white, pentagonal to
rotate. Stamens very characteristic, in a loose barrel-shaped column;
anthers and filaments not well demarcated either in colour or form. Central
Chile, in the region of Valparaiso, near the sea coast to 120 m; Argentina,
provo Mendoza, in dry valleys at 1500 m. 2n=24, 36.
Series XVI TUBEROSA (Rydb.) Hawkes

This series contains all the cultivated potato species as well as the wild and
weed species most closely related to them. It is characterized by impari-
pinnate or simple leaves, forked peduncle, rotate to pentagonal corolla and
round berries. Series Tuberosa is found in the Andes of S. America and the
adjacent coastal belt in temperate and sub-tropical latitudes. One species,
S. verrucosum, occurs in Mexico.
The wild species will be dealt with first, indicating where appropriate
their possible connections with the cultivated ones.
Wild species
1 S. abacayense Ochoa. Leaf 4-jugate, with 12 or more pairs of
interjected leaflets. Stem, inflorescence branches and calyx with long
spreading hairs. Corolla rotate, light purple, quite large. Peru, dept
Apurimac, in humid shrub vegetation at 2900-3600 m. 2n=24; EBN=2.
2 S. acroscopicum Ochoa. Distinguished by the completely glabrous
leaf, narrow lanceolate petiolulate leaflets, many interjected leaflets (some
acroscopic), and rich purple large showy corolla; berry spherical to slightly
ovoid. S. Peru, amongst bushes at 3450 m. 2n=24.
3 S. alandiae Card. Lateral leaflets 2-3-paired, decreasing rapidly in size
from apex to base of leaf; no interjected leaflets; leaves with dense short
hairs. Corolla rotate-pentagonal, pale to medium blue-violet. Bolivia,
depts Cochabamba and Santa Cruz, amongst bushes on the high slopes of
the Andes, at 2300-3000 m. 2n=24.
4 S. ambosinum Ochoa. Leaf 4--5-jugate with up to 8 pairs of interjected
leaflets and short hairs on all green parts; calyx acumens long, 1.5-3 mm;
corolla purple, rotate. Central Peru, on shrubby mountain slopes at 2200-
2500 m. 2n=24; EBN=2.
5 S. andreanum Baker. Characterized by 2-3 pairs of acuminate
decurrent leaflets and purple rotate corolla. S. Colombia, N. Ecuador.
Damp high altitude woods and bushy places at 2100-2900 m. 2n=24.
6 S. avilesii Hawkes et Hjerting. Leaf 2-3-jugate, with few interjected
leaflets and terminal larger than laterals. Corolla rotate-pentagonal, deep
violet-purple. Bolivia, dept Santa Cruz, at 2700-2950 m among shrubs and
degraded forest. 2n=24.
7 S. berthaultii Hawkes. Very insect-resistant species with two types of
glandular hairs, 4--5-jugate leaf and many interjecteds; corolla pale violet-
blue, pentagonal to sub-stellate. Bolivia, eastern slopes of Andes in rather
dry valleys amongst bushes and in waste places from 2400-2750 m. 2n=24;
EBN=2.
8 S. brevicaule Bitt. A low-growing bushy species with 3-4-jugate lateral
and many interjected leaflets. Corolla deep rich violet-purple. Central
Bolivia, dept Cochabamba, amongst grass and shrubs, at 3000-3850 m.
2n=24; EBN =2.
9 S. bukasovii Juz. Low semi-rosette plants with delicate growth; 3-4-
jugate leaves and up to 9-10 pairs of interjected leaflets; pubescent on all
green parts. Flowers purple, rotate. Possibly related to S. multidissectum.
Frost-resistant. Central Peru, in the puna formation (high mountain
pastures) at 3300-4300 m. 2n=24; EBN=2.
10 S. cajamarquense Ochoa. A very distinctive species, the whole plant
possessing a dense coarse pubescence; leaf rough, 3-jugate, with undulate-
crenulate margins and few lateral leaflet pairs (2-3) which are generally
smaller than the terminals; interjected leaflets numerous; corolla white. N.
Peru, dept Cajamarca, on mountain slopes at about 2600 m. 2n=24(?).
11 S. canasense Hawkes. Distinguished by the large rotate corolla, highly
dissected leaves and the soft appressed silky pubescence. Leaflets narrow
lanceolate; interjected leaflets numerous. Flowers large, blue-purple.
Some frost-resistance. S. Peru. Dry stony places, often amongst bushes, at
2900-4100 m. 2n=24; EBN=2.
12 S. candolleanum Berth. Leaf highly dissected, with 5-6 pairs of
narrow lateral leaflets and numerous interjected leaflets; pubescence of
very dense long white appressed hairs. Corolla purple, rotate. N. Bolivia,
dept La Paz, on rocky mountain slopes, at 3500-4000 m. 2n=24.
13 S. chancayense Ochoa. Small glabrous plants with 2-3-jugate lateral
leaflets and no interjected leaflets; corolla white, rotate, with very short
lobes and small acumens. Peru; coastal hills (lomas) of Lima and La
Libertad departments, from 150-550 m. 2n=24; EBN=l.
14 S. chiquidenum Ochoa. Leaves with 1-2 pairs of lateral leaflets,
decreasing in size rapidly towards leaf base; no interjected leaflets; coarse
shining hairs on both surfaces; calyx sub-glabrous, with linear acumens;
corolla white, tinged with lavender, rotate. N. Peru, amongst bushes and in
grass, at 2500-3350 m. 2n=24.
15 S. coelestipetalum Vargas. Leaf 3-4-jugate, with frequent interjected
leaflets; lower surface densely velvety-pubescent; corolla bright sky-blue.
S. Peru, endemic to the Urubamba valley, amongst dry sparse shrubs, at
2400-3600 m. 2n=24.
16 S. x doddsii Corr. Leaf 3-5-jugate, glabrescent, with petiolulate
laterals and very few interjecteds. Rest of green parts also almost glabrous;
corolla sub-stellate, light lavender. This 'species' is a casual hybrid of S.
chacoense with the blue-flowered Bolivian species S. alandiae. Bolivia,
Cochabamba, on a rocky wooded slope at 2600 m. 2n=24; EBN =2.
17 S. gandarillasii Card. A very distinct species, entirely glabrous, with
enlarged terminal and few laterals rapidly decreasing below; flowers small;
calyx with oblong-spathulate, leafy lobes; corolla white. Central Bolivia,
eastern Andes, in dry cactus and scrub region from 1800-2500 m. 2n=24;
EBN=2.
18 S. gourlayi Hawkes. Small plants with 3-4-jugate ovate-Ianceolate to
oblong-lanceolate leaflets which are often slightly decurrent; terminal
leaflet often broader than laterals; flowers pale purple, pentagonal. N. W.
Argentina to central Bolivia. Dry hillsides amongst cactus and scrub
vegetation. 2n=24, 48.
18a Subspecies gourlayi. Terminal leaflet larger than the 3-paired
laterals. Argentina, provo Jujuy at 1900-3600 m. 2n=24; EBN=2. 2n=48;
EBN=4.
18b Subspecies saltense Clausen et Okada. Terminal leaflet the same
size as the 4-paired laterals. Argentina, provo Salta at 3300-3900 m.
2n=24; EBN =2.
18c Subspecies vidaurrei (Card.) Hawkes et Hjerting. Leaflets narrow,
marginally toothed; leaf hairs long and silky. N. W. Argentina and S.
Bolivia, at 2700-3450 m. 2n=24; EBN =2.
18d Subspecies pachytrichum (Hawkes) Hawkes et Hjerting. Leaflets
narrow; leaf hairs short and thick. Central Bolivia. 2n=24.
19 S. hondelmannii Hawkes et Hjerting. Leaf 4-5-jugate with few inter-
jected leaflets and long (5-20 mm) petiolules. Corolla rotate to pentagonal,
blue-violet. Stigma larger than style apex. Related to S. oplocense. Central
Bolivia at 2600-2700 m in dry areas with thorn bushes. 2n=24.
20 S. hoopesii Hawkes et Okada. Leaf 3-jugate with few interjected
leaflets; terminal leaflet broader than laterals. Corolla showy, rich purple.
Bolivia, dept Chuquisaca at 2500-3200 m. 2n=48.
21 S. humectophilum Ochoa. Leaf 2-3-jugate, with few interjected
leaflets, densely pubescent above; terminal larger than laterals. Corolla
rotate, pale-lilac with deep purple star and narrow acumens. Peru, dept
Amazonas, in sub-tropical mountain rain forest at 2875 m.
22 S. immite Dun. Leaflets lanceolate, petiolulate, 4-5-jugate, with few
minute interjecteds; calyx glabrescent; corolla large, white, rotate, very
showy. Central to N. Peru, on stony hillsides, from the coastal lomas
(about 200 m) to 2500 m. 2n=24.
23 S. incamayoense Okada et Clausen. Leaf (2-)3(-4)-jugate with few
interjected leaflets, all thick and fleshy, sparsely hairy. Corolla light blue,
pentagonal. N.W. Argentina, at 2100---2700 m on stony slopes. 2n=24.
24 S. kurtzianum Bitt. et Wittm. Distinguished by the elliptic to oblong
obtuse leaflets with well-marked marginal hairs set on cushions of tissue
and visible without a lens; also by the low pedicel articulation placed in the
lower third. Flowers rotate to sub-stellate, white or with central petal
streak of bright violet on external surface only. Globodera-resistant and
possibly drought-tolerant. W. to N.W. Argentina on dry bushy hillsides,
etc., from 1400---2500 m. 2n=24; EBN=2.
25 S. leptophyes Bitt. A low-growing slender species with 4-jugate
narrow-Ianceolate leaflets 3-4 times as long as broad. Leaf hairs quite
frequent, coarse, subappressed. Corolla pentagonal to rotate, purple. S.
Peru, N. Bolivia. Dry open places or amongst scrub from 3200---3950 m.
2n=24; EBN=2.
26 S. marinasense Vargas. Distinguished by the leaves which are bright
glabrescent and shining above, paler and dull, with densely felted short
hairs below; 2-3 pairs of broadly lanceolate leaflets. Flowers blue, very
showy. A very distinct species, perhaps related to series Piurana. S. Peru,
bushy places and grassy hillsides from 2200---3500 m. 2n=24; EBN=2.
27 S. medians Bitt. Low-growing straggling plants with enlarged terminal
and smaller, often decurrent, lateral leaflets in 1-2 pairs with sometimes 1-
2 pairs of interjected leaflets. Distinguished from S. microdontum by the
rhomboid terminal leaflet and rich violet purple flowers. Central Peru,
coastal belt and mountains, in scrub vegetation, from 200---3400 m. 2n=24
(36); EBN=2.
28 S. microdontum Bitt. Large plants with winged stems and large simple
to 2(-3)-jugate leaves, the terminal generally much larger than the laterals;
calyx acumens linear, unequal; corolla white, pentagonal to rotate;
pubescence coarse, shining, of frequent multicellular hairs.
28a Subspecies microdontum. More slender than subsp. gigantophyllum,
with narrow straight stem wings (0---2 mm wide) and terminal leaflets rarely
more than 8 cm long. N. to S. Bolivia and N.W. Argentina, in humid
forests, etc., at 1800---3100 m. 2n=24; EBN =2.
28b Subspecies gigantophylium (Bitt.) Hawkes et Hjerting. Stem
more robust, 3-20 mm diam.; wings 2-5 mm wide, undulate and crenulate
marginally; terminal leaflet 8-18 cm long. Central Bolivia and N.W.
Argentina, southwards to La Rioja province. High mountain rain forests
and amongst shrubs on the eastern slopes of the Andes, from 1800-3000 m.
2n=24 (36); EBN=2.
29 S. mochiquense Ochoa. Leaflets 3-4(-5)-paired, sessile to petiolulate;
3-12 pairs of interjected leaflets; hairs quite dense below, very few above;
leaf margins undulate, crenulate; corolla rotate, white, showy. N. Peru, on
the lomas (low hills) by the coast, at 250-500 m and in the mountains at
1650-1700 m on rocky hillsides. 2n=24; EBN=l.
30 S. multidissectum Hawkes. Low straggling herbs with generally highly
dissected leaf (6-8-jugate) and numerous interjected leaflets. Leaflets
broadly ovate, the terminal sub-rotund. Long spreading hairs occur on the
stem, rachis and petiolules. Flower fairly large, pale blue. Differs from S.
leptophyes and S. canasense in the wider leaflets and long spreading hairs.
Some resistance to frost and Globodera. S. Peru in high Andean pastures,
amongst stones, by paths, etc., at altitudes of 3100-4175 m. 2n=24;
EBN=2.
31 S. multiinterruptum Bitt. Leaflets sessile to sub-sessile; 4-5 pairs of
laterals which are about twice as long as broad, 5-15 pairs of interjected
leaflets. Peduncle very long. Corolla very large, up to 4.5 cm diam., dark
blue to violet. Central Peru, in high mountain grassland, rocky slopes and
among bushes, etc., at 3300-4000 m. 2n=24; EBN=2.
32 S. neocardenasii Hawkes et Hjerting. Highly insect-resistant. Leaf
delicate, narrow, with ovate-cordate leaflets and 12-20 mm petiolules.
Interjected leaflets few to none. All green parts densely covered with short
and long glandular hairs. Corolla rotate, white. Bolivia, dept Santa Cruz,
at 1400 m in xeric environment. 2n=24.
33 S. neorossii Hawkes et Hjerting. A low-growing herb with small 1-3-
jugate leaves and few interjected leaflets; terminal larger than laterals;
pubescence medium dense, velvety. Corolla rotate, showy, violet-purple,
to 3.5 cm diam. N. Argentina, provs Jujuy and Salta, at 2550-3400 m
amongst grasses, shrubs and rocks. 2n=24.
34 S. okadae Hawkes et Hjerting. Leaf 1-2-jugate, generally without
interjected leaflets; hairs sparse to frequent, short. Calyx acumens linear to
spathulate. Corolla white, pentagonal to rotate, to 3.5 cm diam. N. Bolivia
to N. Argentina, at 2600-3200 m, generally in high mountain rain forest.
2n=24.
35 S. oplocense Hawkes. Leaf 3-4-jugate with petiolulate but decurrent
leaflets and denticulate margin; corolla violet to purple, sub-stellate to
pentagonal; anthers strongly tapered from base to apex, often 4-lobed at
base; style long, curved, exserted 5--6 mm; stigma minute, not thicker than
style apex. S. Bolivia, N.W. Argentina on dry hillsides amongst stones, in
cactus and scrub vegetation, at altitudes of 2600-3500 m. Little is yet
known about the nature of the polyploid races of this species. Resistant to
Globodera. 2n=24; EBN unknown; 2n=48, 72; EBN=4.
36 S. pampasense Hawkes. Leaflets lanceolate, covered with hairs of
varying lengths, some gland-tipped; calyx acumens linear, 2-5.5 mm long;
corolla pale purple to pale blue, showy; anthers long and narrow. Central
to S. Peru (depts Ayacucho, Apurfmac and Cuzco) in dry sub-tropical
interandine valleys at 2000-2900 m. 2n=24; EBN=2.
37 S. sandemanii Hawkes. Leaf imparipinnatisect; laterals 2-3-jugate,
the upper pair widely decurrent; no interjecteds; calyx lobes often sub-
spathulate, very pubescent; corolla showy, bright violet, rotate, 3-3.5 cm
diam.; seems to be related to S. weberbaueri (including S. tacnaense). S.
Peru, dept Arequipa, at 2600-3100 m in sandy soil among bushes. 2n=24;
EBN=2.
38 S. x setulosistylum Bitt. Vegetatively intermediate between S.
chacoense and S. spegazzinii, but with corolla varying from white-stellate
to mauve-stellate or sub-stellate. Style setae sometimes present. Leaf often
highly dissected, with numerous narrow-ovate interjected leaflets. Young
fruits sometimes puberulent. This name has been given to hybrid popu-
lations derived apparently from natural crosses between S. chacoense and
S. spegazzinii. N. W. Argentina in scrub and dry stony lands, waysides,
etc., at 1600-2100 m. 2n=24.
39 S. sparsipilum (Bitt.) Juz. et Buk. A very polymorphic species found
as a weed of cultivated fields and waste places, morphologically very
similar to S. tuberosum subsp. andigena (see below). Distinguished from
that species, however, by the sparse short hairs, smaller more straggling
habit, 2-4-jugate leaves with fewer interjected leaflets and rather smaller
flower. Its wide distribution may be due to the fact that it was carri,ed by
man as a weed with cultivated potatoes. S. sparsipilum seems to have
played an important role in the evolution of S. tuberosum. S. Peru,
southwards to N. Bolivia. Cultivated fields, waysides and waste places,
from about 2400-3800 m. 2n=24; EBN=2.
39a Subspecies sparsipilum. Leaflets narrow ovate, interjected leaflets
frequent. Distribution as for species description.
39b Subspecies calcense Hawkes. Leaflets broad ovate, interjected
leaflets 0-1-paired. Peru, dept Cuzco at 2800-3000 m, as a field weed.
40 S. spegazzinii Bitt. A widespread and phenotypically highly variable
species; leaflets 4-7-jugate, long-elliptic, narrowing to each end, often
decurrent; interjected leaflets narrow, 2-7(-18)-paired. Corolla medium
purple to lilac, sub-stellate to pentagonal. Globodera-resistant. N. W.
Argentina, in dry places in the interandine valleys and basins from 1900-
3100 m. 2n=24; EBN=2.
41 S. x sucrense Hawkes. Leaf 4-5-jugate, with 1-3(-6) pairs of
interjected leaflets; pubescence of sparse short appressed hairs. Corolla
rotate to pentagonal, 2.5-3.5 cm diam., with long acumen. This is a
somewhat fixed natural hybrid of S. oplocense X S. tuberosum subsp.
andigena, growing chiefly as a weed of cultivation and in xeric vegetation
from 2600-3900 m. 2n=48; EBN=4. Globodera-resistant.
42 S. ugentii Hawkes et Okada. Leaf 5-6(-7)-jugate, with up to 20 pairs
of interjected leaflets, including acroscopics and basiscopics; petiolules
5-15(-25) mm long. Corolla deep rich purple, rotate. Stigma large,
capitate. Bolivia, dept Chuquisaca at about 3750 m, amongst rocks.
2n=48.
43 S. venturii Hawkes et Hjerting. Leaf with large terminal and smaller
1-3-paired laterals, occasionally simple, with very sparse closely appressed
short 3-celled triangular hairs, visible under a lens only; calyx glabrescent
or with a few hairs of same type as leaf hairs; corolla white, sub-stellate to
rotate. Related to S. microdontum but easily distinguished by the very
characteristic pubescence. Globodera-resistant. Endemic to N. W.
Argentina in high altitude grasslands (pajonales), at 2000-2800 m. 2n=24
(36); EBN=2.
44 S. vernei Bitt. et Wittm. A tall robust plant with large leaves and 4-{)
pairs of ovate or ovate-lanceolate leaflets covered below with a whitish
cottony or woolly pubescence. Flowers deep purple, pentagonal, often
very large, to 4.5 cm diam. Shows strong resistance to Globodera.
44a Subspecies vernei. Distinguished from subsp. ballsii by the apically
acuminate petiolulate leaflets, acute ovate interjected leaflets and larger
corolla. Argentina, provs Catamarca and Tucuman at 2200-2800 m in
mountain forests. 2n=24; EBN=2.
44b Subspecies ballsii (Hawkes) Hawkes et Hjerting. Leaflets gene-
rally sessile, acute to obtuse apically; interjected leaflets generally shorter,
obtuse and sessile; corolla to 3 cm diam. Argentina, provs Jujuy and Salta
at 2700-3450 m in high rainfall forests and open vegetation. 2n=24;
EBN=2.
45 S. verrucosum Schlechtd. Distinguished by the upright habit,
petiolulate leaflets, terminal larger than laterals, well-defined rounded
corolla lobes whose margins roll inwards, and white-verrucose berry. Is
very probably an ancestral form of all series Demissa species, contributing the
one genome that they seem to possess in common, and was formerly included
in that series. It is the only series Tuberosa species which occurs north of the
Panama isthmus. Shows some blight-resistance. North-east, central and
southern Mexico in pine and fir forests at 2400-3200 m. 2n=24; EBN=2.
46 S. virgultorum (Bitt.) Card. et Hawkes. Leaf 2-3-jugate, with
few interjected leaflets; terminal larger than laterals, apically abruptly
acuminate; pubescence of dense short hairs. Corolla rotate, violet-purple,
with rather flat lobes. N. Bolivia, at 2800-3900 m in high altitude rain
forest and amongst bushes. 2n=24.
47 S. weberbaueri Bitt. Leaf 3-4-jugate, with 0(-2) pairs of interjected
leaflets; laterals abruptly decurrent, the terminal much larger. Corolla
rotate, violet, 3.5-4.0 cm diam. Differs from S. sandemanii in the short
leaflet de currency and larger leaflets, and from S. tacnaense (not included
in this review) in the less conspicuous leaf hairs and marginal teeth. S.
Peru, amongst shrubs and rocks, and on stony hillsides, in the coastal
lomas and higher from 500-600 m. 2n=24.
Cultivated species
A. Diploids
1 S. x ajanhuiri Juz. et Buk. This species is similar in many respects to S.
stenotomum. It differs, however, in the small regular calyx, smaller blue
flower, very high pedicel articulation, and stiff leaves. Agrees with S.
stenotomum in the decurrent bases of the uppermost leaflet pair and in the
form of corolla. It was formed as a natural hybrid of S. stenotomum with S.
megistacrolobum. Frost-resistant. S. Peru and N. Bolivia, at high altitudes.
2n=24.
2 S. phureja Juz. et Buk. Dis6nguished by the sparsely pubescent leaf,
which is shining in the living state, and rather irregular calyx with
lanceolate lobes. Tubers yield in 3-4 months under short day conditions
and possess no dormancy period. The absence of tuber dormancy indicates
that it has become specially adapted to regions that are free from frost.
Venezuela, Colombia, Ecuador, Peru and N. Bolivia. Wet mountain
slopes mostly in eastern Andes, at lower altitudes than the other cultivated
species, but can also be grown at higher altitudes. 2n=24; EBN =2.
3 S. stenotomum Juz. et Buk. Distinguished from S. phureja by the more
densely pubescent leaf which is not shining in the living state, tubers
produced in 5-6 months or longer and with definite dormancy period.
Calyx generally irregular with lanceolate lobes as in S. phureja. A very
variable species which is possibly ancestral to all the other cultivated
potatoes (see Section 2.11). Central Peru to central Bolivia, at very high
altitudes. Some forms are frost-resistant. 2n=24; EBN=2.
Note: The former species, S. goniocalyx, has now been placed as a

subspecies of S. stenotomum. It occurs mainly 10 central Peru, but
intergrades into subsp. stenotomum further south.
B. Trip 10 ids
4 S. x chaucha Juz. et Buk. All those triploid forms that have been
derived from natural crosses between S. tuberosum subsp. andigena and S.
stenotomum are grouped here. Some of these, which are most distinct and
more widely cultivated, were formerly classed by Juzepczuk and Bukasov
(1929) and by myself (Hawkes 1944) as separate species. Many more
collections of triploid cultivated potatoes have been made, each of which
differs in certain points from the micro-species already described. This is
only to be expected when we consider that triploid hybrids could have been
formed many times by the crossing of different clones of the two very
polymorphic species S. stenotomum and S. tuberosum subsp. andigena. I
have retained the name S. x chaucha since it was the first to be applied to
these triploid forms by Juzepczuk and Bukasov. The best way of distin-
guishing S. x chaucha from other cultivated species is by the corolla lobes
which are in general about three times as broad as long when spread out
flat. Central Peru to central Bolivia at high altitudes. 2n=36.
5 S. x juzepczukii Buk. Distinguished by the semi-rosette habit, long
straight leaves, short peduncle (2--4 cm long), pedicels with very high
articulation which is indistinct, and small blue corolla (to 2.5 cm diam.)
with very short lobes and small acumens.
A natural triploid hybrid between S. acaule and S. stenotomum. Central
Peru, southwards to S. Bolivia at very high altitudes. Frost-resistant.
2n=36.
C. Tetraploids
6 S. tuberosum L. Distinguished from other species of cultivated potato
by the pedicel articulation placed in the middle third, short calyx lobes
arranged regularly, leaves often slightly arched, leaflets always ovate to
ovate-lanceQlate, about twice as long as broad, never narrow lanceolate as
in some forms of S. stenotomum and S. phureja. Corolla lobes about half as
long as broad. Tubers with well-marked dormancy period.
Two subspecies are now recognized:
6a Subspecies tuberosum. Originally only from the coastal regions of
South Central Chile (Island of Chiloe and adjacent mainland). Is distin-
guished from subsp. andigena by the less dissected leaves with wider
leaflets, generally arched and set at a wider angle to stem. Pedicel
thickened above; corolla often white or pale coloured. Tubers formed
under long days or under short days in the tropics at lower altitudes only
(500-2000 m). Cultivated primitively on the southern coast and islands of
southern Chile (Chiloe region), now worldwide. 2n=48; EBN=4.
6b Subspecies andigena Hawkes. This subspecies may be distinguished
by the narrower, more numerous leaflets, which are generally petiolulate,
the leaves set at an acute angle to the stem and generally more dissected;
pedicel not thickened at apex; tubers formed at high altitudes only (over
2000 m) under short day conditions. This is undoubtedly the ancestral
subspecies of S. tuberosum. Andes of Venezuela, Colombia, Ecuador,
Peru, Bolivia, N.W. Argentina; also sparingly in Guatemala and Mexico.
2n=48; EBN =4.
Escaped forms of both subspecies have been described under the names
of S. apurimacense, S. molinae, S. leptostigma, S. diemii, etc. They differ
only in the longer stolons and sometimes unpigmented tubers from the
cultivated varieties.
D. Pentaploids
7 S. x curtilobum Juz et Buk. Distinguished by the semi-rosette habit,
straight stiff leaves, very high pedicel articulation and large purple circular
corolla 30-35 mm diam. with very short lobes and acumens. Derived from
natural crosses between S. X juzepczukii and S. tuberosum subsp.
andigena. Several variations in tuber colour and form are known. Central
Peru to S. Bolivia at very high altitudes. Frost-resistant. 2n=60.
Series XVII ACAULIA Juz.
Low rosette-forming herbs (occasionally forming long stems), bearing
stolons and tubers; leaves with typically obtuse leaflets auricled at the base
on the acroscopic side; peduncle very short or absent; pedicel articulation
absent or shown only by a ring of pigment, very rarely well-marked; corolla
small, rotate, with very short lobes. Peru, Bolivia, and N.W. Argentina at
very high altitudes in alpine meadows, field borders, etc.
1 S. acaule Bitt. Characters and distribution as for series. Highly
resistant to frost. The fairly wide intraspecific variation has caused certain
authors to split S. acaule into several microspecies. The fertility and range
of variability between all forms so far studied makes it advisable, however,
to reunite all but one of them into the one original species, divided into
geographical subspecies as follows:
la Subspecies acaule. Lateral leaflets not much shorter than terminal
leaflets, not or only slightly decurrent; pedicel articulation generally
invisible or marked by a difference of colour only; pubescence of fairly
short crisped hairs. Peru to Bolivia and N.W. Argentina, from 2600 to 4650
m, in alpine meadows, by paths, walls, drainage ditches, cultivated fields,
etc. 2n=48; EBN =2.
lb Subspecies aemulans (Bitt. et Wittm.) Hawkes et Hjerting. Differs
from subsp. acaule in the short leaf with much enlarged terminal and 0-4
pairs of decurrent laterals; pedicel articulation well-marked but not
abscissing when mature. Pubescence as for subsp. acaule. N.W. Argentina,
Sierra de Famatina, from 2950 to 4000 m in similar habitats to subsp.
acaule. 2n=48; EBN=2.
lc Subspecies punae (Juz). Hawkes et Bjerting. Leaf like that of
subsp. acaule, but stem very short; pubescence of long weak spreading
hairs; articulation of pedicel never visible except by a change of colour,
observable on living plant only. Central Peru at similar altitudes and
habitats to subsp. acaule. 2n=48; (60); EBN=2.
2 S. albicans (Ochoa) Ochoa. Habit robust, rosetted to semi-rosetted.
Leaf 3-4-jugate with 3-5 pairs of interjected leaflets. All green parts with
long weak hairs. Peduncle forked. Pedicel with clearly marked articulation
in upper third but much lower than in S. acaule. Corolla very pale blue to
white; style exserted 1.5-2.0 mm. Berries not separating at articulation. N.
Peru, depts Ancash, Cajamarca and La Libertad at 3500 m. Andean
grassland region. Probably an amphiploid hybrid of S. acaule subsp. punae
with a diploid species from another series, possibly S. cajamarquense or
some similar species. 2n=72; EBN =4.
Series XVIII LONGIPEDICELLATA Buk.

Herbs with long creeping stolons; leaves with coarse white hairs, or
glabrous. The arched corolla lobes and large acumens give the corolla a
circular appearance with acumens standing out sharply from it. Corolla
occasionally, however, sub-stellate. Central Mexico to S.W. United States
on dry plateaux and mountain slopes, medium altitudes. Most species are
tetraploid.
1 S. fendleri A. Gray. Plant with rather dense coarse hairs on leaf, stem,
peduncles, pedicles and calyx; lateral leaflets sessile to sub-sessile, often
decurrent; all leaflets obtuse; corolla deep purple. Some virus resistance.
S. fendleri grows chiefly under the shade of Pinus ponderosa and other
trees in damp leaf mould at altitudes of 1700-2850 m. 2n = 48; EBN=2.
1a Subspecies fendleri. Corolla acumens large, curving gradually into
the lobes; corolla semi-stellate in outline. USA, States of Arizona,
Colorado, New Mexico and Texas; Mexico, States of Baja California,
Chihuahua and Sonora; at 1600-2800 m in oak-pine forests but not under
dense shade. 2n=48; EBN=2.
1b Subspecies arizonicum Hawkes. Differs from subsp. fendleri by the
rotate corolla with rather small acumens and short lobes, and the small, 3
mm long anthers. USA, Arizona State; Mexico, Chihuahua State in forest
clearings, etc., from about 2000 to 2500 m. 2n=48; EBN=2.
2 S. hjertingii Hawkes. Differs from S. fendleri in the glabrous or
glabrescent stems, leaves, peduncles, pedicels and calyx, in the generally
petiolulate narrow leaflets and the glabrous corolla. Vegetatively ex-
tremely similar to S. cardiophyllum subsp. ehrenbergii, but similar in its
floral morphology to S. fendleri. N .E. Mexico, States of Coahuila and
Neuvo Le6n, in dry pinon scrub (Pinus cembroides, Juniperus mono-
sperma, etc. ,) at 1750-2500 m in rather similar habitats to S. jamesii.
2n=48; EBN =2.
3 S. matehualae Hjerting et Tarn. Leaf 2-3(-4)-jugate with very few
interjected leaflets, glabrous or with a few short hairs on the veins. Corolla
dark purple below, paler above, rotate-pentagonal; style 12-13 mm long,
curved above, exserted up to 8 mm. Mexico, State of San Luis Potosi, at
2740 m along field borders. 2n=48.
4 S. papita Rydb. Plant small and delicate, not more than 10 cm tall; leaf
3-4(-5)-jugate; corolla very small, to about 1.5 cm diam., either very pale
lilac to white, or very dark purple, rotate. Anthers only 4 mm long. N.W.
Mexico, States of Chihuahua, Zacatecas (also Sonora - 'So nannodes').
Growing in dry oak-juniper scrub at 1500-2800 m. 2n=48; EBN=2.
5 S. polytrichon Rydb. Distinguished by the dense spreading pubescence
of thick white hairs on the whole plant, and especially on the pedicel, both
below and above the articulation; corolla generally white, occasionally
mauve; articulation high. Some forms possess sparse leaf pubescence, but
the spreading pedicel pubescence is a constant feature. Some blight-
resistance. Central and N.W. Mexico, in dry stony and shrubby places,
occasionally as a weed of cultivation, at 1800-2500 m. 2n=48; EBN =2.
6 S. stoloniferum Schlechtd. et Bche. A very polymorphic species,
differing in degree of leaf dissection and flower colour (white to purple).
Constant features are the circular corolla outline with large acumens, the
coarse appressed hairs over all green parts (not spreading as in S.
polytrichon) , the pedicel glabrous or only sparsely hairy above articulation,
and the acuminate leaflets. Locally abundant forms (previously described
as species) are found in certain regions (e.g. S. antipoviczii and S.
ajuscoense in the higher regions and S. longipedicellatum in the lower parts
of the Valley of Mexico; S. tlaxcalense in the eastern part of the distri-
bution range). Detailed studies are needed here to elucidate the pattern of
variability. Shows resistance to blight, insects and viruses. Central Mexico
from Michoacan to Orizaba. Dry plateaux, valleys and hillsides, chiefly as
ruderal plants, from 1800 to 3000 m.
6a Subspecies Stoloniferum. More coarsely pubescent than subsp.
moreliae and with white to pale mauve corolla and larger calyx (5-9 mm
long). Widespread in central Mexico. 2n=48; EBN=2.
6b Subspecies moreliae Hawkes. Pubescence rather finer than subsp.
stoloniferum, lateral leaflets slightly decurrent, calyx smaller (5 mm long)
and corolla very rich dark purple. Habit delicate. Mexico, State of
Michoacan. 2n=48(?).
7 S. x vallis-mexici Juz. This hybrid 'species' is distinguished from S.
stoloniferum by the dark purple corolla, and broadly obovate to rhomboid
terminal leaflet which is larger than the laterals. Formed as a hybrid
between S. stoloniferum (2n=48) and S. verrucosum (2n=24), occurring in
the Valley of Mexico and elsewhere, where the altitude ranges of these two
species overlap. Woods, fields, waysides, from 2400 to 3000 m. 2n=36.
Series XIX DEMISSA Buk.

A rather heterogeneous group of tuber-bearing species characterized
chiefly by the high pedicel articulation and rotate corolla with generally
very short lobes, similar to those of series Acaulia. It is probably linked
with Tuberosa through the diploid species S. verrucosum. Mexico and
Guatemala. Only pentaploids and hexaploids are known.
1 S. brachycarpum Corr. Clearly distinguishable from other species in
this series by the upright stems, rather poorly dissected leaves with
petiolulate leaflets and few interjecteds; two uppermost leaflets equal in
size and shape. Corolla large, with short lobes, pale blue-purple with thin
white lines on upper surface. Berries short-{;onica!. West, central and
south Mexico, in high altitude pine and fir forests at 1700--3350 m. 2n=72;
EBN=4.
2 S. demissum Lind!. Grows in rosettes or semi-rosettes but sometimes
produces a long stem. Leaflets sessile to sub-sessile with rounded apex.
Corolla purple, generally with very short lobes. Fruits globular to ovoid.
Highly blight-resistant. Mexico, from Chihuahua State southwards into
Guatemala; pine and fir forests from 2650 to 3800 m. 2n=72; EBN =4.
3 S. guerreroense Corr. Distinguished from S. demissum by the conical
fruits and from S. brachycarpum by the well dissected leaves with 5 pairs of
primary laterals and 8 or more pairs of interjected leaflets. Corolla pale
mauve to lavender, with pale white lines on upper surface, to 3 cm diam.
Distribution and ecology of potatoes 41
and with very short lobes. Some blight-resistance. S. W. Mexico, Guerrero
and Jalisco States in pine-oak forest at 2800-3000 m. 2n=72; EBN=4.
4 S. hougasii Corr. Differs from S. demissum and S. guerreroense in its
tall upright habit, white flowers (often tinged purple between petals)
longer corolla lobes, and petiolulate leaflets. From S. iopetalum, S.
brachycarpum and S. guerreroense it differs in the corolla form and colour,
and in the round berries. Bears some similarities to S. stoloniferum in
series Longipedicellata. Some blight-resistance. West Central Mexico;
high pine forests or even above the tree line in alpine meadows, from
1600 to 2950 m. 2n=72; EBN=4.
5 S. iopetaium (Bitt.) Hawkes. Habit similar to that of S. brachycarpum
but corolla larger and of a richer deep red-purple colour, with short flat
lobes. East Central Mexico, States of Hidalgo, Puebla and Veracruz, at
1900-2360 m, in pine-oak forest. 2n=72; EBN=4.
6 S. schenckii Bitt. Leaf 2-3-jugate, the terminal larger than the laterals,
which are sessile, generally decurrent on to the rachis; interjected leaflets
generally absent. Corolla rotate to pentagonal, dark purple outside, paler
within. Berry conical, white-verrucose. Mexico, States of Oaxaca, Puebla,
Queretaro and Veracruz, from 2600 to 3000 m in pine forests, etc. 2n=72.
7 S. x semidemissum Juz. Differs from S. demissum in the upright habi,t
and petiolulate leaflets, and from S. x edinense* in the large terminal and
poor leaf dissection (2--4-leaflet pairs). Is of undoubted hybrid origin,
derived possibly from a S. demissum x S. verrucosum cross (with the
participation of an unreduced S. verrucosum gamete). Rarely forms
berries. Blight-resistant. Central Mexico; field weed or along hedges and
waysides, sometimes in pine forests, from 2700 to 3500 m. 2n=60.
2.3 DISTRIBUTION AND ECOLOGY OF POTATOES
The common cultivated potato, S. tuberosum, through the efforts of

breeders and agronomists, can now be raised in most parts of the world,
although in the tropics it needs to be grown in the winter season and even
so, cannot be considered at present as well adapted. Work at the
International Potato Center (CIP), however, is in progress to breed
varieties adapted to the humid lowland tropics (CIP, 1972, 1983, 1984);
and already considerable success has been attained by the use of hybrids
between the two subspecies of S. tuberosum. Neverthless, in general,
potatoes are best adapted to the cool temperate zones of the high altitudes
in the Andes (2000--3500 m), at sea level in temperate regions of North
America, Europe, southern Chile and Argentina, and at appropriate
altitudes in intermediate latitudes.
* s. edinense - a hybrid between S. tuberosum and S. demissum - is not included here because
it is not at present available in gene bank seed lists (inventories).
.--+_ _-r- 10
S. slenotomum
S. phureja
S. ajanhuili
(""0- 30
1
;:: UR~'
$ ~--,--
fi
<:(' \
T
60
0
!
400
!
Figure 2.1 Distribution of diploid cultivated potatoes in South America.
The frost-resistant species S. x ajanhuiri, S. x juzepczukii and S. x

curtilobum are grown from about 3000 to 4000 m above sea level, whilst S.
tuberosum subsp. andigena and S. stenotomum yield best at 2500 to 3500 or
a little higher. By human selection in southern Chile and in Europe subsp.
tuberosum has adapted its photoperiodic requirements to long day lengths,
although the temperature conditions cannot be very different from those to
which the original subsp. andigena was accustomed. The quick maturing
and short dormancy species S. phureja was evidently selected for
a warm temperate frost-free climate in the lower valleys of the eastern
(-20
I I
r~-J-i __ ~
I ~~~[; I "0 400

{"1
800
IscALE ~F MILE§_____;~
Figure 2.2 Distribution of triploid, pentaploid and tetraploid cultivated potatoes

in South America.
Andes but can still be grown at fairly high altitudes in Ecuador and Colombia.
By and large, then, all the cultivated potatoes basically evolved in cool
temperate regions even though some have acquired resistance to frost
through hybridization with the frost-tolerant wild species, S. acaule and S.
megistacrolobum. All historical and archaeological evidence points to the
fact that cultivated potatoes were once entirely confined to the high Andes
of South America and the coastal strip of central to southern Chile (see this
volume, Chapter 1 and Figs 2.1 and 2.2).
The wild species, on the other hand, are found in a very much wider
range of habitats, and over a wider latitude range than the cultivated
species, since they are known to occur from the United States (Colorado,
Utah, etc.) southwards through Mexico and Central America into South
America, as far south as latitude 45°S (see Figs 2.3 and 2.4). In both sub-
continents they are chiefly plants of medium to high altitudes, although in
South America they are found in the coastal areas of Peru (growing during
the winter fog period) and Chile, as well as the plains of Argentina,
Paraguay, Uruguay and southern Brazil.
NEBRASKA ,
NEVADA ," UTAH
, NORTH AMERICA
.......--- I Morellilormia
II Bulbocastana and IV Po~adenia
III Pinnalisecta
XII Conicibaccala
.- XVI Tuberosa
•••••••••• XVIII Longipedicellata
XlX Demissa
0 200 400 600 miles

12- I I I
0 200 400 600 600 1000 km
!
112
I
108
I
104 100 96 92
Figure 2.3 Distribution of wild potato series in North and Central America.
The richness of species varies greatly over North, Central and South
America, being greater in central Mexico on the one hand and in the
central Andes of South America on the other, whilst towards both ends of
the range only one or two species are to be found.
Wild species extend through a wide range of plant communities, from
the scrub and cactus deserts of Mexico, the southern United States, Peru,
Bolivia and Argentina to the high rainfall mountain and cloud forests of
50 40
l.--'-..l.3~!;L-l-_ _ +-_+_-t----rl0
AiiI.=--+---40
10-1------t-····-'(~.~~\i~~~" BR~
SOUTH AMERICA '\ ,\';'. __ ,'
V Commersoniana '.~~~;\ '~\I:r / -'~':L"
ilU ., ",'
00."",0""'
/ VI Circaeifolia ,,'( ,I: 'PARAGUAY.
~ :~~=.
." _ X Megislacroloba and " '.
_,,// XVII Acaulia .(
XI Cuneoalala
XII Conicibaccala and
XIII Piurana
••.•• •••• XIV Ingifolia
...-/ XVMaglfa
. XVI Tuberosa
Figure 2.4 Distribution of wild potato series in South America.
the eastern Andean slopes, Mexico and Central America. Others again
occur at altitudes of from 3500 to over 4500 m in the high Andes where
they exhibit strong frost tolerance. The desert and coastal forms seem to be
able to withstand considerable drought and heat, whilst those from the
southern pampas of Argentina and adjacent countries are more tolerant of
long day length than species from the central regions. This range of
tolerance to climate and day length is obviously of interest to the plant
breeder in his search for initial material to help extend the climatic range of
the cultivated potato.
In addition to their wide range of climatic adaptation, the wild potatoes,
in so far as individual species are concerned, show a greater range of
adaptation to pests and pathogens than the cultivated potatoes. This,
again, makes them of interest and value to breeders, although their full
potential has not yet by any means been fully realized. However, as will be
seen in later chapters of this book, many valuable genetic characters of
resistance have been found in wild potatoes and have been incorporated
into the newer commercial varieties.
46 Biosysternatics of the potato
Thus the Mexican S. demissum and allied species have conferred a high
degree of resistance to late blight, Phytophthora infestans. Resistance to
potato virus Y and allied strains as well as to various insect pests has been
found in the Mexican S. stoloniferum and the Argentine S. chacoense and
allied species in each of the two countries mentioned. Frost-resistance has
been found in a range of wild species, that from the Andean S. acaule
having already been incorporated in promising breeding lines as well as at
least one released variety. This resistance also occurs in some of the
primitive cultivated species, as has already been mentioned, as well as
resistance derived from another wild species, S. megistacrolobum.
What has been accomplished by 'nature' (actually, of course, under
cultivation, although not due to conscious direction by man) can also be
carried out by the plant breeder. Another probable example of this process
is the natural transference of resistance to races of Globodera rostochiensis
and G. pal/ida from at least one of the many resistant wild species, S.
oplocense, into tetraploid weed potatoes and even possibly into cultivated
ones in Bolivia (Astley, 1979; Astley and Hawkes, 1979).
Resistance to potato leaf roll virus has been found in S. acaule and in S.
brevidens and S. etuberosum, whilst potato spindle tuber viroid resistance
has been identified in S. acaule (Salazar, 1981).
Insect-resistance has been known for a long time in S. chacoense.
Recently, aphid-resistance has been identified in S. berthaultii, S. neo-
cardenasii and various other glandular-haired species. Resistance to round
cyst and root-knot nematodes (Globodera rostochiensis, G. pallida, and
Meloidogyne incognita) has been identified in very many wild species from
Peru, Bolivia and Argentina. Examples of bacterial-resistance are also
known, particularly Pseudomonas solanacearum resistance, in some forms
of S. phureja and even better resistance in S. sparsipilum and S. chacoense
(see elP Annual Reports), the former species also contributing Meloidogyne
resistance.
General reviews of evaluation research will be found in Hawkes and
Hjerting (1969, 1989) and in Ross (1986). Detailed evaluation results
are listed in Hanneman and Bamberg (1986), Hoekstra and Seidewitz
(1987), Huaman (1987), Radcliffe et at. (1981) and Radcliffe and Radcliffe
(1986).
2.4 SPECIES CONCEPTS IN POTATOES
Most cultivated plants are notoriously difficult to classify. The bewildering

morphological and physiological variation with few crossability barriers or
clearly and easily identifiable discontinuities has more often than not
caused the taxonomist to leave cultivated plants severely alone. Where
cultivated plants have been the object of study, species concepts have
ranged from very narrow ones, in which nearly every variant has been
Species concepts in potatoes 47
given a Linnaean name at the specific or infraspecific level, and many so-
called 'species' have later been found to depend on no more than one or a
few gene differences; or, at the other end of the spectrum, an excessive
'lumping' together (in opposition to the work of the 'splitters' mentioned
above) by some workers has tended to obscure the very real variation that
exists and needs to be analysed. A middle course between these two
extremes can perhaps be steered when biosystematic, chemotaxonomic
and taximetric studies are used, but unfortunately most taxonomists have
on the whole preferred to work with wild species, although there are of
course many notable exceptions.
Although genetic diversity is high both in wild and cultivated plants the
latter seem to possess a complexity of variability which has always been
difficult to analyse. In cultivated plants various types of artificial selection
have been acting, in addition to natural selection, on a series of species
complexes whose history is difficult to unravel. The broad canvas of natural
selection has been overpainted constantly by artificial selection, acting
often in a contrary or quite unknown direction during prehistoric and
historical times from some 7000 years ago right down to the present.
To take an example far removed from potatoes, we have fairly good
documentary evidence from the last two centuries on the history of the
cultivated rose and we therefore know rather accurately how our garden
roses have evolved during this period under the hand of man from sources
in Europe, the Middle East and the Far East. Before that time the
evidence is fragmentary, but what we have available to us, helps us to
understand the pattern of variability and enables us to set out a valid and
useful classification. Had this knowledge not been available our task would
have been so much more difficult that it might not have been possible at all.
How, then, can one follow over a period of perhaps five to seven millenia
the movements of a cultivated plant like the potato or wheat, taken by man
from its centre of origin through his wanderings or passed on from tribe to
tribe and brought under natural and artificial selection pressures to which,
as a wild plant, it had never been subjected? Such a plant might also have
hybridized frequently with related wild species in areas other than that to
which it was originally adapted and hence acquired further variability, and
in fact we have several good examples of this process in many of our major
crops. After periods of evolution under domestication the cultigen may
have renewed genetic contact with the wild prototype and/or related weed
forms; this process also is likely to provide further variation.
So the complex pattern of variability in a cultivated plant derived from
its subjection to natural and artificial selection pressures of a complex
nature about which we at present know all too little except in general
terms, and its acquisition of variability through introgressive hybridization
and often also through polyploidization, all add to the difficulty of the
taxonomists in classifying it and providing himself with a clear concept of it
as a species or a group of species.
A further difficulty with the potato is its high degree of phenotypic
plasticity, as well as its genotypic diversity. This applies equally to the
related wild species as well as the cultivated ones. The potato taxonomist
therefore needs to look for those characters that are genetically stable
within the species and which are less liable to show a wide range of
phenotypic expression. Thus, with potatoes as with other plants we are
looking for discontinuities of a morphological nature rather than very hard
and fast breeding barriers. After several decades of studying potato species
one reaches the conclusion that morphological discontinuities have arisen
from barriers that are ecogeographical as well as cytogenetical; sometimes
the ecogeographical isolation has brought about clear differences worthy of
species rank, and at other times cytological and genetical barriers have
been effective.
2.5 CROSSABILITY, STERILITY AND BREEDING BEHAVIOUR
The pattern of variability iJil the tuber-bearing Solanum species cannot be

adequately understood without mention being made of the two alternative
methods of reproduction available to them. One of these, the sexual,
largely outbreeding reproductive system is particularly common in wild
species, less so in cultivated ones. The other way of reproduction is
asexually, by means of tubers. Theoretically, potatoes should be able to
maintain themselves indefinitely by this means, especially under a favour-
able stable environment with a minimum of competition from other
vegetation. However, if the environment is changing or the competition
becomes more intense, the clonally propagated plants will die out and new
plants will arise from seed elsewhere. Thus a delicate balance is maintained
by means of natural selection between stability of genotype when con-
ditions are stable and a varying series of genotypes for natural selection to
act upon when the environment is changing. Such a dual system must be of
immense value to potatoes, which are not alone amongst plants in devising
a reproductive strategy of this type.
Most potato breeders and geneticists are aware that the majority of wild
and cultivated diploid potato species will cross without difficulty and that
the resulting Frs are fertile and vigorous. Even tetraploid by diploid crosses
are not very difficult to make, in one direction at least, although the
progeny may not be very numerous or indeed fertile. Be this as it may, the
'gene pool' of potatoes is undoubtedly large, which is of considerable
advantage to those who wish to transfer characters or blocks of genes from
one species to another.
This ease of crossing has led some workers to believe that perhaps all or
most potatoes belong to one or a very few species. If crossability is the only
criterion for defining a species this comment would be correct. However,
in nature, each wild potato species forms a series of interbreeding
Crossability, sterility and breeding behaviour 49
populations separated from those of other species by geographical and
ecological barriers. This seems to be the objective situation, matching also
the range of morphological variation and indicating in general that discrete
groups of populations exist which fit reasonably well with generally
accepted species definitions.
One cannot escape the fact, however, that some wild potato species
frequently form hybrids with each other, especially in habitats that have
been influenced by man. We shall mention in a future section (p. 55) that
the genetic architecture of potato species is so distinct as to reduce fertiiity
and viability in artificial Fz hybrids between them. This process probably
reduces the viability of hybrids in nature but would not prevent gene flow
taking place between two or more species. Indeed, we have some evidence
for the formation of a limited number of 'fixed' hybrids (see S. x rechei and
S. x sucrense) or complete sections of introgressed species (mountain
forms of S. chacoense subsp. chacoense) in some instances.
However, field studies strongly indicate that the viability of hybrids in
nature is not as high as one would have expected. Studies by Hawkes and
Hjerting (1969, pp. 459--60) in Argentina have shown clearly that hybrids
are not able to survive poor or adverse conditions since they possess only
portions of the adaptive complexes of the parents. In a good year the
hybrids will evidently survive as well as the parents, but in a poor year
when rainfall is scarce or too abundant the full adaptive mechanism of the
parents may enable them to survive, whilst the less well-equipped hybrids
will not.
Whatever is the final explanation there seems no doubt that species
hybrids, although quite common in some areas and in some years, do not
seem to persist for more than a year or two. Thus, although Edgar
Anderson's hypothesis of hybrids being able to exist in 'hybrid habitats' is
attractive and possibly correct for some groups of plants, it does not seem
to fit the facts with potatoes, since parents as well as hybrids can exist easily
in those habitats in good years, whilst it is the hybrids, not the parents,
which disappear in the bad ones.
We also have some evidence, admittedly needing verification, of species
being kept apart temporally by difference of flowering time. An example of
this is S. microdontum and S. vernei in N.W. Argentina (see Hawkes and
Hjerting, 1969, p. 460).
Apart from these general examples there is evidence of species evolving under
clear cytogenetical barriers amongst the Mexican wild hexaploid species,
where Fl hybrids are completely sterile (Marks, 1955; Hawkes, 1956a).
Furthermore, crossability barriers between Mexican and South American
species also exist (Gell et at., 1960) as will be discussed in more detail below.
The majority of potato species are outbreeders, and indeed the diploids
are mostly self-incompatible, thus rendering outbreeding obligatory. This
outbreeding system helps no doubt to maintain the high degree of
intraspecific variability which has already been commented upon.
The odd-numbered polyploids (triploids and pentaploids) which are the
results of diploid by tetraploid or tetraploid by hexaploid crosses respec-
tively are of course different 'species' from the normally outbreeding
largely sexually reproducing species which we have just been discussing.
Many would argue that taxa such as S. x vallis-mexici, S. x juzepczukii, S.
X chaucha and S. X semidemissum ought not to be called species at all.
The pentaploid S. X curtilobum, formed from a triploid X tetraploid cross
(the first parent providing an 'unreduced gamete') is another similar case.
Here one is probably justified in giving species names for convenience,
even though these taxa are maintained entirely by vegetative reproduction,
since they can be easily identified and have a definite distribution area if
wild and a place in the agricultural economy if cultivated. They may indeed
be continuously replenished by the sort of crossing processes by which they
are originally formed.
Many (if not all) wild and cultivated potato species seem to be able to
produce 2n gametes (diplogametes or unreduced gametes), as shown by
the work of Peloquin and his colleagues in the USA (e.g. Mok and
Peloquin, 1975a, b). This has been thought to be the mechanism respon-
sible for polyploidy in potatoes, but extraordinarily enough, most diploids
have remained as diploids; this would indicate that in general there is a
mechanism selecting against the functioning of diplogametes. However,
diplogametes seem to function occasionally and to be responsible for the
formation of triploid cytotypes in otherwise completely diploid species.
These were examined in S. commersonii and found to be identical with
crosses of diploid by colichicine-tetraploid synthetics (Tarn and Hawkes,
1986). Morphological comparisons also bear out this suggestion. These
triploid clones are sterile and from an evolutionary point of view are 'dead
ends'. From a taxonomic point of view such 3 X forms are generally
given the same species name as the original diploids. Under favourable
conditions, as with S. maglia, S. caivescens, S. cardiophyllum and S.
commersonii, the triploids form very large colonies. In fact, with the first
two species mentioned, they have almost entirely replaced the sexually
reproducing diploids. With other species, such as S. jamesii and S. venturii
the triploids appear to be rare, but the collection of tubers (now not
generally attempted) might reveal that they could be more common than is
generally considered likely.
Triploid hybrids are also known, and their origin has been satisfactorily
explained, as with S. X vallis-mexici (Marks 1958), S. X indunii (Okada
and Clausen, 1982) and S. X viirsooi (Okada and Clausen, 1985).
Pentaploid hybrids have also been noted, as, for instance, S. X edinense (S.
demissum x S. tuberosum) and S. x semidemissum (probably S. demissum
x S. verrucosum).
Returning now to the sexually reproducing species we can see that with
almost all the diploids outbreeding is obligatory, through the functioning of
genetically controlled self-incompatibility mechanisms. In most tetraploids
Endosperm balance numbers 51
and all known hexaploids self-pollination is possible, but insect pollinators
seem to be needed to transfer pollen. The ratio of cross- to self-pollination
is, however, not known, except in S. acaule, which seems to be habitually
self-pollinated. The hexaploid S. demissum also seems to be of this type.
An exception to the rule of self-compatibility in tetraploids is provided by
the curiously self-incompatible S. tuquerrense in series Piurana. Self-
compatible diploids such as S. brevidens, S. etuberosum and S. morelliforme
also provide an exception to the usual rule of self-incompatibility in this
group.
2.6 ENDOSPERM BALANCE NUMBERS
As has already been mentioned, the gene pool of potatoes is extremely

large - a fact which is of the greatest value to the breeder. Most species will
cross with each other, given a certain amount of patience, although one-
way compatibility is not unknown. However, there are two main groups of
diploid potatoes which are very difficult to cross with each other. The first
of these includes diploid species in series Morelliformia, Bulbocastana,
Pinnatisecta, Polyadenia, Commersoniana, Lignicau/ia, and Circaeifolia
(see Table 2.1). These (and one or two others) are not easily crossable with
diploid species in other series such as Megistacroloba, Tuberosa etc. To
account for this Johnston and Hanneman (1978, 1980a, b, 1982) suggested
a modification of the hypothesis concerning the normal 2:1 ratio of
maternal to paternal genomes in the endosperm. They proposed an
Endosperm Balance Number (EBN) of 1 for the first group of series
mentioned above, and 2 for the other diploid series. If the chromosome
numbers of the EBN = 1 species are doubled by colchicine or other
treatment, thus bringing them up to 2n=48 with an EBN of 2, they will
then cross with the other South American diploids. These EBN =2 South
American diploids cross readily with allotetraploids (EBN =2) but not so
readily with autotetraploids such as S. tuberosum (EBN =4). Hexaploid
species, such as S. albicans and the series Demissa hexaploids have an
EBN=4.
The meaning of this in terms of the actual mechanisms involved, is still
obscure, but since series Etuberosa diploids and the Mexican steHate-
flowered diploids all possess EBN = 1, we can tentatively postulate that this
is the primitive condition and that the EBN =2 condition is in some way
advanced (Hawkes, 1990).
These results and others confirm the general thesis that most, if not all,
of the wild species of potato (with the probable exception of series
luglandifolia, which seems more closely linked with tomatoes, and is not
tuber-forming) will sooner or later be brought into the gene pool of the
cultivated potato.
2.7 CYTOLOGY OF POTATO SPECIES
A polyploid series in potatoes has been known for nearly 50 years. With a
base number of 12, the wild species occur as diploids, triploids, tetraploids,
pentaploids and hexaploids, whilst the cultivated species extend to penta-
ploids only. Some of the best-known species are shown in Table 2.1, set out
in columns according to the level of ploidy. An analysis of all those of
which the somatic number is known shows that most (73%) are diploid,
very few (4%) are triploid, rather more (15%) are tetraploid, 2% are
pentaploid and 6% are hexaploid. (This survey excludes triploid and
pentaploid cytotypes of diploid and tetraploid species respectively.)
It is worth noting that (apart from the autopolyploids mentioned above)
there are at least two species known that occur in two or three distinct
cytotypes. These are S. gourlayi (2x, 4X) and S. oplocense (2x, 4x, 6x).
Both of these species occur in Argentina and Bolivia, countries that
have been very intensively studied. One can thus hope to find similar
phenomena in other countries of Latin America in due course.
All the diploid species sp far investigated have shown regular bivalent
pairing at meiosis. This continues in F1 hybrids between them, even when
the hybrids themselves are weak and infertile. However, although the
chromosomes of different species appear to be homologous from this
evidence, there is some reason to believe that homoeologous chromosomes
in polyploids also occur.
We have already discussed the triploid cytotypes and triploid species
hybrids in a previous section and shall refer to them again when cultivated
potatoes are under review. In all triploids so far investigated meiosis has
been shown to be very irregular, and fertility on self-pollination is very
low. Some, however, will produce seeds when used as female parents in
crosses with diploids and tetraploids.
The tetraploid species, with the exception of S. tuberosum, behave as
allotetraploids, with 24 bivalents at meiosis. However, S. tuberosum
exhibits a large range of meiotic anomalies from 1.70 to 5.24 multivalent
frequencies per cell, according to various workers (see Swaminathan and
Howard, 1953 for details). It is therefore generally assumed to be an
autotetraploid (see Section 2.11 below, however).
Pentaploids are of a similar nature to the triploids mentioned above and
are formed by hybridization, although not always as a result of hexaploid x
tetraploid crosses, as might at first sight be supposed. Their origins are
generally agreed to be as follows: S. x edinense (S. demissum X S.
tuberosum; Hawkes, 1963; Ugent, 1967), S. x semidemissum (S. demissum
x S. verrucosum [providing an unreduced gamete] Hawkes, 1963), S. x
curtilobum (S. X juzepczukii [unreduced gamete] X S. tuberosum subsp.
andigena; Hawkes, 1962).
The hexaploids are uncommon in South America (for example, S.
moscopanum and hexaploid cytotypes of S. oplocense) , and their origin
Chemotaxonomic relationships of potatoes 53
and nature is unknown. The Mexican hexaploids, on the other hand, have
received much attention, especially S. demissum, which has been used so
much in breeding for Phytophthora resistance. The formation of 36
bivalents at meiosis is extremely regular but sterility ensues in the
F j hybrids between them. This has considerable bearing on genetic
relationships between them and will be discussed further in Section 2.9.
2.8 CHEMOTAXONOMIC RELATIONSHIPS OF POTATOES
Chemical taxonomy is a discipline that had its origins in the nineteenth

century but which has only recently, within the last 30 years, been used
extensively to help throw light on taxonomic relationships. Most workers
have concentrated their efforts on wild species, but even so a considerable
body of literature is building up on the results of chemotaxonomic studies
of cultivated plants (see Hawkes, 1968 for literature review).
Useful work is reported on potatoes both in the group of substances
sometimes referred to as secondary metabolites and also with proteins,
enzymes, etc. Harborne's papers on potato polyp he no Is are worth noting
here (see Harborne, 1960, 1962). He found (1960) three coumarins and
other major polyphenols in the corolla of the two closely related wild
species S. jamesii and S. pinnatesectum, as well as in S. x sambucinum,
which later (Hawkes and Lester, 1968) was shown to be a hybrid of S.
pinnatesectum with S. cardiophyllum, No other species were found with
these substances in their flowers.
Flavonol glycosides were analysed (Harborne, 1962) in 60 wild species
and 39 cultivated clones. Whilst most pigments are distributed widely a
particular one, kampferol 3-glucoside, was found only in S. santolallae
from Peru. Richness in kampferol glucosides was found in the three series,
Conicibaccata, Piurana and Demissa, which are probably taxonomically
connected. It is interesting to note also that luteolin 7-glucoside was only
found in S. stoloniferum and in the hybrid derived from it, S. x vallis-
mexici.
In a paper chromatography study of leaf alkaloids Marks et al. (1965)
showed the true nature of a presumed hybrid between two wild Solanum
species, S. demissum and S. stoloniferum. Using gas-liquid chromato-
graphy, in a study of inflorescence hydrocarbons, Mecklenburg (1966)
showed interesting taxonomic correlations between her results and those
obtained from other methods. Of the three main similarity groups, the first
included S. demissum, S. verrucosum and S. stoloniferum. Group two
included S. trifidum, S. clarum, S. fendleri and S. polytrichon, all Mexican
species, whilst group three was much more diverse.
Electrophoretic studies of proteins on paper, starch agar or poly-
acrylamide gels have also produced results of interest. The results, in a
series of characteristic band patterns, have still to be fully interpreted, but
on the whole have tended to support taxonomic systems based on
morphology alone. Thus, Desborough and Peloquin (1966) compared disc-
electrophoretic patterns from 26 wild and cultivated species, finding
reasonable comparability between accessions of the same species and clear
between-species differences. Interesting results were also obtained in
hybrids, where there was segregation for the major bands present in each
parental species.
The same workers (Desborough and Peloquin, 1967) studied esterase
isozymes by disc electrophoresis. However, since there were fewer of these
than in the experiments reported above, it was more difficult to draw
taxonomic inferences from the results.
Immunological studies for potato proteins using double diffusion
techniques were reported by Gell et at. (1956, 1960). The results showed
that the tuber-bearing species fell into three groups, according to the type
of precipitin spectrum formed in double diffusion plates "vith S. tuberosum
antiserum. The maximum number of four bands was given by S. tuberosum
itself, together with all the rest of the South American tuber-bearing
species and the Mexican series Demissa and Longipedicellata. Two bands
were shown with Pinnatisecta and Bulbocastana, whilst only one appeared
with Morelliformia. These results agree remarkably well with the main
crossability groups mentioned in Sections 2.5 and 2.6.
Later work by Lester (1965) and Hawkes and Lester (1966, 1968)
showed that S. commersonii could be distinguished serologically from the
rest of the South American species and that with immunoelectrophoretic
techniques many more subtle differences within the Mexican species could
be clarified. For instance, it was possible to demonstrate subspecific
differences in S. bulbocastanum and to show clear segretation of immuno-
electrophoretic characters in seed grown from S. x sambucinum,
thus providing further evidence for its origin as a hybrid between S.
pinnatisectum and S. cardiophyllum.
Much work on isozyme and allozyme analysis has taken place in the last
two decades, too voluminous to include in this chapter (see Hawkes and
Hjerting, 1989, pp. 72-81 for a discussion of Bolivian and Argentinian
species). Chlorophyll DNA work published by Hosaka and colleagues
has differentiated the cultivated species and their allies, separating the
Mexican EBN = 1 species from all the rest (except S. etuberosum) and
clustering the cultivated potatoes together, except for S. tuberosum subsp.
tuberosum (Hosaka et at., 1984; Hosaka, 1986; Hosaka and Hanneman,
1988a, b).
To conclude, it seems quite clear that a wide range of biochemical
techniques have been of considerable value in providing additional
evidence on the biosystematic and evolutionary relationships of wild
potatoes.
Evolutionary divergence 55
2.9 EVOLUTIONARY DIVERGENCE AND GENOME
DIFFERENTIATION IN WILD POTATO SPECIES
Since most diploid potato species hybridize readily (within their EBN
group) and produce fertile Fjs it is clear that genome differentiation in the
conventional sense has not progressed very far. Since F2 progenies from
species crosses often exhibit a range of weak unthrifty plants it seems as
though a cryptic structural differentiation has taken place in the chromo-
somes whereby very small differences (inversions, translocations, etc.)
exist which are too small to affect pairing in species hybrids (Stebbins,
1950). Such F2 breakdown which indicated differences in the genetic
architecture of species is a useful taxonomic tool (Hawkes, 1966).
Although the regular bivalent pairing in the tetraploid species S. acaule
seems to indicate its strong allopolyploid nature, triploid hybrids between
it and diploid species behave cytologically like autopolyploids, thus point-
ing towards a very small degree of difference between its two constituent
genomes (Propach, 1937) and those of the diploid with which it was
crossed. Thus the S. acaule genome could be typified by the formula
A2A2A3A3' that of the diploid species as AI. Such a hypothesis also
accords well with Lester's (1965) conclusions that there are very strong
serological similarities between S. acaule and species in the series Tuberosa
and parts of Yungasensa (S. chacoense, at least). Stebbins points out that
differential pairing may well take place, and to use the terminology of
Huskins (1932) when homologues are absent then homoeologues will pair
(see Table 2.2).
Table 2.2 Suggested genome formulae for certain potato species

Genome Taxonomic series or species
Aj Series Tuberosa, Yungasensa, Cuneoalata, Megistacroloba
A2 A 3 Series Acaulia
A4 B Series LongipediceUata (at least in part)
AjA4 (B, C, D, etc.) Series Demissa
Considering now the other well-known tetraploid, S. stoloniferum, the

results of Matsubayashi (1955) and Kawakami and Matsubayashi (1956,
1958) indicate that in hybrids with S. chacoense something approaching 12
bivalents and 12 univalents are formed. This is thus very different from the
S. acaule situation and would indicate the presence of one genome in
common (designated in Table 2.2 'A') between the two species and a
distinct one in S. stoloniferum (designated 'B') which is absent in S.
chacoense. On the basis of these results one might then postulate a genome
formula for S. stoloniferum of A4A4BB, the A4 being necessary for reasons
discussed above (also shown in Table 2.2). Marks' (1965) results with S.
verrucosum x fendleri, S. verrusom x S. stoloniferum, and reciprocals all
point in the same direction although in every case there is always a small
number of trivalents present, which might thus be taken to indicate that the
genome differentiation was not completely clear-cut, and certainly not
under the degree of strong genetic control as described by Riley et al.
(1959) for Triticum polyploids.
The cytological analyses made by Swaminathan (1953) and by Marks
(1955) on S. demissum and its hexaploid relatives indicate quite a marked
degree of genome differentiation in these species, where there is a
tendency to the formation of 24 bivalents and 24 univalents. If we assume,
as Marks (1955, 1965) does, that all these hexaploids have two genomes in
common, one of these (A4) being somewhat distinct, and the third genome
quite distinct (B, C, D, etc.), then this would accord with the cytology of
their hybrids. Matsubayashi (1981) concluded from a series of research
papers that all South American species possessed a common genome A,
but with certain variants. Series Acaulia was assigned AAAa A a, Cuneoalata
ACAc , Megistacroloba AmA m, Longipedicellata AABB, S. verrucosum
AdAd and S. demissum AdAdC1C1CZCZ. This differs somewhat from the
Hawkes explanation of the hexaploid Mexican genomes, but the rest tallies
fairly well.
2.10 POTATO EVOLUTION
Various lines of evidence point towards the Mexican diploid potatoes, and
especially series Morelliformia as the most primitive of all the tuber-
bearing species, with their small white stellate flowers and EBN of 1.
Morelliformia, with the other related series, Bulbocastana, Pinnatisecta
and Polyadenia are confined to Mexico, the south-west USA and
Guatemala, and it is probably central Mexico that was the centre of origin
of wild potato species.
This group of series, part of the 'Primitive Stellata' (star-shaped corolla)
could have migrated into South America in the early Pliocene, some 3.5
million years ago, when the Panama isthmus was formed, joining North
and South America. Moving southward, it seems that some of these
Primitive Stellata still exist. These are series Olmosiana and Lignicaulia in
Peru, Circaeifolia in Bolivia, and Commersoniana in Argentina and
countries to the east. All possess an EBN of 1, as do the stellate Mexican
species, and do not cross easily with the other diploid species in South
America with an EBN of 2.
From the Primitive Stellata of South America, the Advanced Stellata of
series Yungasensa evolved, still with white stellate flowers but with
EBN=2. From these in turn developed the Primitive Rotata, with sub-
stellate to pentagonal and rotate (wheel-shaped) norolla. The first series to
Evolutionary relationships of cultivated potato species 57
develop were probably those with a sub-stellate to pentagonal corolla
(series Megistacroloba, Cuneoalata, southern forms of Tuberosa and
Conicibaccata and possibly Maglia). The fully rotate corolla was formed in
the central to northern Andes in series Tuberosa, Conicibaccata, Piurana,
Acaulia and Ingifolia.
A return migration northwards over the Panama isthmus was effected by
tetraploid species in Conicibaccata series and the diploid Tuberosa species
S. verrucosum. The Mexican series Longipedicellata may be due to crosses
of earlier diploid series Tuberosa migrants hybridizing with existing
Mexican species, whilst series Demissa hexaploids are very probably
derived from some of these tetraploids or the Conicibaccata tetraploids
crossing with S. verrucosum, the latest arrival from South America, in
post-Pliocene times.
Of course, this scenario contains some guesswork, and further research,
particularly with gene probes, will be needed to verify, modify or refute it.
2.11 EVOLUTIONARY RELATIONSHIPS OF CULTIVATED POTATO

SPECIES
The origins of the potato as a domesticated plant have already been

discussed in Chapter 1. In the present section I shall attempt to look at the
evolution of the whole group of cultivated species subsequent to the time
of domestication (see Fig. 2.5).
The exact nature of the original wild prototype is uncertain, although it
is generally assumed to be a diploid species in series Tuberosa whose
natural distribution area coincides with the high Andes from central Peru
southwards to central Bolivia. On morphological evidence this prototype
may well have been similar to or identical with the present-day species S.
leptophyes, which occurs in the high basin of Lake Titicaca southwards to
central Bolivia at altitudes of 3000-4000 m or more.
1 S. stenotomum Juz. et Buk. The most 'wild-looking', and therefore
presumably primitive, species of cultivated potato is the diploid, S.
stenotomum, which is commonly cultivated at high altitudes but which,
except for a few forms, is not frost-resistant. In several forms the leaflets
are delicate and narrow, quite similar to those of S. leptophyes, although
the tubers of S. stenotomum, like those of all cultivated potatoes, are much
larger, with a wide and interesting range of shape, skin and flesh colour and
skin patterning, whilst the wild ones are all smooth-skinned with shallow
eyes and a white to dingy brown skin.
2 S. phureja Juz. et Buk. An obviously closely-related species to S.
stenotomum is S. phureja, which some workers consider to be no more than
a subspecies or set of special variants of the former species. It has very
probably evolved in response to agricultural conditions in the warmer
eastern valleys of Peru and Bolivia and has a quick growing season of about
Wild S. aeaule S. sparsipilum S. leptophyes S. megistaerolobum
Species (4x) (2X) (2x) (2x)
----l-----r-----r- S. tuberosum S. stenotomum • S. ajanhuiri (Yari)

subsp. andigena (2x) (2x)
(4X)
Cultivated S. tuberosum ......- - - - - - - - - - . (Ajawiri)

Species subsp. tuberosum (3x) ..
Ls~------,
S. eurtilobum S. phureja
(5X) (2X)
S. juzepezukii
......- - - - - -......~
I
(3x) ......- - - - - - - -.......
Figure 2.5 Evolutionary relationships of cultivated potatoes and their ploidy levels.
Evolutionary relationships of cultivated potato species 59
3 months with virtually no dormancy period. Simmonds (1964) believed it
to have been formed many times from S. stenotomum and to be in fact
conspecific with it. It is worthy of note, however, that S. stenotomum does
not occur in Ecuador, Colombia and Venezuela, where S. phureja is very
commonly grown, and it could not therefore have been derived from S.
stenotomum in those countries. Further work is needed, but there is little
reason to disagree with the thesis on present evidence that S. phureja has
certainly been derived from S. stenotomum by mutation and selection.
3 S. x ajanhuiri Juz. et Buk. The nature of this frost-resistant high
altitude species has only recently been elucidated. Experimental studies
have indicated (Huaman et ai. 1980, 1982, 1983; Johns et ai., 1987), that S.
x ajanhuiri was formed from crosses of S. stenotomum with the wild frost-
resistant species S. megistacroiobum. Furthermore, it seems to have
originated in northern Bolivia, not far south of Lake Titicaca. Although
part of S. x ajanhuiri is an F1 hybrid another part seems to be formed as a
series of F2 segregates or back-crosses to the S. stenotomum parent. The
whole process, though occurring under cultivation, must undoubtedly be
considered a 'natural' one, though the selection of this frost-resistant
'species' was obviously accomplished by man.
4 S. x chaucha Juz. et Buk. This name is applied to a series of natural
triploid hybrids of S. tuberosum subsp. andigena and S. stenotomum.
Sometimes this cross gives rise to a tetraploid progeny through the
functioning of unreduced gametes from the diploid parent; at least this is so
under experimental conditions, and we assume it to be so in the Andes.
Triploids are more common than would be expected from the results of
controlled crosses where very few seeds are produced in any case, whether
triploid or tetraploid.
The amount of gene flow from diploid to tetraploid, either directly or
through the medium of the rather sterile triploids, is also a matter of some
interest, although the question is extremely difficult to answer. This subject
has been investigated by Jackson (Jackson et ai., 1977, 1978).
5 S. x juzepczukii Buk. The nature of this high altitude frost-resistant
triploid species was elucidated by Hawkes (1962) and re-worked by
Schmiediche et at. (1980, 1982). It is a natural hybrid, formed under
cultivation, of the wild frost-resistant species S. acauie and the cultivated S.
stenotomum. It is almost completely sterile and has probably been formed
more than once. S. x juzepczukii is grown at higher altitudes than any
other cultivated potato - to over 4000 m. It occurs from central Bolivia to
central Peru.
6 S. tuberosum L. The story of potato domestication in South America
and its spread round the world have already been discussed in Chapter 1.
The tetraploid potato undoubtedly evolved in the central Andes of Peru
and Bolivia, probably in the region of southern Peru and northern Bolivia
where one of its ancestral parent species, S. sparsipiium still exists.
6a Subspecies andigena Hawkes. This is the more primitive of the two
subspecies, formed in the Andes as an amphiploid hybrid of S. stenotomum
and S. sparsipilum (see Cribb and Hawkes, 1986). At one time it was
thought to be a simple autotetraploid of S. stenotomum, and in fact there is
some evidence to support this hypothesis, since it behaves cytologically as
an auto tetraploid. However, since there is virtually no conventional
genome differentiation in the series Tuberosa to which these species
belong one would expect an amphiploid of two genomically similar species
to behave as an autotetraploid in any case. Since the calyx of S. tuberosum
(both subspecies) differs considerably from that of S. stenotomum the
amphiploid hypothesis seemed less certain, until it was shown by Cribb
(Cribb and Hawkes, 1986) that in tetraploid hybrids of S. stenotomum X S.
sparsipilum the calyx shape of S. stenotomum is suppressed. Interestingly,
Howard (1973) and Woodcock and Howard (1975) showed that in S.
tuberosum dihaploids (2n=24) the S. stenotomum calyx form reappeared.
6b Subspecies tuberosum. This subspecies developed in southern Chile
from subsp. andigena where it was evidently taken by Indian tribes several
millenia ago, and where it became adapted to yield under the long-day
conditions at about 45°S latitude. It also developed in Europe during the
seventeenth and eighteenth centuries, also from subsp. andigena stock.
The two subspecies seem reasonably distinct both morphologically and
physiologically (Hawkes, 1956b).
7 S. X curtilobum Juz. et Buk. The nature of this frost-resistant
pentaploid species was also elucidated by Hawkes (1962). It has un-
doubtedly been formed, possibly only once or a very few times, from a
cross between S. X juzepczukii (providing an unreduced gamete) and S.
tuberosum subsp. andigena (see also Schmiediche et at., 1980, 1982).
Thus the cultivated potatoes form perhaps one of the most complex
patterns of species and hybrids of any of the ancient domesticated plants.
They seem all to be derived originally from the primitive diploid S.
stenotomum, either by mutation and selection (S. phureja) , by species
hybridization at the diploid level (S. X ajanhuiri), by diploid X tetraploid
crosses (S. x chaucha, S. x juzepczukii) , by amphidiploidy (S. tuberosum)
or by triploid x tetraploid crosses (S. X curtilobum). It would seem that
four wild species, S. leptophyes, S. megistacrolobum, S. sparsipilum and S.
acaule were involved in the creation of the group, surely a record for any
crop plant. It is also quite possible that cultivated potatoes may have
assimilated genes from other wild potato species by introgressive
hybridization during the course of their evolution.
REFERENCES
Astley, D. (1979) Solanum sucrense Hawkes - a clarification, in Proceedings of the
Conference on Broadening the Genetic Base of Crops, Wageningen (eds A.C.
Zeven and A.M. van Harten), Pudoc, Wageningen, pp. 233-5.
References 61
Astley, D. and Hawkes, J.G. (1979) The nature of the Bolivian weed potato
species Solanum sucrense Hawkes. Euphytica, 28, 685-96.
CIP (1972) Prospects for the Potato in the Developing World, Intc.mational Potato
Center, Lima, Peru.
CIP (1974--1990) Annual Reports, International Potato Center, Lima, Peru.
CIP (1983) Research for the Potato in the Year 2000. Tenth Anniversary, 1972-1982,
International Potato Center, Lima, Peru.
CIP (1984) Potatoes for the Developing World. A Collaborative Experience,
International Potato Center, Lima, Peru.
Cribb, P.J. and Hawkes, J.G. (1986) Experimental evidence for the origin of S.
tuberosum subsp. andigena, in Solanaceae: Biology and Systematics (ed. W.G.
D'Arcy), Columbia University Press, New York, pp. 383-404.
Desborough, S. and Peloquin, S.J. (1966) Disc electrophoresis of tuber proteins
from Solanum species and interspecific hybrids. Phytochemistry, 5, 727-33.
Desborough, S. and Peloquin, S.J. (1967) Esterase isozymes from Solanurtl tubers.
Phytochemistry,6, 989-94.
Gell, P.G.H., Wright, S.T.C. and Hawkes, J.G. (1956) Immunological methods in
plant taxonomy. Nature, London, 177, 573.
Gell, P.G.H., Hawkes, J.G. and Wright, S.T.C. (1960) The application of
immunological methods to the taxonomy of species within the genus Solanum.
Proceedings of the Royal Society, Series B, 151, 364--83.
Hanneman, R.E. and Bamberg, J.B., (1986) Inventory of tuber-bearing Solanum
species. USDA Bulletin, 533, 216 pp.
Harborne, J.P. (1960) Plant polyphenols. 2. The coumarins of Solanum
pinnatisectum. Biochemical Journal, 74, 270-3.
Harborne, J.P. (1962) Plant polyphenols. 6. The flavonol glycosides of wild and
cultivated potatoes. Biochemical Journal, 84, 100-6.
Hawkes, J.G. (1944) Potato Collecting Expeditions in Mexico and South America.
Il. Systematic classification of the collections, Imperial Bureau of Plant Breeding
and Genetics, Cambridge, 142 pp.
Hawkes, J.G. (1956a) Hybridization studies on four hexaploid Solanum species in
series Demissa Buk. New Phytologist, 55, 191-205.
Hawkes, J.G. (1956b) Taxonomic studies on the tuber-bearing Solanums. 1.
Solanum tuberosum and the tetraploid species complex. Proceedings of the
Linnean Society, 166, 97-144.
Hawkes, J.G. (1962) The origin of Solanum juzepczukii Buk. and S. curtilobum
Juz. et Buk. Zeitschrift fur Pjlanzenzilchtung, 47, 1-14.
Hawkes, J.G. (1963) A revision of the tuber-bearing Solanums, (2nd edn). Scottish
Plant Breeding Station Record, pp. 76-181.
Hawkes, J.G. (1966) Modern taxonomic work on the Solanum species of Mexico
and adjacent countries. American Potato Journal, 43, 81-103.
Hawkes, J.G. (1968) (ed.) Chemotaxonomy and Serotaxonomy. Systematics
Association, Special Volume No.2. Academic Press, London, 299 pp.
Hawkes, J.G. (1990) The Potato - Evolution, Biodiversity and Genetic Resources.
Belhaven Press, London and Smithsonian Institute Publishing Division,
Washington DC, 259 pp.
Hawkes, J.G. and Hjerting, J.P. (1969) The Potatoes of Argentina, Brazil,
Paraguay and Uruguay. A biosystematic study, Oxford University Press,
Oxford, 525 pp.
Hawkes, J.G. and Hjerting, J.P. (1989) The Potatoes of Bolivia. Their breeding
value and evolutionary relationships, Oxford University Press, Oxford, 472 pp.
Hawkes, J.G. and Lester, R.N. (1966) Immunological studies on the tuber-bearing
Solanums. II. Relationships of the North American species. Annals of Botany,
30,269-90.
Hawkes, J.G. and Lester, R.N. (1968) Immunological studies on the tuber-bearing
Solanums. III. Variability within S. bulbocastanum and its hybrids with species
in series Pinnatisecta. Annals of Botany, 32, 165-86.
Hoekstra, R. and Seidewitz, L. (1987) Evaluation Data on Tuber-bearing Solanum
Species, (2nd edn), Dutch German Curatorium for Plant Genetic Resources,
Braunschweig, 202 pp.
Hosaka, K. (1986) Who is the mother of the potato? - restriction endonuclease
analysis of chloroplast DNA of cultivated potatoes. Theoretical and Applied
Genetics, 72, 606--18.
Hosaka, K. and Hanneman, R.E. Jr (1988a) The origin of the cultivated tetraploid
potato based on chloroplast DNA. Theoretical and Applied Genetics, 76, 172-6.
Hosaka, K. and Hanneman, R.E. Jr (1988b) Origin of chloroplast DNA diversity
in the Andean potatoes. Theoretical and Applied Genetics, 76, 333--40.
Hosaka, K., Ogihara, Y., Matsubayashi, M. and Tsunewaki, K. (1984) Phylo-
genetic relationship between the tuberous Solanum species as revealed by
restriction endonuclease analysis of chloroplast DNA. Japanese Journal of
Genetics, 59, 349-69.
HO\vard, H.W. (1973) Calyx forms in dihaplotds in relation to the origin of
Solanum tuberosum. Potato Research, 16,43-6.
Huaman, Z. (1987) Inventory of Andean Potato Cultivars with Resistance to some
Pests and Diseases and other Desirable Traits, International Potato Center
(CIP), 22 pp.
Huaman, Z., Hawkes, J.G. and Rowe, P.R. (1980) Solanum ajanhuiri: an
important diploid potato cultivated in the Andean altiplano. Economic Botany,
34, 335--43.
Huaman, Z., Hawkes, J.G. and Rowe, P.R. (1982) A biosystematic study of
the origin of the diploid potato, SoJanum ajanhuiri. Euphytica, 31,
665-75.
Huaman, Z., Hawkes, J.G. and Rowe, P.R. (1983) Chromatographic studies on
the origin of the cultivated potato, Solanum ajanhuiri. American Potato
Journal, 60, 361-7.
Huskins, C.L. (1932) A cytological study of Vilmorin's unfixable dwarf wheats.
Journal of Genetics, 25, 113.
Jackson, M.T., Hawkes, J.G. and Rowe, P.R. (1977) The nature of Solanum
chaucha Juz. et Buk., a triploid cultivated potato of the South American Andes.
Euphytica, 26, 775-83.
Jackson, M.T., Rowe, P.R. and Hawkes, J.G. (1978) Crossability relationships of
Andean potato varieties of three ploidy levels. Euphytica, 27, 541-55.
Johns, T., Huaman, Z., Ochoa, C. and Schmiediche, P.E. (1987) Relationships
among wild, weed and cultivated potatoes in the Solanum X ajanhuiri complex.
Systematic Botany, 12(4), 541-52.
Johnston, S.A. and Hanneman, R.E. (1978) Endosperm balance factors in
some tuber-bearing Solanum species. American Potato Journal, 55, 380.
Johnston, S.A. and Hanneman, R.E. Jr (1980a) Support of the endosperm balance
References 63
number hypothesis utilizing some tuber-bearing Solanum species. American
Potato Journal, 57(1), 7-14.
Johnston, S.A. and Hanneman, R.E. Jr (1980b) The discovery of effective ploidy
barriers between diploid Solanums (Abstract). American Potato Journal,
57(10),484--5.
Johnston, S.A. and Hanneman, R.E. Jr. (1982) Manipulations of endosperm
balance number overcome crossing barriers between diploid Solanum species.
Science, 217, 446-8.
Juzepczuk, S.W. and Bukasov, S.M. (1929) (A contribution to the question of the
origin of the potato). Proc. USSR Congr. Genet. Plant and Animal Breed., 3,
593-611.
Kawakami, K. and Matsubayashi, M. (1956) Studies on the species differentiation
in the section Tuberarium of Solanum. II. Meiotic behaviour in the triploid FJ
from S. longipedicellatum x S. chacoense, with some regards to chromosome
affinity between the parent species. Sci. Rep. Hyogo Univ. Agric.; Ser. Agric.,
2, 143-8.
Kawakami, K. and Matsubayashi, M. (1958) Studies on the species differentiation
in the section Tuberarium of Solanum. VI. Meiotic behaviour of amphiploids
from Solanum longipedicellatum x S. chacoense and its bearing on genomic
affinity between the parent species. Sci. Rep. Hyogo Univ. Agric.; Ser. Agric.,
3, 124--30.
Lester, R.N. (1965) Immunological studies on the tuber-bearing Solanums. I.
Techniques and South American species. Annals of Botany, 29, 609--24.
Marks, E. (1955) Cytogenetic studies in tuberous Solanum species. I. Genomic
differentiation in the group Demissa. Journal of Genetics, 53(2), 262-9.
Marks, E. (1958) Cytogenetic studies in tuberous Solanum species. II. A synthesis
of Solanum x vallis-mexici. New Phytologist, 57, 300-10.
Marks, E. (1965) Cytogenetic studies in tuberous Solanum species. III. Species
relationships in some South and Central American species. New Phytologist, 64,
293-306.
Marks, G.E., McKee, R.K. and Harborne, J.B. (1965) Double chromosome
reduction in a tetraploid Solanum. Nature, London, 208, 359-61.
Matsubayashi, M. (1955) Studies on the species differentiation in the Section
Tuberarium of Solanum. III. Behaviours of meiotic chromosomes in F J hybrid
between S. longipedicellatum and S. schickii in relationship to its parent species.
Sci. Rep. Hyogo Univ. Agric., 2(1), 25-31.
Matsubayashi, M. (1981) Species differentiation in tuberous Solanum and the
origin of cultivated potatoes. Recent Advances in Breeding, 22, 86-106 (in
Japanese).
Mecklenburg, H.C. (1966) Inflorescence hydrocarbons of somc species of Solanum
L., and their possible taxonomic significance. Phytochemistry, 5, 1201-9.
Mok, D.W.S. and Peloquin, S.J. (1975a) Three mechanisms of 2n pollen formation
in diploid potatoes. Canadian Journal of Genetics and Cytology, 17,217-25.
Mok, D.W.S. and Peloquin, S.J. (1975b) Breeding value of2n pollen (diplandroids) in
tetraploid x diploid crosses in potatoes. Theoretical and Applied Genetics, 46,
307-14.
Okada, K.A. and Clausen, A.M. (1982) Natural hybridization between Solanum
acaule Bitt. and S. megistacrolobum Bitt. in the province of Jujuy, Argentina.
Okada, K.A. and Clausen, A.M. (1985) Natural triploid hybrids between Solanum
acaule Bitt. and S. infundibuliforme Philippi in the province of Jujuy,
Argentina. Euphytica, 34, 219-31.
Propach, H. (1937) Cytogenetische Untersuchungen in der Gattung Solanum, Sect.
Tuberarium. II. Triploide und tetraploide Artbastarde. Z. indukt. Abstamm. u.
Vererb.-Lehre, 73, 143-54.
Radcliffe, E.B. and Radcliffe D.G. (1986) Evaluating the U.S. potato germplasm
collection for resistance to Colorado potato beetle. American Potato Journal,
63(8), 448-9.
Radcliffe, E.B., Lauer, F.1., Lee, M. and Robinson, D.P. (1981) Evaluation of the
United States Potato Collection for resistance to Green Peach Aphid and
Potato Aphid. Univ. Minnesota Agric. Exp. Sta. Technical Bulletin, 331.
Riley, R., Chapman, V. and Kimber, G. (1959) Genetic control of chromosome
pairing in intergeneric hybrids with wheat. Nature, London, 183, 1244-6.
Ross, H. (1986) Potato breeding -- problems and perspectives. Supplement 13 to
Journal of Plant Breeding (eds W. Horn and G. R6bbelen), Paul Parey, Berlin
132 pp.
Salazar, F. (1981) Progress in PSTV research. Rep. Plann. Conf. Strategy for Virus
Management (1980), International Potato Center, Lima, Peru.
Schmiediche, P.E., Hawkes, J.G. and Ochoa, C.M. (1980) Breeding of the
cultivated potato species Solanum juzepczukii Buk. and S. curtilobum Juz. et
Buk. 1. Euphytica, 29, 685-704.
Schmiediche, P.E., Hawkes, J.G. and Ochoa, C.M. (1982) The breeding of the
cultivated potato species Solanum juzepczukii Buk. and S. curtilobum Juz. et
Buk. II. Euphytica, 31, 695-707.
Simmonds, N.W. (1964) Studies of the tetraploid potatoes. II. Factors in the
evolution of the Tuberosum Group. Journal of the Linnean Society (Botany),
59(376), 43-56.
Stebbins, G.L. (1950) Variation and Evolution in Plants, Colombia University
Press, New York, 643 pp.
Swaminathan, M.S. (1953) Studies on the inter-relationships between taxonomic
series in the section Tuberarium, genus Solanum. 1. Commersoniana and
Tuberosa. American Potato Journal, 30(11), 271-81.
Swaminathan, M.S. and Howard, H.W. (1953) The cytology and genetics of the
potato (Solanum tuberosum) and related species. Bibliografia Genetica, 16,
1-192.
Tarn, T.R. and Hawkes, J.G. (1986) Cytogenetic studies and the occurrence of
triploidy in the wild potato species S. commersonii Dun. Euphytica, 35,
293-302.
Ugent, D. (1967) Morphological variation in Solanum X edinense, a hybrid of the
common potato. Evolution, NY, 21, 696--712.
Woodcock, K.M. and Howard, H.W. (1975) Calyx types in Solanum tuberosum
dihaploids, S. stenotomum, S. sparsipilum and their hybrids. Potato Research,
18,460-5.
CHAPTER 3
Structure and development of the

potato plant
Elizabeth G. Cutter
3.1 INTRODUCTION
This chapter attempts to provide a description of the structure of the

potato plant, including both external morphology and internal anatomy.
Since the existing literature appeared to be particularly deficient on
developmental aspects, some re-examination of early stages of develop-
ment has been made where possible. Notwithstanding the considerable
morphological and anatomical literature, it is clear that the structure and
development of the potato, admittedly a very complex plant, are by no
means fully understood; strangely enough, this applies particularly to the
development of the tuber, despite its economic importance. There is
considerable scope for further work in this field.
In addition to descriptive studies, experimental work on the control of
development, especially of the stolon and tuber, has been reviewed. It is
clearly of fundamental importance to achieve some understanding of the
factors controlling the development of thes.e organs. Greater association of
anatomical studies with these experimental investigations would clearly be
beneficial, especially in view of the known effects of hormones on structure
in other plants.
Recent developments in the field of tissue culture, a technique (or
techniques) by which potatoes can be rapidly propagated and maintained
for long periods, are briefly considered. The potentialities of genetic
transformation, particularly the possibilities of modifying levels of
endogenous hormones, are also outlined.
In any survey of the morphology of the potato plant, one is struck by the
wealth of varietal differences. Since these often seem important, the
cultivar used in the work cited in this chapter has been stated wherever
possible. In a plant such as the potato, where the development of storage
organs and the movement of assimilates into them are of such paramount
66 Structure and development of the potato plant
agricultural importance, it is remarkable that so much remains unknown,
or incompletely known, in the field of structure and development.
3.2 DEVELOPMENT OF THE SEEDLING
The potato is relatively rarely grown from seed, except for genetical
purposes, although a considerable research effort has been put into
the practicability of propagating the crop in this way (Umaerus 1987).
Germination is epigeal, i.e. the cotyledons are borne above ground by
elongation of the hypocotyl. The radicle emerges from the micropylar end
of the seed and develops as a tap root., which soon forms lateral roots (Fig.
3.1). The first foliage leaves are ovate and hairy. When the young plant is
only a few cm high, stolons develop in the axils of the cotyledons (Fig.
(0)
Figure 3.1 Developing potato seedlings: (a---<:) parts of Figs 272 and 276 of
Hayward (1938); (d) seedling of cv. Desiree. x 1.5. Note developing stolons in
axils of cotyledons in (d), tubers in (c).
The root system 67
3.1d), and after penetrating the soil form small tubers (Fig. 3.1c; Hayward,
1938). The tubers which develop from these stolons or from others in the
axils of foliage leaves are usually small.
3.3 THE ROOT SYSTEM
A much branched fibrous root system is formed either by the seedling tap
root, or by adventitious roots in tuber-grown plants. In early stages of
growth the root system is restricted to surface soil, the roots turning down-
ward after extending for some distance horizontally; this leaves the area of
subsoil directly below a plant almost free of its own roots (Weaver, 1922).
3.3.1 Structure of the root

Longitudinal sections of the root tip show that the root cap and epidermis
have initials in common (Fig. 3.2). Tangential divisions occur in the
protoderm to a point about ten cells proximal to the apex and the epi-
dermis is not clearly differentiated from the root cap until this level. More
proximally the epidermal cells are slightly elongated radially. Starch grains
are abundant in the more mature cells of the columella of the root cap.
Figure 3.2 LS root apex, showing the common initials of the epidermis and root
cap. Cells containing starch grains are shaded; heavy line indicates boundary
between epidermis and root cap. X 80.
The young root may be diarch (Artschwager, 1918; Hayward, 1938) or

triarch (de Vries, 1878) with a wide cortex and an epidermis bearing root
hairs. Older roots may have a larger number of protoxylem points. In the
adventitious roots examined by the writer (cv. Arran Pilot), the number of
Figure 3.3 TS pentarch adventitious root of cv. Arran Pilot. Maturing sieve
elements (se) (arrowed) can be seen at this level. e, Epidermis; rc, root cap. x 150.
protoxylem poles varied from four to six. In the differentiation of the

vascular tissues, the protophloem sieve elements become distinguishable
first, at a distance of 400-500 /lm from the apex. A transverse section of a
pentarch root at this stage is illustrated in Fig. 3.3. The future pericycle and
endodermis can also be discerned. Protoxylem elements become mature
some distance further from the tip than those of the protophloem.
The endodermis later has well developed Casparian strips. A vascular
cambium develops early and the formation of secondary vascular tissues
occurs in the same way as in most dicotyledons (see Fig. 3.4). The
secondary xylem consists of vessels, which sometimes develop tyloses, and
of fibres and parenchyma. The primary phloem becomes crushed; phloem
fibres are not present. The cortex persists for some time (Artschwager,
1918; Hayward, 1938).
Lateral root primordia originate in the pericycle (Esmarch-Bromberg,
1919). Adventitious root primordia are formed in a comparable region in
The root system 69
Figure 3.4 TS older pentarch adventitious root. ca, Cambium; co, cortex; en,
endodermis, showing Casparian strips; ep, epidermis; mx, metaxylem; peI,
pericyeIe; ph, phloem; px, protoxylem; xy 2, secondary xylem. (Fig. 277 from
Hayward, 1938.)
underground portions of the stem of tuber-borne sprouts (Hayward, 1938;

Ernst et aI., 1968). Adventitious roots can also be induced to form on the
basal surface of tuber slices treated with indoleacetic acid (Miedema,
1973), and on leaflet discs treated with naphthalene acetic acid (Webb et
al., 1983). In the tuber slices the root primordia form close to the vascular
tissue, but the exact site of origin has not been established.
3.3.2 Regeneration from roots

Techniques have been devised for inducing the development of adven-
titious buds on potato roots. One purpose of this is to study periclinal
chimeras of buds (see Section 3.4.1(b)). One method involves inducing
abundant adventitious root development from etiolated sprouts, and
subsequently exposing the roots to light after removing the sprout and all
buds. The roots were scraped at a distance of 5 mm from the sprout to a
sufficient depth to expose the vascular tissue on the upper side, and small
buds later developed just distal to the scraped area (Howard, 1964). In one
series of experiments 85% of the scraped roots gave rise to adventitious
buds (Claver, 1967). Using roots induced on de-eyed slices of non-dormant
tubers (cv. Bintje) treated with auxin, Miedema (1967) subsequently
showed that buds developed on both scraped and uninjured roots exposed
to light. The exact site of origin of these buds is not known. Further
experiments demonstrated wide differences in the ability of different
varieties to regenerate buds from roots (Miedema, 1973). Buds formed in
this way develop into apparently normal tuber-bearing plants. Regeneration
from roots induced by Agrobacterium rhizogenes will be discussed in
Section 3.11.
3.4 THE LEAFY SHOOT
The potato is a herbaceous plant with spiral phyllotaxis. Foliage leaves are
imparipinnate, with small additional leaflets or folioles intervening
between the pinnae. The aerial shoot is initially erect but later becomes
partially procumbent (Art schwager , 1918); it dies back at the end of a
single season, but the plant perennates by means of underground tubers.
The stem is winged, the ribs or wings corresponding in position to the
leaves and resulting from unequal decurrence of the edges of the petioles
(de Vries, 1878). The compound leaves subtend axillary buds; the shoot
shows partial apical dominance. The shoot system is sympodial (Danert,
1957).
Specific morphological features of the leafy shoot differ in the various
varieties. British cultivars are described in the British Atlas of Potato
Varieties (Potato Marketing Board, 1965).
An interesting feature of the morphogenesis of the leafy shoot is that the
lateral buds have diverse developmental potentialities. According norm-
ally to their position on the stem, these buds may develop either as
diageotropic stolons with scale leaves (normally from basal, underground
nodes), or as negatively geotropic shoots with foliage leaves (from nodes
on the aerial shoot). The factors controlling these alternative pathways of
development are considered in Section 3.5.3. A great deal of work has
demonstrated both that the habit of the potato plant can be experimentally
altered, and that this may have important consequences related to yield.
The degree of outgrowth and type of development of lateral shoots in
the potato also have important consequences for the final yield. Varieties
differ in both the amount of branching and the pattern of branch
development (Toosey, 1963); thus there may be varietal differences in the
number of stolons per node, the number of tubers per node, and the
number of tubers per stolon (Wurr, 1974). To cite an example of such
differences, Taylor (1953) found an average of 14.2 nodes with axillary
shoots above ground and 11.8 nodes with axillary stolons below ground in
cv. Ulster Chieftain, compared with 17.7 nodes above ground and 9.0
nodes with stolons in cv. King Edward. Early varieties tend to produce
more nodes subtending stolons than maincrop varieties, but the ratio can
be affected by environmental factors. In maincrop potatoes axillary shoots
attain their greater foliar area by growing more rapidly than similar shoots
in early varieties, rather than by beginning to grow earlier in the season
(Taylor, 1953). Lovell and Booth (1969) found that fewer nodes sub tended
stolons under conditions of low nutr~ent level, and stolon length was
reduced. In cv. Arran Pilot, Morris (1966) found that the development of
The leafy shoot 71
axillary shoots was affected both by the size of the tuber and by the number
of sprouts allowed to grow. For example, in small tubers with four sprouts,
leafy branches sometimes developed at the fourth node from the base,
whereas in large, single-sprout tubers at least 12, and sometimes as many
as 20, of the basal nodes subtended stolons, the first leafy shoot occurring
above this level. Morris (1966) suggested that competition between sprouts
for some growth factor, such as gibberellins, might be important. Further
discussion of the factors controlling the development of an axillary bud as a
stolon or a leafy shoot will be deferred until Section 3.5.3.
3.4.1 The shoot apex

The shoot apex of the potato is of peculiar interest, because of the various
phases in its development. The apex of a lateral stolon gives rise to an
elongated, horizontal shoot, forming scale leaf primordia, becomes and
remains dormant during the formation, growth, harvest and storage of the
tuber, and finally after the breaking of dormancy sprouts and gives rise to
an actively growing erect shoot bearing foliage leaves. Physiological
changes may, however, occur in the meristem during this time. For
example, meristems from tubers stored for 19 months gave rise to plants of
lower dry-weight and with a lower capacity to accumulate reduced forms of
nitrogen than those from tubers stored for 7 months (Knowles, 1986).
Steward et ai. (1981) have shown that daylength and temperature have
profound effects on the shoot apical meristem.
(a) Structure and development

Sussex (1955) surveyed the apical anatomy of six varieties of potato, using
dark-grown sprouts. He found that the apical meristem comprised five
layers of cells in cv. Kerr's Pink, the whole meristem being highly
stratified. This stratification was minimal just after the formation of a new
leaf primordium on the apical flank. Of these stratified layers, two
constitute the tunica (Steinberg, 1950). In early stages of development,
apices of axillary buds may have only two stratified layers. Klopfer (1965b)
also found only two layers in apices of embryos and young seedlings.
Sussex (1955) found some differences both in size and structure of apices of
different varieties. For example, in cv. Majestic the diameter of a sprout
apex at the level of the axil of the youngest leaf primordium was 70-80 ).lm,
whereas it was 140-160 ).lm in cv. Kerr's Pink and Ulster Chieftain, and
180-200 ).lm in Arran Banner and two others. The characteristic stratified
appearance of the potato shoot apical meristem is shown for cv. King
Edward in Fig. 3.5.
In cv. Majestic, Leshem and Clowes (1972) found that the apex of a
sprouting eye was flat, whereas the stolon apex was slightly dome-shaped.
In cv. King Edward and Arran Pilot, apices of both stolon and leafy shoot
Figure 3.5 LS apex of axillary leafy shoot of cv. King Edward, showing the
stratified apical dome. A young leaf primordium with associated procambium is
present on the right. x 230.
are dome-shaped (compare Figs 3.5 and 3.18) so this probably depends on
the variety. Pogorelova (1969) noted that the apex of a dormant bud was
flat, but became convex after the onset of active growth. Viewed in the
scanning electron microscope, the apex of the leafy shoot is slightly domed
and differs from that of the stolon largely in the greater thickness of the
surrounding leaf primordia. Figure 3.6 shows the domed apex of a leafy
shoot of cv. Epicure, under SEM.
In the initiation of leaf primordia, periclinal divisions occur in the second
layer of cells on the apical flank. The three outermost cell layers participate
in the formation of axillary bud primordia (Klopfer, 1965b). These first
occur as a tangentially elongated area of cells in the axils of P 3 and P4 - the
third and fourth youngest leaf primordia - (Sussex, 1955), and in
longitudinal section form a 'shell zone', a group or arc of radially elongated
The leafy shoot 73
Figure 3.6 Surface view of an axillary shoot of cv. Epicure, as seen by cryo-SEM.
The spiral arrangement of foliage leaf primordia can be seen; hairs are developing
on the older primordia. x 168.
Figure 3.7 LS shoot apex, showing bud primordia in the axils of P3 and P 4 , the
third and fourth youngest leaf primordia respectively. TI:w bud meristems are
separated from the apical meristem by a shell zone (sz), a zone of periclinally
dividing cells. (Figs 13a and b from Sussex, 1955.)
cells delimited from the apical meristem by a zone of more differentiated

tissue (Fig. 3.7).
(b) Periclinal chimeras
The occurrence of striking morphological differences between varieties of
the potato has already been mentioned. Some of the most divergent types
are, in fact, periclinal chimeras, i.e. bud sports which have originated by a
mutation, probably in a single cell, which has divided to form a whole layer
of mutant cells in the developing shoot apical meristem (Howard, 1970b).
Various methods have been used in attempts to analyse the location of
these cells in the meristem. The surface layer of the apical meristem, which
gives rise to the epidermis, is designated Lf, the next Lz, and the third layer
L3. Studies of periclinal chimeras thus attempt to discover from which layer
of the apical meristem particular structures are formed, and also from how
many cells each layer has itself originated. The subject has recently been
reviewed; in potato, the evidence is thought to support the view that the
shoot apex is three-layered (Tilney-Bassett, 1986).
The first bud sports to be investigated were those involving changes in
pigmentation of the tuber. The colour of the tuber depends on anthocyanin
in the periderm, formed from Ll and Lz (Asseyeva, 1927; Howard, 1970b).
Asseyeva (1927, 1933) devised the technique of eye-excision to investigate
chimeras. If all eyes are removed from a whole tuber, or from half of a
tuber, new adventitious buds may be formed, presumably from deeper
tissues (Howard, 1969). There is evidence that these buds originate from
one or a few cells derived from L3 (Howard, 1970a). Using this technique,
Asseyeva (1927) obtained normal and leaf mutant forms from halves of the
same tuber. She was also able to obtain one variety, for example one which
had pigmentation around the eyes, from another. By the same technique,
Crane (1936) demonstrated that cv. Golden Wonder is a periclinal chimera
with an inner core which corresponds to cv. Langworthy, since some tubers
of Golden Wonder with the eyes excised gave rise to plants which
produced Langworthy tubers. Other examples are cited by Tilney-Bassett
(1986). In most chimeral tubers a mutation has occurred in L 1 , and the
genotype of L] differs from L2 plus L3 (Tilney-Bassett, 1986).
Many leaf-shape bud sports also occur in the potato, for example plants
with entire leaves, ivy leaf, holly leaf, and wildings, which have few
folioles. It is thought that leaf shape is principally affected by changes in
L 2, although mutations in L] and L3 may have a modifying influence
(Howard, 1967). The dock en leaf bud sport in cv. Majestic, which has
entire leaves, is a periclinal chimera with L] unchanged but 1"'2 and L3
showing a mutation. Following X-ray treatment, a very interesting shoot
was obtained which bore not only normal pinnate, but also entire and
sectorial leaves. The arrangement of these leaves on the shoot was such
that it could be postulated that L2 originated from only two initial cells
(Howard, 1966).
As already mentioned, the potato apical meristem is considerably
stratified. Work on periclinal chimeras of potato, as in other species, has
contributed to an understanding of the degree of independence of the
outer three layers. Thus Klopfer (1965a) considers that such studies
support the view that three independent layers are present, but Howard et
al. (1963) conclude that L3 does not have an independent existence. These
The leafy shoot 75
workers suggest that the apex of axillary buds is produced only by L J and
L 2. However, anatomical observations of axillary bud formation suggest a
more deep-seated origin. Simmonds (1965) concluded from eye-excision
experiments that in some varieties ~ in lateral buds contributes both ~
and L 3, but that in other varieties the inner layers remain distinct. Other
observations suggest that occasional tangential divisions of cells in L2 may
contribute cells to the L J or L3 layers (Howard, 1971, 1972). It seems that
here, as in many other facets of work on potatoes, considerable varietal
differences may exist.
3.4.2 Meristem culture

Plant pathologists have used meristem culture, often in conjunction with
heat treatment, as a means of obtaining virus-free stock of infected
varieties. The terminology used in the literature is extremely confused, and
often shows complete lack of understanding of apical morphology. As
Hollings (1965) has pointed out, the term 'meristem culture' has been
applied to pieces of tissue ranging from 0.1 mm to 1 cm or more in size.
Strictly, the term shoot apical meristem relates to that part of the apex
distal to the leaf primordia, but in fact, although it is now possible, at least
in some species, to obtain the development of whole plants from the
cultured apical dome only (Smith and Murashige, 1970), the pieces of
tissue cultured by plant pathologists usually include the meristem together
with at least the two youngest leaf primordia. Hollings (1965) suggests the
term 'meristem-tip', but this is too vague to constitute a substantial
improvement. That the size of the piece of excised tissue is important in
various ways will be shown below.
(a) Methods
Attempts to culture the shoot apex have shown that the potato is
apparently one of the more difficult species to grow, some varieties being
easier than others. Several different methods have been tried. The potato
apex is naturally sterile (Morel et al., 1968), and surface sterilization of the
shoot tip is all that is required (Morel and Martin, 1955). The general
techniques of dissection and excision have been described by Kassanis
(1967).
Stace-Smith and Mellor (1968) compared several of the culture media
that have been employed. Of the four media they used, that of Murashige
and Skoog (1962) was easily the best, both in terms of percentage survival
and rate of development of the explants. This medium, which has a
relatively high concentration of inorganic salts, was also used (sometimes
in a modified form) by Goodwin (1966), Morel et at. (1968), Mellor and
Stace-Smith (1969, 1987) and Sward and Hallam (1976). Liquid medium
was better than agar-solidified medium, both with regard to rate of growth
and the proportion of rooted buds (Stace-Smith and Mellor, 1968; Mellor
and Stace-Smith, 1969). A bridge of filter paper folded into an 'M', dipping
into the culture solution and supporting the explant, is sometimes used
(Goodwin, 1966), and has the advantage that explants can be removed
without damage to the root system (Hollings, 1965).
Morel (1964) showed that an exogenous supply of gibberellin is neces-
sary in the culture of potato apices. Without it, a callus-like growth
develops. In the presence of gibberellic acid (GA), a normal shoot is
formed, but growth stops at 4-5 mm lrnless the explant is given a high
concentration of potassium and ammonium ions (Morel, 1964; Morel and
Muller, 1964). Alternatively, the Murashige and Skoog medium is ade-
quate to support normal growth (Morel et al., 1968). Optimal growth of
meristems of four cultivars into complete plants was obtained with kinetin,
GA and indoleacetic acid (IAA) , although there were some differences
between cultivars (Novak et al., 1980). On the other hand, Sward
and Hallam (1976) obtained better results without IAA, kinetin and
meso inositol. Explants from sprouts formed in the light grew better than
those from dark-grown sprouts (Kassanis, 1967). Both a higher than
normal concentration of micronutrients and the presence of a low concent-
ration of 8-hydroxyquinoline stimulated the growth of excised buds
(Goodwin, 1966). In cv. Desiree, cultured shoot tips survived better than
axillary buds, although when callus was produced and regenerated shoots
these survived best from buds initially taken from the lower nodes (Anwar,
1984). Moving the cultured plantlets into soil is a critical stage (Mellor and
Stace-Smith, 1987).
(b) Size of explant

Various sizes of explant have been used by different workers. Some
examples are: 75-100 11m wide and 150-200 11m long, usually with one leaf
primordium (Morel, 1964); 100-250 11m square, sometimes with one leaf
primordium (Kassanis, 1957); apical dome with two to four leaf primordia
(Mellor and Stace-Smith, 1969; Gregorini and Lorenzi, 1974); meristem
300-500 11m long, with two leaf primordia (Novak et al., 1980); dome plus
two to three leaf primordia, of length 300-1000 11m (Stace-Smith and
Mellor, 1968); dome plus one, two or four leaf primordia, of length, 120,
270 and 600 11m respectively (Pennazio and Redolfi, 1974); and the
terminalS mm of the shoot tip (Thomson, 1956).
The size of the explant is important because it affects both the success of
culture and the percentage of virus-free plants obtained. Larger explants
rooted better and more quickly (Stace-Smith and Mellor, 1968). As
Kassanis and Varma (1967) have pointed out, some cultivars of potato
(e.g. Epicure and Arran Comet) have large, domed apical meristems
which are relatively easy to excise. The size of the explant is of critical
importance in virus eradication (Stace-Smith and Mellor, 1968). For
The leafy shoot 77
example, pieces of apical meristem of less than 200 11m diameter can be
freed from paracrinkle and virus X, whereas plants derived from a large
explant still had virus (Kassanis, 1957). In another investigation tips more
than 1000 11m long failed to give virus-free plants (Sip, 1972). Sometimes
cultured apices are grown into plants, and their apices are again excised
and cultured; this procedure may be repeated several times (Svobodova,
1964).
Thus, explants cultured in order to eradicate viruses should be as small
as is compatible with growth. In this connection, the demonstration that in
other species, including other Solanaceae, excised apical meristems devoid
of leaf primordia can be successfully cultured to yield whole plants (Smith
and Murashige, 1970) is clearly important.
(c) Presence of virus in the apex

For some time it has been a debatable point whether virus particles are
actually absent from the cells of the apical meristem of infected plants or
are merely inactivated by the period of growth in culture. Morel (1964) has
suggested that the virus cannot compete for nucleic acid precursors with
the actively dividing cells of the meristem.
Electron microscopic observation of cells of the apical meristem and
youngest leaf primordia from sprouts of infected tubers revealed bundles
of elongated particles arranged in a parallel fashion. These appeared
paracrystalline in transverse section, and resembled structures found in
infected leaves. It was concluded, therefore, that potato virus X is
present in apices of infected plants but becomes eliminated during culture
(Pennazio, 1971; Appiano and Pennazio, 1972). In another investigation,
homogenates of apical meristems were examined under the electron
microscope. Particles of virus X were found in 98% of preparations from
meristems less than 100 11m long. No particles were found in 50 meristems
of similar size which had been grown for 4 weeks in culture (Krylova et al. ,
1973). Virus particles of potato virus S were also eliminated in heat-treated
and cultured meristems (Sward and Hallam, 1976). Thus, on present
evidence it appears that the meristems become freed from viruses during
culture, although further work will be necessary to establish how this
occurs.
(d) Heat treatment

Potato tubers which are given heat treatment sometimes give rise to virus-
free plants (Kassanis, 1949). This suggested that a combination of heat
treatment and culture of the shoot apex might enhance the degree of virus
elimination, and this was found to be the case (Thomson, 1956; Morel et
al., 1968; Pennazio, 1971; Sip, 1972; Macdonald, 1973; Mellor and Stace-
Smith, 1987). Tubers are usually given fairly high temperatures for periods
of a month or more, and apices excised from these grown in culture. Much
greater success followed after 6 weeks of heat treatment rather than 3, with
subsequent meristem culture (Sward and Hallam, 1976).
(e) Viruses eliminated by culture

Using the technique of meristem culture, with or without heat treatment,
various varieties of potato have been freed of paracrinkle, and potato
viruses X, Y, A and S (see references already cited).
3.4.3 Anatomy of the aerial stem

Transverse sections of the apex of the leafy shoot show its spiral
phyllotaxis, and the rather uniform meristematic cells of the apical dome
(Fig. 3.8a). Owing to the onset of vacuolation in the developing pith and
Figure 3.8 TS apex of a leafy shoot of cv. King Edward. (a) At the junction of P 2
with the apex. The apex itself (a) consists of rather uniform densely staining cells.
P j -P7 are present in spiral sequence. (b) At the junction of Ps with the axis.
Axillary bud primordia (b) can be discerned in the axils of Ps and P6 . Procambium
is present in the axis, and differentiated vascular elements in Ps . X 75.
The leafy shoot 79
cortex, a ring of still densely stammg, meristematic cells, the residual
meristem, becomes delimited about 70 ~m below the tip of the dome
(Fig. 3.8b). Slightly lower down the axis, procambial strands become
differentiated in the residual meristem in positions related to young leaf
primordia.
Using etiolated sprouts, Sussex (1955) found that the first sieve tubes
could be observed in a leaf primordium four or five plastochrones old and
noted that differentiation was continuous and acropetal. The first lignified
xylem elements occurred as a discontinuous strand in the stem below a leaf
primordium three or four plastochrones old and were observed in the leaf
primordia themselves when one or two plastochrones older. Artschwager
(1918) considered that the protoxylem elements were the first elements to
differentiate from the procambium in developing sprouts, whereas at the
tips of older shoots differentiation of the inner phloem occurred before
that of the xylem. He had some difficulty in distinguishing differentiating
sieve elements, and it seems likely that better differential staining tech-
niques were required. Pogorelova (1969) also reported that the protoxylem
differentiated first, groups of external and internal phloem appearing later.
The writer found, on the contrary, that in apices of growing lateral shoots
the first vascular elements to differentiate were the sieve elements of the
abaxial (external) phloem in leaf primordia four or five plastochrones old
(Fig. 3.9); these observations thus agree with those of Sussex (1955), and
of the present author, on sprouts. Immature protoxylem elements can be
observed only slightly later, and sieve elements of the inner phloem in leaf
primordia about six plastochrones old.
Most studies on internal phloem agree that it differentiates later than the
external phloem (see references in Esau, 1938). Some workers regard the
vascular bundles of the Solanaceae as truly biocollateral (Bonnemain,
1970), while others consider that this term, suggesting as it does units of
,one group each of external phloem, xylem and internal phloem, is
inappropriate, since many strands of internal phloem occur in inter-
fascicular regions (Fukuda, 1967). Esau (1938) has described the dif-
ferentiation of internal phloem in tobacco. Here the procambium on the
adaxial side is not sharply delimited from the cells of the ground meristem.
Some of the cells of the latter are converted into procambium by
longitudinal divisions, and differentiate as strands of internal phloem.
Adjacent cells fail to divide, and differentiate as larger parenchyma
cells. Although these cells do not remain meristematic, they retain the
potentiality to divide and some later give rise to additional strands of
internal phloem. Differentiation of internal phloem in the potato appears
to be similar; indeed, this description seems to be true for most Solanaceae
(Fukuda, 1967; Bonnemain, 1970). Extensive anastomosis occurs between
the internal and external phloem, especially by means of more or
less horizontal sieve elements in the young internodes (Sussex, 1955;
Bonnemain, 1970).
Figure 3.9 TS young foliage leaf primordium, showing maturing sieve elements of
the external (abaxial) phloem (arrowed). x 400.
The primary stem has three large and three small bicollateral vascular
bundles in internodal regions (Fig. 3.10) (Artschwager, 1918; Esmarch-
Bromberg, 1919). Artschwager (1918) stated that a cambium was dif-
ferentiated in the young stem even before the sieve tubes of the inner
phloem became evident. The writer's observations suggest that this is
erroneous. However, both fascicular and interfascicular cambium does
become discernible in fairly young stems, and functions normally to give
rise to secondary xylem centripetally and secondary phloem centrifugally.
In interfascicular regions no large vessel members are present (Fig. 3.11)
(Artschwager, 1918). Rays are usually uniseriate. Phloem fibres differ-
entiate on the outer periphery of the vascular cylinder (Fig. 3.11) from
elements of the primary phloem. The cells between the primary xylem [or,
in interfascicular regions, secondary xylem (Fig. 3.11)] and the groups of
internal phloem remain parenchymatous and contribute to a region of
tissue sometimes called the peri medullary zone (Markkrone).
Artschwager (1918) states that a recognizable endodermis, which may
not be distinguishable by its Casparian strips, but is identifiable by its
regular cell shape and by its content of starch grains, is fairly distinct in the
young stem; the writer has not found this evident in the aerial stem. The
The leafy shoot 81
• Q ~
~o
1mm
Figure 3.10 TS internodal region of young stem in primary growth. Three large
and three smaller vascular bundles are present (c, cambium.; p, internal and
external phloem; x, xylem). x 18.
explanation for this may lie in the observation of Venning (1954) that the
starch in the starch sheath was converted to soluble substances during the
day and was only visible at night.
The stem is bounded by a single layer of fairly isodiametric epidermal
cells, with a waxy cuticle, and a distinctive underlying hypodermal layer
(Fig. 3.12). Stomata occur in the epidermis (Artschwager, 1918). Beneath
the hypodermis are three to five layers of angular, or sometimes lacunate,
collenchyma (Fig. 3.12). A parenchymatous cortex lies internal to this.
Starch grains occur in all parenchymatous tissues; those close to sieve
elements are the most developed and remain longest (Yoshida, 1970).
3.4.4 Development and structure of the leaf

The potato foliage leaf is pinnately compound and has small folioles
between the pinnae. There are varietal differences in the number of leaflets
and of folioles, size, colour, etc. (Howard, 1970b). Leaf form can be
profoundly changed by daylength and temperature (Steward et al., 1981).
As already mentioned, leaf primordia are formed by periclinal divisions
Figure 3.11 TS internode of an older stem with secondary thickening, in an

interfascicular region. Small groups of internal phloem elements (ip) are evident.
Phloem fibres (f) are present in both external and internal phloem. Parenchymatous
cells of the perimedullary zone (pz) are present between the secondary xylem and
the internal phloem (c, cambial region; x, xylem). x 160.
in the second layer of cells on the flanks of the apical meristem. Phyllotaxis
is spiral (Fig. 3.6). The young leaf primordium grows apically and
arches over the shoot apex. At an early stage of development, marginal
meristems, which contribute to the growth of the lamina, become evident,
and an adaxial meristem, contributing to increase in thickness of the leaf, is
also discernible (Fig. 3.13). In the compound foliage leaf, development of
the pinnae takes place basipetally, and each leaflet passes through stages
similar to that shown by the young leaf primordium (Artschwager, 1918).
In the young leaf primordium only an arc of procambium is present.
Differentiation of the first phloem and xylem elements takes place as already
described above. Small groups of internal (adaxial) phloem are evident (Fig.
3.14). The vascular tissue of both petiole and midrib forms a semi-circle as
seen in transverse section. A study of the ultrastructure of the phloem gave
a fairly standard picture. Sieve elements were connected to companion
cells by plasmodesmata. Callose and slime (P-protein) were present in the
sieve plate pores. Companion cells had many ribosomes, mitochondria and
dictyosomes, but did not have dense cytoplasm (Tsutsumi et at., 1972).
Artschwager (1918) has described and illustrated the early differentia-
tion of palisade and spongy mesophyll in the leaf lamina. Elongation of the
future palisade cells begins shortly after vascular tissue is differentiated in
The leafy shoot 83
Figure 3.12 TS outer region of a fairly young stem, showing the epidermal layer
(e) with cuticle, the hypodermis (h), and the underlying layers of collenchyma (c).
x 640.
the midrib. The mature leaf has a single layer of palisade tissue, and three
to five rows of spongy mesophyll cells, separated by large intercellular
spaces (Fig. 3.14). Some variation occurs depending on environmental
factors. In a survey of 24 species of Solanum and seven cultivars of S.
tuberosum, a correlation was found between the presence of two palisade
layers (and thicker leaves) and frost-hardiness. It was suggested that this
might serve as a marker for identifying frost-hardy potatoes (Palta and Li,
1979). In leaves of plants grown at high light intensity, the palisade tissue
consisted of large, more elongated cells rich in plastids. There was also
abundant phloem and collenchyma, and many more hairs than in leaves
from plants grown at lower intensities (Scaramella Petri, 1963a). In potato
leaves injured by air pollution, the palisade cells were the first to become
affected and show necrosis, but the spongy mesophyll was also affected
later (Hooker et al., 1973).
The epidermis consists of a single layer of cells with sinuous walls.
Stomata are anomocytic and anisocytic and occur on both surfaces, but are
much more numerous on the abaxial side of the leaf (Ahmad, 1964).
Stomatal number differs in different varieties (Esmarch-Bromberg, 1919).
Figure 3.13 TS developing leaf primordium, showing marginal (mm) and adaxial
(am) meristems. Differentiating vascular tissues are also present. x 200.
Frost-hardy species and varieties had many more stomata on the adaxial
surface than other types (Palta and Li, 1979). Stomatal length and the
number of plastids in the guard cells are positively correlated. Plastid
number can be used as an indicator of ploidy level in hybrids (Frandsen,
1968). Both covering and glandular hairs occur on the epidermis; they are
initially dense, but become separated by growth of the intervening regions
of tissue (Artschwager, 1918). In the genus Solanum each individual bears
two types of hair. As with other structural characters of leaves, these can
be of taxonomic use (Seithe, 1962; Sizova, 1963, 1965a, b). Covering hairs
are uniseriate; glandular hairs have either a small, more or less spherical
head or a rather ovate multicellular head. Uniseriate covering hairs and
two kinds of capitate glandular hairs have been described in cv. Sieglinde
(Bancher and H61zl, 1959). Both types of glandular hair have a single stalk
cell. Hairs with a four-celled head have a neck or collar cell between the
stalk and the head; these hairs apparently secrete volatile oil. The second
type of glandular hair is many-celled and club-shaped; each head cell
con.tains a protein crystal.
The leafy shoot 85
(0)
- - spo
Ph}
mv
- - ob ph
Figure 3.14 TS mature foliage leaf in the region of (a), the lamina, and (b) the
midrib. ab ph, Abaxial phloem; ad ph, adaxial phloem; ca, cambium or cambium-
like region; col, collenchyma; cp, epidermis; hr, hair; my, midvein; pal, palisade
tissue; spo, spongy mesophyll; sto, stoma; v, lateral vein; xy, xylem. (Fig. 288 from
Hayward, 1938.)
Gibson (1971) has described glandular hairs with a four-celled head on

the leaves and stems of certain species of wild potato (Fig. 3.15a). These
hairs secrete a clear exudate which on contact with air changes to black
gummy material. About 30% of aphids placed on these plants became
stuck to the exudate within 24 h, and later died, probably of starvation. It
was thought that if bred into the cultivated potato, in which they are
scarce, these hairs might afford some resistance to aphids (Gibson, 1971;
Tarn, 1971). The hairs of Solanum tuberosum (Fig. 3.15b) do not seem to
have the capacity for capturing and anchoring aphids. However, on hybrids
of S. tuberosum cv. Pentland Crown and S. berthaultii glandular hairs were
numerous and aphids were found gummed down by the exudate (Gibson,
1974). Thus these interesting hairs may constitute at least a partial
resistance mechanism to aphids which can be bred into the cultivated
potato.
Isolated leaves of the potato will form roots, and these can be stimulated
by hormone treatment (Verma, 1966; van Harten, 1972). If the leaf is
separated from the plant, together with its axillary bud, this may develop
as a stolon or, under certain conditions, as a tuber (see Section 3.6.4). This
system has been widely used experimentally in recent years.
3.5 THE STOLON
Potato stolons are lateral shoots, usually those from the most basal nodes
below soil level. Typically they are diageotropic shoots with elongated
internodes, hooked at the tip and bearing spirally arranged scale leaves
(Figs 3.16 and 3.17) (Kumar and Wareing, 1972). When tubers develop,
they do so from the sub-apical region of stolons (Fig. 3.16). However,
tuber formation is regarded as the sum of two separate processes: stolon
Figure 3.15 Glandular hairs as seen with the scanning electron microscope: (a)
single glandular hair with four-celled head on the leaf of Solanum berthaultii. x 350
(Fig. 1 from Gibson, 1971); (b) covering and glandular hairs on S. tuberosum cv.
Arran Pilot. x 170. (By courtesy of Dr R.W. Gibson and Long Ashton Research
Station.)
The stolon 87
Figure 3.16 Base of a young plant of cv. Arran Pilot, showing diageotropic stolons
(s), hooked at the tip, well-formed tubers (t), and young developing tubers Cyt)
visible as a swelling behind the stolon tip. x 0.75.
formation, and tuberization of the stolon tip (Booth, 1959, 1963). These
processes are apparently controlled by different factors.
3.5.1 The stolon apex

Booth (1963) described the stolon tip as an attenuated form of a vegetative
shoot apex, but except for the type of leaf development there was little
evidence of this in the varieties examined by the writer (compare the apices
of vegetative shoot and stolon of cv. King Edward in Figs 3.5 and 3.18).
The apical dome of the stolon is perhaps of smaller diameter. Like the
apical meristem of the leafy shoot, that of the stolon is conspicuously
stratified (Fig. 3.18).
Although no attention seems to have been drawn to this point in the
literature, the hooked part of the stolon tip is striking because of the highly
meristematic nature of the apical and sub-apical regions (Fig. 3.19a, b).
The epidermal cells in this region are radially elongated (Artschwager,
1924) and vertically compressed (Fig. 3.19b). Booth (1963) noted that the
degree of curvature of the stolon tip varied and was related to differences
in cell length on the two sides. Cell lengths became equal on opposite sides
at about 3 mm from the apex.
Figure 3.17 Stolon tips of (a) cv. Arran Banner, and (b) King Edward. Relatively
long internodes are present between the scale leaves (I). x 15.
Leshem and Clowes (1972) studied the duration of the mitotic cycle in
various zones of growing stolon apices by treatment with colchicine, and
found the values obtained to be comparable with those recorded for other
species. The cells situated on the flanks divided 1.6 times as fast as those at
the summit of the apex. As dormancy began, the mitotic cycle lengthened
in all regions of the meristem; this was evident before the onset of tuber
formation, after which the rate became too low to measure. In a comparison
of cell cycling in cv. King Edward, Clowes and MacDonald (1987) found
that, although structurally similar, apical domes of stolons produced cells
at half the rate of those of orthotropic shoots. The greatest difference was
in the young leaf primordia. The rate of mitosis increased within 24 hours
in apices of stolons induced to become erect shoots.
3.5.2 Anatomy of the stolon

As in the leafy shoot, a cylinder of densely staining cells, the residual
meristem, becomes delimited about 80 11m below the tip of the stolon apex,
The stolon 89
Figure 3.18 LS apex of stolon, cv. King Edward. Compare the stratified apical
dome and scale leaf primordia with the section of the apex of a leafy shoot in Fig.
3.5. x 230.
by vacuolation of the cells of the future pith and cortex. At a relatively

early stage of development, intercellular spaces form between the cortical
and pith cells. Starch is also found in cortical cells near the procambium,
and other cells contain many small crystals of calcium oxalate. Sclereids are
reported to occur in the cortex of older stolons (Esmarch-Bromberg,
1919). No collenchyma is present. The pith is of small area compared to the
cortex. Stomata and occasional hairs occur in the epidermis (Artschwager,
1924).
The stolon tip is remarkable for the early differentiation of abundant
sieve elements. In the specimens observed, about 80 J..lm below the tip of
the apex, the first sieve element of the external phloem was differentiated
in relation to the fourth youngest leaf primordium, P 4, and protoxylem
elements below Ps and P 6 , just proximal to their junction with the axis.
Indeed, vascular differentiation in the stolon tip is initially similar to that in
Figure 3.19 LS stolon tip. (a) cv. Arran Pilot. Note the meristematic nature of
that portion of the stolon distal to the hook, and the more elongated, vacuolated
cells proximal to the hook. x 35. (b) Part of LS stolon tip of cv. King Edward,
showing the radially elongated, vertically compressed epidermal cells (e) on the
concave side of the hook. x 200.
the leafy shoot. At about 220 11m below the apex, several sieve elements of
the internal phloem had differentiated in the axis below leaf primordia P r
P6, and by 500 11m below the tip the internal phloem was the conspicuous
tissue (Fig. 3.20). In young stolons subtending developing tubers abundant
phloem is present (Fig. 3.21).
In the slightly older developing stolon, separate bicollateral vascular
bundles are present. There are relatively few xylem elements, but an
extensive development of phloem (Artschwager, 1918). Sieve tube ele-
ments are numerous and of considerable diameter (Artschwager, 1924).
The end walls are transverse, and indeed in a transverse section of a stolon
a number of sieve plates are often seen. Considerable anastomosis takes
place between sieve elements, which form a network (Crafts, 1933).
Phloem fibres differentiate both in the external and internal phloem
(Fig. 3.22). A vascular cambium eventually develops in fascicular and
interfascicular regions, and gives rise to some secondary vascular tissues.
An endodermis consisting of cells smaller than the rest of the cortex,
without intercellular spaces, and possessing characteristic Casparian strips,
is present (Artschwager, 1924).
As in the aerial stem, the elements of the internal phloem, especially in
The stolon 91
Figure 3.20 TS apical region of stolon, showing differentiation of phloem. (a)

Approximately 220 J.lm below the tip of the apex, at the junction of P6 with the axis.
x 75. (b) Part of (a) enlarged, showing differentiated elements of external phloem
(ep), xylem (x), and internal phloem (ip). x 150. (c) Approximately 500 J.lm below
the tip of the apex, below the junction O!f P7• X 75. (d) Part of (c) enlarged, showing
that elements of the internal phloem (ip and marked * on the inner side) are now
conspicuous. x 150.
interfascicular regions, are separated from the xylem by parenchymatous

cells which comprise the Markkrone or perimedullary zone (Fig. 3.22). The
Markkrone is important in tuber development.
The relative amounts of phloem and xylem in a mature stolon, as seen in
transverse section, are shown in Fig. 3.23. Many of the earlier workers
Figure 3.21 TS young stolon below a developing tuber. Abundant internal (ip)
and external phloem (ep) is present. Starch grains occur in the pith and cortex. x
100.
were puzzled by the apparent ability of the rather narrow stolon to

undertake translocation of all the necessary food materials for a growing
potato tuber. Dixon (1922a, b) estimated the area of phloem in a stolon
and the amount of carbohydrate which must be transported along it in a
certain time. From this he calculated that a rate of flow of nearly 50 cm h- 1
would be required. By somewhat similar methods, Crafts (1933) estimated
even higher rates of flow. On the basis of existing knowledge at that time,
Dixon (1922a, b) did not believe that the phloem could possibly support a
rate of flow of this magnitude and therefore argued that transport must
have taken place through the xylem. However, these estimates are well
within the range of more modern calculations of rate of flow through sieve
tube elements (see Crafts and Crisp, 1971). Nevertheless, the potato stolon
is clearly an organ capable of and structurally adapted for unusually
efficient translocation. Indeed, it has been shown that the growth rates of
individual tubers were correlated with the cross sectional area of the
stolons. This area varied ten-fold, but the proportion of it occupied by
external and internal phloem, and by sieve elements, was similar (Engels
and Marschner, 1986).
The stolon 93
Figure 3.22 Part of TS young stolon of cv. King Edward, showing internal (ip)
and external (ep) phloem with fibres (f), the developing cambial region (ca) and the
perimedullary zone (pz) or Markkrone. Strands of internal phlocm are present in
interfascicular as well as fascicular regions. x, xylem. x 160.
3.5.3 Factors controlling stolon development

Stolons normally develop first at the most basal nodes, and then at
progressively higher ones. The number of nodes which subtend stolons,
and the length of the stolons, are adversely affected by low levels of
nutrients (Lovell and Booth, 1969). Booth (1959) showed that stolons
developed at the lower nodes of cuttings taken from the top 1(}-15 cm of
plants of Solanum andigena. Thus any axillary bud along the axis can
develop as a stolon under appropriate conditions, and stolon development
is apparently normally controlled by conditions prevailing in the basal part
of the shoot (Kumar and Wareing, 1972). More recently, the propensity
for any axillary bud to grow as a stolon has been convincingly demon-
strated by the use of cuttings comprising one or more nodes in many
experiments.
Environmental factors may be important. Moist, dark conditions favour
stolon development (Clark, 1921), but only in the presence of a dominant
apex (Booth, 1959; Kumar and Wareing, 1972; Woolley and Wareing,
1972b). For example, Booth (1959, 1963) showed that, whereas in
decapitated shoots of Solanum andigena the bud at the uppermost node
". 0: :'~
1 I I I r !
o 0·1 0·2 0·3 0·4 O,Smm
Figure 3.23 Diagram of TS mature stolon, showing the relative areas occupied by
the various tissues. (Fig. 6 from Artschwager, 1924.)
developed as a leafy shoot, if a combination of IAA and GA was applied to

the cut surface the bud developed as a diageotropic stolon (Fig. 3.24).
After 10-15 days, recovery to a leafy shoot usually occurred, the tip
turning up and giving rise to foliage leaves. If a solution of GA was
introduced into the base of dark-grown sprouts of the cultivated potato (cv.
Majestic) by means of a capillary tube, there was considerable stimulation
of stolon formation, but the pattern of emergence was changed (Booth,
1963). Application of GA to the stems of intact plants also resulted in the
development of aerial stolons in the apical region (Kumar and Wareing,
1972). One-node cuttings cultured in the presence of GA gave rise to thin
shoots with axillary stolons (Woolley and Wareing, 1972b). Single-node
cuttings of cv. Irish Cobbler also gave rise to stolons in media containing
GA, and to tubers in the absence of GA (Iriuda et al., 1983). Thus GA, in
the presence of natural or applied IAA, seems to stimulate stolon
development under a variety of conditions. Lamotte (1983), however,
found that basally-applied abscisic acid (ABA) promoted stolon growth in
decapitated cuttings. In tuber-grown potato plants (cv. Spunta) gibberellin
The stolon 95
activity markedly increased at the beginning of stolon growth, but this was
not true in sprout-grown plants (Abdala de Bottini and Tizio, 1981). From
stolon tips themselves, little or no endogenous gibberellin was isolated
(Obata-Sasamoto and Suzuki, 1979a, b). It is suggested that stolon tips
may produce ethylene as a result of mechanical constraint in the soil
(Vreugdenhil and Struik, 1989).
Figure 3.24 Decapitated plants of Solanum andigena treated at the cut surface
with IAA + GA in lanolin. The uppermost lateral buds have grown out as
horizontal stolons, hooked at the tip. (Plate 6, bottom left, from Booth, 1959.)
Applied cytokinins and gibberellins have antagonistic effects in control-

ling lateral shoot development. If kinetin or benzyladenine (BA) in
carbowax and alcohol was applied to a stolon tip, the apical hook opened
out, and subsequently the shoot became erect and foliage leaves developed
(Fig. 3.25). This could be achieved also with stolons induced by decapita-
tion and treatment with IAA + GA. Basally-applied BA caused stolons to
develop as leafy shoots in cuttings (Lamotte, 1983). Both light and applied
cytokinin were found to be necessary to cause a stolon to develop as a leafy
shoot (Kumar and Wareing, 1972). By the use of radioactive BA, it was
found that the cytokinin accumulated in the stolon tip after the withdrawal
of IAA + GA from a decapitated shoot. It was considered that there was
a causal relationship between this accumulation and the subsequent
conversion of the stolon to a leafy shoot (Woolley and Wareing, 1972a).
(0)
Figure 3.25 Stolon development in Solanum andigetla: (a) naturally developing

stolons emerging from the soil; (b) after kinetin has been applied to the tip, the
stolon turns up and develops as an erect leafy shoot. (Plate 2, Fig. 2 from Kumar
and Wareing, 1972.)
The results of these various experiments indicate that, in the case of

lateral buds which are not subject to apical dominance, a high ratio of
cytokinin to gibberellin favours the development of leafy shoots, and a low
ratio favours stolon development (Woolley and Wareing, 1972b).
Analysis of endogenous hormones in Solanum andigena showed that low
light intensity increased levels of acidic gibberellins in leaves of short day
(SD) plants. Plants grown under low light intensity formed aerial stolons,
and these could be induced in high light intensity plants by application of
GA (Woolley and Wareing, 1972c). Levels of endogenous cytokinins were
lower in SD plants (Woolley and Wareing, 1972c). Thus, to a large extent,
the observations concerning endogenous hormones are compatible with
the view that the control of lateral shoot development depends on the
balance between cytokinins and gibberellins.
As already mentioned, cuttings have often been used in recent investiga-
tions of the control of tuberization. Such studies have shown that there is a
quantitative relationship between the photoperiodic conditions to which
the cuttings are exposed, and the morphological response of the axillary
buds; stolons are produced under minimal inductive conditions (Ewing,
1978,1981; Ewing and Wareing, 1978; see Section 3.6.4(a) and Fig. 3.36).
The structural changes accompanying these changes in external form,
and their control, have not been examined, and this seems a promising
field for future investigation.
3.6 THE TUBER

As already mentioned, stolon formation usually begins at the lower nodes
and progresses acropetally. The first tubers, in turn, usually develop from
The tuber 97
the lower stolons, and tend to become dominant over those formed later
(Plaisted, 1957). The importance of the tubers is indicated by the fact that
75-85% of the total dry matter produced by the plant accumulates there
(Ivins and Bremner, 1964). Tubers which ultimately attain the greatest
weight are usually produced by the lowest stolons, and indeed most tubers
which grew to a diameter of more than half an inch (1.27 cm) were found to
have been formed at the beginning of the period of tuber development
(Clark, 1921).
3.6.1 Morphology of the tuber

As Artschwager (1924) pointed out, the potato tuber is a modified stem
with a shortened (and broadened) axis and rather poorly developed leaves.
The 'eye' of the potato tuber, according to Artschwager, is a leaf scar with
a subtended lateral bud having undeveloped internodes. He also says that
'each eye contains at least three buds arranged in the form of an obtuse
triangle ... ' Both of these statements, although broadly correct, are rather
misleading. Firstly, the leaf component of the eye is not so much a leaf
scar, as a complete scale leaf which has attained little height and which,
because, of the growth in diameter of the tuber, has a base which is greatly
extended tangentially (Fig. 3.26). The 'three buds' in its axil, also
mentioned by Esmarch-Bromberg (1919), are not multiple axillary buds,
but the original axillary bud of the scale leaf and the second order axillary
buds of its first two leaf primordia, which occur (as in the axillary buds of
many species) in nearly opposite positions to right and left of the leaf axil;
Artschwager's Fig. 5 (1924) in fact shows this quite clearly. Again because
of the expansion of the tuber, the first two leaf primordia of the axillary
bud have become tangentially separated (Fig. 3.26). Other workers also
state that 'many axillary buds' are present in each eye (Reeve et aI., 1969).
The eyes are spirally arranged, as were the scale leaves of the stolon tip
from which they have developed. ShaUow and deep-eyed varieties are
known. The potato tuber is thus morphologically straightforward, a stem
with internodes greatly distended in the radial plane, each scale leaf of the
phyllotactic spiral sub tending an axillary bud with its own spiral sequence
of leaf primordia. At the 'heel end' can be found the stolon or the scar of its
Figure 3.26 Diagram of a single potato eye, showing a tangentially expanded scale
leaf subtending an axillary bud (b 1), the first two leaves of which, to right and left of
the scale leaf axil, themselves subtend axillary bllds (b 2).
attachment, and at the 'rose end', slightly offset from the direct axial line,
is the apical bud of the stolon (see Fig. 3.28). Buds towards the heel end
may be smaller than those in median sites (Goodwin, 1967).
In tubers of cv. Arran Pilot, an average of one eye per cm length of tuber
was found, although small tubers had a larger number of eyes per unit
length than larger ones (Goodwin, 1967). This indicates that tubers
continue to elongate, and to incorporate new internodes during a certain
period of development. Went (1959) also found a correlation between
weight of full grown tubers and the number of internodes. More internodes
were present in tubers from plants grown at higher night temperatures.
Upon sprouting of the tuber, the 'apical eye' (i.e. the original stolon
apex) resumes growth first (Artschwager, 1924). Temperature can have an
important effect on apical dominance in the tuber. This is quickly
established by the terminal bud in relatively high temperatures of more
than 15°C (Goodwin, 1963; Ivins and Bremner, 1964). On the other hand,
at 10°C several buds begin and continue to grow. If the tubers are stored at
I-5°C after lifting for several months, and are then kept at high tempera-
tures, all the buds begin to grow, although eventually some are inhibited
(Goodwin, 1963). Thus the temperatures during both storage and sprout-
ing can affect the number of stems per tuber. The factors which lead to
sprouting, after the imposition of dormancy on the bud apices, will be
considered in Section 3.7.1.
3.6.2 Formation and early development of the tuber

Surprisingly little accurate information seems to exist in the literature on
the earliest stages of tuber formation and development, although there
have been a few studies recently on tubers induced in culture.
As has long been known (de Vries, 1878), tuber formation begins in the
sub-apical region of the stolon. It is first evident externally as a radial
enlargement of this region (Figs 3.16 and 3.27; Artschwager, 1924).
Plaisted (1957) states that the first indication of tuber formation is a
thickening of the first internode behind the apical bud of the stolon. This
statement actually refers, as will be seen below, to the youngest elongating
internode of the stolon tip, which is often about the eighth internode
proximal to the apical meristem.
There are conflicting statements in the literature as to whether the initial
radial expansion of this internode is brought about by cell divisi,on or by
cell enlargement. Artschwager (1918, 1924) stated that the initial swelling
was a result of extensive cell division in the pith region. This was followed
by cell division in the cortex and perimedullary zone. Reed (1910) also
reported extensive cell division in the pith, although it is not certain
whether he was referring to the earliest stages of tuber formation. Reeve et
al. (1969) also attributed the initial radial enlargement of the stolon tip
to cell divisions in the pith in random planes and to cell growth.
The tuber 99
Figure 3.27 (a) An incipient tuber, cv. Arran Banner. A swelling has developed
subapically, just proximal to the stolon hook. x 7.5. (b) A comparable stage in
tuber development, cv. Kennebec. In this case, the stolon was cultured for c.6 days
on a medium with high sucrose. (By courtesy of Dr R.L. Peterson.)
However, Booth (1963) considered that the initial swelling was attributable
to cell enlargement.
At a later stage, the cells of the perimedullary zone (Markkrone) divide
and apparently give rise to the bulk of the tuber tissue (Artschwager, 1918,
1924).
Goodwin (1967) states that the continued expansion of the tuber
depends on the production of new internodes from the apical bud. Without
further elaboration this statement, too, is misleading. Goodwin considers
that growth of the buds only stops, and dormancy begins, at lifting or death
of the parent plant, and Moorby and Milthorpe (1975) also state that
growth of the apical bud and of the tuber itself stops simultaneously, but
Leshem and Clowes (1972) have shown that the rate of mitosis in the stolon
apex becomes too low to measure soon after the beginning of tuberization.
In fact, growth of the tuber depends primarily on expansion of internodes
already present in the apical bud, not on the formation of new ones.
Further growth of the tuber gives rise to an elliptical or spherical
structure depending on the balance between growth in length and
thickness (de Vries, 1878). The shape of the tuber is, of course, a varietal
characteristic.
(a) Current observations
These observations do not comprise a full study of tuber development, but
consider early stages in particular.
As already mentioned in Section 3.5.1, the whole of the stolon hook
consists of highly meristematic cells, as judged by their staining properties
(Fig. 3.19). This includes the elongating internode which is incorporated in
the hook itself. Below the hook there is a sudden change to much more
expanded, vacuolated cells (Fig. 3.19).
The first sign of tuber formation, as mentioned by earlier authors, is a
swelling of the sub-apical region of the stolon tip. This occurs at about the
hooked region. Observation of developing tubers of increasing size shows
that initially only one internode is involved in this swelling. In slightly
(0) (b)
lmm
7
5
~
4
'~.(\- .
2
1cm
~
1
(e) (d)
oS
Figure 3.28 Developing tubers, cv. Arran Pilot. (a) Young tuber with the second
internode becoming incorporated. The stolon hook has almost straightened out. x
5. (b-d) Tubers comprising 6, 8 and 11 inte:modes respectively. x !. (s, Stolon or
scar of its attachment. Eyes numbered in order of formation; dotted eyes on other
side of tuber.)
The tuber 101
older, more swollen tubers a second internode, distal to the first, is also
incorporated in the developing tuber, so that a scale leaf may be present at
or near the base of the developing tuber and another about half way along
its length (Fig. 3.28a). At about this stage, because of the considerable
radial expansion of these two internodes, the hook becomes straightened
and the apical bud of the stolon is situated in a more or less terminal
position on the young tuber (Fig. 3.28a). In cv. Arran Pilot, tubers at
this stage measured about 1.5 X 1.0 cm. Some tubers about this size
incorporated four internodes. External examination of tubers of increasing
size revealed that an increasing number of internodes became incorporated
into the tuberous portion of the axis. For example, tubers respectively 4.5
x 2.5, 6.0 x 3.5 and 8.0 x 4.0 cm comprised 6, 7-8 and 11 internodes (see
Fig. 3.28b--d). Internodes are shorter towards the rose end. Tubers
developed in vitro are also formed by the radial expansion of several
internodes (Fig. 3.27b) (Peterson and Barker, 1979). Thus tuberization
proceeds acropetally, involving some longitudinal extension and very
considerable transverse expansion of successive internodes.
Transverse sections of stolon tips show about nine scale leaf primordia
above the hook. Similar sections of very young developing tubers show a
swollen region, with starch-containing parenchymatous cells, distal to the
junction of the eighth or ninth youngest leaf primordium with the axis (Fig.
3.29a). In tubers of this size, about 1.0-1.5 mm in diameter, a cylinder of
differentiating vascular tissue can be seen. Internal and external phloem is
abundant (Fig. 3.29b), xylem elements rather sparse and widely spaced
(Fig. 3.29a). During tuber development lignified fibres are first associated
with external, and later also internal phloem, and an endodermis is present
(Peterson and Barker, 1979). Longitudinal sections of stolon tips and of
developing tubers in early stages seem to show no increase in the number
of files of cortical cells; it was more difficult to be certain in the pith. The
pith cells did seem to be wider, however, and contained starch (Fig. 3.30a).
Thus the writer tends to agree with Booth (1963) that early tuber
development is primarily attributable to enlargement of the pith cells,
although cell division probably rapidly follows. Koda and Okazawa
(1983b) also found that cell enlargement led to initial swelling of the tuber,
followed by cell division. A careful study of the development in vitro of
tubers of cv. Kennebec which developed from excised nodes of stolons,
maintained in the dark and supplied with 12% sucrose, showed that for at
least the first 10 days tuber enlargement was due to cell expansion in the
cortex and pith, and not to cell division in these tissues (Fig. 3.29b);
(Peterson and Barker, 1979). Such tubers usually do not attain more than 1
cm diameter (Peterson et al., 1985). In single-node cuttings of cv.
Katahdin, induced to tuberize by SD, an increase in mitotic index after one
day was reported, whereas cell size did not increase for 4 days (Duncan and
Ewing, 1984). It should be noted that these were sessile tubers (see Section
3.6.4(a». Thus, disagreement still remains concerning the roles of cell
Figure 3.29 (a) TS young developing tuber of cv. Arran Pilot. At junction of ninth scale leaf
primordium and its axillary bud, b. Starch grains are conspicuous in pith and cortex. The scattered
xylem elements (x) are stained dark. Periderm (pe) is beginning to develop by periclinal divisions
in the epidermis and hypodermis. x 75. (b) TS 4-day-old tuber of Kennebec grown in vitro. Cells
in the pith and cortex have enlarged, and contain starch grains. Groups of internal phloem
elements (ip) are evident. Cell division has occurred below a stoma (*) in the epidermis, but not in
the cortex and pith (p). x 136. (Fig. 11 from Peterson and Barker, 1979, © The University of
Chicago.)
Figure 3.30 LS young developing tubers of cv. Arran Pilot. The axis proximal to the stolon hook is
distended radially. In (a) starch grains are present in these cells. At this stage in tuber development the
hook has not yet straightened out. (a) x 25; (b) x 40.
104 Structure and development of the potato pl.ant
division and cell expansion in early stages of tuberization. Deposition of
starch grains varied considerably in the young tubers observed, cv. Arran
Pilot showing conspicuous starch grains (Fig. 3.30a) which were not
present in Arran Banner. In cvs Kennebec and Katahdin, there was an
increase in starch grains 1 to 4 days after tuberization began (Fig. 3.29b;
Peterson and Barker, 1979; Duncan and Ewing, 1986).
Periclinal divisions giving rise to the phellogen occurred in both the
epidermis and hypodermis at an early stage of tuber formation (Fig. 3.31);
Peterson and Barker (1979) consider that the hypodermis gave rise to the
phellogen.
Figure 3.31 Sections of young developing tubers showing the inception of

periderm by periclinal divisions in the epidermis (e) and hypodermis (h): (a) TS;
(b) LS X 300.
The tuber 105
3.6.3 Structure of more mature tubers
In the literature on the more mature tuber there is still disagreement
concerning the relative contributions of cell division and cell expansion.
Reed (1910) found that nearly all the pith cells divided, and that the
parenchyma between the protoxylem and internal phloem (the Markkrone)
also divided actively; as a result the phloem became broken up into many
strands. Lehmann (1926) noted that growth of the tuber mainly depended
on both division and enlargement of the storage parenchyma.
Others consider that cell division in the perimedullary region is more
important than that in the pith (Reeve et at., 1973a, b). Certain quantita-
tive observations have been made. For example, tubers of cvs Kennebec
and Russet Burbank increased in size 225- to 260-fold during their
development. Cell size increased from 15- to 20-fold in the former and 7- to
18-fold in the latter, indicating that there must have been considerable cell
division early in tuber development (Reeve et al., 1973a). These workers
considered that cell enlargement becomes increasingly important in tubers
above 3G-45 g, but Plaisted (1957) found that cell division continued in
tubers up to 200 g (cv. Cobbler). He observed a 500-fold increase in cell
number in tubers between 37 mg and 200 g, and a lO-fold increase in
average cell size. Plaisted calculated that there was a daily increase in cell
number per tuber of 11.7%, and considered that tuber growth was more
directly related to increase in cell number than to increase in cell volume.
Cell size increases with tuber size (Lehmann, 1926) and tubers of 1 cm
diameter have already undergone a considerable amount of cell enlarge-
ment (Reeve et al., 1973a). Cell enlargement is thought to become the
dominant mode of growth in tubers over 3G-40 g (Reeve et al., 1973b).
There is a decreasing gradient in cell size of storage parenchyma from heel
to rose end of mature tubers (Reeve et al., 1971).
The tuber, like the stolon from which it develops, contains an abundance
of internal and external phloem strands (Fig. 3.32). Reeve (1967) considers
that the region of internal phloem comprises at least three-quarters of the
total tuber tissues. Certainly this is an exaggeration if one considers only
the sieve elements and not all the surrounding parenchyma. Some workers,
however, include the perimedullary zone and other tissues in the internal
phloem region (Reeve et al., 1969). The phloem strands in a mature tuber
comprise only about 5% of its volume (Reeve et at., 1973b). As Reed
(1910) pointed out, the phloem becomes 'broken up' into many strands
separated by parenchyma. These strands undergo anastomosis.
Examination of the phloem of cv. Irish Cobbler tubers showed that it
consisted of sieve elements, companion cells, storage parenchyma and
specialized parenchyma cells. The latter contained protein inclusions and
plastids with little or no starch, in contrast to the storage parenchyma
which possessed amyloplasts with large starch grains. The specialized
phloem parenchyma cells were not transfer cells; indeed, it was thought
Figure 3.32 Part ofTS of a mature tuber showing the vascular tissues (p, phloem;
x, xylem). (Fig. 2 from Artschwager, 1924.)
that the abundant mitochondria in the companion cells might indicate that
they functioned in energy-requiring transport (Peterson et ai., 1981).
In the mature tuber the proportion of xylem is small. The protoxylem
elements gradually become separated from the metaxylem (Artschwager,
1924). There is no continuous cambium, although some workers consider
that there is a discontinuous one which produces a few conducting
elements (Lehmann, 1926).
Former views that much of the tuber was of cambial origin seem to be
erroneous. There is rather general agreement that division of the peri-
medullary zone makes a major contribution to the tuber tissues. This
region is said to be of procambial origin (Reeve et ai., 1969, 1970), but see
the discussion of the differentiation of internal phloem in Section 3.4.3.
An endoderm is with Casparian strips is reported in early stages of tuber
development (Reeve et ai., 1969; Peterson and Barker, 1979), but, as a
result of the great expansion of the tissues within it, gaps gradually appear
where the endoderm is has failed to keep pace with this active cell division,
and in later stages no trace of an endodermis remains (Artschwager, 1924).
The cells of the storage parenchyma are more or less isodiametric
The tuber 107
(Lehmann, 1926). In aged tubers callose often develops in the storage
parenchyma cells. It occurs in the form of spherical or elliptical structures,
and various staining reactions demonstrate its similarity to sieve tube
callose (Wodicka and Wenzel, 1971).
(a) Protein
Cubical protein crystals occur in parenchyma cells of the potato, especially
those which are not rich in starch. They occur in cells at the periphery of
the tuber in all varieties, in specialized phloem parenchyma cells (Fig.
3.33) (Peterson and Barker, 1979; Peterson et aI., 1981), and also in the
head cells of glandular hairs (H6Izl, 1965). Pure protein occurs in greater
amount in peripheral regions than in the interior of the tuber (Fischnich
and Heilinger, 1959). Protein crystals do, however, occur in the pith cells,
and these are the first to disappear during sto,rage. They seem to comprise
normal reserve protein in crystalline form (H6lzl and Bancher, 1958).
Figure 3.33 Crystalline protein body from a phloem parenchyma cell. x 1650.
(Fig. 20 from Peterson and Barker, 1979, © University of Chicago.)
The protein crystals of the potato are unique because of their large size -
usually about 10 J.lm, but they may occasionally reach 25 J.lm. These crystals
can now be isolated from potato tissues. They consist of protein with some
amino acid component, chiefly lysine (14%) (Hoff, 1971). A relatively
high-molecular RNA is present (H6Izl, 1965; Hoff, 1971). It has been
suggested that this could be stabilized messenger RNA, required during
the onset of active growth following the breaking of dormancy (Hoff,
1971). The protein crystals are known to disappear during sprouting. Much
of the protein in potato tubers is patatin.
(b) Starch
Storage parenchyma cells contain starch grains with an eccentric hilum and
a series of lamellae, thought to represent layers of starch. These are more
distinct in tubers from early plantings (Salunkhe and Pollard, 1954). Starch
grains may attain a length of up to 100 J.lm. A proportion of compound
grains occurs (Fischnich and Heilinger, 1959; Banks et al., 1973). Even in
developing tubers of about 1 cm in diameter, the fact that the storage
parenchyma cells are filled with starch grains can be well seen in the
scanning electron microscope (Fig. 3.34).
Starch content of tubers is 10-25% of fresh weight, with a maximum of
35%, depending on the variety. Cells with the highest starch content are
those adjacent to vascular bundles in the tuber (Fischnich and Heilinger,
1959). Starch content is reduced by storing tubers at low temperatures
(Ohad et al., 1971). Formation of starch grains proceeds much more
rapidly in early varieties than in late ones (Jager and J akovlev, 1939).
Varieties having tubers with a high starch content were also composed of
larger cells, and there was a relationship between cell size and mealiness
(Barrios et al., 1961). Methods for extracting starch grains from tubers
have been devised (Reeve, 1967). In cvs Russet Burbank and White Rose,
considerable differences in the size of starch grains in different regions of
the tuber were noted (Reeve et aI., 1970). Starch grains are formed by
amyloplasts (Hori, 1954).
Starch grains were formed in tubers when sugars such as sucrose, glucose
or maltose were supplied to the cut ends of stems (Hori, 1954). When
plants were supplied with 14COz, autoradiographs showed the greatest
amount of label along the margin and at the hilum end. This is regarded as
supporting the view that growth occurs by apposition (Badenhuizen and
Dutton, 1956).
The fine structure of potato starch has been investigated by various
techniques. When starch grains were ruptured by ultrasound, they
appeared to have a smooth surface, with internal concentric lamellations.
The grains may be made up of a system of radial submicroscopic units,
probably fibrillar (Gallant et aI., 1972). A structure comprising radially
organized fibrillar units (microfibrils) was previously proposed for
The tuber 109
Figure 3.34 Part of a tuber about 1 cm in diameter, cut across and viewed with
cryo-SEM. (a) Outer tissues, showing the periderm (p) and cortical cells (c)
containing starch grains. Some of the grains have dropped out. x 175. (b)
Individual starch-filled parenchymatous cells from a young tuber. x 640.
Lintnerized potato starch (Sterling and Pangborn, 1960), although it is not
certain whether the micro fibrils are real or artefacts (Leonard and Sterling,
1972). (Lintnerized starch has been treated with cold, dilute mineral acid
for a short time.) Freeze-etching indicated a granular structure of native
starch (Leonard and Sterling, 1972). Other workers propose that potato
starch grains consist of a core consisting of crystalline amylose, surrounded
by a jacket of layers of micelles of amylopectin (Gruber et aI., 1972).
Various models of starch grain structure have been proposed (Sterling and
Pangborn, 1960; Frey-Wyssling, 1974).
(c) Periderm
At an early stage of tuber development, periclinal divisions occur in the

epidermal and hypodermal layers (Fig. 3.31) (Reed, 1910; Esmarch-
Bromberg, 1919; Reeve et aI., 1969; Peterson and Barker, 1979).
Artschwager (1918) considered that the resulting phellogen originated
from the inner row of daughter cells produced by division of the hypo-
dermal layer. He believed that the hypodermis and epidermis give rise to
separate, distinctive periderms.
The cell walls of the phellem autoftuoresce when viewed in ultraviolet or
blue light (Peterson et aI., 1985). Usually 6--10 layers of cork are present
(Fig. 3.34a), but this can be modified by environment. For example, tubers
of cv. Russet Burbank from plants grown at different temperatures had
varying numbers of cell layers in the periderm. At 45-50°F (7.2-10°C) the
periderm consisted of7-12 layers of cells, and was 100-140!lm thick; at 65-
70°F (18.3-21. 1°C), these figures were respectively 9-25 and 120-250, and
at 75-80°F (23. 9-26. 7°C) there were 10-28 layers of periderm, 125-270 !lm
thick. Tubers from plants grown at the higher temperatures were well-
nlsseted, i.e. the periderm was cracked and divided into patches of thick
tissue (Yamaguchi et al., 1964). Perhaps these structural effects were a
response to altered hormone levels in the plant; however, factors
controlling periderm formation are little understood.
The outer phellem cells of the potato tuber, when observed under the
electron microscope, contain numerous ribosomes and mitochondria, but
. no plasmalemma or tonoplast. The cell walls seem to consist of layers of
different electron density, and it is considered that autolysis of the
cytoplasm may yield wall-thickening material (Rainbow and White, 1972).
Using enzymes, it is possible to isolate 'periderm membranes', i.e. several
radial rows of cork cells, from potato tubers (Schmidt and Schonherr,
1982; Vogt et al., 1983). Electron microscopy reveals alternating electron-
dense and electron-lucent lamellae in the suberized walls. It is suggested
that these are layers of suberins differing in polarity, rather than alternat-
ing layers of suberin and wax, as was formerly thought (Schmidt and
Schonherr, 1982). The phellem layers became extremely effective barriers
The tuber 111
to water following several days of storage, which was believed to be an
adaptation to water stress (Vogt et al., 1983).
Lenticels are formed in the periderm at sites below the original stomata
at quite early stages of tuber development (Artschwager, 1918; Adams,
1975; Peterson and Barker, 1979), and provide more permeable regions. It
is possible to estimate the number of lenticels in a tuber by placing it in a
solution of methylene blue under vacuum. The resulting pattern of blue
spots when analysed showed a range between 74 and 1411enticels per tuber
(Wigginton, 1973). Lenticels afford a means of entry for several pathogens
(Adams, 1975).
The number of lenticels per unit of tuber surface can be influenced by
size of tuber, soil type and weather. Moisture conditions in the soil at the
time of tuber formation were particularly important (Meinl, 1966). The
relationship of the lenticels with underlying tissues is also affected by
soil moisture. Under dry conditions a suberized layer forms below the
complementary cells of the lenticel. Increased soil moisture, and water
content of the tuber, lead to swelling of the cortical cells of the tuber and
eventual rupture of the suberized layer, resulting in proliferating open
lenticels (Perombelon and Lowe, 1975). Lenticel development was also
affected by immersing the tubers in GA solutions (Chirilei et aI., 1963).
As seen under the scanning electron microscope, lenticels of potato
tubers are usually circular, forming a crater in the surface of the tuber. In
the centre there may be a raised plateau or a dome. Pores can be seen
between the cells in the centre, where gaseous exchange is thought to
occur. Waxy outgrowths resembling fine threads occurred on cells from the
sides of these pores. It is thought that these outgrowths, which were
observed in 11 varieties, may function in the regulation of water loss
(Hayward, 1974).
(d) Wound periderm

The potato tuber responds to wounding by the formation of periderm, and
this 'wound periderm' confers an important degree of resistance to some
bacterial and fungal diseases. Wounding tissue usually stimulates cell
division (Lipetz, 1970). This is generally preceded by deposition of suberin
at the cut surface. The type of wounding may have an effect, extensive
abrasion resulting in a deep-seated periderm, and smooth peeling giving
rise to a superficial periderm (Artschwager, 1927). The distance of the
phellogen from the wounded surface is also affected by the water content
of the tissue (Rosenstock, 1963). The formation of wound periderm is
affected by temperature and humidity, being more rapid at higher
temperatures and high humidity (Artschwager, 1927; Wigginton, 1974).
Oxygen partial pressures lower than those in the atmosphere are inhibitory
(Wigginton, 1974). Suberization can occur after only one day at 21-35°C,
being followed by periderm formation one or more days later. The layers of
cork cells become impregnated with suberin. At the same time starch
disappears from the layers of cells adjacent to the cut surface (Priestley and
Woffenden, 1923; Artschwager, 1927).
Analysis of the wound periderm showed that hydrocarbons (components
of plant waxes) were present in the suberized layer, increasing consider-
ably 4-7 days after wounding. Treatment with trichloroacetate diminished
the number of electron-lucent bands in the suberin, supporting the
hypothesis that these were waxes (Soliday et al., 1979). Such treatment
decreased the diffusion resistance to moisture (EspeJie et al., 1980),
already known to be much less than that of intact tubers (Vogt et al., 1983).
Priestley and Woffenden (1922) showed experimentally that blocking of
the cut surface was essential for development of a phellogen. More
recently, other experimental studies have succeeded in separating phel-
logen formation and suberin deposition. Mitotic activity resulting from
wounding could be totally suppressed by placing the tissue in tris buffer
(tris-hydroxymethyl-aminomethane), but synthesis of suberin proceeded
normally. If the tris ions were washed out of the tissue slices mitosis could
occur (Kahl et ai., 1969). Using thin slices of tubers of cv. Saskia and
others, it was possible to obtain proliferation in the surface layers of cells
with inhibition of suberin synthesis. The necessary conditions included
separating the tissue without bruising it, rinsing the slices to remove
adhering injured tissues, and maintaining 100% humidity. The inhibition
of suberin synthesis was reversible (Lange et ai., 1970).
Washing tuber discs within 3 or 4 days of cutting severely inhibited
suberization, suggesting that the continued presence of a factor that
induced suberization was needed until induction of the appropriate
enzymes took place. The initial wash water (up to one day after cutting)
contained abscisic acid (ABA), and indeed treatment with ABA increased
the rate (though not the amount) of suberization. Cytokinin and IAA
inhibited suberization (Soliday et ai., 1978). When callus cultures of cv.
Russet Burbank were supplied with ABA, activity of phenylalanine
ammonia lyase and (later) a particular isozyme of peroxidase was much
greater than in controls. Amounts of fatty acids and hydrocarbons, and of
other components of suberin, were all increased in the presence of ABA
(Cottle and Kolattukudy, 1982). It is thought that ABA may trigger the
production of a suberization-inducing factor, which in turn induces the
enzymes involved in suberin biosynthesis (Soliday et al., 1978).
The nature and amount of wound periderm varies in different varieties
(Priestley and Woffenden, 1923; Artschwager, 1927; Nielsen, 1968). In
wounded tubers maintained under the same conditions, cvs Alpha and
Bintje had 10-12 layers of suberized cells, Up-to-Date and Ackersegen 7-8
and Kennebec 5-6. Tubers of Ackersegen formed wound periderm more
slowly than the other varieties (Nielsen, 1968).
Although these and other studies have been carried out in an attempt to
discover the factors leading to development of wound periderm, little or no
The tuber 113
attempt seems to have been made to study the factors controlling the
formation of normal periderm in the potato tuber; indeed, little is known
about such factors in general. Further work is needed to elucidate factors
affecting periderm formation.
3.6.4 Factors controlling tuber formation

Although tubers develop from stolon tips, not all stolons form tubers
(Svensson, 1962). Separate factors regulating tuber formation must there-
fore be involved. It has also been cogently pointed out that tuber formation
does not take place simultaneously in all the stolons of a plant, and indeed
that stolon and tuber formation may overlap (Vreugdenhil and Struik,
1989). Investigation of the effects of various environmental factors led to
the view that a tuber-inducing stimulus was formed in the potato plant. The
nature of this stimulus has been investigated by grafting, by application of
exogenous substances such as various hormones or growth inhibitors, and
by extraction and analysis of endogenous substances from induced and
non-induced plants.
In this section the formation both of underground tubers, and of those
experimentally induced to develop from buds on aerial parts of the plant
will be considered, although it is possible that the processes may not be
identical.
(a) Environmental effects

Tuber formation occurs earlier at low temperatures (Mes and Menge,
1954; Slater, 1963, 1968; Ivins and Bremner, 1964) and is delayed at high
temperatures (Slater, 1963; Menzel, 1980). Gregory (1956) also found,
using cv. Kennebec, that tuber yield was good if plants were grown in short
days (SD) with low night temperature, but no tubers were formed in SD
with high night temperature. Menzel (1980) found that the effects of high
temperature could be reversed by (2-chloroethyl)-trimethylammonium
chloride (CCC) or, to a lesser degree, ABA. Gibberellic acid reduced
tuberization at all temperatures. Menzel (1980, 1985) considers that both
high temperature and low irradiance exert their effects by the production
of a growth substance, possibly gibberellin. High temperatures decrease
partitioning to tubers, and favour haulm growth (Ewing, 1981). Went
(1959) showed that temperature affected not only the yield of the treated
plants, but also that of their progeny. This interesting effect, which Went
compared to that of vernalization, remains unexplained.
Really low temperature treatment may also be effective in inducing
tuber formation. For example, even in young rapidly growing plants,
tubers can be induced to develop by maintaining the plants for 7 days at
7°C or less (Burt, 1964).
There is also a quantitative effect of photoperiod in the potato. Plants
Figure 3.35 Cuttings of Solanum andigena taken from plants grown in (a) long
days, and (b) short days. The lowermost axillary bud has developed as a stolon in
long days, and a tuber in short days. (Plate 1, Figs 1 and 2 from Kumar and
Wareing, 1973.)
grown in SD form tubers earlier than those kept in long days (Chapman,
1958; Madec and Perennec, 1959; Slater, 1963; Ivins and Bremner, 1964).
In Solanum andigena short photoperiods are an absolute requirement for
tuber formation (Fig. 3.35) (Booth, 1959). In the cultivated potato,
however, plants maintained under long days (LD) make active vegetative
growth, but do ultimately form tubers, some 3-5 weeks later than plants
kept under SD (Chapman, 1958). Plants given photoperiods of 9 h formed
tubers in 26-30 days. At least 14 short photoperiods were required for
induction to occur, and the stimulus could also disappear after 14 days of
The tuber 115
non-inductive conditions (Gregory, 1956; Chapman, 1958). The region of
perception of the stimulus was the terminal bud, including leaves less than
5 cm long. The stimulus was apparently only slowly transmitted through
other parts of the plant (Chapman, 1958).
The nature of this photoperiodically controlled stimulus to tuber forma-
tion has been investigated in various ways. Both Gregory (1956) and
Chapman (1958) showed that it could be transmitted across a graft union.
If scions from SD plants were grafted on to LD stocks, the latter began to
form tubers in 14 days; if LD scions were used no tubers were formed. In
single-node segments of stem excised from induced plants and grown in
sterile culture, a tuber developed from the axillary bud; only vegetative
shoots developed in the case of segments from non-induced plants
(Gregory, 1956; Chapman, 1958). This work was later confirmed (Forsline
and Langille, 1976) and has recently given rise to a number of experiments.
Gregory (1956) concluded that a graft-transmissible tuber-inducing
stimulus was formed under specific conditions of photoperiod and
temperature, and occurred throughout the plant. Madec (1963) considers
that the stimulus is produced in tuber tissues, the relative role of tuber and
foliage in the inductive process being variable. In seedlings, however,
tubers must be induced by the leaves only (Madec and Perennec, 1959),
and this must be true also for cuttings. In experiments with cuttings
involving the excision of various parts, Kahn et al. (1983) showed that the
shoot apex alone could induce tuberization, although expanding leaves
were more effective.
It has been shown by further grafting experiments that the foliage of a
tomato scion cannot induce tuber formation in a potato stock, but can
support the development of tubers which have been previously induced
(Madec and Perennec, 1959). More recently, potato stocks grafted to SD-
requiring tobacco scions formed tubers only when the scions were given SD
(Chailakhyan et al., 1981; in Russian, cited by Ewing, 1985). They also
formed tubers when the LD plant Nicotiana sylvestris was the scion, and
was exposed to LD (Martin et aI., 1982). Ewing (1985) points out that this
suggests that the tuber-inducing hormones may not be specific to the
potato but of widespread occurrence, and may even equate to f1origen;
however, some hybrid potatoes flower in LD or SD and form tubers in the
converse conditions (Martin et aI., 1982), which might suggest that the
stimuli are not identical. Steward et al. (1981) showed that in a potato
hybrid SD and low temperatures promoted tuber formation, whereas LD
and high temperatures retarded tuber formation but induced flowering.
Kumar and Wareing (1973) carried out a series of ingenious grafting
experiments on Solanum andigena, in which tuber formation can be
induced by 10 SD, although maximal tuber induction only occurs after
20 SD cycles. This species has an absolute SD requirement for tuber
formation. They showed that, as in the cultivated potato, SD scions grafted
on to LD stocks induced tuber formation in the stocks in 2 weeks, and that
none was formed in LD/LD grafts, Cuttings made from plants maintained
in SD also formed stolons with small tubers at the basal node, whereas no
tubers formed on cuttings of LD plants (Fig. 3.35). Kumar and Wareing
concluded that a stimulus to tuber formation was formed under SD
conditions, and in further experiments showed that the stimulus could
move upwards, and could even move acropetally across a graft union.
Okazawa and Chapman (1962), using the cultivated potato, also concluded
that the movement of the stimulus was not polar; nor was that of
gibberellins, believed to inhibit tuber formation.
Tuber formation may be regulated by the relative activity of several
substances rather than the absolute concentration of a single substance
(Slater, 1968; Hammes and Nel, 1975), and the substances involved may be
the as yet unidentified tuber-forming stimulus, and natural gibberellins
(Okazawa and Chapman, 1962). The point has been made, however, that
some positive stimulus to tuber formation must exist in induced plants,
since it would be difficult to explain the results of grafting experiments
solely in terms of an inhibitor, such as gibberellin, present in non-induced
plants (Gregory, 1956; Kumar and Wareing, 1974). Other experiments
also support the view that SD induction involves increased production of a
factor that promotes tuberization (Wareing, 1982).
Ewing (1978) established the critical photoperiod (CPP) for a number of
clones by growing populations of seedlings under initially long photo-
periods that were progressively shortened, Apical and single-node cuttings
were taken every fortnight, and the photoperiod that first promoted
tuberization in the cuttings (CPP) determined. A quantitative effect of
photoperiod was demonstrated. If the plants from which the cuttings had
been taken had been exposed to photoperiods longer than the CPP, the
basal bud on apical cuttings, buried underground, failed to develop; with
progressively shorter photoperiods it developed as a shoot or stolon, a
stolon with a terminal tuber, or a sessile tuber (Fig. 3.36) (Ewing, 1978,
1981; Ewing and Wareing, 1978). The cuttings themselves were maintained
under LD. Using single-node cuttings, it was found that all the nodes of a
plant responded in much the same way (Ewing, 1978). On the other hand,
Pereira and Vellio (1984) found that single-node cuttings from the apical
and basal regions of the stem of cv, Aracy were less sensitive to SD than
those from the mid-region. Treatment with GA prevented tuber formation
and led to elongated orthotropic shoots (Pereira and Valio, 1984). In
multiple-node cuttings strongest tuberization occurred at the most basal
nodes; it appeared that tuberization was expressed most strongly by buried
buds furthest from leaves or from portions of stem exposed to light (Kahn
and Ewing, 1983; Ewing, 1987). In six-node cuttings with a single bud
remaining, however, the presence of leaves stimulated tuberization (Kahn
et ai., 1983). Ewing (1985) suggests a possible explanation for the effect of
the stem: that the tuberization stimulus might move beyond the bud
into the stem and thus be less concentrated in the bud. The possibility of
B
c o
Figure 3.36 Apical cuttings, kept in a mist bench for 11 days, with the basal node
below soil level (dotted line). A to D were taken from plants which respectively
received increasing numbers of inductive SD. (A) The bud at the basal node failed
to grow. (B) The bud developed as a stolon (or a leafy shoot). (C) A stolon with a
terminal tuber developed. (D) The bud developed as a sessile tuber. (Fig. 1 from
Ewing, 1987). (Reprinted by permission of Kluwer Academic Publishers.)
a gradient of the tuberizing stimulus along the stem should also be
considered.
Supporting the view (Ewing, 1978) that in cuttings stolons represent
buds that are less fully induced than tubers is the observation that 8-h days
stimulated numerous buds that did not form tubers to grow out as leafy
shoots or stolons. A similar effect was obtained in clones grown at day
lengths slightly shorter than their CPP (Ewing and Wareing, 1978).
High nitrogen supply is inimical to tuberization, and can prevent it even
under inductive conditions. It apparently acts by affecting levels of
endogenous hormones (Krauss, 1985).
(b) Experiments in culture

Barker (1953) showed that isolated nodes of etiolated potato sprouts could
be grown in aseptic culture. He obtained outgrowth of roots and stolons,
some of which later gave rise to tubers. Other workers have utilized
this technique for growing isolated nodal pieces (e.g. Gregory, 1956;
Chapman, 1958; Peterson and Barker, 1979), and cultured stolons or
isolated stem segments have been used in studies of the effects of
hormones on tuber formation (Palmer and Smith, 1969b, 1970; Smith
and Palmer, 1970; Garcia-Torres and Gomez-Campo, 1973; Palmer and
Barker, 1973).
Peterson and Barker (1979) showed that the structure of the small tubers
induced in vitro was basically the same as that of field-grown tubers.
However, Ewing (1985) has pointed out that in some experiments where
repeated sub-culturing in the dark is carried out, depletion of endogenous
growth substances may occur.
Single-node cuttings of etiolated shoots grown in vitro without GA
tuberized, whereas in the presence of GA only stolons were obtained
(Iriuda et aI., 1983). In similar experiments with cv. Russet Burbank,
tubers were formed in the presence of coumarin and 6-8% sucrose
(Stallknecht and Farnsworth, 1982a, b). Abscisic acid, and to a lesser
extent GA, inhibited this tuberization (Stallknecht and Farnsworth, 1982a)
as also did auxins (Stallknecht, 1985). Although inhibitors of protein and
nucleic acid synthesis reduced or delayed tuberization (Stallknecht and
Farnsworth, 1982b), they did not seem to prevent it.
Nodes cultured in vitro tuberized more quickly than those left attached
to the tuber (Tizio and Tizio, 1981). Segments from etiolated shoots
tuberized when provided with zeatin riboside, but not in the presence of
ethylene precursor or releasing agent. Tuberization was inhibited by GA.
It was considered unlikely that the cytokinin was the direct cause of
tuberization, partly because it had no effect at low sucrose concentrations,
and partly because the level of cytokinin was no greater than that in
segments which did not form tubers (Koda and Okazawa, 1983a).
However, Mauk and Langille (1978) obtained convincing evidence of tuber
The tuber 119
induction by zeatin riboside. Tuberization occurred in excised stolons from
SD, but not from LD, plants when cultured in medium with BA. Single-
node leafless cuttings from LD plants also failed to form tubers when
cultured with BA or kinetin (Wareing, 1983).
Serial culture of axillary shoots of potato resulted in the formation of
small tubers, minitubers, in senescent cultures (Hussey and Stacey, 1981).
These structures have proved useful for the storage and transport of
germplasm, and can now be induced in a matter of weeks in the presence of
BA and 6% sucrose. Addition of CCC enhanced the effect of the BA (Fig.
3.37) (Hussey and Stacey, 1984). Some of the results obtained in these
experiments were somewhat unusual; for example, GA gave rise to
elongated leafy shoots and BA to stolons. Addition of CCC resulted in
tubers being formed on shorter stolons, and above the level of the agar.
The effect of CCC suggested that formation of a tuber at the tip of a stolon
depends upon a low level of endogenous gibberellin. Tizio (1969, 1972)
Figure 3.37 Shoot cultures of cv. Red Craig'S Royal which have been induced to
form minitubers. (a) Grown in 8-h days with 2.0 mg I-I BA. (b) Grown in 8-h days
with 2.0 mg 1- 1 BA and 500 mg 1- 1 CCC, showing an extreme effect on tubering.
(Fig. 2B and C from Hussey and Stacey, 1984.)
had previously shown that CCC induced earlier tuberization in segments of
sprouts cultured in the dark. However, Abdala de Bottini et al. (1981)
found that treatment with CCC actually enhanced levels·of gibberellins at
the time of stolon formation. Stimulation of tuberization by ABA only
occurred under certain circumstances (Hussey and Stacey, 1984). There
were considerable differences between cultivars in these experiments;
perhaps this suggests differences in their endogenous hormones.
(c) Effects of hormone treatment

Auxins such as IAA, NAA or 2,4-D seem to increase the size and
sometimes earliness of tubers (Madec and Perennec, 1959; Harmey et al.,
1966), but not to have any inductive effect. Indeed, IAA was inhibitory at
certain concentrations (Kumar and Wareing, 1974). However, a mixture of
NAA and BA led to great numbers of tubers as well as to increased tuber
volume (Ahmed and Sagar, 1981).
Treatment of potato plants with gibberellic acid delayed tuber formation
even under SD (Slater, 1963). Fewer tubers developed per plant, and there
was a tendency for tubers to grow out as stolons (Lovell and Booth, 1967).
Cuttings of induced plants of Solanum andigena also failed to form tubers
when treated basally with a solution of GA (Kumar and Wareing, 1974). In
cuttings of S. tuberosum, too, GA reduced or eliminated tuberization
(Menzel, 1980; Pereira and Valio, 1984), even when applied in 10 ~g
quantities to the leaf (Ewing, 1985). Tuberization was retarded or pre-
vented when GA was supplied to nodal segments in culture (Harmey et al. ,
1966; Tizio, 1972); see also above.
This inhibitory effect of GA on tuber formation has led to the experi-
mental application of growth retardants such as CCC, which affect the
synthesis of endogenous gibberellins, to test whether they have a stimu-
latory effect. When tubers were treated with CCC, or the plants were
watered with this substance, tubers were formed about 5-10 days earlier
than in controls, depending on concentration (Gifford and Moorby, 1967;
Tizio, 1969). In other experiments, CCC was either applied as a soil drench
to LD plants, sprayed on the foliage, or applied in solution to plugs of
tuber. Tuber formation was induced (Kumar and Wareing, 1974; Hammes
and Nel, 1975) or enhanced (Shade que and Pandita, 1982).
In plants of Solanum andigena sprayed with ABA, tuber formation was
promoted, and in the cultivated potato, cvs Ulster Premier and Ulster
Prince, a greater weight of tubers was obtained (EI-Antably et al., 1967).
Using S. tuberosum cv. White Rose in LD, Smith and Rappaport (1969)
were unable to induce tuber formation by spraying the plants with ABA or
by treating the stolon tips in culture. In other experiments with cultured
stolons, ABA prevented tuber formation (Palmer and Smith, 1969b). In
cuttings from plants grown at (inhibitory) high temperatures ABA pro-
moted the formation of sessile tubers, but in plants grown at lower
The tuber 121
Figure 3.38 Single node cuttings from induced plants of Solanum andigena, i.e.
grown in short days, maintained in the dark with the leaf removed: (a) control; the
bud grows out as an etiolated shoot; (b) treated with ABA; the bud develops as a
tuber; (c) treated with GA; the bud develops as an elongated shoot with an apical
hook; (d) treated with kinetin; the bud develops as a shoot with slight swelling of
the basal node. (By courtesy of Professor P.F. Wareing.)
temperatures its only consistent effect was on tuber size (Menzel, 1980). In
experiments with S. andigena ABA seemed to be able to substitute for an
induced leaf. In single node cuttings from induced plants with the leaf
attached, the axillary bud rapidly forms a tuber. If the leaf is removed,
however, and the segment is kept in the dark, the bud grows out as an
elongated, etiolated shoot (Fig. 3.38a). When ABA was supplied in
solution through the base of a leafless segment, the axillary bud developed
as a tuber (Fig. 3.38b). When GA was similarly supplied, the bud
developed as an elongated shoot with an apical hook (Fig. 3.38c). Kinetin
caused some swelling of the basal internode (Fig. 3.38d), but did not lead
to tuberization. Under these conditions, therefore, ABA can substitute for
the effect of the SD-induced leaf and cause tuberization of the axillary bud
(Wareing and Jennings, 1979). Further experiments showed that labelled
ABA moved from leaves into the buds, and that both induced and non-
induced leaves (or exogenous ABA) could stimulate tuber formation in
induced stems. It was postulated that a 'second factor' accumulated in
stems of induced plants, so that a non-induced leaf could then promote
tuber formation (Wareing and Jennings, 1979). Melis and van Staden
(1984), however, consider that ABA plays an indirect role, by suppressing
shoot growth. Ewing (1987) has also cogently pointed out that a mutant of
the potato which lacks ABA still forms tubers.
Excised stolons of S. tuberosum grown in aseptic culture formed tubers if
supplied with a cytokinin. Of those tested, kinetin was the most effective,
inducing 80-100% tuberization. Controls did not form tubers (Fig. 3.39)
(Palmer and Smith, 1969a, b). A certain period of continuous treatment
was required. Kinetin-induced tuber formation could be completely in-
hibited by a temperature of 35°C (Palmer and Smith, 1969b, 1970). As
already mentioned, cytokinins induced tuberization in induced, but not
non-induced, stolons of S. andigena (Wareing, 1983). By supplying
kinetin-8- 14C to cultured stolons it 'ivas shown that the labelled material
accumulated at the site of tuberization, although even more label was
found at the base of the stolon, perhaps because of the proximity of the
label in the medium. The extracted labelled material did not have the same
R f value as kinetin (Smith and Palmer, 1970).
Experiments with excised nodal segments of induced and non-induced
potato plants, cv. Katahdin, showed that the presence of kinetin in the
culture medium could substitute for the stimulatory effect of induction on
tuber formation in the axillary buds (Forsline and Langille, 1976). On the
other hand, single-node cuttings from induced plants of cv. Katahdin
expressed (in the morphology of the buds) a lower level of induction than
they had in fact received, when immersed daily in a solution of BA
(McGrady et al., 1986). This was indicated by a lengthening of the bud.
Possibly the cytokinin content of induced cuttings might now have been
supra-optimal, but the results are still surprising. It was suggested that the
mobilizing properties of the cytokinin, applied after 4 days, might have
The tuber 123
Figure 3.39 Cultured stolons of cv. Norgold Russet. Right, grown in control
medium; left, grown for 30 days in the presence of 2.5 Ilg ml- 1 kinetin. Tubers have
developed behind the stolon tips. (Fig. 3 from Palmer and Smith, 1970.)
impeded the flow of substances from the leaf to the bud, or have caused
them actually to move back into the leaf (McGrady et al., 1986).
The results of some of these experiments suggest that a natural cytokinin
could be the tuber-forming stimulus. When applied to the basal end of
cuttings, however, kinetin inhibited tuber development at high concentra-
tions although it slightly stimulated it at the lowest concentration used
(Kumar and Wareing, 1974). Since here, too, the cuttings were from
induced plants, however, these results are somewhat analogous to those
discussed above, and need not be at variance with the postulated occur-
rence of an endogenous cytokinin capable of inducing tuber formation.
Melis and van Staden (1984), however, consider that cytokinins may be
involved in regulating the growth of tubers rather than in inducing their
formation; and Vreugdenhil and Struik (1989) consider that cytokinins
cannot be the only factor involved in tuber initiation.
It is gradually becoming apparent that the site of hormone treatment
may be an important factor affecting the result obtained. Together with the
use of S. andigena, which requires SD for tuber formation, and S.
tuberosum, which has no such absolute requirement, the widely differing
treatments may account for the conflicting results that are sometimes
obtained.
There is some evidence that ethylene, also, may playa role in tuber
formation. When sprouts of cv. Arran Pilot bearing several stolons were
treated with gaseous ethylene, extension growth was inhibited and swelling
of all rapidly growing regions occurred. Sub-apical swellings on stolons
were similar morphologically to normal tubers, but contained no starch
(Catchpole and Hillman, 1969). In sprouts that developed from tubers of
cv. Irish Cobbler treated with ethylene, however, much more starch was
present than in controls (Minato et al., 1979). Ethrel, a substance which
releases ethylene, led to an increase in tuber formation when applied to the
soil (Garcia-Torres and Gomez-Campo, 1972). In cultured stem segments,
also, ethrel enhanced tuber formation and counteracted the effects of
applied GA (Garcia-Torres and Gomez-Campo, 1973). On the other hand,
cultured stem segments supplied with 2-chloroethylphosphonic acid,
another substance which releases ethylene, formed stolons which were
generally swollen, but without localized tuberous swellings (Palmer and
Barker, 1973). Restricted gaseous exchange, believed to result in accumu-
lation of ethylene, inhibited tuberization in cultured nodes (Hussey and
Stacey, 1984). Thus ethylene may play some part in the process of stolon
and tuber formation, but is probably not the sole stimulus involved. These
observations also emphasize the need for accompanying anatomical work,
to establish the degree to which the induced tuber is structurally normal.
(d) Carbohydrate metabolism

A number of workers have concluded that sucrose is required for, but does
not induce, tuber formation (Gregory, 1956; Madec and Perennec, 1959;
Madec, 1963). In cultured stem segments, tubers were formed in the
presence of 8%, but not 2%, sucrose (Harmey et al., 1966). In other
experiments with cultured stolons, however, tubers did not develop in the
presence of sucrose unless kinetin was also supplied. For the induction of
tuber formation by kinetin, 6%, 8% or 10% sucrose was required, 2% or
4% being inadequate (Palmer and Smith, 1970). Sucrose, and especially
mannitol, promoted tuber formation in single node cuttings of Solanum
andigena (Wareing and Jennings, 1979).
Tuber formation seems to be associated with a high concentration of
soluble sugars in the stolon tip, and with conditions which lead to this
(Slater, 1963, 1968; Burt, 1964). In cultured stolons treated with kinetin
the level of reducing sugars decreased, as compared with controls or
ethylene-treated stolons. This coincided with the rapid synthesis of starch
in cultures with kinetin (Palmer and Barker, 1973).
Lovell and Booth (1967) considered that the first sign of starch deposition
The tuber 125
was a reliable indicator of tuber initiation. Starch accumulated in a
localized sub-apical region of the stolon tip prior to any external sign of
tuberization. Palmer and Smith (1969a) noted that starch accumulated in
the apical region of kinetin-treated stolons, but not in control stolons. They
considered that this provided some evidence that cytokinins may act by
mobilizing metabolites to the site of tuber formation. Stolons themselves,
however, have a capacity for storing starch (Engels and Marschner, 1986).
It has already been noted that starch was absent from tuber-like structures
induced by ethylene (Catchpole and Hillman, 1969). Smith and Palmer
(1970) have suggested that kinetin may exert its stimulatory effects on
tuber formation by promoting the activity of starch synthetase and
suppressing that of starch hydrolase. Gibberellic acid, on the contrary, may
have the converse effect on these enzymes, thus inhibiting tuber formation.
In stolons induced by GA, only small starch grains (2-5 /-lm) were present,
in contrast to larger ones (10-25 /-lm in diameter) in developing tubers.
Levels of soluble phosphorylase increased in the stolons along with the
small starch grains (Iriuda et al., 1983). Treatment of cuttings from induced
plants with GA led to a decrease in the amount of starch present in the
buds (Pereira and Vellio, 1984). As stolon tips developed into tubers in cv.
Irish Cobbler, the amount of starch increased rapidly, and also the
activities of bound synthetase and phosphorylase (Obata-Sasamoto and
Suzuki, 1979a). An increase in starch and phosphorylase activity was also
evident in cultured stolons (Obata-Sasamoto and Suzuki, 1979b). It has
been suggested that GA may modify levels of phosphorylase (Pereira and
Valio, 1984).
(e) Analysis of endogenous hormones
Endogenous hormones present in the potato plant have been extracted and
identified, and the effects of various environmental conditions on the levels
of endogenous hormones have been studied.
Indoleacetic acid was extracted from potato tubers and identified by
chromatography (Booth and Wareing, 1958). As will be shown in Section
3.7.2, gibberellins have been extracted from potato tubers. In one
investigation, these increased after planting, and especially at the time of
stolon formation, falling off when tubers were formed in the daughter
plant. It was suggested that the mother tubers supplied a continuous flow
of gibberellins or precursors to the developing plant (Abdala de Bottini
and Tizio, 1981). Under storage conditions, potato tubers also produce
ethylene (Creech et al., 1973). The inhibitor abscisic acid (ABA) has also
been extracted from potato tubers (Cornforth et al., 1966).
Although it is considered that the tuber-forming stimulus is present in
the tuber as well as in aerial parts of the plant, neither seedlings nor
cuttings have attached tubers, yet both can form tubers. Thus there is no
evidence that any of these substances isolated from tubers is the elusive
tuber-inducing stimulus in question.
The hormones extracted from the leaves have chiefly been gibberellins.
By means of thin layer chromatography, at least four zones or peaks of
gibberellin-like activity were found in leaves of Solanum andigena (Railton
and Wareing, 1973a). Levels of gibberellin were significantly lower in SD
than in LD plants (Kumar and Wareing, 1974). When plants which had had
five SD cycles were irradiated with red light for 30 min during the long dark
period of the sixth SD cycle, there was an increase in the content of
gibberellins so that levels approached those of LD plants, apparently
because of an increased rate of synthesis. There were also changes in the
relative amounts of the four peaks (Railton and Wareing, 1973b).
Levels of two out of four cytokinins extracted from above-ground parts
of potato plants were significantly higher in induced than in non-induced
plants, reaching a maximum several days before tuber initiation (Forsline
and Langille, 1975). Levels of zeatin riboside were greatest in both above-
and below-ground parts 4 days after inductive conditions began, and tuber
formation began 4 days later (Mauk and Langille, 1978). Shoots can
evidently synthesize their own cytokinin in the absence of roots (Wang and
Wareing, 1979). A transformed line of cv. Maris Bard containing T-DNA
from Agrobacterium tumefaciens, which showed an exceptionally strong
tendency to tuberize, had very high concentrations of cytokinins in
the shoots. Shoots of the transformed plants had 100-200-fold more
cytokinins, and 3-4-fold less auxins, than normal shoots (Ooms and
Lenton, 1985).
Levels of endogenous ABA were slightly higher in leaves of LD plants of
Solanum andigena than in plants induced with SD, although there may
have been more bound ABA in the latter. In experiments with 14C-Iabelled
ABA, radioactivity was found in the buds, indicating that there was
movement from leaf to bud (Wareing and Jennings, 1979). Interruption of
the supply of nitrogen to the shoot, which favours tuberization, led to a
considerable increase in ABA content, and some decline in gibberellins
(Krauss, 1985).
Recently tuber-inducing substances have been extracted from both old
tubers and leaves and assayed against single-node cuttings. Increasing
concentrations of the extract induced tubers on shorter stolons, similar to
the effects obtained by Ewing (1978) with daylength (Fig. 3.36) (Koda and
Okazawa, 1988). A purified aqueous extract appeared to be a growth
inhibitor, and was thought to be a glycoside (Koda et al., 1988).
More recently, attempts have been made to assess the endogenous
hormones in stolon tips that are undergoing tuberization. In tips that were
just swelling, the content of auxin was high and that of cytokinin low; at
later stages, auxin decreased and cytokinin increased. Gibberellin content
was low at all stages (Obata-Sasamoto and Suzuki, 1979a). In sections of
etiolated shoots grown in vitro, no cytokinin activity was found in the small
The tuber 127
tubers, although it was present in the basal parts of the stolons (Obata-
Sasamoto and Suzuki, 1979b). Since kinetin was supplied in the medium,
this may not be significant. In field-grown stolon tips, however, Koda and
Okazawa (1983b) found that auxin reached a maximum in swelling stolons
and then decreased, cytokinin was initially low and increased as the tips
swelled, and an ABA-like substance followed a similar pattern, but
continued to increase. In other experiments, non-induced stolon tips and
small tubers contained several cytokinin-like compounds at low levels; in
tubers larger than 7.5 mm in diameter the levels, especially of components
resembling zeatin and zeatin riboside, increased (Jameson et al., 1985).
Von Staden and Dimalla (1977) found that stolon tips and small tubers
contained more cytokinin than larger tubers, but the smaller ones were
'little potatoes' and may possibly be somewhat anomalous.
Although the occurrence of higher levels of giberellins in LD plants is
compatible with the failure of such plants to form tubers, in view of the
known inhibitory effects of GA on tuber formation, it must be recognized
once more that a difficulty exists in attributing tuber formation to a
negative condition such as the lower concentration of an inhibitor. As
Kumar and Wareing (1974) have pointed out, low levels of gibberellin in
SD scions are unlikely to lead to reduction of gibberellins in LD stocks, yet
in SD/LD grafts these form tubers. This positive stimulus to tuber
formation as yet remains as elusive as the flowering hormone.
(f) Conclusions
The literature relating to the control of tuber formation, aspects of which
have been reviewed by Wareing and Jennings (1979), Ewing (1985, 1987),
Krauss (1985), Stallknecht (1985) and Vreugdenhil and Struik (1989),
remains confusing. Numerous techniques have been employed to in-
vestigate it in numerous cultivars, and Stallknecht (1985) has pointed out
that comparison of different in vitro methods under similar conditions,
including careful specification of the nutrients in the medium (which may
affect the action of growth substances), would be helpful. The use of a
single cultivar for study by different methods would also be helpful.
Some conclusions, however, may be drawn. It seems certain that
gibberellins promote stolon formation, and inhibit tuberization. Ewing
(1978) has suggested, however, that vegetative buds, stolons, tuberizing
stolons and sessile tubers form a continuum representing increasing levels
of induction. Thus what appears to be (and normally is) a two-stage
sequence, stolon then tuber, can be by-passed, a tuber forming directly.
This may indicate that the balance between environmentally-controlled
growth substances changes, sessile tubers developing when the process is
abnormally hastened. Candidates for the second, or later, substance(s)
are, in the main, cytokinins and abscisic acid. Wareing and Jennings (1979)
have argued that ABA may promote tuberization by inhibiting the growth
of the apical meristem, and that a 'second factor' , a growth promoter, may
exist in the stem and be of a new type, with effects not specific to
tuberization. Having regard to the development of the tuber (see Section
3.6.2), it appears possible, at least, that several known growth substances
could act (normally) in sequence; for example, GA would promote stolon
formation, IAA cell enlargement in the subapical region of the stolon,
ABA cessation of apical growth (perhaps maintained throughout
dormancy of the tuber), and cytokinins cell division in the slightly later
stages of development of the young tuber. This sequence is compatible
with that of the endogenous hormones actually found in tuberizing stolon
tips (Obata-Sasamoto and Suzuki, 1979a; Koda and Okazawa, 1983b).
There might, of course, be as yet unknown substances involved. But if
several substances normally operate in sequence, the somewhat anomalous
structures, recognizable as tubers but not entirely normal, obtained in
some experiments might be more readily interpreted. We have seen
already that at least one phase of the normal development, stolon
formation, has been eliminated in the strongly induced sessile tubers,
which are perhaps the endpoint of an abnormally rapid sequence.
However, this serial sequence of events must be capable of occurring
locally, out of phase with other regions, since, as has been pointed out
(Vreugdenhil and Struik, 1989), different stages of development may occur
in the plant simultaneously. To this must be added, of course, the
stimulation of starch-producing enzymes. Since these putative growth-
regulating substances are affecting very general processes, such as cell
enlargement and cell division, it is of course possible that other substances
or combinations of substances (whether normally endogenous or not),
could mimic their effects, again providing a possible explanation for some
of the experimental results obtained. It would be helpful if the structure of
experimentally induced organs were examined.
3.7 DORMANCY AND ITS CONTROL
Growth of the buds on potato tubers is subject to a number of rather

different controls. During the development of the tuber there is no true
bud dormancy, but correlative inhibition is operative. After harvest other
mechanisms supervene (Madec and Perennec, 1969). One of these is
classed as true dormancy, that is, a state in which the tuber will not sprout
if stored at a temperature below the optimum. According to the variety,
the period of dormancy may be from 18 to 33 weeks, usually 20-23 weeks.
However, immediately after harvest potato tubers cannot be induced to
sprout even under optimum environmental conditions. This is usually
termed the rest period, a somewhat unsatisfactory term, and is of 5-19
weeks duration in different varieties (Emilsson, 1949). It is not always clear
whether authors are referring to rest or true dormancy, as thus defined.
Dormancy and its control 129
Hemberg (1985) has recently reviewed rest in the potato. In the so-called
rest period buds fail to grow even under optimum temperature and light
conditions, and there appears to be an internal 'clock' which triggers
cellular events leading to sprouting (Rappaport and Wolf, 1968a, 1969).
Tuan and Bonner (1964) consider that while the genome of dormant buds
is largely repressed, it is not yet established that this is the primary cause of
dormancy. Others believe that the tuber may be temporarily unable
to supply the buds with metabolites essential for growth (Madec and
Perennec, 1969).
Treatment of excised plugs of tuber tissue with ABA inhibits sprouting,
whereas GA is slightly stimulatory. Maleic hydrazide inhibits sprouting in
all the varieties tested, probably by inhibiting protein and nucleic acid
synthesis. The methyl ester of naphthaleneacetic acid also inhibited sprout
growth (Stallknecht, 1983). Ethylene inhibited growth of the apical sprout,
but led to outgrowth of swollen sprouts from the lateral buds (Minato et
al., 1979), as did thiourea (Stallknecht, 1983). Various other substances,
including a mixture known as Rindite, and several cytokinins, can break
the rest period (Stallknecht, 1979; Hemberg, 1985). Dormancy and
sprouting of the potato tuber are probably controlled by a balance of
inhibiting and promoting substances (Bruinsma and Swart, 1970), perhaps
endogenous ABA and gibberellins (Wareing, 1982); it seems likely that
much more work will have to be devoted to the elucidation of the
mechanisms involved.
3.7.1 Changes occurring at sprouting

When the rest period is terminated and the buds of the tuber resume
growth, various changes occur both in the bud meristem itself and in the
tuber.
(a) Effects on the tuber

When the buds start to grow, starch is broken down in the tuber. This
ceases if the buds are removed. Thus if a hormone is involved the tuber
must require a continuous supply of it to maintain starch breakdown
(Edelman et at., 1969). These workers suggest that such a hormone might
be required for synthesis of an enzyme, and that the rate of its synthesis
might control further sprout growth. In view of the involvement of
gibberellin in the control of sprouting, it is interesting that GA treatment'
did not modify a-amylase activity in buds, although it is suggested that it
may modify levels of phosphorylase (Pereira and Veilio, 1984). Other rest-
breaking substances did increase the activity of a-amylase and other
enzymes (Hemberg, 1985). During dormancy crystals are formed in
peripheral cells of tubers. It is thought that they may be formed within
vesicles of the endoplasmic reticulum. The crystal-containing bodies
Figure 3.40 Autoradiographs of LS shoot apex after being supplied with 3H_
thymidine for 12 h: (a) water control; (b) treated with GA. (Plate 5 from
Rappaport and Wolf, 1969.)
may be sites of protein storage; they disappear after sprouting (Marinos,

1965).
(b) Changes in the bud meristems

Changes in activity of the bud meristems when the rest period is terminated
have been studied principally by labelling with radioactive isotopes, mainly
the precursors of RNA and DNA. Usually 3H-uridine or 3H-thymidine was
Dormancy and its control 131
applied to apices of buds on plugs of tuber tissue, and in some experiments
GA was also applied. Buds were subsequently sectioned and histoauto-
radiographs made. Using this method, RNA synthesis was detected in the
smaller plugs (0.2 X 0.2 cm) 6 h after excision, and a low level of DNA
synthesis 12 h after excision. On larger plugs, RNA synthesis was first
detected at 12 h and DNA synthesis at 24 h after excision. Buds treated
with GA showed more labelling in the first 12 h (Fig. 3.40); DNA synthesis
was accelerated in the period from 6 to 12 h after excision (Rappaport and
Wolf, 1968b, 1969). In all these experiments synthesis of RNA preceded
that of DNA (Rappaport and Wolf, 1968a, 1969), and both preceded cell
expansion and cell division in the buds. Rappaport and Wolf (1969)
consider that since RNA synthesis increased after excision of the plugs
whether GA treatment was given or not, it is doubtful whether gibberellin
plays a primary role in the termination of rest. However, gibberellin may
be synthesized in the cells along the cut surface of tuber tissue (Rappaport
and Wolf, 1968b).
Treatment of buds with ABA considerably reduced the incorporation of
labelled uridine and thymidine. If ABA and GA were applied together,
ABA overcame the ability of GA to promote synthesis of RNA and DNA.
This is thought to indicate that ABA has a direct effect on the synthesis of
nucleic acids, rather than an indirect one such as blocking synthesis of
gibberellins (Shih and Rappaport, 1970).
Treatment of tubers with ethylene chlorhydrin led to incorporation of
label in the buds (Tuan and Bonner, 1964; Rappaport and Wolf, 1968a).
The rate of RNA synthesis doubled within 2 days after the end of
treatment, and by 10 days had risen to 130 times the level in dormant buds.
Rate of DNA synthesis was similar but with a longer lag period (Tuan and
Bonner, 1964).
Tuan and Bonner extracted chromatin from tubers and from dormant
buds or those which had been given ethylene chlorhydrin treatment.
Chromatin from tubers was quite ineffective in supporting DNA-dependent
RNA synthesis, and that from dormant buds was inactive, whereas
chromatin from buds after treatment was ten-fold more effective.
The RNA which is synthesized in buds after treatment with ethylene
chlorhydrin is DNA-dependent, since actinomycin-D inhibits its synthesis
in vivo. Tuan and Bonner concluded that the genome of dormant potato
buds is largely repressed. It is thus rather surprising that RNA synthesis in
activated buds always seems to precede DNA synthesis.
Marinos (1967) studied the fine structure of dormant potato apices,
cv. Sebago. Plastids contain osmiophilic droplets and starch grains.
Intraplastid bodies, believed to be sites of accumulation of nucleic acids
and protein, are prominent in the apical dome and young leaf primordia of
dormant buds, but tend to disappear when sprouting begins, along with
phytoferritin granules. In resting buds of cv. White Rose potatoes, striking
concentric configurations of endoplasmic reticulum (ER) were observed
(a)
(b)
(e)
Figure 3.41 Electron micrographs of part of the apical dome of potato buds 12 h
after they were excised and treated with (a) water; (b) GA; and (c) ABA. Large
vacuoles are present in cells treated with ABA. x 3800. (Fig. 1 from Shih and
Rappaport, 1971.)
Development and structure of the flower 133
(Shih and Rappaport, 1971). These consisted of 7-13 ER lamellae with
associated ribosomes. Such configurations were never observed in cells of
buds on plugs which had been excised for more than 6 h. They tended to
open out and by 12 h after excision only typical ER was observed. Thus this
opening out seems to coincide with the acceleration of RNA synthesis.
Treatment of buds with ABA or GA stimulated the opening and extension
of these concentric structures.
Treatment with ABA led to increased vacuolation (Fig. 3.41c), first notice-
able about 1 h after treatment, whereas GA-treated buds were not strikingly
different in ultrastructure from controls (Fig. 3.41a, b) (Shih and Rappaport,
1971). Rappaport (1972) suggested that ABA might therefore have some
active role, rather than the passive one of merely keeping cells quiescent. It
is also thought that this effect of ABA may explain the failure of GA
completely to reverse its inhibitory action (Shih and Rappaport, 1971).
3.7.2 Endogenous substances in tubers

Gibberellins and gibberellin-like substances have been extracted from
potato peelings. Using the dwarf maize bioassay, Smith and Rappaport
(1961) showed that the level of endogenous gibberellins remained low
during the rest period and increased towards the end of this time. Boo
(1962), on the other hand, found an increase in endogenous gibberellin-
like substances during the last part of the rest period.
In other analyses, gibberellin-like substances were detected in the
neutral fraction obtained from potato buds and peelings; these substances
stimulated elongation of dwarf peas but were inactive on dwarf maize
(Hayashi et al., 1962; Hayashi and Rappaport, 1966). The level of acidic
auxin was found to be very low during the rest period (Hemberg, 1985).
One of the most interesting findings was that gibberellin-like substances
increased markedly following wounding of the tuber tissues. It was
considered that the amount of such substances produced by an excised plug
of tuber tissue would be sufficient to trigger changes in the buds associated
with termination of dormancy (Rappaport and Sachs, 1967). Thus this may
account for the effects of wounding on bud growth.
3.8 DEVELOPMENT AND STRUCTURE OF THE FLOWER
Danert (1957) has described the cymose inflorescence of the potato. The
vegetative shoot is a sympodium; each portion terminates in an inflores-
cence, vegetative growth being continued by the bud in the axil of the last
true foliage leaf. This shoot appears to be laterally displaced. Danert
interprets the last leaf of the main axis as a bract subtending the
reproductive shoot, which is concaulescent with the parent axis. A second
part of the inflorescence is formed below the terminal flower without a
sub tending bract. In this way, helicoid monochasia are formed below the
terminal flower. Flowering was delayed or inhibited by low night temper-
atures (Haynes and Haynes, 1988). The flowers are actin om orphic and
hypogynous (Artschwager, 1918; Jones, 1939). According to Young (1923)
the calyx originates as a marginal ring with five lobes, but Sattler (1973)
reports for the flower of Solanum dulcamara that the sepals are initiated
separately in spiral sequence and the calyx tube is later formed by growth
of the tissue between these primordia; the author found the latter to be the
case also in S. tuberosum cv. King Edward. At maturity the calyx is green
and hairy and consists of five joined sepals. The five petal primordia form
in close sequence and the corolla tube develops later than the calyx tube.
The five stamens alternate with the petals and are borne on the corolla
tube. The anthers fuse postgenitally, enclosing the pistil (Sattler, 1973). At
maturity the stamens have short, stout filaments and long anthers. Pollen is
shed through pores at the tips of the anthers (Jones, 1939).
There are usually two carpels, the primordia of which are formed
simultaneously. These fuse to form a syncarpous, bilocular, superior ovary
with a long style and a bilobed stigma. The mature fruit is a green berry
with axile placentation, which often fails to develop in cultivated potatoes.
The carpels are orientated obliquely to the median plane of the flower (Fig.
3.42) (Jones, 1939).
Figure 3.42 Floral diagram of potato. (Fig. 56F from Jones, 1939.)
The sepals and petals each have from three to five vascular strands, the
median strand being the largest (Fig. 3.43) (Artschwager, 1918; Hayward,
1938). Rather large cells containing crystal sand (numerous small crystals
of calcium oxalate) are common in the calyx and carpels (Young, 1923).
Development and structure of the flower 135
Figure 3.43 TS young infertile flower bud of cv. King Edward. The bilocular
ovary (0) with placentae (pI) can be seen in the centre. Each stamen (st) has four
pollen sacs (ps). Marginal meristems are evident in the petal primordia (p, petal; s,
sepal). x 100.
Marginal meristems are evident in the developing petal primordia (Fig.

3.43).
The stamens have a single vascular strand. In the developing anthers the
sporogenous tissue forms a mass of densely staining cells, horse-shoe
shaped in cross-section (Fig. 3.43). The parietal layers are not modified to
aid in dehiscence, which takes place through terminal pores. The tapetal
cells are large (Young, 1923; Hayward, 1938). A glandular tapetum has
been reported (Davis, 1966). Meiosis in the anther takes place before that
in the ovule (Clarke, 1940). Just after mitosis in the microspore, the
generative cell in Solanum species contained both plastids and mito-
chondria; however, during development the plastids were gradually lost
from the generative, but not the vegetative, cell (Clauhs and Grun, 1977).
The occurrence of stamens bearing naked ovules at the junction of
anther and filament has been reported for quite a considerable proportion
of flower buds collected from crop fields (Pullo and Slusarkiewicz, 1975).
Only shrunken pollen grains were present and no fruit was set.
Pollen sterility occurs quite frequently, and depends partially on
environmental factors. One cause is apparently abnormal division of the
pollen mother cells resulting from higher temperatures. Under such
conditions abortive pollen grains were formed (Stow, 1927). The propor-
tion of abnormalities in meiosis may also be increased by virus and fungus
diseases (Khan, 1951).
Numerous ovules are formed, covering the placenta. A single archesporial
cell develops in the hypodermis, distinguishable by its dense contents and
deep staining (Young, 1922, 1923; Rees-Leonard, 1935). The archesporial
cell develops directly as a megaspore mother cell. Of the four megaspores
formed by meiosis, three degenerate, the megaspore at the chalaza I
end remaining functional (Rees-Leonard, 1935; Hayward, 1938). The
megaspore increases in length and undergoes nuclear division, the nuclei
moving to opposite ends of the embryo sac, which grows rapidly and
becomes curved. These two nuclei divide simultaneously, then the four
resulting nuclei divide again, producing four nuclei at each end of the
embryo sac. Walls are formed and a seven-celled megagametophyte
results. The antipodal cells are usually ephemeral (Lamm, 1937; Davis,
1966), and degenerate early, before fusion of the polar nuclei. The
synergids elongate and develop the filiform apparatus. The polar nuclei
fuse before fertilization. These events seem to be similar in at least three
cultivars, Irish Cobbler, Earlaine and Khatadin (Rees-Leonard, 1935;
Clarke, 1940; Williams, 1955). Ovules with more than one embryo sac
have occasionally been reported (Young, 1922; Lamm, 1937).
Fertilization has been observed 36 h (Clarke, 1940) or 4~5 h
(Williams, 1955) after pollination, doubtless depending on environmental
conditions. The usual type of double fertilization takes place. The endo-
sperm nucleus divides 60-70 h after pollination; an early free-nucleate
stage has not been observed (Clarke, 1940; Williams, 1955).
Various degenerative changes may occur at different stages during ovule
development; it seems likely that this may account in part for the poor
capacity for seed-setting of many commercial varieties, but more work
is required (Rees-Leonard, 1935; Clarke, 1940). Evident degenerative
changes in the developing flower buds, sometimes described as 'blasting',
include cessation of growth, development of a yellowish colour, and
wilting. Both the embryo sac and surrounding nucellus may shrivel, and
degeneration of the mature megagametophyte often occurs (Young, 1923;
Rees-Leonard, 1935). Degenerative changes in the embryo sac and ovule
seem to result from unfavourable environmental conditions, and are much
more uniform than those in the anther (Young, 1923).
Abscission of the flower often occurs before fruits and seeds become
mature. An abscission layer is formed at a region where there is a joint
in the pedicel. Weinheimer and Woodbury (1967) have described the
abscission layer in cvs Russet Burbank and Menominee. They found that
the abscission zone was not well defined. The protective layer developed
Em bryogenesis 137
when the buds were still small, and consisted of a large mass of small
meristematic cells. A separation layer could not be discerned, but at the
time of abscission whole areas of cells disintegrated and collapsed. The
abscission zone can be located externally by a hairy swelling, due to cell
enlargement, just below an indentation in the pedicel. This was evident at
an earlier stage of development of the flowers of cv. Russet Burbank, and
Weinheimer and Woodbury (1967) considered that it might be associated
with their premature abscission. These workers thought that abscission in
Russet Burbank might be by breakdown of the cells in the separation layer
of the middle lamella only, whereas in Menominee there was a complete
dissolution of the cells in this region. When bud blasting occurs the upper
part of the abscission layer splits, and callus is formed over the broken
surface (Hayward, 1938).
3.9 EMBRYOGENESIS
Embryo development is of the Solanad type, in which the terminal cell of

the two-celled proembryo divides by a transverse wall (Fig. 3.44)
(Maheshwari, 1950; Wardlaw, 1955; Davis, 1966).
,a~
CbV
Figure 3.44 Early development of Solanad type of embryo. The zygote divides by
a transverse wall to give a distal cell (ca) and basal c.ell (cb). The latter divides
transversely to give the most basal cell of the tetrad (ci) and an intermediary cell
(m). The distal cell divides transversely to give distal (cc) and subdistal (cd)
segments. When cc divides transversely, the distal segment I and adjacent segment
I' are formed. (Figs 55A, F and G from Wardlaw, 1955.)
The first division of the zygote is transverse and occurs 4 or 5 days after
pollination (Clarke, 1940; Williams, 1955). By further division a four-
celled linear proembryo is formed about 7 days after pollination (Hayward,
1938; Clarke, 1940; Williams, 1955). The basal cell plays a relatively minor
role in subsequent development, but divides to form a short suspensor of
two or more cells (Hayward, 1938; Maheshwari, 1950; Wardlaw, 1955;
Williams, 1955). There is no hypophysis (Soueges, 1922). The location and
planes of subsequent cell division are highly variable in Solanaceous
embryos (Johansen, 1950). Soueges (1922) considers that the terminal cell
of the pro embryo gives rise to the cotyledonary part of the embryo, the
next cell to the hypocotyl and initials of the central cylinder, and so on,
but, in agreement with more recent views on embryology in general,
Bhadurim (1936) found that definite parts of the embryo were not always
formed from a particular cell of the four-celled proembryo.
Whatever the precise sequence of cell divisions, a club-shaped embryo is
formed by 10 days, and a spherical embryo by 12 days, after pollination
(Williams, 1955).
In cv. Chippewa, seed failure is characterized by enlargement and
proliferation of the tissue surrounding the embryo sac; these cells spread
through the area normally occupied by the developing embryo. Placental
tissue also proliferates, increasing the diameter of the ovary, which
develops into a parthenocarpic fruit (Williams, 1955).
3.10 TISSUE CULTURE
Recently, various types of tissue culture have afforded ways of propagating

desirable varieties, of storing germplasm, of increasing variation, and of
obtaining material easy to transport to other laboratories or to other
countries. The topic has been reviewed by Wang and Hu (1985), and only
some examples of the usefulness of tissue culture will be given here.
3.10.1 Somatic tissues

Micropropagation using axillary buds as explants can lead to ten-fold
multiplication per month, and cultures can be stored at low temperatures
for long periods between sub-cultures (Wooster and Dixon, 1986).
Plantlets can be transferred to pots or peat blocks in a greenhouse before
planting out, maintaining high humidity (Wooster and Dixon 1986;
PospiSilova et al., 1988). Any abnormalities occurring during culture
disappeared after transplanting (PospiSilova et al., 1988).
As already pointed out (see Section 3.4.2), shoot tips or meristems can
be used as explants. The apical dome plus two to four leaf primordia of
several clones was successfully cultured by a two-stage procedure, yielding
single-shoot or multiple-shoot cultures (Westcott et at., 1977).
Other types of explant can also be used, such as pieces of leaf, stem or
tuber. Using cv. Superior, Jarret et at. (1980a) tested various regions of the
tuber for regeneration potential. They found that discs of perimedullary
and cortical tissue formed adventitious shoots under appropriate culture
conditions (including the presence of GA and a cytokinin), but those from
the pith region did not. Shoots developed from green protuberances
formed at the periphery of the disc (Fig. 3.45). Auxin was not essential, but
Tissue culture 139
Figure 3.45 Shoots which have formed from green protuberances on a tuber disc
of cv. Oneida after 6 weeks in culture. (Fig. 8 from Jarret et al., 1980b.)
a low concentration of NAA increased the number of shoots. In cv. Irish

Cobbler, zeatin was the most effective of the cytokinins tested, and sucrose
was inhibitory; best results were obtained with no added carbohydrate
(Kikuta and Okazawa, 1982). In cv. Superior, however, sucrose was
essential for shoot formation and BA was better than zeatin (Jarret et al.,
198Gb). These authors succeeded in obtaining adventitious shoots in eight
out of ten cultivars tested. Most of these shoots could then be established
in the greenhouse (Fig. 3.46) and transferred to the field.
Using a two-step protocol, Wheeler et al. (1985) obtained shoots from
the leaf, rachis or stem of 13 cultivars, and from pieces of tuber of seven
cultivars. Explants of Desiree could form the largest number of shoots on
the greatest range of media. There was some indication of chromosomal
instability in plants derived from tuber pieces.
Different culture conditions were optimal for different tissues (tuber
discs, stem pith, sections of leaf lamina), but regeneration could be
obtained from all of them by the same two-step procedure (Ochatt and
Caso, 1986).
A two-stage protocol also led to successful shoot formation in leaflet
discs of six cultivars (Webb et aI., 1983). Both auxin and cytokinin were
Figure 3.46 Plants of cv. Superior grown on in the greenhouse after regenerating
from cultured tuber discs. (Fig. 7 from Jarret et at., 1980b.)
required for the formation of callus; cytokinin was required for shoot
formation but auxin was inhibitory. Callus was first visible after 10-14 days
culture, but cell division began in the palisade and spongy mesophyll after
only 4 days. Shoots were formed in regions of callus at the cut surface of
the explants.
Silva (1985) found that tuber explants could be maintained in a viable,
green but non-regenerating state for up to 18 months on White's medium.
After transfer to modified Murashige and Skoog's medium green globular
structures developed peripherally, and eventually gave rise to plantlets.
Silva (1985) suggested that this technique could be used for germplasm
storage, instead of low temperatures.
The formation of minitubers (see Section 3.6.4(b) and Fig. 3.37) can also
increase the time between sub-cultures (Wooster and Dixon, 1986). Such
mini- or microtubers (about one cm or less in diameter) can be produced
from every axil of cultured shoots. Their formation is promoted by SD, low
night temperatures, high light intensity, the presence of ABA and CCC,
and some other factors (Wang and Hu, 1985). Microtubers will sprout and
give rise to new plants, and thus provide an easy means of transporting
germplasm between countries (Schilde-Rentschler and Roca, 1987).
Tissue culture 141
3.10.2 Anther culture
Considerable work has been devoted in recent years to the experimental
induction of embryo-like structures, often called embryoids, from develop-
ing pollen grains in cultured anthers. These embryoids can develop into
plantlets and eventually mature plants. The first successful reports related
to pollen of Datura (Guha and Maheshwari, 1964, 1966) and Nicotiana
(Nitsch and Nitsch, 1969), both members of the Solanaceae. It might there-
fore be expected that pollen of the cultivated potato would be equally amen-
able to this treatment, but unfortunately this has not proved to be the case,
although greater success has been attained recently. Since haploid embryoids
capable of developing into mature plants can be obtained by this technique, it
is potentially an important one for the plant breeder. To obtain homozy-
gous pure lines by conventional breeding methods takes 6-8 years, whereas
it may be done in 6 months by anther culture (Sopory and Bajaj, 1987).
Dunwell and Sunderland (1973) successfully induced embryoid forma-
tion in pollen of S. tuberosum, cvs Pentland Crown, Record and Maris
Piper, by culturing anthers at or just prior to the stage of the first mitosis in
the pollen grains. The success rate was not high, and in most of these
experiments embryoids did not progress beyond the multicellular stage
(Fig. 3.47). In potato, an average of less than five pollen embryoids were
produced per anther, whereas in tobacco they may be numbered in
thousands. Dunwell and Sunderland (1973) found that the stage of
development was far more critical than the composition of the medium.
Figure 3.47 Pollen embryoids of (a) cv. Maris Piper; (b) cv. Pentland Crown.
Considerable cell division has taken place. Degenera>ting pollen grains can also be
seen. (Fig. 1 from Dunwell and Sunderland, 1973.)
In other experiments with S. tuberosum, cv. Hansa, several embryoids
were obtained from callus originating from one anther (Kohl en bach and
Geier, 1972). However, the callus had proliferated from connective tissue.
Using a dihaploid clone of S. tuberosum, Sopory (1979) found that
optimal results required 6% sucrose, IAA, BA and addition of activated
charcoal to the medium. Wenzel et al. (1979) obtained a regeneration
frequency of 60 embryoids per 100 anthers; 10% of the embryoids grew to
plants. Nearly all the plants regenerated from anthers of dihaploid
potatoes were themselves dihaploid in the mature stage. Monohaploid
plants have been obtained by anther culture from embryo ids (Johansson,
1986).
Wang and Hu (1985) have reviewed the hormone combinations used by
different workers. Johansson (1988) devised an ingenious method of
culturing anthers on cubes of solid medium surrounded by liquid medium;
in this way, the surrounding medium could easily be changed. He found
that the best yield of embryo ids was obtained after culture without
hormones for one week, with the addition of 2, 4- D thereafter. Regeneration
from callus was best on a double-layer medium, with liquid medium on top
of solid medium. In general, successful anther culture depends on the
genotype of the parent plant, the stage of development of the pollen at the
time of culture, and the composition of the medium (Sopory and Bajaj,
1987).
3.10.3 Protopiasts
The technique of isolation (usually by enzymatic means) and culture of
isolated protoplasts has been applied to potatoes with some success. The
topic has been reviewed by Shepard et al. (1980) and Wang and Hu (1985).
Successful regeneration from protoplasts affords a method of rapid
multiplication. Considerable differences in protoplast production, callus
growth, and shoot formation were found in different cultivars (Radke and
Grun, 1986).
Usually the protoplasts regenerate a cell wall, then divide to form a small
callus. This may occur within two weeks (Shepard et al., 1980). In general,
division will only occur if large numbers (e.g. 5 x 103 to 10 X 103 )
of protoplasts per ml of medium are present (Shepard et al., 1980). Usually
the callus is then transferred to solid medium, although it may be
important that this is soft (Bokelmann and Roest, 1983). Both the
protoplasts and tissue regenerating shoots of different cultivars thrive
best on specific media adjusted to their needs (Foulger and Jones,
1986). Regeneration of shoots from the callus may take about 2 weeks
(Bokelmann and Roest, 1983) or longer. In cvs King Edward and Desiree
70% of the calli regenerated shoots (Foulger and Jones, 1986). The process
from protoplast isolation to the formation of plantlets may take 3 to 6
months (Bokelmann and Roest, 1983; Vries and Bokelmann, 1986).
Tissue culture 143
It is now known that the conditions under which the source material for
the protoplasts is grown is important in achieving division of the latter
when cultured. This involves temperature, light intensity, photoperiod and
nutrition (Shepard et aI., 1980; Foulger and Jones, 1986).
One of the aims of protoplast culture is somatic fusion. Some success has
led to the formation of somatic hybrids between the potato and other
species of Solanum (Wang and Hu, 1985). Enhanced vigour may result
(Baer et al., 1984).
3.10.4 Somaclonal variation

Although it was originally believed that vegetative reproduction such as is
induced in tissue culture led to a clone of genetically identical individuals,
it is now known that considerable heterogeneity occurs in the products of
tissue culture. The term 'somaclonal variation' denotes the variation found
in plants derived from cell culture (Larkin and Scowcroft, 1981). This
variation, which can be utilized to select for particular characters, is
greatest in plants derived from callus (and thus from protoplasts). Potato
appears to be quite prone to somaclonal variation (Bajaj, 1987).
For example, polymorphic shoots were obtained from subcultured callus
derived from portions of potato sprouts (Quraishi, 1985). Plants regene-
rated from pieces of rachis of cv. Desiree gave rise to red, white and red-
splashed tubers; it was thought that the latter might be stable periclinal
chimeras (Wheeler et al., 1985). Much phenotypic variability has been
observed in the progeny of protoplasts (Wenzel et aI., 1979; Bokelmann
and Roest, 1983; Sree Ramulu et al., 1984). Plants regenerated from a
single callus, and thus a single protoplast, showed this variation (Thomas et
al., 1982).
Shepard et al. (1980) examined more than 10 000 clones derived from
protoplasts (sometimes called protoclones) of cv. Russet Burbank. Under
field conditions these plants showed variation in morphology, earliness of
tuber set, and skin characters. Tubers with smooth white skin were
produced by some clones. In 65 plants derived from protoclones, a number
of characters were compared (Secor and Shepard, 1981). In 22 characters
the plants showed statistical differences from cv. Russet Burbank, while in
nine they did not.
Thus a somatic cell population may represent a source of potentially
useful genetic variation (Shepard et aI., 1980); it is thought that the
variations may pre-exist in the somatic cells of the explant (Larkin and
Scowcroft, 1981). Unfortunately, it is not yet possible to manipulate this
reservoir of variation at will.
3.11 GENETIC ENGINEERING
By using the techniques of genetic manipulation, however, it is beginning

to become possible to introduce desirable characters at will. In the potato,
the bacterium Agrobacterium rhizogenes has been used as the agent for
introducing specific genes. Transgenic plants, i.e. those which have been
genetically modified, can be obtained within 1 or 2 months (Twell, 1988):
changes in phenotype were maintained in the next generation of plants,
derived from tubers (Ooms et aI., 1986). The possibilities of these
techniques applied to the potato have been reviewed by Bajaj (1987).
Plants regenerated from the roots of plants of cv. Desiree transformed
with A. rhizogenes were uniform and differed in morphology from
untransformed plants. They tuberized quickly and produced longer tubers
with more prominent eyes (Ooms et aI., 1985). It might therefore be
possible to use transferred DNA (T-DNA) involved in the control of
tuberization or other morphological features.
A line of cv. Maris Bard transformed by A. tumefaciens produced much
branched stunted shoots with a basal callus and no roots, when grown in
culture (Fig. 3.48). When grafted on to stocks of normal Maris Bard in
culture, a more normal morphology was induced. Stolons and aerial
mini tubers developed on the transformed scions. Much higher concentra-
tions of cytokinins were present in the transformed shoots (100- to 200-fold
higher than normal); this was somewhat reduced in grafted shoots. There
was a three-fold reduction of IAA in transformed shoots (Ooms and
Lenton, 1985). The changes induced by the T-DNA are highly conserved
both in culture and in the field (Ooms, 1987).
Figure 3.48 Shoot cultures of cv. Maris Bard (left) and line Mb1501B, trans-
formed by Agrobacterium tumefaciens (right). The transformed line has stunted,
much branched shoots and no roots. (Fig. 1 from Ooms and Lenton, 1985.
Reprinted by permission of Kluwer Academic Publishers.)
Conclusions 145
These techniques, when perfected, offer the possibility of studying the
effects of specific genes which may modify the endogenous hormones, and
hence morphological characters such as tuber formation and development.
3.12 CONCLUSIONS
It is clear that the control of growth and development in the potato plant
depends on correlative effects between one part of the plant and another.
As yet, too little is known about the structural relationships of these
various parts, and the possible channels of transport between them.
In the foregoing account of the structure and development of the potato
plant, particular attention has been directed to the stolon and tuber, having
regard to the commercial importance of the latter. Much work has been
devoted to the factors which control stolon and tuber formation and
development. Although some of this work seems far removed from actual
agricultural application, clearly techniques must be devised for enhancing
tuber formation, growth and development which can actually be applied in
the field. One promising current line of investigation seems to be the
application of various hormonal substances, together with a study of
extracted endogenous hormones. It seems clear that gibberellins are
closely involved in the control of tuber formation. Paradoxically,
gibberellins stimulate the development of stolons - the usual site of
subsequent tuber formation - but inhibit tuber formation itself. Tuber
formation seems to be associated with a low level of endogenous
gibberellins, but the results of certain grafting experiments and some
experiments with cuttings make it difficult to accept that this condition
alone is sufficient to lead to tuberization. More probably the balance
between gibberellins and some other substance(s) is important.
For example, Hammes and Nel (1975) have suggested that a balance
between endogenous gibberellins and inhibitors may determine the active
gibberellin level in the plant, and hence affect tuberization. As already
pointed out, both cytokinins and inhibitors of extension growth such as
ethylene and abscisic acid are probably also implicated. Indeed, it has been
suggested in this chapter that in normal tuber development a sequence of
growth regulators may be involved. Various experimental treatments may
mimic part of this co-ordinated sequence of events.
The possibility of introducing foreign T-DNA which can affect or control
levels of endogenous hormones offers a powerful tool for investigating the
factors involved in tuberization.
In the normal development the formation of a tuber just behind the
stolon tip ensures that it is subtended by a stem that possesses a consider-
able complement of phloem, which plays a fundamental role in the ensuing
translocation of food materials into the tuber. Although tubers can be
induced to develop in other locations, such as from axillary buds on the
leafy shoot, it may be questioned whether the development of such tubers
would ultimately be limited by lack of phloem. Thus it may be quite
important to assess the anatomical aspects of responses to hormone
treatments; there is certainly much scope for more work in this field.
ACKNOWLEDGEMENTS
I am greatly indebted to Dr Subha Basu for making some of the

preparations on which some of the original illustrations are based, to
Mr G. Grange and Mr 1. Miller for taking or printing many of the
photographs, and to a number of people for supplying copies of previously
published photographs. The source of these illustrations is acknowledged
in the captions. I am also grateful to Miss Mary Gregory, of the 10drell
Laboratory, Kew, for a number of references to anatomical literature, to
Dr S. Carnegie, Department of Agriculture and Fisheries for Scotland, for
supplying some material with developing tubers, and to Mr N. Claughan
and Mr D. Newton for growing young potato plants.
REFERENCES
Abdala de Bottini, G. and Tizio, R. (1981) Hormonal contribution of the mother

tuber to growth, stolonization and tuberization of the potato plant (Solanum
tuberosum L. cv. Spunta). Phyton (Buenos Aires), 41, 27-32.
Abdala de Bottini, G., Goleniowoski, M. and Tizio, R.M. (1981) Effect of
(2-chloroethyl) trimethylammonium chloride upon gibberellin levels in potato
plants (Solanum tuberosum L.) and influence of these phytohormones on
tuberisation in vitro. Z. Pjlanzenphysiol., 103, 149-55.
Adams, M.J. (1975) Potato tuber lenticels: development and structure. Ann. Appl.
BioI., 79, 265-73.
Ahmad, K.J. (1964) Epidermal studies in Solanum. Lloydia, 27, 243-50.
Ahmed, Ch.M.S. and Sagar, G.R. (1981) Effects of a mixture of NAA + BA on
numbers and growth rates of tubers of Solanum tuberosum L. Potato Res., 24, 267-78.
Anwar, R. (1984) Potato shoot-tip position and its regeneration potential. Pakistan
1. Agric. Res., 5, 107-9.
Appiano, A. and Pennazio, S. (1972) Electron microscopy of potato meristem tips
infected with Potato Virus X. 1. Gen. Virol., 14, 273-6.
Artschwager, E. (1918) Anatomy of the potato plant, with special reference to the
ontogeny of the vascular system. 1. Agric. Res., 14, 221-52.
Artschwager, E. (1924) Studies on the potato tuber. 1. Agric. Res., 27, 809-35.
Artschwager, E. (1927) Wound periderm formation in the potato as affected by
temperature and humidity. 1. Agric. Res., 35, 995-1000.
Asseyeva, T. (1927) Bud variations in the potato and their chimerical nature. 1.
Genet., 19, 1-26.
Asseyeva, T. (1933) Bud mutations in the potato. Trudy Prikl. Bot. Genet. Selek.,
27, 135-218.
References 147
Badenhuizen, N.P. and Dutton, R.W. (1956) Growth of 14C-labelled starch
granules in potato tubers as revealed by autoradiographs. Protoplasma, 47,
156-63.
Baer, M.A., Austin, S. and Helgeson, J.P. (1984) Somatic fusions of protoplasts of
Solanum species. Plant Physiol., 75 (supp!. 1), 133.
Bajaj, Y.P.S. (1987) Biotechnology and 21st century potato, in Biotechnology in
Agriculture and Forestry, 3. Potato (ed. Y.P.S. Bajaj), Springer-Verlag, Berlin,
pp.3-22.
Bancher, E. and Halzl, J. (1959) Uber die Driisenhaare von Solanum tuberosum,
Sorte 'Sieglinde'. Protoplasma, 50, 356-69.
Banks, W., Greenwood, C.T. and Muir, D.D. (1973) Studies on the biosynthesis of
starch granules. 7. Compound granules in potato starch. Die Starke, 25, 331-5.
Barker, W.G. (1953) A method for the in vitro culturing of potato tubers. Science,
NY, 118, 384---5.
Barrios, E.P., Newsom, D.W. and Miller, J.C. (1961) Some anatomical characters
associated with the culinary quality of Irish potatoes. Proc. Amer. Soc. Hart.
Sci., 78, 413-20.
Bhadurim, P.N. (1936) Studies on the embryogeny of the Solanace.ae. I. Bot. Gaz.,
98,283-95.
Bokelmann, G.S. and Roest, S. (1983) Plant regeneration from protoplasts of
potato (Solanum tuberosum cv. Bintje). Z. Pflphysiol., 109,259-65.
Bonnemain, J.-L. (1970) Histogenese du phloeme interne et du phloeme indus des
Solanacees. Rev. Gen. Bot., 77, 5-51.
Boo, L. (1962) Gibberellin-like substances in the potato tuber during and after the
rest period. Svensk Bot. Tidskrift., 56, 193-6.
Booth, A. (1959) Some factors concerned in the growth of stolons in the potato. J.
Linn. Soc. (Bot.)., 56, 166--9.
Booth, A. (1963) The role of growth substances in the development of stolons, in
The Growth of the Potato (eds J.D. Ivins and F.L. Milthorpe), Butterworths,
London, pp. 99-113.
Booth, A. and Wareing, P.F. (1958) Growth-promoting activity of potato tuber
extracts in the coleoptile straight-growth test: quaJiltitative aspects of
indoleacetic acid detection. Nature, Land., 182, 406.
Bruinsma, J. and Swart, J. (1970) Estimation of the course of dormancy of potato
tubers during growth and storage, with the aid of gibberellic acid. Potato Res.,
13,29-40.
Burt, R.L. (1964) Influence of short periods of low temperature on tuber initiation
in the potato. Eur. Potato J., 7, 197-208.
Catchpole, A.H. and Hillman, J. (1969) Effect of ethylene on tuber initiation in
Solanum tuberosum. Nature, Land., 223, 1387.
Chailakhyan, M.Kh., Yanina, L.I., Devedzhyan, A.G. and Lotova, G.N. (1981)
Photoperiodism and tuber formation in grafting of tobacco on to potato. Dokl.
Akad. Nauk SSSR, 257, 1276--80.
Chapman, H.W. (1958) Tuberization in the potaJo plant. Physiol. Plant., 11,
215-24.
Chirilei, H., Chirila, C. and Curticapeanu, G. (1963) Contributiuni la cunoa-
sterea modificarilar morfo-anatomice produse la plantele de soia si de cartof
sub influenta aciduli giberelic. Lucr. Grad. Bot. Bucuresti, 1961-2(2),
651-65.
Clark, C.F. (1921) Development of tubers in the potato. USDA Bulletin No. 958,
27 pp.
Clarke, A.E. (1940) FertiUzation and early embryo development in the potato.
Am. Potato 1.,17,20-5.
Clauhs, R.P. and Grun, P. (1977) Changes in plastid and mitochondrion content
during maturation of generative cells of Solanum (Solanaceae). Am. l. Bot., 64,
377-83.
Claver, F.K. (1967) Formacion de yemas adventicias en tejidos de papa. (Solanum
tuberosum). Rev. Fac. Agron., La Plata, 43, 45-54.
Clowes, F.A.L. and MacDonald, M.M. (1989) Cell cycling and the fate of potato
buds. Ann. Bot., 59,141-8.
Cornforth, 1.W., Milborrow, B.V. and Ryback, G. (1966) Identification and
estimation of (+ )-abscisin II (,DormiIl') in plant extracts by spectropolarimetry.
Nature, Lond., 210, 627-8.
Cottle, W. and Kolattukudy, P .E. (1982) Abscisic acid stimulation of suberization.
Induction of enzymes and deposition of polymeric components and associated
waxes in tissue cultures of potato tuber. Plant Physiol., 70, 775-80.
Crafts, A.S. (1933) Sieve-tube structure and translocation in the potato. Plant
Physiol., 8, 81-104.
Crafts, A.S. and Crisp, C.E. (1971) Phloem Transport in Plants, Freeman, San
Francisco.
Crane, M.B. (1936) Note on a periclinal chimaera in the potato. l. Genet., 32,
73-7.
Creech, D.L., Workman, M. and Harrison, M.D. (1973) The influence of storage
factors on endogenous ethylene production by potato tubers. Am. Potato l. , 50,
145-50.
Danert, S. (1957) Uber den Sprossaufbau und die Blattentwicklung bei der
Kartoffel. Der Ziichter, 27, 22-33.
Davis, G.L. (1966) Systematic Embryology of the Angiosperms, Wiley, New York.
de Vries, H. (1878) Beitrage zur speciellen Physiologie landwirthschaftlicher
Kulturpflanzen. V. Wachstumgeschichte der Kartoffe1pflanze. Landw. lb., 7,
591-682.
de Vries, S.E. and Bokelmann, G.S. (1986) Regeneration of callus and plants from
cell suspension protoplasts of dihaploid potato (Solanum tuberosum L.). l.
Plant Physiol., 122, 199-203.
Dixon, H.H. (1922a) Transport of organic substances in plants. Nature, Lond.,
110, 547-51.
Dixon, H.H. (1922b) Transport of organic substances in plants. Rept. Brit. Ass.
Adv. Sci., Hull, 1922, pp. 192-203.
Duncan, D.A. and Ewing, E.E. (1984) Initial anatomical changes associated with
tuber formation on single-node potato (Solanum tuberosum L.) cuttings. Ann.
Bot., 53, 607-10.
Dunwell, 1.M. and Sunderland, N. (1973) Anther culture of Solanum tuberosum L.
Edelman, J., lefford, T.G. and Singh, S.P. (1969) Studies on the biochemical basis
of physiological processes in the potato tuber. The pathway and control of
translocation from the tuber. Planta, 84, 48-56.
El-Antably, H.M.M., Wareing, P.F. and Hillman, 1. (1967) Some physiological
responses to d,l abscisin (dormin). Planta, 73, 74-90.
References 149
Emilsson, B. (1949) Studies on the rest period and dormant period in the potato
tuber. Acta Agric. Suecana, 3, 189-282.
Engels, Ch. and Marschner, H. (1986) Allocation of photosynthate to individual
tubers of Solanum tuberosum L. III. Relationship between growth rate of
individual tubers, tuber weight and stolon growth prior to tuber initiation. l.
Exp. Bot., 37,1813-22.
Ernst, J.M., Perry, B.A. and Blackhurst, H.T. (1968) Developmental anatomy of
the Irish potato (Solanum tuberosum) during root and stolon differentiation. l.
Rio Grande Val. Hart. Soc., 22, 53-7.
Esau, K. (1938) Ontogeny and structure of the phloem of tobacco. Hilgardia, 11,
343-424.
Esmarch-Bromberg, F. (1919) Beitrage zur Anatomie der gesunden und kranken
Kartoffelpflanze. I. Anatomie der vegetativen Organen. Landw. lb., 54,
161-266.
Espelie, K.E., Sadek, N.Z. and Kolattukudy, P.E. (1980) Composition of suberin-
associated waxes from the subterranean storage organs of seven plants. Parsnip,
carrot, rutabaga, turnip, red beet, sweet potato and potato. Planta, 148, 468-76.
Ewing, E.E. (1978) Critical photoperiods for tuberization: a screening technique
with potato cuttings. Am. Potato l., 55, 43-53.
Ewing, E.E. (1981) Heat stress and the tuberization stimulus. Am. Potato l., 58,
31-49.
Ewing, E.E. (1985) Cuttings as simplified models of the potato plant, in Potato
Physiology (ed. P.H. Li), Academic Press, London, pp. 153-207.
Ewing, E.E. (1987) The role of hormones in potato (Solanum tuberosum L.)
tuberization, in Plant Hormones and Their Role in Plant Growth and
Development (ed. P.J. Davies), Martinus Nijhoff, Dordrecht, pp. 515-38.
Ewing, E. and Wareing, P.F. (1978) Shoot, stolon, and tuber formation on potato
(Solanum tuberosum L.) cuttings in response to photoperiod. Plant Physiol. , 61,
348-53.
Fischnich, O. and Heilinger, F. (1959) Die Kartoffel, Bildung, Erhaltung,
Verwertung ihrer Inhaltsstoffe. Angew. Bot., 33, 49-70.
Forsline, P.L. and Langille, A.R. (1975) Endogenous cytokinins in Solanum
tuberosum as influenced by photoperiod and temperature. Physiol. Plant., 34,
75-7.
Forsline, P.L. and Langille, A.R. (1976) An assessment of the modifying effect of
kinetin on in vitro tuberization of induced and non-induced tissues of Solanum
tuberosum. Can. l. Bot., 54, 2513-16.
Foulger, D. and Jones, M.G.K. (1986) Improved efficiency of genotype-dependent
regeneration from protoplasts of important potato cultivars. Plant Cell Reports,
5,72-6.
Frandsen, N.O. (1968) Die Plastidenzahl als Merkmal bei der Kartoffel. Theor.
Appl. Gen., 38, 153-67.
Frey-Wyssling, A. (1974) Uber die Ultrastruktur der Kartoffelstarkek6rner.
Schweizerische landwirtschaftliche Forschung, 13, 385-93.
Fukuda, Y. (1967) Anatomical study of the internal phloem in the stems of
dicotyledons, with special reference to its histogenesis. l. Fac. Sci. Univ. Tokyo,
Sect. III, Bot. 9, 313-75.
Gallant, D., Degrois, M., Sterling, C. and Guilbot, A. (1972) Microscopic effects
of ultrasound on the structure of potato starch. Die Starke, 24, 116-23.
Garcia-Torres, L. and Gomez-Campo, C. (1972) Increased tuberization in potatoes
by ethrel (2-chloro-ethyl-phosphonic acid). Potato Res., 15, 76-80.
Garcia-Torres, L. and Gomez-Campo, C. (1973) In vitro tuberization of potato
sprouts as affected by ethrel and gibberellic acid. Potato Res., 16,73-9.
Gibson, RW. (1971) Glandular hairs providing resistance to aphids in certain wild
potato species. Ann. Appl. BioI., 68, 113-19.
Gibson, R.W. (1974) Aphid-trapping glandular hairs on hybrids of Solanum
tuberosum and S. berthaultii. Potato Res., 17, 152-4.
Gifford, RM. and Moorby, J. (1967) The effect of CCC on the initiation of potato
tubers. Eur. Potato J., 10, 235-8.
Goodwin, P. (1963) Mechanism and significance of apical dominance in the potato
tuber, in The Growth of the Potato (eds J.D. Ivins and F.L. Milthorpe),
Butterworths, London, pp. 63-71.
Goodwin, P.B. (1966) An improved medium for the rapid growth of isolated
potato buds. J. Exp. Bot., 17,590--5.
Goodwin, P.B. (1967) The control of branch growth on potato tubers. I. Anatomy
of buds in relation to dormancy and correlative inhibition. J. Exp. Bot., 18,
78-86.
Gregorini, G. and Lorenzi, R (1974) Meristem-tip culture of potato plants as a
method of improving productivity. Potato Res., 17,24-33.
Gregory, L.E. (1956) Some factors for tuberization in the potato plant. Am. J.
Bot., 43, 281-8.
Gruber, E., John, K., Alloush, S. and Schurz, J. (1972) Hinweise auf eine Kern-
Mantel-Struktur im Korn der Kartoffelkudlen-Starke. Microscopica Acta, 72,
189-200.
Guha, S. and Maheshwari, S.C. (1964) In vitro production of embryos from anthers
of Datura. Nature, Land., 204, 497.
Guha, S. and Maheshwari, S.C. (1966) Cell division and differentiation of embryos
in the pollen grains of Datura in vitro. Nilture, Land., 212, 97-8.
Hammes, P.S. and Nel, P.C. (1975) Control mechanisms in the tuberization
process. Potato Res., 18, 262-72.
Harmey, M.A., Crowley, M.P. and Clinch, P.E.M. (1966) The effect of growth
regulators on tuberisation of cultured stem pieces of Solanum tuberosum. Eur.
Potato J., 9, 146-51.
Hayashi, F., Blumenthal-Goldschmidt, S. and Rappaport, L. (1962) Acid and
neutral gibberellin-like substances in potato tubers. PI. Physiol., Lancaster, 37,
774-80.
Hayashi, F. and Rappaport, L. (1966) Isolation, crystallization, and partial identi-
fication of potato factor II from potato tubers. Pl. Physiol., Lancaster, 41, 53-8.
Haynes, K.G. and Haynes, F.L. (1988) The effect of day and night temperatures
on bud initiation and flowering in diploid potatoes. Am. Potato J., 65, 589-96.
Hayward, H.E. (1938) The Structure of Economic Plants, Macmillan, New York.
Hayward, P. (1974) Waxy structures in the lenticels of potato tubers and their
possible effects on gas exchange. Planta, 120, 273.
Hemberg, T. (1985) Potato rest, in Potato Physiology (ed. P.H. Li), Academic
Press, New York, pp. 353-88.
Hoff, J.E. (1971) Potato protein crystals. HortScience, 6, 190.
Hollings, M. (1965) Disease control through virus-free stock. Ann. Rev.
Phytopath., 3, 367-96.
References 151
Holzl, J. (1965) Licht- und elektronmikroskopische Untersuchungen an Zwiebel-
und Kartoffelzellen. Protoplasma, 60, 446--79.
Holzl, J. and Bancher, E. (1958) Uber die Eiweisskristalle von Solanum
tuberosum. Osterr. Bot. Ztschr., 105, 385-407.
Hooker, W.J., Yang, T.e. and Potter, H.S. (1973) Air pollution injury of potato in
Michigan. Am. Potato J., 50, 151--61.
Hori, S. (1954) Formation of storage starch in storage organs. 1. The starch
formation in potato tuber. Bot. Mag. (Tokyo), 67, 57--62.
Howard, H.W. (1964) Experimental production of buds on the roots of potatoes.
Nature, Land., 203, 1303-4.
Howard, H.W. (1966) A sectorial entire pinnate leaf chimera in the potato variety
Majestic. New Phytol., 65, 284--7.
Howard, H.W. (1967) Further experiments on the use of X-rays and other methods
in investigating potato chimeras. Radiat. Bot., 7, 389-99.
Howard, H.W. (1969) A full analysis of a potato chimera. Genetica, 40, 233-4l.
Howard, H.W. (1970a) The eye-excision method of investigating potato chimeras.
Potato Res., 13, 220--2.
Howard, H.W. (1970b) Genetics of the Potato Solanum tuberosum, Logos Press,
London.
Howard, H. W. (1971) The stability of Ll-mutant periclinal potato chimeras. Potato
Res., 14, 91-3.
Howard, H. W. (1972) The stability of an L3 mutant potato chimera. Potato Res.,
15, 374--77.
Howard, H.W., Wainwright, J. and Fuller, J.M. (1963) The number of indepen-
dent layers at the stem apex in potatoes. Genetica, 34, 113-20.
Hussey, G. and Stacey, N.J. (1981) In vitro propagation of potato (Solanum
tuberosum L.). Ann. Bot., 48, 787-96.
Hussey, G. and Stacey, N.J. (1984) Factors affecting the formation of in vitro
tubers of potato (Solanum tuberosum L.). Ann. Bot., 53, 565-78.
Iriuda, N., Kikuta, Y. and Okazawa, Y. (1983) Significance of phosphorylase on
starch synthesis of potato tuberization. J. Fac. Agric. Hokkaido Univ., 61,
231-43.
Ivins, J.D. and Bremner, P.M. (1964) Growth, development and yield in the
potato. Outlook on Agriculture, 4, 211-17.
Jager, F.Kh. and Jakovlev, M.S. (1939) Condition of starch in potato at different
stages of tuber ripening. C.R. (Dokl.) Acad. Sci. URSS, 23, 491-4.
Jameson, P.E., McWha, J.A. and Haslemore, R.M. (1985) Changes in cytokinins
during initiation and development of potato tubers. Physiol. Plant., 63, 53-7.
Jarret, R.T., Hasegawa, P.M. and Erickson, H.T. (1980a) Factors affecting shoot
initiation from tuber discs of potato (Solanum tuberosum). Physiol. Plant., 49,
177-84.
Jarret, R.L., Hasegawa, P.M. and Erickson, H.T. (1980b) Effects of medium
components on shoot formation from cultured tuber discs of potato. J. Amer.
Soc. Hart. Sci., 105, 238-42.
Johansen, D.A. (1950) Plant Embryology, Chronica Botanica, Waltham, Mass.
Johansson, L. (1986) Improved methods for induction of embryogenesis in anther
cultures of Solanum tuberosum. Potato Res., 29, 179-90.
Johansson, L.B. (1988) Increased induction of embryogenesis and regeneration in
anther cultures of Solanum tuberosum L. Potato Res., 31, 145-9.
Jones, S.G. (1939) Introduction to Floral Mechanism, Blackie & Son Ltd, London
and Glasgow.
Kahl, G., Rosenstock, G. and Lange, H. (1969) Die Trennung von Zellteilung und
Suberinsynthese in dereprimiertem pflanzlichem Speichergewebe durch Tris-
(hydroxymethyl-) aminomethan. Planta, 87, 365-71.
Kahn, B.A. and Ewing, E.E. (1983) Factors controlling the basipetal pattern of
tuberization in induced potato (Solanum tuberosum L.) cuttings. Ann. Bot., 52,
861-71.
Kahn, B.A., Ewing, E.E. and Senesac, A.H. (1983) Effects of leaf age, leaf area,
and other factors on tuberization of cuttings from induced potato (Solanum
tuberosum) shoots. Can. l. Bot., 61, 3193-201.
Kassanis, B. (1949) Potato tubers freed from leaf-roll virus by heat. Nature, Lond.,
164,881.
Kassanis, B. (1957) The use of tissue cultures to produce virus-free clones from
infected potato varieties. Ann. Appl. Bioi., 45, 422-7.
Kassanis, B. (1967) Plant tissue culture, in Methods in Virology (eds K.
Maramarosch and H. Koprowski), Academic Press, New York and London,
pp. 537--66.
Kassanis, B. and Varma, A. (1967) The production of virus-free clones of some
British potato varieties. Ann. Appl. Bioi., 59, 447-50.
Khan, Shams-ul-Islam (1951) Pollen sterility in Solanum tuberosum L. Cytologia,
16, 124--30.
Kikuta, Y. and Okazawa, Y. (1982) Shoot-bud formation and plantlet regeneration
in potato tuber tissue cultured in vitro. l. Fac. Agric. Hokkaido Univ., 61,
166-79.
Klopfer, K. (1965a) Uber den Nachweis von drei selbstandigen Schichten im
Sprossscheitel der Kartoffel. Z. Pf/Zucht., 53, 67-87.
Klopfer, K. (1965b) Histogenetische Untersuchungen am Sprossscheitel der
Kartoffel. Flora, B, 156, 50-77.
Knowles, N.R. (1986) Differences in nitrogen metabolism during growth of plants
from aged potato (Solanum tuberosum L.) meristems. Ann. Bot., 58, 711-18.
Koda, Y. and Okazawa, Y. (1983a) Influences of environmental, hormonal and
nutritional factors on potato tuberization in vitro. lap. l. Crop. Sci., 52, 582-91.
Koda, Y. and Okazawa, Y. (1983b) Characteristic changes in the levels of
endogenous plant hormones in relation to the onset of potato tuberization. lap.
l. Crop Sci., 52, 592-7.
Koda, Y. and Okazawa, Y. (1988) Detection of potato tuber-inducing activity in
potato leaves and old tubers. Plant Cell Physiol., 29, 969-74.
Koda, Y., Orner, E.-S.A., Yoshihara, T., Shibata, H., Sakamura, S. and
Okazawa, Y. (1988) Isolation of a specific potato tuber-inducing substance from
potato leaves. Plant Cell Physiol., 29, 1047-51.
Kohlenbach, H.W. and Geier, T. (1972) Embryonen aus in vitro kultivierten
Antheren von Datura meteloides Dun., Datura wrightii Regel und Solanum
tuberosum L. Z. PflPhysiol., 67,161-5.
Krauss, A. (1985) Interaction of nitrogen nutrition, phytohormones, and
tuberization, in Potato Physiology (ed. P.H. Li), Academic Press, New York,
pp.209-30.
Krylova, N.V., Stepanenko, V.D. and Reifman, V.G. (1973) Potato virus X in
potato apical meristems. Acta Virol., 17, 172.
References 153
Kumar, D. and Wareing, P.F. (1972) Factors controlling stolon development in the
potato plant. New Phytol., 71, 639--48.
Kumar, D. and Wareing, P.F. (1973) Studies on tuberization in Solanum andigena.
I. Evidence for the existence and movement of a specific tuberization stimulus.
New Phytol., 72, 283-7.
Kumar, D. and Wareing, P.F. (1974) Studies on tuberization of Solanum andigena.,
II. Growth hormones and tuberization. New Phytol., 73, 833--40.
Lamm, R. (1937) A contribution to the embryology of the potato. Svensk Bot.
Tid., 31, 217-20.
Lamotte, C.E. (1983) Exogenous abcisic acid causes reorientation of branch shoots
of Solanum andigena. Test of the Snow hypothesis of apical dominance. Am. 1.
Bot., 70, 88.
Lange, H., Rosenstock, G. and Kahl, G. (1970) Induktionsbedingungen
der Suberinsynthese und Zellproliferation bei Parenchymfragmenten dec
Kartoffelknolle. Planta, 90, 109-18.
Larkin, P.J. and Scowcroft, W.R. (1981) Somaclonal variation - a novel sou!rce of
variability from cell cultures for plant improvement. Theor. Appl. Gen., 60,
197-214.
Lehmann, R. (1926) Untersuchungen iiber die Anatomie der Kartoffelknolle,
unter besonderer Beriicksichtigung des Dickenwachstums und der Zellgrosse.
Planta, 2, 87-131.
Leonard, R. and Sterling, C. (1972) Free.7!e-etched surfaces in potato starch. 1.
Ultrastruct. Res., 39, 85-95.
Leshem, B. and Clowes, F.A.L. (1972) Rates of mitosis in shoot apices of potatoes
at the beginning and end of dormancy. Ann. Bot., 36, 687-91.
Lipetz, J. (1970) Wound-healing in higher plants. Int. Rev. Cytol., 27,1-28.
Lovell, P.H. and Booth, A. (1967) Effects of gibberellic acid on growth, tuber
formation and carbohydrate distribution in Solanum tuberosum. New Phytol.,
66,525-37.
Lovell, P.H. and Booth, A. (1969) Stolon initiation and development in Solanum
tuberosum L. New Phytol., 68, 1175-85.
Macdonald, D.M. (1973) Heat treatment and meristem culture as a means of
freeing potato varieties from viruses X and S. Potato Res., 16, 263-9.
Madec, P. (1963) Tuber-forming substances in the potato, in The Growth of the
Potato (eds J.D. Ivins and F.L. Milthorpe), Butterworths, London, pp. 121-30.
Madec, P. and Perennec, P. (1959) Le role respectif du feuilIage et du tubercule-
mere dans la tuberisation de la pomme de terre. Eur. Potato 1., 2, 22--49.
Madec, P. and Perennec, P. (1969) Levee de dormance de tubercules de pomme de
terre d'age different: action de la rindite, de la gibbe.relline et de l'oeilletonnage.
Eur. Potato 1.,12,96-115.
Maheshwari, P. (1950) An Introduction to the Embryology of Angiosperms,
McGraw-Hill, New York.
Marinos, N.G. (1965) Comments on the nature of a crystal-containing body in
plant cells. Protoplasma, 60, 31-3.
Marinos, N.G. (1967) Multifunctional plastids in the meristematic region of potato
tuber buds. 1. Ultrastruct. Res., 17, 91-113.
Martin, c., Vernay, R. and Paynot, N. (1982) Physiologie vegetale. Photoperiodisme,
tuberisation, floraison and phenolamides. c.R. Hebd. Seances Acad. Sci.,
Paris, 295, 565-8.
Mauk, e.S. and Langille, A.R. (1978) Physiology of tuberization in Solanum
tuberosum L. Cis-zeatin riboside in the potato plant: its identification and
changes in endogenous levels as influenced by temperature and photoperiod.
Pl. Physiol., Lancaster, 62, 438--42.
McGrady, 1.1., Struik, P.C. and Ewing, E.E. (1986) Effects of exogenous
applications of cytokinin on the development of potato (Solanum tuberosum L.)
cuttings. Potato Res., 29, 191-205.
Meinl, G. (1966) Untersuchungen zur sorten und unweltbedingten Variation der
Lentizellenanzahl von Kartoffelknollen. Flora, B, 156,419-26.
Melis, R.l.M. and van Staden, 1. (1984) Tuberization and hormones. Z.
PfiPhysiol., 113,271-83.
Mellor, F.e. and Stace-Smith, R. (1969) Development of excised potato buds in
nutrient culture. Can. 1. Bot., 47, 1617-21.
Mellor, F.C. and Stace-Smith, R. (1987) Virus-free potatoes through meristem
culture, in Biotechnology in Agriculture and Forestry, 3. Potato (ed. Y.P.S.
Bajaj), Springer-Verlag, Berlin, pp. 30--9.
Menzel, e.M. (1980) Tuberization in potato at high temperatures: responses to
gibberellin and growth inhibitors. Ann. Bot., 46, 259-65.
Menzel, C.M. (1985) Tuberization in potato at high temperatures: interaction
between temperature and irradiance. Ann. Bot., 55, 35-9.
Mes, M.G. and Menge, I. (1954) Potato shoot and tuber cultures in vitro. Physiol.
Plant., 7, 637--49.
Miedema, P. (1967) Induction of adventitious buds on roots of the potato.
Miedema, P. (1973) A physiological study of adventitious bud formation in potato.
Thesis, Groningen. Centre for Agricultural Publishing and Documentation,
Wageningen. Also Agricultural Research Reports, 787.
Minato, T., Kikuta, Y. and Okazawa, Y. (1979) Effect of ethylene on sprout
growth and endogenous growth substances of potato plants. 1. Fac. Agric.
Hokkaido Univ., 59, 239--48.
Moorby, 1. and Milthorpe, F.L. (1975) Potato, in Crop Physiology. Some case
histories (ed. L.T. Evans), Cambridge University Press, pp. 225-57.
Morel, G. (1964) Regeneration des varietes virosees par la culture des meristemes
apicaux. Rev. Hort. (Paris), 261, 733--40.
Morel, G. and Martin, e. (1955) Guerison de pommes de terre atteintes de
maladies a virus. CR. Seances Acad. Agr. Fr., 41, 472-5.
Morel, G. and Muller, 1.-F. (1964) La culture in vitro du meristeme apical de la
pomme de terre. CR. Hebd. Seances Acad. Sci. Paris, 258, 5250--2.
Morel, G., Martin, C. and Muller, 1.-F. (1968) La guerison des pommes de terre
atteintes de maladies a virus. Ann. Physiol. Veg., 10, 113-39.
Morris, D.A. (1966) Intersprout competition in the potato. I. Effects of tuber size,
sprout number and temperature on sprout growth during storage. Eur. Potato
1., 9, 69-85.
Murashige, T. and Skoog, F. (1962) A revised medium for rapid growth and
bioassays with tobacco tissue cultures. Physiol. Plant., 15, 473-97.
Nielsen, N.K. (1968) An investigation of the regenerative power of periderm in
potato tubers after wounding. Acta Agric. Scand., 18, 113-20.
Nitsch, 1.P. and Nitsch, e. (1969) Haploid plants from pollen grains. Science, NY,
163, 85-7.
References 155
Novak, F.J., Zadina, J., Horackova, V. and Maskova, I. (1980) The effect of
growth regulators on meristem tip development and in vitro muUiplication of
Solanum tuberosum L. plants. Potato Res., 23, 155-66.
Obata-Sasamoto, H. and Suzuki, H. (1979a) Activities of enzymes relating to
starch synthesis and endogenous levels of growth regulators in potato stolon tips
during tuberization. Physiol. Plant., 45, 320-4.
Obata-Sasamoto, H. and Suzuki, H. (1979b) Activities of enzymes relating to
starch synthesis and endogenous levels of growth regulators during tuberization
of isolated potato stolons cultured in vitro. Z. PflPhysiol., 95, 69-75.
Ochatt, S.J. and Caso, O.H. (1986) Differential requirements among tissue source
in Solanum tuberosum L. spp. andigena callus cultures. Turrialba, 36, 363-8.
Ohad, I., Friedberg, I., Ne'eman, Z. and Schramm, M. (1971) Biogenesis and
degradation of starch. 1. The fate of the amyloplast membranes during
maturation and storage of potato tubers. Plant Physiol., 47, 465-77.
Okazawa, Y. and Chapman, H.W. (1962) Regulation of tuber formation in the
potato plant. Physiol. Plant., 15, 413-19.
Ooms, G. (1987) Genetic manipulation in potato using Agrobacterium, in The
Production of New Potato Varieties: Technological Advances (eds G. JeUis and
D.E. Richardson), Cambridge University Press, Cambridge, pp. 293-308.
Ooms, G. and Lenton, J.R. (1985) T-DNA genes to study plant development:
precocious tuberisation and enhanced cytokinins in A. tumefaciens transformed
potato. Plant Molec. BioI., 5, 205-12.
Ooms, G., Karp, A., Burrell, M.M., Twell, D. and Roberts, J. (1985) Genetic
modification of potato development using Ri T-DNA. Theor. Appl. Gen., 70,
440-6.
Ooms, G., Bossen, M.E., Burrell, M.M. and Karp, A. (1986) Genetic manipula-
tion in potato with Agrobacterium rhizogenes. Potato Res., 29, 367-79.
Palmer, C.E. and Barker, W.G. (1973) Influence of ethylene and kinetin on
tuberization and enzyme activity in Solanum tuberosum L. stolons cultured in
vitro. Ann. Bot., 37, 85-93.
Palmer, C.E. and Smith, O.E. (1969a) Cytokinins and tuber initiation in the potato
Solanum tuberosum L. Nature, Lond., 221, 279-80.
Palmer, C.E. and Smith, O.E. (1969b) Effect of abscisic acid on elongation and
kinetin-induced tuberization of isolated stolons of Solanum tuberosum L. Plant
Cell Physiol., 10, 657-64.
Palmer, C.E. and Smith, O.E. (1970) Effect of kinetin on tuber formation on
isolated stolons of Solanum tuberosum L. cultured in vitro. Plant Cell Physiol.,
11, 303-14.
PaIta, J.P. and Li, P.H. (1979) Frost-hardiness in relation to leaf anatomy aoo
natural distribution of several Solanum species. Crop Sci., 19, 665-71.
Pennazio, S. (1971) Terapia di virosi della patata (Solanum tuberosum L.): cultura
di apici meristematici abbinata a termoterapia. Riv. Ortoflorofrut. Ital., 5, 446-52.
Pennazio, S. and Redolfi, P. (1974) Potato virus X eradication in cultured potato
meristem tips. Potato Res., 17, 333-5.
Pereira, M.F.A. and Valio, I.F.M. (1984) Gibberellic acid and the inhibition of
aerial tuberisation in Solanum tuberosum L. Plant Growth Reg., 2, 41-7.
Perombelon, M.C.M. and Lowe, R. (1975) Studies on the initiation of bacterial
soft rot in potato tubers. Potato Res., 18, 64-82.
Peterson, C.A., Peterson, R.L. and Barker, W.G. (1981) Observations on the
structure and osmotic potentials of parenchyma associated with the internal
phloem of potato tubers. Am. Potato J., 58, 575-84.
Peterson, R.L. and Barker, W.G. (1979) Early tuber development from explanted
stolon nodes of Solanum tuberosum var. Kennebec. Bot. Gaz., 140, 398-406.
Peterson, R.L., Barker, W.G. and Howarth, M.J. (1985) Development and
structure of tubers, in Potato Physiology (ed. P.H. Li), Academic Press, New
York, pp. 123-52.
Plaisted, P.H. (1957) Growth of the potato tuber. Plant Physiol., 32, 445-53.
Pogorelova, O.V. (1969) Differentiation of tissues in the Solanum tuberosum L.
shoot. Farmacija, SSSR, 18, 52--6 (in Russian with English summary).
PospfSilova, J., Solarova, J., Catsky, J. and Ondl'ej, M. (1988) The photosynthetic
characteristics during the micropropagation of tobacco and potato plants.
Photosynthetica, 22, 205-13.
Potato Marketing Board (1965) British Atlas of Potato Varieties.
Priestley, J.H. and Woffenden, L.M. (1922) Physiological studies in plant
anatomy. V. Causal factors in cork formation. New Phytol., 21, 252--68.
Priestley, J.H. and Woffenden, L.M. (1923) The healing of wounds in potato
tubers and their propagation by cut sets. Ann. Appl. Bioi., 10, 96-115.
Pullo, E. and Slusarkiewicz, A. (1975) Development of ovules on the stamens in
flowers of Solanum tuberosum variety F1isak. Acta Soc. Bot. Pol., 44, 519-27,
Quraishi, A. (1985) Tissue culture studies as a means for exploring variability in
potato. Pakistan J. Agric. Res., 6, 20-2.
Radke, S. and Grun, P. (1986) Isolation, culture and regeneration of leaf
mesophyll protoplasts of selected clones of Solanum. Potato Res., 29, 451--62.
Railton, I.D. and Wareing, P.F. (1973a) Effects of daylength on endogenous
gibberellins in leaves of Solanum andigena. I. Changes in levels of free acidic
gibberellin-like substances. Physiol. Plant., 28, 88-94.
Railton, I.D. and Wareing, P.F. (1973b) Effects of daylength on endogenous
gibberellins in Solanum andigena. III. Gibberellin production by the leaves.
Physiol. Plant, 29, 430-3.
Rainbow, A. and White, D.J.B. (1972) Preliminary observations on the ultra-
structure of maturing cork-cells from tubers of Solanum tuberosum L. New
Phytol., 71, 899-902.
Rappaport, L. (1972) Mechanism of dormancy in storage organs. Proc. 18th
Internat. Hart. Congr., 5, 143--55.
Rappaport, L. and Sachs, M. (1967) Wound-induced gibberellins. Nature, Land.,
214, 1149-50.
Rappaport, L. and Wolf, N. (1968a) Regulation of bud rest irn tubers of potato,
Solanum tuberosum L. III. Nucleic acid synthesis induced by bud excision and
ethylene chlorohydrin. Proceedings of the International Symposium on Plant
Growth Substances (ed. S.M. Sircar), Calcutta, pp. 79-88.
Rappaport, L. and Wolf, N. (1968b) Regulation of bud rest in tubers of potato,
Solanum tuberosum L. IV. Gibberellins and nucleic acid synthesis in excised
buds, in Biochemical Regulation in Diseased Plants or Injury (eds H. Tokazo, J.
Hidaka and I. Uritani), Phyto-pathological Society of Japan, Tokyo, 1968, pp.
203-11.
Rappaport, L. and Wolf, N. (1969) The problem of dormancy in potato tubers and
related structures. Symp. Soc. Exp. BioI., 23, 219-40.
Reed, T. (1910) On the anatomy of some tubers. Ann. Bot., 24, 537-48.
References 157
Rees-Leonard, D.L. (1935) Macrosporogenesis and development of the macro-
gametophyte of Solanum tuberosum. Bot. Gaz., 96, 734-50.
Reeve, R.M.(1967) Suggested improvements for microscopic measurement of cells
and starch granules in fresh potatoes. Am. Potato J., 44, 41-50.
Reeve, R.M., Hautala, E. and Weaver, M.L. (1969) Anatomy and compositional
variation within potatoes. I. Developmental histology of the tuber. Am. Potato
J., 46, 361-73.
variations within potatoes. III Gross compositional gradients. Am. Potato J.,
47,148-62.
Reeve, R.M., Timm, H. and Weaver, M.L. (1971) Cell size in Russet Burbank
potato tubers with various levels of nitrogen and soil moisture tensions. Am.
Potato J., 48, 450-6.
Reeve, R.M., Timm, H. and Weaver, M.L. (1973a) Parenchyma cell growth in
potato tubers. I. Different tuber regions. Am. Potato J., 50, 49-57.
Reeve, R.M., Timm, H. and Weaver, M.L. (1973b) Parenchyma cell growth in
potato tubers. II. Cell divisions vs. cell enlargement. Am. Potato J., 50, 71-8.
Rosenstock, G. (1963) Physiologische und anatomische Studien zum Problem der
Wundheilung. I. Untersuchungen tiber Zusammenhange zwischen dem
Wassergehalt verletzten Kartoffelparenchyms und dessen Vernarbungsreak-
tionen nebst Bernerkungen zur Theorie des Wundreizes. Beitr. BioI. Pfl., 38,
275-319.
Salunkhe, D.K. and Pollard, L.H. (1954) Microscopic observations of starch grains
in relation to maturity of potatoes. Proc. Amer. Soc. Hart. Sci., 64, 331--4.
Sattler, R. (1973) Organogenesis of Flowers, Univ. of Toronto Press.
Scaramella Petri, P. (1963) Influenza dell 'intensita' luminosa sulla morfologia del
fusto e delle foglie de Solanum tuberosum. Agric. Ital., 12, 481-90.
Schilde-Rentschler, L. and Roca, W.M. (1987) Tissue culture for the international
exchange of potato and cassava germplasm, in Biotechnology in Agriculture and
Forestry, 3. Potato (ed. Y.P.S. Bajaj), Springer-Verlag, Berlin, pp. 453-65.
Schmidt, H. W. and Schonherr, J. (1982) Fine structure of isolated and non-isolated
potato tuber periderm. Planta, 154, 76-80.
Secor, G.A. and Shepard, J.F. (1981) Variability of protoplast-derived potato
clones. Crop Sci., 21, 102-5.
Seithe, A. (1962) Die Haararten der Gattung Solanum L. und ihre taxolilomische
Verwertung. Botanische Jb., 81, 261-335.
Shadeque, A. and Pandita, M.L. (1982) Effect of cycocel (CCe) as foliar spray on
growth, yield and quality of potato (Solanum tuberosum L.). J. Res. Assam
Agric. Univ., 3, 34-9.
Shepard, J.F., Bidney, D. and Shahin, E. (1980) Potato protoplasts in crop
improvement. Science, NY, 208, 17-24.
Shih, C.Y. and Rappaport, L. (1970) Regulation of bud rest in tubers of potato,
Solanum tuberosum L. VII. Effect of abscisic and gibberellic acids on nucleic
acid synthesis in excised buds. Plant Physiol., 45, 33-6.
Shih, C.Y. and Rappaport L. (1971) Regulation of bud rest in tubers of potato,
Solanum tuberosum L. VIII. Early effects of gibberellin A3 and abscisic acid on
ultrastructure. Plant Physiol., 48, 31-5.
Silva, G.H. (1985) In vitro storage of potato tuber explants and subsequent plant
regeneration. HortSci., 20, 139--40.
Simmonds, N.W. (1965) Chimeral potato mutants. J. Hered., 56, 139--42.
Sip, V. (1972) Eradication of potato viruses A and S by thermotherapy and sprout
tip culture. Potato Res., 15, 270-3.
Sizova, M.A. (1963) Anatomical studies of leaves in some wild potato species
Trudy Prikl. Bot. Genet. Selek., 35, 118-33 (in Russian with English summary).
Sizova, M.A. (1965a) Anatomical leaf structure of some potato species. Trudy
Prikl. Bot. Genet. Selek., 37, 51-108 (in Russian with English summary).
Sizova, M.A. (1965b) Potato leaf pubescence as a systematic character. Trudy
Prikl. Bot. Genet. Selek., 37,109-28 (in Russian with English summary).
Slater, J.W. (1963) Mechanisms of tuber initiation, in The Growth of the Potato
(eds J.D. Ivins and F.L. Milthorpe), Butterworths, London, pp. 114-20.
Slater, J.W. (1968) The effect of night temperature on tuber initiation of the
potato. Eur. Potato J., 11, 14-22.
Smith, D.E. and Palmer, C.E. (1970) Cytokinin-induced tuber formation 00
stolons of Solanum tuberosum. Physiol. Plant., 23, 599-606.
Smith, D.E. and Rappaport, L. (1961) Endogenous gibberellins in resting and
sprouting potato tubers. Adv. Chem. Ser., 28, 42-8.
Smith, D.E. and Rappaport, L. (1969) Gibberellins, inhibitors and tuber formation
in the potato, Solanum tuberosum. Am. Potato J., 46, 185-9l.
Smith, R.H. and Murashige, T. (1970) In vitro development of the isolated shoot
apical meristem of angiosperms. Am. J. Bot., 57, 562-8.
Soliday, C.L., Dean, B.B. and Kolattukudy, P.E. (1978) Suberization: inhibition
by washing and stimulation by abscisic acid in potato discs and tissue culture.
Soliday, c.L., Kolattudy, P.E. and Davis, R.W. (1979) Chemkal and ultra-
structural evidence that waxes associated with the suberin polymer constitute
the major diffusion barrier to water vapor in potato tuber (Solanum tuberosum
L.). Planta, 146, 607-14.
Sopory, S.K. (1979) Effect of sucrose, hormones and metabolic inhibitors on the
development of pollen embryoids in anther cultures of dihaploid Solanum
tuberosum. Can. J. Bot., 57, 2691--4.
Sopory, S.K. and Bajaj, Y.P.S. (1987) Anther culture and haploid production in
potato, in Biotechnology in Agriculture and Forestry, 3. Potato (ed. Y.P.S.
Bajaj), Springer-Verlag, Berlin, pp. 89-105.
Soueges, R. (1922) Recherches sur l'embryogenie des Solanacees. Bull. Bot. Soc.
Fr., 69, 163-78, 236--41, 352-65, 555-85.
Sree Ramulu, K., Dijkhuis, P. and Roest, S. (1984) Genetic instability in
protoclones of potato (Solanum tuberosum L. c.v. 'Bintje'): new types of
variation after vegetative propagation. Theor. Appl. Genet., 68, 515-19.
Stace-Smith, R. and Mellor, F.C. (1968) Eradication of potato viruses X and S by
thermotherapy and axillary bud culture. Phytopathology, 58, 199-203.
Stallknecht, G.F. (1983) Application of plant growth regulators to potatoes,
production, and research, in Plant Growth Regulating Chemicals (ed. L.G.
Nickell), CRC Press, Boca Raton, Florida, Vol. 2, pp. 161-78.
Stallknecht, G.F. (1985) Tuber initiation in Solanum tuberosum: effect of
phytohormones and induced changes in nucleic acid and protein meta-
bolism, in Potato Physiology (ed. P.H. Li), Academic Press, London, pp.
231-60.
Stallknecht, G.F. and Farnsworth, S. (1982a) General characteristics of
References 159
coumarin-induced tuberization of axillary shoots of Solanum tuberosum L.
cultured in vitro. Am. Potato J., 59, 17-32.
Stallknecht, G.F. and Farnsworth, S. (1982b) The effect of the inhibitors of protein
and nucleic acid synthesis on the coumarin-induced tuberization and growth of
excised axillary shoots of potato sprouts (Solanum tuberosum L.) cultured in
vitro. Am. Potato J., 59, 69-75.
Steinberg, C. (1950) Ricerche sull'istogenesi dell'apice vegetativo di alcune specie
del genere Solanum. Nuovo G. Bot. Ital., 57, 319-34.
Sterling, C. and Pangborn, J. (1960) Fine structure of potato starch. Am. J. Bot.,
47,577-82.
Steward, F.e., Moreno, U. and Roca, W.M. (1981) Growth, form and com-
position of potato plants as affected by environment. Ann. Bot., 48 (supp\. 2),
1-45.
Stow, I. (1927) A cytological study on pollen sterility in Solanum tuberosum L. Jap.
J. Bot., 3, 217-37.
Sussex, I.M. (1955) Morphogenesis in Solanum tuberosum L.: apical structure and
developmental pattern of the juvenile shoot. Phytomorphology, 5, 253-73.
Svensson, B. (1962) Some factors affecting stolon and tuber formation in the potato
plant. Eur. Potato J., 5, 28--39.
Svobodova, J. (1964) Freeing potato plants from virus. Abstr. 10th Internat. Bot.
Congr., pp. 485--6.
Sward, R.J. and Hallam, N.D. (1976) Changes in fine structure of the potato
meristem following heat treatment for virus eradication. Aust. J. Bot., 24,
597--605.
Tarn, T.R. (1971) Protecting potatoes. Nature, Lond., 234, 425--6.
Taylor, e.E. (1953) Vegetative development of the potato plant. Ann. Appl. BioI.,
40,778-88.
Thomas, E., Bright, S.W.J., Franklin, J., Lancaster, V.A. and Miflin, B.J. (1982)
Variation amongst protoplast-derived potato plants (Solanum tuberosum cv.
'Maris Bard'). Theor. Appl. Genet., 62, 65-8.
Thomson, A.D. (1956) Heat treatment and tissue culture as a means of freeing
potatoes from virus Y. Nature, Lond., 177,709.
Tilney-Bassett, R.A.E. (1986) Potato Chimeras, Edward Arnold, London.
Tizio, R. (1969) Action du CCC (chlorure de (2-chloroethyl)-trimethylammonium)
sur la tuberisation de la pomme de terre. Eur. Potato J., 12,3-7.
Tizio, R. (1972) Effet de la lumiere sur la tuberisation de la pomme de terre. Potato
Res., 15, 257--62.
Tizio, R. and Tizio, e.R. (1981) Role du tubercule-mere dans la tuberisation de la
plante de pomme de terre (Solanum tuberosum L., variete Bintje). Phyton,
Buenos Aires, 40, 49-59.
Toosey, R.D. (1963) The influence of sprout development at planting on sub-
sequent growth and yield, in The Growth of the Potato (eds J.D. Ivins and F.L.
Milthorpe), Butterworths, London, pp. 79-95.
Tsutsumi, F.V., Olivas, E.E. and Alvarez-Fuertes G. (1972) Ultrastructura del
floema de las hojas de papa (Solanum tuberosum) I. (Eng\. abstr.). Revista
latinoamericana de Microbiologia, 14, 153--64.
Tuan, D.Y.H. and Bonner, J. (1964) Dormancy associated with repression of
genetic activity. Pl. Physiol., Lancaster, 39, 768-72.
Twell, D. (1988) Genetic engineering of potato. Plants Today, 1, 191--6.
Umaerus, M. (1987) True potato seed. Proceedings of the 10th Triennial Conference
of the European Association for Potato Research, Aalborg, Denmark.
van Harten, A.M. (1972) A suggested method for investigating L J constitution in
periciinal potato chimeras. Potato Res., 15, 73-5.
Venning, F.D. (1954) The relationship of illumination to the differentiation of a
morphologically specialized endodermis in the axis of potato, Solanum
tuberosum L. Phytomorphology, 4, 132-9.
Verma, S.e. (1966) Induction of stolons and tubers by the leaves of the potato
(Solanum tuberosum L.). Eur. Potato 1.,9,259-60.
Vogt, E., Schonherr, J. and Schmidt, H.W. (1983) Water permeability of periderm
membranes isolated enzymatically from potato tubers (Solanum tuberosum L.).
Planta, 158, 294-301.
von Staden, J. and Dimalla, G.G. (1977) The distribution of cytokinins in
tuberizing potatoes. Ann. Bot., 41, 741-6.
Vreugdenhil, D. and Struik, P.e. (1989) An integrated view of the hormonal
regulation of tuber formation in potato (Solanum tuberosum). Physiol. Plant.,
75,525-31.
Wang, Po-jen and Hu, Ching-yeh (1985) Potato tissue culture and its applications
in agriculture, in Potato Physiology (ed. P.H. Li), Academic Press, New York,
pp. 503-77.
Wang, T.L. and Wareing, P.F. (1979) Cytokinins and apical dominance in Solanum
andigena: lateral shoot growth and endogenous cytokinin levels in the absence
of roots. New Phytol., 82, 19-28.
Wardlaw, C.W. (1955) Embryogenesis in Plants, Methuen & Co. Ltd, London.
Wareing, P.F. (1982) The control of development of the potato plant by endo-
genous and exogenous growth regulators, in Chemical Manipulation of Crop
Growth and Development (ed. J.S. McLaren), Butterworths, London, pp.
129-38.
Wareing, P.F. (1983) Hormonal control of stolon and tuber development,
especially in the potato plant, in Strategies of Plant Reproduction (ed. W.J.
Meudt), BARC symposium no. 6. Allanheld, Osmun, Totowa, pp. 181-95.
Wareing, P.F. and Jennings, A.M.V. (1979) The hormonal control of tuberisation
in potato, in Plant Growth Substances (ed. F. Skoog), Springer-Verlag, Berlin,
pp. 293-300.
Weaver, J.E. (1922) Development and Activities of Roots of Crop Plants, Carnegie
Institute, Washington.
Webb, K.J., Osifo, E.O. and Henshaw, G.G. (1983) Shoot regeneration from
leaflet discs of six cultivars of potato (Solanum tuberosum subsp. tuberosum).
Plant Sci. Lett., 30, 1-8.
Weinheimer, W.H. and Woodbury, G.W. (1967) The anatomy and morphology of
the abscission layer in two cultivars of Solanum tuberosum L. Am. Potato 1. , 44,
402-8.
Went, F. W. (1959) Effects of environment of parent and grandparent generations
on tuber production by potatoes. Am. 1. Bot., 46, 277-82.
Wenzel, G., Przewozny, T., Sopory, S.K. and Melchers, G. (1979) Comparison of
single cell culture derived Solanum tuberosum L. plants and a model for their
application in breeding programs. Theor. Appl. Genet., 55, 49-55.
Westcott, R.J., Henshaw, G.G. and Roca, W.M. (1977) Tissue culture storage of potato
germplasm: culture initiation and plant regeneration. Plant Sci. Lett., 9, 309-15.
References 161
Wheeler, V.A., Evans, N.E., Foulger, D., Webb, K.J., Karp, A., Franklin, J. and
Bright, S.W.J. (1985) Shoot formation from explant cultures offourteen potato
cultivars and studies of the cytology and morphology of regenerated plants.
Ann. Bot., 55, 309-20.
Wigginton, M.J. (1973) Diffusion of oxygen through lenticels in potato tuber.
Potato Res., 16, 85-7.
Wigginton, M.J. (1974) Effects of temperature, oxygen tension and relative
humidity on the wound-healing process in the potato tuber. Potato Res., 17,
200-14.
Williams, E.J. (1955) Seed failure in the Chippewa variety of Solanum tuberosum.
Bot. Gaz., 117, 10-15.
Wodicka, B. and Wenzel, H. (1971) Auftreten von Callose im Speicherparenchym
der Kartoffelknolle. Potato Res., 14, 150-3.
Woolley, D.J .. and Wareing, P.F. (1972a) The role of roots, cytokinins and apical
dominance in the control of lateral shoot form in Solanum andigena. PMnta,
105,33-42.
Woolley, D.J. and Wareing, P.F. (1972b) The interaction between growth
promoters in apical dominance. I. Hormonal interaction, movement and
metabolism of a cytokinin in rootless cuttings. New Phytol., 71, 781-93.
Woolley, D.J. and Wareing, P.F. (1972c) The interaction between growth pro-
moters in apical dominance. II. Environmental effects on endogenous cytokinin
and gibberellin levels in Solanum andigena. New Phytol., 71, 1015-25.
Wooster, P. and Dixon, T.J. (1986) Micropropagation - an aid in the production of
new varieties, in The Production of New Potato Varieties: Technological
Advances (eds G.J. Jellis and D.E. Richardson), Cambridge University Press,
Cambridge, pp. 142-5.
Wurr, D.C.E. (1974) Individual tuber growth. National Vegetable Research Sifltion.
24th Annual Report, 1973, pp. 65--6.
Yamaguchi, M., Timm, H. and Spurr, A.R. (1964) Effects of soil temperature on
growth and nutrition of potato plants and tuberization, composition, and
periderm structure of tubers. Proc. Amer. Soc. Hart. Sci .• 84, 412-23.
Yoshida, M. (1970) Physio-ecological studies in potato plant. V. On the starch
grains found in the stem tissues. Mem. Fac. Agric. Hokkaido Univ., 7, 209-16
(in Japanese with English summary).
Young, W.J. (1922) Potato ovules with two embryo sacs. Am. J. Bot., 9, 213-14.
Young, W.J. (1923) The formation and degeneration of germ cells in the potato.
Am. J. Bot., 10, 325-35.
CHAPTER 4
Mineral nutrition
P.M. Harris
4.1 INTRODUCTION
When considering the mineral nutrition of a crop, there are a number of

questions the grower attempts to answer. For the immediate requirements
of yield, it must be known what nutrients are limiting, and how much needs
to be applied. Since the price of the product is dependent upon its quality,
either in terms of its size, its dry matter content, its content of reducing
sugars or some other characteristic required for specific market outlets,
then the grower should know how these qualities may be affected by the
fertilizers applied. He will usually know the price to be paid for the
nutrient carriers used, but he should also know whether the form in which
specific nutrients are applied affects the response to it in terms of yield and
quality. Since he has to make a choice of how and when to apply the plant
nutrients, he should also know how the placement and timing of the
nutrient carriers he intends to use might be expected to influence yield and
possibly tuber quality.
In many parts of the world, particularly in the tropics and sub-tropics, the
grower may not have access to or be able to afford fertilizers, and must
consider the use of other materials such as animal manures, or green manures
to supply essential minerals. Similar problems, but for different reasons, may
confront 'organic farming' practitioners in temperate agriculture, who from
conviction, or from the incentive of attractive market prices, may reject the
use of inorganic fertilizers, in order to satisfy the small but growing market
for food produced by so called 'organic' farming methods.
It is frequently stated, and usually true, that no two farms are alike. The
nutrient status of the soil is in a state of flux, under the influence of
changing cropping sequences, fertilizer use and climatic conditions, while
biotic hazards such as weeds, pests and diseases may affect yield and
nutrient demand. It is not surprising that to give an accurate prediction of
the nutrient requirement for a specific crop of potatoes would require a
reasonable to assume that yield, price and the cost of fertilizer inputs are
Nutrient accumulation and use 163
degree of clairvoyance which few have been able to demonstrate. It is
the major factors determining the grower's potential use of fertilizers, since
his major objective must be to maximize his profit. However in many
countries environmental and health concerns may increasingly be expected
to modify decisions.
It is therefore assumed that the grower's major objective with respect to the
mineral nutrition of any of his crops is, subject to economics, environmental
and health concerns, to ensure that yield is unlimited by mineral elements.
The mineral element demand is dependent upon total dry matter yield, which
in turn is dependent upon the amount of radiation intercepted (IR) by a
crop and the efficiency with which it is utilized (e) (Monteith, 1977). In
temperate regions, it has been argued, tuber yield is primarily determined
by the total amount of radiation intercepted, since e and the partitioning of
dry matter between tubers and the rest of the crop appear to be relatively
constant (Allen and Scott, 1980). It might therefore be expected that the
optimal quantity of nutrients to apply would be that which would sustain a
closed crop canopy for the duration of the potential growing season.
The mineral element supply will in total be determined by amounts
available from natural resources, such as rainfall, symbiotic and non-
symbiotic nitrogen fixation, weathering of rocks, and residues from
previously applied fertilizers and manures. While aequate for some
elements, these sources are usually insufficient to meet the demand for all
the essential elements, and may be supplemented by the use of fertilizers
applied to the crop. The proportion of these applied nutrients taken up by
the crop, and therefore effective in meeting the crop's demands, will
depend on the specific nutrient in question, on the form in which the
nutrient is supplied, its time of application, the placement method used
and soil conditions.
These questions can, in principle, be resolved by field experiments in which
output/input relationships are established for specific nutrients or nutrient
combinations. The optimal level of fertilizer application, subject to cost
price relationships, will then depend upon the form of the output/input
relationship and the amount of nutrients supplied from the soil (Fig. 4.1).
These are a formidable set of variables to resolve, and clearly any
answers will be specific for a particular site, season, possibly genotype and
related to a specific end product, such as the yield of particular size grades,
perhaps with specified dry matter contents or other quality characteristics.
In this chapter the basic objective is to consider what general principles
may be established which can assist decision making.
4.2 NUTRIENT ACCUMULATION AND USE
There are some 13 essential mineral elements, which are taken up in

various quantities by all crops (Table 4.1). Table 4.2 gives a representative
Distribution Farmer's objective Fertilizer
subject to economic,
of dry matter
environmental &
health concerns
Carrier Input/output I
Timing relations
Placement
Total dry Mineral Yield Mineral Mineral

matter nutrient unlimited by nutrient nutrient
yield demand mineral uptake supply
nutrient
supply
I Environment I Level of nutrient

supplied by the
t------tl soil: starting
position of field
on the response curve
i
Intercepted Efficiency Natural I Residues
radiation of conversion r sources from
(IR) of intercepted fertilizers
radiation into & manures
dry matter
(e)
Figure 4.1 Factors influencing decisions about fertilizer use.

Table 4.1 Concentrations of mineral nutrient
elements in plant material at levels considered adequate
(from Epstein, 1972)
Element Concentration in Relative number of
dry matter atoms with respect
to molybdenum
ppm
Molybdenum 0.1 1
Copper 6 100
Zinc 20 300
Manganese 50 1000
Iron 100 2000
Boron 20 2000
Chlorine 100 3000
%
Sulphur 0.1 30000
Phosphorus 0.2 60000
Magnesium 0.2 80000
Calcium 0.5 125000
Potassium 1.0 250000
Nitrogen 1.5 1000000
Reproduced by permission of John Wiley and Sons.
selection of data for the quantity of essential mineral elements taken up by

potatoes and removed in the tubers.
Some data showing the removal of 12 essential mineral elements in fresh
tubers are given in Table 4.3. The data from the different sources agree
quite well in terms of the orders of magnitude, but a considerable degree of
variability must be accepted. For example, nutrient concentration is not
independent of yield as shown in Table 4.4. Yield increases induced by N
fertilizer resulted in a decrease in K and P and an increase in N removed
per tonne of tuber fresh weight. In data calculated from Gunasena (1969)
the effect of season, variety and the rate and timing of Nand K fertilizer
resulted in the following ranges in the removal of mineral nutrients in fresh
tubers: N, 2.28-3.57; P, 0.40-0.62; K, 3.70--5.41 kg t- 1 .
To resolve all the questions implied in Fig. 4.1 would clearly be beyond
the capacity of field experiments should all essential elements be limiting in
some degree. Fortunately for the grower and the sanity of the applied
scientist, in many soils, trace element supply is not a problem, or if
identified may often be corrected fairly inexpensively, since the quantities
involved are small. Indeed, in many temperate areas the major nutritional
problem to be resolved is how much N to apply, since this nutrient is the
one which is most likely to be limiting on most soil types and in most
seasons. Unlike P or K, it is rarely possible to build up in the soil through
Table 4.2 Selected characteristics of various potato cultivars grown in various parts of the world (adapted from Ezeta and
McCollum, 1972)
Location Peru Colombia Utah California Maine Maine North Netherlands England
Variety Renacimiento 3 varieties Russet White Green 4 varieties Carolina Mentor Craig's
Burbank Rose Mountain 4 years Pungo Royal
Plant density plants

ha·! x 103 33.3 45.6 ? 43.0 35.9 35.9 37.7 40.0 35.9
Planting to tuber
initiation 125 44-51 60 60--65 60 48-60 50 48 58
Planting to harvest 180 109 138 119-124 112 89-97 100 120 142
Total dry matter

production t ha·! 12.6 11.9-18.6 9.9 10.6-12.7 7.9 4.5-8.8 7.5 10.2 12.7
Tuber dry matter/
total dry matter 0.70 0.77-0.82 0.77 0.75-0.85 0.75 0.60-0.80 0.83 0.95 0.96
Rate of tuber growth
kg·! ha·! day·! 224 280-360 170 180-290 141 132-204 350 184 144
Rate of uptake N 2.5 2.5-4.0 2.2 3.2-3.3 4.2 1.7-5.1 3.3 1.7 2.2
kg ha·! day·! P 0.22 0.38-0.45 0.25 0.3-0.48 0.39 0.14-0.50 0.22 0.30
K 6.6 4.7-5.6 2.6 3.8-6.3 5.8 2.6-6.9 4.5 3.34
% tuber dry matterN 0.9 1.5 1.4 1.1-1.3 1.8 1.1-1.8 1.6 1.1 1.2
at harvest P 0.18 0.2 0.14 0.15 0.18 0.17-0.26 0.25 0.18
K 2.6 1.3 1.4 1.5-1.7 1.8 1.5-2.5 2.1 1.8
Nutrients removed N 80 107 107 100 107 147

in tubers kg ha·! P 16 11 11 16 22
K 230 107 107 131 220
Table 4.3 Mineral nutrient elements
removed t- 1 of fresh tubers
Element 2 3
kg
Nitrogen 2.68 2.36

Phosphorus 0.62 0.53
Potassium 3.93 4.67 4.55
Calcium 0.07 0.2
Magnesium 0.22 0.13
Sulphur 0.21 0.48
Zinc 0.001 8 0.005
Copper 0.00143 0.00219
Manganese 0.00134 0.0021
Iron 0.0042
Boron 0.00062
Molybdenum 0.000037
Sodium 0.23
Data calculated from the following sources: (1)
Kunkel et al. (1973); (2) Cooke (1967); (3)
Gunasena (1969).
Table 4.4 The effect of N on yield and

the removal of N, P and K t- 1 of tltber
fresh weight
Treatment Yield N P K
N tubers
(kg ha- 1 ) (t ha- 1) (kg ta- 1 )
0 11.95 2.81 0.50 5.41
94 19.57 3.26 0.46 5.24
188 22.74 3.57 0.45 4.73
Data calculated from Gunasena (1969).
the use of fertilizer, reserves of available nitrogen which will be carried

over from season to season, and the amounts required for satisfactory
yields tend to be relatively large (Table 4.1).
In the light of this it is perhaps not surprising that much of the
experimental work on the nutrition of the potato is biased towards N.
Perrenoud (1983) has presented data for fertilizer use on potatoes in
various countries, while Fig. 4.2 shows the trend in the use of N, P and K
on the potato crop in England and Wales over the period 1958-86. It shows
a rising trend over most of the period, although there was a slight decline at
the end. The ratio of N:P2 0 5 :K2 0 was 1.00:1.07:1.65 at the beginning of
the period narrowing, by virtue of a reduction in the proportion of K2 0, to
168 Mineral nutrition
240
00
00 OK
000 •
200
o
o o 00
.... ·.N
•• •
0
• •
';"eo 160
..r::
•
:::.
0)
"0
CD
•
'5. 120
a.
eo
Q;
~
1:: 80
CD
u..
/:,
40
1958 1963 1968 1973 1978 1983 1986
Figure 4.2 Trends in fertilizer use in England and Wales 1958-86. (PMB, ADAS/
RothamstedIFMA surveys.)
1.00:1.04:1.35 at the end. Fertilizer was used on practically all the farms
growing potatoes and the average quantities used corresponded closely to
the average amounts recommended for the crop.
4.3 RELATIONS BETWEEN MINERAL NUTRIENTS, GROWTH AND

YIELD
Tuber yield is considered to be a product of three major processes:

radiation interception, conversion of intercepted radiation to dry matter
and the partitioning of dry matter between tubers and the rest of the plant.
It is most easy to demonstrate the effects of nutrient supply on radiation
use and dry matter partitioning where the nutrient in question is in short
supply. Since this is most frequently N, for reasons given in Section 4.1, the
following example is concerned with this nutrient and is illustrated by data
from a field experiment carried out on Reading University Farm in 1987
under the temperate climatic conditions of S.E. England and on a fine
sandy loam soil (Harris, unpublished). Two contrasting cultivars were
grown with and without supplementary irrigation at six rates of nitrogen:
0, 30, 70, 150, 340 and 750 kg ha- 1 . Both cultivars were planted on 8 and
9 April. The second-early variety Wilja had completed its life cycle in early
Relations between mineral nutrients, growth and yield 169
August. The late-maturing variety Cara still retained foliage on the high N
treatments when the final harvest was taken on 12 to 14 October. Forty-
eight mm of irrigation was applied to Wilja and Cara in early July and a
further 20 mm to Cara on 11 August. The total amounts of radiation
intercepted by the variously treated crops were estimated from the
proportion of radiation intercepted by the crop canopy, derived from
measurements of percent ground cover (Burstall and Harris, 1983) and
incident radiation obtained from a Kipp solarimeter.
4.3.1 Total dry matter yield

Figure 4.3 shows the relationships between total dry matter yield and
accumulated intercepted radiation for the first part of the growing season,
before losses of dry matter through leaf senescence became important. For
Wilja (Fig. 4.3a), up to 103 days after planting (dap) , accumulated dry
matter was linearly related to the amount of radiation intercepted by the
crop, which accounted for 98% of the variation in dry matter yield. The
effects of nitrogen and water supply could be almost entirely attributed to
their effects on the size of the crop canopy. For Cara (Fig. 4.3b), up to
131 dap, total dry matter yield was linearly related to accumulated
2000 (a) (b)

•
1800
1600 •
cf
V
~ 1400
E
~Iia/
• 00
31200 o •
CD 0
•
:I" ~
~ 1000
Cara
• 750 kg N ha- 1
~
"0 800
..s
[]
~
I- 600
•
~
400
200
0 200 400 600 800 1000 0 200 400 600 800 1000 1200 1400
Intercepted radiation (MJm- 2 )
Figure 4.3 Relationships between total dry matter yield and intercepted radiation
for two contrasting potato cultivars: (a) Wilja; (b) Cara. Open symbols =
unirrigated, closed symbols = irrigated (Harris, unpublished).
intercepted radiation for levels of N up to 340 kg ha- 1 , although etota ! was
apparently 22% smaller than for Wilja. With 750 kg N ha- 1 however, the
efficiency of use of intercepted radiation was lower and became progres-
sively worse as the level of IR increased (Table 4.5). The small positive
intercept was probably due to an overestimation of the amount of crop
cover in the early part of the season, and some contribution of dry matter
from the parent tubers.
Table 4.5 Relations between intercepted radiation and

total dry matter yield (Ytot) in two contrasting potato
cultivars, before leaf senes,cence became important
(Harris, unpublished data)
Cultivar Level of N ,.z
(kg ha- 1)
Wilja 0-750 Ytot = 139.3 + 1.23IR 0.98
Cara 0-340 Y tot = 186.4 + 0.96 0.95

750 Ytot = 75.2 + 1.17IR - 3.94IR2 0.96
Thus, except for the highest level of N applied to Cara, and while noting
that for this cultivar etota ! is lower than that of Wilja, it is clear that there
was a direct relationship between IR and total dry matter yield and that
N affected total yield through its effect on the amount of radiation
intercepted.
Millard and Marshall (1986) showed that with the maincrop cultivar
Maris Piper grown in northern Scotland, yield increases from N fertilizer
were due to increased radiation interception and that the efficiency of
conversion of radiation into dry matter was little affected. Using direct
measurements of photosynthesis on a range of varieties grown in the field
in eastern England, Firman and Allen (1988) demonstrated that the rate of
photosynthesis in the leaves of plants grown without applied N reached
levels as high as those receiving N in all experiments,. In some cases the
photosynthetic rate was reduced by N fertilizer, but this was attributed to
the effect of N on the size of the canopy which in turn increased water
stress and stomatal resistance. They concluded that any increase in dry-
matter production by the addition of N fertilizer was achieved through
effects on leaf area duration rather than on the efficiency of conversion of
incident radiation.
There would therefore seem to be no reason to disagree with the
conclusion of Gregory et al. (1981), that the available N in many arable
soils is not low enough to limit leaf photosynthesis of crop plants directly
although, as shown for the cultivar Cara, application rates far in excess of
those normally applied may depress etota!'
The effect of the level of fertilizer N applied on the total amount of
Relations between mineral nutrients., growth and yield 171
radiation intercepted throughout the life of the crop is shown in Fig. 4.4 for
the two cultivars with and without irrigation. The figures clearly show the
diminishing returns to increasing levels of N, but there is no evidence to
suggest that very high rates led to a reduction in the total amount of
radiation intercepted for either cultivar with or without irrigation. It is also
demonstrated that for any given level of N, Cara was abk to intercept
more radiation than Wilja. Since Cara utilized radiation apparently less
efficiently than Wilja, both this observation and the ability of the crop to
extract more N from the soil may be accounted for by the larger root
system of this cultivar (Harris, 1983). Roots were not included in the
estimate of total dry matter yield, which would therefore bias the estimate
against Cara, while the larger root system could permit the more complete
extraction of soil N.
4.3.2 Tuber dry matter

In Figure 4.5a, the relationship between IR and tuber dry matter yield is
shown for Wilja. Tuber dry matter yield was linearly related to the amount
2000
1800
0
•
1600 •
1400
0
0 •
N
...,E
6 1200
c
0
•
~ 1000
~ ~~ ••
?
"0
.'!l
g. 800
~
~
600
400
200
II I ,
03715 34 75 03715 34 75
Fertilizer (N gm-2 )
Figure 4.4 Relationships between applied N fertilizer and the total amount of
radiation intercepted during the course of growth for two contrasting cultivars
(Wilja = 0, Cara = D) grown with (closed symbols) and without (open symbols)
supplementary irrigation (Harris, unpublished data).
of radiation intercepted by the crop canopy, but the rate of N did affect the
relationship. At levels of N exceeding 70 kg ha- 1 , the initial growth of
tubers was delayed relative to the accumulation of radiation, but etuber was
increased somewhat (Table 4.6). Since total dry matter yield was directly
proportional to the amount of radiation intercepted and the relation was
independent of the amount of N applied, increasing the level of N diverted
more dry matter to canopy production, which eventually resulted in an
increase in the total amount of radiation intercepted and a net increase in
tuber dry matter yield.
2200
.0
2000
1800
1600
if'
• •
E 1400
El
:§, 1200
"iii
;: ••
~ 1000
"0
Q;
.0 ;800
:J
I-
600
400
200
•
0 ~~ ~ ~1~1~ 0 ~ ~~~1~1~1~1~1~
Intercepted radiation (MJm- 2 )
Figure 4.5 Relationships between intercepted radiation and tuber dry matter
yield at different levels of N in (a) the cultivar Wilja, N kg ha- 1 : 00-70; e 150-750
and (b) the cultivar Cara, 0, 0-70, e, 150; D, 340 and ., 750 kg N ha- 1 (Harris,
unpublished data).
Figure 4.Sb shows how tuber growth in the cultivar Cara was delayed
relative to the interception of radiation, as the level of N fertilizer
increased, a phenomenon which was much more exaggerated in this
cultivar.
The relationships between IR and tuber yield given in Table 4.6 have
been used to calculate the amount of radiation intercepted up to the point
where a significant amount of tuber growth has occurred (Table 4.7),
arbitrarily taken to be when tuber dry matter yield reached 200 g m- 2 • It
Table 4.6 Relations between tuber dry matter yield (Ytub ) and IR for
two contrasting potato cultivars (Harris, unpublished data)
Cultivar Level of N
(kg ha- 1 )
Wilja 0-70 Ytub = -23.95 + 1.20IR 0.93

150-750 Ytub = -166.32 + 1.26IR 0.93
Cara 0-70 Ytub = -102.51 + 1.04IR 0.93

150 Ytub = -460.95 + 1.36IR 0.89
340 Y tub = 268.38 - 0.846IR + 0.001066R2 0.91
750 Y tub = 567.10 - 1.692IR + O.001284R 2 0.96
Table 4.7 The effect of cu-Ltivar

and level of fertilizer N on the
amount of radiation intercepted
before the accumulation of 200g
m-2 of tuber dry-matter (Harris,
unpublished data)
Cultivar Level of N (kg ha· l )
0-70 150 340 760
Wilja 190 300 300 300
Car a 300 500 700 1040
illustrates the very marked contrast between the cultivars in their response
to the effects of N on early tuber growth.
For Wilja, the efficiency of conversion of radiation into tuber dry matter
(etuber) was only slightly affected by the level of N applied. For Cara, etuber
changed with the level of N applied and for anyone level at rates in excess
of 150 kg N ha- 1 it changed dramatically as the amount of radiation
intercepted increased (Fig 4.5). Estimated values of etuber at selected levels
of accumulated IR are given in Table 4.8.
Table 4.8 The change with N fertilizer and
amount of IR in etub for two contrasting potato
cultivars (Harris, unpublished data)
Cultivar Level of N Accumulated IR (MJ m- 2)
(kg N ha- 1 )
600 1000 1400 1800
Wilja 0-70 1.2 1.2
150-750 1.26 1.26
Cara 0-70 1.04 1.04 1.04
150 1.36 1.36 1.36
340 0.43 1.29 2.14 2.99
750 0.88 1.90 2.93
2200
2000
1800
1600
1400
1200
1000
Wilja 03 Cara
2
800
600
400
200
200 400 600 800 1200 1400 1600 1800 2000

Intercepted radiation (MJm-2 )
Figure 4.6 Relationships between final total tuber dry matter yield and inter-
cepted radiation for two contrasting cultivars, Wilja (open symbols), Cara (closed
symbols), grown without (0, e) and with (0, _) irrigation. Numbers == levels ofN,
1 == 0 and 6 == 750 kg N ha- 1 (Harris, unpublished data).
The exceptionally high values of etuber recorded for the cultivar Cara at
the latter end of the growing season, can only be attributed to a major
redistribution of dry matter from stems and leaves to tuhers. The optimal
rate of N to apply will clearly depend upon the extent to which this
redistribution of dry matter can offset the delay in tuber growth which
increased progressively with the rate of N applied. Such compensation is
almost complete in the case of Wilja since there was a very close relationship
between final tuber dry matter yield and total accumulated IR (Fig. 4.6). It
can be seen that each increment of N has increased yield by increasing IR
both in the presence and absence of irrigation. While there was a good
linear relationship between IR and tuber dry matter yield in the cultivar
Cara (Fig. 4.6), the data are more variable as might be expected from the
effects of N on the distribution of dry matter. However they do demon-
strate how the increase in etuber in the latter stages of growth can
compensate for the delay in tuber growth due to N in the earlier part of the
season.
Millard and Marshall (1986) showed that increases in light interception
in response to applied N in the variety Maris Piper, were also partially
offset by a concomitant decrease in the proportion of the dry weight
partitioned to tubers, particularly during the initial period of bulking. The
effect of N application on tuber yield was therefore dependent on the date
of harvest. Dyson and Watson (1971) also demonstrated that N reduced
the early rate of growth of tubers.
While the rate of tuber growth early in the life cycle may be reduced by
increasing levels of N, as shown here, there is little evidence to suggest that
the actual time of tuber initiation in the field is affected. This can be seen in
Fig. 4.7, where the effect of N on the number of tubers counted during the
period of tuber initiation is given for two cultivars. While in solution
culture experiments (Kraus, 1978; Sattlemacher and Marschner, 1979) the
presence of N may completely inhibit tuber initiation, it would seem
difficult to detect under normal field conditions.
tubers/plant
14 12 June
Desiree
12
10
8
King Edward
~a,
12 June
6
25 May
~
30 May
o 50 100150200250 0 50 100150200250
Fertilizer N (kg ha- 1 )
Figure 4.7 The effect of level of N fertilizer on the number of tubers during tuber
initiation in two cultivars (Harris, unpublished data).
Thus the effect of N on tuber yield can be ascribed to two processes: a
positive effect on the size of the crop canopy which directly affects the
amount of radiation intercepted and thus the accumulation of dry matter,
and a negative effect on diverting dry matter away from tuber growth, but
without necessarily inhibiting tuber initiation. From the data discussed
above, it is clear that the importance of the latter effect is variety
dependent and may be expected to increase as the tendency for varieties to
invest more dry matter in foliage production increases.
4.3.3 Effect of N, P and K on leaf area development

It is difficult to find any experiments in which the effects of P and K
fertilizer have been measured in similar terms to those described for N
above. The effects of these nutrients on leaf area index (L) measured in
pot experiments (Watson and Wilson, 1956) showed that N, P and K
increased dry matter yield mainly by increasing leaf area per plant, while
effects on leaf efficiency as measured by net assimilation rate (NAR, the
net increase in dry matter per unit area of leaf per unit of time), were
relatively small and erratic. It was noted that the stage of growth at which
L was most affected differed with nutrient: N greatly increased L through-
out the life of the crop; P increased L in the early stages of growth, but
hastened the senescence of leaves, thus depressing L towards the end of
the growing period. K, on the other hand, had little effect in the early
stages of growth, but increased L considerably later in the season and
delayed the senescence of leaves (Fig. 4.8).
In the field, the level of soil nutrient supply can be less readily controlled
than in pots, and as mentioned earlier, in British, and presumably in other
intensively farmed temperate agricultural conditions, N is usually the most
important nutrient limiting yield, owing to the build-up of P and to a lesser
extent K reserves in the soil following intensive fertilizer use. Thus in
16
o
May June July May June July May June July
Figure 4.8 The effects of N, P and K on leaf area of potatoes grown in pots with
(e) and without (0) the nutrient. (Redrawn from Watson and Wilson, 1956).
Fertilizer requirements 177
southern England, Gunasena and Harris (1968, 1971) showed that N
fertilizer applied at the time of planting substantially increased L over most
of the growing season, while the effect of K was much smaller and was
confined to the middle of the season. Dyson and Watson (1971) working on
a heavier soil at Rothamsted in southern England also showed that
increases in L due to P and K were small compared with the response to N
(Fig. 4.9). As in the pot experiments N increased L throughout growth, P
only increased L up to 4 weeks after emergence, while the effect of K was
only evident in the middle of the growing season. On P-deficient volcanic
soil in Rwanda, Haverkort (1985) showed that N fertilizer had the
strongest influence on ground cover early and late in the season. P and K
led to earlier closure of the crop canopy, but did not lengthen the growing
period.
3
N4
~2 K1
"tl
.!:
C\l
.Q)
Cd
1ii
Q)
1
-J
0
2 6 10 14 2 6 10 14 2 6 10 14
Number of weeks after emergence
Figure 4.9 The effects of N, P and K on changes with time in leaf area index of
potatoes, cv. King Edward. No: 0, Nl:94, N2:188 and N4:376 kg N ha-\ Po: 0,
PI: 156 kg P ha- 1 ; Ko:O, Kl:82 kg K ha- 1 . (Redrawn from Dyson and Watson,
1971).
4.4 FERTILIZER REQUIREMENTS
In order to apply optimal rates of plant nutrients, usually N, P and K, the

basic biological information required is the relationship between nutrient
supply and tuber yield and an estimate of the nutrient status of the
particular field in which the crop is to be grown. The effect of the nutrient
supply on tuber quality must also be taken into account. It should be
possible to define the general form of the relationship between plant
nutrients and yield from well conducted field trials, but the nutrient status
of the soil must be determined or estimated for each site. Unpredictable
events occurring during the growing season, such as the incidence of pests
and diseases, or the incidence and quantity of rainfall or variation in
radiation receipts may alter the yield - nutrient relationships, although
some opportunities may exist for assessing and correcting the nutrient
status of the crop during the course of growth.
4.4.1 Relationships between fertilizer supply and yield

In a classic paper, Crowther and Yates (1941) examined all the experimen-
tal results available to them with nutrients at three or more levels. They
found that the response curve was an exponential, following the pattern
proposed by Mitscherlich (1913) whose formula relating level of fertilizer
to yield they adopted with slight modifications, namely:
Yl = Yo + d(l - lO-kX)
where Yo = yield without the nutrient in question, d = the limiting
response, Yl = the yield with a dressing of the nutrient in x amount, k = a
value which determines the rate at which response falls off with successive
increments of nutrient and which was assumed to be constant for each of
the three principal classes of fertilizer. In essence, yield bears an
asymptotic relationship to nutrient supply.
A weakness of the experimental data on which the response curve was
based was that the levels of nutrients tested were too low, and that
interactions between nutrients had to be ignored because they could not be
estimated satisfactorily. In a later review, Boyd (1961) presented evidence
which indicated that the interactions between nutrients were larger and
more consistent than had been supposed and that they affected both the
magnitude of the response to a given level of nutrient and the degree to
which the response falls off with increasing dressings. Boyd concluded that
the concept of an exponential response curve did not fit the facts and that
for each nutrient the response curve rises to a maximum and then falls off
again. The level of a nutrient at which the response is maximized would
vary, Boyd argued, with many factors, but particularly with the amount of
other nutrients in the soil and applied as a basal dressing.
Similarly Holliday (1963) commented that 'anyone who has handled
comprehensive fertilizer data will realize that the actual curves show a point
of inversion'. Holliday suggested that it is usua~ to use a quadratic curve of
the form: Y = a + bx - cx2 or a square root function y = a + bx - CXO. 5
where y = yield, x = the level of plant nutrient, and a, band care
constants, the latter equation having the merit that it is not symmetrical
about the point of inversion.
Summarizing 51 experiments carried out between 1956 and 1962 in
Britain, Birch et al. (1967) found that square root functions fitted their
data better than quadratic functions. Sharpe (1969) compared these two
functions using data from 95 experiments reported by Boyd and Dermott
(1964), and found that on statistical grounds it was difficult to select which
function best fitted the experimental data, but concluded that on biological
grounds, the square root function was more appropriate; presumably
because there is little biological evidence to suggest that after an optimal
level of fertilizer has been applied, yield would decrease at the same rate as
it increased before the optimum was reached.
Boyd (1970) and Fitts (1974) suggested that, compared with a curve, as
good or better fit to data relating yield to fertilizer level might be obtained
by two straight lines. Boyd reviewed the evidence for this type of response
for a number of crops. The investigation was prompted by experiments
with sugar beet, where the general form of the relationship between
fertilizer N and sugar yield had a steeply-rising portion with a fairly abrupt
transition to an almost linear and horizontal portion when larger amounts
of N were applied. The relationships only became evident when experi-
ments were grouped according to the amount of N required to reach the
transition point.
The failure of voluminous experiments with fertilizer to characterize the
form of the response curve to N, P and K was probably due to the paucity
of levels tested and the relatively narrow range of the response curve
investigated. At that time Boyd commented that there was little informa-
tion for more than four amounts of N on potatoes, but that many tests with
four amounts indicate that this crop would be expected to behave similarly
to the sugar beet crop. Data presented for the response to Nand P from
two sets of experiments (Fig. 4.10) gave some support for this view.
In a series of 98 experiments carried out on potatoes (cv. Bintje) in The
Netherlands between 1972 and 1982 (Neeteson and Zwetsloot, 1989), the
form of the response curve was adequately described by a common exponen-
tial curve (f(N) = Bo + B1eaN), where feN) describes fresh tuber yield (t ha- I ),
(a) (b)
34 Series 2 55
• -<)
~ ~
-
E 30
32 50 Newehu''''
Bridgenorth
Ol 45
~~oor
.paswell
'Qi
;: 28
~'
.s:::. 35
rJ)
~ 26
30
~ 24
.a
"iii 22
(5
25
(
•
I- 20 20
0 125 250 375 500 0 50 100 150 200 250
Fertilizer N (Kg ha- 1 ) Fertilizer P (kg ha- 1 )
Figure 4.10 The relationship between (a) applied N and tuber fresh weight yield
(drawn from data of Holliday et al., 1965), and (b) P fertilizer and tuber fresh
weight yield (from Boyd, 1970).
70
~ 60
40~------~------~------~----~
o 100 200 300 400
Figure 4.11 The average response curve of potatoes (cv. Bintje) to fertilizer N
estimated from 98 experiments carried out in The Netherlands between 1973 and
1982. (Redrawn from Neeteson, 1989a).
and Bo, Bl and a are coefficients estimated by non-linear analysis. The

average response curve is shown in Fig. 4.11.
Fitting functions to fertilizer responses provides a useful means of
summarizing data, but the calculated constants are usually unique to the
data from which they are derived and cannot be interpreted biologically.
For total harvestable dry matter yield, and allowing for loss of dry matter
through the senescence of leaves and stems, it would seem that the most
logical relationship between yield and N fertilizer would be an exponential
curve. This is based on the observation (Fig. 4.3 and the data reviewed by
Allen and Scott (1980» that there is a linear relationship between total dry
matter yield and IR and that IR is asymptotically related to the level of N
fertilizer applied (Fig. 4.4).
It is of course the effect of nutrients on the yield of tubers that is of
economic interest, and the interpretation of the effects, at least of N,
would appear to be dependent upon two processes, (a) the effects of the
nutrient on the interception of radiation and dry matter production, as
discussed for total dry matter yield, and (b) its effects on the distribution of
dry matter within the plant. This primarily takes the form of a delay, not
necessarily in the time of tuber initiation, but of a depression of the initial
tuber growth rate, and affects the relationship between IR and tuber dry
matter yield during the course of the growing season as shown in Fig. 4.5
for the cultivars Wilja and Cara. These cultivars may be considered to be
representative of the extreme effects of N on these relationships. Thus for
Wilja, there was a close linear relationship between IR and yield which
was only slightly affected by the level of N to the extent that rather more
light was intercepted before a significant yield of tubers had been obtained
(Fig. 4.5), while this was compensated by a slightly higher value of etuber.
Thus for this type of cultivar one would expect to obtain an asymptotic
response of tuber yield to the additions of N fertilizer, since this was the
form of the relationship between the addition of N fertilizer and IR (Fig.
4.4). For cultivars such as Cara, where etuber is profoundly affected by the
rate of fertilizer N applied (Fig. 4.5 and Table 4.8) the form of the response
curve could be expected to show a point of inversion under conditions
where the initial depression of etuber is not offset by its increase as the level
of IR increases, i.e. as the crop ages.
For a cultivar of the Wilja type, it might be predicted that under most
circumstances, tuber yield would be asymptotically related to the level of N
fertilizer, since the tendency for this cultivar to resist the deflection of
assimilate away from tuber growth is very strong. Thus, even if this cultivar
was harvested at an immature stage, then tuber yield would still tend to be
asymptotically related to N, except at a very early stage in its life cycle.
This is illustrated in Fig. 4.12a. On the other hand, for a cultivar of the
Cara type, characterized by a strong proclivity to invest assimilate in stems
and leaves, harvesting the crop before natural senescence permits the
transfer of dry matter from stems and leaves to tubers (Fig. 4.5) would give
rise to a response with a point of inversion and a marked decline in yield as
the level of N increased beyond the optimum. This is illustrated in Fig.
4.12b.
20 (a) (b)
14
18 o
16
~ 14 14
Wilja 0- 0 Cara
~ 13
~~
4
o
o 100 200 300 400 500 600 700 0 100 200 300 400 500 600 700
Figure 4.12 The relationship between N and total tuber dry matter yield at
intervals during the growing season for two contrasting cultivars: (a) Wilja and (b)
Cara (Harris, unpublished data).
The essential difference between these cultivars is possibly determined
by the differences in the size of their root systems (Table 4.9). The larger
root system, by reducing the resistance to flow of water from the soil to. the
leaf, would give rise to a substantial increase in leaf water potential. Since
leaf expansion is critically dependent on leaf water potential, then it is
possible to see the connection between the size of the root system and the
size of the crop canopy. The improved water status of the plant creates the
conditions which provide the necessary sink for assimilates to be used for
the production of leaves and supporting structures.
Table 4.9 Root length densities of two

contrasting cultivars grown on a sandy loam
soil (Simmonds, unpublished data)
Cultivar N (kg ha- J ) Root length density
(km m- 2)
Wilja o 8.57
250 11.90
Cara o 15.30
250 14.66
It would be interesting, and possibly useful, to extend this type of

analysis to the effects of other essential nutrients on growth and yield.
4.4.2 Soil nutrient status and fertilizer requirements

As discussed in Section 4.4.1, the fertilizer trial forms the only basis for
obtaining relationships between the additions of fertilizer and tuber yield.
It provides the information against which any other method of estimating
nutrient requirement must be assessed. The most commonly used aids for
adjusting fertilizer recommendations derived from mean response curves
are measurements of soil and/or plant nutrient content and soil type.
(a) Soil analysis

Assuming that the best possible estimate has been made of the probable
form of response to a particular nutrient, then, since the nutrient status of
the soil may vary considerably between years, due to the previous treat-
ment of the land, or to climatic conditions (Fig. 4.1), it is imperative that
the nutrient status of a site is estimated as accurately as possible in order to
determine the position of the site on the response curve at zero nutrient
input, since unless there is a linear relationship between fertilizer input and
yield, then the greater the nutrient status, the less will be the response per
unit of nutrient supplied. In the case of a 'bent stick' type of response, it
will determine how rapidly the transition point is approached.
The most obvious way of estimating soil nutrient status is to measure it
by some appropriate chemical analysis. In Britain, for example, the levels
of P, K and Mg recommended are modified by taking soil type and
chemical estimates of these elements which are adjudged to indicate their
availability for crop growth. One set of data relating 'available' soil P
and yield is shown in Fig. 4.13a and is derived from two long-term
rotational trials carried out in Britain. The yield response from a dressing
of 55 kg P ha- 1 on soils of differing P status is shown in Fig. 4.13b.
Rothamsted Saxmundham
~ 60
:t::-
j' f
1: -0
.~ 40
;;:
.s:::
Vl
Q)
~ 20
Oi
.0
2
(ij 0
(a) (5
I- 0 20 40 60 0 20 40 60
NaHC03 - soluble P (mg P kg·' soil)
t ha- 1
0
';" 12
.s:::eIl
Vl.s:::
~ a.. 10
~Cl
Q)::£
.cL() 8 0
2L()
.!: E 6
Q)
Vl-
e
eIl_ 0
Q).s::: 4 00
~Cl
0·-
c: Q)
- ;;: 2
0
o
.0
(b) 10 20 30 40 50
NaHC03 - soluble P (mg P kg-' soil)
Figure 4.13 (a) The relationship between tuber fresh weight yield and NaHC0 3
soluble P in soils at Rothamsted and Saxmundham. (b) The increases in yield from
a dressing of 55 kg P ha- 1 on soils of differIng P status. (From Mattingly and
Johnston, 1976).
Residues of P may be particularly long-lasting, and in an experiment

carried out on a poor nutrient-deficient moss peat/sand soil in Holland,
Boswijk (1976) showed how the use of town refuse applied to the soil until
1900, resulted in such a high level of soil P that there was virtually no effect
of P fertilizer on the tuber yield of intensively cropped potatoes after some
60 years without fertilizer P input. The experiment also showed that a high
phosphate status of the soil had no adverse effect on tuber yield. Tuber
yield was depressed if the level of K in the soil was too high.
The effect of soil P and K status on the response of tuber yield to the
additions of P and K fertilizers is also illustrated in Fig. 4.14 by data from
Birch et at. (1967).
(b) Tuber fresh weight

(a) Tuber fresh weight 4.48 0
:: 3.36 <70 ppm K2 0

jg <160 pm P20S 3.36
OJ 2.24 o
'"co 2.24
c.
~ 1.12 1.12
32
'iii
>-
o 45 67 90 90 179 269
K2 0 applied (kg ha· 1 )
Figure 4.14 (a) The effect of soil phosphate status (citric-soluble P20S) on the
response of potatoes to phosphate on 46 mineral soils. (b) The effect of soil potash
status (citric-soluble K 20) on the response of potatoes to potash on 42 mineral
soils. (Redrawn from Birch et at. 1967).
Chemical methods of analysing soil N status have not proved satisfactory

under British conditions, but Birch et ai. (1967) showed that the nitrogen
requirements of potatoes were influenced by the previous cropping and by
the amount of rain during the preceding winter (Fig. 4.15).
Tuber fresh weight

: : 4.48 Mainly cereal rotations
-
ctl
.r::
>305 mm
~3.36
.8 Arable rotations
3l2.24 Grass & mixed c > 3 0 5 mm
c
o rotations <305 mm
c.
~ 1.12 >305 mm
• <305 mm
L-~ __ ~~ I_ _- L_ _d-~
o 67 134 202 0 67 134 202 0 67 134 202

Fertilizer N (kg ha· 1 )
Figure 4.15 The effect of previous cropping and of winter rainfall (mm,
October-March) on the response of potatoes to N on 46 mineral soils. (Redrawn
from Birch et al. 1967).
The effect of previous cropping and management is currently used in
Britain to obtain a qualitative index of the likely contribution of soil N
reserves to crop growth. This is shown in Table 4.10 together with the
associated recommendations for N manuring for maincrop potatoes grown
on most mineral soils.
Table 4.10 (a) Soil nitrogen index, based on last crop grown, and
(b) recommended fertilizer N rates for second early and maincrop
potatoes grown on most mineral soils (MAFF, 1985)
(a)
N index 0 N index 1 N index 2
Cereals Beans Any crop
Forage crops removed Forage crops grazed receiving large
frequent amounts
of farmyard
manure or slurry
1-2 yearly leys cut 1-2 yearly leys grazed
high N
1-2 yearly leys low N Long leys, low N Long leys, high N
Maize Oilse.ed rape Lucerne
Permanent pasture, Peas Permanent pasture,
poor quality average
Sugar beet - tops Potatoes Permament pasture,
removed high N
Vegetables receiving Sugar beet tops
<200 kgN ha- 1 ploughed in
Vegetables receiving
>200 kgN ha- 1
(b) N kg ha- 1
220 160 100
In The Netherlands, N recommendations for arable crops are normally

based on numerous field trials. However on each experimental site, soil
mineral N is measured in early spring, and a range of fertilizer N doses are
applied to determine the optimum application rate from the response
curve. Recommendations for potatoes are derived from the linear relation-
ship between the amount of mineral N (ammonium N + nitrate N) present
in the soil in early spring and the economically optimum application rate of
fertilizer N. The larger the amount of soil mineral N, the lower the
recommended application rate of fertilizer N (Neeteson and Zwetsloot,
1989). However, while the linear relationship between soil mineral Nand
the optimum application rate of fertilizer N was significant, the variation
about the regression line was considerable, only 25% of the variation in
response to N being explained by soil nitrate content.
(b) Modelling
Neeteson et al. (1987) suggested that with simulation models it should be

possible to calculate, on a daily basis, the growth and the nitrogen uptake
of a crop and the nitrogen supply to a crop, using standard weather data
and field and crop parameters as inputs, thus adding the 'time' element to
N fertilizer advice as well as providing a better basis for it. The model was
used to predict the tuber yields obtained at seven N fertilizer levels in 61
field experiments, and to derive for each experiment an N fertilizer
response curve, from which an optimum N fertilizer application rate could
be deduced.
In 53% of the simulations the differences between measured and
simulated tuber dry weights at ·final harvest was less than 0.5 t ha- 1 and
in 76% less than 1 t ha- 1 . Since the measured tuber dry weight averaged
12.6 t ha- 1 , the deviations from the average yield were 4 and 8%
respectively, and therefore considered to be quite satisfactory.
Since weather conditions during the growing season cannot be taken into
account when the fertilizer is applied in the spring, the model was used to
predict yields using information available just before the time the fertilizer
was to be applied. This involved estimating the expected maximum tuber
dry weight, the moisture content of the soil at field capacity and standard
weather data derived from 30 years' weather records. It was assumed that
the rate of apparent N mineralization was 1 kg ha- 1 day-l at 12°C, the
maximum rooting depth was 60 cm and that growth ceased on 1 September.
Nitrogen fertilizer response curves were fitted to the predicted tuber dry
weights and optima calculated. The deviation of the predicted optimum
from the measured optimum was less than 50 kg N ha- 1 in 29% of the
experiments and 100 kg in 66%. The authors acknowledge that this is not
very satisfactory. However, since the confidence limits for measured
optimum rates are very large, this was considered to be an unsatisfactory
method of testing the accuracy of the model, which they then tested on the
basis of tuber dry weight deficits, that is the deviation from the tuber dry
weights obtained with the measured optimum application rate from those
predicted by the model in each individual experiment. In 68% of the
experiments the deficit was less than 1%. They concluded that the model
was reasonably sound and may be used as a tool to predict the optimum
application rate of N. However applying a fixed rate of 300 kg N ha- 1
almost always gave near maximum yields and the probability of getting
tuber dry weight deficits of less than 2% was 84%.
The model just described has been worked out for the cultivar Bintje
under Dutch conditions and, as described earlier, may not be applicable to
cultivars with appreciably different dates of maturity, and particularly for
cultivars such as Cara, where the effects of N on the distribution of dry
matter within the plant may have large effects on the form of the response
curve.
(c) Plant analysis
As an alternative to soil analysis, or to modelling the availability of N to

the plant from standard climatic data prior to the application of fertilizer
N, it may be both simpler and more practical to take advantage of the fact
that at least part of the fertilizer may be withheld at the time of planting,
without detriment to yield (Gunasena and Harris, 1968, 1969, 1971), and
that the nutritional status of the crop may be characterized by chemical
analysis.
The most common application of plant analysis depends upon the
detection of a critical nutrient content (Ulrich, 1952), or range (Dow and
Roberts, 1982), above which crops are not expected to respond to the
addition of specific nutrients, and below which the nutrient requirement
would be expected to bear a fairly close relationship to the decrement from
the critical level or band. It is widely appreciated (e.g. Mackay et al., 1966)
that establishing an optimum content or critical nutrient range is fraught
with difficulty because nutrient concentrations vary with plant part and
stage of development (Fig. 4.16), while optimum nutrient concentrations
may vary with the influence of factors other than the nutrient supply
(Smith, 1962).
The strict sampling procedures required and the time lag between
sampling and obtaining results may limit the practical utility of the
procedure, at least where action is required to rectify a nutrient deficiency
in the current season.
~7 0.7
N p
()
"0
ffi 6 0.6, g
z 2
:::J
'0 5 0.5 ~.
~E 4 0.4 5'
o
:::J
~ a.
"0 -<
.s: 3 0.3 3
c
o
~
g2 0.2 Q.
*
c
~ K
"U
c
8 1 0.1
o , , , , ! • , I , , I , , • , •
44 72 100 128 .44 72 100128 44 72 100 128

Days after planting
Figure 4.16 The N, K and P content of the dry matter of leaves (0), stems (ct)
and tubers (e) of the cv. Craig's Royal. (Drawn from data of Gunasena, 1969).
Since nutrient contents change with time, Prummel and Barnau-Sijthoff
(1984) in investigating the feasibility of using chemical plant analysis for
the early diagnosis of P and K deficiencies in potatoes in The Netherlands,
suggested that the problems of variation in critical levels associated with
the plant part sampled and the time of sampling might be overcome by
sampling the whole plant tops (leaves + stems) and· correcting the
concentrations to a standard N concentration based on the inverse relation-
ship between plant N concentration and plant age. Maximum tuber yields
were associated with leaf P above 0.5 and leaf K above 5% when the N
content was standardized at 5% of the dry matter of the tops. With each
1% decrease in N concentration the critical value for P decreased by about
0.1 % and K by 0.5%. Serious yield reduction could be found below 0.3% P
and 3% K at the standard N level. The estimation of crop age by N content
was obscured if the N supply was low or excessive, and the correction was
only valid if no abnormal conditions concerning N nutrition during growth
occurred. Rhue et al. (1986) in N.E. Florida found it advantageous to time
leaf sampling by reference to a temperature time scale. Methods of
standardizing nutrient levels with respect to stage of development would
seem to merit further research.
Remedial applications of the limiting nutrient must also be effectively
applied. Thus if applied in solid form, the nutrient must be applied in such
a way as to reach a moist layer of soil where active roots are present. This
may be impossible if the soil is dry, or the crop is too advanced to allow
nutrients to be incorporated by cultivations. In situations where the crop is
frequently irrigated, additional nutrients may be incorporated in the
irrigation water (e.g. Lauer, 1986). In other cases, nutrients may be
applied via a foliar spray, although because of the low concentrations
required to avoid damage, this may require a number of applications.
A more complex method of foliar analysis has been proposed by Beaufils
(1973), which recognizes that interactions among nutrients may cause shifts
in the optimum concentration of one nutrient from changes in concentra-
tions of others, and that the precise control of sampling data is difficult in
practice. The Diagnosis and Recommendation Integrated System (DRIS)
proposed by Beaufils, emphasizes the importance of balance among
nutrients rather than individual nutrient sufficiencies. The method requires
yield and leaf analysis data from a large number of sites, whi,ch can be
separated into desirable and undesirable populations on the basis of yields.
Nutrient ratios whose variances are significantly less in high- than in low-
yielding populations are considered to be important contributors to
optimum nutrient balance as required for maximum yields . Mean values of
these ratios are designated as standard 'norms' for assessment of locations
with unknown nutrient status. It has been claimed that the standard norms
developed for any given crop are applicable at any stage of development
and in any environment.
Meldal-Johnsen and Sumner (1980) published data for the application of
the DRIS method to the potato crop based on a survey of 745 commercial
fields in South Africa, while the only other published data appear to be
those of MacKay et at. (1987) in which nutrient ratio norms indicative of
high-yielding populations were derived from extensive data sets available
for two production areas in Canada with distinctly different soil and
climatic conditions. These norms were compared with derived and pub-
lished norms in diagnosing nutrient deficiencies in potatoes grown under
independent experimental conditions.
MacKay et ai. (1987) confirmed that the growth stage of sampling was
less crucial than found necessary for the critical nutrient level method.
Thus in one experiment, P deficiencies were diagnosed at three different
stages of growth, based on a common set of standard norms. On Boroll soil
in southern Alberta, deficiencies of Nand P were predicted reasonably
well, and on Spodosal soil sites in humid Nova Scotia, the indices correctly
predicted deficiencies in N, P and K, although N deficiencies were under-
estimated and deficiencies of P and K over emphasized. However, the
standard values differed from those developed in South Africa and
therefore cannot be universally applied. Furthermore, while deficiencies
were generally identified, the method could not predict accurately the yield
responses obtained nor the quantities of fertilizer nutrients required at
various sites in different years.
Conventional methods of sampling plant material and laboratory
methods of analysis tend to be slow and expensive, and these drawbacks
are not necessarily offset by an increase in sensitivity compared with rapid
tests on sap which can be carried out in the field. In these tests, sensitivity
may be obtained by measuring inorganic fractions such as NOrN, P0 4-P
and S04-S rather than total amounts of elements present (Table 4.11).
Table 4.11 Ratio of mean %N (dry

matter basis) in high N table beet
plants to that in control (zero N)
plants (Scaife and Bray, 1977)
Plant part Total N
Leaf blades 8.8 1.4
Petioles 16.0 2.1
These represent reserves of unassimilated nutrients and show large

fluctuations in response to changes in supply and demand. In contrast, total
amounts of elements include a substantial background of proteinaceous or
other structural matter of fairly constant composition. Analyzing petioles
or stems which contain relatively large proportions of these inorganic
fractions and little cytoplasmic material is an advantage over using leaves
and also improves sensitivity (Table 4.11).
It may be argued that tissue analysis is particularly suited to solving
problems of nitrogen fertilizer use, since at the time of planting it is difficult
to obtain reliable estimates of the amount of N likely to be mineralized and
available for crop uptake over the growing season. There is also much to be
said for withholding at least part of the nitrogen supply at the time of
planting in order to avoid leaching losses. In humid temperate climates,
soils are frequently close to field capacity at the time of planting, and
because of the relatively slow rate of canopy development due to low
spring temperatures, water loss is mainly by evaporation from the soil
surface, which once dry, substantially reduces the rate of evaporation from
the soil thus keeping the soil close to field capacity. Should there be
substantial rainfall during this period, drainage and therefore leaching of
nutrients such as Nand K may occur (Fig. 4.17).
110
, .. 90
-i
c:
0"
m
CD 90
I
~
70 ;
CD
"
I
Ol I 0--.1) CD
~
1968 en
:::I"
';"ca
(leaching) IJf " :E
.s::: 70 1/
50 ~.
~
I
t' ;:r
:E I -:::.
Cl I
0Qi
:: 50 I
30 ~
.s::: ,l 1969
en
d
I
(no leaching) .....
-=Cii
Q)
CD
10
30
.s
Ol
.0
::l
I-
10
Time in 14 day intervals
Figure 4.17 The effect of time of application of N on tuber yield in a year when
leaching occurred after planting (---) and when no leaching occurred (--).
Closed symbols = N applied part at planting and part after tuber initiation, open
symbols = N applied at planting time only. (Drawn from data of Gunasena, 1969
and Ngugi, 1972).
There is ample evidence that tuber growth and yield will respond to N
applied after planting (Gunasena and Harris, 1968, 1969, 1971). Further-
more it is common practice in some potato growing systems to apply part of
the nitrogen during the course of growth (Roberts et aZ., 1989). The major
attraction of applying only a part of the N in the seedbed is to reduce the
risk of leaching N into the drainage water, which is not only wasteful, but
will increase the possibility of exceeding limits set out for the nitrate
content of drinking water.
The commercial availability of a nitrate test strip has made the rapid
assessment of the nitrate content of sap expressed from stems or petioles a
practical possibility. Rapidity, simplicity and reliability are the essential
ingredients of a practical method for use in the field and all three have been
claimed for a variety of test crops. Papastylianou (1989) claimed that the N
fertilizer requirements of cereals could be diagnosed in situ in less than
30 s. Jemison and Fox (1988) found that using test strips and a hand-held
refiectometer gave results which were highly correlated with laboratory
results for both plant tissue nitrate (r = 0.87) and soil nitrate (r = 0.98).
They concluded that test strips provide a rapid, reasonably accurate and
precise method to determine nitrate concentrations in both soil and plant
material. van Loon et al. (1987) obtained a good correlation between leaf
petiole nitrate contents determined by Merckoquant test strips and by
laboratory methods (Fig. 4.18b). The petiole nitrate contents associated
with maximum yield levels are given in Fig. 4.18a. There was a slight, but
non-significant yield benefit from applying N fertilizer on the basis of
delaying part of the N fertilizer and top dressing on the basis of petiole
analysis.
9000 (a)
7000
E ~
~ 5000 "-
C') "-
~
0
z 3000
1000
30 50 70
Days after emergence
45 (b)
000
:; 35 COO
0
Cl
-"" o
.9 30
C')
0
z 25
I 20 o
z o
VJ 15
a.
.;::
1ii 10
o 5 10 15 20 25 30 35
N - N03 (g kg'1)
Laboratory analyses
Figure 4.18 (a) Petiole nitrate contents with maximum yield levels: --,1984; ---,
1985. (b) Relation between leaf petiole nitrate contents as determined by Merckoquant
test strips and by laboratory methods. (Redrawn from van Loon et al. 1987).
The method is least likely to be of use for a nutrient such as P which can
be built up in the soil without risk of leaching and which is likely to be
needed in the very early stages when, because of the lack of mobility of
the element and because of the limited exploration of the soil by roots in
the early stages of growth, a readily available supply of P is needed in the
vicinity of the plant. However, MacKay et al. (1988) have demonstrated
that P deficiency detected at the early bloom stage by leaf analysis can be
corrected by the immedate application of fertilizer P, and yields increased,
provided there is adequate soil water.
4.4.3 Economic considerations

On the basis of 99 fertilizer trials with potatoes carried out at various
locations in The Netherlands between 1973 and 1982 Neeteson and
Wadman (1987) calculated the optimum economic application rate of N on
the basis of the best-fitting model for each trial. The optima covered the
entire range (0--400 kg N ha- 1 ) of the fertilizer levels tested. Disappoint-
ingly, the magnitude of the confidence intervals (p>95%) was higher than
300 kg N ha- 1 in 60% of the potato trials.
The confidence intervals for optimum N application rates were wide
when differences among replicates were large andlor the curves were flat or
did not decline significantly beyond their maximum. The former obviously
depends on the homogeneity of the experimental site, while flat response
curves can be expected where crops are not protected from diseases, under
extreme weather conditions, such as drought in the early stages of growth,
or where heavy rain may occur after fertilizer application. However
Neeteson and Wadman point out that an important reason why the potato
has a higher proportion of very wide confidence intervals for the optimum
N-fertilizer application rate as compared with sugar beet, which also
featured in their trials, was that the N-response curves of the potato crop
generally do not decline beyond their maximum, and that applying too
much fertilizer N to potato seldom had a distinctly negative effect on tuber
yield.
The implications of a response curve without a point of inversion for the
profitable use of fertilizer by the farmer are considerable. This can be
illustrated by data taken from the response of Wilja to nitrogen fertilizer
discussed in Section 4.3.
The yield of tubers >40 mm obtained with fertilizer application over
the range 0-750 kg ha- 1 was fitted with an inverse polynomial of the form
Y = a + lIb + d + N, where Y = yield of tubers> 40 mm, a, band dare
constants with values of 66.11, - 2.06 and 0.008 37 respectively. In Fig.
4.19, the returns from the yield increase due to fertilizer, the cost of the
fertilizer and the margin over fertilizer cost are plotted against fertilizer
rate. It is assumed that the price of the potatoes is £70 per tonne and the
cost of the nitrogen £0.3333 per kg. The salient characteristic of this type of
£ ha- 1
1800 Return
1600
1400 Margin over cost of N
1200
1000
800
600
400
200
03070 150 340 750

Figure 4.19 The effect of N fertilizer on the returns and 'profit' when applied to
the cv. Wilja in 1987 (Harris, unpublished data).
response curve is that the returns to the addition of fertilizer rise steeply
with the first increments, but that over a wide range of fertilizer use, the
margin over fertilizer costs vary little. Thus in the example given, within an
application rate of 300 to 750 kg N ha- 1 the margin over the cost of N varied
by only 7.6% of the maximum profit. Thus not only is it impossible to
determine with any acceptable degree of precision the amount of fertilizer
to apply with a response curve of this type, but as far as the grower's
returns are concerned this lack of precis,ion is of little consequence
financially.
Since however under certain circumstances the response curve may show
a point of inversion (Section 4.3, and Neeteson and Wadman, 1987), and
since there is considerable pressure both from environmental and health
concerns to restrict the amount of fertilizer N applied, then it would seem
to be a sensible policy to apply N fertilizers conservatively. The possible
effects of cultivar on the shape of the response curve needs also to be taken
into account, since while comprehensive fertilizer trials are usually carried
out with the most widely grown cultivar (e.g. Bintje in the case of the
experiments in The Netherlands) new cultivars are introduced, and may
respond in different ways as discussed in Section 4.3.
Using the same experimental data Neeteson (1989) calculated how far
fertilizer N could be reduced before serious yield deficits occur. Table 4.12
shows the effect of applying various proportions of the recommended
application rates. Thus cutting the fertilizer N applied by 50%, only gave
rise on average, to.a 4.2% drop in yield. On the basis that a 2% reduction
in yield is acceptable, it was estimated that fertilizer N could be reduced by
32%; however the chance of serious yield deficits was then about 11%.
Neeteson (1989b) assumed that serious yield depressions should not
exceed 5% in which case the optimal rate of N could be reduced by 27%,
and this would give rise to an average yield deficit of only 1.7%.
Table 4.12 Yields of potatoes for various reductions in the

recommended fertilizer N rates. Average values of 98 potato
trials (Neeteson, J989b).
Fertilizer N Tuber fresh weight Yield deficit
(% recommended rate) (t ha- 1) (t ha- 1)
100 56.8 0
90 56.5 -0.3
75 55.9 -0.. 9
50 54.4 -2.4
0 45.1 -11.7
The effect of increasing the price ratio (cost of 1 kg N fertilizer N to the

price of 1 t potatoes) was to reduce the optimal level of N application by
50%.
4.5 SOME FACTORS MODIFYING THE YIELD RESPONSE TO

FERTILIZERS
4.5.1 Organic manures

A wide range of organic manures may be applied to crops, ranging from
materials such as hoof and horn meal, which have a high N content and
behave in a similar way to inorganic fertilizers, to bulky organic manures
resulting from a mixture of dung, urine and bedding or, more commonly,
to slurries of dung, urine and water, where animals are housed without the
possibility of bedding material becoming mixed with excreta. Here atten-
tion is focused on 'farmyard manure', the traditional bulky organic manure
resulting from cattle over-wintered in yards bedded with straw. Such
manures have been held in high esteem by farmers, and in Britain, when
the material is available, the potato crop received priority. However, while
Some factors modifying the yield response to fertilizers 195
the importance of such materials has declined in Britain, with the increased
use of inorganic fertilizers and the changes in animal husbandry, they may
still be of considerable significance elsewhere. According to Sokolov and
Orlovskii (1971), organic fertilizers, of which farmyard manure is the
principal one, play an 'extraordinary role' in the cultivation of the tundra
and taiga virgin soils in the USSR, not only as a source of nutrients, but
also in improving the physical and chemical properties of soil. In temperate
regions, the material is of considerable importance to the small, but
expanding group of farmers practising 'organic' farming methods. It is
widely believed that the benefits of bulky organic manures are derived
from their physical rather than from their nutritional properties.
Crowther and Yates (1941) showed that the responses of crops to
fertilizer N, P and K were considerably reduced when grown in the
presence of 25 t ha- 1 of farmyard manure; however, they pointed out that
the most remarkable feature concerning the effect of farmyard manure was
the small difference between the responses to N in the absence of farmyard
manure and its presence. Boyd (1959) reviewed the evidence and con-
cluded that the response to N in the absence of farmyard manure was very
dependent on the basal supply of P and K; in the presence of farmyard
manure it is less dependent. Boyd concluded that the effect of farmyard
manure could be adequately accounted for in terms of the nutrients N, P
and K without having recourse to any other effects. The value of 10 t of
the material was estimated to be equivalent to about 15 kg N, 9 kg K and
31 kg P.
Some experiments in which the level of N, P and K supplied as inorganic
fertilizer was increased in the presence and absence of farmyard manure
have shown that, given a sufficiently high level of fertilizer, the response to
farmyard manure could be eliminated (Table 4.13). However in a series of
ten experiments carried out on the Leeds University farm in one of the
drier parts of England (long term mean rainfall 610 mm per annum) it was
shown that the effects of farmyard manure could not always be eliminated
by high levels of inorganic fertilizer (Holliday et al., 1965). In all experi-
ments, treatments consisting of 0 and 25 t ha- 1 of farmyard manure were
factorially combined with 0, 79 and 158 kg N ha- 1 . A uniform rate of P and
K was applied to all treatments
The responses to N in the presence and absence of farmyard manure are
shown in Fig. 4.20a for the means of the five wettest and five driest years .
In the five wettest years the two N response curves converged, indicating
that the effect of farmyard manure could be eliminated by inorganic
fertilizers. However, in the five driest years, the curves were approximately
parallel, indicating that farmyard manure was having an effect which could
not be reproduced by inorganic fertilizers. Internal evidence suggested that
there was no response to levels of N in excess of 158 kg N ha- 1 • Given the
high level of P and K basal to all treatments any response to farmyard
manure at this high level of N could be considered to be due to effects
Table 4.13 Total tuber yield
(t ha· l ) response to 37.5 t ha- l of
farmyard manure at various levels of
N, P and K supplied by fertilizers
(Reith and Inkson, 1958)
N P K Tubers
(t ha- 1)
kg ha- 1
0 0 0 7.0
34 20 47 1.2
68 39 93 0.5
which are not readily explained in terms of quantity of N, P and K supplied

by the organic manure. The term 'extra effect' was coined to describe this
effect and values plotted against the maximum calculated potential soil
moisture deficit occurring in each of the 10 years (Fig. 4.20b) show a
positive relationship between the extra effect of farmyard manure and
increasing dry soil moisture conditions.
Pot experiments (Harris, 1960) suggested that the beneficial effect of
farmyard manure in dry years could not be attributed to any significant
increase in available soil moisture, for the equivalent of 25 t ha- 1 of
farmyard manure in pots increased available soil moisture by only 2 mm.
35 (a)
+FYM r---
+FYM
~
30
-FYM /
25 mean, 5 wettest~n, 5 'd;i~st
years years
~ 20
.~ o 79 158 0 79 158
.r: Fertilizer N (kg ha- 1 )
&10.0
CD
.c
(b) •
.a 7.5
~ 5.0
•
2.5
•
o I-I-,.L..,..----
o 100 200 300
Maximum potential soil water deficit (mm)
Figure 4.20 (a) The effect of N fertilizer with and without farmyard manure on
tuber yields in wet and dry seasons. (b) The effect of soil water conditions on the
response of potatoes to farmyard manure at levels of N, P and K believed to be
sufficient for maximum yield. (From Holliday et at. 1965).
In these experiments the manure was applied in the bottom of the
ridges and the inorganic fertilizers then broadcast before the tubers were
planted and the ridges split to cover the seed. The explanation advanced
for the beneficial effect of the manure in dry seasons, was that nodules of
farmyard manure in the soil were capable of retaining sufficient moisture to
permit roots to absorb N and other nutrients. If correct, it would be
expected that the extra effect of farmyard manure could be reproduced by
inorganic nutrients equivalent to those available in the former, provided
some means of making them more available under dry conditions could be
devised.
Holliday (1970) summarized experiments which tested the above
hypothesis. Two methods were attempted: the first, a foliar application of
N applied at pre-determined soil moisture deficits, proved to be unsuccess-
ful in the one exceptionally dry year in which it was tested. The second
method adopted was to place nutrients at a greater depth into the soil
profile where the loss of moisture is less rapid. This was achieved by
applying liquid fertilizer 15-18 cm below the seed tuber in two bands, each
20 cm from the centre of the row at the time of planting. The deep-placed
fertilizer largely simulated the effects of farmyard manure. The result was
in agreement with an experiment reported by Warren and Johnson (1962)
in which fertilizers which were dug in substantially reduced hitherto
unexplained yield advantages found when farmyard manure was applied to
sugar beet. Further experiments (Holliday and Draycott, 1968), showed
that advantages accrued from deep placed fertilizer (18 cm) under a dry
soil moisture regime but not under a wet one. Garwood and Williams
(1967) demonstrated similar benefits from the deep application of N to a
grass sward in periods without rain.
Modern methods of housing livestock ensure that slurries of organic
manures are more likely to be available than traditional farmyard manure.
In The Netherlands in the series of trials already referred to in Section 4.4
(Neeteson and Zwetsloot, 1989), organic manures were applied to 50
trials. The manures were applied in the autumn or early winter preceding
the potato crop and consisted of green manures (Lolium multiftorum
Lamk. or Vicia sativa) and/or slurries (cattle, pig or poultry at 50 t ha- 1).
The optimum amount of N was reduced by 15-50 kg N ha- 1 in the presence
of the organic manures.
4.5.2 Fertilizer placement

The most efficient placement of fertilizer is that which provides for an
adequate supply of soluble nutrients in a well-aerated zone of moist soil
occupied by actively absorbing roots at periods of growth when the
demands for plant nutrients are most acute.
Batey and Boyd (1967) have reviewed experiments in which various
methods of placement have been compared. They pointed out that
considerable attention has been given to the placement of fertilizer for the
potato crop because the efficiency of fertilizer use is partly determined by
the ultimate position of the fertilizer in the ridge, which in tum depends
upon how and when the fertilizers are applied and the planting method.
The placement methods they described, which are applicable to crops
grown in single row ridges, include the following systems.
The fertilizer may be broadcast 'on the flat', i.e. before the land has been
ridged and the crop planted. The earlier the application relative to planting
and the 'rougher' the soil conditions, the greater the depth of incorporation
of the fertilizer into the final seed bed, and subsequently through the ridge.
Fertilizer applied in this way is more likely to be localized around and
above the seed tuber. A traditional method of application is to broadcast
the fertilizer over ridged land before planting. On splitting the ridges, the
fertilizer is concentrated mainly just below the tuber. Various attachments
to planting machines are available which place the fertilizer in various
zones in the ridge. With contact placement much of the fertilizer is located
in contact with the tuber or developing sprout, whereas with band
placement it is separated from the fertilizer by a zone of soil; the width of
this buffer between fertilizer and tuber and its orientation may be varied
according to the design of the placement mechanism. It is also possible to
inject fertilizer solutions or anhydrous ammonia into the ridge after
planting.
It has been shown that the form of the yield response to fertilizer can be
altered by the method of placement. The response curve may differ for
fertilizer broadcast on the flat and broadcast over the ridges; the degree of
curvature appears to be more pronounced for placed than for broadcast
fertilizer. Thus small and moderate amounts of fertilizer tend to be utilized
more efficiently when placed than when broadcast, and less placed
fertilizer is needed to reach maximum yield. Where conditions are
unfavourable to placement, yield may decrease sharply at high levels of
fertilizer application, for example on sandy soils deficient in Nand K in low
rainfall areas; here the high level of fertilizer required can, in conjunction
with low soil moisture content, damage germinating plants and delay
emergence. Working the fertilizer well into the seed-bed would appear to
be the best method at high application rates.
It was reported in the previous section that placing fertilizer at some
depth below the seed tuber has been shown to be benefi;cial where the
drying out of the ridge or surface soil layers may limit the uptake of
fertilizer located there. However, where crops are irrigated, deep place-
ment would not be expected to be benefi.cial, and could well result in a
greater risk of leaching Nand K below the depth of rooting.
4.5.3 Timing fertilizer application
The usual practice is to apply most or all of the fertilizer at or just before
the time of planting. However Radley (1963) suggested that under good
environmental conditions, e.g. a plentiful supply of plant nutrients,
assimilates tend to be used in the production of foliage rather than tubers
and that this accounts for the apparent delay in tuber initiation associated
with the use of fertilizers applied in the seed-bed. Ivins (1963) suggested
that it might be possible to increase the duration of tuber bulking without
sacrificing bulking rate by withholding N, or part of it, until tubers had
been initiated, and in this way increase final tuber yield.
Experiments testing this hypothesis have been reported on a sandy loam
soil in the south of England (Gunasena and Harris, 1968, 1969, 1971).
Large benefits in delaying the application of all or part of the N fertilizer
were associated with seasons when rainfall after the application of all or
part of the N in the seed-bed was sufficiently high to bring about leaching.
The beneficial effects of delaying the application of both Nand K fertilizers
were associated with increased recoveries of these nutrients in the crop
attributed to the avoidance of leaching losses. In subsequent experiments
on the same site where leaching did not occur, delaying the application of
all or part of the fertilizer until after tuber initiation did not result in any
marked effects on the response to N (Ngugi, 1972). There was little
evidence to suggest that delaying the application of N had any marked
effect on the period between the apparent initiation of tubers and the
establishment of a high bulking rate.
In view of the concern over the leaching of nitrate from arable soBs,
there is justification for withholding part of the N supply until the time of
tuber initiation, although for later applied N to be effective could depend
upon the provision of irrigation. As discussed in Section 4.4.2, when it is
difficult to predict how much N may be mineralized from the soil organic
matter, it could be sound policy to apply a conservative dressing in the
seed-bed until later evidence suggests that the level should be increased.
There is little evidence to suggest that the application of N at later stages
of growth, with or without the use of growth regulators, has any significant
benefits on yield (Holliday, 1970; Kerketta, 1976). On the other hand
Roberts et al. (1989) found that the lowest tuber yields resulted where a
relatively high and uniform level of petiole NOrN was maintained during
the growing season.
The use of slow release forms of N, such as nitroform or sulphur-coated
urea, or organic forms such as hoof and hom, might be considered to
achieve some of the benefits of delaying the application of fertilizer N.
However such forms have compared unfavourably with quicker acting
sources of N. Lorenz et al. (1972) showed that slow-release N sources did
not increase yield or N absorption during the latter stages of growth and
Roberts et al. (1984) found no yield advantages from the use of slow
release N from methylene urea or sulphur-coated urea· compared with
ammonium nitrate and urea on sandy soils in central Washington.
4.5.4 Rainfall and irrigation during the growing season

Rainfall may interfere with the uptake of plant nutrients in several ways:
for example through leaching of soluble nutrients below rooting depth or
through limiting uptake because of dry soil conditions. Where the top soil
is kept moist by frequent rainfall, it may be expected that more nitrogen
would be mineralized from the soil organic matter, while lack of rain may
limit the response of a crop to nutrients which have already been taken up
by the plant. Rainfall is also an important factor affecting the incidence of
disease, particularly such important diseases as Phytophthora infestans L.
The relationship between rainfall during the growing season and crop
response to fertilizer may therefore be expected to be complex.
(a)
180
Normally treated
arable soil
After
ploughing
a ley
80 180 240
Total rainfall: JunEkluly (mm}
(b)
';"I\l
.c
2.5
•
~
Ol~
• •
-'£ ';"I\l
~.c
C')~
2.0
••
• • • •• •
o (/)
; Q; 1.5
0).0
(/) ::J
c-
•
o
••
c.
(/)
0) 1.0
a:
40 50 60
Number of rainless days May-June
Figure 4.21 (a) The relation between June-July rainfall and the optimal
application rate of N for potatoes. (From Kuipers, 1962.) (b) The relation between
the number of rainless days and the response of potato tubers to 398 kg K ha- I .
(From van der Paauw, 1958).
The response of potatoes (cultivar King Edward) over a period of 10
years on glacial drift soils overlying magnesium limestone in central
England (Holliday et ai., 1965), varied from 11.7 t ha- 1 when the maximum
calculated soil moisture deficit was less than 25 mm to 2.7 t ha- 1 when it was
250 mm.
While the response to N would appear to be greater in wet than in dry
years, the optimal rate of application has been shown (Kuipers, 1962) to
increase with a decrease in the June-July rainfall (Fig. 4.21a).
Similarly Evans (1974) reported that with the early cultivar Home Guard
harvested before reaching maturity, where lack of moisture did not affect
growth, the optimum level of N was less than at non-irrigated sites in dry
seasons.
van der Paauw (1958) showed that potash fertilizers had the biggest
effect on yield in dry years (Fig. 4.21b) which was attributed to a reduction
in the availability of soil K. Hackett (1968) has shown with barley, that
when K is deficient, root growth and especially the development of
laterals, may be markedly retarded, which could affect the ability of the
crop to remove water from the soil. Potassium may also influence the water
40 r (a)
30~ 60 (b) 50mm~
20
40 r ~
v
(5/0
88 mm
E
"f 10
20
2:-
, , I I ! , I I
"0 o 50 200 ha- 1 0 200 0 200

N kg
~ 60 (c)
100mm~o-~
"'- .~mm 50 68mm (y)
~ 40 ~ 30
E Sandy loam
0>20
"iii Sandy clay 10
== loam
..c: ,~
~ 02000200
'i-§ 10 (e) .dw t haV-1
.---- 100 mm
- 8 • ~
19 cr- 100 mm
~ 6-
4 -
20 -
2
o '----' L--------l o ~~--I
o
0--,
o 150 o 150 200
Figure 4.22 The effect of fertilizer on tuber yield in the presence and absence of
irrigation. Figs a-f horizontal axis = N kg ha- 1 • (a) Price and Harvey (1962); (b)
Wellings (1972); (c) and (d) Clutterbuck and Simpson (1978); (e) Asfary (1981);
(f) Gerdes et at. (1975).
economy of crops through its influence on stomatal movements (Peaslee
and Moss, 1966).
A problem related to the effect of summer rainfall on the response to
and optimal application rate of fertilizers is whether there is a need to
modify the application rate of fertilizers for irrigated crops in comparison
to practice on unirrigated crops. This is an important problem in areas,
such as Britain, where only certain valuable crops, such as potatoes, are
considered economically worth irrigating. However the proportion of
the crop irrigated in Britain has increased considerably. Harris (1985)
reviewed the available evidence and figures showing the response to
fertilizers in the presence and absence of irrigation are shown in Fig. 4.22.
It was concluded that irrigation will improve the availability of P and K
and might well reduce the quantity needing to be supplied as fertilizer.
Rather more information was available for N, but while the response to N
may be enhanced by irrigation, there was little evidence to suggest that the
maximum rates needed to be increased. More recent data in which the
interactions between cultivar, nitrogen and irrigation have been measured
have become available from two experiments carried out on a sandy loam
100 (a) (b)
80
~
60
';"tt!
.c
~
40
E
Ol
'w 20 CAR A
3:
.c
F:
en
r
~ 0
CD
.0
.2 ~
...
(ij 60 -0
(5
f-
40
20 WILJA
1986 1987
0
0 200 400 600 800 0 200 400 600 800
Figure 4.23 The effect of N on the tuber fresh weight yields of contrasting potato
cultivars (D = Cara, 0 = Wilja) in the presence (closed symbols) and absence (open
symbols) of irrigation in (a) 1986 and (b) 1987 (Harris, unpublished data).
soil in southern England (Harris, unpublished: see Section 4.3). The
effects of the treatments on final total tuber yield are shown in Fig. 4.23.
There was no significant interaction between irrigation and nitrogen in
either year or for either cultivar.
Winter rainfall
Large fertilizer dressings leave residues of nitrate in the soil and more is
released as crop residues decay. Reserves of nitrate are easily leached from
the topsoil and the residue of nitrate left to benefit a following crop
depends on the amount of winter rainfall. Birch et al. (1967) pointed out
that there are two types of variation in winter rainfall: the inherent regional
variations in climate and the year-to-year variations in weather conditions.
Allowance for the first can take the form of regional recommendation,
although there is likely to be a degree of association between rainfall
and cropping system which affects the organic matter status of the soil
and N requirements (see Fig. 4.16). Allowance for seasonal variation
requires assessment and decision in late winter. Where winter rainfall
(October-March) exceeded 300 mm, about 30--40 kg more N ha- 1 was
required for a potato crop grown when winter rainfall was less than
300 mm. Similarly in The Netherlands, it has been shown (van der Paauw,
1963) that 40 kg ha- 1 more soil N is available in early spring after a dry
winter than after a wet one. The measurement of N0 3 -N in the soil profile
would take the effects of winter rainfall into account and its use was
discussed in Section 4.4.2.
4.5.5 Plant density

It has been pointed out (van Burg, 1967) that most experiments on the
effect of plant density have been conducted at one rate of fertilizer
application, but that where the interaction between fertilizer and plant
density had been investigated, some had shown that an increase in the level
of N or of N, P and K had a greater effect at high than at low crop densities,
while in other instances no interaction had been found.
In the rather special circumstances of seed-potato production in Holland
where haulms were destroyed in mid-July, and using seed 40-50 mm
diameter (van Burg, 1967), the response to N increased with an increase in
plant density from 40 000 to 50 000 plants ha- 1 , and the response to N was
brought forward in time by close planting. In a second experiment where
plant density was varied by seed size as well as spacing, the interaction was
marked only with small seed (35-40 mm) and was absent with larger tubers
(50-55 mm). There was an overall positive interaction between numbers of
stems and N supply.
In a further experiment with a maincrop cultivar, where the foliage was
allowed to die naturally, Fig. 4.24a shows that with respect to final tuber
Figure 4.24 (a) The effect of plant density and N fertilizer on tuber dry matter
yield. (b) The effect of N and plant density on haulm senescence: 10 = no die back,
1 = haulms completely dead. (From van Burg, 1967).
yield, the response to N was dependent on plant density. At high plant

densities, extra N seemed to permit the investment of more dry matter in
leaf production which, given a long enough growing season, eventually
increased tuber yields. van Burg was able to show that N delayed the
senescence of foliage, an effect particularly marked at the highest plant
density, which treatment otherwise tended to advance the senescence of
the crop (Fig. 4.25b).
Bodlaender and Reestman (1968) found that with the late cultivar
Alpha, the optimal plant density was increased with 100 kg N ha- 1
compared with no fertilizer N, but that a further increase in N supply did
not change the optimal plant density. Increasing the N supply was most
effective at high plant densities, especially after a long growing period.
Favourable interactions between N and plant density were not usually
obtained at early lifting, and this would apply to later cultivars where the
growing season was prematurely curtailed.
With a second early cultivar (Maris Page), Ngugi (1972) failed to detect
any significant interaction between N and plant density, although the
optimal tuber density was 46000 ha- 1 with 200 kg N ha- 1 ; with 300 kg N ha- 1
the highest yield was achieved with a density of 65 000 tubers ha- 1 • Ifenkwe
and Allen (1983) reported that increasing plant density to levels above
those used in commercial practice increased nutrient (N, K) accumulation,
but had only small effects on yields. They pointed out that while high yields
may be associated with large accumulations of nutrients, they are not
necessarily caused by them.
Since positive effects of plant density on leaf area index (L) tended to be
confined to the early stages of growth and gave way to negative effects as
Fertilizer use and potato quality 205
the canopy senesced, and as N tended to delay the senescence of foliage
(van Burg, 1967), then it would seem that for maximum yields, a positive
interaction between N and plant density might be expected. However,
since in temperate regions the amount of incident radiation is declining
rapidly at the end of the growing season, this would tend to reduce the
importance of effects of N on delaying senescence at high populations.
4.5.6 Soil pH
Apart from the effect of soil pH on soil-borne diseases such as potato scab
(Streptomyces scabies), the major effect on yield is probably associated
with the availability and uptake of plant nutrients. Bolton (1971) found in
liming experiments carried out on the contrasting soil types at Rothamsted
and Woburn in England, that yields of potatoes were similar at all pH
values above 5 when P and K were sufficient, but when K was not applied,
the more acid soils grew the larger crops. In both experiments the response
to K was least on the most acid plots because liming decreased yields when
K was not given. This lime-induced K deficiency was explained by liming
increasing the proportion of K adsorbed on the exchange complex, and
thus decreasing the concentration of K in the soil solution; K ions can
displace adsorbed Ca from cation exchange sites on the soil more easily
than Al which is the predominant exchangeable ion in acid soils. At
Woburn, but not at Rothamsted, the response to P was large on the
unlimed plots.
That the potato crop can tolerate very low pH values was demonstrated
in experiments on a sphagnum peat soil (Maclean et al., 1967), where 6.7 t
limestone ha- 1 increased yield in the year of application on a soil with a pH
of 3.7, but where in three subsequent years, yields were as satisfactory on
the unlimed plots.
4.6 FERTILIZER USE AND POTATO QUALITY
There is a voluminous literature on the relationship between fertilizer use

and potato quality and there are useful reviews by Perrenoud (1983),
Lisinska and Leszczynski (1989), and Wolfe (1987). See also Chapter 12.
Quality in potatoes depends upon the purpose for which the crop is
intended, which can vary considerably, from the provision of potatoes for
home cooking to the perhaps more exacting standards required by crops
used for processing into products such as crisps (USA chips) or French fries
(UK chips). Quality characters are perhaps primarily associated with
genotype, but there is no doubt that the environment, and especially the
nutritional environment experienced by a crop, can have important
consequences for quality.
Perhaps the simplest components of quality to appreciate (but not
40 (a) (b) 1.110 (c) -K
•
~
r
30
35 • •
30
•
~ ~ 1.100
.s;
.- , ....+K-.. " •
• 20 ~
Ol
~ 25
()
~
'0 -K
Q)
q ~.
• c% 1.090 \"(i>
\ ~
I:
dm % 10 \+K
A A A A 30 \
! to. A to. , ):)--<>--0 0- -.0- -0---0 \

b.. """0
A /20 0
5 0 0 0 0 o 10 1.080
, I I I
0 2 3 o 801602400 80160240 o 80160240

Compound fertilizer (t ha· 1 ) Kg N ha· 1 kg K ha- 1 kg N ha· 1
Figure 4.25 (a) The effect of a complete N:P:K fertilizer on the fresh weight yield
of large (e) and small (0) tubers and on the dry matter yield (.) and content of
large (.6.) tubers. (From Rodger and Robertson, 1970.) (b) The effect of Nand K
on the dry matter yield (0) and yield of water (e) of the cv. Aristo. (c) The effect
ofN and K (nil and 166 kg K ha- 1) on the specific gravity of the two potato cultivars..
(b and c from Schippers, 1968).
necessarily to understand) are those associated with size, shape and

external appearance such as colour and superficial blemishes.
Through the ability of fertilizers to influence tuber number and yield,
fertilizers can have a profound effect upon tuber size. In general fertilizers
will tend to increase tuber size. Thus in Fig. 4.25a it is clear that the use of
increasing quantities of a complete fertilizer increased the yield of large
tubers while the yield of small tubers remained fairly constant. In most of
the experiments reviewed by Perrenoud (1983), Nand K increased the
yield of large tubers, while P tended to increase tuber number and have
rather less effect on tuber size.
Tuber shape is largely under genetic control, but may be influenced by
the physical characteristics of the soil and the extent and timing of water
stress (see Chapter 5). Fertilizers would seem to have little influence on
tuber shape. Since however the external appearance of the crop is affected
by the amount of damage experienced, it is perhaps important to note that
mechanical damage to tubers has been shown to be decreased on crops well
supplied with K. There is also some evidence that an important cause of
superficial blemishes, Streptomyces scabies, may be affected by fertilizers,
but the effects are inconsistent and may be associated with changes in soil
pH.
Perhaps the most important internal quality factors, particularly for
potatoes intended for processing by frying, are dry matter (DM) and
reducing sugar content. DM content tends to be decreased by Nand K
Conclusions 207
fertilizers; consequently DM yield is usually less responsive than fresh
weight (FW) yield for Nand K (Fig. 4.26b). For processing into aips or
French fries, high dry DM contents, or high specific gravities, are pre·
ferred, since this improves the economics of the frying process. Nitrogen
and K are particularly important in this respect. Perrenoud (1983) for
example found that with the exception of only one report, all the other
papers reviewed showed that N decreased DM content. Similarly K
generally reduced DM content, although its effect was less marked than
that of N. The effect of N, K and cultivar on tuber specific gravity was
investigated by Schippers (1968). He reported that there may be an
interaction between these nutrients, in that the effect of N was curvilinear
at low levels of K and approached linearity at high levels. The effect of N
was also dependent on cultivar (Fig. 4.26c). The effects of fertilizer on
reducing sugar content and on the production of undesirable colours after
frying appear to be rather small and probably unimportant (Hughes, 1986).
Another important internal quality characteristic of potato tubers is
internal blackening, a character which seems to be promoted by increasing
N fertilizer and reduced by K (Perrenoud, 1983).
There has been a mounting interest in the quantity of nitrates consumed,
primarily brought about by a concern for the nitrate content of drinking
water, but also some concern with the nitrate content of foods. Carter
and Bosma (1974) showed that N fertilizer applied at rates between 0 and
720 kg N ha- 1 increased the tuber nitrate concentration from values of
25-36 ppm (wet weight basis) without N fertilizer to 50-131 ppm at the
highest rates. Concentrations depended on moisture level and year, and
also on the type of fertilizer, being reduced where slow release forms
(e.g. ureaform) were applied. The nitrate level was related to the nitrate
level in leaves sampled at the near full-bloom stage. Greater concentra-
tions of nitrate were located just below the skin, and removing the peel
(about 5% of the tuber) reduced the nitrate content of the whole tuber by
12%. However, the authors point out that about 80% of the nitrate content
of tubers is removed if potatoes are boiled and the water drained. Augustin
et al. (1977) concluded that nitrate levels in potato tubers are not normally
a problem in N. W. Pacific states of the USA, even with the excessive rates
of N often used by growers.
Schippers et al. (1969) concluded that the amount of fertilizer applied
to the seed potato crop had little effect on the productivity of the progeny.
4.7 CONCLUSIONS
Quite large quantities of the major nutrients are taken up by the potato
crop during the course of growth, and there can be few sites where
potatoes are grown where yield responses to at least one of the three major
nutrients are not obtained. In many soils where potatoes are grown in
temperate capital-intensive farming systems, it is likely that the most
immediately limiting nutrient is N.
The relationship between the input of the major nutrients and tuber
yield is most likely to be asymptotic, although examples of relationships
with a point of inversion are not infrequently found with N and possibly
with K, but probably rarely with P. At least for N, an asymptotic
relationship between total dry matter yield and the rate of N fertilizer
would be expected from the similar relationships between N supply and the
total amount of radiation intercepted by the crop. and the linear relation-
ship between IR and total dry matter yield. There appears to be little
evidence that N fertilizer affects the photosynthetic efficiency of leaves,
although such an effect has been demonstrated with other crops for K.
Relationships between tuber yield and N supply are more complex and
while in general the relationship is asymptotic, response curves with a point
of inversion have been frequently reported. For N this may be attributable
to its effects on initial tuber growth rates, although this is probably cultivar-
dependent. This effect may also be completely or partially offset by
increases in the efficiency of conversion of IR into tuber dry matter as the
season progresses. Such analyses appear to have been restricted to N
fertilizer.
Since relationships between tuber yield and nutrient supply tend to be
asymptotic, and since the cost of fertilizer may be low relative to the value
of the crop, in terms of profitability, predicting the optimal amount of N to
apply to a particular field does not seem to be very critical. Furthermore
the confidence with which optimal rates of N can be predicted appears to
be low. However, concern with the potential contribution of excess
fertilizer nitrogen to the content of nitrate in water draining from agri-
cultural land provides an increasingly important incentive to avoid wastage
of N fertilizer through the accurate prediction of optimal levels. The use of
soil and plant analysis, and modelling all have use.ful roles to play in this
respect. The influences of other factors such as the timing and placement of
fertilizer, and modifications for factors such as cultivar, plant density and
irrigation and the use of organic manures may all affect the efficiency of
fertilizer use. However, the understanding of these influences does not
appear to have advanced substantially since the first edition of this book.
This may in part be due, not to a lack of importance attached to them, but
to problems attached to their funding and, where funding is private rather
than public, with lack of general access to the data.
Fertilizers do have a role to play in determining tuber quality, but it is
difficult to avoid the conclusion that if the quantities of fertilizers applied
are chosen carefully with respect to yield, then they should not have
unacceptable effects on most aspects of tuber quality.
References 209
REFERENCES
Allen, E.J. and Scott, R.K. (1980) An analysis of growth of the potato crop.
Journal of Agricultural Science, Cambridge, 92, 151-{i3.
Asfary, A.F. (1981) Water use and nitrogen uptake in relation to the growth of the
potato crop, Ph. D. Thesis, Reading University.
Augustin, J., McDole, R.E. and Painter, G.C. Influence of fertilizer, irrigation and
storage treatments on nitrate-N content of potato tubers. American Potato
Journal, 54, 125-36.
Batey, T. and Boyd, D.A. (1967) Placement of fertilizers for potatoes. NAAS
Quarterly Review, No. 78, 47-56.
Beaufils, E.R. (1973) Diagnosis and recommendation integrated system (DRIS).
Soil Science Bulletin, No.1, University of Natal, Pietamaritzburg, South Africa.
Birch, J.A., Devine, J.R., Holmes, M.R.J. and Whitear, J.D. (1967) Field
experiments on the fertilizer requirements of maincrop potatoes. Journal of
Agricultural Science, Cambridge, 69, 13-24.
Bodlaender, K.B.A. and Reestman, A.J. (1968) The interaction of nitrogen supply
and plant density in potatoes. Netherlands Journal of Agricultural Science, 16,
165-76.
Bolton, J. (1971) Rothamsted Report for 1970, 2, 98-110.
Boswijk, A. (1976) The effect of fertilizers on the yields of industrial potatoes in an
intensive cropping plan on sandy peat. Annales Agronomiques, 27,771-80.
Boyd, D.A. (1959) The effect of farmyard manure on fertilizer reponses. Journal of
Agricultural Science, Cambridge, 52,384-91.
Boyd, D.A. (1961) Current fertilizer practice in relation to manurial requirements.
Proceedings of the Fertilizer Society, 65, pp. 3-30.
Boyd, D.A. (1970) Some recent ideas on fertilizer response curves. Potash
Symposium, 9, 461-73.
Boyd, D.A. and Dermott, Q. (1964) Fertilizer experiments on maincrop potatoes.
Journal of Agricultural Science, Cambridge, 63, 249-{i3.
Burstall, L. and Harris, P.M. (1983) The estimation of percentage light intercep-
tion from leaf area index and percentage ground cover in potatoes. Journal of
Carter, J.N. and Bosma, S.M. (1974) Effect of fertilizer and irrigation on
nitrate-nitrogen and total nitrogen in potato tubers. Agronomy Journal, 66,
263-{i.
Clutterbuck, B.J. and Simpson, K. (1978) The interactions of water and fertilizer
nitrogen in effects on growth pattern and yield of potatoes. Journal of
Cooke, G.W. (1967) The Control of Soil Fertility, Crosby Lockwood, London.
Crowther, E.M. and Yates, F. (1941) Fertilizer policy in wartime. Empire Journal
of Experimental Agriculture, 9, 77-97.
Dow, A.1. and Roberts, S. (1982) Proposal: critical nutrient ranges for crop
diagnosis. Agronomy Journal, 74,401-3.
Dyson, P.W. and Watson, D.J. (1971) An analysis ofthe effects of nutrient supply
on the growth of potato crops. Annals of Applied Biology, 69, 47-{i3.
Epstein, E. (1972) Mineral Nutrition of Plants: Principles and Perspectives, John
Wiley and Sons, New York, London, Sydney, Toronto.
Evans, C. (1974) Effect of nitrogen on yield of early potatoes. Experimental
Husbandry, 27,99-103.
Ezeta, F.N. and McCollum, R.E. (1972) Dry matter production and nutrient
uptake and removal by Solanum Andigena in the Peruvian Andes. American
Potato Journal, 49, 151-63.
Firman, D.M. and Allen, E.J. (1988) Field measurements of the photosynthetic
rate of potatoes grown with different amounts of nitrogen fertilizer. Journal of
Fitts, J.W. (1974) Proper soil fertility evaluation as an important key to increased
crop yields, in Fertilizers, Crop Quality and Economy (ed. V. Hernando
Fernandez), Elsevier, Amsterdam, pp. 5-30.
Garwood, E.A. and Williams, T.E. (1967) Growth, water use and nutrient uptake
from the subsoil by grass swards. Journal of Agricultural Science, Cambridge,
69, 125-30.
Gerdes, K., Koppen, D., Neubauer, W. et al. (1975). Einfluss hoher Mineralizcher
Stickstoffdungung und Beregnug auf Ertrag und qualitat der Kartoffel. 1.
Quoted by Perrenoud, S. (1983), Potato. International Potash Institute Bulletin,
No.8.
Gregory, P.J., Marshall, B. and Biscoe, P.V. (1981) Nutrient relations of winter
wheat. 3. Nitrogen uptake, photosynthesis of flag leaves and translocation of
nitrogen to the grain. Journal of Agricultural Science, Cambridge, 96, 539-47.
Gunasena, H.P.M. (1969) Studies o,n the growth of the potato with particular
reference to the efficient use of nitrogen and potassium, Ph.D. Thesis, Reading
University.
Gunasena, H.P.M. and Harris, P.M. (1968) The effect of time of application of
nitrogen and potassium on the growth of the second early potato, variety Craig's
Royal. Journal of Agricultural Science, Cambridge, 71,283-96.
Gunasena, H.P.M. and Harris, P.M. (1969) The effect of CCC and nitrogen on the
growth and yield of the second early potato variety Craig's Royal. Journal of
Agricultural Science, Cambridge, 73,245-9.
Gunasena, H.P.M. and Harris, P.M. (1971) The effect of CCC, nitrogen and
potassium on the growth and yield of two varieties of potatoes. Journal of
Hackett, C. (1968) A study of the root system of barley. 1. Effects of nutrition on
two varieties. New Phytologist, 67,287-99.
Harris, P.M. (1960) Investiga,tion into the mode of action of farmyard manure with
particular reference to the potato crop, M.Sc. Thesis, Leeds University.
Harris, P.M. (1983) The use of root data in some agronomic research, in Root
Ecology and its Practical Application (eds W. Bohm, L. Kutschera and E.
Lichtenegger). Internationales Symposium vom 27-29 Sept. 1982, veramstaltet
von der Bundesanstalt fur alpen landische Landwirtschaft, Gumpenstein, pp.
525-33.
Harris, P.M. (1985) Irrigation and the fertilizer requirements of the potato crop, in
Irrigating Potat()res. UK Irrigation Association Technical Monograph 2 (eds
M.K.V. Carr and P.J.C. Hamer), Cranfield Press, Silsoe, Bedford, UK.
Haverkort, A.J. (1985) Relationships between intercepted radiation and yield of
potato crops under tropical highland conditions of Central Africa, Ph.D.
Thesis, Reading University.
Holliday, R. (1963) Effects of fertilizers upon potato yields and quality, in The
Growth of the Potato (eds J.D. Ivins and F.L. Milthorpe) Butterworths,
London, pp. 248-64.
References 211
Holliday, R (1970) Soil profile moisture and nitrogen availability, in Nitrogen

Nutrition of the Plant (ed. E.M. Kirkby), University of Leeds, pp. 189-200.
Holliday, Rand Draycott, A.P. (1968) Effect of placement of liquid or solid
fertilizer on the growth and yield of potatoes. Journal of Agricultural Science,
Cambridge, 71,413-18.
Holliday, R., Harris, P.M. and Baba, M.R (1965) Investigations into the mode of
action of farmyard malilure. Journal of Agricultural Science, Cambridge, 64,
161-6.
Hughes, J.e. (1986) The effects of storage temperature, variety and mineral
nutrition on sugar accumulation. Aspects of Applied Biology, 13, 443-56.
Ifenkwe, O.P. and Allen, E.J. (1983) Nitrogen and potassium uptake by high
yielding potato crops. Journal of Agricultural Science, Cambridge, 101,103-11.
Ivins, J.D. (1963) Agronomic management of the potato, in The Growth of the
Potato (eds J.D. Ivins and F.L. Milthorpe) Butterworths, London, pp. 303-10.
Jemison, J.M. and Fox, R.H. (1988) A quick test procedure for soil and plant tissue
nitrates using test strips and a hand-held retle.ctometer. Communications in Soil
Science and Plant Analysis 14, 1569-82.
Kerketta, R. (1976) Growth, nitrogen nutrition and leaf senescenc.e in the potato,
Ph.D. Thesis, Reading University.
Krauss, A. (1978) Tuberization and abscisic acid cont,ent in Solanum tuberosum as
affected by nitrogen nutrition. Potato Research, 21, 183-93.
Kuipers, S.F. (1962) Factors determining the nitrogen fertilization of arable land.
Stikstof, 6, 3-19.
Kunkel, R., Holstad, N. and Russell, T.S. (1973) Mineral element content of
potato plants and tubers vs. yields. American Potato Journal, 50, 275-83.
Lauer, D.A. (1986) Russet Burbank yield response to sprinkler-applied nitrogen
fertilizer. American Potato Journal, 63, 61-9.
Lisinska, G. and Leszczynski, W. (1989) Potato Science and Technology, Elsevier
Applied Science, London and New York.
Lorenz, O.A., Weir, B.L. and Bishop, J.e. (1972) Effect of controlled-release
nitrogen fertilizers on yield and nitrogen absorption by potatoes, cantaloupes,
and tomatoes. Journal of the American Society of Horticultural Science, 97,
334-7.
MAFF (1985) Fertilizer Recommendations 1985~6. Ministry of Agriculture Fisheries
and Food, Reference Book 209, HMSO, London.
MacKay, D.C., MacEachern, e.R and Bishop, RF. (1966) Optimum nutrient
levels in potato leaves (Solanum tuberosum L.). Soil Science Society of America
Proceedings, 30, 73-6.
MacKay, D.e., Carefoot, J.M. and Entz, T. (1987) Evaluation of the DRIS
procedure for assessing the nutritional status of potato (Solanum tuberosum L.).
Communications in Soil Science and Plant Analysis, 18, 1331-53.
MacKay, D.C., Carefoot, J.M. and Entz, T. (1988) Detection and correction of
mid-season P deficiency in irrigated potatoes. Canadian Journal of Plant
Science, 68, 523-34.
MacLean, A.J., Jasmin, J.J. and Halstead, R.L. (1967) Effect of lime on potato
crops and on properties of a sphagnum peat soil. Canadian Journal of Soil
Science, 47, 89-94.
Mattingly, G.E.G. and Johnston, A.E. (1976) Long-term rotation experiments at
Rothamstead and Saxmundham Experimental Stations: the effects of
treatments on crop yields and soil analyses and recent modifica,tions in purpose
and design. Annales Agronomiques, 27, 74~9.
Meldal-Johnsen, A. and Sumner, M.E. (1980) Foliar diagnostic norms for
potatoes. Journal of Plant Nutrition, 2, 569-76.
Millard, P. and Marshall, B. (1986) Growth, nitrogen uptake and partitioning
within the potato (Solanum tuberosum L.) crop, in relation to nitrogen
application. Journal of Agricultural Science, Cambridge, 107,421-9.
Mitscherlich, E.A. (1913) Bodenkunde fur Land-und Forstwirte, Berlin 1913.
[Quoted by E.W. Russell (1961) Soil Conditions and Plant Growth, 9th edn,
Longmans, London].
Monteith, J.L. (1977) Climate and the efficiency of crop production in Britain.
Philosophical Transactions of the Royal Society of London, 281, 277-94.
Neeteson, J.J. (1989a) Evaluation of the performance of three advisory methods
for nitrogen fertilization of sugar beet and potatoes. Netherlands Journal of
Agricultural Science, 37, 143-55.
Neeteson, J.J. (1989b) Effect of reduced fertilizer nitrogen application rates on
yield and nitrogen recovery of sugar beet and potatoes. Netherlands Journal of.
Agricultural Science, 37, 227-36.
Neeteston, J.J. and Wadman, W.P. (1987) Assessment of economically optimum
application rates of fertilizer N on the basis of response curves. Fertilizer
Research, 12, 37-52.
Neeteson, J.J. and Zwetsloot, J .C. (1989) An analysis of the response of sugar beet
and potatoes to fertilizer nitrogen and soil mineral nitrogen. Netherlands
Journal of Agricultural Science, 37, 129-41.
Neeteson, J.J., Greenwood, D.J. and Draycott, A. (1987) A dynamic model to
predict yield and optimum nitrogen fertilizer application rate for potatoes.
Proceedings of the Fertilizer Society, No. 262.
Ngugi, D. (1972) A study of the physiological basis of yield in the potato crop, with
particular reference to th.e efficient use of nitrogen, Ph.D. Thesis, Reading
University.
Papastylianou, I. (1989) Diagnosing nitrogen fertilization requirements of cereals
in less than 30 seconds. Communications in Soil Science and Plant Analysis, 20,
(11-12), 1247.
Peaslee, D.E. and Moss, D.H. (1966) Photosynthesis in K- and Mg-deficient maize
(Zea mays L.) leaves. Soil Science Society of America Proceedings, 30,220-3.
Perrenoud, S. (1983) Potato: fertilizers for yield and quality. International Potash
Institute Bulletin, No.8, 84 pp.
Price, T.J.A. and Harvey, P.N. (1962) Effect of irrigation on sugar beet and
potatoes. Experimental Husbandry, 7, 1-7.
Prummel, J. and Barnau-Sijthoff, P.A. von (1984) Optimum phosphate and
potassium levels in potato tops. Fertilizer Research 5, 203-11.
Radley, R.W. (1963) The effect of season on growth and development of the
potato, in The Growth of the Potato (eds J.D. Ivins ahd F.L. Milthorpe)
Butterworths, London, pp. 211-20.
Reith, J.W.S. and Inkson, R.H.E. (1958) Effects of fertilizer and dung on
potatoes. Journal of Agricultural Science, Cambridge, 51, 218-24.
Rhue, R.D., Hensel, D.R. and Kidder, G. (1986) Effect of K fertilization on yield
and leaf nutrient content concentrations of potatoes grown on sandy soil.
American Potato Journal, 63, 665-81.
References 213
Roberts, S., Cheng, H.H. and Farrow, F.O. (1989) Nitrate concentration in
potato petioles from periodic applications of 15-N-labeled ammonium nitrate
fertilizer. Agronomy Journal 81, 271-4.
Roberts, S., Dow, A.!. and Cline, T.A. (1984) Slow release nitrogen evaluations
and phosphorus and potassium requirements for potatoes on sandy soil.
Agricultural Research Center, Washington State University.
Sattiemacher, B. and Marschner, H. (1979) Tuberization in potato plants as
affected by application of nitrogen to the roots and leaves. Potato Research, 22,
49-57.
Scaife, M.A. and Bray, B.O. (1977) Quick sap tests for improved control of crop
nutrient status. ADAS Quarterly Review, 27, 137-45.
Schippers, P.A. (1968) The influence of rates of nitrogen and potassium application
on the yield and specific gravity of four potato varieties. Europecl'fl Potato
Journal, 11, 23-33.
Schippers, P.A., Hoogland, R.F. and Krijthe, N. (1969) Influence of NPK-
application to seed-potato crops on the productivity of the progeny. European
Potato Journal, 12, 251-63.
Sharpe, P.R. (1969) The determination and economic analysis by regression
techniques of functional relationships in the potato crop, Ph.D. Thesis, Reading
University.
Smith, P.F. (1962) Mineral analysis of plant tissues. Annual Review of Plant
Physiology, 13, 81-108.
Sokolov, A.V. and Orlovskii, N.V. (1971) Acad. Sci. USSR. Translated 1974,
Israel Program for Scientific Translations.
Ulrich, A. (1952) Physiological basis for assessing the nutritional requirements of
plants. Annual Review of Plant Physiology, 3, 207-28.
van Burg, P.F. (1967) Relation of rate of nitrogen fertilization, seed spacing and
seed size to yield of potatoes. Netherlands Nitrogen Technical Bulletin, 4, 1-30.
van Loon, C.D., Slangen, J.H.O. and Houwing, J.H. (1987) Nitrate content of leaf
petioles as a guide to optimalization of N-fertilization of ware potatoes. 10th
Triennial Conference of the European Association for Potato Research, Aalborg,
1987,146-7.
van der Paauw, F. (1958) Relation between the potash requirements of crops and
meteorological conditions. Plant and Soil, 9, 254-8.
van der Paauw, F. (1963) Residual effect of nitrogen fertilizer on succeeding crops
in a moderate marine climate. Plant and Soil, 19, 324-31.
Warren, R.O. and Johnston, A.E. (1962) Rothamsted Report for 1961, pp. 45-8.
Watson, D.J. and Wilson, J.H. (1956) An analysis of the effects of infection with
leaf-roll virus on the growth and yield of potato plants, and of its interactions
with nutrient supply and shading. Annals of Applied Biology, 44, 390-409.
Wellings, L. W. (1972) The effect of seed rate, nitrogen and irrigation on the yield
of Pentland Crown potatoes. Experimental Husbandry, 22, 39-50.
Wolfe, J.A. (1987) The Potato in the Human Diet, International Potato Center and
Cambridge University Press, Cambridge.
CHAPTER 5
Water relations and growth of

potatoes
P.J. Gregory and L.P. Simmonds
5.1 INTRODUCTION
It is well recognized that an adequate water supply is essential for high yields
of potatoes (e.g. Evans and Neild, 1981) so that year to year and site to site
variability of yields is frequently associated with differences in availability and
accessibility of soil water. For example, McDermott and Ivins (1955) found
that fresh weight yields of Majestic potatoes grown in the East Midlands of
England were directly proportional to the amount of rainfall received
between May and September; over an 8-year period, yield increased by
about 140 kg ha- 1 mm- 1 (Fig. 5.1). Similarly, differences in soil water storage
can have marked effects on yields on sites where irrigation is unavailable.
This chapter examines the relations between growth and water use, and
yield and water supply. Rainfall can be supplemented by irrigation and the
quantity and timing of irrigation is of major interest to producers. Water is
extracted from the soil by roots and their depth and distribution is a key
factor influencing the accessibility of water and hence of yields. However,
yield alone is frequently not th.e prime consideration of many growers and
tuber size and quality are important determinants of the economic return
given by crops. For this reason we shall also examine aspects of the timing
of irrigation and the resulting size distribution of tubers.
5.2 GROWTH AND WATER USE
Transpiration by plants is an inevitable consequence of carbon dioxide

exchange between the atmosphere and the plant; if stomata are open to
allow carbon dioxide in then they will also allow water vapour to pass out.
This association between transpiration and photosynthesis has been ex-
pressed using an Ohm's Law analogy. Transpiration (Td and photo-
synthesis (Nd of individual leaves can be described in terms of the
Growth and water use 215
50
o
o 0
40 0
o
10
o 100 200 300

Rainfall (mm)
Figure 5.1 The relation between yield and rainfall for Majestic potatoes in central
England. (From McDermott and Ivins, 1955.)
differences in water vapour and CO 2 concentrations between the inter-

cellular spaces of the leaf and the outside, ambient air and the resistances
to diffusion of these gases (Bierhuizen and Slatyer, 1965):
(5.1)
Ca - Ci
NL = - - - (5.2)
r's + r' a
where Xi is the vapour concentration at saturation in the substomatal
cavity, Xa is the concentration of water vapour in the ambient air, Ca and Ci
are the concentrations of CO 2 in the air and intercellular spaces respec-
tively and rs and ra are the stomatal and boundary layer resistances to water
vapour (r's and r' a for CO 2 ), Assimilation of CO 2 (photosynthesis) does
not occur in the intercellular spaces so that strictly there are other
resistances of both physical and chemical nature (the mesophyll and
carboxylation resistance) to be overcome before assimilation. These addi-
tional resistances are typically of the same magnitude as (r's and r' a). The
ratio of photosynthesis to transpiration can be written:
216 Water relations and growth of potatoes
(Ca - Ci)(rs + ra)
(5.3)
(Xa - Xi)(r's + r' a)
This expression can be simplified considerably. First, the resistances
to both CO 2 and water vapour exchange are related to the diffusion
coefficients for the two gases which are approximately inversely pro-
portional to the square root of their molecular weights. This means that the
ratio (rs + ra)/(r' s + r' a) is approximately constant with a value of 1.56.
Second, several researchers have shown that the ratio of C/Ca is approxi-
mately constant although slightly affected by factors such as leaf age and
leaf water potential (Vos and Oyarzun, 1987); c/ca has a value of about 0.7
for species such as potatoes which photosynthesize by the C 3 pathway
(Wong et aI., 1979; Pearcy and Ehleringer, 1984). Finally, over extended
periods of time, leaf temperature is frequently close to air temperature so
that Xa - Xi can be approximated by XS - Xa where XS is the vapour
concentration at saturation of the ambient air; XS - Xa is proportional to
the saturation deficit of the air, D (D = 0.135(Xs - Xa) at 20°C, where
concentrations are in g m- 3 and D is in kPa). Making these assumptions and
assuming also that crops behave similarly to individual leaves (see
Tanner and Sinclair, 1983 for discussion of this), equation (5.3) can be
approximated for crops as:
N k
(5.4)
T D
where N is the dry weight gain by the crop, T is the transpiration by the
crop, and k is a crop-specific constant (with units of g m- 3 ).
Figure 5.2a shows the relation between Nand T obtained by Rijtema
and Endrodi (1970) using four cultivars of potato over a 6-year period in
The Netherlands. Clearly the scatter is large but when the differences in D
between years were taken account of, a good linear relation was obtained
(Fig. 5.2b) which was independent of the cultivar used. Similarly, Tanner
(1981) found a linear relation between N and TID for the cultivar Russet
Burbank grown for three seasons in Wisconsin, USA.
There is good experimental evidence to suggest that k is a conservative
parameter for many crops (see Tanner and Sinclair (1983) and Monteith
(1990) for discussion) with differences between crops primarily related to
the photosynthetic pathway. Table 5.1 shows values of about 55-75 g m- 3
for C4 cereals such as maize, and millet, and about 20--30 g m- 3 for C 3
species such as barley and soyabeans. However, for potatoes, substantially
different values of k have been reported. Rijtema and Endrodi (1970)
found a value of 7 g m- 3 (the gradient of the line in Fig. 5.2b) whereas
Tanner (1981) found values between 40 and 44 g m- 3 and Bunyolo
(1987) found values between 20 and 30 g m- 3 . The reason for these
differences is not known with certainty but reflects, in part, the
Growth and water use 217
15 (a)
•
DO
,.
••
0
0
0
0
10 0
0 0
0
5
I
.r:
co 0 100 200 300
~
Transpiration (mm)
CD
==co
E 15 (b)
2:-
"0
ro
(5
f-
10
o 10 20
Transpiration (mm)
X. - Xa (91m3 )
Figure 5.2 The relation between total dry matter production and (a) transpiration
and (b) transpiration normalized for the deficit of atmospheric water vapour
concentration. The results are for four genotypes (0, Ackersegen; ., Libertas; D,
Surprise;., Daleo) grown over seven seasons. (From Rijtema and Endrodi, 1970.)
uncertainties of calculating D; Rijtema and Endrodi used a 24-h mean

value whereas the other authors used a value for daylight hours only. A
further factor that may contribute to the variation is that potatoes are
known as a drought-sensitive crop and even in wet soils stomatal closure
and partial wilting may occur when plants are exposed to a large atmo-
spheric demand for water. In such circumstances the difference in
temperature between the leaf and air may be large. Finally, in contrast to
Table 5.1 Values of the crop coefficient, k, for potatoes in
comparison with other species
Crop Lo~ation k Author
(g m- 3)
Potato Wisconsin, USA 44 Tanner (1981)
Netherlands 7 Rijtema and Endrodi (1970)
Reading, UK 21-30 Bunyolo (1987)
Millet Various 70 Monteith (1989)
Maize Mid-West USA 70 Tanner and Sinclair (1983)
Maize Ontario, Canada 55 Walker (1986)
Barley Rothamsted, UK 21 Day et al. (1987)
Soyabean Kansas, USA 30 Tanner and Sinclair (1983)
Rijtema and Endrodi, Bunyolo found differences in k between two

cultivars used in their experiments (Wilja averaged 30 and Cara 20 g m- 3 ).
The ratio NIT is frequently termed the transpiration efficiency and
despite the apparent conservatism of k, there is variation between culti-
vars. Differences between cultivars in mesophyll and carboxylation
resistances (assumed to be constant in the preceding discussion) may
contribute substantially to the observed differences between cultivars.
Transpiration efficiency may also increase with increasing stomatal resis-
tance (Vos and Groenwold, 1989a). Bodlaender (1986) showed differences
in NIT between cultivars of potato ranging from 5.4 g kg- I to 7.9 g kg- I and
Vos and Groenwold (1989a) confirmed Bodlaender's results for two of the
cultivars to show a difference between Bintje and Saturn a of 0.8 g kg-I.
Such measured differences have led to speculation as to whether useful
genetic differences exist that might be usefully used by plant breeders. An
indirect method for screening for NIT has been devised based on a theory
developed by Farquhar et al. (1982). During photosynthesis, the naturally
occurring stable isotope 13C is discriminated against relative to I2C because
of its greater mass. This means that plants have a slightly smaller ratio of
13c;tzc than the ambient air and this can be measured by mass spectro-
scopy. Farquhar et al. (1982) predicted that the discrimination would be
least in those plants with the greatest transpiration efficiency (i.e. discrimi-
nation should be related to NI1) and this was subsequently proven in
experiments with wheat (Farquhar and Richards, 1984) and groundnut
(Hubick et aI., 1986). Vos and Groenwold (1989b) have shown similar
genetic diversity in potatoes (Fig. 5.3) and further work may be expected in
an attempt to identify high NIT especially in conditions where water is
limited.
Yield and water supply 219
i -18
c:
o
~ o
c:
o
~
.:= -20
OJ o
i
c:
00
o
o
0
~
8 -22 L-_--'-_-L-_~___I
8 10 12
Transpiration efficiency (g kg-')
Figure 5.3 The relation between carbon isotope discrimination and transpiration
efficiency. The results are for six genotypes, three of which were grown in
droughted conditions (upper 3 points). (From Vas and Groenwold, 1989b.)
5.3 YIELD AND WATER SUPPLY
Section 5.2 clearly showed the relation between total dry matter pro-
duction and transpiration yet growers are concerned only with the yield of
tubers and in the production of these, water is lost by means other than
transpiration alone. Two questions therefore arise: first, is there any
relation between yield of tubers and the total dry matter production of the
crop? And second, is there any relation between the quantity of water
transpired and the total quantity of water used by a crop which may include
evaporation directly from the soil surface, drainage, and, in the event that
the intensity of rainfall or irrigation exceeds the rate of infiltration into the
soil, runoff?
In an early attempt to define the partitioning of dry matter between
tubers and leaves and stems, Ivins and Bremner (1965) developed an
hypothesis that the growth of the crop involved a conflict between tubers
and shoots such that anything that promoted the growth of one part was at
the expense of growth of the other. However, more recent experiments
suggest that, in general, this is not the case. For example, Fig. 5.4 shows a
linear relation between tuber dry weight and total plant weight irrespective
of season, nitrogen application, and irrigation treatment. Similar results
have been reported by Rijtema and Endrodi (1970), Allen and Scott (1980)
and Millard and Marshall (1986) although the relation is not constant
during the growing season and may be affected by severe drought and
fertilizer applications (see Chapter 4). However, the general relations
described in Section 5.2 seem equally applicable to tuber yield as to total
dry matter production; differences in partitioning are, in general, small.
There will, of course, be specific circumstances when partitioning will not
be constant (if, for example, early growth is promoted by the application of
fertilizers causing exhaustion of soil water before tuber growth can occur)
but these are unlikely when adequate fertilizer and water are supplied.
.
15
o
~
~ 10
o 5 10 15 20
Total plant dry weight (t ha- 1 )
Figure 5.4 The relation between tuber dry weight and total plant dry weight for
potato cultivar Cara; 0, +nitrogen +irrigation; .,+nitrogen -irrigation; 0,
-nitrogen +irrigation; • -nitrogen -irrigation. (From Bunyolo, 1987).
Water is lost from potato crops by transpiration and evaporation directly

from the soil surface so that the relation between yield and total water use
(evapotranspiration) may not be linear especially if large amounts of water
evaporate directly from the soil surface and do not contribute directly to
growth. The amount of water that evaporates is controlled by several
factors (for a discussion see Ritchie, 1972) but the two principal factors are
the wetness of the soil surface and the amount of radiation reaching the
soil. Both of these quantities are likely to be greatest during the early part
of the growing season and to decrease as the crop grows. Although the soil
surface may remain damp throughout the growing season, the interception
of radiation by a closed canopy will ensure that rates of evaporation from
the soil surface are low. MacKerron and Waister (1985) suggest that
canopy closure is achieved in potatoes with a leaf area index (LAI) of 3 and
that this LAI will intercept 95-98% of the incident radiation. Burstall and
Harris (1983), however, indicate that an LAI of 6 may be necessary. These
values of LAI will be achieved easily in well fertilized and irrigated crops in
the UK but in adverse conditions (e.g. water, nutrient or disease limited) a
substantial part of the incident radiation may not be intercepted by the
crop canopy. For example, Millard and Marshall (1986) grew crops without
nitrogen fertilizer and found only 79-85% of the incident radiation was
intercepted. Similarly, compaction of sub-soil will often reduce leaf growth
and increase evaporation from the soil surface.
Irrigated crops rarely lose a large proportion of their total water use
directly from the soil surface so that yield and the amount of irrigation and
rainfall received are frequently linear (Fig. 5.5) provided drainage is small.
The gradient of the relation is dependent on many factors not least the soil
Yield and water supply 221
o
~'" 4
.<::
o
o o
o
o
o
o
300 400 500 600 700
Irrigation (mm)
Figure 5.5 The relation between tuber dry weight and the amount of irrigation
applied to potato cultivar Revolucion during the summer at Lima, Peru. (From
International Potato Center, 1986.)
water-holding capacity, the saturation deficit of the atmosphere, and the

timing and management of the irrigation system. Indeed, in wet years,
yield may not respond to irrigation or may become a negative response.
It is widely accepted that potatoes are very sensitive to drought and large
responses to irrigation are common; the reasons for this sensitivity to
drought will be explored later. Two major questions for any irrigated crop
are how much water should be applied and when? Attempts to answer
these questions led to the development of the concepts of Field Capacity
(FC), Permanent Wilting Point (PWP) and Available Water Capacity
(AWC). The total amount of available moisture (TAM) within the soil
profile is:
TAM = oszrFC - PWP (5.5)
where zr is the depth of rooting. These simple concepts have been widely
used and assume that all of the water held in the soil between FC and PWP
is available to the plant and that growth will continue until it has all been
used. Tables of A WC are widely available (see, for example Wild, 1988)
and values typically range from 13.6 to 22.8 mm water per 100 mm soil as
texture changes from medium sand to fine sandy silt loam; soils with
appreciable clay content typically fall mid-way between these limits.
Assuming a rooting depth of 0.5 m, the soils referred to above would
require irrigation after 19 and 32 days respectively in order to maintain a
supply of available water in a typical English summer with evaporation
rates of 3.5 mm- 1 day. Such simple calculations are useful for showing the
important effects of soil texture but in most soils not all of the available
water is equally available and the proportion depends upon the crop,
climate and soil. For example, results of Peeler et al. (1966) suggest that
maximum yields are obtained by irrigating when only 50% of the available
water has been depleted. Moreover, the calculation depends on a
knowledge of rooting depth; a depth that is rarely known with certainty.
Another approach to the problem, and one that has been widely
accepted by farmers and their advisors, is to base requirement and timing
of irrigation on calculation of atmospheric demand for water and an
experimentally determined value of the amount of readily available water
that the soil can hold. Penman (1970) suggested that unrestricted crop
growth continues until the profile water is depleted to a certain value (the
limiting deficit, D,). Beyond D, both further water loss and growth are
deemed to have ceased. To quote Penman (1970): 'This, though obviously
too drastic a division, has the merits of being simple, meaningful, applic-
able to field results, and it seems to work'. It is assumed that at some time
before the crop starts to grow the soil is at, or close to, field capacity.
Thereafter, the crop uses water at a rate determined by the potential rate
of evaporation (Ep) until the limiting deficit is reached. The value of Ep can
be calculated from standard meteorological measurements (e.g. MAFF,
1967). Calculation of Ep assumes complete ground cover and a freely
evaporating surface; conditions that are the aim of irrigation. When the
deficit exceeds D" the actual amount of water estimated to be lost by
evaporation is calculated using the simple assumptions stated above: i.e.
(5.6)
where Dp is the potential soil water deficit (i.e. the deficit that would have
been achieved if evaporation had proceeded at the potential rate) given by:
Dp = Ep - (R + l) (5.7)
where R and I are rainfall and irrigation.
Combining equations (5.6) and (5.7) leads to the result:
Ea = R+I+D, (5.8)
The aim of irrigation is to ensure that D, is not exceeded but, because of
the costs of equipment and its management, to time applications of
irrigation as close to D, as possible. Penman proposed that yield (Y) was
proportional to the amount of water evaporated:
Y = KEp when Dp < D,
and
Y = K(Ea) = K(Ep - (Dp - D,» (5.9)
where K is a constant. Estimates of Ea and of the timing of irrigation both
require values for D, which varies both with crop (a function of rooting
depth and morphology) and site (a function of soil type). French and Legg
(1979) compared values of D, for potatoes at two sites and found that it
varied from about 35 mm on the sandy loam at Woburn to about 80 mm on
the clay loam at Rothamsted (Fig. 5.6). Table 5.2 compares values of D,
for potatoes with other crops and indicates that potatoes are less tolerant of
deficits than most other common crops.
Before Dp can be calculated for potato crops there are two changes that
Root growth 223
E 1.5
g D,
"0
a;
':;" 0..
§
I..~.
'0
5"0
t~
1.0
--~
~'E
a:~ 0.5
o
"''''
o 50 100 150 200
Maximum potential soil
moisture deficit (mm)
Figure 5.6 The relations between yield of tubers as a proportion of yield from a
fully irrigated crop and the maximum potential soil water defi-cit for potato crops
grown at Woburn (0) and Rothamsted (e). The points Dl represent the limiting
deficit at either site. (From French and Legg, 1979.)
Table 5.2 Comparison of the limiting

deficit, Dl (mm), for potatoes with other
crops (data from Penman, 1971 and
French and Legg, 1979)
Crop Rothamsted Woburn
(clay loam) (sandy loam)
Potato 80 35
Barley 100 40
Beans 80 30
Sugar beet 100
Wheat 140 30
must be made to Penman's original hypothesis. First, crop canopies do not

completely cover the soil surface until LAI is >3; prior to this, evaporation
may be <Ep especially if the soil surface is dry. Generally, Ep can be used
when ground cover is >0.5 but when cover is smaller, Ep must be
multiplied by an appropriate fraction. The second change allows for the
fact that potato crops have a rougher canopy than short grass so that
exchange of water vapour is enhanced. French et at. (1973) found that the
actual evaporation from irrigated crops was 1.08 Ep.
5.4 ROOT GROWTH
The supply of water to potato crops is dependent not only on the storage
capacity of the soil but also on the size and extent of the root system.
Measurements on roots are difficult to make and consequently there is a
dearth of information about the spatial and temporal changes in root
distribution and about how the distribution is affected by environmental
and genotypic factors.
The temporal pattern of root growth of potatoes is less clearly known
than that in other crops. In temperate cereals, for example, it has been
found that the size of the root system does not increase after about anthesis
(Gregory et al., 1978; Barraclough and Leigh, 1984); some root growth
continues but this is balanced by the decay of older roots and in dry
conditions, root death may exceed growth (Mengel and Barber, 1974). For
potatoes, the pattern is less certain largely because there have been fewer
studies. Lesczynski and Tanner (1976) found that the root system (length
and mass) of a frequently irrigated (3-5 day intervals) crop of potatoes
grown on a loamy sand continued to increase in size until 89 days
after emergence when senescence of the whole plant commenced. As
senescence occurred, the root system decreased in size so that by 107 days
after emergence it was 30-50% smaller, varying with the cultivation
practice. In contrast, Asfary et al. (1983) found that root growth occurred
early in the life of crops grown on a sandy loam and root length hardly
changed between 14 days after emergence and harvest some 11 weeks
later. Irrigation in this latter study was infrequent and only given when the
soil moisture deficit approached 100 mm. The sequential measurements of
root growth reported by Vos and Groenwold (1986) for unirrigated crops
grown on marine clay show different patterns in two consecutive years. In
1982, root length and mass increased rapidly during early growth and were
maximal at 45 days after emergence. Thereafter there was a slight decrease
(about 10%) in the size of the system which then remained constant until
101 days after emergence. In the wetter, duller season of 1983, planting
occurred very late (late June) and root growth continued throughout the
period of sampling up to 80 days after emergence. The different temporal
patterns of growth reported may be associated with differences in soil
wetness and irrigation practice. Continued root growth is apparently
associated with wet soils and almost continuous irrigation but this associa-
tion is based on a very limited number of observations and the factors
regulating differences in root length density between sites and seasons are
poorly understood.
Table 5.3 summarizes information on rooting depth and the maximum
length and mass of root systems of potato crops. The depth of rooting of all
potato crops never exceeded 1 m even on deep, uniform soils; this is similar
to the behaviour of some temperate legume crops (e.g. field beans) but
roots of many temperate cereals have been found at depths of almost 2 m.
It is difficult to reach firm conclusions about the depth of rooting of potato
crops in comparison with other crops because comparative measurements
on deep soils are rare. On the deep Sassafras loam of New Jersey, USA,
Corey and Blake (1953) found that the rooting depth of potatoes was 56 cm
compared with 77 cm for maize and >90 cm for tomatoes. Similarly, Ovaa
and de Smet (1984) observed a shallower rooting depth for potatoes (50
Root growth 225
cm) compared with winter wheat, spring barley and sugar beet (>1 m)
grown on a clay overlying a sandy subsoil. Durrant et al. (1973) compared
the growth of roots of potato, sugar beet and barley on a sandy clay loam
overlying chalk at a depth of about 1 m. Root growth was observed
through glass panels and related to measurements of water extraction
made with a neutron probe. Over the 3 years of measurements, the
observed depth of rooting was closely related to, but some 10--15 cm
greater than, the depth of soil water extraction. The mean maximum depth
of water extraction in mid-July was 77 cm for potato, >92 cm for sugar beet
and > 100 cm for barley. These mid-season differences in rooting depth
were also reflected in the amounts of water extracted from deeper soil
layers (Fig. 5.7) particularly later during the season. By mid-August, the
soil water deficit in the 50-60 cm layer was 12, 10 and 8 mm for barley,
sugar beet and potatoes respectively, but in the 9Q-.100 cm layer the
differences between species were much greater at 10, 7 and 3 mm
respectively.
Table 5.3 Comparison of results of depth and size of root systems of potato crops
with other temperate crops. Where studies involved different cultivations, the results
from conventional cultivation (usually ploughing) have been adopted
Soil Cultivar Rooting Root Root Author
depth mass length
(m) (g m- 3 ) (km m- 2)
Potatoes
Loamy sand Vanessa 0.9 12.0 Asfary et al. (1983)
Sandy clay
loam Majestic 0.9 Durrant et al. (1973)
Loamy sand Russet 0.7 100 8.4 Lesczynski and
Burbank Tanner (1976)
Sandy loam Record 0.8 10.0 Parker et al. (1989)
Sandy loam K. Edward 0.47 Steckel and
Majestic 0.55 Gray (1979)
M. Piper 0.50
P. Crown 0.60
Sandy clay Desiree 1.0 Stone (1982)
loam
Marine clay Bintje 0.8-1.0 77 7.1 Vos and
Groenwold (1986)
Wheat
Sandy loam M. Huntsman 2.0 110 23.5 Gregory et at. (1978)
Field beans
Sandy loam 0.8 30 1.7 Gregory (1988)
(a)
..... '.:- - - --
, •••• O' •
...,, .
10
, .'I
.'
i 0
!E . - ... --- --
10
--
........ e· ••••
'
.s " ..'
,,~.
'" .'
ti . . . . . . .-:
~.'
".'
o
~ (c)
~ 10
.~
Q) ... ' " " . , - - - - - ........
.~ .' ;'"
1ii 0 •.r. ....;-..::.::,.:,,-
::;
~ 10 [ f d ) ..•...•......
....... ,""'-----
a -----..:~- ..
I I I I
June July Aug Sept
Figure 5.7 Cumulative soil drying in four 10-cm layers (a) 30--40 cm; (b)
50--60 cm; (c) 70--80 cm; and (d) 90--100 cm beneath crops of barley (--), potato
(--), and sugar beet ( ... ). (From Durrant et al., 1973).
The quantity of roots produced by potato crops is generally similar to or

greater than that produced by many temperate legume crops (Gregory,
1988) but smaller than that produced by temperate cereals. Table 5.3
suggests that root systems of potatoes have about one-third the dry weight
and one-third the length of cereal crops grown in comparable regions.
Moreover, as several workers have suggested, the root system of potatoes
often appears to be more surface orientated than other crops. Figure 5.8
compares the root distribution of crops of winter wheat and potatoes
grown on a sandy loam. Not only is the rooting depth of potatoes shallower
but a greater proportion of the root length (82%) is contained in the upper
30 cm compared with winter wheat (73%). Lesczynski and Tanner (1976)
also found that the roots of Russet Burbank potatoes grown on a loamy
sand were restricted to the upper 30 cm and a maximum of only 15% of
roots were found below this depth. A similar result was found by Parker et
al. (1989) on a sandy loam soil.
The apparent inability of potato root systems to root as deeply and as
densely as other crops means that potatoes have proportionately more of
Root growth 227
Root length (cm/cm 3 )
o 2.0 4.0 6.0
I
30 I
I
1
E
.3-
-=c. 60
al
D hi-
90
120
Figure 5.8 Comparison of root distributions of winter wheat (--) and potato
(--) grown on a sandy loam at Sonning Farm, University of Reading.
their root system close to the soil surface. Greenwood et al. (1982) have
shown that the distribution of root length in many vegetable crops changes
logarithmically with depth such that the root length density (Lv) at depth z
is given by:
Lv = LvO e -qz (5.10)
where LvO is the root length density at depth z = 0 and q is the slope of a
plot of In Lv against z. This equation implies that if the rooting depth of a
crop is shallow then a greater proportion of the roots will be in the surface
layers unless compensated for by a change in q. The shallower rooting
depth of potatoes compared with cereals is apparently not compensated for
by changes in q. However, an alternative explanation for the greater
surface orientation might be the greater sensitivity of potato roots to
unfavourable subsoil conditions and pans caused by cultivation practices
compared with other crops such as cereals. In The Netherlands, several
studies have shown the limited extension of potato roots in fields with a
ploughpan in a marine loam soil (Boone et al., 1985) and in soils
comprising a clay layer overlying a sandy or loamy subsoil (Ovaa and de
Smet, 1984). Removal of ploughpans by deep cultivation will only increase
rooting depth if pores with a diameter equal to or larger than the diameter
of potato roots are present in the subsoil. The results of deep cultivation
differ between seasons and sites depending on the inherent porosity of the
soil, the distribution of pore sizes, and the conditions necessary for large
pores to form (Stone, 1982; Ovaa and de Smet, 1984; Parker et aI.,
1989).
Genotypic differences in root growth have rarely been investigated
although Steckel and Gray (1979) showed small differences in the rooting
depth of four varieties of potato. However, neither these differences nor
the small differences in water extraction between varieties could be related
consistently to the differences in yields observed.
5.5 THE PHYSIOLOGICAL BASIS FOR THE DROUGHT-

SENSITIVITY OF POTATO GROWTH
The small values for limiting soil water deficit (Table 5.2) suggest that the
transpiration and growth of potatoes may be particularly sensitive to soil
water deficit compared with many other major food crops. This section
considers some of the factors that might be responsible for the drought-
sensitivity of potatoes.
The rate of transpiration and the water status of the plant tissue are
influenced by both the evaporative demand of the atmosphere and the
availability of soil water. Increasing the rate of evaporation from the sub-
stomatal cavity causes the water content of leaves to decrease, which, at
least in the short term, results in a decrease in both the hydrostatic pressure
and osmotic components of the water potential within the leaf ('P leaf ).
Because there is hydraulic continuity between the water within leaves, and
because water has to flow through semi-permeable membranes in order to
pass from soil to leaf, a reduction in these components of 'Pleaf increases
the rate of water flow from the soil to the leaf. The magnitude of the
difference in water potential between soil and leaf depends on the rate at
which water is drawn through the plant, and hence the rate of evaporation
from leaves.
If evaporation were unrestricted, then tissues would dehydrate severely
under conditions of large evaporative demand, low soil water potential, a
large hydraulic resistance in the flowpath from soil to leaf, or a combina-
tion of these. In practice, excessive desiccation is avoided by stomatal
closure which increases the resistance to gas exchange between leaf and air
(both water vapour and CO2 ), and thereby restricts both transpiration and
photosynthesis, and limits the reduction in leaf water potential.
In this section, an attempt is made to examine the extent to which the
drought-sensitivity of potatoes is attributable to stomatal responses to
water deficits (restricting growth and transpiration as stomata close) and to
the plant hydraulic factors that govern the uptake of water from soil.
5.5.1 Stomatal responses to water deficits

The stimuli responsible for triggering stomatal closure are the subject of
much debate. Much work in past years has been directed towards
establishing relationships between stomatal conductance and various inter-
dependent indices of leaf water status (such as leaf water potential, turgor
pressure and relative water content). One of the motivations for much of
this work was the appeal of using leaf water potential as a hydraulic 'signal'
Physiological basis for the drought-sensitivity of potato growth 229
to couple stomatal control of evaporation with the movement of water
from soil to leaf. However, any relationship established between stomatal
conductance and leaf water potential is highly empirical, and is inevitably
complicated because stomata respond to many environmental factors (e.g.
temperature, irradiance and humidity). The analyses of Shimshi et al.
(1983) and Levy (1983) suggest that increasing stomatal resistance in
potatoes is associated with values of 'Pleaf less than about -0.6 MPa. This is
consistent with the observations that the rates of photosynthesis (Munns
and Pearson, 1974; Meijer, quoted by van Loon, 1981) and transpiration
(Campbell et al., 1976) of potato plants grown in phytotrons were reduced
when leaf water potential fell below -0.4 or -0.5 MPa. In contrast,
Ackerson et al. (1977) observed that stomatal resistance measured on the
adaxial and abaxial surfaces of potato leaves was little affected until leaf
water potential fell below -0.8 MPa or -1.2 MPa respectively.
These observations suggest that transpiration in potatoes is limited by
stomatal closure at relatively high leaf water potentials in comparison with
many other species. For example, threshold values of 'Pleaf for stomatal
closure in wheat range from -1.1 to -2.5 MPa, depending on the degree
of osmotic adjustment (Miller and Denmead, 1977; Seaton et al. 1977):
approximate corresponding values for beans, lucerne and soybeans are
about -0.8, -1.0 and -1.2 MPa respectively (see the appropriate chapters
in Teare and Peet, 1983).
In recent years doubt has been cast on the premise that stomatal closure
is triggered by leaf water status alone (Kramer, 1988; Passioura, 1988;
Schulze et al., 1988; Boyer, 1989). Passioura (1988), for example, has
pointed out the danger of mistaking empirical correlations such as those
described above for cause and effect. Critical experiments with a number
of species in which ingenious methods have been used to break the
correlation between the water status of soil and shoots have shown that
stomatal conductance and growth appear to respond to the water status of
the soil, rather than leaf turgor. Evidence is accumulating that root-shoot
communication at the biochemical level may be implicated in plant
responses to water deficits, though it may be that such root-shoot
interactions may complement, rather than exclude, a theory based on plant
hydraulics.
Whatever the mechanism responsible for triggering stomatal closure,
measurements of leaf water potential in potato crops provide evidence that
stomatal control of transpiration is such that potatoes, in comparison with
some other species, have limited ability to maintain a large difference
between soil and leaf water potential. Leaf water potentials in droughted
potato crops rarely fall below about -1.0 to -1.5 MPa, even when soil
water deficit and atmospheric demand for water are large (Rutherfoord
and de Jager, 1975; Gandar and Tanner, 1976; Shimshi et al., 1983; Wolfe
et al., 1983; Shimshi and Susnoschi, 1985; Stark et al., 1987; Stark and
Wright, 1985; Jefferies, 1989), although there are some reports of minimum
values of 'Pleaf approaching -2MPa (Ackerson et al., 1977). In
contrast, leaf water potentials in many other crops readily fall below
-2MPa (see Cary and Wright, 1971, for examples for maize, wheat,
lucerne and sugar beet). One interpretation of these observations is that
potatoes avoid leaf water potentials below about -1.5 MPa by limiting
transpiration through stomatal closure, whereas more drought-resistant
species maintain low stomatal resistance until larger leaf water deficits are
experienced.
Split root system experiments such as those described by Blackman and
Davies (1985) have demonstrated that if part of the root system is in dry
soil than stomata may close even though leaves remain turgid as a result of
water uptake from roots in wet soil. These observations support the
concept of non-hydraulic root-shoot communication as a stomatal control
mechanism. Passioura (1988) suggests that such phenomena may occur
often in the field when the upper part of the profile dries to lower soil water
potentials than occur in the plant. We might speculate that the propensity
for stomatal closure in potatoes at modest soil and leaf water deficits might
be due, at least in part, to the shallow rooting habit which results in a
substantial proportion of the root system being close to the soil surface and
exposed to extreme drying. However, such phenomena would not explain
the poor ability of potatoes to maintain rapid growth when exposed to high
evaporative demand under conditions where the whole soil profile is
relatively wet - to explain this characteristic of potatoes we need to
consider the hydraulics of the potato crop.
5.5.2 Water uptake from soil

There is much evidence that a gradient in water potential (or the
appropriate components of water potential) provides the driving force for
water flow from soil to leaf (Boyer, 1985; Kaufmann, 1976). The relation-
ship between the rate of water uptake and the water potentials in soil and
leaf is shown schematically in Fig. 5.9, where line AB shows the rate of
water uptake from soil as a function of 'P leaf . The intercept at A represents
the 'effective mean soil water potential' ('P.oil), which is equivalent to the
value of '!'leaf for a non-transpiring plant in static equilibrium with the soil.
When plants are transpiring, the energy status of water in leaves will be less
than that in soil because energy is dissipated in overcoming the frictional
forces that resist the movement of water from soil to leaf. By analogy with
Ohm's Law, the resistance (R) to water flow in the pathway from the bulk
soil to the leaf can be defined as:
R = ('Psoil - 'Pleaf)IT (5.11)
This simple model provides the basis for two hypotheses to explain the
limited ability of the potato crop to maintain transpiration close to the
A
Leaf Water Potential 0
Figure 5.9 The rate of uptake of water from soil as a function of leaf water
potential. The intercept at A indicates the effective soil water potential.
potential rate as the soil profile dries or the evaporative demand is

increased. These are as follows.
1. Potato plants are relatively poor conductors of water, possibly as a
result of having a relatively small root length per unit land area by
comparison with more drought-resistant species. This would result in
the slope of line AB (Fig. 5.9) being shallow for potatoes compared with
more densely-rooted species.
2. The shallow rooting habit may result in potatoes experiencing a lower
effective mean soil water potential than crops with a larger proportion
of the root system extending into wetter soil at depth. Reducing 1f1soil
causes line AB to be displaced to the left, with the result that the rate of
transpiration achieved by a given leaf water potential is reduced.
These hypotheses are examined in more detail below.
5.5.3 The influence of root characteristics on the resistance to water flow

from soil to leaf
The relation between transpiration rate and the difference in water
potential between soil and leaf (line AB, Fig: 5.9) is complex. Studies of a
number of species (e.g. Kaufmann, 1976) have shown that the relationship
may be non-linear (with hydraulic resistances commonly increasing
dramatically at low flow rates). Also, the hydraulic resistance to flow
through the soiVplant system will tend to increase as the soil around a root
dries, because of decreasing soil hydraulic conductivity and the develop-
ment of a significant resistance at the root/soil interface (Bristow et aL,
1984).
Table 5.4 illustrates the effect of a substantial hydraulic resistance in the
flowpath from soil to leaf. In this example (where the soil is relatively wet),
the difference in water potential between the bulk soil ('I'soil) and the
xylem at the base of the stem ('I'stem) was about 0.4 MPa, whereas
'l'stem-'I'leaf was much smaller ("'" 0.1 MPa). These measurements suggest
that in potatoes, as in many other species (Newman, 1976; Landsberg and
Fowkes, 1978), the major resistance to water flow between soil and leaf lies
below ground. Recent studies at the University of Reading have shown
that 65-75% of the overall resistance to flow between soil and leaf is found
below ground for potato crops growing in wet soil. In veery dry soil, this
figure exceeds 80%, because of a reduction in soil hydraulic conductivity
and, possibly, loss of hydraulic contact between root and soil.
Table 5.4 An example of the gradient in water potential through the soil/plant
system for potatoes growing in uniformly wet soil (see also Vas and Groenwold,
1988, for an example of gradients in water potential through the canopy of a potato
crop)
Location Water Method of measurement
potential
(MPa)
Soil ('I'soil) -0.01 Tensiometer

Base of stem ('I'stem) -0.38 Water potential of covered (i.e. non·
transpiring) lower leaf measured using
a pressure chamber
Uppermost fully-expanded leaf
('I'leaf) -0.50 Pressure chamber
Bristow et al. (1984) presented evidence that in wet or moist soil (where
30% or more of the soil pore space is filled with water) the resistance to
water flow within the root system is likely to dominate the below-ground
resistance. The principal resistance to water flow within roots is probably
associated with the radial flowpath from the root surface to the xylem,
rather than with axial flow along the xylem (Taylor and Klepper, 1971;
Sands et al., 1982; Hamblin and Tennant, 1987). If so, the rate of water
uptake (per unit soil volume) achieved per unit difference in water
potential between soil and stem would be expected to vary proportionally
with the root length density. Figure 5.10 presents evidence to support this
contention, showing that the 'below-ground' conductance (gbg) in stands of
well-watered potatoes was approximately proportional to the length of
root per unit land area (La). The values of gbg used in Fig. 5.10 were
calculated using:
(5.12)
where T is in units of mm transpiration per hour, and water potentials are
expressed in MPa. The data presented in Fig. 5.10 were obtained when the
soil throughout the root zone was uniformly wet (\fsoil > -20 kPa) , so that
resistance to water flow through the soil to the root surface was probably
negligible, and there was little ambiguity about the value for soil water
potential to use in equation (5.12).
~
~ 2r---------------~
g
~ 1.5
•
§.
~ • I
~
i
1
• • • ,
-g
•
0.5
~
f
(Il
OL-·~--L-~--L-~~
5 10 15
root length per unit land area (km/m2)
Figure 5.10 The influence of the overall length of the root system on the
conductance to water flow between the bulk soil and the base of the stem. Based on
measurements made in a well-watered crop (unpublished results).
The conductances recorded in Fig. 5.10 are sufficiently small to suggest

that the drought-sensitivity of potatoes is due in part to the limited ability
of the root system to convey water. The slope of Fig. 5.10 (the conductance
per unit root length) has a value similar to that reported for other species
(e.g. Bristow, 1984, for sunflowers). Hence we might speculate that the
low values of gbg are due primarily to the relatively small root length
compared with, say, cereals (d. Table 5.3), rather than an inherently small
conductance per unit length of root.
The value of gbg for a relatively well-rooted crop (say, La = 10 km m- 2)
was approximately 1 mm h- 1 MPa- 1 , implying that a transpiration rate of
1 mm h- 1 would cause 'Pstem to fall to about 1 MPa below 'P soil ' For a
sparsely-rooted crop (say, La = 4 km m- 2 ), 'Pstem would have to decrease to
-2.5 MPa in order to sustain the same rate of transpiration. In practice, it
is unlikely that 'Pstem would fall so low, as evidence was presented earlier
that the minimum leaf water potential observed in potato crops rarely falls
below about -1.5 MPa.
The concept of a 'minimum leaf water potential' has led to speculation
that there is, in effect, an upper limit to the rate of transpiration (Tmax) that
potato crops can sustain (Campbell et al., 1976). An extension of this idea
is that the maximum sustainable rate of transpiration for a well-watered
crop depends on the ability of the plant to conduct water, and, therefore,
on the length of the root system. For example, a 'fi'lst approximation'
calculation based on the values for gbg presented in Fig. 5.10 suggests that
if 'Pstem is effectively restricted to a minimum of, say, -1.0 MPa (bearing in
mind that 'Pstem is about two-thirds of 'P leaf), then Tmax would be
about 1 mm h- 1 in a crop with La of 10 km m- 2 , growing in wet soil (where
'P soil =0). If La were restricted to 4 km m-2 , the value of Tmax would be
only 0.4 mm h- 1 , which would correspond typically to the mid-day potential
evaporation during a day with about 4 mm mean daily potential evapora-
tion (assuming a peak:mean ratio of 2.4).
The calculations above were based on a nominal minimum stem water
potential of -1 MPa. Evidence was presented earlier that incipient
stomatal closure in potatoes is often observed at much higher tissue water
potentials (e.g. when 'l'leaf < ':"'0.6 MPa). Hence stomatal closure might be
expected as a consequence of inadequate leaf water supply in plants
exposed to quite modest potential rates of evaporation, even if the soil is
kept wet. Any impediment to root growth (e.g. through soil compaction or
the presence of a plough pan) is likely to impose a serious restriction to
transpiration (and, hence, growth) as a direct consequence of the increased
resistance to water flow through the plant, especially in environments with
large evaporative demand.
5.5.4 Root distribution and soil water stress

Soil water potential is rarely uniform throughout the profile, so j,t is not
immediately obvious how to evaluate 'effective mean soil water potential'
('I'soil in equation (5.11); equivalent to the intercept at point A in Fig. 5.9)
when roots in different parts of the profile are exposed to widely differing
soil water potential. Campbell (1985, Chapter 11) discusses how 'l'soil can
be evaluated from knowledge of the distribution of roots and soil water
potential through the soil profile. His analysis suggests that in soil which is
not too dry (i.e. in which low soil hydraulic conductivity is not restricting
water flow severely), a 'first approximation' for 'l'soil is given by:
'l'soil = L 'l'si Lfi (5.13)
where 'l'si is the soil water potential in layer 'i' of the soil profile and Lfi is
the fraction of the total root length present in layer 'i'. Hence the effective
soil water potential experienced by the crop is strongly weighted in favour
of the value of soil water potential in the zone of most dense rooting. The
use of equation (5.13) is illustrated in Table 5.5, which shows the values of
'l'soil calculated for hypothetical crops with contrasting root distributions in
the same 'two-layer' soil profile. 'Crop l' has the whole of the root system
confined to the upper (drier) layer, and experiences a very much lower
value of 'l'soil than 'Crop 3', which has most roots in layer 2.
The susceptibility of shallow-rooted crops to low 'l'soil is exacerbated by
the tendency for the soil water potential to decrease most rapidly in parts
Table 5.5 Example calculations to illustrate the impact of root

distribution on the effective mean soil water potential (ljIsoil)
Layer Soil water Fraction of total root length in each layer
potential
(MPa) Crop 1 Crop 2 Crop 3
1 -0.20 1.0 0.7 0.3
2 -0.02 0.0 0.3 0.7
'l'soil (MPa) -0.2 -0.146 -0.074
of the profile where roots are concentrated. Since van den Honert (1948).
there has been much refinement in the use of electrical analogues to
simulate water fluxes and potentials in the soil/plant/atmosphere con-
tinuum. Figure 5.11 shows the result of one such set of simulations in which
the decrease in 'I'soil as soil water is depleted below a nominal field capacity
is compared for three crops with contrasting distributions of roots. These
simulations demonstrate the tendency for 'I'soil to decrease rapidly as the
profile dries when roots are concentrated in the upper part of the profile.
Figure 5.11 also illustrates that it is the relative amounts of roots in each
layer that controls the progress Of'l'soil as the soil water deficit increases.
For example, 'I'soil decreases most rapidly in 'Crop 3' (which has the
greatest proportion of roots in the upper 25 cm), even though the overall
length of the root system is substantially greater than for 'Crop 2', and the
maximum rooting depths of Crops 2 and 3 are similar.
Section 5.4 presented evidence that potatoes tend to produce shallow,
surface-oriented root systems, and the analysis above illustrates how this
Cumulative water extraction (mm)
o 10 20 30 40 50
........ :;-.:;.::.::.~............ "
................ ""'"
-100
Root length density (cm cm-3 ) ...•.. '"
o 2 4 ..... ",
...... "
~.<.d.········ ...•... ''
.,,'" \.... ", , ®
'
00 "/
-200 50 ~I .....
\\
I
E
-S
100 '\\\
'\
.t::
a.
Q)
Cl
-300 150
\
Figure 5.11 The results of a simulation to demonstrate the iUlpact of root

distribution on the change in 'l'soil as water is depleted from a profile that has an
initial soil water suction of 20 kPa. The inset shows the root length density profiles
for the simulated crops. The model was based on that described by Campbell (1985,
Chapter 11), and simulates a crop with complete ground cover (where evaporation
from the soil surface is negligible) growing in a soil with the water retention
properties of a loamy sand.
will result inevitably in plants being exposed to relatively severe soil water
stress at modest soil water deficits. Also, the shallow rooting habit may be
implicated in direct responses of stomata to soil drying, as discussed earlier
in this section.
5.5.5 Water stress and canopy expansion

Water deficits influence strongly the production of leaf by affecting the
numbers, size and duration of leaves (Jefferies, 1989). In potatoes, as in
many other crops (Boyer, 1970) the processes of leaf and stem extension
are highly sensitive to tissue water status by comparison with photo-
synthesis and stomatal conductance. The widely-quoted work of Gandar
and Tanner (1976) showed that the rate of leaf extension decreased linearly
as leaf water potential decreased, with negligible extension occurring when
'P leaf fell below about -0.5 MPa: hence leaf growth occurred pre-
dominantly at night. A similar linear relation was reported by Jefferies
(1989). His results suggest that the average daily leaf extension did not fall
to zero until midday values of 'Pleaf reached about -1 MPa, though care
has to be taken when interpreting this result because diurnal changes in
leaf water status are not accounted for. The extreme sensitivity of leaf
expansion to tissue water deficit means that the initial response to mild
water stress is a reduction in the rate of production of leaf area (see, for
example, Bunyolo, 1987), so that the leaves that remain are s.till able to
operate near maximum efficiency (Jefferies and MacKerron, 1989).
Gander and Tanner emphasized the importance of the rate of nocturnal
rehydration of tissue as a mechanism controlling the production of leaf
area, although this is a subject that has since received little attention. It is
notable that the few measurements made of leaf water potentials during
the night suggest that 'Pleaf usually fails to rise above about -0.2 MPa,
and is often much lower (Campbell et al., 1976; Gander and Tanner, 1976;
Vos and Groenwold, 1988), even though 'Pson is much closer to zero.
Hence even in wet soil it is likely that leaf area production is mediated to
some extent by tissue water status. The reasons for the limited ability of
crops to rehydrate fully at night are poorly understood. It has been
suggested that a contributory factor is probably that the resistance to water
movement through plant tissue increases by a factor of 10 at very low flow
rates, and that this interfres with nocturnal rehydration (Campbell et al.,
1976).
5.6 WATER SUPPLY AND TUBER QUALITY
There is an extensive literature on the responses of potatoes to irrigation

both in relation to yield and to yield quality (for reviews see Salter and
Goode, 1967; van Loon, 1981). Harris (1978) reviewed several experiments
Water supply and tuber quality 237
conducted in North America and Europe to determine the yield response
to irrigation and the efficiency of water use (dry matter production per unit
of irrigation water applied). The experiments selected were those in which
an irrigated control was included and were mainly confined to sandy-
textured soils. Table 5.6 shows the frequency with which the yield and the
efficiency of water use were maximized by a particular irrigation treatment;
the treatments correspond approximately to irrigating when 75%, 50% and
25% of the available soil water had been depleted. Although total yield
was greatest with most frequent irrigation, ware yield was most frequently
greatest at the intermediate level of irrigation. The efficiency of water use
was also greatest at the intermediate level of irrigation and thus irrigating
when about 50% of the available soil water has been depleted most
frequently maximized the return per unit of water applied. Harris's review
also pointed out that the number of irrigation applications needed to
replenish water depleted to 25%, 50% and 75% of that available was 6.7,
4.4 and 3.5 per season respectively, so that irrigation at 50% depletion
offered considerable practical and economic advantages by reducing the
number of irrigations without substantially reducing marketable yield.
Table 5.6 The frequency with which yield or water use efficiency
was maximized by irrigation regime (from Harris, 1978)
Number of Frequency of irrigation
comparisons
Most Intermediate Least Shared
a b c
Number of occasions maximized
Yield
Total 22 11 7 4 0
Ware 16 4 7 4 1(alb)
Water use efficiency

Total 22 6 11 4 1(a/c)
Ware 16 3 7 5 1(alb)
Despite the many experiments of the type reported by Harris, there have
been few attempts until recently to distinguish the underlying processes
leading to the yields measured at harvest. Too frequently, irrigation
experiments have treatments that are not strictly related to either atmo-
spheric demand for water or. soil water storage nor are the treatments
applied at defined stages of crop development. These failings mean
that results are often apparently contradictory making generalization
impossible. During the 1980s, however, several workers have sought
to understand the effects of drought on the underlying processes
affecting the formation of tubers, the numbers of tubers produced and
the growth of tubers so that yield and yield quality might be predicted
better.
A key problem in relation to water supply is to understand its effect on
the number of tubers produced. For the reasons outlined previously, the
results are often contradictory if irrigation experiments alone are relied on
although the consensus is that dry soil conditions at or about the time of
tuber initiation reduce the number of tubers per plant (van Loon, 1981).
MacKerron and Jefferies (1986) conducted experiments in partially con-
trolled conditions to identify the period when tuber number was sensitive
to drought and to quantify the relation between tuber number and the
intensity of drought. In glasshouse studies conducted over 2 years, Maris
Piper potatoes were grown in compost maintained by daily irrigation at a
soil matric potential of -10 kPa (water content of 42% v/v) except when
drought was imposed. Water was withheld for varying periods at three
principal developmental stages, namely 50% emergence, tuber initiation,
and the s'inall tuber stage. Only drought imposed at 50% emergence had an
effect on th~umber of tubers present at harvest and even severe drought
had no effect when imposed at tuber initiation. The number of tubers per
stem (N) was linearly related to the number of days (D) when the soil
matric potential was <-25 kPa by N = 7.20(±0.21) - 0.16(±0.02) D
(,-2 = 0.85); the use of a simple count of time and a threshold value for soil
matric potential was found to be less complicated than, and as good as, a
time-integrated measure of soil matric potential. Some caution is necessary
in applying these results to other soils: -25 kPa in the compost equates
with a volumetric water content of about 33% and represents a depletion
of about 82% of the available soil water. In relation to other soils this
seems a severe depletion. Moreover, as MacKerron and Jefferies recog-
nized, the choice of a single value of soil matric potential ignores the fact
that the water stress experienced by a plant depends on the balance
between water supply and the demand for water by the atmosphere. Their
results on the effects of drought on yield and frequency of distribution of
tuber size were less clear cut although the depression of yield was greatest
in those plants that were largest when the drought was imposed.
Struik and van Voorst (1986) demonstrated that drought in the environ-
ment of the roots or of the stolons might produce different effects and
hence contribute to an explanation for some of the contradictory results
found by other workers. In their study stolons and tubers were grown in
sand/perlite mixture and the roots in nutrient solution. Drought was
effected by varying the water content of the sand/perlite and the osmotic
potential of the nutrient solution with polyethylene glycol. Their results
indicated that 'drought' in the root environment had no effect on the
numbers of stolons or tubers but the yield was reduced whereas drought
around the stolons enhanced stolon and tuber initiation and gave a minor
reduction in yield. Drought in both media slightly stimulated the initiation
of stolons and tubers and reduced yields. Although changing the osmotic
Water supply and tuber quality 239
potential around roots is not strictly the same as drought and may cause
other physiological changes, the findings suggest that shortage of water
around the stolons should be distinguished from that within the root zone
as a whole.
Once tubers have been initiated, their subsequent growth may also be
affected by drought so that not only are yields reduced but the quantity of
marketable tubers is also decreased. For example, Robins and Domingo
(1956) found reductions of 30% in total yields but of 58% in US No.1
yields of Russet Burbank potatoes when crops were subjected to drought.
Field experiments at the Scottish Crop Research Institute in two consecutive
years with up to five cultivars of main crop potatoes were undertaken to
determine the effects of drought on yields and the distribution of tuber
sizes (Jefferies and MacKerron, 1987a; MacKerron and Jefferies, 1988).
Irrigation was scheduled to maintain the soil moisture deficit at <30 mm
and in the droughted treatments soil moisture deficits were 107 mm and 94
mm in mid-August in the 2 years. Drought reduced the total dry matter
production and tuber yield of all cultivars and increased the concentration
of dry matter in the tubers. For example, tuber yield of Maris Piper was
reduced by an average of 47% over the 2 years while the concentration of
dry matter in the tubers was increased by 19% (see also Shimsi and
Susnoschi, 1985). The droughts also decreased the size oftubers so that the
centres of distribution of tuber sizes were moved in the direction of smaller
potatoes by about 7-10 mm (Fig. 5.12). For example, in 1984 only 30% of
Maris Piper yield was in tubers <45 mm (45 mm has been the limit of ware
grade in the UK since 1985) when the crop was irrigated but drought
resulted in 85% <45 mm. Drought, then, not only reduced the tuber yield
but also reduced the fraction of the yield reaching ware (>45 mm) and
table (40-80 mm) grades. Further analysis of their results (MacKerron et
al., 1988) showed that yield and number of tubers gave a unique tuber-size
distribution for each cultivar irrespective of the soil moisture deficit
experienced. They concluded that because the proportion of the tuber
yield which reaches a marketable size is dependent on the mean tuber size
(and thus on both total tuber yield and the number of tubers) then, where
drought is common, it would be most sensible to grow those cultivars that
produce smaller numbers of tubers (assuming all other factors affecting
choice of cultivar are equal).
Other defects such as knobbly or dumb-bell shaped tubers and cracked
tubers can also be caused by short periods of water deficit although the
reasons for this are only partially understood (van Loon, 1986). Moorbyet
al. (1975) found that defects in shape were induced after the water
potential of the tubers was -50 kPa for a period of 3 days and that tubers
of droughted plants had higher contents of total sugars and reducing sugars
compared with those of well-watered plants. Cracking of tubers during
growth is not found in all cultivars and is sometimes associated with
drought in mid-season. Jefferies and MacKerron (1987b) sought to relate
1.0 o ••
o •
o •
o •
o ••I i .
o. •
20 60 100
Tuber size (mm)
Figure 5.12 The effect of drought on the distribution of yield in size classes as a
cumulative proportion of total tuber yield for cultivar Maris Piper; irrigated CO),
and droughted ce). CFrom MacKerron and Jefferies, 1988.)
the cracking of tubers of two cultivars known to be susceptible to cracking

to the weather conditions over a 3-year period. In the first 2 years, one
cultivar (Guardian) had few cracked tubers «1%) whereas the other
(Record) had 4% and 12% which appeared with the ending of severe soil
moisture deficit (99 mm and 123 mm in late August/early September). The
cracks in tubers of Record were extensive but rarely more than 5 mm deep
and tended to fill slowly so that the incidence of cracking appeared to
decrease with time. In the final year which was wet (maximum soil
moisture deficit 20 mm), few tubers of Record cracked but up to 14% of
Guardian tubers had cracks which differed from those of Record in that
there was usually only one large cleavage 10 mm across, 15-25 mm deep,
which did not re-fill. On the basis of these observations, Jefferies and
MacKerron (1987b) concluded that two forms of cracking can occur, one
associated with re-wetting of the soil after growth had temporarily ceased
(cv. Record) and the other with rapid tuber growth and high turgor
pressure (cv. Guardian). The causes of other forms of growth defect
remain to be elucidated.
There is widespread acceptance of the notion that some potato varieties
are more tolerant of drought than others and many national organizations
such as the National Institute for Agricultural Botany in the UK rank
cultivars on a scale of drought tolerance (based on ware yield only and
without account of the number or size of tubers). However, the experi-
mental evidence for such differences is frequently inconsistent or valid only
in particular circumstances. Steckel and Gray (1979) found that responses
varied markedly from year to year although the variety Pentland Crown
consistently gave slightly greater yields of tubers (but much greater yields
of marketable (>40 mm) tubers) than the variety King Edward over a
range of soil water conditions. This confirmed the difference in drought
tolerance commonly ascribed to these cultivars although results for two
others (Majestic and Maris Piper) were inconclusive. The results
References 241
of Jefferies and MacKerron (1987a) in which the so-called drought-
susceptible cultivars Maris Piper and Record were compared with the
drought-tolerant cultivars Pentland Crown, Pentland Dell and Desiree
were inconclusive with tuber yield ranging from 70% to 81 % of the
irrigated crops and marketable tubers (>40 mm) from 60% to 75% of the
irrigated crops (Table 5.7). One reason for the apparent inconsistency of
results is that few experiments have determined the processes contributing
to yield under dry conditions. Lynch and Tai (1989) studied the response of
eight cultivars to water stress and found significant differences in yield but
also significant differences in the components of yield between cultivars.
Shortage of water at tuber initiation had a greater influence on yield than
stress during tuber growth in three cultivars but three others were more
sensitive during tuber growth. These results suggest that optimum manage-
ment of limited supplies of water may be cultivar dependent. Despite many
reviews of the subject (e.g. van Loon, 1986) much work remains to be done
to understand the responses of cultivars to drought before reliable pro-
cedures are available to breeders to enable them to screen for drought
resistance.
Table 5.7 Differences between cultivars in their response to drought (from

Jefferies and MacKerron, 1987a)
Yield of Pentland Pentland Desiree Maris Record
tubers Crown Dell Piper
Total (t ha- 1) 49.7 47.0 43.8 53.4 47.2
(irrigated)
Total (droughted as 75 81 74 70 77
% irrigated)
Ware (droughted as 75 75 66 60 73
% irrigated)
ACKNOWLEDGEMENTS
We are grateful to Dr R. Jefferies and Dr J. Vos for their constructive

criticism of an earlier draft of this chapter.
REFERENCES
Ackerson, R.C., Krieg, D.R., Miller, T.D. and Stevens, R.G. (1977) Water
relations and physiological activity of potatoes. J. Amer. Soc. Hort. Sci., 102,
572.
Allen, E.l. and Scott, R.K. (1980) An analysis of growth of the potato crop. J.
Agric. Sci., Camb., 94,583-606.
Asfary, A.F., Wild, A. and Harris, P.M. (1983) Growth, mineral nutrition and
water use by potato crops. I. Agric. Sci., Camb., 100,87-101.
Barraclough, P.B. and Leigh, R.A. (1984) The growth and activity of winter wheat
roots in the field: the effect of sowing date and soil type on root growth of high-
yielding crops. I. Agric. Sci., Camb., 103,59-74.
Bierhuizen, J.F. and Slatyer, R.O. (1965) Effect of atmospheric concentration of
water vapour and CO 2 in determining transpiration-photosynthesis
relationships of cotton leaves. Agric. Meteorol., 2, 259-70.
Blackman, P.G. and Davies, W.J. (1985) Root to shoot communication in maize
plants of the effects of soil drying. 1. Exp. Bot., 36, 39-48.
Bodlaender, K.B.A. (1986) Effects of drought on water use, photosynthesis and
transpiration of potatoes. 1. Drought resistance and water use, in Potato
Research of Tomorrow (eds A.G.M. Beekman and K.M. Louwes), Pudoc,
Wageningen, pp. 36-43.
Boone, F.R., de Smet, L.A.H. and van Loon, C.D. (1985) The effect of a
ploughpan in marine loam soils on potato growth. 1. Physical properties and
rooting patterns. Potato Res., 28, 295-314.
Boyer, J.S. (1970) Leaf enlargement and metabolic rates in corn, soybean and
sunflower at various leaf water potentials. Plant Physiol., 46, 1056-62.
Boyer, J.S. (1985) Water transport in plants. Ann. Rev. Pl. Physiol., 36, 473-516.
Boyer, J.S. (1989) Water potential and plant metabolism: comments on Dr PJ.
Kramer's article 'Changing concepts regarding plant water relations', volume
11, number 7, pp. 565-8, and Dr J.B. Passioura's response, pp. 569-71. Plant
Cell Env., 12, 213-16.
Bristow, K.L., Campbell, G.S. and Calissendorf, C. (1984) The effects of texture
on the resistance to water movement within the rhizosphere. Soil Sci. Soc. Am.
I., 48,266-70.
Bunyolo, A.M. (1987) Effects of fertilizer nitrogen and water supply on the growth
and yield of potato crops. PhD Thesis. University of Reading.
Burstall, L. and Harris, P.M. (1983) The estimation of percentage light inter-
ception from leaf area index and percentage ground cover in potatoes. I. Agric.
Sci., Camb., 100, 241-4.
Campbell, G.S. (1985) Soil Physics with BASIC, Elsevier, Amsterdam.
Campbell, M.D., Campbell, G.S., Kunkel, R. and Papendick, R.1. (1976) A
model describing soil-plant-water relations for potatoes. Am. Potato I., 53,
431-41.
Cary, J.W. and Wright, J.L. (1971) Response of plant water potential to the
irrigated environment of southern Idaho. Agron. I., 63, 691-5.
Coleman, W.K. (1986) Water relations of the potato (Solanum tuberosum L.)
cultivars Raritan and Shepody. Am. Potato I., 63, 263-76.
Corey, A.T. and Blake, G.R. (1953) Moisture available to various crops in some
New Jersey soils. Soil Sci. Soc. Am. Proc., 17,314-17.
Day, W., Lawlor, D.W. and Day, A.T. (1987) The effect of drought on barley
yield and water use in two contrasting years. Irrig. Sci., 8, 115-30.
Durrant, M.J., Love, B.J.G., Messem, A.B. and Draycott, A.P. (1973) Growth of
crop roots in relation to soil moisture extraction. Ann. Appl. BioI., 74, 387-94.
Evans, S.A. and Neild, J.R.A. (1981) The achievement of very high yields of
potatoes in the U.K. I. Agric. Sci., Camb., 97, 391-6.
Farquhar, G.D. and Richards, R.A. (1984) Isotopic composition of plant carbon
References 243
correlates with water-use efficiency of wheat genotypes. Aust. J. Plant. Physiol.,
11,539-52.
Farquhar, G.D., O'Leary, M.H. and Berry, J.A. (1982) On the relation between
carbon isotope discrimination and the intercellular carbon dioxide concentra-
tion in leaves. Aust. 1. Plant. Physiol., 9, 121-37.
French, B.K. and Legg, B.J. (1979) Rothamsted irrigation 1964-76. J. Agric. Sci.,
Camb., 92, 15-37.
French, B.K., Long, J.F. and Penman, H.L. (1973) Water use by farm crops. II.
Spring wheat, barley, potatoes (1969); potatoes, beans, kale (1968). Report
Roth. Exp. Stn for 1972, part 2, pp. 43-61.
Gandar, P.W. and Tanner, C.B. (1976) Leaf growth, tuber growth and water
potential in potatoes. Crop Sci., 16, 534-8.
Greenwood, D.J., Gerwitz, A., Stone, D.A. and Barnes, A. (1982) Root
development of vegetable crops. PI. Soil., 68, 75-96.
Gregory, P.J. (1988) Root growth of chickpea, faba bean, lentil and pea and effects
of water and salt stresses, in World Crops: Cool season food legumes (ed. RJ.
Summerfield), Martinus Nijhoff, Dordrecht, pp. 857-67.
Gregory, P.J., McGowan, M., Biscoe, P.V. and Hunter, B. (1978) Water relations
of winter wheat. 1. Growth of the root system. J. Agric. Sci., Camb., 91,
91-102.
Hamblin, A. and Tennant, D. (1987) Root length density and water uptake in
cereal and grain legumes: how well are they correlated? Aust. J. Agric. Res., 38,
513-27.
Harris, P.M. (1978) Water, in The Potato Crop: The scientific basis for improve-
ment (ed. P.M. Harris), Chapman and Hall, London, pp. 244-77.
Hubick, K.T., Farquhar, G.D. and Shorter, R. (1986) Correlation between water-
use efficiency and carbon isotope discrimination in diverse peanut (Arachis)
germplasm. Aust. J. Plant Physiol., 13, 803-16.
International Potato Center (1986) Annual Report C1P 1985, Lima, Peru, 86pp.
potato. Outlook on Agric., 4, 211-17.
Jefferies, R.A. (1989) Water-stress and leaf growth in field-grown crops of potato
(Solanum tuberosum L.). J. Exp. Bot., 40, 1375-81.
Jefferies, RA. and MacKerron, D.K.L. (1987a) Aspects of the physiological basis
of cultivar differences in yield of potato under droughted and irrigated
conditions. Potato Res., 30,201-17.
Jefferies, RA. and MacKerron, D.K.L. (1987b) Observations on the incidence of
tuber growth cracking in relation to weather patterns. Potato Res., 30,
613-23.
Jefferies, R.A. and MacKerron, D.K.L. (1989) Radiation interception and growth
of irrigated and droughted potato (Solanum tuberosum). Field Crops Res., 22,
101-12.
Kaufmann, M.R (1976) Water transport through plants: current perspectives, in
Transport and Transfer Processes in Plants (eds J.F. Wardlaw and J.B.
Passioura), Academic Press, New York, pp. 313-27.
Kramer, P.J. (1988) Changing concepts regarding plant water relations. Plant Cell
Env., 11, 565-8.
Landsberg, J.J. and Fowkes, N.D. (1978) Water movement through plant roots.
Ann. Bot., 42,493-508.
Lesczynski, D.B. and Tanner, C.B. (1976) Seasonal variation of root distribution
of irrigated, field-grown Russet Burbank potato. Am. Potato J .., 53, 69-78.
Levy, D. (1983) Varietal differences in the response of potatoes to repeated short
periods of water stress in hot climates. 1. Turgor maintenance and stomatal
behaviour. Potato Res., 26, 303-13.
Lynch, D.R. and Tai, G.C.C. (1989) Yield and yield component response of eight
potato genotypes to water stress.. Crop Sic., 29, 1207-11.
MacKerron, D.K.L. and Jefferies, R.A. (1986) The influence of early soil moisture
stress on tuber numbers in potato. Potato Res., 29,299-312.
MacKerron, D.K.L. and Jefferies, R.A. (1988) The distributions of tuber sizes in
droughted and irrigated crops of potato. I. Obs.ervations on the effect of water
stress on graded yields from differing cultivars. Potato Res., 31, 269-78.
MacKerron, D.K.L. and Waister, P.D. (1985) A simple model of potato growth
and yield. I. Model development and sensitivity analysis. Agric. For. Meteorol.,
34,241-52.
MacKerron, D.K.L., Marshall, B. and Jefferies, R.A. (1988) The distributions of
tuber sizes in droughted and irrigated crops of potato. II. Relation between size
and weight of tubers and the variability of tuber-size distributions. Potato Res.,
31,279-88.
McDermott, N. and Ivins, J.D. (1955) Rainfall as a factor influencing the yields of
potato crops. NAAS Quart. Rev., 27, 106-8.
Mengel, D.B. and Barber, S.A. (1974) Development and distribution of the corn
root system under field conditions. Agron. J., 66, 341-4.
Millar, B.D. and Denmead, O.T. (1976) Water relations of wheat leaves in the
field. Agron. J., 68, 303.
within the potato (Solanum tuberosum L.) crop, in relation to nitrogen
application. J. Agric. Sci., Camb., 107,421-9.
Ministry of Agriculture, Fisheries and Food (1967) Potential Transpiration,
Technical Bulletin No 16, HMSO, London.
Monteith, J.L. (1990) Steps in crop climatology, in Proc. Int. Conf. Dryland Agric.
(eds P.W. Unger, W.R. Jordan, T.V. Sneed and R.W. Jensen) Texas
Agricultural Experimental Station, pp. 273-82.
Moorby, J., Munns, R. and Walcott, J. (1975) Effect of water deficit on photo-
synthesis and tuber metabolism in potatoes. Aust. J. Plant Physiol., 2,323-33.
Munns, R. and Pearson, c.J. (1974) Effect of water deficit on translocation of
carbohydrates in Solanum tuberosum. Aust. J. Plant Physiol., 1, 529-37.
Newman, E.1. (1976) Water movement through root systems. Phil. Trans. R. Soc.
Land. B., 273, 463-78.
Ovaa, I. and de Smet, L.A.H. (1984) Root growth in relation to soil profile and
tillage system, in Experiences with Three Tillage Systems on a Marine
Loam Soil II. 1976-1979, Agric. Res. Rep. 925, Pudoc, Wageningen, pp.
72-88.
Parker, C.J., Carr, M.K.V., Jarvis, N.J. et al. (1989) Effects of subsoil loosening
and irrigation on soil physical properties, root distribution and water uptake of
potatoes (Solanum tuberosum). Soil Tillage Res., 13,267-85.
Passioura, J.B. (1988) Response to Dr P.J. Kramer's article 'Changing concepts
regarding plant water relations', volume 11, number 7, pp. 565-8. Plant Cell
Env., 11, 569-71.
References 245
Pearcy, R.W. and Ehleringer, J. (1984) Comparative ecophysiology of C3 and C4
plants. Plant Cell Env., 7, 1-13.
Peeler, e.H., Harvey, P.N. and Rosser, W.R. (1966) Effect of irrigation on yield
and tuber diseases of maincrop potatoes. Expl. Husb., 14, 30-42.
Penman, H.L. (1970) Woburn irrigation, 1960-8. IV. Design and interpretation. 1.
Agric. Sci., Camb., 75, 69-73.
Penman, H.L. (1971) Irrigation at Woburn. Report R,oth. Exp. Stnfor 1970, part 2,
pp.147-70.
Rijtema, P .E. and Endrodi, G. (1970) Calculation of production of potatoes. Neth.
1. Agric. Sci., 18,26-36.
Ritchie, J.T. (1972) Model for predicting evaporation from a row crop with
complete cover. Water Resources Res., 8, 1204-13.
Robins, J.S. and Domingo, C.E. (1956) Potato yield and tuber shape as affected by
severe soil-moisture deficits and plant spacing. Agron. I., 48, 488-92.
Rutherfoord, R.J. and de Jager, J.M. (1975) Water s,tatus and stomatal behaviour
of BPI potatoes (Solanum tuberosum L.) and their effect upon yield. Crop
Production, 4, 125.
Salter, P.J. and Goode, J.E. (1967) Crop responses to water at different stages of
growth. Commw. Bureau Hort., East Mailing, 2, 93-7.
Sands, R., Ficus, E.L. and Reid, C.P.P. (1982) Hydraulic properties of pine and
bean roots with varying degrees of suberization, vascular differentiation and
mycorrhizal infection. Aust. 1. Plant Physiol., 9, 559--69.
Schulze, E.D., Steudle, E., Gollan, T. and Schurr, U. (1988) Response to Dr P.J.
Kramer's article 'Changing concepts regarding plant water relations', volume
11, number 7, pp. 565-8. Plant Cell Env., 11,573--6.
Seaton, K.A., Landsberg, J.J. and Sedgely, R.H. (1977) Transpiration and leaf
water potentials of wheat in relation to changing soil water potential. Aust. 1.
Agric. Res., 28, 355-367.
Shimsi, D., Shalhavet, J. and Meir, T. (1983) Irrigation regime effects on some
physiological responses of potato. Agron. I., 75, 262-7.
Shimsi, D. and Susnoschi, M. (1985) Growth and yield studies of potato develop-
ment in a semi arid region. 3. Effect of water stress and amounts of nitrogen top
dressing on physiological indices and on tuber yield and quality of several
cultivars. Potato Res., 28, 177-91.
Stark, J.C. and Wright, J.L. (1985) Relationship between foliage temperature and
water stress in potatoes. Am. Potato I., 62, 59--68.
Stark, J.C., Ojala, J.e. and McCann, I.R. 1987. Estimation of diurnal changes in
potato leaf water potential under irrigated conditions. 1. Amer. Soc. Hort. Sci.,
112,825.
Steckel, J.R.A. and Gray, D. (1979) Drought tolerance iiil potatoes. 1. Agric. Sci.,
Camb., 92, 375-81.
Stone, D.A. (1982) The effects of subsoil loosening and deep incorporation of
nutrients on yield of broad beans, cabbage, leek, potatoes and red beet. 1.
Agric. Sci., Camb., 98, 297-306.
Struik, P.C. and van Voorst, G. (1986) Effects of drought on the initiation, yield
and size distribution of tubers of Solanum tuberosum L. cv. Bintje. Potato Res.,
29, 487-500.
Tanner, C.B. (1981) Transpiration efficiency of potato. Agron. I., 73, 59--64.
Tanner, C.B. and Sinclair, T.R. (1983) Efficient water use in crop production:
research or re-search, in Limitations to Efficient Water Use in Crop Production
(eds H.M. Taylor, W.R. Jordan and T.R. Sinclair), American Society of
Agronomy, Madison, pp. 1-27.
Taylor, H.M. and Klepper, B. (1971) Water uptake by cotton roots during an
irrigation cycle. Aust. 1. BioI. Sci., 24, 853-9.
Teare, I.D. and Peet, M.M. (1983) Crop Water Relations, John Wiley and Sons,
New York.
van Loon, C.D. (1981) The effect of water stress on potato growth, development,
and yield. Am. Potato I., 58, 51~9.
van Loon, C.D. (1986) Drought, a major constraint in potato production and
possibilities for screening for drought resistance, in Potato Research of
Tomorrow (eds A.G.B. Beekman, K.M. Louwes, L.M.W. Dellaert and A.E.F.
Neele), Pudoc, Wageningen, pp. 5-16.
van den Honert, T.H. (1948) Water transport in plants as a catenary process. Disc.
Farad. Soc., 3, 146--53.
Vos, J. and Groenwold, J. (1986) Root growth of potato crops on a marine-clay
soil. Pl. Soil, 94, 17-33.
Vos, J. and Groenwold, J. (1988) Water relations of potato leaves. 1. Diurnal
changes, gradients in the canopy, and effects of leaf-insertion number, cultivar
and drought. Ann. Bot., 62,363-71.
Vos, J. and Groenwold, J. (1989a) Characteristics of photosynthesis and conduct-
ance of potato canopies and the effects of cultivar and transient drought. Field
Crops Res., 20,237-50.
Vos, J. and Groenwold, J. (1989b) Genetic differences in water-use efficiency,
stomatal conductance and carbon isotope fractionation in potato. Potato Res.,
32, 113-21.
Vos, J. and Oyarzun, P.J. (1987) Photosynthesis and stomatal conductance of
potato leaves - effects of leaf age, irradiance and leaf water potential. Photosyn.
Res., 11, 25~4.
Walker, G.K. (1986) Transpiration efficiency of field-grown maize. Field Crops
Res., 14, 29-38.
Wolfe, D.W., Fereres, E. and Voss, R.E. (1983) Growth and yield response of two
potato cultivars to various levels of applied water. [rrig. Sci., 3, 211-22.
Wong, S.c., Cowan, I.R. and Farquhar, G.D. (1979) Stomatal conductance
correlates with photosynthetic efficiency. Nature, Land., 282, 42~.
CHAPTER 6
Seed tuber production and

management
E.]. Allen~ P.}. O'Brien and D. Firman
6.1 INTRODUCTION
The usual means of propagation for potato crops throughout the world is
the vegetative seed tuber which may be planted whole, as in most of
Europe, or after cutting into pieces, as in North America and parts of
Spain. Seed tubers are variable in size ranging from 25 to 60 mm (10-150 g)
when planted whole and up to 100 mm or more (300-400 g) when cut. They
are predominantly water (around 80%) and do not keep even at low
temperatures from one crop year to another, although in favourable
climates two crops can be produced within one year from one seed lot.
Seed tubers can be infected with numerous virus diseases and carry many
fungal diseases which may affect ware crop growth and the health of the
progeny tubers. For effective potato production the availability of an
annual supply of healthy tubers is an essential requirement. In this chapter
methods of seed production and storage and statutory regulation relating
principally to the UK industry are discussed . References are also made to
other seed production problems in contrasting environments. A brief
consideration is also given to the use of true potato seed (TPS) both for
primary propagation and as a means of producing relatively disease-free
seed tubers in some environments as the majority of virus and fungal
diseases are not transmitted through true seed.
6.2 GROWTH AND DEVELOPMENT OF THE SEED TUBER
At harvest seed tubers are usually dormant (rest period of Emilsson, 1949)
and do not show any bud growth even in favourable temperatures.
Dormancy ends at some stage during winter storage in N. W. European
latitudes for all commercial varieties and sprout growth begins if temper-
atures are high enough: >2 to 4°C according to variety. The timing of the
248 Seed tuber production and management
end of dormancy is affected by seed-crop husbandry and has a major
influence on the duration of sprout growth which together with storage
temperatures determines the amount of sprout growth at re-planting. This
affects the number and type of stems produced (Allen et al., 1979), the
extent and duration of the leaf surface (O'Brien et al., 1983), number of
tubers and ultimate yield. Thus, in potatoes there can be large influences of
the seed production year and storage te.mperatures on the following ware
crop and in many ways the crop should be considered as biennial.
An understanding of the re-growth of seed tubers is fundamental to
analysis of almost all aspects of potato production. It is logical to begin
with the basic morphology of the seed tuber and follow the processes of
growth and development through to final yield. The implications of some
of these effects for other aspects of potato agronomy are considered in
other chapters, particularly the following chapter on plant density.
6.2.1 Seed tuber morphology

Seed tubers show a wide variation in size and possess a variable number of
growing points (buds) arranged in groups (eyes) over their surface. The
eyes are arranged in the phyllotactic spiral over the tuber's surface (Fig.
6.1). Milthorpe (1963) explained that the apical bud is part of the primary
vegetative axis of the stolon and the lateral buds arising from it are
secondary axes, some of which become main buds in the eyes of the tuber
as the wave of expansion described by Krijthe (1946) passes from the basal
to the apical end. The number of lateral buds which separate from the
apical bud and form independent eyes depends upon the degree of
development of the tuber and Fig. 6.2 shows that the number of eyes per
seed tuber increases with tuber size but at a decreasing rate. There are
Rose end
Insert of an eye
Apical bud --F----4L~1 ~ Lateral buds. Main B d I
complex Leaf scar ~ ~ u sea es
i ~~t3ral __~J7 k~~eral
Leafscar---------- .-~
,,,--,\
I '• .,., ,
\~
, __ "I Insert
_ _.r----Stolon scar
Heel end
Figure 6.1 A potato tuber showing anatomical details. (After Gray and
Bleasdale, 1972).
Growth and development of the seed tuber 249
differences between varieties in number of eyes and Cara appears to have
substantially more eyes per seed tuber than most varieties. Each main bud
subtends a number of other buds, part of the tertiary axis, so that each eye
contains many buds, each of which is a potential growing point. Overall,
there are at least ten times more potential growing points on seed tubers
than mainstems which ultimately grow.
14
12
10
In
Q)
>-
Q) 8
'0
Cii
.0
E 6
:0
Z
20 40 60 80 100 120 140

Tuber weight (g)
Figure 6.2 Number of eyes per seed tuber for different tuber weights, Home
Guard (Firman, unpublished).
15
ell
~
0
E
'§" 10
(ij
..9:!
'0
(p 5
.c
E
;j
z
0 ,
0 5 10 15 20
Node number
Figure 6.3 Number of leaf primordia on lateral buds at successive nodes of the
primary tuber (50 g) axis. Nodes numbered from the dome of the apical bud. (From
Goodwin, 1967).
The main buds differ in their developmental stage, judged by number of
leaf primordia for as Fig. 6.3 shows dormant, apical and fully developed
lateral buds consist of about 12 leaf primordia while younger lateral buds
nearer the apical bud and tertiary axis buds have fewer primordia
(Goodwin, 1967).
6.2.2 Dormancy
For most varieties seed tubers are dormant at harvest and for a period
afterwards even in temperatures conducive to sprout growth. A few
early varieties have particularly short dormancies which may end before
September and therefore occur before harvest. Generally, early varieties
have shorter dormant periods than maincrop varieties (Table 6.1) but there
are numerous exceptions. Overall, seed potatoes of early varieties may
have 6 months of potential sprout growth before planting while some
maincrop varieties can have as little as 1 to 2 months of sprout growth.
Table 6.1 Dormant periods (weeks) of some British varieties harvested third
week in September 1957 and stored at 1(f'C (from Burton, 1963)
Dormant Arran Arran Golden Home King Majestic
period Consul Pilot Wonder Guard Edward
(maincrop) (early) (maincrop) (early) (maincrop)( maincrop)
After
tuber 26 23 27 24 21 28
initiation
After
harvest 12 5 12 5 6 12
In practical terms, the ending of dormancy in chronological time is the

important event as it marks the onset of the period of sprout growth,
whose duration, together with temperature, determines sprout growth at
planting. The timing of this OCCUffence is influenced by seed-crop hus-
bandry but the causes of these effects remain obscure. It is not certain
whether growth does entirely cease during the dormant period (Emilsson,
1949) or continues extremely slowly (Davidson, 1958). There have been
suggestions that the onset of the dormant period is at tuber initiation
(Burton, 1963) which if substantiated would allow study of the duration of
the period and factors affecting it. No substantial data validating the claim
are published and no serious study of the biological aspects of dormancy in
the potato has been produced. There are many reports of agronomic
factors influencing the chronological ending of dormancy - date of planting
(Wright and Peacock, 1934; Emilsson, 1949; Burton, 1963); timing of
defoliation (Holmes and Gray, 1972); fertilizers (Walker, 1968; Thow,
1970) - as well as purported effects of location (Broadbent et at., 1961;
Iritani, 1967) and altitude (Kozlowska, 1963) which illustrate practical
effects. However, few of these reports have used carefully controlled
experiments with concern for causal factors and hence the repeatability of
their observation. From these reports a number of generalizations have
been accepted, the most important of which is that dormancy ends earlier
in chronological time in warmer conditions. The evidence for this vi,ew is
scant (Went, 1959) while evidence for the view that temperature has little
effect, while not conclusive, is more substantial (O'Brien and Allen, 1986a;
Allen et aI., 1991). In their experiments Allen et al. (1991) found that the
duration of dormancy from tuber initiation was reduced by early defolia-
tion of crops with full leaf canopies (i.e. before commercial defoliation
is usually considered). However, for later defoliations covering the com-
mercial period, in several seasons at different sites, no effects on the
duration of dormancy were found. Even the effects of early defoliation on
dormancy break were quite small especially in relation to the period of
sprout growth. The results did not therefore refute Emilsson's suggestion
that dormancy is largely a fixed period of time from harvest nor Burton's
that it begins at tuber initiation. As the timing of harvesting in relation to
defoliation had no effect on dormancy (Allen et al., 1991), the severance
of the haulm from tubers appeared to be the main influence on the
duration of dormancy. Overall, appreciation of the factors influencing
dormancy is still poor and commercial manipulation largely unknown. The
controlling mechanisms in dormancy are biochemical and it is probably
unrealistic to expect agronomically useful general effects to be established
readily from simple observation of the visible appearance of sprouts. A
detailed biochemical and agronomic study is needed using tubers
from carefully monitored seed crops and strictly controlled storage
environments.
The ending of dormancy is of particular importance where two crops can
be grown each year. Kawakami (1973) has described the difficulties in
central and southern Japan where crops are grown in spring and autumn.
Seed for the spring crop is mainly produced by summer crops in northern
Japan and is freely available but for the autumn crop this seed is too old
and has sprouted extensively. Seed from the spring crop may still be
dormant and not allow emergence in time to give a viable yield. This latter
problem exists in many other regions and can be solved through use of
various chemicals which end dormancy, e.g. thiourea (Denny, 1926);
ethylene chlorhydrin; gibberellic acid (van Hiele, 1961; Holmes et al.,
1970); Rindite (a mixture of ethylene chlorhydrin, ethylene dichloride
and carbon tetrachloride), methyl bromide and potassium thiocyanate,
although many of those chemicals are dangerous to handle and may not be
available in the less-developed countries. Even storage at low tempera-
tures over long periods does not prevent some chronological deterioration,
devoid of the influence of sprout growth. In such countries which need
seed tubers throughout the year, liaison with technically advanced
seed-producing areas would allow non-dormant seed to be available for
planting at any time.
In developed countries some varieties have been shown to be sensitive to
low temperatures during dormancy and end the dormant period earlier
than from ambient storage (Wurr and Allen, 1976; Allen et al., 1978).
6.2.3 Sprout growth

At the end of the dormant period tubers begin to show bud growth if
placed in suitable temperatures. This growth leads to the appearance of
sprouts and while their appearance is well-known, definitions of what is a
sprout have been few yet varied. Thus, discussions of sprout growth taken
from the literature are prone to misconceptions as the description of a
sprout may vary. Bates (1935) defined a sprout as 'an eye showing viable
sprouts' and although used and largely unchallenged until recently, such a
definition is clearly unsatisfactory. Moorby (1967) called sprouts 'the
lateral shoots present on the tuber at planting'. Wurr (1975) defined a
sprout as 'bud development on the seed tuber which had leaf initials
present and at least partly open' and added the term sproutlet to refer to
'bud development on the seed tuber which had leaf initials present and at
least partly open irrespective of whether it was derived directly from the
seed tuber or whether it was a sprout branch'. Wurr's definitions do appear
to be definitive (and therefore repeatable) and are the basis of comments
about sprouts in this review.
The number of sprouts which grow per seed tuber is principally
determined by the size of the tuber and the temperature and duration of
storage. Figure 6.4 shows that the number of sprouts per seed tuber
increases with increasing seed tuber size but the number of sprouts on seed
tubers is also much influenced by storage conditions. Tubers placed in
temperatures suitable for growth at dormancy break produce few, or even
single sprouts, as the growth of the apical bud occurs rapidly and
suppresses the growth of the other buds, producing the apically dominant
(or single sprout) condition. The use of low storage temperatures, later
transfer of tubers to sprouting temperatures (Table 6.2) or damage to the
apical bud all result in more sprouts growing. Thus, prolonged, low
temperature storage will result in large numbers of sprouts growing once
the tubers are transferred to temperatures suitable for growth. Tubers with
many sprouts may be referred to as multi-sprouted, in comparison with the
single-sprout condition.
In general, the increase in number of sprouts per seed tuber with
increasing seed tuber size is restricted where the seed is sprouted for a long
period for even the largest seed may produce only one or two sprouts.
Early varieties such as Home Guard and Arran Comet which break
dormancy in early autumn will frequently show complete apical dominance
in the ambient conditions of autumn. Most maincrop varieties have longer
o
8
E
.3
~
E 7
E
C')
1\
o
~a. 6
oo
'0
E
15 5 o
z
o
4
25 35 45 55 65 75 85 95 105 115
Seed weight (±5g)
Figure 6.4 Relationship between number of sprouts per tuber and tuber weight in
Home Guard (Allen, unpublished). Regression equations, y = 2.93 + 0.055x.
R2 = 0.99.
Table 6.2 Number of sprouts >3 mm per tuber at planting in Home

Guard following different dates of entry of tubers into sprouting
temperatures of 8 and 13°C (Allen et aI., 1979).
Sprouting Seed sprouted from
temperature eC) 15 Sep 15 Nov 15 Jan Mean SE
8 2.8 7.6 7.3 5.9
13 2.1 7.7 6.9 5.6
Mean 2.5 7.7 7.1 0.30
SE 0.43 0.24
dormant periods and do not begin to sprout until ambient temperatures are
lower and usually develop several sprouts. It should not, however, be
imagined that all early varieties have apically dominant seed and all
maincrop varieties have seed in a multi-sprout condition for with suitable
adjustments to storage conditions the converse of this may be produced.
Extreme fluctuations in autumn temperatures can also alter the number of
sprouts per tuber in any variety.
Sprouting of seed tubers may be regarded as a means of restricting the
number of potential growing points which actually grow and the longer the
period of sprouting the greater is the effect. As sprouting affects plant size
and longevity, it has profound implications for considerations of plant
density in potatoes. Unfortunately, in general, variations in the numbers of
sprouts (stems) produced by seed tubers as a result of sprouting have
received limited attention in this context (see Chapter 7).
The length of the sprouting period also affects the morphology of the
sprouts, for single sprouts often exhibit swollen bases and tend to branch,
often following tip necrosis or damage. This sprout branching results in
secondary stems as well as mainstems emerging above ground. Generally,
large seed produces more branches than small and with long sprouting
periods this tends to compensate for the limited number of sprouts.
Goodwin et al. (1969) found that th.e number of mainstems in the fieJd
was proportional to the number of sprouts at planting. However, not all
sprouts> 1 cm at planting emerged because of the re-imposition of apical
dominance referred to by Morris (1967). It is probable that such effects of
apical dominance occur in all sprouted tubers and result in fewer buds
developing into stems than develop sprouts. Morris (1967) suggested that
only those sprouts which were 'actively growing' at planting developed into
stems but it is clearly necessary to know how to recognize such sprouts and
this is not known although sprout length is a useful indicator in some
cases. The relationship between number of 'actively-growing sprouts' and
number of stems is variable e.g. Jarvis and Rogers-Lewis (1974) (Table
6.3). That many sprouts present at planting do not develop into mainstems
is illustrated by the results of Schepers and Hoogland (1968) who found
that although the number of mainstems in the field was proportional to the
number of sprouts at planting, only 30% of sprouts had developed. This
compares with 75% reported by Bleasdale (1965) for Majestic and 64%
which he reported Bates (1935) to have achieved with King Edward.
Schepers and Hoogland used marked sprouts and harvested all plants to
determine whether sprouts observed at planting actually developed into
mainstems. Such an approach is essential if the developmental pattern of
sprouts is to be recorded but only limited data of this type are available.
From the limited data it is clear that there is considerable variation in the
proportion of sprouts which develop into stems and much of this variation
cannot be partitioned. It is essential that effort is directed towards
establishing reliable relationships between a sprout character recorded at
planting and the stem density resulting in the field, for without such
relationships it is impossible to predict the number of stems produced by
seed stocks. A number of possible sprout characters have been suggested.
Schepers and Hoogland (1968) reported that sprouts with root primordia
showed an excellent linear relationship with mainstems for 98% of such
sprouts developed into mainstems. No other reports of this relationship
have been found but it appears worthy of testing for, like sproutlets, root
primordia are an easy character to record. One obvious difficulty would be
the variation in sprout size at which varieties first show root primordia.
Sproutlets, which include all bud growth on tubers, were shown by Wurr
(1975) to be closely related to number of above-ground stems (mainstems
and secondary stems) but the validity of these relationships over a
wider range of varieties and environments has not been established.
Subsequently, Wurr and Morris (1979) found sproutlets less effective in
predicting number of stems than tuber weight. Thomas (1988) also found
closer relationships between number of stems and tuber weight than with
any other seed-tuber characteristic. As Fig. 6.5 shows he obtained separate
relationships for unsprouted and sprouted seed in three varieties which
appeared sufficiently consistent over two seasons for him to argue that
prediction of stem densities appeared to be possible with acceptable
accuracy for commercial use. This work was only over two seasons and the
robustness of the relationships over sites, seasons and seed stocks has not
been tested sufficiently to agree with Thomas' expectations. Such work is
clearly urgently needed and the subject is further discussed in Chapter 7.
Table 6.3 Relationship between number of actively-growing sprouts and number of

mainstems in two potato varieties (Jarvis and Rogers-Lewis, 1974)
Within-row spacing (cm)
Set
size (g) 66 33 22 16.6
Pentland Ivory
Number of actively-growing sprouts 36.3 86.2 129.2 172.1
planted (ooOs ha- 1) 63.5 49.4 98.8 148.1 197.7
101.5 54.2 108.4 162.6 216.7
Number of mainstems in June 36.3 78.5 109.6 137.2

(OOOs ha- 1) 63.5 51.7 106.2 146.9 185.5
101.5 69.9 127.0 185.1 138.0
Record
Number of actively-growing sprouts 36.3 82.4 123.1 164.3
planted (OOOs ha- 1) 63.5 48.7 97.4 146.0 194.7
101.5 53.7 107.3 160.9 214.6
Number of mainstems in June 36.3 92.7 127.7 157..2

(OOOs ha-) 63.5 68.0 124.0 168.4 215.1
101.5 80.6 145.7 210.0 259.7
Rate of sprout growth increases with temperature, over the range

normally found in winter storage in temperate latitudes, 2-16°C (Fig. 6.6).
The number of sprouts which grow at high temperatures increases with
delay in entry to the temperature so rate of total sprout growth is very high
at high temperature after prolonged storage at low temperature. The
increases in rate may continue well past the normal time of planting as
shown by Krijthe (1962). She measured the weight of sprouts which would
Estima Mainstems Above-ground stems

6 6 ,
, •
....
- ,
.n 4
Q)
!
iii 4 , !
,• ,
~
••
~
:::J
I ! I IJ
i ~
••
"0
Q) 2 2
Q)
en i' •
i'C'
....
Q) 25 45 65 85 110 25 45 65 85 110
a.
en
--
E
Q)
en
0
.... Pentland Squire
Q) 6
.n
•
~
E
,• ,A ••
:::J ~
z
,
4
• i •, ,•
•
0
i •
0
• •
•
~
2 ~ • I
"
25 45 65 85 110 25 45 65 85 110
Seed tuber weight (±5g)
Figure 6.5 Relationship between number of stems per seed tuber and seed tuber
weight for three physiological ages of seed tubers of two varieties . Symbols -
number of day-degrees >4°C accumulated by seed tubers; 6., zero; D, 400; 0, 800.
Closed symbols, 1984; open symbols, 1985. (Thomas, 1988).
---0 4°e
15
160 --0 1Qoe
_15°e
50 ~ 20 e0
~
D
~ 40
.s
:5
ec. 30
(J)
'0
£; 20
0>
c
OJ
...J
10
4
_--e-----e
~ _---e--
o Oct. Jan. Apr.
Months - Monate - mois
Figure 6.6 Length of apical sprouts of Arran Pilot tubers stored at four constant
temperatures. (From Sadler, 1961).
be increased by more sprouts of greater length and found increases in
sprouting capacity up to a year after harvesting the seed (Fig. 6.7). Rate of
increase of length of the longest sprout with delayed onset of sprouting can
also occur but the effect is smaller than with total sprout growth. Prolonged
sprouting at high temperatures can lead to a reduction or even cessation in
growth which is probably associated with irreversible changes at the sprout
apex which may result in its death. Such effects are of great practical
significance for early varieties, with short dormant periods, which are
frequently stored in areas of high ambient winter temperatures.
§ 10
:E
Ol
.~
Q;
.0
.a
'0
?!< 5
~
§
:E
Ol
.~
S
e
c%°NDJ FMAMJ J ASO N
Time (months)
Figure 6.7 Seasonal trend in sprouting capacity of Bintje in 1958. (From Krijthe,
1962).
6.2.4 Sprout development and floral initiation

Until recently sprout and stem growth has been considered without
recognition of the potential significance of the developmental sequence
which unfolds during growth. The main buds which grow from eyes
ultimately result in the complete stem structure which encompasses
subterranean scale and above-ground leaves, stolons and tubers and the
terminal floral structures. As crops grown from unsprouted and heavily-
sprouted seed can both achieve complete ground cover, the origins of the
leaves in the two canopies are likely to be different as apical differentiation
continues during pre-planting sprout growth but the fate of the primordia
has been ignored. Recently, Firman et al. (1991a) have studied the effects
of temperature and length of sprouting period on the number of leaf
primordia in sprouts and ultimately stems of several varieties. Number of
primordia increased in some varieties over long periods of sprout growth,
while in other varieties apical differentiation was arrested after a period of
sprout growth (Fig. 6.8). Even at low temperatures «4°C) a slow increase
in number of primordia occurred and low temperatures prior to sprouting
frequently led to floral initiation in sprouts before planting. Thus, the
complete mainstem structure can be determined before planting in some
varieties. Elucidation of the developmental sequence in the potato is likely
to help understanding of variation in growth of canopy and tubers and in
the number of tubers ultimately found on each stem.
50
40
Jl20
E
:::l
Z 10 .--- ---
o 40 80 120 160 200 240

Days at 13°C
Figure 6.8 Number of nodes on the longest sprout per tuber of Home Guard and
Maris Piper stored at 13°C (Firman et aI., 1991a). x, Home Guard; e. Maris Piper.
6.2.5 Emergence and stem development

The number of leaf primordia present in sprouts at planting increases with
increasing sprout length and delay in planting (Table 6.4). Where flowers
are not initiated in long sprouts before planting, in some varieties further
apical differentiation may occur after planting. This may represent
continued differentiation as in Home Guard or resumed differentiation as
in Maris Piper. The consequence is that in many varieties the total number
of nodes (primordia) to the first flower (the mainstem structure) is
extremely variable while in others, notably Estima, the number is less
variable. There is no constancy of number of nodes to the first flower, as
suggested by Taylor (1953). Estima is particularly interesting as similar
numbers of nodes are produced in old and young seed tubers. In the
former, all nodes may be present in the sprout at planting while in the
Table 6.4 Number of nodes on the longest sprout of

tubers of Arran Comet stored at ]3°C (Firman,
unpublished)
Date
4 Dec 26 Feb 18 Apr 16 Jun 15 Aug
Number of nodes 11.6 24.2 34.6 37.8 42.6
latter only a quarter of the nodes are present in the bud and the remainder
are produced during pre- and post-emergence growth.
Increasing sprout length and/or late planting result in more nodes per
mainstem and this increase is mainly in the number of below-ground nodes
(Table 6.5). For all varieties this results in similar numbers of leaves on
mainstems from different sprouting regimes although the number of leaves
to the first flower has a considerable range across varieties. Thus, in any
variety the storage conditions and timing of planting can affect the
environmental conditions experienced by the individual primordia which
ultimately form leaves. The number of below-ground nodes may well
influence the number of stolons and potential tuber sites and the effects of
storage conditions of seed on these aspects of growth and development
require much greater attention. Only through greater understanding of
these processes is real progress in control of growth, number of tubers and
hence value of final yield, likely to be achieved.
Table 6.5 Effect of number of day-degrees (>4°C) accumulated by seed

tubers at planting and date of planting on sprout length and number of below
and above-ground nodes in Maris Piper (Firman et ai., 1991a)
Number of nodes
Date of Length of longest Below-ground Above-ground
planting sprout/tuber at
planting (mm)
Number of
day-degrees 0 750 0 750 0 750
>4°C at
planting
14.iv 1.5 11.0 8.8 12.7 17.3 18.7
24.v 2.7 15.8 8.7 13.7 19.3 18.0
29.vi 6.6 17.2 10.2 16.0 20.3 21.3
9.vii 8.0 18.2 11.3 16.3 19.2 20.5
SE 1.17 0.76 0.56
Environmental conditions after planting influence the rate of emergence

and subsequent growth of the leaf canopy. Rate of sprout elongation
increases with temperature over the range 4-20°C (Fig. 6.6), but there is
usually a lag phase before linear growth with temperature occurs (Firman
et al., 1991b). This lag is shorter at higher temperatures and for longer
sprouts and decreases with delay in planting (increasing sprout length) even
from cold storage. This effect is consistent with the continued differentia-
tion found in such sprouts stored at similar low temperatures by Firman et
al. (1991a). Rate of emergence is slower in dry soil, and especially so at
high temperatures. Prolonged intervals from planting to emergence in low
soil temperatures frequently lead to shorter stems and lower leaf area
indices than later plantings and in varieties such as Estima this can result in
markedly lower yields (Table 6.6). As Estima has relatively few mainstem
leaves, any interference with their functioning and architecture leads
almost directly to effects on yield. Thus, better understanding of the stem
development of individual varieties will directly influence many agronomic
practices, e.g. dates of planting and seed rates.
Table 6.6 Effect of date of planting on

tuber yield >30 mm (t ha- l ) in two
varieties harvested on 20 August 1984
(Allen, unpublished)
Variety Date of planting SE
7 Mar 7 May
Estima 45.1 55.6
4.69
Maris Piper 50.3 40.5
Data for six other varieties are not presented.
6.2.6 Leaf growth and yield

Sprouted seed usually emerges before unsprouted seed and for an initial
period has a higher leaf area index. However, this advantage is gradually
eroded and frequently unsprouted seed produces a higher peak leaf area
index of greater persistence. The general pattern is shown in Fig. 6.9 and
the previous sections have indicated that the histories of the component
leaves of the two canopies may be very different where seed of contrasting
sprout length produces complete ground cover. The two canopies may
comprise leaves of quite different architectures especially if internode
and petiole lengths are affected but little is known of these effects. The
advantages of extra leaves and continued differentiation of seed with
limited sprout length can be nullified if the tum-over of leaves is acceler-
ated through shading or lodging. These effects have not been seriously
studied but are fundamental to effective use of seed in specific systems as
yields are determined by the amount of radiation intercepted and the
efficiency of its conversion (Allen and Scott, 1980). Either component
could be altered by the storage treatment of seed.
6.2.7 Physiological age
( a) Concepts
As dormancy normally ends before planting, recognition that sprout
growth may influence field growth is long established and preceded
Unsprouted seed
Sprouted seed
Time
Figure 6.9 Schematic representation of leaf area index over time for sprouted and
unsprouted seed.
research into such effects. Thomas and Eyre (1950) had clearly observed
important effects of sprouting in early potatoes and from this time attempts
to understand the effects of sprouting on field growth and yield have been
numerous. The term physiological age was introduced by Madec and
Perennec (1962) to cover the effects of the seed tuber on subsequent crop
growth. They believed that these effects were not wholly the consequence
of sprout growth, arguing the effects were carried in the tuber. This
question has rarely been addressed and the literature on physiological age
or sprouting is characterized by confusion as to whether ageing is sprouting
or sprouting plus something else. Both in UK and Europe the ageing
process has been illustrated through sprout growth as its visible manifesta-
tion. Thus, Toosey (1964) described physiological age as 'the physiological
state of the tuber at any given time - which is illustrated by the degree of
visible sprout development'. Krijthe (1962) suggested that changes in
sprouting capacity with time could be used to measure the physiological
age of tubers. She reported the sequence of aging phenomena to be:
1. single sprout stage;
2. multiple sprout stage;
3. branching stage;
4. small tuber formation stage.
Such a sequence involves a decrease in apical dominance with older tubers
producing more stems in the field. This suggestion was misleading as it
failed to recognize the essential progression of an ageing process. A tuber
may be young at the end of dormancy on entering stage one but it cannot
remain so if allowed to sprout. Such a tuber will not experience the second
stage and will frequently be planted before the third stage. Krijthe's
sequence is essentially a combination of the chronological progression in
number of growing sprouts (stages one and two, which cover a range from
one to several dozen sprouts over many months for tubers stored at 2°C
until sprouted) and the final developmental stages of growing sprouts
(stages three and four which can occur following onset of sprouting at
either of the first two stages). In both parts many of the effects occur after
seed tubers are normally planted (as also in Fig. 6.7) and therefore have
limited practical significance. The latter has been the primary considera-
tion of recent research on physiological age in the UK which has recognized
the necessity of separating chronological and physiological effects in
obtaining a practical measure of age and resolving whether effects are
sprout or tuber borne. Of overall importance is the recognition that unless
physiological age can be measured, it can have no practical utility nor
any scientifiic validity. The vast literature which uses the term without
definition is testimony to self-deception among researchers.
Few British research workers have found the incubation period - 'the
time between the onset of sprouting of the seed tube:r (desprouted if
sprouts were present) and tuber formation on the new sprouts, when
tubers are stored in darkness at IS-20°C and 90% RH' (Hartmans and van
Loon, 1987) - useful in studying effects of physiological age as it ends after
seed has usually been planted. Consequently, any differences are only of
academic interest and the whole period is incapable of practically discrimi-
nating between stocks of tubers at different points in the ageing process.
As few tubers are deliberately desprouted in the UK, research has concen-
trated on tubers planted with their initial sprout growth intact. This is
in marked contrast to many authors' work where tubers have been
desprouted at some stage before planting (Madec and Perennec, 19S6;
Reust and Munster, 1974; Hartmans and van Loon, 1987). Such an
experimental approach inevitably confounds effects of sprout growth with
de sprouting and both may be expected to influence the re-growth of
sprouts and field growth. Where desprouting has been studied (Allen and
O'Brien, 1991) it has been possible to establish where the effects of
sprouting are determined. In some varieties, where long, old sprouts were
planted, 'little potato disorder' occurred in many plants but the removal of
such sprouts at planting resulted in the later emergence of full plant stands
and greatly increased yield (Table 6.7). Little potato disorder is the
culmination of a developmental sequence leading to tuberization and is
consequent upon changes at the sprout apex. Thus, the effects of ageing
Table 6.7 Effect of date of desprouting on total tuber yield (t ha- I ) in Red
Craigs Royal in 1980 (Allen and O'Brien, 1991)
Date of desprouting
Temperature of 11 Jan 13 Feb 14 Mar Undesprouted SE
seed storage eC) (planting)
6 29.9 22.9 30.3 31.8
3.59
12 29.6 31.1 26.3 11.0
(which are consistent with the changes in the apex discussed in Section
6.2.4) can be removed in the sprout which leads to the conclusion that
ageing is simply sprout growth. All effects of physiological age can
therefore be studied in relation to the number and rate of growth and
development of sprouts without recourse to contrived and unmeasurable
factors such as growth vigour (van der Zaag and van Loon, 1987).
As rate of sprout growth is linearly related to temperature over the range
usually experienced in stores (4-16°C) and emergence and field growth are
25
-
,.- I
E
E
~
20
-, ,
,
,~
o
o
...
C>l
-...
.D
~
C>l
-
a. 15
~
0
L-
a.
II!
iii
C>l
en
c:
0 10
-
C>l
.I:
15
-
.I:
en
c:
C>l
-' 5
I
4 I
I
o ~I--------~,--------~i----------,r_--------Ti---
o 200 400 600 800
Number of day-degrees >4°C
Figure 6.10 Relationship between length of the longest sprout at planting and
number of day-degrees >4°C in four varieties (O'Brien, unpublished). 0,
Maris Bard, y = 3.26 + 0.017x, R2 = 0.97; D, Pentland Javelin, y = 2.43 + 0.0189x,
R2 = 0.98; 6., Desiree, y = 2.52 + 0.027x, R2 = 0.99;., Cara, y = 0.61 + 0.0266x,
R2 = 0.97; I, SE of treatment means.
(a)
50
~
.c
i::. 40
E
u
'"II
N
30
"0
a;
0;;'
ffi 20
.0
::l
t-
o ~!------~------~--------~------~-
o 200 400 600 800
Number of day-degrees >4°C
Figure 6.11 Relationship between (a) leaf area index and (b) tuber yield and
number of day-degrees >4°C accumulated by seed tubers of Pentland Javelin
at planting (Allen and O'Brien, 1986). (a) 0, 20 May, y = 0.82 + 0.0023x - 1.66 x
1O·6x2, R2 = 0.97; D, 5 Jun, y = 2.86 - O.0007x, R2 = 0.74; 6,25 Jun, y =
3.62 - O.0013x, R2 = 0.70. (b) 0, 7 Jun, y = 11.34 + 0.00382x - 2.59 x 1O-5x2,
R2 = 1.00; D, 25 Jun, y = 37.5 + 0.0223x - 2.00 x 1O-5x2, R2 = 0.93; 6,4 Aug,
y = 60.7 - 0.0127x, R2 = 0.76. t, optimum number of day-degrees.
affected by sprout growth at planting, a quantitative measure of sprout

growth and its effect on field growth may be achieved through accumulated
day-degrees. Figure 6.10 shows that sprout length in many varieties is
closely related to number of day-degrees experienced by the seed and Fig.
6.11 shows that leaf growth and tuber yield are also closely related to
number of day-degrees experienced by the seed. The relationships are very
close which suggests a causal influence of temperature rather than an
association and since the initial findings of the mid-1970s increased
understanding of growth and development allow explanation of these
effects. Increasing physiological age increases sprout lengths at planting,
hastens emergence, tuber initiation and early leaf growth, produces a
lower peak leaf area index and often leads to earlier senescence (Figs 6.10
and 6.11). Consequently, yields increase with increasing age at early
harvests, may be unaffected by age at harvests in the middle of the season
and decrease with increasing age at the end of harvesting period. As many
early varieties have short dormancies, an extremely wide range of ages can
be produced and over their period of harvesting the whole range of effects
can be demonstrated. In many instances leaf area indices do not exceed 3
so complete ground cover is not achieved and all differences in leaf cover
are directly linked to yields. In maincrop varieties with longer dormancies
and longer growing seasons effects are rarely so dramatic but usually of
commercial significance. In such varieties most ages of seed will achieve
complete ground cover and effects on yield are determined by effects on
emergence and time course of leaf growth in relation to incident radiation.
Effects of physiological age on tuber yields are caused by effects on the
timing of emergence, leaf growth and senescence of the different crops. If
these do not occur or do not alter the amount of light intercepted by crops
grown from different ages, there will be no effects of physiological age nor
should any be expected. Firman et al. (1991a) have shown that in some
varieties a marked slowing of apical differentiation occurs after long
periods of sprouting and such seed may show reduced rates of sprout
elongation after planting (Jones and Allen, 1983). Consequently, effects of
age on emergence may be reduced or eliminated. If such seed does not
resume apical differentiation it will produce determinate stems and a small
leaf canopy compared with younger seed and lead to large effects on yield.
However, a resumption of differentiation may lead to comparable leaf
areas from different ages and different effects on yield. For any variety
there can be no single effect of age for all conditions, as environmental
conditions influence these effects (O'Brien et al., 1983) but the general
pattern shown in Fig. 6.11 occurs in all situations with the magnitude and
duration of the effects changing.
The concept of measuring physiological age as the number of day-
degrees above a base temperature from onset of sprouting (end of
dormancy) to planting is quantitative and practical. O'Brien et at. (1983)
established that the base temperature could be 1 to 4°C with little effect on
the fit of the relationship and chose 4°C as this was frequently the lowest
temperature used for storage and little sprout growth occurred. Sprout
growth will occur in some varieties at 4°C and lower base temperatures
would be appropriate. Dormancy is generally taken to have ended when
the mean length of the longest sprouts on a sample of tubers is 3 mm. This
length is easy to record but shorter lengths could be used. For widespread
practical use it is essential that all unsprouted seed, for whatever reason,
has the same growth pattern and that the sequence of temperatures leading
to similar final numbers of day-degrees has little effect. Seed planted
without sprouts after several desproutings (Table 6.8) and from different
Table 6.8 Effect of frequency of desprouting and seed storage temperature
on final tuber yield >23 mm (t ha-l ) in Desiree (Allen and O'Brien, 1991)
Number of desproutings
Temperature of
seed storage (0C) 5 4 3 2 1 Undesprouted SE
6 52.7 49.3 45.5 53.5 53.5 51.5
3.27
12 41.1 44.3 52.3 51.7 49.3 53.4
Table 6.9 Total tuber yield (t ha- l ) in ware crops of Pentland Dell from ten seed
stocks planted and harvested at different times in Scotland and stored at two
temperatures (Allen and Wormington, 1991)
Seed stock number
Temperature of
seed storage ("C) 2 3 4 5 6 7 8 9 10 SE
4 50.9 44.8 49.2 47.4 48.4 48.4 50.2 45.8 40.6 44.6
3.07
4/12 53.9 49.7 50.3 55.7 55.4 54.1 52.4 56.2 52.3 48.7
seed stocks stored at <4°C (Table 6.9) produced similar growth patterns
and yields. There may be varieties which respond differently to desprout-
ing e.g. laerla (van Loon, 1987) but for the UK the effect seems minimaL
Temperatures differ between seasons and areas, so unless controlled
temperature storage is practised, the sequence of temperature experienced
by seed in individual seasons over similar periods of sprouting can vary
considerably. In combination with the effects of the timing of onset of
sprout growth on number of sprouts there may be effects on emergence
and number of stems and hence time course and extent of leaf growth. The
results of Wurr (1979), and O'Brien (unpublished) (Table 6.10) show that
there were few important effects of sequence of temperature. Gillison et al.
(1987) found some effects of sequence using temperatures of 4°C and 14°C
as high temperatures close to dormancy break sometimes reduced number
of stems and yields in some cultivars at early harvests. Whether these
effects would occur in practice is debatable. The branching of stems on the
plants with fewer stems as a result of higher initial temperature is
frequently a reason for the lack of effects as differences in ground cover are
minimized. Thus, there seems every justification for measuring the effect
of the seed production phase and pre-planting storage environment on field
growth through the number of day-degrees above a base temperature
experienced by the seed tuber during the growth of the sprouts actually
planted (Allen and O'Brien, 1986).
Table 6.10 Effect of different sequences of seed storage
temperature on tuber yield >25 mm (t ha· l ) in Arran Comet
(O'Brien, unpublished)
Sequence of temperature CC) resulting
in 834 day-degrees >4°C
Date of
harvest 4112 6/8 8/6 12/4 12/6/4 SE
26 Jun 22.7 19.6 21.2 20.1 22.3 1.22
11 Jul 40.8 36.0 39.2 38.6 35.3 3.23
15 Aug 60.9 57.1 56.3 55.8 62.7 4.48
(b) Application in ware production

The type of relationships between tuber yield and age in Fig. 6.11 allow the
determination of optimum ages for different dates of harvesting in each
variety. These ages (Table 6.11) should then be achieved through appro-
priate storage duration and temperature in each season. As onset of
sprouting and ambient temperatures vary seasonally, there may be
variation in age from year to year even from similar periods of sprouting.
This has to be accepted by growers unless some control of temperature can
be achieved. If age is simply recorded it is helpful in explaining seasonal
effects but more positive management is needed to use the increased
knowledge of effects of age for commercial advantage.
Table 6.11 Derived optimum number of day-degrees >4°C accumulated

by seed tubers at planting for tuber yield in four varieties for different
periods of harvesting (O'Brien, unpublished)
Harvest period
Variety Early June Late June Early July Late July August
Home Guard 1944 1112 1014
Pentland 737 558 0
Javelin
Desiree 560 466
Cara >840
In early potato production traditional varieties have short dormancies

and achieve very advanced age (>1000 day-degrees) by planting even in
ambient conditions. Some seasonal variation in age would occur but only
rarely would ageing be seriously restricted. Until relatively recently,
considerable harvesting began at low yields (5-8 t ha- 1) but very high prices
and such old seed achieves this yield before any other. Figure 6.11 shows
that such seed is only suitable for harvesting for a short period (1-2 weeks)
as more rapidly bulking younger seed soon achieves· a higher yield.
However, for most growers there is only old seed and consequently many
later harvested crops only achieve a small fraction of the potential yield
and hence growers' returns are lower than they could be. It would be
beneficial to use younger seed of the variety or change to a variety which
does not achieve such extreme age like Pentland Javelin or Maris Bard.
Relatively few growers in some early arenas e.g. Kent and Pembroke shire
(Dyfed) have appreciated the need for such changes but in some other less
early areas (Shropshire, Cheshire and Suffolk) appreciation has been
greater and a general advance in earliness and increase in acreage of early
crops has occurred (Table 6.12). These differences are an interesting
comment on the effective development of research findings as the original
work was carried out in Pembrokeshire. Substantial improvements in the
yields of most crops harvested during June and early July could be
achieved if the appropriate age of seed for the most popular varieties were
planted. As much of the information is already available, the deficiency
reflects the inadequacies of application of existing knowledge.
Table 6.12 Area (hectares) of early potatoes planted in different

counties of the UK, 1979--89 (PMB statistics)
County
Year Kent Suffolk Cornwall Cheshire Shropshire Dyfed
1979 2920 1475 1655 1610 2145 2410
1980 2920 1460 1745 1605 2130 2290
1981 2635 1415 1615 1755 2120 2185
1982 2530 1435 1645 1830 2125 2030
1983 2810 1635 1715 1965 2315 2175
1984 2785 1795 1835 2275 2585 2310
1985 2960 1985 2050 2420 2810 2465
1986 2535 1990 1940 2260 2650 2190
1987 2540 2080 1915 2280 2750 1970
1988 2415 2205 2030 2320 3100 1830
1989 2372 2188 2150 2363 3190 1748
The traditional use of aged seed with the belief that the 'older the better'
was the principal reason for the widespread use of home-grown seed in
early areas. It is now clear that the source of the seed is of little importance
if physiological age at planting is the same. However, this approach
resulted in the demise of the variety Red Craigs Royal as greater age
increased the prevalence of 'little potato disorder' to uneconomic levels.
Although effects are rarely so dramatic this emphasizes that the perfor-
mance of varieties is considerably influenced by the age of seed used and
their relative performance greatly influenced by time of harvesting. For
much of the harvesting period older varieties benefit from restriction in
their physiological age while many newer varieties benefit from increase in
their physiological age. Thus, the appropriate storage management for
varieties can be very diverse and frequently difficult to achieve in one
storage environment. Utilization of vertical temperature differentials in
store can be a simple way of achieving the necessary differences. Where
seed tubers are stacked to heights of 2-4 m the temperature of the upper
boxes may be 2-3°C higher than the lower boxes and allow greater
physiological age to be achieved at the top of the stack. As these boxes are
unloaded first, there is no inconvenience at planting time.
For longer season crops the effects of age of seed are greatly influenced
by the pattern of leaf senescence which largely determine whether the
advantages of earlier emergence and tuber initiation of old seed are
maintained, eliminated or reversed. The maincrop varieties used rarely
achieve great age and for the earliest harvest during August ageing seed
will give higher yields. As harvesting is delayed the effect is invariably
reduced and may be reversed in a very long growing season. There is a
need to advance harvest to ensure high quality of tubers from storage
(Wilcockson et at., 1985) so the available growing season is likely to shorten
and consequently the likelihood of yield advantages from unsprouted seed
will be reduced. Only c. 25% of maincrop seed is aged and this seems to be
Table 6.13 Effect of date of planting and

physiological age on tuber yield (t ha·1) in
Desiree (Allen and O'Brien, 1990)
Physiological age
o 300 600
Date of planting Yield >30 mm on 30 May
20 March 7.9 13.6 12.6
6 April 0.3 1.9 2.9
SE = 0.67
Yield >40 mm on 21 June

20 March 32.6 32.9 33.7
6 April 12.2 18.6 22.1
24 April 0.9 5.0 7.8
SE = 1.16
Yield >40 mm on 10 July

20 March 46.9 45.6 43.7
6 April 26.0 34.0 38.9
24 April 12.0 25.4 19.8
SE = 1.23
decreasing. The main reasons for this are the availability of second early
varieties such as Estima, Wilja, Marfona and the practical difficulties in
planting sprouted seed mechanically. Second early varieties produce much
higher yields than maincrops during August and early September and can
be harvested for storage. High yields and good quality can be achieved
with relatively little ageing although the earlier the harvesting of such
varieties the greater the optimum age. The yield advantage of these
varieties results from reduced dry matter content but its existence has
reduced interest in advancing yields of maincrop varieties through sprout-
ing. In cases such as potatoes for crisping or pre-packing where specific
maincrop varieties are used, increases in yield due to ageing can be usefully
obtained at early harvests (Table 6.13).
The advantages of ageing seed can only be obtained if the tubers can be
planted with intact sprouts and the almost universal adoption of mechani-
cal planting makes this more difficult than in the past, especially for
early potato production. Sprouted seed does not 'flow' through planting
mechanisms as well as iisprouted seed and, if the sprouts become
entwined, damage is inevitable in separation, even before the effects of
delivery into the soil. As a result, observation suggests that sprout damage
occurs to a greater extent than before although effects and their con-
sequences are poorly documented. In order to achieve high rates of
planting, rapid loading of planters is essential and the traditional sprouting
(chitting) tray with 16-18 kg of seed is small and hence slow and labour-
intensive. There has been considerable interest in using larger, bulk
containers for sprouting but their exclusion of light from the majority of
tubers leads to more rapid elongation of sprouts and it has to be accepted
that such containers are only suitable for short periods of growth just prior
to planting, in effect the mini-chitting system of the 1960s and 1970s.
Sprouts from a period of 3-5 weeks' growth immediately prior to planting
are short, pale and often suffer more from damage during planting than
longer sprouts (Jarvis and Palmer, 1973). Yields are usually greater than
from unsprouted seed but often less than from older seed (Table 6.14).
Table 6.14 Effect of length of sprouting period

and type of planter on yield (t ha' z) of King
Edward, 1969-71, (Jarvis and Palmer, 1973)
Sprouting
Type of planter
period
(weeks) Hand-fed Cup-fed BelMed
Long (11-15) 42.4 42.3 41.3
Short (4-6) 42.5 39.8 39.7
SE 0.62
Seed multiplication and certification 271
6.2.8 Utility in seed production
The complexities of the growth and development of the seed tuber are
considerable and still imperfectly understood. However, it is quite clear
that the seed tuber exercises a major role in the growth of the crop and
therefore the relationship between seed production and storage and ware
production is crucial to successful production of potatoes for consumers.
As yet this relationship is not as close as it should be and opportunities of
commercial value to both seed and ware growers such as manipulation of
dormancy are being missed. The majority of seed is still purchased (and
traded) as a standard commodity with little regard to its biological qualities
and hence value. Closer links between seed and ware growers are likely to
be the mechanism by which seed of the desired biological quality is
produced for specific ware growing circumstances.
6.3 SEED MULTIPLICATION AND CERTIFICATION
The aim of a seed certification system is to maintain the varietal purity of

seed stocks and to control the spread of disease by prescribing tolerances
and cultural conditions. The starting point is disease-free tubers from
genetically pure stems which may be tuberized in glasshouses as stem
cuttings or as plantlets in micro-propagation. Until recently the former
system, generating clonal tubers from Virus Tested Stem Cuttings (VTSC),
had been the only primary source but despite higher cost the higher
multiplication rates of micro-propagation are releasing increasing numbers
of mini-tubers into the multiplication chain. As a large number of mini-
tubers can be produced very quickly they can reduce the first three or four
years of clonal, outdoor multiplication to only one indoor year. Whatever
the source of the initial disease-free tubers, multiplication proceeds from
small, often hand-worked, plots to field-scale crops in areas deemed
suitable for seed production.
Most countries are selective in their choice of seed area and in the UK
great advantage is taken of the relatively cool, windy and comparatively
aphid-free conditions in Northern Ireland, Scotland and northern England
to produce seed with minimal virus infection. In many countries the
climate is such that this is more difficult. In the Netherlands, for example, a
low level of virus infection is attainable only by making early defoliation of
the haulm in late July or early August obligatory, the precise date
depending on the variety, the grade of seed and seasonal aphid activity.
This system produces seed tubers which are relatively small in size with
moderate levels of latent fungal disease infection because the skins are well
set at harvest and the tubers are harvested in relatively warm, dry
conditions. By comparison, planting and consequently haulm destruction
and harvesting in the seed areas of the UK take place late in the season,
with attendant higher risks of tubers becoming infected with fungal
diseases.
6.3.1 Health
The potato suffers from many virus diseases and latent tuber diseases
which are considered in detail in Chapter 10. Virus diseases are seed-
borne, most are spread by aphids and the use of infected tubers leads to
smaller plants with mottled or malformed leaves and reduced yields. The
multiplication process begins with virus-free tubers and the tolerances for
infection are severe during multiplication. There are aphids in the seed
areas of Scotland and therefore there is a risk of virus transmission because
a reservoir of infection exists in gardens and commercial ware crops which
can be proximal to crops entered for certification (Ted, 1983). Despite
these difficulties the effects of virus diseases in ware crops are currently
limited and the last serious infection occurred in 1975-77. In some parts of
the world seed crops can be isolated from ware crops but this does not
occur in UK. Recently, it was agreed that ware growers in Scotland must
purchase new certified seed every second year in an attempt to reduce the
planting of uncertified seed in seed-growing areas.
There are many fungal diseases which infect tubers and their influence
covers both field growth and storage since their presence affects tuber
appearance which increasingly determines crop value. These diseases, skin
spot, gangrene, dry rot, silver scurf, are absent from the extremities of
stems plantlets so tuberization of such material in sterile conditions
produces progeny tubers which are free from latent tuber diseases. Such
tubers can become re-infected from soil inoculum and from spores in stores
and grading areas (Hide, 1989). The former is likely to be more important
as healthier seed enters th,e multiplication chain nearer the ware producer.
Soil inoculum can re-infect the highest grades and knowledge of its effects
and especially its distribution is limited. The introduction of mini-tubers
as the initial tubers for field-scale multiplication over a restricted number
of generations has been suggested as a means of reducing or eliminating
some diseases from seed used by ware producers. Similar claims were
made for the introduction of clones for VTSC and the effect has been
modest (Carnegie et al., 1981). Re-infection in field scale production seems
inevitable at present, regardless of source of initial material, and Hide's
(1989) recent evidence of contamination of early generation tubers through
contact with infected lower-grade tubers in communal stores and grading
areas suggests that progress may require a major re-think on storage of
individual crops in the multiplication chain. If isolation of tubers from
individual years is necessary substantial capital investment in new stores
will be required.
As some of these diseases, skin spot, and silver scurf, can be introduced
to and grow on tuber surfaces during storage, imposition of tolerances at
grading early in storage may not reflect the state of the tubers some months
later at planting. In addition some diseases can be carried on tubers as tiny
amounts which may ultimately infect the plant but which give no micro-
scopic symptoms on seed tubers. Thus, some seed could be infected in the
ware growers' store after delivery and lead to diseases in ware crops and
their progeny tubers apparently in contradiction of the health status given
in the certification. There is a great need for discussion of these possibilities
to be broadened so that the most effective testing of tuber health can be
developed and all growers, seed and ware, become aware of their
opportunities for preserving health. As restrictions on the use of fungicides
may increase, the preservation of health will increasingly rely on manage-
ment throughout the multiplication process.
6.3.2 Sizes
As tuber initiation takes place over a period of 14-21 days (Bean and
Allen, 1978) and there are hierarchies of tuber positions for growth, all
crops contain a range of sizes. Where whole seed tubers are replanted an
upper limit is usually specified for seed tubers and, as in the UK, a lower
limit may also be specified. For use as cut seed the seed tubers may differ
little in size from ware tubers. Chapter 7 deals with the effects of seed size
and subsequent growth and generally seed is separated into several sizes by
square mesh graders. Each size is described by its upper and lower limits
e.g. 35-55 mm and the tuber count, the number of tubers per 50 kg.
Much seed is sold over a lower limit of 35 mm so there is a considerable
quantity of unused seed between 25 and 35 mm and in many crops more
usable seed below 25 mm which is legally unsaleable. As this seed is aU as
productive as larger seed (Allen et al., 1991), changes in the legislation
relating to size are necessary. The production of seed crops should involve
much shorter growing seasons than ware crops as seed is smaller than ware
and of much greater value. A long growing season would produce
decreasing quantities of seed and increasing quantities of ware of lower
value. Much seed however is still sold on the open market and if not
purchased frequently has no real value for human consumption because of
its small size. As a consequence many seed crops are allowed to grow for
much of the season and therefore contain considerable quantities of over-
sized ware tubers. Such long growing seasons and a generally long period
from haulm defoliation to harvesting also allow considerably more disease
infection than earlier defoliation and harvesting. The latter could be
achieved by statutory defoliation dates but these have been avoided in the
UK, although essential in Holland to protect the virus health of seed in
their climate. The wide range of dates of planting and varieties used in
individual areas of the UK would make the administration of statutory
defoliation difficult and would most likely lead to considerable cost (and
argument). The increasing demand for disease-free seed is likely to
accelerate earlier defoliation and harvesting as dramatic reductions in
many diseases can be achieved in seed multiplied in the traditional manner
(Tables 6.15 and 6.16). Such changes also reduce mean tuber size in the
crops. However, smaller seed is now known to be as productive as larger
seed, so specialist seed crops defoliated before any over-sized tubers are
present would produce the required sizes of tubers and also achieve
substantial improvements in tuber health. Since such crops require no
more than 12 weeks from planting to harvesting, they could be grown in
near ideal conditions in any season as planting could be ddayed as late as
June and still allow the growth of a full crop.
Table 6.15 Incidence of black scurf (index 0-100) at intervals after

various treatments to the crop on day 0 (from Dijst et aI., 1986)
Number of days between
treatment and harvest
LSD
Cultivar treatment* 0 3 10 17 24 P = 0.05
Prominent S+CHD 11 14 32 42 12.7
S+RC+CHD 8 19 30 43
RC+6h+CHD 7 20 25 32
Haulms pulled 10 15 15 31
RC 13 15 20 29
Control (untreated) 10 14 14 16 26
* Treatments included: S = shoots reduced by stripping; CHD = chemical haulm

destruction; RC = roots cut through; 6h = shoot destroyed chemically in the
afternoon, six hours after root severing.
Table 6.16 Percentage infection of eyes following early defoliation and

harvesting (luly-early August) of Record crops in Scotland in 1989 (Allen
and Hide, unpublished)
Crop Helminthosporium Polyscytalum Rhizoctonia Number
solani pustulans solani of eyes
1 3.7 1.0 1.9 107

2 2.0 0 1.0 98
3 1.9 0 0 105
4 1.9 0 8.3 108
5 1.0 0 1.9 105
6 1.8 0 0 110
7 0 0 0 114
8 1.9 0 13.0 108
9 2.0 0 0 100
10 8.1 0 0 111
No eye showed black dot, Colletotrichum atramentarium.

6.3.3 Systems
England and Wales, Scotland, and Northern Ireland administer their own
separate certification schemes for seed potatoes but the requirements for
certification are basically similar and are defined by consultation between
the three Departments concerned. A unified approach to certification is
clearly desirable in order that the potato directives of the EEC can be
interpreted on a UK basis.
The Directive on the Marketing of Seed Potatoes lays down minimum
requirements for seed certification and marketing. The main details of the
directive are as follows.
1. There shall be two categories of seed potatoes: basic seed which is
intended for further seed production; and certified seed which is to be
used for the production of ware. These categories are defined in terms
of minimum quality standards but may be subdivided into grades of
different standard provided that these are higher than the minimum laid
down.
A protected region was established in terms of the Prevention of
Spread of Pests (Seed Potatoes) (Great Britain) Order 1974 which
comprises Scotland, the English counties of Northumberland (excluding
the district of Blyth Valley and Wansbeck) and Cumbria (excluding the
districts of Barrow-in-Furness and South Lakeland). Within this region
certification schemes include only grades within the basic seed category
and imports of any other seed from outside, whether for planting as
seed or ware, are banned except under licence. Certified seed (Grade
CC) will be produced only in the remainder of England and Wales.
2. Only potatoes certified within the categories basic seed and certified
seed may be marketed and this must be done in homogeneous batches
in new bags, sealed and officially labelled. Re-sealing m.ay only be done
officially and the facts of re-sealing must be recorded on the label.
3. Minimum sizes and permitted size ranges within a seed l.ot are defined
and tolerances for out-sizes are prescribed.
4. The treatment of seed for sale with germination inhibitors is prohibited
and any chemical treatment of the seed must be declared.
In the UK basic seed incorporates the three grades: VTSC (seed which is
multiplied from tubers produced by microplants either at the Department
of Agriculture, Fisheries for Scotland or in approved laboratories), SE
(Super Elite) and grade E (Elite) - all of which are currently produced in
Scotland. Stocks of VTSC in Scotland are grown by 'VTSC growers' on
land which has not produced potatoes for at least 7 years and clonal stocks
are subject to stringent schemes of testing for the presence of viruses, fungi
and bacteria. In general, crops cannot be entered for certification at VTSC
level before their third year of multiplication from tubers from microplants
nor after their fifth. Stocks of SE must be derived from VTSC grades and
Foreign cultivars Foreign cultivars

(low disease-risk) (high disease-risk)
! ~
New cultivars
(approved stocks)
+
Tubers/in vro cultivars
Nucleus Stock Production

Unit (DAFS)
Cultivar collection:
A: Field grown
\ '
I
I
B: in vitro
cultures
j
Tubers
~;r:PR(LA~ON Year in which
L"' ,:£:ropl'om
planted
o
I
~ pre-claJification
clonal stocks
1-2
!
VTSC 1 3
VTSC
grower
VTSC2
! 4
l
SE 1 5
!
Commercial
seed
grower
srSE 3
6
! 8
Figure 6.12 Pathway for the entry of cultivars into the Scottish Classification
Scheme. (After Jeffries, 1986).
are eligible for certification at SE level for only 3 years. Grade E seed can
be derived from E or higher grades and is without time restriction.
The possible pattern of seed movement which is illustrated diagramatic-
ally in Fig. 6.12 shows that a producer of SE stocks must plant VTSC seed
in order to produce SE stocks - the 1 designating the first year in tbe SE
grade. SE1 stocks may then be sold to the ware grower or to produce SE2
or E seed, or even CC seed in the certified seed area. The possible uses of
SE2, SE3, and E are similarly shown in Fig. 6.12.
Seed crops are certified following field inspections and have to meet
strict regulations concerning crop separation, the length of the rotation,
and the permitted levels of: gaps, groundkeepers, purity and trueness-to-
Table 6.17 Visual tolerances at final field inspection of UK seed potato

crops (Jeffries, 1986)
Seed classes
Disease/disorders VTSC SE E AA
}
Tobacco veinal necrosis virus 0.00 0.00 0.00
0.25
}
Severe mosaic 0.00 0.00
0.10
Leaf roll virus
Mild mosaic 0.00 0.05 0.50 1.00
Blackleg (Erwina carotovora
subsp, atroseptica) 0.00 0.25 0.50 1.00
Purity and trueness to type 100 99.95 99.95 99.90
Table 6.18 Visual tolerances (%) for tuber-borne fungal diseases for
certified seed potatoes (except VTSC grade) produced in the UK
(MAFF, 1984)
Individual Group Collective group
tolerances tolerances tolerances
Group ii
}
Blight
Dry rot 1
I
Gangrene 1
Group iii
I
Skin spot 2 6
Group iv
Black scurf 3 4
Common scab 4
Powdery scab 4
type, virus diseases, blackleg and potato cyst eelworm. The permitted
tolerances are progressively increased for lower grades of seed and are
shown in Tables 6.17 and 6.18.
Crops are certified provisionally on the basis of field inspections because
in terms of the new procedure following accession to the EEC, require-
ments for certification include fulfilling tuber health standards. Hence after
field inspection only provisional certificates are issued and their terms do
not require the issue of a separate final certificate since this will automatic-
ally depend on the consignment not being rejected at the time of
preparation for sale. However, the stringent UK virus health standards, in
conjunction with the climatic conditions in the main seed areas, mean that
there is no need for post-harvest virus health tests, which are an integral
part of the certification schemes of other EEC countries.
Almost half the seed planted in England and Wales is uncertified and the
proportion of seed in this category has risen in recent years for a number of
reasons: savings in seed cost; availability of the varieties such as Pentland
Crown, which are resistant to severe mosaic and moderately resistant to
leaf roll; increasing use of systemic insecticides and in early areas the belief
that home-saved seed is earlier. However, the sale of uncertified seed is
banned.
Growers obtain their own once-grown uncertified seed either as small
tubers from ware crops or from small areas devoted to their own seed
production. The latter approach allows greater control and should allow
production of healthy seed. The greatest attraction to the production of
once-grown seed is economic; the cost of seed to the ware grower can be
markedly reduced by one multiplication in England and the general
availability of virus-free certified seed allows the ware grower this oppor-
tunity. The seed grower does not see this as anything other than competi-
tion and likely to reduce the sales of certified seed. He must, however, face
the economic reality that the premium paid for certified seed is for health
(and restricted size) and such produce may be multiplied again somewhere
else. Any developments such as virus resistance which prolong and protect
the health of seed will inevitably reduce the demand for certified seed.
6.4 SEED PRODUCTION AND UTILIZATION
In the first part of this chapter consideration was given to seed growth and
development and concluded by emphasizing the important connection
between seed and ware production. A concept of physiological age was
proposed which measured the influence of seed production and storage
environment on ware crop growth through the duration of sprout growth
and storage temperature, as number of day-degrees above a base tempera-
ture. Thus, the components of a seed production system themselves can
affect physiological age at re-planting by influencing the onset of sprout
Seed production and utilization 279
growth, whose timing may influence the temperatures experienced by the
seed. Equally, manipulation of storage temperature can ensure that
variation in onset of sprout growth does not affect physiological age at re-
planting. This approach provides a framework for studying the voluminous
reports of effects of aspects of seed-crop husbandry on progeny tuber
growth, many of which provide contradictory conclusions. An appreciation
of the relationships between yield and physiological age acknowledges the
possibility of such variation and emphasizes the need for identification of
the circumstances in which the different effects would occur.
In addition to physiological effects, the methods of harvesting, storage
and marketing of seed crops are important and are considered in the final
part of this section.
6.4.1 Location
There have been reports from many countries of seed from different
sources giving different levels of sprout development, foliage size and yield
(Kawakami, 1952; Madec and Perennec, 1956; Broadbent et ai., 1961;
Iritani, 1967). These comparisons have usually been made between
imported seed stocks from different origins or between imported seed and
home-produced seed, although in general little has been known about the
way in which the imported seed has been produced and stored prior to
reaching the experimental centre.
In the UK a comparison of the growth of plants from 'once grown' seed
and from Scottish seed usually shows that 'once grown' seed emerges
earlier and forms tubers earlier (Burton, 1966). It gives a greater yield than
imported seed early in the season although the reverse may be true if crops
are allowed to grow on to maturity. Clearly the two types of seed stock
have different physiological ages which must be attributable to environ-
mental and cultural factors during the seed production year and in storage.
Where seed of both early and maincrop varieties has been produced with
similar husbandry at contrasting sites (O'Brien and Allen, 1986a) few
differences in growth and yield have been found (Table 6.19). Contrary to
many reports, seed from sites differing in temperature broke dormancy
at similar times and consequently achieved similar physiological ages at
re-planting. Similarly, attempts to detect differences in growth between
stocks of different grades of several varieties from several areas of the UK
also failed to detect any consistent effects (Allen and Wormington, 1991;
Table 6.20). Seed from crops grown in England from widely different dates
of planting were also found to produce similar yields (Table 6.21). Thus, it
is apparent that there is no specific attribute of a location which is crucial to
the subsequent performance of seed if it is healthy. However, as sites
normally use different husbandry the range in husbandry across sites in
some varieties will be very large and probably capable of explaining many
small reported effects of sites in the literature (Madec and Perennec, 1956;
Goodwin et at., 1969). Differences in husbandry almost certainly contri-
bute to effects of altitude (Kozlowska, 1963), soil type (Bardeeva et at.,
1955) and temperature (Went, 1959) which have also been reported. As
few of these authors reported much about seed crop growth and storage
the causes of the effects are difficult to establish with certainty.
Table 6.19 Effect of site of seed production on (a) date of ending of dormancy and
(b) tuber yield >38 mm (t ha- I ) on 12 July in Home Guard (O'Brien and Allen,
1986a)
Site of seed production and, in parenthesis, mean soil temperature (OC)

during seed crop growth
Abbey Rhayader Tenby Gower Rosewar Wellsbourne
Cwm-Hir
Powys Powys Dyfed West Glam Cornwall Warwickshire SE
(16.5) (18.0) (19.4) (18.5) (19.0) (21.2) days
(a) 27 Oct 23 Oct 24 Oct 4 Nov 3 Nov 28 Oct 1.9

(b) 26.1 25.4 26.6 25.0 24.7 21.4 1.32
Table 6.20 Total tuber yield (t ha- I ) in ware crops of Record from twenty seed
stocks planted and harvested at different times in Scotland
Seed stock number
2 3 4 5 6 7 8 9 10 II 12 13 14 15 16 17 18 19 20 SE
57.4 54.1 53.655.1 52.953.651.2 52.6 54.4 56.049.3 52.7 55.3 55.0 59.7 53.4 54.3 53.3 54.0 56.51.66
Table 6.21 Effect of seed source on total tuber

yield (t ha- I ) in four varieties (O'Brien and Allen,
1991)
Seed source
Variety Once-grown Scottish or
Dutch
Planted Planted
April/May July
Wilja 56.1 51.4 57.4
Maris Piper 55.8 59.8 54.7
Record 39.7 36.8 43.6
Saturna 48.0 42.5 48.2
Mean 49.9 47.6 51.0
SE 3.05
6.4.2 Husbandry
(a) Time of planting

Variation of the time at which the seed crop is planted and consequently
emerges effectively produces seed of different chronological ages by
moving the whole growth cycle of the crop. Late-planted seed crops
produce tubers which break dormancy later and have shorter sprouts and
younger physiological age at replanting than seed from earlier plantings.
This technique is used to limit the physiological age of seed in areas where
seed tends to get too old (Dubuffet, 1954). In one area of France crops of
some early varieties intended for the production of high quality basic seed
cannot be planted before a specified date in order to prevent that seed
becoming too old before it is used the next year. If time of planting does
not affect time of emergence, there are no effects on the ending of
dormancy or subsequent yield (Allen et ai., 1991).
In UK conditions the maximum variation in the time of planting is likely
to be found between early crops in Cornwall, Kent or Pembrokeshire
which produce their own uncertified seed and the same varieties planted in
Scotland to give certified seed. Scottish seed crops probably initiate tubers
2 months later and early growers have tended to prefer their own old,
'acclimatized' seed. They are observing effects of physiological age as their
own seed has a longer period for sprout growth which begins in higher
temperatures in late summer and early autumn. Such production systems
are most extreme in Kent where seed crops are planted and harvested
immediately after the early ware crops in March/April and July respec-
tively. The variety used, Arran Comet, has a short dormant period and
may be sprouting by the end of August, thus allowing 6-7 months of sprout
growth. This system produces extremely old seed which is only suitable for
early harvests of varieties such as Arran Comet which do not suffer from
'little potato disorder' at extreme ages. In other varieties this system of
seed production would restrict yields through increasing occurrence of
'little potato disorder' or in extreme cases result in complete crop failure
e.g. Red Craigs Royal.
In many varieties manipulation of the time of planting is potentially
extremely effective in altering the onset of sprouting favourably in relation
to the intended physiological age at planting. In circumstances where
increasing age is desired, a longer sprouting period resulting from an
earlier onset is a much cheaper way of achieving this than by increasing
storage temperature. In conjunction with the locality of production and
time of defoliation there is much scope for manipulating the end of
dormancy in all varieties whether produced for domestic production or
export. Development of this potential requires ware growers to perceive
the advantages and seed growers to utilize the opportunity.
(b) Time of defoliation and harvest
Most seed crops are defoliated and then left for about 3 weeks so that
tubers have set skins when harvested. Thus, any manipUlation of the time
of defoliation usually, but not invariably, involves a similar manipulation
of date of harvest. In Kent, defoliation and harvesting of Arran Comet
seed crops are frequently consecutive operations on the same day (even the
same machine) but this is unusual. Allen et at. (1991) found a small but
consistent effect of early defoliation, when canopies were approaching
peak leaf area, in achieving an earlier start to sprout growth (Table 6.22).
Few seed crops are defoliated so early and defoliations after peak leaf area
do not affect the onset of sprout growth. As the intervals between onset of
sprout growth are always less than the delays in defoliation the effects on
age at re-planting are small. They are only likely to have any commercial
significance alone in early crops where increases in age may be disadvan-
tageous after the initial harvest, especially if increased incidence of 'little
potato disorder' is caused, as found by Allen et al. (1991).
Table 6.22 Effect of date of defoliation and interval to harvesting seed

crops on date of onset of sprouting of Home Guard (Allen et aI.,
1991)
Interval to
harvesting
Date of defoliating seed crop
(weeks after
defoliating) 27 Jun 11 Jul 25 Jul 8 Aug Mean
1 23 Sep 16 Oct 25 Oct 2 Nov 17 Oct
3 4 Oct 2 Oct 22 Oct 1 Nov 15 Oct
5 13 Oct 15 Oct 26 Oct 31 Oct 21 Oct
7 30 Sep 20 Oct 27 Oct 30 Oct 19 Oct
SE 3.0 1.5
Mean 2 Oct 13 Oct 25 Oct 1 Nov
SE 1.5
From the more detailed recent experiments the effects of defoliation can
be seen to be small and influenced by the timing of defoliation in relation to
seed-crop growth, so it is not surprising that older references show no large
or consistent effects of defoliation on onset of sprout growth. Emilsson
(1949) and Burton (1963) found that seed from crops defoliated at different
times but harvested at the same time began to sprout at the same time.
Table 6.22 shows that Allen et al. (1991) also found that varying the
interval from defoliation to harvesting had little effect on onset of sprout
growth irrespective of the timing of defoliation. The severance of the link
between foliage and tubers can influence the subsequent onset of sprout
growth but only if carried out earlier in the life of the foliage than is usually
practised in commerce. However, the effects are so small that any move
towards earlier defoliation in attempts to control diseases and tuber size is
unlikely to cause problems in age of seed at re-planting.
Although the effects of defoliation are small they are additive to effects
of date of planting and therefore can contribute usefully to manipulating
age. Table 6.23 shows the range in onset of sprout growth for combinations
of date of planting and defoliation found by Jones (1981). The range is
large and even in constant temperatures would produce a wide range of
ages at planting. In the varying regional and seasonal temperatures it is
clear that a massive range in ages of any variety should be expected.
Table 6.23 Range in physiological ages of

seed at replanting and, in parenthesis, date
of ending of dormancy in Desiree from
extreme combinations of dates of planting
and dates of defoliating seed crops following
storage at 12°e (from Jones, 1981).
Planting/harvesting combination
Early planted/ Late planting/
Year early defoliation late defoliation
1975 1043 (18 Nov) 812 (17 Dec)
1976 1196 (25 Oct) 868 (5 Dec)
(c) Other husbandry factors

A number of factors have been reported to influence sprout and plant
growth but effects are invariably small and little consistency is apparent.
Although most reports of application of fertilizer to seed crops found no
effects, there are some reports of increases in sprout growth (Walker,
1968; Thow, 1970) which have also been found in early field growth but few
effects by final harvest. O'Brien and Allen (1986b) found no effects of N
over the range 0--300 kg ha- 1 and concluded that commercial application
rates were unlikely to influence seed re-growth.
Increasing the age of seed used in seed crops has been found slightly to
advance onset of sprout growth in progeny tubers of several varieties but
effects were too small to influence field growth (O'Brien et al., 1986). In
the same experiments, the authors studied the effects of two generations of
multiplication in the early area (Pembroke shire ) at different storage
temperatures on seed re-growth. The two years of multiplication were
warm and the spring of the ware year was cold and late yet few differences
were found between seed from local multiplication and certified seed. The
predominant effect was the age of the seed at re-planting and all effects
conformed to the relationships described in Fig. 6.11.
These results do not suggest that physiological deterioration of seed is
likely through repeated annual multiplication in any environment. If the
storage temperature is too high it may create 'little potato disorder' in one
storage phase for any source of seed. Observation suggests that tubers
from little potato plants will begin sprout growth earlier in the year than
tubers from normal plants and in principle there is every reason to expect a
cold storage phase to allow typical seed to be produced for the following
crop. Where seed is kept for longer than 7-8 months which is usual in
Europe, the effects of chronological deterioration may be considerable
(Kawakami, 1952) and will be greater if high storage temperatures allow
extensive sprout growth.
6.4.3 Harvesting, storage, grading and management

Seed-potato crops should be grown at higher densities than ware crops and
the principles involved in manipulating density are discussed in the
following chapter. In the past, in the UK many seed crops were grown at
relatively low densities but recently densities have been increased in an
attempt to increase the yield of seed-sized tubers. As seed tubers are
smaller and more valuable than ware tubers it might be expected that
defoliation would be timed to maximize seed yield (and minimize oversize
tubers) and therefore be relatively early in the crop's life. This is not so in
the UK and surprisingly there are few published papers from anywhere
which are concerned with the yields of seed. Many seed crops contain
substantial quantities of over-size tubers (Allen and O'Brien, 1987) and
defoliation is frequently late in the crop's life. This approach derives from
the marketing of seed being still a largely trading (non-contract) operation.
The small seed tubers have limited value if not purchased as seed so their
yield is not maximized through early defoliation once the largest tubers
approach the upper size limit (Allen and O'Brien, 1987) and a substantial
quantity of large tubers is frequently allowed to form before defoliation.
Saleable yields of seed are therefore relatively low and the rate of
multiplication (ware acreage -:- previous year's seed acreage) has not
materially changed in 25 years (Allen and O'Brien, 1987). The attitude to
defoliation in the UK contrasts with Holland where the need to avoid
aphid-transmitted virus diseases requires early defoliation and necessarily
produces lower total yields of mainly seed-sized tubers. Their harvesting is
also early in warm conditions and any skinning of tubers heals rapidly and
leads to tubers relatively free from virus and fungal diseases.
The ingress of disease inoculum is minimized by short intervals from
defoliation to harvesting and Table 6.16 shows the near disease-free state
of tubers from early-defoliated crops harvested within 2 weeks in Scotland.
Seed growers in the UK have been slow to appreciate the significance of
this aspect of their husbandry in the overall disease status of their seed. As
with ware growers, the traditional view of seed producers is that crops must
be 'mature' before they can be harvested. The basis for this view has been
seriously challenged in ware crops (Wilcockson et al., 1985) and the need
for earlier harvest is at least as great in seed crops as ware crops.
Innovations in harvesting, such as windrowing, may contribute to success-
ful harvesting in August and the crucial starting point is that fungal diseases
can be restricted most effectively by early defoliation and harvesting. The
seed grower cannot continue to defoliate in late August or September and
harvest in October if seed health is to be significantly improved.
In most crops harvested tubers are transferred to store, usually in boxes,
and cured. The temperature will be allowed to decrease after curing to
4-5°C and the tubers either remain in boxes until grading for delivery or
are treated with 2-amino-butane after 2-4 weeks. After the latter treat-
ment, tubers remain in store until grading. Generally early varieties are
delivered in OctoberlNovember and maincrops during the remainder of
the year. Price of seed increases with delay in delivery to reflect storage
costs but the quantities of seed delivered in spring have increased recently
raising problems with transport and general organization. This results from
fewer growers sprouting their seed and many wishing to avoid storage by
having seed delivered as they plant. In most main crop varieties dormancy
ends some time before delivery and frequently before grading. Cons-
equently some element of sprout damage and/or desprouting is common in
the seed used in the UK and earlier delivery of seed would reduce this
factor whose significance has not been seriously studied.
The seed crop was traditionally graded into one seed grade 35-55 mm
(or 35-60 mm for earlies) which contained a wide range of individual tuber
sizes. As seed rate is determined by seed size and mechanical planters
require seed of similar size for efficient operation the grading of seed crops
is having to change. More grades are being used and for each the tuber
count (number per 50 kg) is the effective measure of tuber size for
determining seed rates (see Chapter 7). The grading should involve 5 mm
increments especially for smaller seed in order to reduce the variation in
tuber counts per grade. Increases in the extent of grading increase the cost
of seed but accuracy in planting and economic use of seed require seed
grading more closely than 35-55 mm. Seed growers quite understandably
expect high prices for higher tuber counts but frequently underestimate the
size of the increase necessary to benefit the ware grower. This must be
determined by the minimum adjustment on the planter and equates to
about 100 tubers per 50 kg. Simply stabilizing tuber count within a
relatively wide grade is not sufficient as it necessarily results in a range of
sizes to be planted. Allen and O'Brien (1987) recognized the many
weaknesses in the current production and utilization of seed and argued
that seed crops should be valued in relation to the number of replantable
ware hectares obtained from one hectare of seed crop. This may be
calculated by dividing the seed yield of each grade by the intended ware
seed rate for the grade and they showed that up to 20 ware hectares could
be obtained from one hectare of seed of varieties with many tubers, if all
seed down to 20 mm were used. They showed that maximum value of a
seed crop occurred around the time when the first tubers approached the
upper size limit as few tubers subsequently entered the seed fraction from
below and lower value ware tubers were the result of further growth. If
seed growers are to receive a premium for seed over ware then defoliation
at maximum seed yield and value must become the norm. Close collabora-
tion between ware and seed growers is necessary to develop the ideas of
Allen and O'Brien (1987) and this is likely to occur initially where seed is
produced on contract as in the processing section of the industry. The
ultimate end point will be the purchase of whole seed crops graded into as
many grades as seed rate changes demand and valued as the number of
ware hectares X a price per hectare which accurately reflects current ware
costs. This will achieve substantial increases in multiplication rate and
inevitably reduce the number of seed hectares unless additional markets
are established. For the remaining seed growers each hectare will be more
profitable than before and other sectors of the industry will follow thereby
reducing and ultimately eliminating the trading element in seed marketing.
At grading tubers may be treated with fungicides e.g. thiabendazole but
problems with resistance are causing changes. As Hide (1989) has shown,
one application of thiabendazole may cause resistance in populations of
silver scurf, so it is advisable to use such fungicides only on seed to be used
for ware crops so that any effects due to resistance are minimized in the
seed multiplication chain.
6.5 TRUE SEED
In many developing countries high aphid populations and continuous

presence of many fungal and virus diseases make it difficult to maintain the
health of seed tubers despite seed certification programmes (Golmirzaie
and Mendoza, 1988). As relatively few diseases are transmitted through
the seed, true potato seed (TPS) produces disease-free plantlets which can
be transplanted as the planting material for the ware crop or used to
produce relatively disease-free seed tubers. In the first system, major
savings in storage of seed are achieved as a few kilograms of TPS can
replace tonnes of seed tubers. The use of TPS in potato production is not
new as it has been used by the Incas (Malagamba and Monares, 1988) and
was introduced to China from the UK in 1906 (Geddes, 1988). The
usefulness of TPS in aiding plant health in tropical environments is clear
and systems for generating large numbers of relatively healthy seed tubers
have been developed (Wiersema, 1986). However, as TPS progeny are a
collection of genotypically different individuals which show considerable
diversity in many of the important quality traits of potatoes in developed
economies, there is little prospect of TPS being used in the UK in the
near future. Considerable effort is being made at the International Potato
References 287
Center to improve TPS through selection of parental lines which produce
progenies with improved uniformity in plant and tuber characteristics.
Attention is also being paid to the factors influencing the yield of seed in
tropical latitudes as sparse flowering and reduced seed set are serious
limitations to TPS production in the tropics (Malagamba, 1988).
ACKNOWLEDGEMENTS
The authors wish to thank members of the Potato Research group and
office staff at Cambridge University Farm for their help in the preparation
of this chapter.
REFERENCES
Allen, E.J., Bean, J.N. and Griffith, R.L. (1978) Effects of low temperature on
sprout growth of several potato varieties. Potato Res., 21,249-55.
Allen, E.J., Bean, J.N., Griffith, R.L. and O'Brien, P.J. (1979) Effects of length of
sprouting period on growth and yield of contrasting early potato varieties.
J. Agric. Sci., Camb., 92, 151-63.
Allen, E.J. and Scott, R.K. (1980) An analysis of growth of the potato crop. J.
Agric. Sci., Camb., 94, 583-606.
Allen, E.J. and O'Brien, P.J. (1986). The practical significance of accumulated
day-degrees as a measure of physiological age of seed potato tubers. Field Crops
Res., 14, 141-51.
Allen, E.J. and O'Brien, S.A. (1987). An analysis of the effects of seed weight,
seed rate and date of harvest on the yield and economic value of seed-potato
crops. J. Agric. Sci., Camb., 108, 165-82.
Allen, E.J. and O'Brien, S.A. (1990) Cambridge University Potato Growers
Research Association. Annual Report 1990.
Allen, E.J. and O'Brien, S.A. (1991) Effects of timing and frequency of desprout-
ing of seed tubers on sprout and field growth of three potato varieties. J. Agric.
Sci., Camb., (in press).
Allen, E.J., Jones, J.L. and O'Brien, P.J. (1991a). Effects of date of planting and
harvesting seed potato tubers on sprout and ware crop growth. J. Agric. Sci.,
Camb. (in press).
Allen, E.J., O'Brien, P.J. and Firman, D. (1991b). An evaluation ofsmall seed for
ware potato production. J. Agric. Sci., Camb. (in press).
Allen, E.J. and Wormington, E.G. (1991). Comparisons of commercial seed stocks
of maincrop potato varieties. J. Agric. Sci., Camb. (in press).
Bardeeva, A.S., Bychkov, A.A., Mozhaev, G.!. et al. (1955). Growing seed
potatoes on peat soils. Zemledelie (1) 122-23. Abstracted in Soils and Fertilizers,
19, 103.
Bates, G.H. (1935). A Study of the factors influencing size of potato tubers.
J. Agric. Sci., Camb., 25, 297-313.
Bean, J.N. and Allen, E.J. (1978). The relationship between tuber initiation and
subsequent plant growth in field grown potatoes. Proc. 7th Triennial Con! Eur.
Ass. Potato Res. Warsaw.
Bleasdale, J.K.A. (1965). Relationships between set characters and yield in
maincrop potatoes. J. Agric. Sci., Camb., 64, 361-66.
Broadbent, L., Brown, P.H. and Wright, R.C.M. (1961). Home production of
seed for early potatoes. 2. Comparison of home produced and imported seed.
Expl. Hort., 5, 37-9.
Burton, W.G. (1963). Concepts and mechanism of dormancy, in The Growth of the
Potato, (eds J.D. Ivins and F.L. Milthorpe), pp. 17-41, Butterworths, London.
Burton, W.G. (1966). Effect of source of seed upon yield. The Potato, 2nd Ed.
Chapter 6, 130--1, Wageningen: European Association for Potato Research.
Carnegie, S.F., Adam, J.W., MacDonald, D.M. and Cameron, A.M. (1981).
Contamination by Polyscytalum pustulans and Phoma exigua var. foveata on
seed stocks derived from stem cuttings in Scotland. Potato Res., 24,389-97.
Davidson, T.M.W. (1958). Dormancy in the potato tuber and the effects of storage
conditions on initial sprouting and on subsequent sprout growth. Am. Potato J.
35,451-6.
Denny, F.E. (1926). Effect of thiourea upon bud inhibition and apical dominance
of potatoes. Contributions. Boyce Thompson Institute PI. Res. 1, 154-68.
Dijst, G., Boyman, A., Mulder, A. and Roosjen, J. (1986). Effect of haulm
destruction supplemented by cutting off roots on the incidence of black scurf
and skin damage, flexibility of harvest period and yield of seed potatoes in field
experiments. Neth. J. of Plant Path., 92,287-303.
Dubuffet, G. (1954). Experiments in improving potato growing by late planting..
C.R. hebd. Seane. Acad. Agric. Fr. 40, 84-8. Abstracted in Field Crops
Abstracts, 7, 16l.
Emilsson, B. (1949). Studies on the rest period and dormant period in the potato
tuber. Acta Agricultura Suecana, 3, 189-284.
Firman, D., O'Brien, P.J. and Allen, E.J. (1991a). Leaf and flower initiation i,n
potato (Solanum tuberosum) sprouts and stems in relation to number of nodes
and tuber initiation. J. Agric. SeL, Camb., 117, 61-4.
Firman, D., O'Brien, P.J. and Allen, E.J. (1991b) Predicting the emergence of
potato sprouts. J. Agric. Sci., Camb. (in press).
Geddes, A.M.W. (1988). Introduction of potatoes to remote mountain area of
China through TPS in 1906. Asian Potato Association Proc. 2nd. Triennial
Conference, Kunming, China. Manila: Asian Potato Association.
Gillison, T.C., Jenkins, P.D. and Hayes, J.D. (1987). Some factors affecting the
expression of the physiological age of potato seed tubers. J. Agric. Sci., Camb.,
108, 437-5l.
Golmirzaie, Ali M. and Mendoza, H.A. (1988). Breeding strategies for true potato
seed production. c.I.P. Circular Vol. 16. No.4, Dec. 1988. Lima, Peru.
Goodwin, P.B. (1967). The control of branch growth on tubers. I. Anatomy of
buds in relation to dormancy and correlative inhibition. J. of Exp. Bot., 18,
(54), 78-99.
Goodwin, P.B., Brown, A., Lennard, J.H. and Milthorpe, F.L. (1969). Effect of
centre of production, maturity and storage treatment of seed tubers on growth
of early potatoes. II. Field growth. J. Agric. Sci., Camb., 73, 167-76.
Gray, D. and Bleasdale, J.K.A. (1972). The production of potatoes for canning.
Bull. Potato Marketing Board. London.
References 289
Hartmans, Klaasje J. and van Loon, e.D. (1987). Effect of physiological age on
growth vigour of seed potatoes of two cultivars. I. Influencc of storage period
and temperature on sprouting characteristics. Potato Res., 30, 397--409.
Hide, G.A. (1989). Practical progress in potato production - Disease control. Proc.
V.T.S.C. Growers Conf. Aviemore. Nov.
Holmes, J.C., Lang, R.W. and Singh, A.K. (1970). The effect of five growth
regulators on apical dominance in potato seed tubers and on subsequent tuber
production. Potato Res., 13, 342-52.
Holmes, J.e. and Gray, D. (1972). Carry over effects of sprouting and haulm
destruction in the potato seed crop. Potato Res., 15,220-35.
Iritani, W.M. (1967). Some factors affecting productivity of Russet Burbank seed
potatoes. Am. Potato J., 44, 153-8.
Jarvis, R.H. and Palmer, G.M. (1973). Effect of type of planter on the growth and
yield of maincrop potatoes. Expl. Hush., 24,29-36.
Jarvis, R.H. and Rogers-Lewis, D.S. (1974). Population studies with Pentland
Ivory and Record Potatoes. Expl. Hush., 27,23-30.
Jeffries, e.J. (1976). Symposium on healthy planting material. B.C.P.C. Mono.
No. 33. 239--47.
Jones, J.L. (1981). Effect of date of planting on contrasting potato varieties. PhD
Thesis. University College of Wales, Aberystwyth.
Jones, J.L. and Allen, E.J. (1983). Effects of date of planting on plant emergence,
leaf growth and yield in contrasting potato varieties. J. Agric. Sci., Camh., 101,
81-95.
Kawakami, K. (1952). Physiological aspects of potato seed tubers. Mem. Hyogo
Univ. Agric., 2, 1-114.
Kawakami, K. (1973). Proc. Fifth Triennial Conf. Eur. Ass. Potato Res. 1972.
151-2. Norwich, England.
Krijthe, N. (1946). The influence of storing seed potatoes on the structure of the
buds and the development to a full-grown plant. Meded. Landh Hoogesn
Wageningen, 47, (6).
Krijthe, N. (1962). Observations on sprouting of seed potatoes. Eur. Potato J., 5,
316-33.
Kozlowska, A. (1963). Differences in growth and metabolism of potatoes grown in
mountains and in the lowlands. Eur. Potato J., 6, 143-59.
Madec, P. and Perennec, P. (1956). Influence of the origin on the behaviour of
potato plants. Annis. Amelioration des plantes, 6, 5-26.
Madec, P. and Perennec, P. (1962). The relationship between the induction of
tuberization and plant growth in the potato (Solarum tuherosum L). Annis.
physiologie vegetale, Paris, 4, 5-84.
Malagamba, P. (1988). Potato production from true seed in tropical climates. Hort.
Science, 23, 495-500.
Malagamba, P. and Monares, A. (1988). True potato seed: past and present uses.
Lima: International Potato Centre. 37pp.
Milthorpe, F.L. (1963). Some aspects of plant growth, in The Growth of the Potato,
(ed. J.D. Ivins and F.L. Milthorpe), pp. 3-16. Butterworth, London.
Ministry of Agriculture, Fisheries and Food. (1984). Control of diseases of
potatoes. Bulletin 2388.
Moorby, J. (1967) Inter-stem and inter-tuber competition in potatoes. Eur. Potato
J., 10, 189-205.
Morris, D.A. (1967). Intersprout competition in the potato. II. Competition for
nutrients during pre-emergence growth after planting. Eur. Potato J., 10,
296-311.
O'Brien, P.J. and Allen, E.J. (1986a). Effects of site of seed production on seed
yields and regrowth of progeny tubers in potatoes. J. Agric. Sci., Camb., 107,
83-101.
O'Brien, P.J. and Allen, E.J. (1986b) Effects of nitrogen fertilizer applied to seed
crops on seed yields and regrowth of progeny tubers in potatoes. J. Agric. Sci.,
Camb., 107, 103-11.
O'Brien, P.J. and Allen, E.J. (1991) Effects of seed crop husbandry, seed source,
seed tuber weight and seed rate on the growth of ware potato crops. J. Agric.
Sci., Camb. (in press).
O'Brien, P.J., Jones, J.L., Allen, E.J. and Raouf, G.S.M. (1986). Effects of
physiological age of seed tubers on seed yield and regrowth of progeny tubers in
potatoes. J. Agric. Sci., Camb., 107,307-27.
O'Brien, P.J., Allen, E.J., Bean, J.N., Griffith, R.L., Jones., S.A. and Jones, J.L.
(1983). Accumulated day-degrees as a measure of physiological age and the
relationships with growth and yield in early potato varieties. J. Agric. Sci.,
Camb., 101, 613-31.
Potato Marketing Board (1979-83). Potato Statistics in Great Britain. London:
PMB pub!.
Reust, W. and Munster, J. (1974). Pregermination des plants de pomme de terre
avec ov sans choc thermique. Revue Suisse Agric., 6, 9-12.
Sadler, E. (1961). Factors influencing the development of sprouts of the potato.
PhD Thesis. University of Nottingham.
Schepers, A. and Hoogland, R.F. (1968). Relation between sprout development in
seed potatoes and number of mainstems in the crop. Jaarb. [nst. bioi. Scheik.
Onderz. Landb Gewass. 47-53.
Taylor, C.E. (1953). The vegetative development of the potato plant. Ann. appJ.
Bioi., 40, 778-88.
Thomas, M.N. (1988). The control of tuber size in maincrop potatoes. PhD Thesis.
University of Cambridge.
Thomas, W.L. and Eyre, R.W. (1950) Early potatoes. Faber and Faber, London.
Thow, R.F. (1970). Second generation effect of NPK on the potato. PhD Thesis.
University of Edinburgh.
Toosey, R.D. (1964). The pre-sprouting of seed potatoes: factors affecting sprout
growth and subsequent yield. Part 1. Field Crop Abstracts, 17, 161-8.
Turl, L.A.D. (1983). The effect of winter weather on the survival of aphid
populations on weeds in Scotland. EPPO Bulletin No. 13, pp. 139-43.
van Hiele, F.J.H. (1961) Un sprouted potato tubers treated with gibberellic acid
(GA3). Eur. Potato J., 4, 26-39.
van Loon, C.D. (1987). Effect of physiological age on growth vigour of seed
potatoes of two cultivars. 4. Influence of storage period and storage tempera-
ture on growth and yield in the field. Potato Res., 30,441-50.
van der Zaag, D.E. and van Loon, C.D. (1987). Effect of physiological age on
growth vigour of seed potatoes of two cu.ltivars. 5. Review of literature and
integration of some experimental results. Potato Res., 30, 451-72.
Walker, M.G. (1968). The effects of nutrient treatments of potato plants on the
performance of their progeny. PhD Thesis. Bath University of Technology.
References 291
Went, F.W. (1959). Effects of environment of parent and grandparent generations
on tuber production by potatoes. Am. J. Bot., 46,277-82.
Wiersema, S.G. (1986). A method of producing seed tubers from true potato seed.
Potato Res., 29,225-37.
Wilcockson, S.J., Allen, E.J., Scott, R.K. and Wurr, D.C.E. (1985). Effects of
crop husbandry and growing conditions on storage losses of Pentland Crown
potatoes. J. Agric. Sci., Camb., 105,413-35.
Wright, R.C. and Peacock, W.M. (1934). Influence of storage temperatures on the
rest period and dormancy of potatoes. U.S.D.A. Technical Bull. 424.
Wurr, D.C.E. (1975). Relationships between sprouting characters and stem
development in two maincrop potato varieties. Potato Res., 18, 83-91.
Wurr, D.C.E. (1979), The effect of variation in the storage temperature of seed
potatoes on sprout growth and subsequent yield. J. Agric. Sci., Camb., 93,
619-22.
Wurr, D.C.E. and Allen, E.J. (1976). Effects of cold treatments on sprout growth
of three potato varieties. J. Agric. Sci., Camb., 86, 221--4.
Wurr, D.C.E. and Morris, G.E.L. (1979). Relationships between the number of
stems produced by a potato seed tuber and its weight. J. Agric. Sci., Camb., 93,
403-9.
CHAPTER 7
Plant density
E.J. Allen and D. C.E. Wurr
7.1 INTRODUCTION
At the end of dormancy some buds on seed tubers grow into sprouts and
after planting a variable proportion of these sprouts develop into main
stems. A potato plant consists of a variable number of stems, the exact
number depending upon the size and treatment of the plant's parent tuber.
The individual stems of a plant are largely dependent upon their parent
tuber for growth until emergence and a considerable time may elapse
after emergence before the stems become fully independent. Thus, the
competitive interference between stems of a potato plant changes con-
siderably with time and this, together with variation in the number of stems
per plant, necessarily results in the effects of plant density being complex.
As variation in the number of stems per plant may be associated with
variation in their size and performance it is clearly unsatisfactory to assess
density in this crop on the simple basis of number of plants per unit area.
The establishment and use of a sound unit of plant density is particularly
important in potatoes for seed costs still amount to between 30% and 50%
of total growing costs (Potato Marketing Board, personal communication)
and because plant density affects the number of tubers set, which in turn
affects the size distribution of those tubers. For all outlets of potatoes, the
size of the tubers is an important quality criterion and it is therefore
important to obtain the correct plant density. In practice, plant density in
the potato crop is manipulated through the number and size of the seed
tubers planted. This sounds simple yet there is a need to establish the most
appropriate unit of plant density which should be capable of explaining the
effects of all its component parts, namely seed size, within and between-
row spacing (and their combination in seed rate), number of stems and
spatial arrangement, on total and graded yields of potatoes. There is also a
need to consider whether the response to plant density or indeed the most
appropriate unit of plant density is affected by other agronomic factors
such as the physiological age of the seed tubers because tuber age markedly
Units of density 293
affects stem growth and development, as shown in Chapter 6. It is,
therefore, logical to begin a consideration of the effects of plant density in
the potato crop with the possible units of density and then use the most
appropriate unit to illustrate the effects of plant density.
7.2 UNITS OF DENSITY
The establishment of accurate relationships between tuber yield per unit area
and plant density requires a definitive, repeatable unit of plant density to be
identified. A number of possible units have been suggested and discussion of
their merits must precede any discussion of the effects of plant density.
7.2.1 Number of eyes

Our knowledge of the growth of individual eyes is poor, yet it is important
to know whether there are differences between eyes in the growth of and
the yield from the stems which comprise the popUlation. Birecki and
Rostropowicz (1963) found no differences in the performance of excised
eyes of uniform weight from different sizes of seed tuber. De Carvalho
(1975) stated that eyes from large seed tubers were larger than eyes from
small seed tubers and the differences were affected by variety. Data,
however, were not presented. Allen (1979) examined the performance of
different eyes within tubers of the same size and total eye number (8) by
dissecting out eyes to produce single-eye tubers. As can be seen in Table
7.1, differences between eye positions in number of stems and tubers and
tuber yield were small. Single-eye tubers did not differ significantly from
intact tubers in number of above-ground stems which suggests that few, or
indeed only one eye, developed in the intact tubers and observation of the
few such tubers which survived until harvest confirmed that most stems
arose from one eye, that at the apex. As Allen and Wurr (1973) have
observed that most stems in commercial crops arise from the 'apical
complex', there is evidence to suggest that the majority of eyes on seed
tubers do not grow. From the limited data available, it appears that there
are only small differences between eyes which do grow. Experiments which
report high proportions of eyes producing stems have not usually explained
how the figures were obtained and have probably regarded one sprout
(stem) as the growth of one eye. This indirect assessment is probably
invalid in view of the evidence above and detailed study of the growth and
development of eyes is long overdue.
The seed tuber is clearly wasteful of growing points, especially when
larger seed tubers are used and it is surprising that little effort has been
directed towards increasing the proportion of eyes which do grow into
stems. In addition to seed sprouting, which was discussed in the previous
chapter, cutting of the seed tubers is one possibility. In North America this
is standard practice, for seed tubers are rarely sprouted and the varieties
294 Plant density
Table 7.1 Effect of eye positioTllon performance of single eye tubers
(Allen, 1979). Variety - King Edward
Eye position Number of above- Number of tubers Total yield
ground stems
(000s ha- 1) (OOOs ha- 1) (t ha- 1)
(Apical) 1 103.2 1421.8 35.1
8 86.2 1277.3 35.8
7 87.1 1312.3 36.4
4 102.3 1156.1 32.3
3 92.5 1295.2 39.6
(Heel) 2 86.2 1203.7 35.0
Control (all) 96.0 1490.0 45.5
SE ±6.13 ±69.49 ±2.25
produce relatively few, large tubers. In the UK it has been used for early
potato production (lng, 1966) but is of no practical significance at present.
The possibilities for increasing the number of eyes which develop and
thereby the stem density are illustrated in Table 7.2. Cutting large seed
tubers into various sized pieces produced more stems than whole tubers
and the effect increased as the size of seed piece was reduced. Cutting of
the seed tuber breaks the dormancy imposed on many eyes by the apical
bud, probably aided by wound-induced gibberellins (Rappaport and Sachs,
1967). The small seed pieces were grown using normal ridge culture which
retarded their emergence and usually resulted in reduced final yields; much
of this deleterious effect could be removed by more appropriate planting
depths. Table 7.2 also shows that half a large (113 g) tuber was equivalent
to a whole tuber of the same size, a finding reported by other workers in
the absence of disease (Stuart et al., 1924; Brandreth and Bryan, 1937).
This should allow for considerable economy in the use of larger seed for, as
will be discussed later (Section 7.4.7), the seed rate required for small seed
is lower than that for large seed. Machinery for cutting seed tubers is
available and a detailed study of their cutting should be undertaken,
particularly as the maincrop acreage in the UK is increasingly being
planted with unsprouted seed and the seed is free from virus diseases.
Use of number of eyes as the scale of plant density was suggested by
Bleasdale (1965) but, as our understanding of the growth and development
of eyes is poor and the number which develop is known to be affected by
the sprouting regime, the scale has considerable deficiencies. There are
also serious practical difficulties, for it is extremely difficult to count eyes
because such counts depend upon the ability to recognize the lateral buds
which have separated sufficiently from the apical bud to form individual
eyes. For eyes close to the apical bud this is extremely difficult, especially
in varieties with either rather deep-seated eyes or where there is minimal
physical separation away from the apical eye, e.g. Maris Piper and
Table 7.2 Effect of seed cutting on number of above-ground
stems and tubers (OOOs ha·1 ) (Allen, 1979)
(a) Planting material
Whole Sixteenths Peel SE
1969 Stems 186.0 346.5 244.5 ±15.54
Tubers 784.5 938.6 823.8 ±57.72
1970 Stems 151.4 207.7 ± 7.67
Tubers 400.6 552.0 ±19.19
1971 Stems 144.4 191.0 ±28.1O
Tubers 482.2 550.0 ±80.40
Variety - Majestic
(b) Data for equivalent plant (whole tuber) densities

Whole Half (56 g) Half cut Whole
(113 g) cut laterally longitudinally (56 g)
1971 Stems 301.6 184.7 172.7 179.4
Tubers 750.7 692.3 617.9 607.2
Variety - Pentland Crown
Pentland Crown. Nonetheless the number of eyes has been used as the unit
of plant density but the proportion of eyes has not been constant over seed
sizes and unequal stem densities have resulted (Thompson and Taylor,
1974). Eyes may also fail to grow because of the effects of pathogens.
Skinspot (Polyscytalum pustulans) has been shown to kill up to 25% of
eyes on tubers of Majestic stored in bulk until planting (Hide et al., 1969)
and this pathogen is so widely distributed on stocks that its effect on the
number of eyes which grow is still important. Griffith et al. (1974) showed
that gangrene (Phoma exigua) killed eyes and also increased the number of
sprouts and stems which grew from the remaining eyes. Such effects of
disease upon the number of sprouts and stems growing from seed stocks
are likely to influence responses to plant density, especially as the stems
from seriously-infected tubers may be less productive than stems from
healthy tubers. For general application, the number of eyes as a scale of
density has limited usefulness.
7.2.2 Number of sprouts

The difficulties in defining sprouts have already been discussed (Chapter 6,
Section 6.2.3). As the majority of the maincrop acreage is planted with
unsprouted seed, and for sprouted seed the number of sprouts growing into
stems is variable, the use of number of sprouts as the unit of plant density is
impractical.
296 Plant density
7.2.3 Number of seed tubers
The use of such a scale assumes that all sizes of seed have similar numbers
of stems and since this has already been shown to be untrue (Chapter 6,
Section 6.2.3) the scale is unjustified. However, for practical purposes the
number of seed tubers has great relevance provided that recommendations
(see Section 7.4.7) take account of seed tuber size.
7.2.4 Surface area of seed tuber

Reestman and de Wit (1959) found a linear relationship between the
number of mainstems in the crop and the surface area of the seed tuber.
They proposed the surface area of the seed tuber as a general unit of plant
density. Such a scale is unlikely to be of practical value and there are
reservations about the accuracy of the methods used for calculating the
surface area of the tuber (Bleasdale, 1965).
7.2.5 Seed rate

This is the weight of seed planted per unit area and is calculated as the
product of the number of seed tubers planted and their average weight.
Growers are familiar with this unit of plant density because they buy seed
by weight, appreciate variation in average tuber size and for every crop
must decide upon the within and between-row spacing and hence the
weight of seed to be planted. The use of any other unit of plant density
must necessarily be achieved by planting the appropriate number of tubers
of the particular seed size.
In view of the great practical relevance of seed rate it is not surprising
that the majority of reported experiments on potato densities have used
seed rate as the plant density scale; those experiments which have not used
the scale can be assessed by it since the number and weight of the seed
tubers used are usually recorded. These experiments may be summarized
by examining the component effects of seed size and within-row spacing
(number of tubers) and then discussing the general applicability of seed
rate as the plant density scale.
(a) Effect of seed size

The results of a number of relatively old experiments examining the effects
of seed size on total and graded yields were summarized in Table 7.7 of the
previous edition of this book (Allen, 1978) and the conclusions were:
1. total yield and yield over small riddle sizes increase with increase in seed
tuber size;
2. as the riddle size over which yield is considered increases, there is a
tendency for yield to increase with seed size to a maximum and then
decrease. Further increase in the riddle size may completely reverse the
effect of seed size, so that more large ware is produced by small seed
than by large seed.
(b) Effect of within-row spacing

The effects of within-row spacing were also summarized in Table 7.7 of the
first edition (Allen, 1978). The effect of reducing spacing is similar to the
effect of increasing seed size and similar conclusions to those made for seed
size apply.
In very few experiments has a significant interaction between seed size
and spacing been reported, although with large riddle sizes or small
fractions of yield the tendency towards interaction increased. Thus, the
merits of seed rate as the plant density scale may be assessed without
consideration of possible interaction effects.
In 1954 Boyd and Lessells used data from many British experiments to
examine the effect of seed rate on the yield of maincrop varieties. They
showed that both total and ware yield increased over the range of com-
mercial seed rates, while yield >51 mm reached a maximum and then
decreased. By constructing curves showing the effect of altering seed rate
by variation in seed size or within-row spacing they found that increasing
seed tuber size had a smaller effect on total yield but a larger effect on
percentage ware than reducing within-row spacing. They concluded,
however, that provided the optimum seed rate was attained, the precise
combination of seed size and spacing was of minor importance in determin-
ing ware yield over 38 or 45 mm riddles. This conclusion led to the view
that seed rate was a reliable and accurate plant density scale for potatoes.
The findings of Boyd and Lessells have profoundly influenced the con-
sideration of potato density effects in the UK for the acceptance of seed
rate as the plant density scale has been difficult to alter. Their conclusions
were not generally shared by growers who until recently have considered
large seed to be more productive than small seed.
Since 1954 a number of workers have reported a superiority of small
seed over large seed at equal seed rates, particularly at seed rates outside
the usual commercial range. The results of these experiments were
summarized in Table 7.10 of the first edition (Allen, 1978) and show for a
range of varieties that small seed out yielded large seed at the same seed
rate. Holliday (1960), EI Saeed (1963) and Holmes (1966) explained that
these effects resulted from the higher stem densities established from small
seed at equal seed rates, for the relationship between number of stems and
seed size is frequently asymptotic and where linear requires large increases
in seed tuber weight to achieve small increases in number of stems per seed
tuber. Thus, with the larger seed sizes, large increases in seed tuber size
produce small increases in the number of stems. As a consequence of these
results and the advocacy of Holliday that the stem was the true unit of plant
298 Plant density
density for potatoes, a detailed examination of stems has been made by
several workers.
7.2.6 Stems
Although the interest in stems as the unit of plant density is of recent origin
it is notable that Arthur (1892) reported that increases in 'stalks' (stems)
per hill gave increases in total and graded yields and Bates (1935) also
appreciated the importance of stems. There are three types of stem which
may emerge above ground. Mainstems grow directly from the seed tuber
and therefore may be derived from main or lateral buds (Krijthe, 1955).
Secondary stems do not grow from the seed tuber and may arise either as
sub-surface branches from buds on the underground part of a mainstem or
where stolons emerge above ground and bear leaves. Mainstems always
bear tubers but the two types of secondary stem may not bear tubers (Allen
and Wurr, 1973).
Differences in sprouting regime cause differences in the proportions of
the different types of stem produced by seed tubers and this affects the
number of tubers set and the growth and longevity of the haulm. Table 7.3
shows that a short sprouting period produces densities comprising main-
stems while a long sprouting period produces relatively few mainstems but
many secondary stems resulting from lateral branch development on
the sprouts. Generally, stems produced by maincrop varieties are pre-
dominantly mainstems because seed tubers have either no sprouting period
or only a short sprouting period while early varieties produce a mixture of
main and secondary stems. Thus, sprouting affects the number and the
type of stems which occur and it may be anticipated that there will be
differences in the growth and yield of individual stems derived from
different sprouting treatments.
Whatever type of stem density is involved, in experimental work it is
essential that it is recorded accurately but data on both differences in field
stem densities and the life of individual stems are limited. Allen and Wurr
Table 7.3 Effect of length of sprouting period on number and types of stems
produced in the field (Allen et aI., 1979)
Sprouting period
From From From SE
harvesting mid-November mid-January
Above-ground stems per
plant 4.71 4.79 4.75 ±0.441
Mainstems per plant 1.88 3.04 3.66 ±0.202
Secondary stems per plant 2.83 1.75 1.08 ±0.453
Variety - Home Guard. Data are means of two temperatures and two seed sizes.
(1973) recorded the stem densities of a sample of commercial crops in East
Anglia by harvesting plants in mid-season in order to determine the origin
of the stems. They found that mainstems accounted for >99% of the
above-ground stem density and suggested the latter as an easy way of
recording the effective plant density in a growing crop. Many of th,e other
published reports of stem densities have not indicated how or when the
counts were made and as the definition of a mainstem involves the seed
tuber, some assumptions may have been made in making these counts.
Without actually harvesting plants or removing soil covering the seed
tuber, accurate determination of mainstems is difficult, if not impossible
and must involve either the acceptance of all above-ground stems as
mainstems or some subjectivity in deciding upon the origin of stems. For
example, Thompson and Taylor (1974) called mainstems 'those likely to
bear tubers', an unsatisfactory basis for recording, especially as Allen and
Wurr (1973) had reported that the size of a stem was a poor guide to the
number and size of the tubers carried. It is likely that many published
estimates of mainstems contain such assumptions. In some cases there are
no references to secondary stems which may indicate wholly mainstem
densities or the dismissal of secondary stems as unimportant. It is desirable
to have much more detailed information on stem density and on the factors
affecting the proportions of the different types of stem. In this context it is
of interest that Allen (1977) found that secondary stems were more
common on maincrop varieties grown in Pembrokeshire than he had found
in East Anglia and their number increased over a long period. This
suggests environmental influences during crop growth on the proportion of
each type of stem and is worthy of greater study.
The timing of stem counts is also important for it is known that stems die
and may disappear as a result of competition before the harvest of the crop
(Bleasdale, 1965; Wurr, 1971). Counts made at haulm death or after
defoliation are likely to be prone to underestimation, while counts taken
early in the season must be taken after complete emergence. Scott and
Younger (1972) reported that the number of mainstems per seed tuber was
determined at an early stage but Ifenkwe (1975) found that it increased
slowly for almost 2 months after emergence. Much more information is
required on the number, type, emergence pattern and longevity of stems
for efficient stem recording to be practised especially where few counts are
to be made. At present it appears that maincrop varieties should be
counted in mid-season and in the absence of definitive evidence that only
mainstems are present, all experimental counts should involve harvesting
of the plants.
Sometime after emergence the mainstems arising from the seed tuber
assume an independent existence and the plant resulting from one seed
tuber becomes a collection of competing stems. As large seed tubers
usually produce more stems than small seed tubers, the spatial arrange-
ment of a given stem density is quite different from different seed
300 Plant density
Variety - Majestic
60
•• • • 60
• • •
50 t t • 50 tr
40 40
30 30
.,_ 20 20
2 10 10
~ O~--~----~---r---'--~ O+----r--~--~----r_--~
1::J 0 1·5 3 4·5 6 7·5 o 50 100 150 200 250
+> Seed rate (t ha- 1 ) Number of mainstems
'0 (000'5 ha- 1 )
"0
]i
Variety - King Edward
c
+>
{!.
30
• • 30
•
20 20
10 10
0,4------r----~----~--
o 1 2 3 o 37·5 75 112·5
Seed rate (t ha- 1 ) Number of mainstems
(000'5 ha- 1)
Figure 7.1 Relationships between tuber yield, seed rate and stem density
(Bleasdale, 1965) . • , seed weight 32g; . , seed weight 64g; ., seed weight 128g.
sizes. This effect is termed 'clumping' and is discussed later in Section

7.3.4.
Bleasdale (1965) produced a detailed examination of the mainstem as
the unit of plant density. Using data from several sources he plotted yields
against seed rate and the number of mainstems. When plotted against seed
rates, seed sizes appeared to give independent curves but when plotted
against mainstems the curves became indistinguishable. Bleasdale concluded
that total yield and the proportions in the size grades were a function,
independent of seed size, of the number of mains terns per unit area (Fig. 7.1).
Sharpe and Dent (1968) agreed with Bleasdale's findings and found that a
square root function gave the best fit to their data for both total and graded
yields. Subsequently, a number of workers including Gray (1972) using
mainstems and Wurr (1974) using above-ground stems have found that
stems are capable of relating yields by a single line for a range of seed sizes.
Close relationships between tuber yields and the number of stems have
not always been found. Goodwin et al. (1969) found no relationship
between yield and mainstems, for at the same above-ground stem density,
seed-tubers with many mainstems and seed-tubers with single well-branched
mainstems gave similar yields. The authors considered there was confusion
over the term mainstem and many other workers had included sub-surface
branches in their counts. Nevertheless, these results do not discredit the
use of above-ground stems as the plant density scale. Holmes (1973)
rejected stems as a useful plant density scale for, in experiments where
stem density was varied by sprouting regime, such a scale proved quite
inadequate to relate yields. In these experiments the stem densities ranged
from wholly mains terns (from short sprouting periods) to mainly secondary
stems (from long sprouting periods) and in these circumstances it is likely
that number of stems will not be capable of relating all treatments on a
common scale. The performance of mainstems may be affected by the
presence or absence of branches but more importantly the effects of long
sprouting periods are to increase the physiological age of the seed tubers,
resulting in earlier senescence and lower yields, compared with seed from
shorter sprouting periods (Chapter 6). Thus, the growth and longevity of
the stems is affected by the means of manipulating plant density. In
practice crops would be grown from one seed type or the other, not a
mixture of both, but it is of importance to know whether the response to
increasing plant density is the same in two such contrasting types of seed.
It must be emphasized that the evidence which most strongly supports
the use of stems as the plant density scale is still from experiments with
maincrop varieties sprouted in such a way as to produce almost entirely
mainstems. Even with maincrop varieties it has been found by Jarvis (1971)
and Jarvis and Rogers-Lewis (1974) that the varieties Pentland Crown and
Record show larger responses to increasing seed size than would be
expected if stem density eliminated differences between seed sizes. The
reasons for this are not clear though Section 7.4.7 presents a possible
explanation. Other varieties, King Edward, Pentland Dell and Pentland
Ivory, have been found to respond to seed rate in a way consistent with the
use of stems as an overall unit of plant density. Maincrop varieties do
produce secondary stems but little is known of the factors affecting their
production and even less about the influence of such stems on effects of
plant density.
7.2.7 Use of stems as the density scale

It is with early potato varieties that the greatest ranges of type and number
of stems per seed tuber are produced, for early ware crops of many
varieties are produced from seed with few or single sprouts which
commonly branch while late harvested ware and seed crops are produced
from seed which usually has several sprouts which develop into mainstems.
302 Plant density
Unfortunately the effects of plant density in such varieties are sparsely
reported in the literature. The use of higher seed rates than used for
maincrop varieties is commonly noted (e.g. Cole, 1973) and the merits of
large seed, usually in direct comparison with small seed, are indicated (e.g.
Scott and Younger, 1972), although the data from which these comments
derive are limited. Bean (1981) studied the effects of plant density on the
early variety Home Guard in Pembrokeshire and found that there was
little difference between seed rate and above-ground stems as the plant
density scale for apically-dominant seed while both scales were better
than mainstems (Fig. 7.2). All seed sizes produced similar numbers of
mainstems but the larger seed produced more secondary stems and the
number of above-ground stems was similar at any seed rate for all seed
sizes. Mainstems were a poor scale, as seed tuber size had the same
number of mainstems but widely different tuber yields. There was little
evidence to support the use of large seed for early production. With seed
from short sprouting periods Bean found that differences between seed
rate, main stems and above-ground stems as plant density scales were very
small at early harvests (June). Above-ground stem densities were again
similar for all seed sizes at any seed rate although this seed produced
few secondary stems. The number of mainstems increased linearly with
increasing seed tuber size principally because the smallest seed produced
only one stem per tuber. As Fig. 7.2 shows, the relationship between total
yield and any plant density scale was quite different for the two types of
seed. These data support the findings of Holmes (1973) that marked
differences in the sprouting of seed may prevent tuber yields being related
to number of stems by a single line but they also indicate that seed rate is
equally inadequate in this respect. Where marked differences in sprouting
occur, it is essential to restrict plant density responses to a specific type of
seed. Number of above-ground stems and seed rate appear to be equally
useful as plant density scales within a specific type of seed of Home
Guard and other early varieties when harvested from May to July (Allen,
unpublished) .
It is desirable to establish whether such effects of sprouting on the
response to plant density and on the most appropriate plant density scale
exist in other varieties. Even if the response to plant density is not affected,
large differences in numbers of stems per seed tuber may be expected to
affect the combination of seed size and spacing (i.e. seed rate) required to
achieve the optimum stem density. At present it may be concluded that a
'stem' scale is usually as effective as any other and in the case of the
reported responses of maincrop varieties, rather better.
Acceptance of the stem as the most appropriate theoretical unit of plant
density in the potato allows for effects of plant density to be assessed
through the effective unit rather than through seed rate which has
limitations. A scale based on stems allows easy interpretation of the effects
of seed size, within-row spacing and seed rate via their effects on stem
3 30 (b)
20
10 •
,/.~
v·-
E
O~~----~--~---c----
~ 01 2 3 4 5 6 7 8 9 10 1 234 5
lTd
~ Seed rate It ha- 1) Number of main stems x 105 ha- 1
:J I
~ 2
/' .~'
~A
o
o 1 2 3 4 5
Number of above-ground stems x 105 ha- 1
Figure 7.2 Relationships between tuber yield >25 mm and (a) seed rate, (b)
mainstems and (c) above-ground stem densities for apically dominant (open
symbols) and multi-sprouted (closed symbols) seed of the variety Home Guard
(Bean, 1981).0, seed weight 40g; !::" seed weight 70g; D, seed weight 100g.
(a) y = - 9.88 + 17.3 Yx - 2.57x, R2 = 98% for multi-sprouted seed and
y = - 5.96 + 8.55 Yx - 1.11x, R2 = 87% for apically dominant seed;
(b) y = - 7.48 + 2.05 Yx - 0.04x, R2 = 97% for multi-sprouted seed and
y = - 5.04 + 1.53 Yx - 0.04x, R2 = 61% for apically dominant seed;
(c) y = - 9.95 + 2.26 Yx - 0.04x, R2 = 97 % for multi-sprouted seed and
y = - 10.93 + 1.63 Yx - O.03x, R2 = 89% for apically dominant seed.
density though it cannot be used directly in practice as yet. Figure 7.3

shows the relationship between total and ware tuber yields and the above-
ground stem densities for two main crop varieties in two seasons (Wurr,
1974). The relationship between the total yield curve and the ware yield
curves and the point of maximum ware yield vary according to variety and
season. Thus, establishment of more than the general principles of plant
density-yield relationships in this crop must be specific to variety, probably
location and age of seed. This is discussed in detail in Section 7.4.7.
The curves shown in Fig. 7.3 may be taken to illustrate the effects of seed
size and within-row spacing. Increasing seed size or decreasing within-row
spacing or both (and therefore increasing seed rate) result in more stems,
304 Plant density
50~ 50~
25 .... . ..................... .............. ........ ..........
25 .~~......................... :~._..............~
'"
ro O~--~~~~--~~~ OL-__- L____L-__- L____L-~.j

~ 0 100 200 300 400 0 100 200 300 400 500
~ Number of above-ground stems (OOO's ha- t )
(e) 30 (d)
20 20
~ --------
:;ioI"._o_._._
';"t'.~.- ........... ..... ~
,/
...... ............. .....
....... .......
,
10 10 ".
00L---t.....L0-0--20.L0--3~00 00 tOO 200 300 400

Number of above-gr.ound stems (OOO's ha- t )
Figure 7.3 Relationship between stem density and tuber yields for (a) Pentland
Crown, 1969; (b) Maris Piper, 1969; (c) Pentland Crown, 1970; (d) Maris Piper,
1970. - - , total yield; - - - - yield >38 mm; -'-'-' yield >44 mm; ...... yield
>S1mm (Wurr, 1974).
with reduced spacing having the greater effect. The optimum stem density
for ware yield is lower than for total yield, so that increases in the number
of stems are more likely to exceed the optimum stem density for ware yield
than total yield and reductions in ware yield from large seed sizes and close
within-row spacing would therefore be expected. Figure 7.4 presents
model responses of tuber yields to changes in seed sizes and within-row
spacings and expressed as number of tubers and seed rate which for seed
rate may be compared with the results of recent individual experiments in
Fig. 7.5. At equal within-row spacings large seed produces a greater total
yield than small seed until the spacing is reached at which the small seed
achieves the optimum stem density. This within-row spacing will be much
closer than that at which large seed achieves the same stem density. At
closer within-row spacings there will be no differences in total yield
between seed sizes. A similar situation exists for ware yields but the actual
values change. In addition, ware yield decreases at close within-row
spacings and for large seed this will occur at a wider spacing than for small
seed. The relationship between large and small seed is therefore dependent
upon the range of within-row spacings (i.e. stem densities) considered. If
seed sizes are compared at equal seed rates, their relative positions are
reversed, for the small seed produces more stems per unit seed weight as
already described. Any comparison between treatments is determined by
their stem densities and the relative effects of treatments may change if
they are compared at widely different parts of the yield-stem density
response curve. There is no convincing evidence that seed sizes differ in
total yield when each is grown at the optimum stem density. The extensive
and often confusing literature on plant density-yield relationships in the
potato crop is made easier to interpret when it is appreciated that the stem
is the basic unit of plant density. It is no longer necessary to make the
comparisons detailed above, for treatments should be compared over a
range of stem densities and when treatments produce different densities no
valid comparisons can be made.
(0)
// .. /-_.
"",,< /
//
/'>
~ ,'/~
]! _ _ _ Small
>- Numbers of tubers -
.... seed
QI _._._. Medium
.0 seed
~ (b) ------------ Lorge
seed
/
/
The decline in tuber yields
where large numbers of seed
/ ./ tubers and high seed rates
are used applies primarily to
/." wore yields. Total yields may
;/ decline where growing
conditions are poor (see text)
~/
','
Seed rate~
Figure 7.4 Schematic representation of effects of seed size on relation:ship

between tuber yield and (a) number of seed tubers planted; (b) seed rate. (Not to
scale.)
In conclusion it may be argued that a plant density scale involving

stems has been shown to relate tuber yields better than any alternative.
Differences between treatments and the changes in their relationships can
be explained via their stem densities and the yield-density response curve.
It is intended simply to refer to stems from hereon in this review. It must be
reiterated, however, that stem densities cannot be used in a predictive or
advisory context without the detailed knowledge of seed tuber develop-
ment already found lacking in Chapter 6 and considered more extensively
in Section 7.4.7. This deficiency in knowledge is capable of remedy and the
306 Plant density
70
f'
o
60
• •
0'
o
/
50
C?
f' 40
•
""E
E
A70
0
•
32
Q)
>=
60
•
50
40
0 2 3 4 5 6
Seed rate (Vha)
Figure 7.5 The relationship between yield >40 mm (t ha- 1) and seed rate (t ha- 1)
in two experiments carried out on silt soils (variety, Cara). 0, 35g seed; e, 105g seed.
continued statement of its existence ought to stimulate its solution rather

than inhibit the use of the stem as the unit of plant density.
7.3 SPATIAL ARRANGEMENT
The discussion of plant density so far has only considered the number of
plants or stems per unit area, but the way in which this plant density is
arranged may affect its performance. In potatoes the spatial arrangement
of the density of stems may be affected by changes in row width,
irregularity of within-row tuber spacing, failure of plants to emerge and by
clumping of stems resulting from the use of seed of different sizes.
7.3.1 Row width

The traditional row widths used in the UK are from 61 to 76 cm, with the
early crop being produced on the narrowest rows. However, wider rows
are required to aid clod and stone separation and mechanical harvesting of
the main crop and row widths up to 90 cm are now in commercial use. At
similar plant densities, changes in row widths within the range 60--90 cm
have not been found to affect tuber yields markedly (Patzold, 1964;
Spatial arrangement 307
Ziegler, 1968; North and Proctor, 1971; Jarvis and Shotton, 1972) for any
slight reduction in total yield has usually been compensated by improved
saleability through reduced greening and cracking. Studies on quite
marked deviations from square plant arrangement by Saunt (1960) and
Eckersall (1965) have also shown that the potato is remarkably tolerant of
changes in its plant arrangement. Attempts to grow the crop in beds
(Nelson, 1967; Jarvis and Shotton, 1968) with narrow row widths have not
shown any advantage and have revealed serious obstacles in this type of
spatial arrangement with respect to soil cover and tuber growth.
Few attempts have been made to establish the response of plants to
changes in spatial arrangement but recently Fowler (1988) studied
the effects in two contrasting varieties Estima and Pentland Dell. He
compared square planting with a rectangularity of four at a range of
densities and dates of planting. Square planting increased early leaf growth
and yields and reduced stem length but by final harvest differences in yield
were small because plants on wider rows senesced more slowly and
achieved similar yields to square planting. Fowler considered that effects
could be explained in terms of amount of intercepted radiation. He argued
that in practice determinate varieties which produce a fixed number of
leaves per mainstem, such as Estima, were less suited to wide rows than
indeterminate ones because their limited number of leaves made them
vulnerable to any restriction on leaf expansion (e.g. limited water avail-
ability) leading to inadequate leaf cover. The use of such varieties on 90 cm
rows without irrigation is likely to produce variable results as has been
found on the silt soils around The Wash in the UK.
Although the evidence is not extensive, it suggests that for maincrops the
choice of row width within the range considered is not an important factor
in determining yields. There is limited information on the effects of spatial
arrangement on early potatoes but since these crops are of shorter duration
than maincrops and are composed of much smaller plants it is suggested
that the narrower the row width the better. Information on this question is
required for it is intimately connected with attempts to mechanize all
aspects of early potato production.
The cultural advantages which accrue from widening rows may not be
maximized at the 90 cm row width and it is relevant to ask whether even
wider rows could be used. Studies on the response of potatoes to row
widths beyond this level are limited. Allen (1972) compared 69 with 138 cm
rows and found that although the use of the wider row width significantly
reduced yields, the reduction was smaller than expected. Ifenkwe (1975)
continued this work in the wetter environment of Pembrokeshire, compar-
ing 66 and 132 cm row widths at a range of plant densities for two maincrop
varieties. The wider rows reduced tuber yields early in the season but this
difference became smaller as the season progressed so that differences in
total and ware yields at the end of the experiments were small. Greening
and growth disorders of tubers were always reduced by wider rows so that
308 Plant density
saleable yields were unaffected by the change in row width (Fig. 7.6). At
the end of the experiments tuber yield could be satisfactorily related to
stem density by a single line independent of row width (Fig. 7.7). This
evidence from an environment not normally associated with maincrop
potatoes would suggest that wider rows than 90 cm should be seriously
examined, especially where irrigation is available, for in Ifenkwe's
experiments water was only seriously limiting once in three years.
Fowler (1988) and Ifenkwe (1975) found that increasing row width
reduced number of tubers. The effect was usually small but sufficiently
consistent to be of importance in affecting the graded yields from similar
stem densities. Where number of tubers is of greatest importance, as in
seed crops, the narrowest row widths are to be preferred.
_60 Desiree p60 Maris Piper

,/
'0 7
....
J:: /'
!! 50 ./' 50 r'
/' 7
...g"". 40
II
/'
/
P
40
~ 30 30
"
:is
0
- - - 66 em rows
.....
.!! 20 20 0-·-0-·-0132 em rows
0
0
.7 " '/
~ 10 10 ~/
Ii
>=
0
June July I August I Sept. June I July I August I Sept.
Figure 7.6 Effect of row width on yield of saleable ware tubers (>38 mm less
greened and cracked tubers) in two maincrop potato varieties. Data averaged over
five planting densities (Ifenkwe, 1975). Vertical lines are LSDs.
~ 60 Desire. 60 Maris Piper

CD
I"l o
.66 em rows
0132 em rows
.
~ 50 o • 50
.Q"
:::I
"40
'0
:!i! 0
40 ~
.!!30L---~--~--~--~ 30~~~~~--~--~~~--~
>- 50 100 150 200 250 100 150 200 250 300 350 400
Numbers of above-ground stems (000'5 ha- 1 )
Figure 7.7 Relationships between tuber yield and numbers of above-ground stems
for combinations of five planting densities and two row widths of two maincrop
potato varieties, Desiree and Maris Piper (Ifenkwe, 1975). Desiree: y = 26.2 +
O.l1x, R2 = 87%; Maris Piper: y = 15.63 + O.236x - O.0004x2 , R2 = 81 %.
7.3.2 Irregularity of within-row spacing
While experimenters plant their seed tubers uniformly down the row, the
adoption in commercial practice of mechanical planters has resulted in
increased irregularity of spacing in farm crops. Planters of the belt-fed type
are prone to extremes of irregularity unless very carefully set and operated.
Table 7.4 shows the variation in within-row spacing found in a range of
commercial crops planted with cup-feed planters. Such crops are clearly
suffering considerable irregularity and the data also show that the mean
spacing differs appreciably from that intended in most cases. The seed rate
(i.e. plant density) is usually lower than intended which is highly relevant
to the discussion of the practical application of our knowledge of effe.cts of
density in Section 7.4.7. The effects of irregularity are that tuber yields are
relatively unaffected by moderate irregularity (20-40%) but may some-
times be reduced by extreme irregularity especially after a short period of
growth (Table 7.5). Irregularity reduces early leaf growth in a similar way
to increased row widths but, over time, effects become small. Early potato
crops should in principle be planted as regularly as possible.
Table 7.4 The intended, mean, maximum and minimum (em) values
and the coefficient of variation (CV) of seed spacing of surveyed potato
crops (Booth and Allen, unpublished)
Seed size Intended Mean Maximum Minimum CV
(mm) (em) (em) (em) (em) (%)
Site 1 35-55 20 26 60 8 39
29 33 90 11 38
35-40 25 32 72 12 38
45-50 29 32 75 11 34
50-55 29 35 74 5 41
Site 2 35-55 28 26 69 5 39
45-50 32 32 77 7 38
50-55 36 35 83 12 34
Site 3 40-45 28 26 87 7 41
45-50 31 30 80 9 4{)
50-55 37 32 75 0 35
Site 4 25-35 14 19 66 5 40
7.3.3 Missing plants

The failure of planted seed tubers to emerge because of disease andlor
damage will affect the spatial arrangement of the remaining plants. In
North America the use of cut seed frequently results in losses of 10-15% of
seed pieces and protracted emergence (Chase et at., 1989). It has been
310 Plant density
Table 7.5 The effect of within-row spacing and irregularity
of planting on total, and ware yield (40-65 mm) (Booth and
Allen, unpublished)
Total yield
CV(%)
Spacing (cm) 0 20 40 60 Mean
23 34.6 31.9 34.1 34.6 33.8
27 32.7 30.8 34.4 31.0 32.3 SE
35 35.1 32.7 32.2 30.0 32.5 0.56
Mean 34.1 31.8 33.6 31.9
SE 0.65
Ware (40-65 mm)

CV(%)
Spacing (em) 0 20 40 60 Mean
23 25.2 23.7 25.9 26.9 25.4
27 25.1 23.9 27.0 24.0 25.0 SE
35 28.7 26.0 26.1 23.5 26.1 0.63
Mean 26.3 24.5 26.3 24.8
SE 0.72
shown by Reestman (1970) that at high plant densities, missing plants have
relatively little effect, a not unexpected result in view of the flat-topped
nature of the yield-stem density relationship. At very low plant densities
there may be no compensation and missing plants will directly reduce
yield. As the proportion of missing plants increases the compensation by
the remaining plants eventually becomes insufficient and yield decreases.
The incidence of virus and fungal diseases such as gangrene (Phoma
exigua) , skinspot (Polyscytalum pustulans) and black scurf (Rhizoctonia
solani) and variation in depth of planting affects the emergence and growth
of stems considerably and creates heterogeneous stem densities in respect
of leaf growth, number of tubers and canopy longevity. The influence of
these diseases upon responses to plant density is worthy of greater
attention. The effect of depth of planting has hardly been studied but Table
7.6 shows the range in depth in many farm crops and suggests that variation
in soil conditions and planter operation frequently create substantial
effects. The use of seed of wide-ranging sizes e.g. 35-55 mIn will exacer-
bate the situation.
7.3.4 Clumping
The number of stems per seed tuber increases with increase in seed size, so
that similar stem densities have quite different spatial arrangements when
produced from small and large seed tubers. The existence of a relationship
Table 7.6 The intended, maximum and minimum (mm) values of seed
depth and the coefficient of variation (ev) of surveyed potato crops (Booth
and Allen, unpublished)
Seed size (Within-row Intended Maximum Minimum CV
(mm) spacing) (mm) (mm) (mm) (%)
Site 1 35-55 (20 cm) 167 236 94 20
35-55 (29 cm) 177 235 102 12
35-40 164 245 102 15
45-50 164 234 98 13
50-55 165 228 117 12
Site 2 45-50 111 157 60 17
50-55 112 176 32 28
Site 3 40-45 143 197 103 14
45-50 151 214 101 17
50-55 176 266 115 17
between yield and stem density through a single line, suggests that the
effect of this stem clumping is unimportant. Nonetheless it is essential to
consider whether this situation, implicit in the adoption of stems as the
density scale, is justified. Bleasdale and Thompson (1965, 1966) planted
one, two and three seed tubers per plant position using the variety Majestic
at similar stem densities for two years. They found that stem clumping had
no effect in one year but, in the other, yields from the single seed tubers
were superior to those from the clumped seed tubers. However, in view of
the good relationship between tuber yield and stems, Bleasdale concluded
that for practical purposes clumping has only a small effect. Reestman and
de Wit (1959) considered that large seed were inferior to small seed
planted at the same skin surface area because of clumping effects (although
Bleasdale (1965) considered that this was caused by errors in calculating
the surface areas). In Sweden, Svensson (1966, 1972) has examined the
effect of stem arrangement on yields using single sprout pieces to build lip
differing stem arrangements. Although he has shown that yields may be
reduced by clumping, the growing period was usually short, not more than
12 weeks and it seems likely that the greatest effects of clumping will be
early in the season with compensation occurring later. Clumping may
affect the numbers of tubers set and consequently have a much greater
effect on yields of specific grades. Data on this point are lacking; any
marked effects of clumping on tuber size distribution would favour the use
of smaller seed.
312 Plant density
OL-----~----~----~--_,
100 200 300 400
Number of stems (OOO's he-I)
Figure 7.8 Relationship between number of tubers per stem and number of stems
per unit area (after Scott and Younger, 1972).
7.4 EFFECTS OF PLANT DENSITY
Following the discussion in Section 7.2 it is intended to consider stems as

the effective unit of plant density and to discuss the effects of stems on the
growth, development and yield of the crop. It is accepted that these stem
densities may be achieved in different ways and no mention of this is made
unless essential to understanding.
7.4.1 Number of progeny tubers

The number of tubers set by plants is determined by stem density, spatial
arrangement, variety and environment. Any increase in the stem density
over the commercial range results in a reduction in number of tubers set
per stem (Fig. 7.8). If density is increased by planting larger seed tubers
then number of tubers per plant will increase despite the reduction in
number of tubers per stem (Fig. 7.9). Where stem density is increased by
planting more seed tubers, number of tubers per plant will decrease as
number of tubers per stem decreases and number of stems per seed tuber
remains unaffected (Fig. 7.10). Increasing stem density over a wide range
for all varieties, and the whole explored range in some varieties, results in
increased numbers of tubers per unit area (Allen, 1972; Gray, 1972). As
tuber yield is relatively constant over much of this stem density range the
increase in number of tubers allows for manipulation of tuber size. Those
Effects of plant density 313
17
..
c
16
.
.
-£15
II
Q.
f14
II
.Q
::J
..
A
:: 13
0
A
II
~ 12 A
::J
z
11
56 lt2 168 224

Size of seed tuber (9)
Figure 7.9 Relationship between number of tubers per plant and size of seed
tuber (after Scott and Younger, 1972).
100
..
c
0
80
Ci.
~ 60
.
...
Q.
II
~40
::J
z
20
0 • •
. •
12 27 47 109
Number of seed tubers plonted (000'5 ha- 1 )
Figure 7.10 Relationship between number of tubers per plant (,6.), number of
stems per plant (e) and number of seed tubers planted (after Scott and Younger,
1972).
varieties such as Pentland Crown (Wurr, 1971) which do not increase in

number of tubers beyond a certain stem density are restricted in their
control of size. Spatial arrangement affects number of tubers in a similar
manner to density, for increasing rectangularity (which reduces within-row
spacing) reduces number of tubers set per stem.
These general effects of density on numbers of tubers are widely
314 Plant density
accepted but the absolute values show considerable variation. Table 7.7
shows the range in number of tubers in different seasons for a number of
older experiments. These data were used in the first edition of this book
and much of the variation was ascribed to availability of water. In the
intervening 14 years two aspects have changed. First, it is now recognized
that most of such variation is in the tuber initials and tubers <10 mm which
contribute little to yield or size grading. Methods of counting tubers based
on swellings twice the diameter of the stolon lead to such variation.
Number of tubers> 10 mm from the same seed planted at similar densities
at different locations are frequently much more stable (Table 7.8).
Secondly, the causes of the remaining variation are much less clear than
was suggested previously. Severe restriction of incident radiation (Thomas,
1988) and water availability (Thomas, 1988; Stalham, 1990; Firman, pers.
comm.) have been shown to produce only small reductions in the number
of tubers> 10 mm. The treatments producing these effects were so extreme
that they are unlikely to occur in practice, as with other published effects of
water supply (MacKerron and Jefferies, 1986), the causes of variation in
numbers of tubers in practice remain obscure and therefore at present
uncontrollable. For a process long regarded as central to the growth and
development of the crop, tuber initiation has received remarkably little
serious study in field-grown plants. The literature is full of comments about
it but in most cases it was never measured and inferences were made whose
relevance can never be properly assessed. Unfortunately, many of these
inferences have entered the conventional wisdom of the crop and inhibited
creative thought and measurement. For example, there are many
references to the sensitivity of tuber initiation to daylength but no
published evidence showing the effect of a factor which may affect crops
over the range of normal dates of planting within the UK and would
certainly be important if varieties were grown extensively throughout
Europe over the whole year, as is likely. Although varieties usually
produce the same ranking order for number of tubers >10 mm, the
components, number of stems per seed tuber and number of tubers> 10
mm per stem, are diverse (Wurr et al., 1991). As Table 7.9 shows, varieties
with few or many tubers can arise from disparate components. Serious
study of the whole process of development leading to initiation of tubers in
relation to environmental conditions is essential for practical progress in
control of plant density and hence number of progeny tubers.
Because increase in number of progeny tubers continues beyond the
stem densities giving maximum ware and total yield, it is poss,ibl:e to reduce
average tuber size greatly by using stem densities in excess of those
required for maximum ware yields. Crops for canning and seed which
require small tubers with precise upper and lower riddle sizes are grown at
stem densities above those for ware production. The control of tuber size is
not very precise, for, apart from seasonal fluctuations in the number of
tubers, it seems likely that all tubers are not equally affected by increasing
Table 7.7 Effect of season on numbers of above-ground stems and tubers produced
from similar seed rates of different varieties
Number of Number of
Author Variety Location Season above-ground tubers
stems
(ooos ha- 1) (OOOs ha- 1)
Allen King Edward Cambridge 1969 299.0 1173

(1972) 1970 233.5 522
Up-to-Date 1969 365.5 1045
1970 275.5 544
Majestic 1969 228.5 732
1970 209.5 430
Jarvis and King Edward Terrington 1965 88.7
Shotton 1966 148.3
(1971) 1967 103.3
Bremner and King Edward Nottingham 1959 123.4 1542.5
Taha (1966) 1960 243.5 971.6
Majestic 1959 111.4 1136.3
1960 141.9 691.8
Bean Arran Comet TreHoyne 1975 146.7 552.9
(1978) Pembs 1976 282.7 1007.5
King Edward 1975 146.7 1279.3
1976 254.6 1332.4
Gray Maris Peer Welles bourne 1969 590.0 2660.0
(1972) 1970 550.0 1490.0
Ifenkwe Maris Piper TreHoyne 1972 230 821
(1975) Pembs 1973 235 780
Desiree 1972 182 580
1973 148 520
Table 7.8 Number of tubers >10 mm (OOOs ha- l ) in

Estima for different N applications on six mineral soils in
1989 (Allen and Fowler, unpublished)
N Site
(kg ha- 1) 2 3 4 8 10
0 484 476 417 479 462 744
60 549 540 498 532 533 837
120 512 630 498 537 538 821
180 540 637 495 542 584 852
240 557 653 528 580 572 766
SE 16.1 19.8 11.3 13.2 12.7 22.0
316 Plant density
Table 7.9 Varietal mean data adjusted to a common seed-tuber
density of 40000 ha-l (Wurr et aI., 1991)
Variety Stems per Daughter tubers Total number
seed tuber per stem of tubers (ha- 1)
Pentland Squire 3.9 3.6 450
SE 0.12 0.14 18.9
Cara 6.0 2.8 526
SE 0.33 0.08 25.4
Estima 3.9 4.4 580
SE 0.12 0.09 16.2
Maris Piper 5.2 3.2 612
SE 0.18 0.08 25.8
King Edward 4.7 5.4 788
SE 0.23 0.28 32.0
Table 7.10 Effect of very high seed rates on yields of large-

sized tubers
Author Seed rate Yield (t ha- 1 ) Number of
(t ha- 1) >51 mm tubers >51 mm
(OOOs ha- 1)
Allen (1972) 4.06 20.1 117

8.11 18.1 115
16.22 13.0 85
Data are mean of three varieties.
competition and little is known about the factors involved. As Table 7.10
shows, stem densities vastly in excess of those required for ware yield do
not eliminate yields in large-sized fractions. This suggests that tubers differ
in their response to competition and some may be preferentially positioned
in relation to supply (Gray, 1973; Gray and Smith, 1973; Oparka, 1987) so
that they remain largely unaffected by competition while others are 'labile'
and with increased competition may suffer and perish. These differences in
tuber position may be reflected in different tuber growth rates and patterns
of growth. Wurr (1973) and Ahmed and Sagar (1981) have reported that
individual tubers do differ in their growth rates and elucidation of the
causal factors and the extent of their effects would greatly help the
manipulation of tuber size through number of progeny tubers, for it may
enable the range of tuber sizes per stem (plant and crop) to be reduced.
The more regular spatial arrangements of stems resulting from the use of
small seed are usually reflected in slightly enhanced tuber set which does
not influence total or ware yields but can affect the yield of the finer grades.
Bean (1981) found that small seed increased the number of tubers set
compared with larger seed at similar stem densities and this affected the
grading of the yield from similar total yields. The significance of this effect
would depend upon the financial aspects of producing for outlets with
apparently precise size limits.
7.4.2 Total yields

Figure 7.3 shows the relationship between total yield and stem density for
harvests at, or approaching, complete senescence. Total yield increases
to a maximum with increasing stem density and then either remains
unchanged with further increase in density or eventually begins to decline.
The relationship appears to be 'flat-topped' in most reported cases (e.g.
Sharpe and Dent, 1968) and any decline in yield at high stem densities
seems to occur when growing conditions are poor, especially where water
is limiting (Wurr, 1971; Ifenkwe, 1975). The absence of a marked point of
inversion in the curve is important, for small changes about the optimum,
which may result from the practical methods of establishing crops, are
unlikely to affect yields markedly.
It is not intended to review the functions which have been used to relate
tuber yield to plant density in the potato, but some comment is necessary.
The fitting of quadratic and especially square root functions (Sharpe and
Dent, 1968; Wurr, 1974; Wurr et al., 1990) has been found satisfactory and
optimum stem densities have been derived, but such functions have no
biological significance. Other functions have been suggested, notably those
of Bleasdale and Nelder (1960), but no definitive evidence of their
biological validity has been produced either.
The general relationship between stem density and total yield would
appear to hold, for all varieties, for seed produced under specific sprouting
regimes. Whether marked changes in sprouting and seed preparation aHect
the optimum stem density or only the seed rate required to achieve the
single optimum still needs to be established. Variety markedly affects the
optimum stem density and although the relative positions of varieties may
remain the same from year to year, the absolute values change (Wurr,
1974). These differences in optimum stem densities between varieties are
of great importance and are amplified in the following section on ware
yields.
Within any variety, the length of the growing season may affect the
optimum stem density, for the longer the season the lower is the stem
density required for maximum total yield. Thus, over the range of stem
densities explored in most experiments, total yield is commonly linearly
related to stem density early in tuber growth and the relationship shows a
more pronounced optimum stem density as harvesting is delayed. The
highest stem densities increase leaf area early in the season and thereby
light interception and improve early tuber growth but this advantage may
subsequently be counterbalanced by increased leaf senescence, especially
if water becomes limiting and lodging occurs (Ifenkwe, 1975).
318 Plant density
7.4.3 Graded yields
The relationships between graded tuber yields and stem density (Fig. 7.3)
show more pronounced optima than the curve for total yield. The larger
the size grade considered, the more pronounced is the optimum and the
lower is the optimum stem density (Wurr, 1974). The relationship between
the stem densities for optimum total and graded yields depends upon the
proportion of the total yield which falls into the grade concerned. This is
determined by the number of tubers per stem; where this is low and inter-
tuber competition limited, most of the yield will be in the larger sizes and
differences in optimum stem densities for ware and total yields will be
small. Where the number of tubers per stem is large, inter-tuber com-
petition will reduce average tuber size and the difference in optimum stem
densities will be large. Thus varieties which set many tubers per stem require
very much lower stem densities, for graded yields than for total yields. Such
varieties, e.g. King Edward and Maris Piper, are inappropriate for the
production of the largest size grades. Notwithstanding this basic character-
istic, Maris Piper is widely grown to produce baking-sized potatoes
although yields are frequently unsatisfactory as would be expected.
The large differences between varieties in their response to stem density
must be appreciated if appropriate varieties are to be selected for produc-
tion for specific outlets. Further, the responses to increasing stem density
explain why crops for differing outlets are grown at widely different stem
densities. Table 7.11 illustrates the differences in stem density required for
early seed, canning and maincrop ware production. For the first two outlets
very small tubers are marketable and high stem densities are established; in
the case of the canning crop the variety Maris Peer is used, which sets
many tubers per stem so that very high numbers of tubers are produced
and the resulting inter-tuber competition restricts tuber size. For seed
crops, stem densities intermediate between those for canning and ware are
required, because seed tubers are larger than canners but smaller than
ware. It is remarkable that few studies of the effects of stem density for
seed production have been produced and Allen and O'Brien (1987)
showed that high densities were justified for maximum returns from the
crop. If the objective is to produce large ware then low stem densities
especially of varieties with relatively few tubers per stem are required. It is
essential to realize that optimal yields of tubers of specific sizes can only be
achieved by appropriate stem densities which may be quite different from
the densities required for normal ware yields.
The length of growing season affects the optimum stem density for
graded yields considerably. For early harvests of any size grade, lower
densities are required which result in limited inter-tuber competition and
rapid attainment of the required size. Yields are, however, low. For longer
growing seasons, higher and higher stem densities are justified and higher
yields will result. This situation has been demonstrated for canning
Table 7.11 Optimum stem densities for different types of crop
Optimum stem
Author Type of crop Variety population Comments
(Ooos ha- I )
Bean (1981) Early ware Home Guard 400 For harvests
in early June
Wurr (1974) Maincrop ware Maris Piper 286 Data of year
Pentland Crown 270 1969
Large ware Maris Piper 10
>51 mm Pentland Crown 102
Jarvis and Maincrop ware King Edward 150
Shotton (1971)
Sharpe and Maincrop ware Desiree 263
Dent (1968)
Gray (1972) Canning Maris Peer 700
Allen and Seed Record 300-400
O'Brien (1987) Maris Piper 500
potatoes by Gray (1972). For production of graded tubers over a period of

time, crops grown at low stem densities are harvested first and larger stem
densities later. A similar situation exists in choosing varieties for very early
production. Ulster Prince which sets very few tubers per stem (sometimes
only one) rapidly achieves a marketable sample of very small tonnage but
does not bulk rapidly unless very high and uneconomic plant densities are
established in comparison with the more favoured varieties. Consequently
this variety is only used for the very earliest harvest in early areas. In
earlies, the rapid fall in market price in mid-season results in lower
economic optimum stem densities at late harvests (Bean, 1981).
During the season the minimum riddle changes through both statutory
control from the Potato Marketing Board and the commercial specifica-
tions of buyers. For varieties which are harvested over long periods for
different purposes a considerable range of stem densities will be required
for the individual crops. In most cases the data do not exist to calculate the
precise optima but the principles explained here can be applied to help
maximize yield in the required grades. As market specifications are
changing so frequently and new products appearing, it is inevitable in
future that choosing densities for crops will require informed application of
basic principles.
7.4.4 Quality and saleability

High stem densities produce tubers with higher dry-matter contents than
similar-sized tubers from low stem densities and the size of progeny tubers
320 Plant density
also affects tuber dry-matter content (Wurr and Allen, 1974). As stem
density affects tuber size distribution its effect is considerable and probably
deserving of further examination. Detailed consideration of quality is given
in Chapter 12. Stem density affects the saleability of the crop by influencing
the prevalence of tuber greening and growth disorders. Unfortunately
there are few quantifiable reports of effects which separate these two
factors as they are usually considered together. Ridges of satisfactory size
and construction greatly reduce the incidence of tuber greening and this is
one of the advantages of using wide rows. Large numbers of stems per
plant, as from the use of large seed, often inc.rease greening, for the growth
of large numbers of tubers in a restricted soil volume results in tubers being
forced through the upper surface of the ridge and exposed to light (Scott
and Younger, 1972; Ifenkwe, 1975). Effects of stem density on tuber
secondary growth are small but the incidence of such tubers is usually
decreased by increasing stem density (Jarvis and Shatton, 1968; Scott and
Younger, 1972). The prevalence of such disorders in the variety Majestic
was as much a reflection of inadequate stem densities as varietal propensity
for secondary growth.
The effects of stem density on saleability are composed of several
opposing factors and a detailed examination of them has not been
reported. Their overall effect is usually small and occasional inconsistency
in reports should not be viewed with concern; for example Gray (1972),
who never had more than 6% unsaleable tubers, found that unsaleable
tubers increased with increasing stem density. Of greater importance are
the effects of stem density and row width on the damage caused to tubers
during the harvesting operation. Detailed data on this aspect are scarce
and difficult to obtain but effort should be made to examine this question.
The work of Jarvis (1972) has shown that the adoption of 90 as opposed to
76 cm rows allowed for faster rates of harvesting with slightly lower levels
of tuber damage. The former point is important as it should allow more of
the crop to be harvested in early autumn when conditions for harvesting
are more favourable. Stem density and row width also affect tuber size and
unpublished work by Jarvis has shown that smaller tubers are less liable to
damage than larger tubers. Thus, increasing stem density and widening
row width exert opposing effects upon the number of tubers and therefore
the tuber size distribution and possibly upon damage during harvesting. A
more detailed study of these effects is still required.
The value of saleable potatoes is increasingly affected by the prevalence
of skin diseases e.g. silver scurf, and the transmission of inoculum from
seed tuber to progeny tuber is an important component in disease
expression. Table 7.12 shows that large seed tubers result in more disease
in progeny tubers than small seed tubers and thus the quality of tubers
resulting from similar stem densities can be affected by the size of tuber
used to establish the required density.
Table 7.12 Effect of seed size on silver
scurf (% surface infected) of progeny tubers
in Estima (Firman, unpublished)
Seed size SE
Small Large
Date of harvest: 22 Aug 88
After storage
at Sutton Bridge 26.4 30.2 1.52
at Lords Ground 19.8 23.6 1.76
Date of harvest: 26 Sep 88
At harvest 4.2 9.7 1.58
After storage
at Sutton Bridge 28.7 33.1 1.56
At Lords Ground 17.9 23.3 0.99
Date of harvest: 7 Sep 89
At harvest 2.2 3.4 0.33
Date of harvest: 5 Oct 89
At harvest 4.9 7.7 0.54
7.4.5 Plant morphology and growth

Increasing stem density results in taller plants with much reduced axillary
branching (Ifenkwe, 1975). In the same experiments it was shown that the
use of wider rows at the same stem density resulted in taller plants and
restricted axillary branches. These gross changes in plant morphology have
been known in general terms for some time (Taylor, 1953) but their
implications in the study of effects of stem density on crop growth have
been neglected. These changes in density affect the standing ability of the
crop, and the tendency of crops grown at high stem densities to lodge
deserves greater attention, for it affects the ability of the crop to intercept
and utilize light. It probably represents a major factor restricting the
response to increased stem density in those crops where foliage collapse
occurs early in the season. No studies appear to have considered the types
of stem and leaf arrangement which would fit the plant for growth at high
stem densities. Discussion of desirable plant forms are conspicuously
absent from the literature of this crop. It is remarkable that since Bremner
and Taha (1966) reported their work on the growth and development of
different stem densities, no further detailed developmental study of
different stem densities has been published in the UK. Much work on tuber
growth has been carried out to which reference has already been made but
information on the whole crop is sadly deficient. As stem density affects
the morphology of stems and leaves throughout the life of the crop, it
322 Plant density
should be used to examine the influence of these factors on growth
and yield formation. A beginning should be made by carefully record-
ing the canopy structure of selected crops of contrasting densities
throughout their life, together with their light interception, growth and
development.
7.4.6 Nutrient supply and uptake

A positive interaction between stem density and N supply was reported by
van Burg (1967) and Bodlaender and Reestman (1968) but few other
convincing examples of this have been reported. The optimum stem
density was higher when fertilizer N was used than when none was
applied but differences in optimum stem densities at different levels of
applied N were small. Nitrogen uptake has been shown to be an important
factor in determining leaf longevity (Dyson and Watson, 1971) and as N
concentration within the plant usually declines with increasing density
(Scott and Younger, 1972), it might be expected that the larger canopies of
high plant densities would justify more N, especially in long season crops.
However, effects of stem density on root morphology and efficiency of
nutrient uptake are largely unknown and a complete examination of the
interaction between stem density and nutrition, especially N, is still
awaited.
7.4.7 Economic and advisory considerations

The general principles of effects of stem density presented in the preceding
sections are widely known and generally accepted. In order to be of
practical value they have to lead to the availability (and use) of accurate
recommended economic optimum densities for specific circumstances.
However, the prediction of number of stems per seed tuber and progeny
tubers per stem is still inadequate to allow seed tuber spacing to be
adjusted in relation to achieving an optimum number of stems with a
particular seed lot. There are examples of close relationships between
number of stems and seed tuber sizes for specific storage environments
which appear stable over seasons (Thomas, 1988) but the data are not
extensive enough for use in recommendation. It is clearly unfortunate that
little attention has been directed to this subject in recent years in the UK
and this failure illustrates a growing difficulty in applied research. The
general principles are known but they have not been used, primarily
because their application requires a co-ordinated national approach as no
individual researcher can have the resources adequately to test the
robustness of the relationships on which national recommendations must
be based.
Table 7.13 Estimated number of tubers formed per stem at a
planting density of 40 000 seed tubers ha- J for the potato variety
Record (Wurr et al. 1990)
Seed-tuber weight
planted (g)
Site Year 35 70 105 SE
DF = 6
Walkers (Notts) 1982
1983 8.5 7.9 6.9 0.89
1984 5.0 4.5 3.9 0.29
United Biscuits (Lines) 1982 3.0 3.2 0.23
1983 5.3 5.0 3.9 0.21
1984 4.3 4.5 3.6 0.49
Smiths (Suffolk) 1982 4.2 2.5 0.09
1983 5.0 3.6 4.2 0.27
1984
Golden Wonder (Cambs) 1982 1.9 1.5 1.18
1983 3.9 3.5 3.3 0.27
1984 5.6 5.0 4.8 0.66
(a) (b)
i
"C
Q)
';;'
....
r
a.> (c) (d)
.0
::::s
I-
72
Number of sets planted/ha (OOOs) ~
Figure 7.11 Optimum seed tuber popUlations from four experiments of the same
series.
324 Plant density
Without the necessary in~ormation on factors influencing number of
stems (and tubers per stem) effects of density have still to be considered
through either seed rate or number of seed tubers planted for use by
growers. Using either of these scales for establishing recommendations
also requires careful consideration to be given to the quality of the data, its
suitability for analysis leading to economic optima and the range of
circumstances in which the optima may be justified. There are few
examples of such comprehensive analyses being carried out anywhere and
generally individual experiments or a local series of experiments are
analysed with little regard to the principles of the analyses involved. The
large variation which can occur in number of stems per seed tuber and
number of tubers per stem has already been mentioned (Section 7.2.7) and
the latter variation is illustrated for a series of density experiments in Table
7.13. This variation is often associated with a large range in average effects
of increasing density in different experiments, as shown in Fig. 7.11, and
which comprises even greater variation in effects in individual replicates.
Consequently, in economic analysis, and especially for series of experi-
ments, simple averaging of responses is likely to lead to large inaccuracies
where the recommendations are used in specific circumstances. Wurr et
at. (1990, 1991) have presented methods of analysis which reject con-
siderable data as too inaccurate to use and which stabilize variation
through weighting procedures based on the reciprocal of the vari-
ance. These techniques should lead to more accurate and appropriate
recommendations.
Historically, economic analyses have simply been concerned with the
gross margin produced by densities, measured as seed rates, for saleable
yield above the minimum grade size. For the silt land area of East Anglia
the work of Jarvis and Shotton (1968, 1972), Jarvis (1971) and Jarvis and
Rogers-Lewis (1974) has resulted in recommendations of seed rates for
maximum gross margins for a number of varieties. These results are
illustrated in Table 7.14. It can be seen that the optimum seed rate is
dependent upon seed size, being lower for smaller seed. This would be
expected from the relationship between seed size and number of stems
discussed in Section 7.2.5 if there is one overall relationship between tuber
yield and stem density. It also applies to those varieties, Pentland Crown
and Record, in which there was evidence in these experiments of differen-
tial seed size effects. It was assumed that seed is purchased largely on a
weight basis, irrespective of tuber size, and other costs are little affected by
seed size, so it follows that the largest financial returns usually accrue from
the use of smaller seed, for the achievement of the optimum stem density
will produce similar yields. Table 7.15 illustrates this effect and shows
that in Pentland Crown and Record the advantage of small seed is not
apparent unless seed costs become very high in relation to ware prices.
With all varieties any increases in seed costs relative to ware prices
improve the relative profitability of small seed. There is very little seed of
Table 7.14 Seed rates for maximum gross margins for different seed sizes
Author Variety Seed size Optimum seed Gross margin
(g) rate (t ha· 1) £ ha- 1 at
optimum seed
rate
Seed at £30 t-1
Ware at £15 rl
Jarvis (1971) Pentland Dell 31.4 1.87 222
62.8 3.12 200
94.2 3.07 185
Pentland Crown 31.4 2.09 314
62.8 3.33 331
94.2 4.15 324
Seed at £25 t-1
Ware at £18 rl
Jarvis and King Edward 31.4 1.64 346
Shotton (1971) 62.8 2.55 311
94.2 3.05 277
Seed at £30 t- 1
Ware at £15 rl
Jarvis and Pentland Ivory 36.3 2.44 339
Rogers-Lewis 63.5 3.44 324
(1974) 101.5 4.23 285
Record 36.3 230 225
63.5 2.98 230
101.5 3.22 191
Table 7.15 Gross margins (£ ha- l ) at optimum seed rates with

different seed prices
Author Variety Seed Seed price (£ t- 1 )
size
(g) 20 30 40
Ware at £15 rl
Jarvis (1971) Pentland Crown 31.4 336 314 294
62.8 366 331 299
94.2 368 324 284
Pentland Dell 31.4 242 222 205
62.8 235 200 170
94.2 222 185 161
Ware at £18 t- I
Jarvis and King Edward 31.4 356 336 316
Shotton 62.8 321 294 269
(1971) 94.2 299 259 215
Ware at £15 rl
Jarvis and Record 36.3 255 255 201
Rogers-Lewis 63.5 262 230 198
(1975) 101.5 235 191 156
326 Plant density
Pentland Crown available as small as 31 g so that commercially there are
greater returns from the smallest readily available seed than from larger
seed.
These results are from sound well-executed experiments and have
formed the basis for national recommendations (MAFF, 1982, 1987)
although they were only from one quite distinct soil type. In the presenta-
tion of these national recommendations no indication is given as to the
justification for this extrapolation or for the methods of analyses (and
combination) used nor for the statements that small seed is inappropriate
for early harvests in some varieties. Variation in number of tubers from
similar stem densities is frequently associated with soil type (site) and
season (Table 7.13) so the general application of these average values must
involve serious inaccuracy. There must also be concern that an area such as
the silts uses distinct husbandry which may not be used elsewhere. The use
of 91 cm rows with consequent narrower within-row spacings than with
narrower rows will reduce number of tubers in most years. The wide rows
use substantial ridges to minimize greening and this probably contributes
to slower emergence of smaller seed sizes as all seed was planted at the
base of similar sized ridges. Small seed, therefore, had further to grow to
emerge than large seed and probably took longer. This probably explains
the effects of seed size found in Pentland Crown and Record and which
have not been found elsewhere (Wurr et al., 1990; Allen et al., 1991).
There is clearly a need for national recommendations to be re-thought so
that they are specific to the recognizable circumstances in which growers
operate. No single recommendation for a size and cost of seed is usable by
all growers.
Future recommendations must also accommodate the changes in grading
which are constantly occurring in market requirements and the increases in
yield which occur. Sales of tubers into markets with upper and lower riddle
limits are increasing and for many crops separation of the whole crop into
component grades, each with different values, is normal. Thus, the va,lue
of the crop is more difficult to establish and the effects of density are only
just beginning to be studied in this new context. As many of the desired
grades are new, existing experimental evidence based on measurement of
these grades is scant. Fortunately, the derivation of mode,ls characteriz-
ing the size distribution of tubers for diameter-limited grades (Travis,
1987) and weight-limited grades (Sands and Regel, 1983) through two
parameters, ~ the size with most yield and e a measure of the spread of
yield across size grades, are effective in allowing calculation of yields
of grades not measured from the yield of measured grades. Wurr et at.
(1990, 1991) have used this technique to study effects of density in the
processing variety Record and a range of ware varieties. In the latter a
wide range of seed costs and values of individual grades were used. As the
cost of seed increased the optimum planting density (as number of seed
tubers) was reduced especially where all ware potatoes had the same value
(Table 7.16). The effect was much larger in Pentland Squire than other
varieties such as Cara where the effect was quite small. As the value
of larger tubers increased the optimum seed density was little affected
in Pentland Squire and reduced in the other varieties. Where large-
sized tubers had increased value, the effect of increasing seed cost was
reduced and the effect on net returns at the optimum density was also small
(Table 7.17). Therefore, for the production ofbigh-value crops the cost of
seed is not very important and, for the future, increased returns to the seed
industry can be readily justified by ware producers aiming for the highest
quality.
Table 7.16 The optimum seed tuber planting density

(OOOs/ha) for different ratios of seed cost: small ware value:
large ware value. No optima were evident for seed tubers of
varieties Pentland Squire and Cara of 1430 count. (DF = 133,
residual from model fit) (Wurr et aI., 1991)
Monetary Pentland Cara Estima Maris King
ratio Squire Piper Edward
Tuber count (1430/50 kg)
1:1:1 63.3 75.2 44.3
2:1:1 51.9 61.0 37.6
3:1:1 39.9 46.2 30.1
1:1:2 51.3 57.3 31.7
2:1:2 43.8 48.7 28.1
3:1:2 35.4 39.0 23.7
1:1:3 45.3 49.3 25.9
2:1:3 39.7 43.0 23.5
3:1:3 33.0 35.7 20.5

1:1:1 57.4 46.9 38.7 44.0 32.1
2:1:1 43.0 36.5 30.6 34.5 26.0
3:1:1 29.9 26.3 22.6 25.1 19.7
1:1:2 58.5 42.7 33.2 36.9 26.2
2:1:2 46.8 35.9 28.4 31.3 22.9
3:1:2 35.2 28.3 23.0 25.1 18.9
1:1:3 60.6 41.0 31.0 34.1 23.8
2:1:3 50.3 35.8 27.5 30.1 21.5
3:1:3 39.5 29.7 23.3 25.4 18.6
In the future, the use of models characterizing tuber size distribution for
both diameter and weight-limited grades will be invaluable as optical
size grading and weight grading of tubers become more common. The
recommended optimum densities produced by Wurr et al. (1990, 1991) are,
328 Plant density
in many cases, quite different from those reported by MAFF and the data
also provide optima for other varieties, seed sizes and economic circum-
stances not previously available. The derivation of more accurate seed rate
recommendations for specific circumstances assumes that these rates will
be achieved in practice. The results of the surveys in Table 7.4 show
dramatically that growers frequently do not achieve the intended seed rate
and the most frequent error is to plant at wider spacing than intended. This
error is also associated with considerable irregularity in spacing and depth
of planting. The development of more efficient planters with greater
adjustment and automatic monitoring of spacing will ultimately offer
means of achieving the density intended. In the short term growers must
be more aware of the inaccuracies frequently created and their con-
sequences. Few growers check their spacings by uncovering a length of row
Table 7.17 Net return at the optimum seed tuber planting

density in small ware equivalents for different ratios of seed
cost: small ware value: large ware value. No optima were
evident for seed tubers of varieties Pentland Squire and Cara of
1430 count. (DF = 133, residual from model fit) (Wurr et aI.,
1991)
Monetary Pentland Cara Estima Maris King

ratio Squire Piper Edward
1:1:1 43.6 54.1 50.9
2:1:1 43.1 53.2 50.5
3:1:1 41.6 51.0 49.3
1:1:2 58 . 1 68.9 66.2
2:1:2 58.0 68.7 66.0
3:1:2 57.2 67.5 65.4
1:1 :3 73.7 85.8 83.3
2:1:3 73.9 85.9 83.3
3:1:3 73.5 85.4 82.9

1:1:1 53.2 61.8 47.0 57.6 56.2
2:1:1 51.2 59.9 45.7 55.7 55.1
3:1:1 46.5 55.4 42.2 51.5 52.3
1:1:2 77.3 88.2 63.0 73.0 72.7
2:1:2 77.6 87.2 62.3 72.3 72.2
3:1:2 74.5 84.1 60.2 69.8 70.5
1:1:3 100.7 115.1 79.8 90.0 90.5
2:1:3 102.8 114.6 79.6 89.7 90.2
3:1:3 101.7 112.4 78.1 88.2 89.1
and this simple observation would help many to avoid major inaccuracies
in their densities.
Since size distribution is determined by the number of tubers and the
yield for each crop once the number of tubers can be established by
sampling it is possible to predict changes in yield of individual grades as
overall yield increases. This may be helpful to large growers in planning
the harvesting, grading and marketing of their crops. However, until the
causes of variation in number of tubers at similar stem densities are
understood it is not possible to use prediction of yield of grades to choose
the most appropriate seed rates before planting.
ACKNOWLEDGEMENTS
The authors wish to thank all members of the potato group at Cambridge
University Farm and the office staff for their help in the preparation of this
chapter.
REFERENCES
Ahmed, Ch M.S. and Sagar, G.R. (1981). Volume increase of individual tubers of
potatoes grown under field conditions. Potato Res., 24, 279-88.
Allen, E.J. (1972). The effect of row width on the yield of three potato varieties. f.
Agric. Sci., Camb., 79, 315-21.
Allen, E.J. (1977). Effects of date of planting on growth and yield of contrasting
potato varieties in Pembrokeshire. 1. Agric. Sci., Camb., 89, 711-35.
Allen, E.J. (1979). Effects of cutting seed tubers on number of stems and tubers
and tuber yields on several potato varieties. f. Agric. Sci., Camb., 93, 121-8.
Allen, E.J. and Wurr, D.C.E. (1973). A comparison of two methods of recording
stem densities in the potato crop. Potato Res., 16, 10-20.
Allen, E.J. and O'Brien, S.A. (1987). An analysis of the effects of seed weight,
seed rate and date of harvest on the yield and economic value of seed-potato
crops. f. Agric. Sci., Camb., 108, 165-82.
Allen, E.J., O'Brien, P.J. and Firman, D. (1991). An evaluation of small seed for
ware-potato production. f. Agric. Sci., Camb. (in press)
Allen, E.J., Bean, J.N., Griffith, R.L. and O'Brien, P.J. (1979). Effects of length
of sprouting period on growth and yield of contrasting early potato varieties.
f. Agric. Sci., Camb., 92, 151-63.
Arthur, J.e. (1892). The relation of number of eyes on the seed tubers of potatoes
to the product. Indiana Sta. Bulletin., 42, 105-18.
Bates, G.H. (1935). A study of the factors influencing size of potato tubers. f.
Agric. Sci., Camb., 297-313.
Bean, J.N. (1981). Density studies with early potato cuItivars. Ph.D. Thesis.
University College of Wales, Aberystwyth.
330 Plant density
Birecki, M. and Roztropowicz, S.T. (1963). Studies of seed potato size and
productivity. Eur. Potato J., 6, 1-13.
Bleasdale, J.K.A. (1965). Relationships between set characters and yield in
maincrop potatoes. J. Agric. Sci., Camb., 64,361--6.
Bleasdale, J.K.A. and Nelder, J. (1960). Plant population and crop yield. Nature,
Lond., 188,342.
Bleasdale, J.K.A. and Thompson, R. (1965). Rep. Natn. Veg. Res. Station for
1964,37-8.
Bleasdale, J.K.A. and Thompson, R. (1966). Rep. Natn. Veg. Res. Station for
1965, 39-40.
Bodlaender, K.B.A. and Reestman, A.J. (1968). The interaction of nitrogen
supply and plant density in potatoes. Neth. J. Agric. Sci., 16, 165-76.
Boyd, D.A. and Lessells, W.J. (1954). The effect of seed rate on the yield of
potatoes. J. Agric. Sci., Camb., 44, 465-76.
Brandreth, B. and Bryan, H. (1937). 1. The effect of size of 'seed' on the yield of
the larger grades of ware potatoes. J. Natn. Inst. Agric. Bot. for 1937. IV, (2),
183-92. Potato Trials 1936.
Bremner, P.M. and Taha, M.A. (1966). Studies in potato agronomy. 1. The effects
of variety, seed size and spacing on growth, development and yield. J. Agric.
Sci., Camb., 66, 241-52.
van Burg, P.F.J. (1967) Relation of rate of nitrogen fertilizer, seed spacing and
seed size to yield of potatoes. Neth. Natn. Tech. Bulletin 4 Oct. 1967.
de Carvalho, T. (1975). Potato growing. Tech. Bull., 11, (1) 1-19. Publ. Rhod.
Ministry Agric.
Chase, RW., Silva, G.H. and Kitchen, R.B. (1989). Pre-cutting of seed potatoes.
Am. Potato J., 66, 723-30.
Cole, H. (1973). Early potatoes in West Cornwall 1962-1972. An economic survey,
Report 19. Agric. Economics Unit, University of Exeter.
Driver, C.M. and Hawkes, J.G. (1943). Photoperiodism im the potato. Bulletin of
the Imperial Bureau of Plant Breeding and Genetics, Cambridge.
Dyson, P.W. and Watson, D.J. (1971). An analysis of the effects of nutrient supply
on the growth of potato crops. Ann. Appl. Bioi., 69, 47--63.
Eckersall, R.N. (1965). Implications of herbicide use on plant population, plant
arrangement and cultivations in the potato crop. Ph.D. Thesis. University of
Nottingham.
EI Saeed, A.K. (1963). Aspects of plant population in the potato crop. Ph.D.
Thesis, University of Nottingham.
Fowler, J. (1988) Effect of plant arrangement and density on growth develop-
ment and yield of two potato varieties. Ph.D. Thesis. University of
Cambridge.
Goodwin,P.B., Brown, A., Lennard, J.H. and Milthorpe, F.L. (1969). Effect
of centre of production, maturity and storage treatment of seed tubers on
the growth of early potatoes. II. Field growth. J. Agric. Sci. Camb., 73,
167-76.
Gray, D. (1972). Spacing and harvest date experiments with Maris Peer potatoes.
J. Agric. Sci., Camb., 79, 281-90.
Gray, D. (1973). The growth of individual tubers. Potato Res., 16, 80-4.
Gray, D. and Smith, D.J. (1973). The pattern of assimilate movement in potato
plants. Potato Res., 16, 293-5.
References 331
Griffith, R.L., Hide, G.A., Hirst, J.M. and Stedman, O.J. (1974). Effects of
gangrene (Phoma exigua) on potatoes. Ann. Appl. BioI., 77,237-50.
Hide, G.A., Hirst, J.M. and Mundy, E.J. (1969). The phenology of skin spot
(Oospora pustulans, Owen and Waket) and other fungal diseases of potato
tubers. Ann. Appl. Biol., 64,265-79.
Holliday, R. (1960). Plant population and crop yield: Part II. Yield and plant
population in British crops. Fld. Crop Abstr., 13, 247-54.
Holmes, J.c. (1966). Seed size and spacing for seed and ware potato crops. Scottish
Agric., 45, 176-80.
Holmes, J.C. (1973) Relationship between stem number and tuber number in the
potato crop. Proc. 5th Trien. Cont Eur. Ass. Potato Res. Norwich.
Ifenkwe, D.P. (1975). Effects of row width and plant density on growth and
development of two maincrop potato varieties. Ph.D. Thesis. University
College of Wales, Aberystwyth.
lng, E.G. (1966). Cutting of Arran Pilot seed potato and size of seed. Expl. Hort.,
14,39-42.
Jarvis, R.H. (1971). Population studies with Pentland Crown and Pentland Den
potatoes. Expl. Husb., 20,84-92.
Jarvis, RH. (1972). Comparison of 30 in. and 36 in. rows for maincrop potatoes.
Part II. Soil factors, workrates and mechanical damage. Expl. Husb., 21,
85-92.
Jarvis, R.H. and Rogers-Lewis, D.S. (1974). Population studies with Pentland
Ivory and Record potatoes. Expl. Husb., 27,23-30.
Jarvis, RH. and Shotton, F.G. (1968). Population studies with Majestic potatoes
in rows and in beds. Expl. Husb., 16, 73-92.
Jarvis, RH. and Shotton, F.G. (1971). Population studies with King Edward
potatoes. Expl. Husb., 20, 12-29.
Jarvis, R.H. and Shotton, F.G. (1972) Comparison of 30 in. and 36 in.
fOWS for maincrop potatoes. Part I. Effects on yield. Expl. Husb., 21,
78-84.
Krijthe, N. (1955). Observations on the formation and growth of tubers on the
potato plant. Neth. J. Agric. Sci., 3, 291-304.
Krug, H. (1960). The photoperiodic behaviour of a number of potato varieties.
Eur. Pot. J., 3,47-79 and 107-36.
McKerron, D.KL. and Jefferies, R.A. (1986). The in,fluence of early soil moisture
stress on tuber numbers in potato. Potato Res.. , 29,299-312.
Ministry of Agriculture, Fisheries and Food (1982). Seed rate for potatoes grown as
maincrop. Leaflet 653 19 pp.
Ministry of Agriculture, Fisheries and Food (1987). Seed rate for potatoes grown as
maincrop. Leaflet P653.
Nelson, D.C. (1967). Effects of row spacing and plant populations on yields and
tuber-size of potatoes. Am. Potato J., 44, 17-21.
North, J.J. and Proctor, J.M. (1971). Row widths for King Edward potatoes. Expl.
Husb., 22, 99-103.
Oparka, KJ. (1987). Influence of selective stolon removal and partial stolon
excision on yield and tuber size distribution in field-grown potato cv. Record.
Potato Res., 30,477-83.
Piitzold, D. (1964). Studies of the effect of row width in the potato crop. II. Plant
development. Eur. Potato J., 7, 1-12.
332 Plant density
Rappaport, L. and Sachs, M. (1967). Wound-induced gibberellins. Nature, Lond.,
214, 1149-50.
Reestman, A.J. (1970). Importance of the degree of virus infection for the
production of ware potatoes. Potato Res., 13, 248--68.
Reestman, A.J. and de Wit, C.T. (1959). Yield and size distribution of potatoes as
influenced by seed rate. Neth. J. Agric. Sci., 7,257--68.
Sands, P.J. and Regel, P.A. (1983). A model of the development and bulking of
potatoes (Solanum tuberosum L.) V. A simple model for predicting graded
yields. Fld. Crops. Res., 6, 25-40.
Saunt, J.E. (1960). Plant population studies with the potato crop, M.Sc. Thesis.
University of Leeds.
Scott, R.K. and Younger, A. (1972). Potato agronomy in a changing industry.
Outl. Agric., 7,3--9.
Sharpe, P.R. and Dent, J.B. (1968). The determination and economic analysis of
relationships between plant population and yield of maincrop potatoes. J.
Agric. Sci., Camb., 70, 123--9.
Stalham, M.A. (1990). Growth and water use of the potato variety Record on
contrasting sites. Ph.D. Thesis. University of Cambridge.
Stuart, W., Lombard, P.M., Vosbury, M.C., Corder, G., Edmundsen, W.C.,
Clark, C.F. and Dewey, G.W. (1924). Size of potato sets: comparison of whole
and cut seed. Bull. 1248 U.S. Dep. Agric.
Svensson, B. (1966). Seed tuber - stand - yield. Viixtodling, 21, 1-86.
Svensson, B. (1972). Influence of the place of a stem in the hill on the weight and
dry matter content of its tubers.. Potato Res., 15,346--53.
Taylor, C.E. (1953). The vegetative development of the potato plaQt. Ann. Appl.
BioI., 40, 778-88.
Thomas, M.N. (1988). The control of tuber size in maincrop potatoes. Ph.D.
Thesis. University of Cambridge.
Thompson, R. and Taylor, H. (1974). Stem density and maturation studies wi,th
the potato cultivars Maris Peer and Pentland Marble. Potato Res., 17,
51--63.
Travis, K.Z. (1987). Use of a simple model to study factors affecting the
size distribution of tubers in potato crops. J. Agric. Sci., Camb., 109,
563--71.
WUIT, D.C.E. (1971). Some effects of seed size and spacing on the yield and grading
of two maincrop potato varieties. Ph.D. Thesis. University of Cambridge.
Wurr, D.C.E. (1973). Individual tuber growth .. Rep. Natn. Veg. Res. Stn. (1972)
65.
Wurr, D.C.E. (1974). Some effects of seed size and spaciftg on the yield and
grading of two maincrop potato varieties. I. Final yield and its relationship to
plant population. J. Agric. Sci., Camb., 82, 37-45.
Wurr, D.C.E. and Allen, E.J. (1974). Some effects of planting density and variety
on the relationship between tuber size and tuber dry-matter percentage in
potatoes. J. Agric. Sci., Camb., 82, 277-82.
Wurr, D.C.E., Fellows, Jane R., Sutherland, R.A. and Allen, E.J. (1990).
Determination of optimum tuber planting density for production of tubers in
processing ware grades in the potato variety Record. J. Agric. Sci., Camb., 114,
11-18.
Wurr, D.C.E., Fellows, J.R. and Allen, E.J. (1991). Determination of optimum
References 333
tuber planting density in the potato varieties Pentland Squire, Cara, Estima,
Maris Piper and King Edward. J. Agric. Sci., Camb. (in press)
Zeigler, G. (1968). Influence of changed row spacing on potato yields. Albrecht-
Thaer-Archiv., 12, 155-72.
CHAPTER 8
Breeding new varieties

P.D.S. Caligari
8.1 INTRODUCTION
8.1.1 General considerations

The methods employed and the approaches taken in breeding any plant
species are largely dictated by the natural reproductive mechanisms that
the species possesses and how these are exploited under domestication. In
this way plant breeding methods. are usually categorized into distinct
groups, the most obvious distinction being made between those capable of
self-pollination (thus being readily inbred and allowing the derivation
of genetically homozygous, true-breeding lines) and those which for a
number of reasons have to be cross-pollinated to give s.eed (thus they are
usually marketed as out-bred populations or as hybrids in order to retain
genetical heterozygosity). This distinction is, however, often more a
theoretical convenience than a practical reality since other factors, usually
biological, agronomic or economic, influence the methods used. Another
plant breeding and selection system commonly identified is where the plant
is readily reproduced by asexual means. In such cases clonal reproduction
can be used as a method to exploit the species in terms of production.
Again, however, the possibilities within this simple heading are many.
For example, species such as grasses are capable of being multiplied
extensively by asexual means but are sown by gardeners and farmers alike
as seed. Their potential for asxual reproduction is often exploited,
however, in the breeding programme itself. The potato is, of course, one of
the most familiar examples of a crop species that is not only capable of
asexual reproduction but has largely been exploited in horticulture and
agriculture via its potential for clonal propagation.
8.1.2 Potato breeding in general

The production of potatoes is in the process of being re-examined,
particularly in terms of the potential to exploit sexual rather than the more
Published in 1992 by Chapman & Hall, London. ISBN 0412296403
Introduction 335
traditional asexual propagation (Ross, 1986). Whichever method is used
there are, as with all biological systems, benefits and drawbacks to be
considered. The advantages of particular features are present to be taken
advantage of, if possible, while the minimization of the drawbacks provides
the challenges that need to be faced. The obvious advantage of clonal
reproduction is that any genotype, irrespective of its genetical com-
position, can be multiplied and perpetuated. By this means any plant, even
if only initially present as a single individual, if perceived as desirable by
the breeder, can be multiplied to levels which can satisfy the commercial
market. The major drawback to this mode of reproduction is the ease with
which diseases, particularly virus diseases, can become established in a
stock. Once plants are infected, the virus is readily transmitted to the
daughter tubers and the disease is maintained and multiplied along with
the potato stock. Such diseases are not usually transmitted to the true,
botanic seeds produced by an infected plant, although there are excep-
tions. In potato an exception is provided by virus-like organisms, such as
the spindle tuber viroid (Ross, 1986), which seem to be readily transmitted
via true, botanic seed. Thus generally sexual reproduction when used as a
method to multiply stocks has great advantages from the point of view of
readily maintaining health status. The complicating factor for exploiting
this, however, is producing parental genotypes which give progenies with
uniform phenotypic expression, a point which will be returned to later.
8.1.3 History of potato breeding

We have no real idea when potatoes were first the subject of conscious
selection by man. Hawkes (this volume, Chapter 1), however, cites
evidence for the production of seeds and the selection for desirable
characters at the start of the seventeenth century. It would be naive to
suppose that considerable selection had not been practised in earlier times.
Indeed, although the science underlying plant breeding has only recently
(twentieth century) been acquired, much of the selection required to
produce most of the present day crop plants was carried out during the
early stages of the potatoes' domestication. As soon as man started to
practise a settled form of agriculture it is probable that growers started to
exercise choice among the different 'types' available to them. If a particular
plant grew well, or seed collected from a particular area had desirable
characteristics, it is unlikely these would not be used preferentially in the
following years. Thus. by the time that we have written records, a large and
very effective part of the domestication and subsequent selection of
potatoes had taken place in South America.
In Europe the introduction of the potato was much later (sixteenth
century) and hence more details are available about the spread of potatoes
for domestic consumption and the problems encountered. However, the
exact details of the introduction into Europe are still unknown and mostly
based on circumstantial information. It seems fairly certain that the first
336 Breeding new varieties
introductions were in Spain, in the second half of the sixteenth century.
The origin of these potatoes is still uncertain (Hawkes, this volume,
Chapter 1; Glendinning, 1983). What is clear is that they were tetraploids
from the species Solanum tuberosum. The main controversy is from which
country they originated and hence from which subspecies of Solanum
tuberosum they were derived. Much of the current evidence points to their
Columbian (subspecies andigena) origin and hence not from the Chilean
subspecies tuberosum. If this is the case then the potato breeders in Europe
would have had to practise fairly stringent selection for adaptation to the
longer daylight conditions of the growing season here, and hence mimicked
the derivation of tuberosum in Chile. The selection for such adaptation
seems to have been successful relatively quickly since potatoes were
reported as being grown commercially in Ireland in the seventeenth
century (Salaman, 1949) and were a major food source in England by the
middle of the eighteenth century (Davidson, 1934; Glendinning, 1983). It
also seems to be the case that potatoes spread to much of the rest of the
world from this introduction into Europe.
As was pointed out by Glendinning (1983) the propagation of potatoes
in the eighteenth century was by a mixture of methods. New varieties were
produced, often in abundance, from true botanic seed particularly as a
result of field-set berries. This was popular in many regions in order to
overcome the effects of the gradual accumulation of various diseases. The
sale of seed tubers was also practised and allowed the dissemination of
particularly desirable agronomic genotypes (which we would now term
cultivars or varieties). At this early stage there was already recognition of
geographical areas that were more appropriate for seed tuber production
than others (Glendinning, 1983). By this mixture of rapid variety pro-
duction alongside seed tuber distribution there appeared to be consider-
able turn-over in the varieties that were grown and since large numbers of
seedlings were being raised, considerable scope for artificial selection to be
practised. What had happened prior to this is uncertain; the earlier
reference (Hawkes, this volume, Chapter 1) that we noted only takes us
back to the sixteenth or seventeenth century. It is interesting that it has
proved possible to demonstrate the potential to recapitulate the supposed
selection of subspecies andigena not only simply to tuberize relatively
early, under conditions of long days, but to exhibit many useful and
commercially acceptable characters. This is remarkable especially when
the time scale taken, only some 20 years, is considered (Simmonds, 1966;
Glendinning, 1975a, b). Indeed, a similar picture also emerged when
primitive cultivated diploid potatoes from South America were selected
under modern European conditions (Carroll, 1982).
At present, and in the recent past, most commercial potato breeding
programmes have concentrated on the production of potato varieties that
will be multiplied asexually, i.e. as clones, and distributed in the form of
seed tubers. Since such activities have generally centred on countries which
Breeding methods 337
introduced the potato from Europe these have mostly used the tetraploid
genotypes of S. tuberosum. Such programmes therefore provide an
obvious starting point in terms of describing the breeding of potatoes,
although other possibilities and developments will be considered.
8.2 BREEDING METHODS
Having set out the case for describing the general features of potato
breeding programmes aimed at the clonal propagation of varieties the
actual starting point for any breeding programme is the generation or
release of genetic variation. There exists the potential to exploit mutations
as a source of variation and these can occur naturally or their rate of
appearance enhanced artificially. Although such methods have been used
in potatoes, for instance in obtaining different tuber skin colours, they
have not been used extensively. The major source of variation, upon which
selection is to be practised, is generated by the segregation exposed by
sexual reproduction. The crossing of different genotypes and the selfing of
heterozygotes have been carried out with some confidence since the
rediscovery of Mendel's work at the turn of the twentieth century. With
crops which are destined to be reproduced clonally, as is the present case
with potatoes, once this variation is released there is no problem with
stabilizing ('fixing') any desirable combination that arises, as any clone can
be multiplied unchanged by asexual reproduction..
8.2.1 Advantages and disadvantages of clonal reproduction

Asexual reproduction, in the form of tubers, or more recently by means of
meristem culture and micropropagation, allows the multiplication of any
genotype without affecting its genetic constitution (apart from the effect of
rare mutations), as has already been discussed. The advantages of being
able clonally to multiply any genotype extend beyond the simply com-
mercial exploitation. It enables individuals with low fertility, such as often
arise when wide or interspecific crossing is carried out, to be maintained,
tested under varying conditions, or replicated for experimental investiga-
tions. It also means that selection in the progenies does not suffer the
complications that can arise, particularly in inbreeding crops where selec-
tion in early generations can be distorted or biased by the effects of
dominance which will not be present in the final inbred lines that are being
produced.
The disadvantages of exploiting clonal reproduction are also evident.
The first and major one has already been noted and relates to the difficulty
of keeping stocks free of diseases, particularly viruses, during multi-
plication. This is now partially capable of being overcome by the use of in
vitro propagation (including micropropagation and micro tuber production)
and minituber production (Ross, 1986; Chandra et al., 1988). Neverthe-
less, unless such propagules can be planted directly for commercial ware
production some risk of accumulating disease is still present during field
multiplication. Another feature which has been a disadvantage in the past,
but which again has been somewhat lessened by the techniques noted
above, is the relatively low multiplication rate via tubers. This has
generally not affected breeding programmes per se so directly as the
subsequent multiplication of breeders' clones for commercialization,
where it has been claimed to limit the spread of new varieties. A further
disadvantage is that vegetative propagules, such as tubers, are generally
not readily stored over extended periods of time. This has traditionally
meant that breeders' lines, useful germplasm, etc. have had to be grown
each year in order to maintain them with, of course, an ever present risk of
disease infection or loss due to handling errors. This has entailed large
amounts of time and effort being expended and, at the same time, limited
the amount of material that has been preserved. Various methods of
preserving potato germplasm have been investigated in order to circum-
vent this problem, including cryopreservation (Withers, 1986) and slow-
growth conditions in tissue culture (Powell and Caligari, 1989).
There are also factors that arise indirectly from the fact that clonal
reproduction has been used primarily in potato production. The first of
these is that because any genotype can be successfully multiplied, there has
been no advantage or pressure to produce inbred, homozygous material.
This has meant that the effects of many of the alleles at any locus are
hidden by the dominance effects of others. It has also led to the inefficient
use in breeding schemes of major genes, such as those for disease and pest
resistance. In most varieties and clones used as parents such genes are
present as only one allele (one 'dose', called simplex); by selfing or
controlled crossing it is possible to produce and identify resulting in-
dividuals with two (duplex), three (triplex) or four (the homozygous,
quadruplex) copies of the dominant alleles present. An example is shown
in Table 8.1, giving the case of a dominant advantageous allele. The table
shows the outcome of using parents with different numbers of copies of the
allele, when used in crosses with an hypothetical other parent with no
copies of the advantageous allele (nulliplex). This is, of course, not the
only possibility but the one most commonly of concern. As can be seen the
case normally used by breeders, simplex crossed nulliplex, means 50% of
the progeny will be discarded as susceptible before selection for any other
character is considered, while with a triplex or quadruplex parent all
progeny are resistant and therefore acceptable! This lack of attention in the
past to parental clone production is puzzling; the basic philosophy that has
been adopted is that a good variety or clone (i.e. good phenotypic
expression) is somehow equivalent to good genetic composition (Le.
good genotype) while evidence, including empirical results, shows this is
commonly an unrealistic assumption. When it is considered that the
Table 8.1 The expected ratios in the progenies of crosses
between genotypes (Parent 1) carrying varying numbers of
copies of a dominant advantageous allele (A) with one
(Parent 2) containing only recessive disadvantageous alleles
(aaaa) at a single locus (the possible effects of double reduction
are ignored)
Cross ResistaRt % resistant
:susceptible in progeny
Simplex (Aaaa x aaaa) 1:1 50
Duplex (AAaa x aaaa) 5:1 83
Triplex (AAAa x aaaa) 1:0 100
Quadruplex (AAAA x aaaa) 1:0 100
example only deals with one character controlled by one gene, when there
are many characters to be considered and most are controlled by many
genes, it seems irrational not to try to handle the few characters that are
controlled simply in an efficient manner.
The second indirect effect that reliance on clonal reproduction has had,
combined with a situation where the products of sexual reproduction are
not of commercial value, is that it is common for infertility to be a feature
of varieties or clones. Not only can such sterility be tolerated in a crop such
as potatoes but it was actually a breeding objective in many programmes.
This followed from the belief that the formation of berries and yield of
tubers were negatively correlated (Bartholdi, 1942). This has meant that
there is considerable infertility present in much of the germplasm used by
breeders and has restricted the crosses that can be carried out. The use of
protoplast fusion in the future (see p. 363) may provide a potential solution
to this restriction but the methodology is far from routine at present and
has other potentially complicating factors such as the increased ploidy of
the resultant fusion products.
8.2.2 Controlled pollinations

The carrying out of controlled pollinations in potatoes is essentially a
simple operation, in that:
1. the anthers are large and therefore easy to remove;

2. the anthers do not dehisce before or soon after flower opening and
hence give easy access and timing for removal;
3. the pollen is not easily distributed by wind and thus if plants are raised
in insect-proof houses, flowers need not be bagged to prevent illegitimate
pollinations;
4. the ovary starts to swell quite quickly after pollination, while failure
results in flowers dropping off, thus the initial success or failure can be
judged relatively quickly and repeat crosses made;
5. each flower stem has a number of flowers and thus gives the potential
for a number of pollinations;
6. each pollination, if fully successful, results in a large number of seeds
(usually more than 50);
7. developing berries can be bagged readily and allowed to ripen on the
plant;
8. seed is sometimes difficult to extract but the use of a domestic blender
(if used cautiously) allows the seed to be fairly readily separated, dried
and packeted;
9. seed can be stored over a number of years or used fairly quickly (there
are some suggestions of innate seed dormancy with freshly harvested
material (Simmonds, 1963) but this is also affected by the length of
ripening time of the berries and their subsequent storage before seed
extraction).
8.2.3 Problems in producing seed from crossing

There are, however, other complications which disturb this picture of an
easy species to cross pollinate. In the first place it has already been noted
that many varieties and clones do not set selfed seed. This failure can be
attributed to a number of causes but can be traced not just to the tolerance
of a lack of sexual fertility in clonally reproduced crops but, as we noted, to
the deliberate action of breeders to select for such infertility. The first, and
obvious feature which immediately strikes anyone trying to cross potatoes
is that many clones and varieties do not normally flower at all. It is often
the case that inflorescences are formed but they cease growing after a short
period and soon fall off at the abscission layer. Even when flowers are
formed and pollinated with known viable pollen there are frequent cases of
flower drop and no berry formation.
The above features have led most breeders to adopt various techniques
to try to reduce these effects as much as possible. The first important
point is that temperatures above about 22°C lead to lack of seed set;
therefore temperature control via shading in glasshouses is often impor-
tant. Secondly, it has been found that if the daughter tubers are removed as
they form, the plants will tend to flower more profusely and retain flowers
more readily. This has led to the technique of removing developing
daughter tubers (Knight, 1807; Thijn, 1954). The technique commonly
employed is one in which the mother tuber is placed on a house-brick and
covered with soil; as the plant develops the soil is washed away to expose
the roots, which are trained into the lower soil level. As stolons develop and
tubers form these are readily removed without further disturbance to the
plant. Another technique often employed with varieties that do produce
flowers is to decapitate them and place the stems, with attached flowers, in
jars of water (Peloquin and Hougas, 1959) often with an anti-bacterial
agent in solution to reduce contamination. It also provides a particularly
useful way of exploiting flowers produced in field grown material. Another
popular method of affecting flowering is to graft potato stems onto tomato
stocks (Ross, 1986). Other factors affecting successful seed production are
planting under conditions of long-days (Patterson, 1953; Steineck, 1957),
late planting (Rothacker, 1957) and spraying with auxin analogues
(Wiersema, 1950).
A further complication in potatoes is the occurrence of both pollen and
ovule sterility. Several early studies on these characters suggested various
underlying determinants of this sterility with major genes, gene-cytoplasm
interactions and polygenes all being implicated (for some reviews see
Krantz, 1946; Swaminathan and Howard, 1953; Ross, 1986). The inter-
pretation of the position in most cases is, however, complicated. The use
by many breeders of various wild species introduces a number of factors
which are likely to affect either or both pollen and ovule fertility.
8.2.4 Choice of parents

Clearly the choice of parents used will be restricted by effects such as those
noted above but, leaving this aside, it will depend largely on the aims and
objectives of the breeder. If the breeder is primarily concerned with the
production of varieties with commercial attributes then slhe will try to
choose pairs of varieties which have complementary sets of characters with
desirable levels of expression. For instance, slhe may wish to try to
combine the yield potential of one with the tuber shape of another. In
general, of course, there are a large number of characters to be considered,
usually accepted at around 50, but by using varieties as parents many of
these are taken to be of acceptable standard. In other cases the breeder
may be specifically interested in improving pest or disease resistance and
this will then dominate the choice of parent. Mostly, whatever the breeding
objectives, the choice of parent is based on the phenotypic expression of
the characters it displays and, as noted, little effort has been expended in
producing parental clones specifically. Breeders, however, do use the
empirical evidence of previous crossing programmes and identify clones or
varieties which appeared to give a higher than expected acceptability of
clones among their progenies and thus practice some selection based on
this empirical, genotypic information. The major exception to this type of
approach is that taken by workers at the International Potato Center (CIP),
Peru who progeny test their potential parent lines for use in breeding
programmes in a range of countries (Mendoza, 1987). Recently Brown and
Caligari (1989) examined experimentally several methods of making predic-
tions of the 'parental value' of clones and concluded that this could readily be
achieved. They proposed that the methods developed for cross-prediction
provided a suitable practical method for use in potato breeding programmes.
8.2.5 Traditional variety breeding programmes
The exact details of the procedures and practices of traditional breeding
programmes differ, as might be expected, but in basic outline they share
much in common. The scheme used at the Scottish Crop Research Institute
(SCRI) will serve to demonstrate the basic outline. The scheme used in
1982 is outlined in Table 8.2 (Mackay et a{, 1986; Mackay, 1987) as far as
the assessment and selection for agronomic characters is concerned. The
size of the initial population of seedlings is, at 115 000, fairly typical of
large breeding programmes and it can be seen that this is rapidly reduced
over the first three generations such that by the start of the fourth year only
1000 clones remain (less than 1%). Clearly the early clonal generations,
and how they are handled and selected, are crucial to the success of the
final outcome.
Some examples of the numbers of seedlings raised initially each year
and the subsequent reductions in these numbers (given as cumulative
percentage reduction) are shown in Table 8.3. Probably the most striking
feature of this table is the last column of figures. These represent the
Table 8.2 A summary of the breeding scheme at the Scottish Crop Research
Institute in 1982 (adapted from Mackay, 1982, 1987)
Year No. of clones Sites Selection for agronomic characters
1 115000 Glasshouse Visual selection on tubers from individual
plants in pots
2 40000 Seed site Visual selection on tubers from individual

plants
3 4000 Seed site Visual selection on tubers from a

three-plant plot
4 No selection at seed site but yield trials at

1 ~~ ) Seed site ware site with single 12-plant plots.
5 500 Ware site Selection for yield, quality, storage
characteristics, etc
6 200
7 60 Seed site Selection much as for years 3, 4, 5 but with

Ware site larger plots and based on wider geographic
8 10 UK trials spread of sites
Overseas
9 5
10 5 National List Trials

11 5
12 ? Named varieties for commercialization
proportion of the initial population of seedlings which have been discarded
by the end of the second clonal generation. What the figures show is that by
the third year of all breeding programme a minimum of 96%, and generally
99% of the sample of genotypes has been eliminated. Clearly this phase of
the breeding programme is critical since most of the selection will be
achieved during this period. It is, however, one which is still subject to
discussion and controversy.
Table 8.3 The numbers of seedlings grown to initiate a number of traditional potato
breeding programmes and the percent of this initial number that have been discarded
by the end of each growing season
Number of Seedling First clonal Second clonal
Country Reference* Year seedlings generation generation generation
Canada 1 1984 40000 0 85 96
England 2 1978 25000 68 96 99
Germany 3 1984 140000 39 95 99
Ireland 4 1982 100000 64 98 99
N. Ireland 5 1985 30000 85
Scotland 6 1982 115000 65 96 99
* 1, Tai and Young (1984); 2, Howard (1978); 3, Ross (1986; from Fitschen (1984»; 4, Kehoe
(1982); 5, Lee (1985); 6, Mackay (1987).
8.2.6 Selection in the seedling and first two clonal generations of traditional
breeding programmes
(a) Production and selection of seedlings

The current procedure for raising potato seedlings is to grow them in
aphid-proof houses or glasshouses. Seeds are sown in pans and the
seedlings transplanted to individual lO-cm pots when sufficiently large to
be handled. These pots are generally watered from beneath to avoid
problems of disease associated with moist foliage and kept in as disease-
free conditions as possible. They are grown to maturity, the water supply
discontinued and the foliage removed. When the compost has dried out the
tubers are harvested from each pot individually. It should be remembered
that as the parents are themselves heterozygous these seedlings will show
the effects of genetic segregation and hence each one will be genetically
unique.The question that arises at this stage is should selection be practised
on these seedlings? If it can then the amount of material that has to be
handled in the next clonal generation is dramatically reduced and much
time and effort saved. However, potatoes grown from seed, especially in
small pots in the glasshouse, would hardly seem likely to be representative
of later field performance when grown from tubers.
The question of selection at the seedling stage is still controversial but
most breeders recognize the inefficiency of selecting on the basis of single
plants at this stage. Having pointed this out, however, the figures given for
many breeding programmes (Table 8.3) still show a reduction in numbers
between seedlings raised and plants in the next year. It can be seen that the
reduction in numbers varies between programmes from 0% to 60%.
Various suggestions have been made as to what characters might respond
to selection at this stage, namely irregular tuber shape, deep eyes and
excessive numbers (Stuart, 1937; Howard, 1963), maturity (Krantz, 1938;
Maris, 1964; Tellheim, 1975), low yields (Howard, 1963; TeUheim, 1975),
growth cracking (Kichefski et al., 1976). Many of the experiments involved
comparisons between the seedling year and the subsequent clonal field
year (Pfeffer, 1963; Niopek, 1967; Tellheim, 1975; Swiezynski, 1978, 1984;
Anderson and Howard, 1981). Clearly performance in the next clonal
generation is a partial criterion but it is, by definition, likely to be affected
by the health, size, condition, etc. of the planted tuber (Brown et al., 1984;
Maris, 1986). It therefore follows that any association observed between
seedling and the directly subsequent first clonal generation is likely to be
biased by mother tuber effects (Blomquist and Lauer, 1962). It is not
surprising therefore that the answers obtained appeared equivocal as to the
desirability of carrying out such selection (Maris, 1964; Howard, 1978). In
these cases it was suggested that the correlations observed did not warrant
positive selection being practised but they did recommend that the very
poorest phenotypes should be discarded (i.e. negative selection practised). A
few studies have extended over further years (e.g. Maris, 1969) but more
recently a series of experiments have been published covering an extensive
investigation of selection in the seedling and subsequent generations (Brown
et al., 1984, 1987a,b; Brown and Caligari 1986a; Caligari et al., 1986).
The general conclusion from the work at SCRI was that although there
were some indications of slight improvements in the mean of the popula-
tions after such selection, either using visual appraisal or measured
characters, there were large penalties to be paid in terms of discarding the
most commercially desirable individual clones. Indeed, it was shown
(Brown and Caligari, 1986b) that when comparisons were made between
selected and unselected groups at a later stage the unselected group
produced the higher proportion of commercially acceptable clones!
Selection for characters that are controlled by major genes can, of course,
be successfully carried out at this stage (Plaisted et al., 1984; Swiezynski,
1984). What was also shown by the experiments noted above was that
although individual clones could not be selected data collected on such
clones could be used as the basis for uni- and multi-variate cross predic-
tions (Caligari and Brown, 1986; Brown and Caligari, 1987; Brown et aI.,
1988). It was shown that this methodology (pioneered by Jinks and his
colleagues e.g. Jinks and Pooni, 1976; Pooni and Jinks, 1978) provided an
efficient approach to identifying the crosses which had the most potential
for producing what the breeder was aiming for.
Another question that has been the subject of debate in the literature
has been concerned with the use of visual appraisal as opposed to objective
measurement of characters in breeding programmes (Tai, 1975). Most
breeders use visual appraisal (breeder's preference) in this stage of the
breeding programme since it is the only method by which they can take
account of the numerous characters such as tuber shape, tuber number,
eye depth, stolon persistence, skin appearance, etc. while handling the
large numbers they are growing. In other words by either recording or
merely using a select/reject system for clones they can very quickly and
easily handle the large number of seedlings grown. Some of the debate is
noted by Tai and Young (1984) who make clear that much of the earlier
concern about its efficacy is not a reflection of the method of assessment
but of the low heritabilities of the underlying characters, a result also
supported by Caligari and Brown (1986).
(b) First clonal year plants

Common practice has been to take a single tuber from each selected
seedling and plant this in the field in the following year (as is shown for the
scheme outlined in Table 8.2). These individual tubers are generally
planted at a seed-growing site, since they form the basis of any future
stock, and selected on a visual basis for agronomic characters. Some
programmes observe foliage characters but mostly selection is on the basis
of the tubers. It should be noted at this stage that plants at a seed site are
generally grown under conditions of a short season, with the haulm
removed before maturity, and often at sites where ware potatoes would not
generally be grown (in order to have isolation from potential disease
sources). These are not therefore normally the conditions under which the
material is destined for, and for which it is supposedly being selected.
The effect of selection at this stage is, like that with the seedlings, subject
to debate. More breeders will put forward positive views to support
selection at this stage and virtually all programmes drastically reduce the
number of clones in the programme based on their observations on this
first clonal year. Indeed, it can be seen in Table 8.3 that the percentage of
the original population which has been discarded by this stage is becoming
more uniform and varies in a narrow range between 85 and 98%. Thus all
these programmes subject the population to selection at this stage and
those that relaxed selection in the seedling stage generally practise even
more severe selection in the first clonal year. This generation has been
even less thoroughly investigated than the seedling one but again formed
part of the large study carried out at SCRI (Caligari et al., 1986; Brown et
al., 1987a, b). Again the evidence for an overall and clearly beneficial effect
of selection at this stage has been lacking (Maris, 1966). Indeed there was
even an indication of a potential negative effect of selection (Brown et al.,
1987b).
(c) Second clonal year plants
The number of plants per clone grown at this stage varies between
programmes but is usually in the order of 3 - 7 and generally these are
grown in a single plot and again at a seed site rather than a ware one. The
methods of assessment and selection also vary, but this does not usually
include objective yield assessments and often relies, as in the first clonal
year, on visual assessment in the field. The efficacy of selection at this stage
seems slightly better (Caligari et ai., 1986) but questions about the effect of
using a seed site as opposed to a ware one in terms of overall selection
effects have been raised (Brown et al., 1987b). Reference once more to
Table 8.3 shows that the percentage of the population discarded by this
stage is really very uniform and only varies between 96 and 99%. Clearly
the main effects of selection in all the programmes must have taken place
by this stage as there is little of the original variation persisting. It therefore
is clear that uncertainties about the efficiency of selection during these
early generations need to be resolved.
(d) A suggested scheme for selection in the early generations

The experiments of various groups have pointed to the inefficiencies of the
widely practised methods of selection in the seedling and early clonal
generations; a particularly comprehensive set of experiments have been
published from work carried out at SCRI (given as references above and a
large part of it is described by Brown, 1988). These experiments were
carried on in the context of the breeding programme described in Table 8.2
and showed very clearly the inefficiencies of the early generation selection
that was practised. As has been stressed these practices were those which
were commonly adopted in potato breeding programmes worldwide. Also
it should be noted that this particular breeding programme at SCRI
(formerly known as the Scottish Plant Breeding Station) had been very
successful in producing commercial varieties and breeding lines which
had, and still do, command large acreages of potato production and were
in demand for breeding and research work worldwide (MacArthur, 1970).
Thus this inefficiency of selection did not prevent the production of
commercially acceptable varieties in the past. This inefficiency almost
certainly applies in other breeding programmes and has probably meant
that such practices have not been examined in the depth they would have,
had the products been less successful. What it effectively has meant is that
much of the hard work and energy expended, in the early generations of
many breeding programmes, was unnecessary (perhaps slightly deleterious
in terms of the direction the population was moved) and that the effective
population sizes, i.e. the later generations, being handled were therefore
much smaller than the breeders appreciated. In other words, selection
could simply have started on a population of the size that could be handled
in the later generations by starting at this size, growing and multiplying it in
a simple manner without assessment or selection.
It would seem inefficient, however, not to try to use the potential for
handling larger numbers in the early generations if an effective method can
be found. What has been shown to provide such a potential is where
attempts to select individuals are abandoned and attention is focused on
choosing the most promising crosses or progenies. Such progeny assess-
ment can take various forms but it is at its most powerful when allied to the
cross-prediction methods (noted earlier as originally suggested by Jinks
and colleagues). The methods they propose use parameters which are
readily calculated and then used to provide probabilities of obtaining a
given value (target value set by the breeder) or, better, based on the
properties of the normal probability integral (Jinks and Pooni, 1976,
1986). The methodology was readily extended to allow simultaneous
prediction over a number of characters (Pooni and Jinks, 1978). Once a
probability has been estimated for a number of crosses or progenies then
Table 8.4 Proposed scheme for the early generation of a potato breeding
programme, based on replacing the scheme in Table 8.2
Year Numbers handled Sites Selection for agronomic characters
1 200 progenies Glasshouse Vis",al assessment of tubers from
(30--60 seedlings of each) individual plants in pots. Data
used give cross-predictions to
allow selection of best progenies
(the number taken depends on
facilities, etc. but perhaps the best
10%. might be reasonable)
2 (a) 20 progenies Ware site Visual assessment of tubers from

(3~0 randomly selected individual plants. Data used to
clones planted in two carry out cross-prediction and
replicates) reduce number of progenies
further: perhaps half of them
(b) 20 progenies Glasshouse No assessment made; three tubers

(400 seedlings of each) harvested for planting in
subsequent year from those ten
progenies identified in the field
trials as having the best potential
3 4000 Seed site Visual selection on tubers from

a three-plant plot to eliminate
grossly uncommercial clones. Also
tubers can be used for some
disease tests, etc.
these can be ranked and the best (i.e. with highest probability of producing
genotypes having the desirable expressions for the characters in question)
can be identified. Progeny assessments have been found to be remarkably
versatile and in many cases robust. They have been investigated, deve-
loped and implemented for a range of characters, not only agronomic ones
but also disease resistance and quality (see p. 358). These investigations led
to cross-prediction methods being tested on a scale equivalent to a major
breeding programme (Brown et ai., 1988) and being implemented as
providing a practical and viable alternative (Mackay et al., 1986). This
implementation can be in various forms but one possible scheme, to
replace the early generations of Table 8.2, is shown in Table 8.4.
The success of such methods has not yet been fully recognized in that not
only can such methods identify the crosses or progenies which have the
greatest potential to produce commercial varieties, they can also be used to
judge the merits of the parents used (Brown and Caligari, 1989). In other
words, if the progenies being examined arise from crossing schemes where
each parent is either crossed to a number of tester lines, or simply used
in a reasonable number of crosses, then the probabilities attached to its
progenies can be used to provide a relative ranking of one parent to
another. This can then form the basis for deciding which parents should be
used in further rounds of crossing.
8.2.7 Selection in the intermediate clonal generations of traditional

breeding programmes
In most breeding programmes the 4th, 5th and 6th years are those in which
the clones are grown in yield trials and where the first main assessments of
quality, storage, etc. are made (see Table 8.2). Traditionally these have
tended to be in non-replicated plots sometimes of increasing size in
advancing years. A decision to be made at this stage is whether to grow the
material at a single site or use several different locations. Many breeders
now recognize the necessity to try to balance the need to be able to
estimate error variances, with the desire to make plots as commercially
relevant as possible, and to cover as representative a range of environ-
ments as possible. For instance, it has been argued that small plots will
have excessive numbers of plants which are at the edge of the plot and thus
make their performances untypical of their potential when subject to
agricultural practice. This again is an area where the evidence is not
complete and it is often recognized that the breeder is less concerned with
absolute performance but must simply judge the entries in a trial relative to
one another, and of course the controls (Caligari et ai., 1985a). Factors
other than simple considerations of increasing the accuracy of estimation
need to be considered when deciding on whether to replicate or not and if
so to what level. Without replication at some level there is no method of
judging whether the trial is basically satisfactory i.e. whether the level
of uncontrolled environmental effects is acceptable or is a cause for
concern. One proposal is to include some varieties in the trial and replicate
these to provide just this sort of measure. This does, however, potentially
have some risks attached unless the vadeties are a representative sample in
terms of characters of interest, and even then the varieties are likely to be
a biased sample. They are biased in the sense that they have been
highly selected to give commercial performances under a wide range of
conditions. They may therefore be regarded, for example, as having been
selected as showing less sensitivity to environmental changes. Thus it
would seem reasonable to replicate the trial material in order to obtain
estimates of the level of certainty with which any mean of a character is
known. This allows the level of confidence to be judged in any selection
procedure undertaken. However, since most commercial cultivars are
destined to be grown over a wide geographic area, it is sometimes argued
that the replication should be over different trial sites.
Selection is applied in each of the years and the numbers of clones is
reduced, as shown in the example in Table 8.2. The scope for selection
during this stage is much more restricted than in the early generations and
thus provides the opportunity for many of the clones to be grown in a
number of years. This is also argued to be a practical level of replication
over a representative range of climatic variables.
8.2.8 Selection in the later clonal generations of traditional breeding

programmes
The number of years that clones are trialled and the number of clones left
in these later stages of the breeding programmes depend largely on the
wishes and desires of the particular breeder, the perceived potential of the
clones present, the market forces operating at that time, etc. Typically, the
number of clones left in the 7th, 8th and 9th generations are those shown in
Table 8.2, where they start at 60 and reduce to about five. These five are
then considered to have fairly clear commercial potential. Because of th.e
relatively small numbers of clones involved and the proximity to com-
mercial exploitation, the trials are usually more sophisticated in terms of
plot sizes and trial designs. They also usually are carried out on a much
wider geographic basis in order that their potential can be assessed on a
practical basis.
8.2.9 Statutory trials and commercialization

Any clones that emanate from the breeding programmes need to be of
value to the customer. Who this is varies with the aims of the programme.
To safeguard the consumer, especially in those countries where Plant
Breeders' Rights have been established, there is a requirement for
statutory trials to be carried out on the clones before they can be given
variety status and commercialized. Such trials cover the obvious areas of
Distinctness, Uniformity and Stability which are clearly required in at-
tempts to control, identify and collect revenue on each variety. They are
often, however, also subject to tests of their commercial attributes (Value
for Cultivation and Use) where they are compared with existing varieties
and expected to show specific improvements over them. Thus the clones
that have survived the various years and trials of the breeding programme
need to fulfil these final requirements before being classified as commercial
varieties (Biitz, 1987; Richardson, 1987; Woude, 1987).
8.2.10 Other breeding methods
(a) Population breeding

This has been carried out in a number of instances, mostly where the aim
has not been to produce varieties for release, but to improve the stocks
from which the selection of varieties can occur. The general philosophy has
been to increase the frequency of desirable alleles within the population
while still retaining genetic diversity. One well known example of this has
already been noted and is that provided by the work at CIP. 'The mandate
of the breeders at the International Potato Center (CIP) is to increase
potato production by helping to develop varieties better adapted to the
growing conditions of developing countries and improving agronomic, seed
production, and storage technologies' (Mendoza, 1987). They therefore
decided that the work at CIP should concentrate on providing populations
of breeding and parental material which could be supplied to developing
countries and then be selected, or crossed and selected, in the more
realistic context of that country. Another example of population selection
is provided by the work quoted earlier relating to the reselection of
spp. tuberosum type clones from spp. andigena forms (Simmonds, 1966;
Plaisted, 1972; Glendinning, 1975b). Indeed a similar philosophy was
employed with diploid potatoes and is described by Carroll (1982).
The methods employed in schemes of population, or 'pre-breeding',
often use, as a basis for decision making, the assessment of individual
clones which are then crossed to produce a new population of seedlings.
Some schemes simply make use of the assessments to identify, in an
empirical manner, the individuals for further crossing (Carroll, 1982).
Others, however, have attempted to gain some element of prediction, for
example by using specific crossing designs and estimating genetical or
statistical parameters (Mendoza, 1987).
The reasons behind the use of wild and primitive species and the
advantages perceived in their use when breeding cultivated potatoes have
been discussed thoroughly elsewhere and a short but comprehensive
review is given by Hermsen (1987).
(b) True potato seed (TPS)
Although, in China, production of potatoes using true seed has been
practised for many years, for most of the rest of the world true seed has
only been used within the context of breeding programmes. However, in
1978, CIP adopted the potential utilization of TPS as a major research
objective and since then it has attracted considerable interest and activity
(Mendoza, 1980; Wiersema, 1983; Jackson, 1987).
Some of the disadvantages of asexual, clonal reproduction have been
noted earlier and the attraction of using true seed must be seen in the light
of these. A major attraction is the fact that most diseases are not
transmitted via true seed and this reduces the difficulties and expense of
maintaining healthy material for multiplication and production. Another
factor is simply related to the bulk of seed tubers that need to be
transported from multiplication sites to ware growers compared with the
packets of seed when TPS is used. As might be expected TPS also has its
drawbacks. First, TPS derived plants require a longer growing season to
reach full maturity, a factor which can present problems in countries with
large seasonal differences in climate. Secondly, the true seed from hetero-
zygous parents will, as would be expected, show the result of genetic
segregation following meiosis. In many countries this potential lack of
uniformity in the tuber crop will provide a barri,er to its use. Such problems
can, however, be solved in a number of ways: (a) parents which produce
progenies with reasonable small phenotypic variances can be selected on
an empirical basis; (b) parental material can be inbred, at least partially,
in order to make it more genetically homozygous; or (c) true breeding
material can be produced. This later possibility is becoming an ever
increasing reality by the development of a variety of techniques, some of
which are described below.
(c) Dihaploids, monohaploids and inbred lines

The first report of the production of diploids from cultivated tetraploid
tuberosum was in 1957 (Hougas and Peloquin, 1957) who used a diploid
species, Solanum phureja, to induce the production of parthenogenetic
diploid plants. Later anther culture was successfully carried out (Dunwell
and Sunderland, 1973; Sopory et al., 1978) and gave an alternative method
for producing plants with the diploid number of chromosomes, which will
be referred to here as dihaploids. These techniques opened the possibility
of breeding at the diploid level (Hougas and Peloquin, 1958) and thus
circumventing some of the problems associated with breeding a tetraploid
crop (Hermsen and Ramana, 1981). This has led to assessments of the
potential for using dihaploids in breeding programmes (e.g. De Maine,
1982) and proposals to increase the efficiency of production (Uhrig and
Salamini, 1987; Caligari et al., 1988).
Further work on these techniques allowed an even more exciting
possibility to emerge; this was the production of monohaploid plants. In
other words by repeating the types of techniques used at the tetraploid
level, but now on the dihaploids, it was possible to obtain plantlets with the
haploid number of chromosomes (Breukelen et ai., 1975; Foroughi-Wehr
et ai., 1977; Uhrig, 1985; Uijtewaal et ai., 1987a). Clearly, doubling the
chromosome number of the mono haploid plants obtained will give
homozygous genotypes at the diploid level and doubling these, homo-
zygotes at the tetraploid level. This therefore gives a route to producing
true breeding lines for use as parents or directly in the form of TPS. The
problem frequently stated, however, is that many of the homozygous
genotypes produced in this way are not very high yielding and usually have
very poor fertility (Uijtewaal et ai., 1987b). This is, of course, expected
when normally outbred material is made homozygous and is usually
attributed to inbreeding depression. The causation of this depression is,
however, sometimes ascribed to the exposure of deleterious recessive
alleles maintained in the population which are masked in heterozygotes
and sometimes simply to a reduction in heterozygosity per se. Inbreeding
depression is the counterpart to heterosis, which is seen as the excess
performance of heterozygotes over their homozygous parents, and is often
described with the implication of the presence of overdominance being the
major factor. Little evidence exists for the widespread occurrence of
overdominance and a more realistic interpretation lies in the dispersion of
favourable genes and epistatic interaction (Jinks, 1983). This then means
that it will be possible to derive and select inbred lines that will perform as
well as any outbred clone and is more a matter of the expenditure of time,
resources and energy than underlying biological limitations. In any case,
this genetical basis also means that crosses between even mediocre inbred
lines will show heterosis and allow acceptable levels of performance to be
achieved. Thus the use of F} hybrids in potatoes could allow the potential
of TPS to be exploited readily. It suffers one drawback, however: to
produce the F} seed requires, at present, hand pollination which is time-
consuming and expensive and so a method of encouraging cross-pollination
is required for a practical scheme and none is available at present. The
potential of inbred material is therefore perhaps a sensible objective to
pursue. There are indications already that this may not be necessarily
too distant an objective in that inbred lines of sufficient performance
may be obtained by judicious choice of parental material, as might be
expected (Jackson, 1987). Indeed the use of the techniques cited for
producing monohaploids means that inbred lines might be derived in two
generations from a tetraploid genotype and once this is done in large
numbers successful homozygous genotypes could quickly be identified if
present.
Breeding objectives 353
8.3 BREEDING OBJECTIVES
8.3.1 Overall aims and objectives

The aim of all potato breeding programmes has been to supply the growers
and potato industry with improved varieties which make their task easier
or more profitable. This over-simplified statement, however, hides a
multitude of factors. In South America the Indians not only eat fresh
potatoes but also a type of dried potato which they can store and use when
required (Hawkes, this volume, Chapter 1; Anon., 1980). In many of the
developing countries potatoes are used both in a fresh form as well as in
processed products such as French fries, chips (crisps), dehydrated powder
or canned, small potatoes. In addition potatoes are also used as (a) a
source of starch, (b) for distillation and (c) for fodder (Anderson and
Horton, this volume, Chapter 16). The characteristics required for all these
uses are not identical but there are, of course, overlaps in the require-
ments. Thus, for instance, all users are benefited by increasing disease
resistance in the crop and stored product, at least to some extent. Thus
while French fry producers generally prefer a long-oval tuber of a
moderate size, chip (crisp) producers like a high dry matter potato of a
small round shape, and for domestic, fresh use the general preference is for
a medium size, smooth skinned round-oval tuber. It is therefore important
in any breeding programme to determine the market, or markets, that are
to be the targets of the selection. To add to these complications, it is also
very clearly necessary to decide what is the geographical area that is being
bred for. Not only are there great differences in the climate, agronomic
practices in different areas, disease spectra and priorities for use but also
different consumer preferences (e.g. skin and flesh colour).
A generalized list covering some of the characters that are typically
taken into account in breeding programmes is given in Table 8.5. As can be
seen there are a substantial number of characters noted and the list is not
exhaustive. Thus, although any individual breeding programme does not
necessarily select for all these, and not all at the same time, they
must consider a majority of them and this, of course, makes breeding
particularly demanding in this crop.
8.3.2 Breeding for agronomic characters, yield and tuber morphology

Some of the characters considered specifically under this category are given
in Table 8.5. However, as was noted earlier, in the early generations most
breeding programmes rely on an overall visual appraisal, which would take
many of these characters into account within a single score. This score is
either recorded for future use or simply forms the basis for a decision to
select, and therefore retain a clone, or discard it from the programme. It is
only in the later generations (from about year 4, Table 8.2) that more
Table 8.5 Some typical characters to be considered in breeding programme
(1) Agronomic and yield Bulking rate, heat tolerance, marketable yield,
maturity, number of tubers, tuber size and
distribution, yield of tubers
(2) Tuber morphology Eye depth, flesh colour, growth cracks, hollow heart,
regularity, secondary growth, shape, skin colour
(3) Quality After-cooking blackening, dry matter, enzymic

browning, glycoalkaloid level, reducing sugar
content, sloughing, storage characters, taste, texture
(4) Resistances Disease resistance: Bacterial soft rot and black

leg, black scurf, common scab, dry rot, early blight,
gangrene, late blight (foliage and tuber), powdery
scab, silver scurf, spraing, verticillium wilt,
viruses (PVX, PVY, PLRV, etc), wart
Pest resistance: Aphids, colorado beetle, cyst
nematodes (G. rostochiensis, G. pallida), root-knot
nematodes.
Mechanical damage: Brittle cracking, bruise
resistance, scuffing
Miscellaneous: Herbicides, internal rust spot, wind
damage.
objective methods are used to assess the relevant characters. At these later
stages the produce of the plot is graded, weighed, tubers are cut and
internal condition examined. However, many of the characters are still
only sensibly assessed on the basis of visually appraising the material, but
now on the basis of individual characters, and recording the score on a
predefined scale. (A 1-9 scale seems a particularly appropriate one where
only the odd numbers are commonly used, i.e. 1,3,5,7 and 9, giving five
scoring points but when averaged over replicates, etc. providing an easily
appreciated impression. It is also helpful to standardize the direction of the
scales and one in which 1 is the least desirable in agronomic terms and 9 the
most desirable has been found to work very well in a practical context.) A
good example of why such visual appraisal is needed for many of the
characters is provided by the consideration of eye depth. It would, of
course, be possible to measure the depth of a number of eyes on a range of
tubers and record the result. It would, however, be extremely time
consuming. Other less direct methods of assessing this character might also
be possible but have not yet been shown to work well in practice or
correlate well with the one of interest. Thus a single score, based on visual
appraisal of eye depth, is used.
Thus the various characters can be assessed and recorded but there is a
complication that must be considered. The breeder wishes to carry out
trials in the most agronomically representative manner possible such
that the assessment will most accurately reflect the clones' commercial
potential. However, the material that is handled will reflect a wide
diversity for most of the characters, for instance foliage maturity, bulking
rate, root spread, etc. In other words in agronomic reality these different
clones would be handled very diffe:ently with respect to control of their
sprouting, their time of planting, their tuber spacing when planted, their
harvest date, etc. The breeder is faced with material with unknown
characteristics and an almost continuous range of expression for most of
the characters. All that is possible, except towards the very end of the
programme, is to grow all the material under 'average' conditions and
make some allowance for this in the assessment. It is also possible to
impose a few rather major differences in treatment for a few of the
cultivation variables and thus assess the material under some more extreme
conditions. For example, it is possible once the material has been reduced
somewhat in number to plant trials on different dates, one early and one
late, the early trial material perhaps being pre-sprouted and treated as if it
all had the ability to become 'early varieties'. Ones with a potential to bulk
early could thus be identified. Nevertheless, it generally means that it is not
possible to handle the individual clones in terms of agronomic practice and
husbandry in an individual way as is possible with a variety.
8.3.3 Breeding for quality characters

The characters listed in Table 8.2 affect, to varying degrees, aspects of
potato quality when eaten fresh. They are generally difficult to assess as
characters except by actually cooking mature tubers. This has to some
extent limited the potential for selecting during the breeding programme to
a stage where the numbers have been reduced to manageable levels and
sufficient tubers can be spared for destructive testing (generally after year 4
onwards, Table 8.2). However, the success of using cross-prediction to
assess progenies rather than individual clones in the early generations (see
p. 347), has been successfully adopted for some of the quality characters
(Caligari and Brown, unpublished), particularly for sloughing, after-
cooking blackening and crisp colour (largely a reflection of reducing sugar
content, Storey and Davies, this volume, Chapter 12). The predicted
ranking of six crosses based on data collected on the progenies grown in
1984 and the observed results based on tests carried out in the subsequent
year, 1985, on all the individual clones within each of the progenies are
shown in Table 8.6. The data show the rankings of the progenies based on
the combined probability of obtaining clones with a better expression of
the characters crisp colour and after-cooking blackening than the overall
mean of the popUlation of clones observed. As can be seen the sample of
Table 8.6 Rankings of six crosses based on
the observed and predicted proportion of
clones having better than the overall progeny
average expression of the characters crisp
colour and after-cooking blackening
Progeny Predicted 1984 Observed 1985
1 4 2
2 2 1
3 6 5=
4 3 4
5 1 3
6 5 5=
progenies shown is small but nevertheless there is clearly reasonable

agreement betwen observed and expected. There is therefore the potential
to include the quality characters when identifying progenies in the early
generations using cross prediction. The quality characters could thus be
selected equally vigorously as other aspects of importance.
When processing quality is considered specifically then, as noted earlier,
the characters which are of primary importance vary with the processing
that is to be carried out (Ross, 1986). Dehydration as a process does not
have very specific requirements for its potato raw product, whereas for
French fries the general requirements are for modest dry matter (prefer-
ably specific gravity just greater than 1.08) and not too high levels of
reducing sugars. For chips (crisps) the specifications are more exact and the
requirements are for tubers with a high dry matter (specific gravity in
excess of 1.085) and with low reducing sugar levels. The developments
noted above in terms of cross-prediction can be used to help select for
these requirements. A method that could be used as an alternative or in
conjunction with such prediction has been proposed by Louwes and Neele
(1987). They used salt solutions to assess dry matter content and glucose
strips to indicate levels of reducing sugars.
Not only do chip (crisp) manufacturers require low levels of reducing
sugars at harvest but also after storage. Indeed, they like to store potatoes
for lengthy periods so that they can spread their manufacturing more
evenly over time. Unfortunately at low temperatures, at which storage is
best in terms of lessening tuber sprouting, loss of water, etc., reducing
sugars are accumulated in all commercially suitable varieties available at
present. However, it was discovered that variants did occur among the
germplasm at SCRI; indeed they were present among the early generations
of material in the commercial programme (Mackay et al., 1985) at a
frequency of about 2-3%. The way has thus been opened to increase the
range of selection to include characteristics of reducing sugar accumulation
at low temperatures.
8.3.4 Breeding for resistances
(a) Diseases and pests

The potato is no exception to other crop plants in suffering from attack by
numerous pests and diseases (a sample is shown in Table 8.5). The
prospect of handling all these with equal effort is not only daunting but
probably impossible. Nevertheless breeding for resistance potentially
provides the most economical, in the long term, and moreover the most
environmentally sympathetic approach for controlling the effects of pests
and diseases on crop plants. It is interesting to note that in many ways the
history of potato breeding has been largely determined by the effects of
pests and diseases. The early growers found that their stocks degenerated
over time to become less vigorous and productive. As was noted earlier
this was largely due to the effects of virus infection which was then
perpetuated by the clonal reproduction of the potato. They soon dis-
covered that this could be ameliorated by starting new varieties from seed
and limiting their lifespan before replacement, since most viruses are not
transmitted through seed. Thus the role of the breeder was established.
Another major landmark in the history of potatoes in Europe was the
potato blight (caused by Phytophthora infestans) in the nineteenth century.
This led to a renewed upsurge in potato breeding which continued into the
twentieth century.
The many and different diseases and pests of potatoes have differing
priorities in breeding programmes depending on the countries in which the
varieties produced are to be grown. On average, pests and diseases are
estimated to cause in the region of a 22% yield loss.
As with the majority of crops the early breeding work of the twentieth
century concentrated mainly on resistance mediated by major genes .. These
genes could be readily manipulated by the breeder once Mendel's work
was rediscovered in 1900 and led Bateson (1908) to comment on Biffen's
finding of a single, major gene conferring resistance to rust in wheat as
being 'of fundamental importance to the physiology of disease'. It became
clear however, that in many cases such resistance was short lived because
of the ability for the causal organism to evolve to overcome the resistance.
A good example of this is provided by the early work on late blight.
In about 1909 resistance to late blight was demonstrated in wild species
by Salaman and also by Wilson working with S. demissum. Using various
crossing schemes the resistance was crossed into S. tuberosum and showed
clear segregation ratios. These resistance genes became known as the R
genes and were used widely in breeding programmes. Their use was,
however, short lived since once into commercial production they soon
succumbed to late blight. Indeed in 1951 a seedling was registered by the
Scottish Plant Breeding Station (later to become SCRI) and called
'Pentland Ace'. The records show that the variety was 'immune to several
biotypes of the blight fungus ... In 1953, when the total acreage was only
nine acres, the crop suffered a catastrophic blight attack, the ace had been
trumped .. .' (MacArthur, 1970). As we now know pathogens, particularly
ones with airborne spores, can evolve rapidly to overcome such major gene
resistance. The breeders then turned their attention to 'field resistance'
which was also a feature present in S. demissum (Toxopeus, 1964; Black,
1971). This resistance was controlled by many genes each of rather small
effect (Killick and Malcolmson, 1973) and was typical of polygenic ally
controlled resistance. Because of its polygenic nature and range of
phenotypic expression such resistance was more troublesome to handle.
The early tests involved empirical field trials (Malcolmson, 1976) but
glasshouse screens were also devised to provide some preliminary screen-
ing. Further modification included the use of leaf discs, detached leaves or
leaflets. More recently seedling screens have been proposed which allow
large numbers to be screened under glasshouse conditions and hence
enable the breeders to apply a higher selection pressure for suitable field
(horizontal) resistance. It can also be used to assess progenies and thus
form an integral part of the cross-prediction approach noted earlier
(Caligari et ai., 1984, 1985b). While at present the disease is managed by
preventive spraying with chemical fungicides to complement host resistance
(Fry, 1981), the new-found ability to apply much higher selection pressure
allied with an effective methodology for handling the polygenic variation
gives hope for the future. The position with respect to tuber blight is in
many ways similar and, although its routine testing is more troublesome,
progeny tests have now been devised (Wastie et ai., 1987).
There are, however, examples of the very effective use that can be made
of major genes in other circumstances. The resistance for potato virus X
(PVX) , which is already widely distributed in commercial varieties, is
controlled by major genes. There were, in fact, two sources of major gene
resistance, S. tuberosum ssp. andigena and S. acauie; these genes have
been incorporated into commercial backgrounds and are proving success-
ful in providing immunity to PYX. A somewhat similar picture emerges in
relation to potato virus Y (PVY) in that major gene resistance is available
(Cockerham, 1970). Some of these major genes are strain specific while
others give comprehensive resistance to all strains of PVY (e.g. those
derived from S. chacoense, S. demissum, S. microdontum and S.
stoioniferum). These genes conferring comprehensive resistance to PVY
are however only just reaching the stage of being incorporated into
backgrounds which are sufficiently good to have commercial status. They
have therefore to be tested in their practical context of wide commercial
distribution.
A very different picture emerges when resistance to Potato Leaf Roll
Virus (PLRV) is considered. No major gene has been identified which
confers resistance (Davidson, 1980). The breeder is thus faced with having
no choice but to try to exploit the polygenic resistance for this virus. This
is made more troublesome by the nature of infection which, at least until
recently, involved exposing the plants to the infector vector (aphids) in one
year and screening the resultant infection in plants from daughter tubers
the next year. The process is made even more cumbersome by the uneven
rates of infection due to environmental causes and the variability in
expression of the symptoms (Davidson, 1973). As might be expected,
methods have been developed to try and reduce such problems of handling
the screening and underlying genetic determination. One development has
seen the use of cDNA probes to detect the presence of PLRV and hence
eliminate the need for elaborate field trials in the second year (Boulton et
al., 1987) while another has tried to use primary symptom expression and
enzyme-linked immunosorbent assay to allow the development of a rapid
progeny test (Soloman et al., 1987).
Another example where the use of major genes has proved beneficial for
providing resistance to one species, while only polygenic variation has
been found to be available for another, is with respect to Potato Cyst
Nematodes (PCN). These are discussed by Evans and Trudgill (this volume,
Chapter 11) but in outline there are two species of cyst nematode that are
found on potatoes, Globodera rostochiensis and G. pallida (Stone, 1972)
and the chemical control of these soil-borne pests is an expensive variable
cost to the grower. Resistance to G. rostochiensis was found by Ellenby
(1948, 1954) as well as Mai and Peterson (1952). The material screened by
Ellenby included Commonwealth Potato Collection accession CPC 1673 of
the ssp. andigena which has been subsequently used widely in breeding as
the source of the dominant single major gene H j conferring resistance to
pathotypes Ro-1 and Ro-4 of G. rostochiensis. Although only effective
against these two pathotypes, they are commonly occurring ones and in fact
in the United Kingdom only Ro-1 is present. This gene has been incor-
porated in a number of breeding programmes and resulted in commercial
varieties with resistance but as far as the breeder is concerned there is a
further complication. Such resistance operates after the nematode has
invaded the root and caused damage to the plant; thus a crop planted with
a variety carrying this gene still suffers a yield loss unless it is tolerant to
attack (Evans and Trudgill, this volume, chapter 11). This tolerance is
controlled by other genes independent of those conferring resistance
(Holden, 1977; Dale, 1987) and those are not of the simple major type.
There are several sources of resistance to the other species of cyst
nematode, namely G. pallida, particularly ssp. andigena and S. verneii; the
former was at first thought to give resistance controlled by a single major
gene but was later taken to be polygenic and treated accordingly. The
resistance from S. verneii was also the subject of debate but is now
generally accepted to be polygenic in determination. The variations from
both sources, even if some lingering uncertainties remain, are handled as if
they were polygenically inherited as far as practical breeding is concerned
(Forrest and Holliday, 1979; Phillips et al., 1980; Phillips and Dale, 1982).
The results of using this type of resistance are just beginning to produce
results in terms of finished varieties (Anon., 1989).
The above are given in some detail but still only treated in a superficial
manner and the genetics and breeding of each could easily become a
chapter in its own right. There are many other diseases that should be
drawn to attention and some of these are listed in Table 8.5. Most of them
have been found to have, associated with them, variation in the potato and
this mainly, as expected, of a polygenic type. Many of the relevant aspects
have been reviewed by Ross (1986) and hence this will not be attempted
here. However several additional points can perhaps be usefully added to
Ross's review. One is that over the last few years much effort has been
expended in attempts to devise progeny tests for many of the diseases that
can be carried out on seedlings or 1st clonal year plants (Phillips, 1981;
Plaisted et al., Lacey et al., 1987; Soloman et al., 1987; Wastie et al., 1987,
1988a). This effort has been considered worthwhile since the potential
return in terms of the increase in selection efficiency and hence variety
production is large. It would seem that many such tests are indeed possible,
as we have already noted for late blight, and they can be combined with the
cross-prediction used for other characters. Another area of endeavour has
been continued work on increasing the potential of field and glasshouse
trials to handle large numbers of entries (Bourne et al., 1981; Caligari and
Wastie, 1985; Caligari and Nachmias, 1988; Nachmias et al., 1988; Wastie
et al., 1988b). Many of the more traditional tests for pests and diseases
have been devised to give a clear and absolute value to the resistance
or tolerance of established varieties. This has usually meant handling
known material with a reasonable supply of tubers, etc. With breeder's
clones, however, the material is usually of unknown behaviour for most
characters, there are large numbers of them and the amount of each is
severely restricted. In addition the breeder is only generally interested
in the relative performances, or rankings, and not so concerned with
absolute value. The main concentration of many pathologists has been in
the first sort of testing, and this has therefore led to the somewhat slower
development of suitable methods for screening larger numbers of breeder's
clones.
(b) Mechanical damage

A major component of quality as far as the domestic consumer is
concerned is the overall appearance of the tuber in terms of freedom from
greening, disease and damage. These are the characters which are most
readily seen and which first affect the choice of whether or not to buy. Not
only is freedom from damage, both external and internal (bruising or black
spot), therefore important for the domestic consumer but also for the
processor. It causes increased waste, particularly in terms of extra peeling,
and a need for more intensive inspection and handling. Having noted this,
however, there is little information available about the potential for
breeding resistance to such damage.
There are several problems as far as the breeder is concerned with
selecting for increased resistance. The first is that it is a character which is
highly dependent on other variables, such as temperature, tuber size, cell
turgidity, husbandry, soil type and, of course, harvesting techni'ques
(McRae and Fleming, 1989). Potatoes can always be damaged, and
damaged quite easily, if handled badly. The second problem is that there
is not a completely satisfactory way of measuring resistance on small
numbers of tubers using a test which is quick and easy and yet still
correlates well with field performance (Hughes, 1980; De Maine, 1986,
1988; De,Maine and Caligari, 1988).
Despite these difficulties most breeders consider resistance to be im-
portant (Scholtz, 1987) and usually take it into consideration when scoring
the produce from their trials. Thus they generally select against susceptible
clones during the later stages of their programmes when they are handling
reasonable size plots which are mechanically harvested. Such plots are thus
more likely to be subjected to the type of damage that might be exerted
under agricultural practice. The effects of such selection are becoming
apparent even if progress has not been very pronounced (Munzert, 1987).
8.3.5 Combining characters to produce commercial varieties

The various characters that have been considered in this section, when
taken with others that breeders may consider, mean that some 50 or more
need to be taken into account when trying to select the one or two clones
that are likely to achieve variety status. Clearly not all the characters can
be considered at the same time or with an equal priority; nevertheless the
breeder has always to choose between clones with an array of different
advantages and disadvantages. Clearly the breeder can select for a few
characters at a particular stage and ignore others until a later clonal
generation. Sometimes this is unavoidable because of the number of tubers
available, or the expression of the character requires a particular minimum
plot size, etc. At other times the breeder will deliberately concentrate on a
particular attribute in order to make progress with it and relegate other
attributes to a more secondary role until it is firmly established in a
background that makes full commercial selection realistic. For example,
when disease resistance is introduced from a wild species or primitive
varieties it is this resistance that is of primary concern; having established
its expression, then a choice between remaining clones is based on other
characters.
The combining of characters is always problematical; the decision about
the relative weightings to give to various characters is never an easy and
straightforward one. In the early generations the breeders often avoid
having to make an explicit weighting between characters but rely, as noted
earlier, on an overall visual assessment (breeder's preference). Such an
assessment enables the breeder to take into account any of the characters
that can be seen and allow the integration, in his mind, of these to give an
overall measure of its commercial worth. In later generations this becomes
less easy or acceptable. Nevertheless having scored the characters
objectively the ultimate decision about the acceptability of the combina-
tion of character expressions recorded is one for the breeder to make. The
use of predefined indices, based on various factors such as economic
weights, accuracy of measurement, etc. is also a possibility. This effectively
provides weights by which the characters are multiplied; then they are
added to give an overall score on which the clones are ranked. The top
scoring ones are then selected and the only decision, in theory, is how far
down the list it is acceptable to go. Another approach, which fits well with
the methods of cross-prediction mentioned earlier, is simply to set target
values for each character. These target levels would usually be set initially
at the minimum level that would be acceptable, in the view of the breeder,
for a commercial variety to emerge from the programme and, as for the
index method, they can take into account information about importance,
repeatability, etc. The target values for each character are then applied to
each clone and if it falls below anyone of them it is rejected. If this leads to
an excessive number of clones being retained the target values can be re-
assessed. This still leaves the problems associated with how a very good
performance in several years or at several sites is balanced against a poor
one at another. In the end the breeder must face a practical decision based
on his experience and knowledge.
8.4 THE POTENTIAL ROLE OF OTHER BREEDING TECHNIQUES
The potential for techniques other than recurrent phenotypic selection in

the breeding of a clonally reproduced potato crop has already been noted.
The change to using true seed to grow potatoes has clear implications for
breeding objectives and methodologies. The use of dihaploid or mono-
haploid clones to reduce ploidy levels or even produce true-breeding
genotypes has also been mentioned.
The potential to multiply any clone or variety from one plantlet to 1
million in 1 year using in vitro micropropagation has interesting potential
but, as noted earlier, is not of such direct consequence to the breeder.
However, another aspect is that it can also be adapted to give slow-growth
cultures and hence allow the maintenance of healthy stocks, from which
plants can be produced for trialling. These cultures can be used to store or
multiply the number of plantlets and then used to give a disease-free nuclear
stock for the few clones that are successful. However, the performance
of plants raised from micropropagated plantlets is not always predictable
and does not necessarily correlate directly with the performance of plants
The potential role of other breeding techniques 363
raised from conventional tubers (Caligari and Powell, 1989; Powell et al.,
1989). Leaving this possibility aside the multiplication rate of potatoes does
not generally restrict the work within the breeding programme itself. It
does, of course, mean that any clone which achieves variety status can be
multiplied much more rapidly for commercial exploitation. The same
considerations basically apply to the potential use of microtuber produc-
tion (Chandra et al., 1988) in culture or mini tubers in the glasshouse (Ross,
1986).
It has been found by empirical means that the Solanaceae, including
potatoes, are relatively amenable to various in vitro tissue culture techniques
and manipulation. Potatoes have thus been used fairly widely in the develop-
ment work underlying many biotechnology techniques. The application of
these techniques is, however, only just reaching the stage of being readily
available to the breeder and hence being subject to direct comparison with
the more traditional methods used. It is clear that these newer techniques,
such as gene transfer, will increase the scope of the breeders' 'armory of
techniques' but will be used alongside the rest of their traditional range of
techniques rather than replacing them.
The induction of somaclonal variation has been widely reported as
providing a useful source of variation for breeders (Larkin and Scowcroft,
1981; Evans, 1989). Such variation has been shown to be produced in
potatoes regenerated from protoplast culture by Shepard et al. (1980).
Subsequently, its production has been demonstrated whenever a callus
phase is involved in going between an explant and a regenerated plant. The
underlying mechanisms are still not fully understood, although it has now
been clearly shown that at least some of the original variation reported
could have been ascribed to gross cytological changes (Karp and Bright,
1985). In practice it has yet to be shown to be of major significance and is
likely to find only a minor role in breeding potatoes in an overall sense
(Thompson, 1987).
Potato breeding has been characterized by the frequent use of wild
species and primitive varieties to increase the variability available to the
breeder, especially in terms of disease resistance. However, it is not always
easy to cross sexually other species with ssp. tuberosum (Hermsen, 1987).
Indeed it is not always easy to cross within ssp. tuberosum because of the
inherent lack of fertility in many of the desirable clones and varieties. The
production of somatic hybrids via protoplast fusion has been proposed as a
useful method of overcoming these barriers to normal crossing in potatoes
as well as other species (Evans, 1983; Jones, 1987) and has been success-
fully carried out in a few cases (Austin et al., 1985; Haberlach et al., 1985;
Fish and Jones, 1988; Fish et al., 1988). The potential for such fusions is
promising but will, of course, be only a small part of the overall breeding
strategy; once such fused products have been obtained they will still need
to be crossed and their progeny selected unless asymmetric fusion is used
(Scheider et al., 1985). Even with this modification it is unlikely that
somatic hybridization will form the routine method of further transfer after
the initial, essential introduction.
An even more refined level of transfer of genetic material into com-
mercial species and varieties is now possible in terms of single, specific
genes. This has been shown to be possible in potatoes and has the potential
not only to allow transfer from species that are related to commercial
potatoes but from plants in other genera, indeed in principle from any
other organism (Ooms, 1987). The methods used and the details of such
technology are beyond the scope of this chapter but the potential has now
been raised and it is likely that this technique will form a very powerful
additional source of variation for the breeder. The ability to transfer a
desirable gene, without the need subsequently to eliminate all the undesir-
able other genes that are usually a consequence of sexual crossing from a
wild ancestor, is useful in its own right. The ability to incorporate genes
from even less closely related species is even more exciting, including the
transfer of genes such as those coding for virus coat protein to provide
novel forms of resistance (e.g. Hoekema et at., 1989). The strength and
attraction of such techniques are also a restriction. They allow single genes
to be transferred and as such are not able to handle characters controlled
by polygenes in any ready manner.
The new techniques will thus find a very useful place in breeding
schemes and should be welcomed as increasing the possibilities that the
breeder can use. They will, however, be used as adjuncts to the more
classical methods of breeding, which will themselves continue to evolve in
efficiency as the underlying mechanisms and properties of genetic systems
become more clearly known. It will be informative to see how much our
knowledge of the genetic map of potatoes is increased, particularly as it
becomes possible to exploit new techniques, and how this effects breeding
strategies.
REFERENCES
Anderson, J.A.D. and Howard, H.W. (1981) Effectiveness of selection in the early
stages of potato breeding programmes. Pot. Res., 24, 289-99.
Anon. (1980) History of the potato, in Potato Handbook (ed. D. Anderson),
Prince Edward Island Potato Marketing Board, pp. 7-18.
Anon. (1989) Classified list of potato varieties, England and Wales, 1989. Nat. [nst.
Agric. Bot., 16pp.
Austin, S., Baer, M.A. and Helgeson, J.P. (1985) Transfer of resistance to potato
leaf roll virus from Solanum brevidens into Solanum tuberosum by somatic
fusion. Pl. Sci., 39, 75-82.
Bateson, W. (1908) The Methods and Scope of Genetics (An Inaugural Lecture),
Cambridge University Press, Cambridge.
Bartholdi, W.L. (1942) Influence of flowering and fruit upon vegetative growth and
tuber yield in the potato. Tech. Bull. Minn. Exp. Sta., 450, 20pp.
References 365
Batz, W. (1987) Potato variety assessment in the Federal Republic of Germany, in
The Production of New Potato Varieties (eds G.J. Jellis and D.E. Richardson),
Cambridge University Press, Cambridge, pp. 110-13.
Black, W. (1971) Researches in potatoes at the Scottish Plant Breeding Station.
Ann. Rep. Scot. Pl. Breed. St., 1970-1, pp. 52--60.
Blomquist, A.W. and Lauer, F.I. (1962) First clonal generation potato progeny
performance at two Minnesota locations. Am. Pot. J., 39, 460-3.
Boulton, R.E., Jellis, G.J. and Squire, A.M. (1987) Breeding for resistance to
potato viruses with special reference to cDNA probes, in The Production of
New Potato Varieties (eds G.J. Jellis and D.E. Richardson), Cambridge
University Press, Cambridge, pp. 86-7.
Bourne, W.F., McCalmont, D.C. and Wastie, R.L.W. (1981) Assessing potato
tubers for susceptibility to bacterial soft rot (Erwinia caratovora ssp. atroseptica).
Pot. Res., 24, 409-15.
Breukelen, E.W.M. van, Ramana, M.S. and Hermsen, J.G.Th. (1975) Monoha-
ploids (n=x=12) from autotetraploid Solanum tuberosum (2n=4x=48) through
two successive cycles of female parthenogenesis. Euphytica, 24, 567-74.
Brown, J. (1988) The efficiency of selection in the early generations of a potato
breeding programme. Ph.D Thesis, University of St Andrews, UK.
Brown, J. and Caligari, P.D.S. (1986a) The efficiency of seedling selection for yield
and yield components in a potato breeding programme. Ziet. fur Pjlanzenz., 96,
53--62.
Brown, J. and Caligari, P.D.S. (1986b) Cross prediction methods in potato
breeding programmes. Proc. of Eucarpia (Biometrics Section), Birmingham,
1986, pp. 348-55.
Brown, J. and Caligari, P.D.S. (1987) The use of multivariate cross prediction
methods in the breeding of a cion ally reproduced crop (Solanum tuberosum).
Heredity, 60, 147-53.
Brown, J., Caligari, P.D.S., Mackay, G.R. and Swan, G.E.L. (1984) The
efficiency of seedling selection by visual preference in a potato breeding
programme. J. Agric. Sci., Camb., 103, 39--46.
Brown, J. and Caligari, P.D.S. (1989) Cross pfediction in a potato breeding
programme by evaluation of parental material. Theor. Appl. Genet., 77,
246-52.
Brown, J., Caligari, P.D.S. and Mackay, G.R. (1987a) The repeatability of
progeny means in the early generations of a potato breeding programme. Ann.
Appl. Bioi., 110, 365-70.
Brown, J., Caligari, P.D.S., Mackay, G.R. and Swan, G.E.L. (1987b) The
efficiency of visual selection in early generations of a potato breeding
programme. Ann. Appl. Bioi., 110, 357--63.
Brown, J., Caligari, P.D.S., Dale, M.F.B., Swan, C.E.L. and Mackay, G.R.
(1988) The use of cross prediction methods in a practical potato breeding
programme. Theor. Appl. Genet., 76,33-8.
Caligari, P.D.S. and Brown, J. (1986) The use of univariate cross prediction
methods in the breeding of a cion ally reproduced crop (Solanum tuberosum).
Heredity, 57, 395--401.
Caligari, P.D.S. and Nachmias, A. (1988) Screening for field resistance to early
blight (Alternaria solani) in potatoes. Pot. Res., 31,451--60.
Caligari, P.D.S. and Powell, W. (1989) Variability in response of potato cultivars
to micropropagation. I. In vitro performance. Ann. Appl. Bioi., 115,
115-21.
Caligari, P.D.S. and Wastie, R.L. (1985) Assessment of a glasshouse test for
measuring the resistance of potato cultivars to common scab. Pot. Res., 28,
379-87.
Caligari, P.D.S., Mackay, G.R., Stewart, H.E. and Wastie, R.L. (1984) A
seedling progeny test for resistance to potato foliage blight. (Phytophthora
infestans (Mont.) de Bary). Pot. Res., 27,43-50.
Caligari, P.D.S., Brown, J. and Manhood, C.A. (1985a) The effect of varying the
number of drills per plot and the amount of replication on the efficiency of
potato yield trials. Euphytica, 34, 291-6.
Caligari, P.D.S., Mackay, G.R., Stewart, H.E. and Wastie, R.L. (1985b)
Confirmatory evidence for the efficacy of a seedling progeny test for resistance
to potato foliage blight (Phytophthora infestans (Mont.) de Bary). Pot. Res., 28,
439-42.
Caligari, P.D.S., Brown, J. and Abbott, R.J. (1986) Selection for yield and yield
components in a potato breeding programme. Theor. Appl. Genet., 73,218-22.
Caligari, P.D.S., Powell, W., Liddel, K. et al. (1988) Methods and strategies for
detecting Solanum tuberosum dihaploids in interspecific crosses with S. phureja.
Ann. Appl. BioI., 112, 323-8.
Carroll, C.P. (1982) A mass-selection method for the acclimatization and im-
provement of edible potatoes in the United Kingdom. J. Agric. Sci., Camb., 99,
631-40.
Chandra, R., Dodds, J.H. and Tovar, P. (1988) In vitro tuberization in potato
(Solanum tuberosum L.). Int. Ass. Pl. Tiss. Cult. News., 55, H~20.
Cockerham, G. (1970) Genetical studies on resistance to potato viruses X and Y.
Heredity, 25, 309-48.
Dale, M.F.B. (1987) Breeding for resistance to and tolerance of potato cyst
nematode, in The Production of New Potato Varieties (eds G.J. Jellis and D.E.
Richardson), Cambridge University Press, Cambridge, pp. 91-3.
Davidson, T.M.W. (1973) Assessing resistance to leafroll in potato seedlings. Pot.
Res., 16, 99-108.
Davidson, T.M.W. (1980) Breeding for resistance to virus disease of the potato
(Solanum tuberosum) at the Scottish Plant Breeding Station. Ann. Rep. Scot.
Pl. Breed. Stn., 1979-80, pp. 100-8.
Davidson, W.D. (1934) History of potato varieties. J. Dept Rep. Ire., 33, 57-81.
De Ma,ine, M.J. (1982) An evaluation of the use of dihaploids and unreduced
gametes in breeding for quantitative resistance to pathogens. J. Agric. Sci.,
Camb., 99, 79-83.
De Maine, M.J. (1986) A method of testing the resistance of potato cultivars to
tuber damage caused by squeezing. J. Agric. Sci., Camb., 106, 255-8.
De Maine, M.J. (1988) Testing the tuber deformation resistance of tetraploid and
dihaploid potatoes. J. Agric. Sci., Camb., 110,445-9.
De Maine, M.J. and Caligari, P.D.S. (1988). Assessments of the resistance of
potato cultivars and breeders' clones to external damage. J. Agric. Sci., Camb.,
110, 527-30.
Dunwell, J.M. and Sunderland, N. (1973) Anther culture of Solanum tuberosum
L. Euphytica, 22, 317-23.
Ellenby, C. (1948) Resistance to the potato root eelworm. Nature, 162, 704.
References 367
Ellenby, C. (1954) Tuber forming species and varieties of the genus Solanum tested
for resistance to the potato root eelworm Heterodera rostochiensis Woll.
Evans, D.A. (1983) Agricultural application of plant protoplast fusion. Biotech., 1,
253--6l.
Evans, D.A. (1989) Somaclonal variation - genetic basis and breeding applications.
Trends in Genet., 5, 46-50.
Fish, N. and Jones, M. (1988) Engineering somatic hybrid potatoes. Pl. Today, 1,
47-9.
Fish, N., Steele, S.H. and Jones, M.G.K. (1988) Field assessment of dihaploid
Solanum tuberosum and S. brevidens somatic hybrids. Theor. Appl. Genet., 76,
880-6.
Fitschen, H.J. (1984) Ziichtung neuer Kartoffelsorten. Kartoffelbau, 35,52-5.
Foroughi-Wehr, B., Wilson, H.M., Mix, G. and Gaul, H. (1977) Monohaploid
plants from anther culture of dihaploid genotypes of Solanum tuberosum L.
Forrest, J.M.S. and Holliday, J.M. (1979) Screening for quantitative resistance to
the white potato cyst nematode Globodera pallida. Ann. Appl. Bioi., 91,371--4.
Fry, W.E. (1981) Management of late blight, in Advances in Potato Pest Manage-
ment (eds J.H. Lashomb and R. Casagrande), Hutchinson Ross, Pennsylvania,
USA, pp. 244--60.
Glendinning, D.R. (1975a) Neo-tuberosum: a new potato breeding mate.rial. 1. The
origin, composition and development of the Tuberosum and Neo-tuberosum
gene pools. Pot. Res., 18, 256-61.
Glendinning, D.R. (1975b) Neo-tuberosum: a new potato breeding material. 3.
Characteristics and variability of Neo-tuberosum, and its potential value in
breeding. Pot. Res., 18, 351--62.
Glendinning, D.R. (1983) Potato introductions and breeding up to the early 20th
century. New Phytol., 94, 479-505.
Haberlach, G.T., Cohen, B.A., Reichert, N.A., Baer, M.A., Towill, L.E. and
Helgeson, J.P. (1985) Isolation, culture and regeneration of protoplasts from
potato and several related Solanum species. PI. Sci. 39, 67-74.
Hermsen, J.G.Th. (1987) Efficient utilization of wild and primitive species in
potato breeding, in The Production of New Potato Varieties (eds G.J. Jellis and
Hermsen, J.G.Th. and Ramana, M.S. (1981) Haploidy and plant breeding, in The
manipulation of genetic systems in plant breeding. Phil. Trans. Roy. Soc.,
Lond., B. 292, 499-507.
Hoekema, A., Huisman, M.J., Molendijk, L., van den Elzen, P.J.M, and
Cornelissen, B.J.C. (1989) The genetic engineering of two commercial potato
cultivars for resistance to potato virus X. Biotech., 7, 273-8.
Holden, J.H.W. (1977) Potato breeding at Pentlandfield. An. Rep. Scot. Pl. Breed.
Stn., 1976-77, pp. 66-97.
Hougas, R.W. and Peloquin, S.J. (1957) A haploid plant of potato variety
Katahdin. Nature, 180, 1209-10.
Hougas, R.W. and Peloquin, S.J. (1958) The potential of potato haploids in
breeding and genetic research. Am. Pot. J., 35, 701-7.
Howard, H.W. (1963) Some potato breeding problems. Pl. Breed. Inst. Camb.
Ann. Rep., 1961--62, pp. 5-21.
Howard, H.W. (1978) The production of new varieties, in The Potato Crop, 1st edn
(ed. P.M. Harris), Chapman and Hall, London, pp. 607-46.
Hughes, J.e. (1980) Potatoes. 1. Factors affecting susceptibility to damage. Span,
23,65-7.
Jackson, M.J. (1987) Breeding strategies for true potato seed, in The Production of
Jinks, J.L. (1983) Biometrical genetics of heterosis, in Heterosis (ed. R. Frankel),
Springer-Verlag, Berlin, pp. 1-45.
Jinks, J.L. and Pooni, H.S. (1976) Predicting the properties of recombinant lines
derived from single seed descent. Heredity, 36, 253-66.
Jinks, J.L. and Pooni, H.S. (1986) Description and illustration of the practical
applications of biometrical genetics to plant breeding. Proc. of Eucarpia
(Biometrics Section), Birmingham, 1986, pp. 1-20.
Jones, M.G.K. (1987) Use of protoplast fusion and somaclonal variation in potato
breeding, in The Production of New Potato Varieties (eds G.J. Jellis and D.E.
Karp, A. and Bright, S.W.J. (1985) On the causes and origins of somaclonal
variation, in Oxford Surveys of Plant Molecular and Cell Biology, Vol. 2 (ed.
B.J. Miftin), Oxford University Press, Oxford, pp. 199-234.
Kehoe, H.W. (1982) Potato breeding. An Foras Talun. Res. Rep., 1982, 42-8.
Kichefski, D.F., Quinn, A.A. and Peloquin, S.J. (1976) The e.ffectiveness of
selection during early clonal generations in varietal breeding. Am. Pot. J., 53,
370-1.
Killick, R.J. and Malcolmson, J .F. (1973) Inheritance in potatoes of field resistance
to late blight (Phytophthora infestans (Mont.) de Bary). Physiol. Pl. Path., 3,
121-31.
Knight, T.A. (1807) On raising of new and early varieties of potato (Solanum
tuberosum). Trans. Hort. Soc. Land., 1,57-9.
Krantz, F.A. (1938) Maturity of potato seedlings in the glasshouse and their later
behaviour in the field. Am. Pot. J., 15, 153-7.
Krantz, F.A. (1946) Potato breeding methods. III. A suggested procedure for
potato breeding. Tech. Bull. Univ. Minn., 173, 24pp.
Lacey, C.N.D., Jellis, G.J., Currell, S.B. and Starling, N.C. (1987) An
early generation screen for combined cyst nematode (Globodera spp.) and
blight (Phytophthora infestans) resistance in potatoes. Pot. Res., 30,
59-69.
Larkin, P.J. and Scowcroft, W.R. (1981) Somaclonal variation - a novel source of
variability from cell cultures for plant improvement. Theor. Appl. Genet., 60,
197-214.
Lee, H.C. (1985) Breeding new export potato varieties. Agric. North. Irel., 60 (6),
198-201.
Louwes, K.M. and Neeie, A.E.F. (1987) Selection for chip quality and specific
gravity of potato clones: possibilities for early generation selection. Pot. Res.,
30,241-51.
McRae, D.e. and Fleming, J. (1989) Potato Damage - Where it occurs and how to
avoid it. Potato Market. Board, Oxford, 28pp.
MacArthur, A.W. (1970) Fifty years of potato breeding at the Scottish Plant
Breeding Station. J. Nat. Assoc. Seed Pot. Merch., 10 (3), 1-12.
References 369
Mackay, G.R. (1982) Breeding for resistance to pests and diseases. Scot. Crop Res.
Inst. Bull., 1, 27-35.
Mackay, G.R. (1987) Selecting and breeding for better potato cultivars, in
Improving Vegetatively Propagated Crops (eds A.J. Abbott and R.K. Aitken),
Academic Press, London, pp. 181-96.
Mackay, G.R., Caligari, P.D.S., Dale, M.F.B., Brown, J., Torrance, C.J.W.,
Swan, G.E.L. and Spence, J.S. (1985) Low temperature storage for crisping
cultivars. Ann. Rep. Scot. Crop Res. Inst., 1984, 69pp.
Mackay, G.R., Caligari, P.D.S., Dale, M.F.B., Brown, J., Torrance, C.J.W.,
Swan, G.E.L. and Spence, J.S. (1986) Early generation selection. Ann. Rep.
Scot. Crop Res. Inst., 1985, 63pp.
Mai, W.F. and Peterson, L.C. (1952) Resistance of Solanum ballsii and Solanum
sucrense to the golden nematode, Heterodera rostochiensis Woli. Science, 116,
224-5.
Malcolmson, J.F. (1976) Assessment of field resistance to blight (Phytophthora
infestans) in potatoes. Trans. Brit. Mycol. Soc., 67,321-5.
Maris, B. (1964) Studies concerning the relationship between plant height of
potatoes in the seedling year and maturity in the clonal generations. Euphytica,
13,130--8.
Maris, B. (1966) The modifiability of characters important in potato breeding.
Maris, B. (1969) Studies on maturity, yield, under water weight and some other
characters of potato progenies. Euphytica, 18,297-319.
Maris, B. (1986) The effect of seed tuber weight on characters in the first and
second clonal generation of potato populations. Euphytica, 35, 465-82.
Mendoza, H.A. (1980) Preliminary results on yield and uniformity of potatoes
grown from true seed, in Report of a Planning Conference on 'Production of
Potatoes grown from True Seed', Manilla, Philippines, 1979, pp. 156-72.
Mendoza, H.A. (1987) Advances in population breeding and its potential impact
on the efficiency of breeding potatoes for developing countries, in Production of
Munzert, M. (1987) Potato breeding strategy in the Federal Republic of Germany,
in The Production of New Potato Varieties (eds G.J. Jellis and D.E. Richardson),
Nachmias, A., Caligari, P.D.S., Mackay, G.R. and Livescu, L. (1988) The effects
of Alternaria solani and Verticillium dahliae on potatoes growing in Israel. Pot.
Res., 31, 443-50.
Niopek, J. (1967) Bewertung und auslese der kartoffel-samlingsklone nach dem
gebrauchswert der knollenform. Kartoffelbrau, 18, 344-8.
Ooms, G. (1987) Genetic manipulation in potato using Agrobacterium, in The
Production of New Potato Varieties (eds G.J. Jellis and D.E. Richardson),
Patterson, C.F. (1953) A method of obtaining fruits in the potato variety Russet
Burbank. Am. Pot. J., 30, 89-91.
Peloquin, S.J. and Hougas, R.W. (1959) Decapitation and genetic markers as
related to haploidy in Solanum tuberosum. Eur. Pot. J., 2, 176-83.
Pfeffer, C. (1963) Vergleichende untersuchungen uber auslesmoglichkeiten von im
freiland und in topfen kultivierten kartoffelsamlingen. Ziichter, 33, 6-11.
Phillips, M.S. (1981) A method of assessing potato seedling progenies for
resistance to the white potato cyst nematode. Pot. Res., 24, 101-3.
Phillips, M.S. and Dale, M.F.B. (1982) Assessing potato seedling progenies for
resistance to the white potato cyst nematode. J. Agric. Sci., Camb., 99,67-70.
Phillips, M.S., Forrest, J.M.S. and Wilson, L.A. (1980) Screening for resistance to
potato cyst nematode using closed containers. Ann. Appl. Bioi., 96, 317-22.
Plaisted, R.L. (1972) Utilization of germ plasm in breeding programs, use of
cultivated tetraploids, in Prospects for the Potato in the Developing World, CIP,
Lima, Peru, pp. 90-9.
Plaisted, R.L., Thurston, H.D., Brodie, B.B. and Hoopes, R.W. (1984) Selecting
for resistance to diseases in early generations. Am. Pot. J., 61, 395--403.
Pooni, H.S. and Jinks, J.L. (1978) Predicting the properties of recombinant
inbred lines derived from single seed descent for two or more characters
simultaneously. Heredity, 40, 349--61.
Powell, W. and Caligari, P.D.S. (1989) The use of hormonal and osmotic growth
retardants for the storage of potato germplasm in-vitro. Pot. Res., 32,57--64.
Powell, W., Brown, J. and Caligari, P.D.S. (1989) The field performance of potato
cultivars derived from micropropagated donor plants. Ann. Appl. Bioi., 115,
123-8.
Richardson, D.E. (1987) Potato variety assessment in the UK, in The Production
of New Potato Varieties (eds G.J. Jellis and D.E. Richardson), Cambridge
Ross, H. (1986) Potato Breeding - Problems and perspectives, Parey, Hamburg,
132pp.
Rothacker, D. (1957) Untersuchungen iiber den einfluss von kreuzungs-zeitpunkt
und pfropfung aug den bastard-samenertrag bei kreuzungen zwischen
knollentragenden Solanum-arten. Ziichter, 27,232-8.
Salaman, R.N. (1949) The History and Social Influence of the Potato, Cambridge
University Press, Cambridge.
Scheider, 0., Hein, T. and Kohn, H. (1985) Plant cell fusion as a tool for genetic
manipulation, in Genetic Manipulation in Plant Breeding (eds W. Horn, C.J.
Jensen, W. Odenbach and O. Scheider), W. de Guyter, Berlin, pp. 641-51.
Scholtz, M. (1987) Potato breeding strategy in the German Democratic Republic,
in The Production of New Potato Varieties (eds G.J. JeUis and D.E. Richardson),
Shepard, J.F., Bidney, D. and Shahin, E. (1980) Potato protoplasts in crop
improvement. Science, 208, 17-24.
Simmonds, N.W. (1963) Experiments on the germination of potato seed. I and II.
Eur. Pot. J., 6, 45--60, 69-76.
Simmonds, N.W. (1966) Studies of the tetraploid potatoes. III. Progress in the
experimental recreation of the Tuberosum Group. J. Linn. Soc. (Botany), 59,
279-88.
Soloman, R.M., Brown, J. and Mackay, G.R. (1987) Progeny testing for resistance
to potato leaf roll virus. Pot. Res., 30, 161.
Sopory, S.K., Jacobsen, E. and Wenzel, G. (1978) Production of monohaploid
embryoids and plantlets in cultured anthers of Solanum tuberosum. Pl. Sci.
Lett., 12, 47-54.
Steineck, O. (1957) Photoperiodismus und kartoffelziichtung. Bodenkultur, 9,
263-74.
References 371
Stone, A.R. (1972) Heterodera pallida N. sp. (Nematoda: Heteroderidae), a

second species of potato cyst nematode. Nematolog., 18, 591-606.
Stuart, W. (1937) The Potato: Its culture, uses, history and classification, J.B.
Lippincott, Chicago.
Swaminathan, M.S. and Howard, H.W. (1953) The cytology and genetics of the
potato (Solanum tuberosum) and related species. Bibliog. Genet. (reprinted by
Martinus Nijhoff, The Hague), pp. 1-192.
Swiezynski, K.M. (1978) Selection of individual tubers in a potato breeding. Theor.
Appl. Genet., 53, 71-80.
Swiezynski, K.M. (1984) Early generation selection methods used in Polish potato
breeding. Am. Pot. J., 61, 385-94.
Tai, G.C.C. (1975) Effectiveness of visual selection for early clonal generation
seedlings of potato. Crop Sci., 15, 15-18.
Tai, G.C.C. and Young, D.A. (1984) Early generation selection for important
agronomic characters in a potato breeding population. Am. Pot. J., 61,419-34.
Tellheim, E. (1975) Untersuchungen zur friihdiagnose aug knollenertrag und
qualitiitsmerkmale in der kartoffelziichtung. Arch. Zuchtungsforschg., 5,
165-73.
Thijn, G.A. (1954) Observations of flower induction with potatoes. Euphytica, 3,
28-34.
Thompson, A.J. (1987) The potential value of somaclonal variants in potato
improvement, in The Production of New Potato Varieties (eds G.J. Jellis and
Toxopeus, H.J. (1964) Treasure digging for blight resistance in potatoes..
Uhrig, H. (1985) Genetic selection and liquid medium conditions improve the yield
of androgenetic plants from diploid potatoes. Theor. Appl. Genet., 71,455-60.
Uhrig, H. and Salamini, F. (1987) Dihaploid plant production from 4X-genotypes
of potato by the use of efficient anther culture plants producing tetraploid
strains (4XEAPP-clones) - proposal of a breeding methodology. Pl. Breed., 98,
228-35.
Uijtewaal, B.A., Huigen, D.J. and Hermsen, J.G.Th. (1987a) Production of
potato monohaploids (2n=x=12) through prickle pollination. Theor. Appl.
Genet., 73, 751-8.
Uijtewaal, B.A., Jacobsen, E. and Hermsen, J.G.Th. (1987b) Morphology and
vigour of mono haploid potato clones, their corresponding homozygous diploids
and their tetraploids and their heterozygous diploid parent. Euphytica, 36, 745-53.
Wastie, R.L., Caligari, P.D.S., Stewart, H.E. and Mackay, G.R. (1987) A
glasshouse progeny test for resistance to tuber blight (Phytophthora infestans).
Pot. Res., 30, 533-8.
Wastie, R.L., Caligari, P.D.S., Stewart, H.E. and Mackay, G.R. (1988a)
Assessing the resistance to gangrene of potato (Solanum tuberosum) progenies
from parents differing in susceptibility. Pot. Res., 31, 355-65.
Wastie, R.L., Caligari, P.D.S. and Wale, S.J. (1988b) Assessing the resistance of
potatoes to powdery scab (Spongospora subterranea (Wallr.) Lagerh.). Pot.
Res., 31, 167-71.
Wiersema, H.T. (1950) Groeistoffen en plantenveredeling naar aanleiding van een
onderzoek bij aardappels. Studiekring voor Plantenveredeling, Wageningen,
pp. 338-41.
Wiersema, S.G. (1983) Potato seed-tuber production from true seed, in Research
for the Potato in the Year 2000 (ed. W.J. Hooker), CIP, Lima, pp. 95-116.
Withers, L.A. (1986) In vitro approaches to the conservation of plant genetic
resources, in Plant Tissue Culture and Its Agricultural Applications (eds L.A.
Withers and P.G. Alderson), Butterworths, London, pp. 261-76.
Woude, K. van der (1987) Variety assessment in the Netherlands, in The
Production of New Potato Varieties (eds G.J. Jellis and D.E. Richardson),
CHAPTER 9
Weed control
P.J. W. Lutman
9.1 WEEDS IN POTATOES
9.1.1 The potato crop in the rotation

When first introduced as a food crop to Great Britain, the potato was
grown mainly for local consumption rather than as a saleable cash crop, as
it is today. The need to grow potatoes on a more extensive basis only
emerged following the industrial revolution, when cheap food sources were
required to feed rapidly expanding urban populations. The Norfolk four-
course rotation, developed in the nineteenth century, based on roots,
barley, seeds (e.g. clover) and wheat, provided an opportunity for the
production of potatoes as the 'roots' component of the rotation, although
initially potatoes formed only a small proportion of the roots grown.
During the early part of the twentieth century this basic rotation was
modified, particularly in arable areas where the 'seeds' component was
replaced by another cereal or root crop.
In 1963 the survey conducted by the Potato Marketing Board (1963)
showed that the majority of potato crops were sown after cereals, parti-
cularly in the east of England where 70-80% followed cereals. In the west,
more than 25% of the crops followed a ley or permanent grass. This
distribution of preceding crops changed little in the following 15 years
(Potato Marketing Board, 1979). However, during this period the number
of producers declined appreciably and although the national area planted
also declined, the average area sown by each producer increased. This
trend has continued in the 1980s with potatoes being grown by fewer
specialists, mainly on more easily managed lighter soils. In 1955, when
there were 90000 producers, each grew on average 3.3 ha; in 1987, 21400
producers each grew on average 7.8 ha (Potato Marketing Board, 1963,
1987). This average shows the increase in area grown by each producer but
does not show the increase in the percentage of the national crop produced
by the large growers. In 1963 producers growing over 20 ha grew 42% of
Published in 1992 by Chapman & Hall, London. ISBN 0 412296403
374 Weed control
the total area but by 1987 this had increased to 52%. Consequently,
potatoes feature as an important part of the rotation of an increasing
number of specialist growers, who often also grow other root and vegetable
crops. These changes in cultural patterns have influenced the weed flora,
their potential effects on the crop and the importance of the weed
problems caused by the potatoes left in the field after the potato harvest.
These effects will be discussed in more detail later in the chapter.
9.1.2 The potato as a cleaning crop

One function of crop rotation in farming has always been that of control-
ling weeds. Before the advent of herbicides, cereals permitted weeds to
proliferate. Hoeing was impractical because of the closeness of the cereal
rows and, consequently, weeds brought about serious loss of yield. In
contrast, it was possible to control weeds in the 'roots' part of the rotation,
as these crops were grown in wide rows that enabled both hand and
mechanical cultivations to be carried out. Potatoes had other advantages;
the need for a fine deep seedbed at planting and for re-ridging after
planting gave a number of opportunities to control weeds and so increased
the value of potatoes as a cleaning crop. In addition, the crop's vigorous
growth and dense canopy, once established, enabled it to suppress many of
the late emerging weeds. However, with the universal use of herbicides to
control weeds in cereals and other crops, the value of potatoes as a
cleaning crop is now much diminished. Indeed, some growers now use the
previous crop to minimize weed infestations in the potatoes. For example,
it is much better to control cleavers (Galium aparine) in the cereal crop
prior to potatoes, because of the ineffectiveness of most of the potato
herbicides on this weed.
9.1.3 Adverse effects of weeds
(a) Competition
Weeds compete with the potato crop for light, nutrients and water. Yields
of tubers can be severely reduced, the size of the reduction depending on
the density and competitive ability of the weed species present and the
availability of light, nutrients and water. A series of trials in the early 1960s
Neild and Proctor (1962) showed that, on average, weeds reduced tuber
yields by 36%. In the trial with the densest weed infestation (900-1200
weeds m- 2 ) yields were reduced by 80%, whilst less extreme weed
populations of just over 100 weeds m- 2 reduced yields by 14%. Similarly,
Nelson and Thoreson (1981) found that weeds emerging with the crop (58-
311 weeds m -2) reduced yields by up to 54%. Although there is a lot of
simple information available from herbicide evaluation trials on the effects
of weeds on the growth of potatoes in non-herbicide treated plots, there is
Weeds in potatoes 375
very little detailed information on how weeds affect the growth and yield
of potatoes. Saghir and Markoulis (1974) reported the results of an
experiment on critical periods for competition (Nieto et al., 1968), where
plots were either left weedy for increasing periods before removal, or were
kept weed free for increasing periods. This showed that weeds could be left
in potatoes for 6-9 weeks after planting and that they should then be kept
weed free for the next 3 weeks. Other experiments by Vitolo and Ilnicki
(1985) demonstrated that grass weeds could be left in potatoes for 6-8
weeks after planting but must be controlled thereafter. Similarly Thackral
et al. (1989) have shown that weed control in potatoes could be delayed for
4--6 weeks after planting. In both these latter examples one weed removal
at the appropriate time was adequate to prevent loss of yield; later
emerging weeds were unimportant. This markedly reduced competitive
effect of later emerging weeds was also described by Nelson and Thoreson
(1981). As well as affecting overall yields, weed competition can influence
tuber quality. Competition may increase the percentage of smaller tubers,
and Saghir and Markoulis (1974) found that competition affected the
specific gravity of the tubers, thus altering their suitability for processing.
Experience in the UK is that the majority of weeds emerge just before or
as the potato shoots emerge. Obviously this will be dependent on adequate
availability of moisture, as very dry conditions will delay the germination
of weeds. These early emerging weeds are the most competitive. Later
emerging annual species, such as black nightshade (Solanum nigrum), or
perennials such as creeping thistle (Cirsium arvense) can penetrate the crop
canopy but, although they are difficult to control, they probably do not
have a great effect on potato yields.
(b) Other effects of weeds

In addition to affecting yields, weeds can interfere with harvesting. This
has become particularly important in recent years as the majority of potato
crops are now mechanically harvested. The presence of large amounts of
weed on the webs of the harvesters reduces the efficiency of soil separation
and slows the harvesting operation. Late emerging weeds, like those
mentioned in the previous paragraph and fat hen (Chenopodium album),
can produce large plants as the potato haulm collapses in late summer. The
use of a haulm desiccant will minimize the effects of these late growing
weeds (see Section 9.5.2). Rhizomes of perennial grass weeds will grow
through potato tubers, reducing the quality of the crop. Weeds can also be
alternate hosts for some potato pests and diseases. Aphid vectors of potato
virus diseases can live on a wide range of weed species and research in
Germany has shown that the presence of weeds increased infection of
potato crops with black scurf (Rhizoctonia solani) (Griesbach and Eisbein,
1975).
376 Weed control
9.2 WEED BIOLOGY
The time of planting of the potatoes, the timing, type and frequency of
cultivations, soil type, weather conditions and the types of crops grown in
the rotation with the potatoes can all have a profound effect on the weed
flora. If winter cereals predominate in the rotation, the weed flora will be
biased in favour of cereal weeds. Thus, annual grass weeds will be
common. If the rotation includes a high percentage of vegetable and root
crops, broad-leaved weeds will predominate. Cultivations will stimulate
the emergence of weed seedlings but only those species with the appro-
priate innate germination pattern will emerge (Roberts et al., 1982). Thus,
as the majority of potatoes are planted in April, weed species that have an
emergence peak in the spring will be the most common.
9.2.1 Perennial weeds
(a) Grass weeds

Common couch grass (Elymus (Agropyron) repens) and black bent (Agrostis
gigantia) can be very aggressive weeds. They produce rhizomes or under-
ground stems which spread the plants rapidly within fields. Normally, the
majority of buds on these rhizomes are dormant but cultivation will break
up the rhizome system and many buds will be released from dormancy and
sprout to make new plants, thus increasing the distribution and frequency
of these weeds. However, repeated cultivations can have a beneficial
effect, by exhausting the food reserves of the rhizomes, through prevention
of the establishment of new plants. Both weeds are favoured by minimum
tillage, so if potatoes are grown after a series of direct drilled, or minimally
cultivated, cereal crops they can pose serious problems. Prior to the
development of glyphosate their control was extremely difficult, even in
cereal crops. Even now their control within the potato crop is not easy (see
Section 9.4) and is best achieved in the previous crop, or in the autumn
prior to the potatoes. Cultivations carried out prior to and after the
planting of potatoes can help to break up rhizome clumps and slow down
the growth of the plants, giving the potato haulm time to close over the
rows and smother the weeds.
Other perennial grass weeds (e.g. creeping bent - Agrostis stoloni/era)
occur in potatoes but are not as widespread as common couch and black
bent. Common reed (Phragmites communis) is found on the edges of
potato fields in low-lying areas, as in the fenlands, where it grows out into
the fields from ditch banks.
(b) Broad-leaved weeds

Several perennial broa.d-Ieaved weeds cause problems in potatoes, the
commonest being thistles, particularly creeping thistle (C.arvense) and
Weed biology 377
perennial sowthistle (Sonchus arvensis). Docks (Rumex spp.) were
formerly common perennial weeds but the increase in the number of
specialist potato growers and the decline in the number growing potatoes
in mixed rotations with grass leys has decreased the frequency in these
weeds. Field bindweed (Convolvulus arvensis) and amphibious bistort
(Polygonum amphibium) can also cause problems, the latter being
frequently found on the fenland soils of East Anglia. All these weeds, with
the exception of thistles, are now less common than in the recent past. The
widespread use of glyph os ate against perennial weeds in cereals has also
reduced the frequency of these weeds in potatoes.
The maintenance or increase in creeping thistle populations indicates
that glyphosate may not be as effective on this species as it is on other
perennial weeds. It spreads by brittle roots that break up readily on
disturbance and produces fresh plants from adventitious buds. Thus,
intensive cultivations prior to planting potatoes will spread existing pa.tches
and increase plant density appreciably. In undisturbed soil large quantities
of roots can remain hidden for many years, with only the occasional above-
ground shoot. Although large numbers of seeds (thistle-down) are pro-
duced, few of the seeds are viable and so perennation of the plant is almost
solely dependent on the roots. Seed production is more important in the
distribution of perennial sowthistle.
Field bindweed, although less common than creeping thistle, is also
mainly distributed from fragments of broken root. Similarly, it is very deep
rooted and is difficult to destroy by cultivation. Its scrambling habit results
in the production of mats of foliage that can interfere with harvesting,
making it a serious weed where it does occur.
Another perennial broad-leaved weed to cause problems in potatoes is
the potato itself. The occurrence of volunteer potatoes from the preceding
crop poses a risk to the health of the new crop and may prevent the crop
being sold for seed. This 'weed' problem is discussed in more detail later in
the chapter.
Although it is possible to minimize the effects of perennial grass weeds in
potato crops by using a graminicide (see Section 9.4), it is much more
difficult, or even impossible, to suppress perennial broad-leaved weeds
with herbicides, once the potato crop has emerged. As with the perennial
grass weeds, it is better to minimize their presence prior to the potato crop
by using appropriate control measures earlier in the rotation.
9.2.2 Annual weeds

The commonest annual weeds to occur in potatoes are either those that
have a very wide period of emergence, such as common chickweed
(Stella ria media) or mayweeds (Matricaria recutita, Matricaria matri-
carioides, Tripleurospermum inodorum), or those with an emergence peak
in the spring to early summer period like knotgrass (Polygonum aviculare)
378 Weed control
and fat hen (c. album). Autumn emerging weeds, such as ivy-leaved
speedwell (Veronica hederifolia) , do not usually occur in potatoes.
However, cleavers (G. aparine), a mainly autumn germinating species, is
becoming increasingly common in potatoes, suggesting that the emergence
pattern of this weed is changing. Similarly, the common autumn germinat-
ing winter cereal weed, black-grass (Alopecurus myosuroides) , can also
occur in potatoes, particularly after wet, cool autumns when dormancy is
enforced on this weed, so increasing spring germination. Weed species that
require warm temperatures to germinate (e.g. black nightshade, S.
nigrum) pose particular problems. Except in late sown crops, they do not
germinate prior to the emergence of the potatoes and so are often not
well controlled by standard pre-emergence herbicide treatments. Once
established, they grow vigorously and can cause problems at harvest.
Although the composition of the annual weed flora is controlled by the
inherent characteristics of weed seeds present in the soil, the number of
weeds emerging in each potato crop will be influenced by several factors.
The timing, type and frequency of cultivation, fertilizer use, soil type,
preceding cropping and weather conditions are all important. Cultivations
encourage the germination of weed seeds, as the change in the micro-
climate around the seeds caused by the cultivator, coupled with physical
abrasion of the seed coat, breaks dormancy and stimulates germination. So
the timing of these cultivations can influence the weed flora. If the deep
cultivations employed prior to potatoes follow a series of minimally tilled
cereal crops, weed species with a longer dormancy, such as poppy (Papaver
rhoeas) will be favoured over species with a shorter dormancy. Conversely,
if intensive cultivations have been practised in previous crops, as when
potatoes are grown in rotation with vegetable crops, the weed species
favoured will be more ephemeral, such as groundsel (Senecio vulgaris).
Heavy soils favour some species (e.g. cleavers), whilst light soils favour
others (e.g. corn marigold - Chrysanthemum segetum).
Thus, a wide range of annual broad-leaved weed species can occur in
potato crops. Their relative importance will vary from field to field and
from year to year. Nationally, the following species seem to be the most
important: charlock (Sinapis arvensis), wild radish (Rap han us raphanistrum) ,
field pennycress (Thlaspi arvense), mayweeds, black bindweed (Fal/opia
convolvulus), knotgrass, redshank (Polygonum persicaria) , fat hen, orache
(Atriplex patula) , common chickweed, common fumitory (Fumaria
officianalis) , cleavers, black nightshade and volunteer oilseed rape.
Although broad-leaved weeds are the predominating annual weed species,
some annual grass weeds do occur, the commonest being annual meadow-
grass (Poa annua), wild oats (Avena fatua) and volunteer cereals. These
are not excessively competitive and so will only reduce yields if present in
large numbers.
Weed control 379
9.3 WEED CONTROL
9.3.1 Cultural weed control

Traditionally, weed control in potatoes was carried out by cultivation,
supplemented by hand labour. The land was cultivated prior to planting
and then recultivated several times thereafter to control weeds until the
density of the potato foliage was adequate to suppress weed growth.
Normally, the potato ridges were harrowed down and then re-ridged two
or three times. Although this system enabled the farmer to achieve high
levels of weed control there were a number of disadvantages. Each pass of
the cultivator disturbed the soil, brought clods of soil to the surface and
stimulated a further flush of weed seeds to germinate. The soil disturbance
also damaged the roots of the potato plants and increased water loss from
the soil. When more mechanized and less labour intensive husbandry
operations were introduced, cultivation remained an important method for
controlling weeds, but the nature of the harrows and cultivators changed.
Harrowing tended to be lighter to minimize clod production, and specialist
helical harrows which fitted the shape of the ridge were introduced from
The Netherlands. These harrows controlled weeds and maintained a fine
tilth more suited to the performance of modem complete harvesters. Jarvis
and Shotton (1972) clearly showed that the traditional cultivation systems,
although giving excellent weed control, had a greater adverse effect on
saleable ware tubers than did a reduced system of cultivation based on
Dutch methods.
As long ago as 1941, Pereira compared hand weeding with mechanical
weed control and showed that the latter reduced yields by about 8%. It was
only with the use of herbicides for weed control in potatoes in the early
1960s that it became possible to study, in detail, the effects on the crop of
post planting cultivations. Bremner (1966) in reviewing the results of a
series of experiments comparing herbicidal and mechanical weed control,
concluded that cultivation reduced yield by about 5%. Similarly, Elliott
(1965) showed that potato yields following the application of paraquat
were 4 t ha- 1 higher than those following cultivation. However, the precise
effect of cultivation on yield will depend on the nature, frequency and
timing of the operations, the type of potato crop, weather conditions, soil
type and moisture level. For example, early potatoes are likely to be more
adversely affected by root pruning from cultivations than are maincrops,
because of the former's shorter growing season and more vigorous early
root growth. It must also be remembered that the compaction caused by
the repeated passage of tractor wheels can itself reduce crop growth and
yield.
Some form of post-planting cultivation may be necessary to create a
good tilth for the production of ridges of the required size and shape, even
if weeds are to be controlled by herbicides. However, most of the available
380 Weed control
information shows that repeated cultivations reduce yields. The trend
towards increasing row widths from 76 cm to 91 cm has made post-planting
cultivation less necessary, as there is enough tilth between the wider rows
for the final ridge to be made at planting, without further cultivation.
9.3.2 Weed control with herbicides

Herbicides were first introduced for weed control in potatoes at the end of
the 1950s. The Potato Marketing Board (PMB) recorded in 1963 that
approximately 1% of potato crops were treated with diquat and a further
1% with either dinoseb or MCPA (Hampson and Taylor, 1970). No
herbicides had been recorded in their previous survey in 1958. Over the
following 20 years herbicide use increased dramatically, so that by 1980
the Potato Marketing Board survey showed 69% of crops received
herbicides (Taylor, 1976, 1980) (Fig. 9.1). This figure for 1980 conflicts
with that produced independently by the British Agrochemical Association
(BAA) which claimed that herbicides were applied to 174 000 ha of the
189 000 ha crop area (92%). This inconsistency probably relates to the
difference in assessment methods as BAA recorded all herbicides used and
the PMB recorded fields treated. So if a field received two herbicide
treatments it would be counted twice by BAA and once by PMB. It would
appear from the figures that approximately 30% of crops received no
herbicides and those treated received 1.3 treatments. Most recent figures
from the arable crops survey in 1988 by Davis et al. (1990) show that 92%
of potato crops received a herbicide and each field received 1.5 products.
Thus, weed control in a small proportion of crops is still done by cultivation
alone. Some of these crops are grown on small areas, where herbicide
application is difficult or uneconomic, or for the organic food market.
Others are grown on organic soils where herbicide activity is much reduced
and herbicidal weed control can be expensive.
Many experiments were carried out in the 1960s to compare the growth
and yields of potatoes following weed control with herbicides and by
cultivation. In general, weed control with herbicides was shown to increase
yields over weed control by cultivation (see Section 9.3.1). However, these
comparisons were sometimes confounded by differences in weed control
between the different treatments and by differing intensities of cultivation.
Although herbicides seemed to give superior yields to cultivations, there
were adverse effects from some herbicide treatments. For example, late
applications of pre-emergence treatments, such as paraquat, will damage
emerging potato shoots and can reduce yields (Ivany, 1974).
The first herbicides to be recommended were primarily those used in
other crops whose recommendations were extended to include potatoes
(e.g. TCA, MCPA). The use of herbicides increased rapidly following the
development of products that were more truly selective. The majority of
these herbicides have to be applied before the emergence of potato shoots.
Weed control 381
100
80
60
%
40
20
o*-~~~ __ L-~~-L~~L-~~-L~~W
58 60 62 64 66 68 70 72 74 76 78 80 82 84 86
Year
Figure 9.1 Change in the percentage of potato crops treated with herbicide in
Great Britain 1958-86. * Potato Marketing Board: % of cropped area treated. 0
British Agrochemicals Association: % hectares treated.
They are either applied as contact, non-selective treatments to control

emerged weed seedlings prior to potato emergence (e.g. paraquat), or as
selective, persistent, soil-applied treatments to prevent the emergence of
newly germinated weed seedlings (e.g. linuron). These two types are
frequently used together, so that weeds present at spraying and those
that would have emerged thereafter are controlled. The introduction of
metribuzin in 1973 (see Mannall et al., 1972) was a considerable step
forward, as this could be used before or after the emergence of many
potato cultivars, thus increasing flexibility in the timing of application. This
greater flexibility is reflected in the widespread usage of this product.
Unfortunately, it cannot be used after the potatoes exceed 150 mm in
height, so the crop still lacks a fully selective, broad spectrum, post-
emergence herbicide for the control of broad-leaved weeds. However,
the development of the selective grass weed herbicides alloxydim and
sethoxydim for broad-leaved crops, in the early 1980s, has resulted in
recommendations for the post-emergence control of A. fatua and other
grasses in potatoes (Ingram et al., 1978; Formigoni et al., 1979).
9.3.3 The interaction between herbicides and cultivations

Weed control with herbicides or by cultivation alone, will not achieve
optimum weed control every time. The soil conditions may be too dry for
382 Weed control
the residual herbicides to work effectively, or the weed species present may
not be sensitive to the herbicide products available. Cultivations may be
ineffective in wet conditions, simply transplanting rather than killing the
hoed weeds. Often a combination of the two will give the best results. It
must be remembered that the ridge should not be recultivated once a
residual herbicide has been applied, as this will destroy the uniform
herbicide layer, at or near the soil surface, that is vital to ensure good
activity on subsequently germinating weeds. As residual herbicides work
best if applied to a finely cultivated soil and if the potatoes are grown in low
well rounded ridges, it is important that pre- and post-planting cultivations
create these conditions. Cultivations that create cloddy ridges will reduce
the performance of residual herbicides, by preventing the herbicide
achieving an even cover of the soil surface. Cultivation immediately before
herbicide application can have a positive effect, eliminating the need for a
contact herbicide such as paraquat.
9.4 RECOMMENDATIONS FOR THE USE OF HERBICIDES
9.4.1 General
The use of herbicides III UK crops is controlled by the Food and
Environmental Protection Act and by the Control of Substances
Hazardous to Health. This new legislation makes 'approval' a legal
requirement and it is an offence to sell, store, advertise, or use unapproved
pesticides. All products must be 'approved' under the Control of Pesticides
Regulations. The approved label for each product contains all the informa-
tion required for the user to use the product legally, safely and effectively.
Consequently, the label must be read thoroughly before use.
Herbicides may act on the weeds after entry through the roots, shoots or
foliage, or any combination of these. They may be applied before the crop
is planted, after planting but before it emerges, or after crop emergence.
Selectivity may depend on separation in time and/or space, or on physio-
logical differences between weeds and the crop. Four main types of
herbicides are used for weed control in potatoes.
1. Pre-planting treatments in which the herbicide is applied before the

potatoes are planted. Selectivity depends on the disappearance of any
herbicide residues before planting. The herbicides may be active
through the foliage and/or roots.
2. Contact pre-emergence treatments in which the herbicide is applied, after
the crop has been planted but before it has emerged, to the foliage of
emerged weed seedlings. They are non-selective, depending for their
selectivity on the absence of emerged potato shoots and on their lack of
activity through the soil.
Recommendations for the use of herbicides 383
3. Soil-acting pre-emergence treatments in which the herbicide is applied
after planting but before both potatoes and weeds have emerged. They
are absorbed by the roots or emerging shoots of sensitive weed species
and remain active for several weeks or months after application.
Selectivity may be due to physiological differences in sensitivity
between potatoes and weeds, or to depth protection, where the shallow
rooted weeds are killed but the deeper-sown potatoes are not.
4. Post-emergence treatments in which the herbicide is applied after both
the crop and weeds have emerged. Activity may be through the foliage
or soil. Selectivity depends on physiological differences between
potatoes and the weeds to be controlled.
In general, herbicides are applied to weeds in potatoes with conventional

hydraulic sprayers, using hollow cone or flat fan jets. The size of the orifice
of the jet, the forward speed and the pressure will influence the droplet
spectrum and the volume rate applied. Small weeds will be controlled more
effectively with foliage-acting herbicides if the droplets are 'fine' but
'coarse', less drift sensitive sprays are acceptable for applications to the
. soil surface prior to weed emergence. If bare soil or small weeds are to be
treated, a low volume of approximately 100 I ha- 1 will be adequate.
However, once the crop or weeds become larger a higher volume rate may
be more appropriate, to increase spray penetration through the canopy. As
the majority of potato herbicides are applied to the soil surface, the quality
and volume of the spray solution will not be very critical, provided an even
cover of the soil surface is achieved. Before deciding on which jet or
volume rate to use, it is important to read the instructions on the herbicide
label.
When choosing a herbicide treatment both the weed species that are to
be controlled and the risk of unwanted damage to the crop must be
considered. It is vital that the herbicide chosen will control the species
present or expected in the potato field. In addition, particularly for seed
crops and for earlies, it is important that the herbicide does not adversely
affect the potato crop. Seed crops may be rejected for certification, if, as a
result of herbicide treatment, the leaves exhibit abnormal symptoms that
could mask those caused by disease. For example, some herbicides can
cause transitory yellowing of the foliage, blotchy yellowing, or viral-like
leaf curling (Lawson and Wiseman, 1985a, b). Early potatoes are only
grown for a short period and there is insufficient time for them to recover
from early inhibitions to growth caused by inappropriate or mis-timed
herbicide treatments. Consequently, some herbicides are not recom-
mended for early potatoes and others should not be used on seed crops. In
addition, some herbicides are only recommended for use on some potato
cultivars. A further factor to consider is persistence, an important attribute
of soil-acting herbicides. Poor persistence will result in the emergence of a
flush of weeds before the canopy of potato haulm is sufficiently dense to
384 Weed control
suppress weed growth. Excessive persistence may also cause problems, as

the herbicide residues may become translocated into the daughter tubers,
may remain in the soil and affect the health of subsequent crops and may
contaminate groundwater. All these three factors are of considerable
concern to potato growers in the 1990s. Recent Ee and UK environmental
legislation, aimed at reducing pesticide usage in arable crops, is particu-
larly relevant to soil-acting herbicides and several of those used in potatoes
are currently being reviewed. Particular precautions with persistent products
should be taken in dry summers, when persistence tends to be greater. For
example, in such conditions it is essential to plough after the use of
metribuzin, before the planting of a crop of winter cereals.
9.4.2 Specific herbicide recommendations

This section includes details of all herbicides currently approved for use in
UK potato crops. Information is given on some of the more important
attributes of each product but it is not possible to include all the informaton
that is included on the label, which should be consulted before use.
Alterations, additions and deletions are continually being made to these
recommendations, so as a consequence some of the information in this
section will become inaccurate with the passage of time. All doses are
given as active ingredient and not as product, as for some herbicides a range
of products, with different formulations, are available. At the end of this
section reference is made to products that are currently being developed
but are not yet available.
(a) Pre-planting herbicides

These herbicides are most commonly used for the control of perennial
weeds and are applied either in the autumn or spring prior to the potato
crop.
Amitrole
As this herbicide is mainly taken up by the leaves of plants it is important
that the target weeds have produced adequate foliage at the time of
treatment and must be growing actively. It is translocated from the leaves
to the roots, rhizomes, stolons, etc., of perennial weeds. In the autumn
weeds must be at least 100 mm tall and in the spring at least 75 mm high.
Because amitrole persists in the soil, the land should be ploughed 3 weeks
after application. A dose of 4.5 kg ha- 1 is recommended.
Dalapon
Dalapon can be used to control annual and perennial grass weeds,
especially common couch (E. repens). As with amitrole, it is taken up
mainly by the leaves and translocated to roots and rhizomes. Thus, it is
important that the weeds have produced adequate foliage at the time of
treatment. It is recommended that the weeds should be 100-150 mm high
prior to treatment. In the autumn a dose of 14.4 kg ha- 1 should be used and
the land ploughed deeply 3-6 weeks after treatment. In the spring a lower
dose of 11.0 kg ha- 1 should be used and the land ploughed not less than
2 weeks after treatment. Potatoes can be planted immediately after
ploughing. Tubers of ed skinned varieties may suffer some loss of
pigment.
EPTC
EPTC is a soil-acting herbicide which enters the plant through the roots
and/or shoots. As it is volatile, it must be incorporated into the soil within
15 min of application. Although a number of grass and broad-leaved weeds
are susceptible, its main use is for the control of common couch. To
achieve best activity from the recommended dose of 4.54 kg ha- 1 , it should
be applied in the spring when temperatures have started to rise and the
common couch rhizomes have started to grow. Potatoes should be sown
within 2 weeks of application.
Glyphosate
This herbicide controls many annual and perennial weed species, especially
common couch and, when applied to the actively growing foliage, is
translocated through the leaves, stems, roots, rhizomes and other peren-
nating organs. It can be applied to foliage of the weeds at any time prior to
planting the potatoes, providing the weeds are growing actively, and may
even be used prior to the harvest of previous crops of cereals, oilseed rape,
field beans and peas. Allow at least 5 days to elapse after application, for
the herbicide to achieve optimum activity on perennial weeds, before
cultivation prior to planting potatoes. Only 1 day need elapse for the
control of annual weeds. When treating perennial weeds in the spring, the
plants should be allowed to produce at least 21 days new growth prior to
treatment. Doses of 0.36-1.44 kg ha- 1 may be used, the lower rates for
annual weeds, the higher ones for perennials.
TCA
TCA is a soil-acting herbicide that is taken up by plant roots and will
control a number of annual and perennial grass weeds. It can be used in
autumn or spring up to 8 weeks prior to planting potatoes. The herbicide
should be incorporated into the soil immediately after application to
improve activity. Doses from 16.1 to 31.3 kg ha- 1 are recommended. As
this is a persistent herbicide, cereals should not be sown until at least 16
weeks after application.
386 Weed control
(b) Contact pre-emergence herbicides
These herbicides are applied, after planting potatoes but normally before
all the shoots have emerged, to control emerged annual weed seedlings.
The soil should not be recultivated after planting as this will affect
the emergence of weed seedlings prior to treatment. Nor should it be
cultivated, for example by re-ridging after treatment, as this will stimulate
a further flush of weeds. Because of the absence of persistent effects from
these herbicides, they are frequently applied in mixtures with herbicides
that are primarily soil-acting and persistent, killing seedlings as they start
to germinate.
Paraquat and paraquat + diquat

These two herbicides are the constituents of the principal contact pre-
emergence products. Doses of 0.6 and 1.1 kg ha- 1 are recommended, the
higher for larger weeds, the lower for smaller ones. Neither has any activity
through the soil but both are rapidly absorbed by the foliage of treated
weeds, often killing them within 48 h. Speed of action is greatest in warm
conditions. As they are poorly translocated within the plant they are less
effective on perennial weeds. On early varieties of potato and crops grown
for seed, application can be delayed until 10% of the crop has emerged.
With main crop varieties application may be delayed until 40% has
emerged, provided no plants are bigger than 150 mm. Although emerged
shoots can be severely scorched, recovery is normally rapid and yields are
not affected. Crops growing from diseased or very small tubers, or in very
hot and dry conditions, should only be treated before they have emerged.
It must be remembered that although the recommendations state that
potatoes can be treated after they have emerged it is very easy to misjudge
the percentage crop emergence, thus jeopardizing yields. Headford (1968),
Ivany (1974) and Lawson et al. (1985) have all shown that applications at
100% emergence reduced haulm growth and sometimes reduced tuber
yields.
(c) Translocated pre-emergence herbicides

Dalapon
This is the only herbicide in this group. As well as being used pre-planting
(see above) it can be used at lower rates (2.89-4.67 kg ha- 1) before crop
emergence to control emerged grass weeds. As with the pre-planting treat-
ments, tubers of red skinned potato varieties may suffer a loss of colour.
(d) Contact and soil-acting pre-emergence herbicides

These herbicides, applied after planting but before crop emergence, have
some foliar activity and so kill some emerged seedlings. However, their
major role is to persist in the soil and kill weeds as they germinate. Several
of the products available are mixtures of two components, one with most
activity through the soil, the other with more foliar activity. Single
herbicides are often mixed with the contact herbicides paraquat and diquat
to improve control of larger emerged weeds. Most of the products are
members of the triazine or substituted urea groups of herbicides and, as
they are mainly active through the soil, their activity is influenced by soil
texture, organic matter and moisture. If these fa,ctors are not taken into
consideration poor weed control, or conversely crop damage, can result.
As these herbicides are more active in light soils than heavy ones
recommended doses are often adjusted to take account of soil type. Their
activity will also be lower if the soil is dry and/or cloddy. The soil surface
must not be disturbed after application.
Linuron
Linuron is probably the soil-acting herbicide that is most widely used in
potatoes and a large number of different formulations are available (13-
50% a.i.). It should be applied before the crop and weeds emerge,
particularly as some weed species are less sensitive after the seedling stage.
These include the important species annual meadow-grass (P. annua) ,
cleavers (G. aparine), corn marigold (c. segetum), groundsel (S. vulgaris),
knotgrass (P. aviculare), mayweeds (e.g. M. recutita) and speedwells (e.g.
Veronica persica). Common fumitory (F. officinalis) is resistant. The
difficulty of controlling emerged weeds can be overcome by mixing linuron
with a contact herbicide such as paraquat. If this is done some formulations
permit its use up to 20% crop emergence. The recommended doses vary
from 0.5 kg ha- 1 on very light soils to 2.2 kg ha- 1 on heavy and organic soils.
Although some products are recommended for use on sandy soils many are
not, because of the risk of causing crop damage. The performance of
linuron on organic soils is much poorer than it is on mineral soils and it only
controls small emerged seedling weeds. Performance on organic soils has
been improved by treating the weeds twice with a 50% dose of some
formulations, prior to crop emergence. Umbelliferous crops may be sown
at any time after the application of linuron but other crops should not be
sown within 2 or 3 months of treatment. Lettuce is particularly sensitive
and should not be sown in the same season.
Linuron + terbutryn
It is recommended that this product be used until 10% of the crop plants
have emerged. If emerged weeds are large it should be mixed with a
contact herbicide such as paraquat. A dose of 1.2 kg ha- 1 is recommiended
on very light soils growing early potatoes whilst up to 1.8 kg ha- 1 should be
used on heavy and organic soils. Although the product can be used on soils
with more than 10% organic matter, its residual life will be limited.
Linuron + terbutryn should not be used on sand soils. This mixture
388 Weed control
is more effective on common fumitory than linuron alone. As these
herbicides persist in the soil, no other crops should be planted for 3 months
after application. If a prolonged drought occurs this should be extended to
4 months. Lettuce should not be sown in the same season.
Metribuzin
Metribuzin is one of the most effective herbicides available for weed
control in potatoes (Mannall et al., 1972; Orson, 1986). First and most
second earlies should only be treated before they emerge but most
maincrop varieties and some second earlies can be treated pre- or post-
emergence (see p. 392). Some maincrop varieties, including Maris Piper.
should only be treated pre-emergence. Most weeds, including volunteer
oilseed rape, are sensitive to both pre- and post-emergence treatments but
black bindweed (F. convolvulus) and black nightshade are less sensitive if
treated pre-emergence, and field pansy (Viola orvensis) and knotgrass are
less sensitive if treated after they emerge. Black bindweed is particularly
difficult to kill but this herbicide will give better control of volunteer
oilseed rape than other potato herbicides. Doses vary from 0.52 to 1.05 kg
ha- 1 , according to soil type. On organic soils a pre-planting incorporated
application is recommended to improve activity, especially with the
variety Maris Piper, where a post-emergence treatment is not possible.
A programme of repeated low dose applications of metribuzin can be
as effective as a full single dose treatment (Askew and Flint, 1985).
Residues of metribuzin can cause damage to subsequent crops, and so it is
important that fields are ploughed to at least 150 mm after the potato
harvest, prior to sowing the next crop. Ryegrass, cereals and winter beans
may be sown in the same season, provided at least 16 weeks have elapsed
since treatment.
Monolinuron
Like linuron, this is a mainly soil-acting herbicide but it does have some
activity on seedling weeds. As the majority of weed species quickly become
resistant once emerged, it is frequently mixed with a contact herbicide such
as paraquat. Application of these mixtures can be delayed until 20%
emergence of maincrop potatoes and 10% emergence of earlies. It is
absorbed through the roots and leaves of the germinating weeds and
remains active in the soil for some weeks. The weed spectrum of mono-
linuron is very similar to linuron, but it is marginally more active on
knotgrass and annual meadow-grass and less effective on groundsel. The
broad-leaved weed species that are not controlled post-emergence by
linuron are also not controlled at this stage by monolinuron. Common
fumitory is resistant. The doses recommended vary from 0.84 kg ha- 1 for
early potatoes on light soils to 1.68 kg ha- 1 on heavy and organic soils. This
herbicide should not be used on very light soils because of the risk of crop
damage, but can be used on seed potato crops. It is recommended that
other crops should not be sown for 2-3 months after application, depend-
ing on the dose used. Lettuce should not be sown in the same season.
Prometryn
This is another pre-emergence herbicide with mainly soil activity but with
some effect on emerged weeds. Many species are susceptible until they are
50 mm high but knotgrass, mayweeds and corn marigold must be treated
pre-emergence. Unlike treatment with the urea herbicides (linuron, mono-
linuron) common fumitory is susceptible. It is only recommended for use in
early potatoes and should be applied at 1.7 kg ha -1 before 10% of the
potatoes have emerged. On highly organic soils it should only be used post-
weed emergence as soil activity on these soils is minimal. Any crop may be
grown 8 weeks after application but peas or umbelliferous crops (e.g.
carrots) can be sown earlier.
Terbutryn + prometryn
This formulated mixture may be used on early and maincrop potatoes,
until 10% of crop plants have emerged, on all soil types except sands and
those with more than .10% organic matter. Seed potato crops should not be
treated. A lower dose (1.39 kg ha- 1) is recommended for early potatoes on
light soils and higher ones, up to 1.82 kg ha- 1 , on heavy soils growing
maincrop potatoes. Many broad-leaved weed species are susceptible up to
the fully expanded cotyledon stage of growth, including common fumitory
and mayweeds. Any crop may be sown after the potato harvest provided
that the land is thoroughly cultivated or ploughed to 150 mm prior to
sowing.
Terbutryn + terbuthylazine
This mixture of triazine herbicides is again mainly soil acting. It does have
some effect on emerged seedling weeds, but only whilst they are at the
cotyledon stage of growth. Cleavers plants are even more resistant to this
mixture than they are to the urea herbicides but common fumitory is
susceptible. Both maincrop and early potatoes should be treated before
10% of the potato plants have emerged. If weeds are emerged this
formulated mixture can be mixed with paraquat. Doses are related to the
soil type, 1.15 kg ha- 1 being recommended for light soils and early potatoes
and 1.7 kg ha- 1 on heavy and organic soils. Potatoes on sandy soils should
not be treated because of the risk of crop damage. Land should be
ploughed or cultivated to 150 mm before sowing the next crop, which
should not be planted within 12 weeks of application (14 weeks if soil
conditions have been very dry).
Terbutryn + trietazine
As with the previous terbutryn mixture, this product is mainly soil acting
but does have some activity on weeds with emerged cotyledons. A wide
390 Weed control
range of broad-leaved weeds, including common fumitory, are susceptible

but cleavers is not reported to be sensitive. All varieties of early and
maincrop potatoes can be treated until 10% of the shoots have emerged. A
dose of 1.5 kg ha- 1 is recommended on all soils for early potatoes and for
maincrops on very light soils. Maincrops on heavy or organic soils can be
treated with up to 2.5 kg ha- 1 . Before sowing another crop, the land should
be ploughed to a depth of 150 mm. Any crop can be planted 12 weeks after
application, except bras sic as where the interval should be 14 weeks.
(e) Soil-acting pre-emergence herbicides

Herbicides in this group must be applied before the weeds emerge and so
should be applied fairly soon after the crop is planted, or in association
with a contact herbicide, such as paraquat, if application is delayed.
Pendimethalin
Pendimethalin may be used only on maincrop and some second early
potatoes but not on seed or first early potatoes. It is only recommended in
mixtures with low rates of cyanazine or metribuzin and it is important that
it is applied as soon as possible after planting and reridging. It must be
applied at least 7 days before the emergence of the most advanced potato
shoots. The activity of pendimethalin will be reduced if prolonged dry
weather follows application. Best weed control will be achieved if rain
follows within 7 days of treatment, or if the crop is irrigated. If emerged
weeds are present at application, pendimethalin with cyanzine may be
mixed with paraquat to improve control. The pendimethalin with metribuzin
mixture will control small emerged weeds. The dose is not influenced by
soil type and the pendimethalinlcyanazine mixture can be used on all
mineral soils, except sandy and gravelly soils. The metribuzin mixture
should not be used on stony and gravelly soils. There is no recommenda-
tion for use on organic soils. Performance is less good if the soil is cloddy
when the herbicide is applied, or if the soil is disturbed after treatment.
A dose of 1.32 kg ha- 1 is recommended in both mixtures, together with
0.75 kg ha- 1 cyanazine or 0.35 kg ha- 1 metribuzin. The weed susceptibilities
of the two mixtures differ. For example, corn marigold and parsley piert
(Aphanes arvensis) are more sensitive to the cyanazine mixture and black
bindweed and field pennycress (T. arvense) to the metribuzin mixture.
Cleavers, a weed that is particularly difficult to control in potatoes, may be
suppressed by the metribuzin mixture.
(f) Post-crop emergence

Few truly selective post-emergence herbicides are available for use in
potatoes. Consequently, it is advisable to ensure that weeds are controlled
mainly before the crop has emerged. A number of post-emergence
herbicides are now available for the control of grass weeds in broad-leaved
crops (Knott, 1985). At present, not all these are approved for use in
potatoes. The availability of selective herbicides for post-emergence
control of broad-leaved weeds is even more limited.
Alloxydim-sodium
This translocated herbicide can be used in potatoes to control many annual
grass weeds and for the suppression of common couch. A lower dose of
0.94 kg ha- 1 is recommended for annual grasses and a higher one for
common couch (1.87 kg ha- 1 ). The annual grasses should have at least
three leaves and the common couch be 300 mm high. It may be used in all
potato crops, once they have emerged until just before the crop canopy
closes. Annual meadow-grass is not susceptible. Alloxydim may be pre-
ceded by linuron and monolinuron and by pre-emergence metribuzin,
provided there are at least 3 weeks between the treatments. At least 4
weeks should elapse between treatment and harvest.
Bentazone
Bentazone can be used at 1.44 kg ha- 1 to control some broad-leaved weeds,
especially fat hen, redshank (Polygonum persicaria) and cleavers which
compete with the crop late in the season. Two lower rate applications (e.g.
0.72 + 0.72 kg ha- 1) have been found to be more effective than one full rate
treatment. Fat hen is controlled more effectively if oil is added to
bentazone. This herbicide can be applied after pre-emergence herbicides,
but not after post-emergence metribuzin, to some varieties of maincrop
and second early potatoes, but not seed potato crops. Some leaf chlorosis
or scorch may follow application, especially if temperatures are high when
it is applied. Do not treat potatoes when temperatures are at or above
21°e, in the middle of the day. The potatoes should not be more than
150 mm high when treated and the weeds should be at the cotyledon to
seedling stages. This treatment is particularly useful when dry soil con-
ditions have caused the failure of pre-emergence herbicides.
Cycloxydim
This is another translocated post-emergence herbicide for the control
of grass weeds. Doses vary according to the species to be controlled,
being lower (0.15 kg ha- 1 ) for annual grasses and higher for perennials
(0.45 kg ha- 1). Annual weeds should be treated when they are between
the two-leaf stage and the end of tillering and perennials when they are
150 mm high. Annual meadow-grass is resistant. The adjuvant oil Actipron
should be added to this herbicide for all treatments. At the moment it
should not be applied in sequence with other post-emergence herbicides,
with the exception of bentazone, which can be followed with cycloxydim
after 14 days.
392 Weed control
Diclofop-methyl
Diclofop is a translocated post-emergence herbicide for the control of
some annual grass weeds, especially wild oats (A. fatua) , young black-grass
(Alopecurus myosuroides) and rye-grasses. Potatoes can be treated from
100% emergence onwards. The weeds should be treated at 1.14 kg ha- 1
when most plants have three leaves and the largest not more than four
leaves plus one to two tillers. At least 6 weeks should elapse between
spraying and harvest.
Metribuzin
This herbicide can be applied after the emergence of maincrop potatoes as
well as pre-emergence (see p. 388). until the shoots are 150 mm high (variety
Maris Piper and some others should only be treated pre-emergence). The
doses used and the other conditions associated with the use of this
herbicide are similar to those required for pre-emergence applications.
Sethoxydim
Most grass weeds, except annual meadow-grass, are sensitive to this
herbicide. It is most effective when applied to actively growing weeds in
warm weather. Annual grass weeds are sensitive from the two-leaf stage to
the end of tillering to 0.34 kg ha- 1 and common couch should be treated,
when the largest leaves are at least 300 mm long, with 0.87 kg ha- 1 . With
the perennial grasses it is important that the plants have adequate foliage
to permit good translocation to the rhizomes. The weeds should not be
treated before the crop is 200 mm high, nor after the potato canopy
becomes too dense to permit penetration of the spray.
.
9.4.3 Herbicides under development
A number of other herbicides are being developed for use in potatoes but,
although some are being used elsewhere in Europe, none is yet approved
in the UK. A non-selective foliar-acting herbicide, glufosinate (Schwerdtle
et al., 1981), is currently available in Germany as an alternative to
paraquat/diquat. It is slightly more systemic than paraquat and so can be
more effective on weeds. However, because of its greater translocation, it
can be more damaging to potato shoots if applied after they have emerged
(Lawson et al., 1985). Recent research in Belgium has demonstrated that it
is possible to transfer genes for resistance to glufosinate into potato plants,
thus making it possible to use this herbicide after potato emergence
(Botterman and Leemans, 1988). Several alternative post-emergence grass
weed herbicides appear to have potential for use in potatoes. These include
ftuazifop-butyl (Plowman et al., 1980) and quizalofop (Sakata et al., 1983),
both of which are widely used in other broad-leaved crops, such as sugar
beet and oilseed rape. Three pre-emergence broad-leaved and grass weed
herbicides were discovered in the early 1980s: aclonifen (Buck et al., 1983),
fomesafen (Colby et at., 1983) and fluorochloridone (Pereiro et at., 1982;
Forbes and Matthews, 1985). All three appear to be selective in potatoes
and have a wide weed spectrum that includes for example, cleavers and
volunteer oilseed rape. Fluorochloridine is already widely used in France,
especially for weed control in sunflowers.
9.4.4 Herbicide residues

The potential problems to subsequent crops caused by soil residues of
potato herbicides have already been mentioned in the sections on the
relevant herbicides. It must be emphasized that the instructions on the
label regarding following crops and post-potato harvest cultivations must
be followed. Current concerns over the presence of herbicides in ground-
water also reinforce the need for care when using the more persistent soil-
acting products. Indeed in the foreseeable future it seems likely that some
herbicide uses will be restricted or withdrawn, as has already occurred in
Germany. This may have serious implications for the control of weeds in
this crop. A number of incidents of crop damage occur each year because
of failure to follow the label instructions (Eagle, 1981; Caverley, 1987). A
further aspect of problems caused by residues is the damage caused to
potatoes by herbicides used in other crops. For example, Caverley (1987)
has reported damage to potatoes from isoproturon applied to previous
crops of cereals. Recently, problems have arisen in potato seed crops from
the application of blight sprays that were contaminated with herbicides.
Low concentrations of clopyralid and of metsulfuron methyl, present in
spray tanks because of inadequate washing, can affect the appearance and
vigour of daughter tubers of potato plants that have themselves exhibited
few or no symptoms of damage (Lawson and Wiseman, 1986, 1987;
Lawson et at., 1990). Similarly, low concentrations of glyphosate, con-
taminating blight sprays, can have a devastating effect on both the potato
plants and on the vigour of the daughter tubers. These problems have
arisen because of the increase in late applications of herbicides to cereal
crops, which are carried out at the same time as early blight sprays. The
need for thorough washing of spray tanks is clear.
9.4.5 Weed control under plastic mulch

In recent years there has been increased interest in the production of early
potatoes under plastic mulches (Askew, 1986). This system has implica-
tions for weed control, as once the mulch (plastic film) is in place no further
weed control is possible. In addition, the warm conditions under the mulch
can affect the performance of previously applied soil-acting herbicides and
will increase the vigour of the weeds. If the herbicide treatments and the
plastic mulch are applied immediately after planting, the herbicide will
have to remain active for several weeks longer than normal. In addition,
394 Weed control
the warm temperatures under the mulch are likely to dry the soil surface,
rendering the herbicide less effective. These problems will be reduced if
the application of the plastic mulch is delayed, allowing perhaps a week for
natural rainfall to incorporate the herbicide into the soil. Mulches protect
the soil from rainfall and so the leaching of the herbicides may be less and,
consequently, persistence higher. This may result in higher herbicide
residues in tubers and problems with soil residues affecting subsequent
crops.
9.5 POTATO HAULM DESTRUCTION
The ease with which a potato crop is harvested is influenced by the health
and vigour of the potato plants as they approach maturity, the presence of
weeds and the condition of the soil. Haulm destruction often simplifies
harvesting, although modern machines are capable of harvesting plants
with healthy green haulm. However, large amounts of green haulm can
seriously impede harvesting, reducing soil separation, blocking the webs
and making sorting of tubers more difficult. In addition, haulm destruction
will kill any surviving weeds that would otherwise also slow harvesting
operations. Haulm destruction can also be used to manipulate tuber size
distribution and will slow the development of diseases such as potato blight
(Phytophthora infestans). The latter feature is particularly important in
seed crops.
In maincrop potatoes, natural senescence will start in late summer and,
especially in dry summers or where the haulm has been infected with
disease, the haulm will be partially senescent at harvest. In wetter summers
the haulm can continue to grow and will remain green. The onset of
senescence is accompanied by changes in the skin of the tubers (skin set),
making them more resistant to mechanical damage and disease. Haulm
destruction can hasten senescence and skin set in maincrops. Early
potatoes do not normally have their haulm destroyed prior to harvesting,
as maximizing yield is of prime importance and skin set is a marketing
disadvantage. In addition, early potatoes have less haulm when harvested.
Early potato seed crops may be desiccated for the same reason as maincrop
seed crops.
9.5.1 Haulm destruction by mechanical methods

Mechanical destruction by pulverizing the haulm with metal or rubber flails
revolving at high speed is used in ware crops. This method has the
advantage of allowing 'instant' access for the harvester but has the
disadvantage of permitting the continued persistence of diseases such as
potato blight and blackleg, which may then spread to the tubers via the
incompletely killed haulm. Specially designed machines are available
Potato haulm destruction 395
where the flails are arranged to follow the contours of the ridge, but it is
possible to use standard forage harvesters. These are less effective and may
also blow the flailed haulm onto other ridges.
9.5.2 Haulm destruction by chemical methods

The use of chemical desiccants to destroy the potato haulm was quickly
adopted by growers as a quick and effective method of killing potato haulm
and weeds prior to harvest. The technique had the added advantage that
the use of a desiccant minimizes the spread of diseases from infected haulm
to the tubers. Thus, in practice, desiccants are often used if blight becomes
established in a crop, to minimize tuber infection. Although photosyn-
thesis stops soon after application of the desiccant, the tubers continue to
absorb water and so in wet years there can be an appreciable increase i.n
tuber size, perhaps affecting the crop's marketability for seed. This uptake
of water will also reduce tuber dry matter which may affect the suitability
of the tubers for processing.
A range of chemical desiccants were used in the 1950s, including
sulphuric acid, sodium arsenite, tar oil, sodium chlorate, DNOC and
copper sulphate + salt. Most of these are no longer available. In the 1980s
diquat, metoxuron, dinoseb and sulphuric acid were the most widely used,
but dinoseb is now no longer available in the UK.
(a) Diquat
Diquat is the most widely used potato desiccant, giving a quick kill of the
haulm. It should be applied at 0.8 kg ha- 1 , preferably in bright light and in
low humidity. If it is applied in dry soil conditions it can be translocated
into the tubers causing vascular browning and, in extreme cases, stem end-
rot. Potatoes grown on light soils are more likely to be affected in this way
than those on heavier soils, as are cultivars that are drought susceptible.
Precise details as to conditions when the product should or should not be
used are given on the product label.
(b) Metoxuron
A mixture of metoxuron and fentin hydroxide is recommended at
2.2 kg ha- 1 , for ware potato haulm desiccation. It should be used as soon as
the haulm has begun to senesce naturally and may take 7-10 days for
symptoms to appear. It will desiccate some weeds, but not dense infesta-
tions of large, late germinating species, such as fat hen. There is little risk
of vascular browning in dry conditions.
396 Weed control
(c) Sulphuric acid
This causes very rapid kill of the haulm and so is especially useful to
prevent further spread of disease. However, as it is applied as diluted
Brown Oil of Yitriol (BOY), a strong acid, it is very corrosive and so it
must be applied by specialist contractors and not by individual farmers.
Doses vary according to growing conditions and the degree of haulm
senescence at application. It is particularly favoured under dry conditions
when diquat can cause some tuber damage (see above).
9.6 POTATOES AS WEEDS
Potatoes are generally grown as one constituent of a crop rotation,

occurring only once every 4 or 5 years. This length of rotation is favoured
because it minimizes the risk of build-up ·of potato pests and diseases,
especially eelworm. However, in recent years, with the development of
eelworm-resistant potato varieties, there has been a tendency to shorten
rotations and grow potatoes more often, particularly on favoured fields. In
some areas of Holland potatoes are grown every other year. It is an
inevitable consequence of harvesting potatoes that some tubers will be left
in the field after harvest. When potatoes were hand harvested and there
was a plentiful supply of labour, the number of tubers left behind was
small. The development of complete harvesters, and the decline in labour,
has resulted in many more tubers being left behind after harvest. The
problems caused by this have been exacerbated by the shortening of
rotations. Plants from groundkeeper tubers can cause weed problems in
subsequent crops and may affect the health of the next potato crop.
As long ago as 1948, Doncaster and Gregory suggested that volunteer
potatoes could affect virus levels in potato crops. A further problem
has developed in recent years. Although traditional potato varieties,
such as King Edward, set few seeds, modern varieties, such as Maris Piper
and Desiree, produce many berries and seeds. In consequence, weed
problems have arisen from the true seedlings that emerge in subsequent
crops.
The potatoes discarded from processing lines can also act as sources
of several potato diseases. Therefore, it is of utmost importance that
the clamps of these potatoes are destroyed before the tubers start to
grow in spring. Dichlobenil granules can be used for this purpose in winter,
but as this is a very persistent herbicide it can leave high soil residues
that could affect crops planted after the removal of the clamp. If these
residues are likely to cause problems a foliage acting herbicide, such as
glyphosate, should be used in the spring. Spring treatments are not as
effective at preventing disease spread, as it is possible for the foliage to be a
source for disease in neighbouring healthy crops before it is treated and
killed.
Potatoes as weeds 397
9.6.1 Volunteer potatoes
(a) Biology
The origin of the problems caused by volunteer potatoes, apart from those
derived from true seed (see Section 9.6.2), lies in the inability of potato
harvesters to remove all potato tubers from the field. Surveys have shown
that populations in excess of 100000 groundkeeper tubers ha- l were
frequently left in the field after harvest and as many as 370 000 ha- l have
been recorded (Lutman, 1977). The majority of these tubers were small,
less than 40 mm in diameter and had fallen through the webs of the
harvester. Inefficient harvesting, leaving ware-sized tubers, exacerbates
the problem. Surveys have shown that only 30-50% of ground keeper
tubers were on the soil surface, so post-harvest gleaning of the fields after
harvest to remove surface tubers is only partially effective. During the
winter, tubers on or near to the soil surface will be killed by frost and eaten
by birds and animals. Buried tubers will often survive, unless frost
penetration of the soil is appreciable. Experiments in Holland have shown
. that tubers must be exposed to at least 50 frost hours with temperatures
below -2°C (e.g. 25 h at -2°C, 10 h at -5°C) (Lumkes and Sijtsma, 1972)
before they are killed. In England, deeply buried tubers will be exposed
only rarely to these temperatures.
Shoots from the surviving tubers emerge in late spring and summer
(May-July). There tend to be fewer plants in very competitive crops (e.g.
winter barley), than in more open row crops (e.g. sugar beet). Emergence
also tends to be later and tuber production lower, in competitive crops.
Hence the competitive effects of volunteer potatoes are much more severe
in row crops such as sugar beet and field vegetables, than they are in
cereals. The tubers produced by these plants ensure the continuation of the
weed into the following year. It has been calculated that tuber populations
will decline by approximately 60% following a winter cereal crop (Lutman,
1986). This suggests that there would be a 99% decline over 5 years.
Observations made in seed potato growing areas have demonstrated that
potatoes can survive 5 year rotations in appreciable numbers and may
survive for as long as 12 years. These surviving plants may cause the
rejection of seed potato crops for certification, because of the presence of
'rogues'. Pieces of potato haulm and berries can also cause the rejection of
processing crops such as peas and beans, because of their potential toxicity
and the difficulty of removing them from the harvested products. Not only
are the surviving volunteer potatoes serious contaminants of other crops,
they are also very competitive, reducing yields of the crops in which they
occur, and are potential foci for potato diseases and pests. Den Ouden
(1967) suggested that volunteer potato plants could increase the persis-
tence of potato cyst nematode (Globodera rostochiensis, G. paUida) and
other research has suggested that they may increase gangrene (Erwinia
spp.) and virus diseases (Fox, 1983; Thomas, 1983).
398 Weed control
(b) Control
The control of volunteer potatoes is not easy, although a number of
potential opportunities exist. As the origin of this weed problem is tubers
left in the field after the potato harvest, it is vital that crops are harvested as
efficiently as possible, to minimize leavings. Over-winter mortality of
ground keepers will be maximized if the land is not ploughed, as this will
result in the greatest number of tubers being killed by frost. Ploughing
buries many tubers below normal freezing depth. One of the best ways of
reducing the effects of volunteer potato plants is to ensure that the crops
succeeding potatoes are as competitive as possible. Winter cereals and
grass leys minimize the production of daughter tubers and so maximize the
rate of decline in the populations. In subsequent crops of cereals it is
possible to achieve some control of haulm and tubers with fluroxypyr
(Graham et al., 1987; Ogilvy et al., 1989), but often the potato shoots do
not emerge until the cereals are beyond the growth stages that are safe to
be sprayed. Pre-harvest applications of glyphosate can be successful in
winter barley but performance is affected by the vigour of the potato plants
when treated. Applications in the later harvested winter wheat tend to be
less effective because of the greater senescence of the potato plants
(Lutman, 1986). Selective control of potatoes in broad-leaved crops is
difficult, but metoxuron in carrots and dendritic salt in sugar beet can have
some effect (Williams, 1986). Autumn applications of glyph os ate in cereal
stubbles will kill tubers attached to growing potato plants. Unfortunately,
many tubers fail to initiate regrowth, thus reducing the effectiveness of the
treatment. The application of glyphosate through selective applicators to
potato plants growing in shorter crops has been investigated but the results
have been generally disappointing. Thus, the control of potatoes with
herbicides is not easy, although glyphosate can be effective if applied to
potato plants growing in the absence of a crop. One further method of
control has been developed for ware potato growers. Application of the
growth regulator maleic hydrazide to the potato crop just prior to the onset
of senescence will inhibit the sprouting of the daughter tubers and has been
reported to reduce the numbers of volunteers in the following year (Peddie
et al., 1986; Ogilvy et al., 1989).
9.6.2 Potato seedlings
Over the last 20 years there has been a dramatic increase in the cultivation
of potato cultivars that produce large numbers of berries and, con-
sequently, seeds. Lawson (1986) has calculated that some cultivars will
produce over 100 million seeds ha- 1 . These seeds can remain viable for at
least 7-8 years. Potato seedlings are difficult to control and give rise to
tubers that perpetuate the weed in the same way as volunteers. It is
possible that the occurrence of volunteer potato plants in seed potato crops
References 399
grown in very long rotations (up to 12 years) may be due to plants arising
from true seed. Germination takes place in spring and summer and,
consequently, true seedlings cause weed problems mainly in spring-sown
crops. The performance of herbicides on potato seedlings in these spring
crops is variable and, although they are easier to control than volunteer
potatoes, control is not simple.
REFERENCES
Askew, M.F. (1986) Weed control in potatoes. Aspects of Applied Biology 13, Crop
Protection of Sugar Beet and Crop Protection and Quality of Potatoes, pp. 227-38.
Askew, M.F. and Flint, C.E. (1985) Trials with metribuzin applied as a low dose
programme. Proceedings 1985 British Crop Protection Conference - Weeds, pp.
819-26.
Ball, A.P. (1972) The use of a mixture of trietazine and linuron for weed control in
potatoes. Proceedings 11th British Weed Control Conference, pp. 511-18.
Botterman, J. and Leemans, J. (1988) Engineering herbicide resistance in plants:
status and perspectives. Proceedings EWRS Symposium Factors Affecting
Herbicidal Activity and Selectivity, pp. 331-6.
Bremner, P.M. (1966) The effect of cultivation on yield of the potato crop.
Proceedings 8th British Weed Control Conference, pp. 1-7.
Buck, W., Friedlander, H., Linden, G. and Schneider, G. (1983) CME 127 a new
pre-emergence herbicide. Proceedings 10th International Congress of Plant
Protection, pp. 307-14.
Caverley, D.J. (1987) Advisory problems with residual soil herbicides. Proceedings
1987 British Crop Protection Conference - Weeds, pp. 601-10.
Colby, S.R., Barnes, J.W., Sampson, T.A., et al. (1983) Fomesafen - a new
selective herbicide for post-emergence broad-leaved weed control in soybean.
Proceedings 10th International Congress of Plant Protection, pp. 295-302.
Davis, R.P., Garthwaite, D.G. and Thomas, M.R. (1990) AD AS Pesticide Usage
Survey Report 78. Arable Farm Crops 1988, Ministry of Agriculture, Fisheries &
Food, Reference Book, 578, 56 pp.
Den Duden, H. (1967) The influence of volunteer potato plants in oats on the
population density of Heterodera rostochiensis. Nematologica, 13, 325-35.
Doncaster, J.P. and Gregory, P.H. (1948) The survival of volunteer potatoes in
arable fields, in The Spread of Virus Diseases in the Potato Crop. UK
Agricultural Research Council Report, 7, 121-8.
Eagle, D.J. (1981) Residue problems encountered in England. Proceedings
EWRS Symposium. Theory and Practice of the Use of Soil Applied Herbicides,
pp.201-7.
Elliott, J.G. (1965) Development of chemical weed control in potatoes. 1st Annual
Report ARC Weed Research Organisation, 1960-1964, pp. 10--14.
Forbes, G.R. and Matthews, P.R. (1985) Pre-emergence R-40244 for early weed
control in potatoes and carrots. Proceedings 1985 British Crop Protection
Conference - Weeds, pp. 797-804.
Formigoni, A., Iwataki, I. and Ishihara, H. (1979) Selective post-emergence grass
control in different broadleaf crops with the new herbicide NP55. Proceedings
400 Weed control
EWRS Symposium. The Influence of Difference Factors on the Development and
Control of Weeds, pp. 403-10.
Fox, R.A. (1983) Potatoes: problems, progress and future research. Proceedings
10th International Congress of Plant Protection, pp. 1157-64.
Graham, J.C., Bunn, F.E. and Jeffery, P.J. (1987) The control of volunteer
potatoes with fluroxypyr in UK cereals. Proceedings 1987 British Crop
Protection Conference - Weeds, pp. 241-8.
Griesbach, E. and Eisbein, K. (1975) The importance of weeds for the transmission
of Rhizoctonia solani (Kuhn). III The influence of weeds on the infestation of
potatoes by Rhizoctonia solani. Zbl. Bakt. Abt. II Bd., 130, 745--60.
Hampson, c.P. and Taylor, J.A.H. (1970) Herbicide usage on maincrop potatoes
in Great Britain. Proceedings 10th British Weed Control Conference, pp.
1128-31.
Headford, D.W.R. (1968) The effect of time of paraquat application on weed
control and the growth of potatoes. Proceedings 8th British Weed Control
Conference, pp. 547-52.
Ingram, G.H., Turner, M.T.F., Hirono, Y. and Iwataka, I. (1978) Alloxydim-
sodium for grass weed control in broad leaved crops in the UK. Proceedings
1978 British Crop Protection Conference - Weeds, pp. 761-8.
Ivany, J .A. (1974) Effects of delayed paraquat application on Sebago potato yields.
Can. J. Plant Sci., 54, 853-4.
Jarvis, R.H. and Shotton, F.E. (1972) Cultivation systems for maincrop potatoes.
Experimental Husbandry, 22, 61-8.
Knott, C.M. (1985) Grass-weed control in broad-leaved crops - the options.
Proceedings 1985 British Crop Protection Conference - Weeds, pp. 429-40.
Lawson, H.M. (1986) Potato seedlings: a review of the current situation. Aspects of
Applied Biology 13, Crop Protection of Sugar Beet and Crop Protection and
Quality of Potatoes, pp. 187-94.
Lawson, H.M. and Wiseman, J.S. (1985a) Tolerance of seed potato crops to a
range of selective graminicides. Proceedings 1985 British Crop Protection
Conference - Weeds, pp. 457--62.
Lawson, H.M. and Wiseman, J.S. (1985b) Evaluation of new residual herbicides
for use in potato seed crops. Proceedings 1985 British Crop Protection
Conference - Weeds, pp. 805-10.
Lawson, H.M. and Wiseman, J.S. (1986) Contamination of seed potato crops.
Annual Report Scottish Crop Research Institute, 1986, p. 147.
Lawson, H.M. and Wiseman, J.S. (1987) Contamination of seed potato crops
by herbicides. Annual Report Scottish Crop Research Institute, 1987, pp.
159--60.
Lawson, H.M., Wiseman, J.S., Davies, D.H.K. and Richards, M.C. (1985)
Tolerance of potato crops to glufosinate-ammonium applied as an early post-
emergence herbicide. Proceedings 1985 British Crop Protection Conference -
Weeds, pp. 811-7.
Lawson, H.M., Wiseman, J.S. and Wright, G. MeN. (1990) Tolerance of seed
potatoes to contamination with two sulfonyl urea herbicides. Annals of Applied
Biology, 116, (supp!.), Tests of Agrochemicals & Cultivars, 11, 76--7.
Lumkes, L.M. and Sijtsma, R. (1972) Mogelijkheden aardappelen als onkruid in
volgewassen to voorkomen enlof te bestrijden. Landbouw en Plantenziekten,
May 1972, 17-36.
References 401
Lutman, P.J.W. (1977) Investigations into some aspects of the biology of potatoes
as weeds. Weed Res., 17, 123-32.
Lutman, P.J.W. (1986) The biology and control of groundkeeper potatoes: a
review. Aspects of Applied Biology 13, Crop Protection of Sugar Beet and Crop
Protection and Quality of Potatoes, pp. 177-85.
Mannall, H.G., Davies, M.E. and Whitworth, S.H. (1972) The development of
metribuzin in the UK for weed control in potatoes. Proceedings 11th British
Weed Control Conference, pp. 519-27.
Neild, J.R.A. and Proctor, J.M. (1962) Chemical weed control in potatoes.
Proceedings 6th British Weed Control Conference, pp. 697-712.
Nelson, D.C. and Thoreson, M.C. (1981) Competition between potatoes (Solanum
tuberosum) and weeds. Weed Sci., 29, 672-7.
Nieto, J.H., Brondo, M.A. and Gonzalez, J.T. (1968) Critical periods of the crop
growth cycle for competition from weeds. PANS (C), 14, 159--66.
Ogilvy, S.E., Cleal, R.A.E. and Rogers-Lewis, D.S. (1989) The control of potato
groundkeepers in cereal crops. Proceedings 1989 Brighton Crop Protection
Conference (Weeds), pp. 205-12.
Orson, J.H. (1986) Chemical control of weeds in maincrop potatoes; ADAS
Agriculture Service and Terrington EHF trial results, 1983-1986. Aspects of
Applied Biology 13, Crop Protection of Sugar Beet and Crop Protection and
Quality of Potatoes, pp. 253--6l.
Peddie, A.S., Bartlett, D.H. and Taverner, P.B. (1986) Pre-harvest application of
potassium maleic hydrazide to potatoes to suppress volunteers in succeeding
crops. Aspects of Applied Biology 13, Crop Protection of Sugar Beet and Crop
Protection and Quality of Potatoes, pp. 219-25.
Pereira, H.e. (1941) Studies in soil cultivation ix. The effect of interrow tillage on
the yield of potatoes. J. Agric. Sci., Camb., 31, 212.
Pereiro, F., Ballaux, J.e. and Beraud, J.M. (1982) R 40244, a new herbicide for
weed control in potatoes, sunflower and winter wheat. Proceedings 1982 British
Crop Protection Conference - Weeds, pp. 225-30.
Plowman, RE. Stonebridge, W.C. and Hawtree, J.N. (1980) Fluazifop-butyl - a
new selective herbicide for the control of annual and perennial grass weeds.
Potato Marketing Board (1963) Report on the Survey of Maincrop Potatoes, 1963,
18 pp.
Potato Marketing Board (1979) Report on the Survey of Maincrop Potato
Production Techniques 1977-78, 84 pp.
Potato Marketing Board (1987) Potato Statistics in Great Britain 1983-1987, 25 pp.
Roberts, H.A. Chancellor, RJ. and Hill, T.A. (1982) The biology of weeds, in
Weed Control Handbook: Principles (ed. H.A. Roberts), Blackwell Scientific
Publications, Oxford, pp. 1-36.
Saghir, A.R and Markoulis, G. (1974) Effects of weed competition and herbicides
on yield and quality of potatoes. Proceedings 12th British Weed Control
Conference, pp. 533-9.
Sakata, G., Makino, K., Kawamura, Y. etal., (1983) NCI-96683, a new selective
herbicide for annual and perennial grass weed control in broad-leaf crops.
Proceedings 10th International Congress of Plant Protection, pp. 315-24.
Schwerdtle, F., Bieringer, H. and Finke, M. (1981) HOE 39866 - ein neues nicht
selectives blattherbizid. Z. PflKrankh. PflSchutz. Sonderh., IX, 431-40.
402 Weed control
Taylor, J.A.H. (1976) Herbicide usage on maincrop potatoes in Great Britain.
Taylor, J.A.H. (1980) Herbicide usage on potatoes in Great Britain in 1980.
Thackral, K.K., Pandita, M.L., Khurana, S.c. and Kalloo, G. (1989) Effect of
time of weed removal on growth and yield of potatoes. Weed Res., 29, 33-8.
Thomas, P.E. (1983) Sources and dissemination of potato viruses in the Columbian
basin of the Northwestern USA. Plant Disease, 67,744--7.
Vitolo, D.B. and Ilnicki, R.D. (1985) Grass competition in white potatoes.
Abstracts Weed Science Society of America, 25, 30.
Williams, P.G. (1986) Some practical experiences of controlling volunteer potatoes
in sugar beet and some other crops. Aspects of Applied Biology 13, Crop
Protection of Sugar Beet and Crop Protection and Quality of Potatoes, pp.
195-9.
CHAPTER 10
Disease aspects of potato production

G.A. Hide and D.H. Lapwood
10.1 INTRODUCTION
Disease control is a prerequisite for improving and maintaining yield and

quality of the potato crop and since the potato became widely grown
serious outbreaks of disease and crop failures and consequent social and
economic effects have repeatedly provided incentive for improvement. For
example, in the mid-eighteenth century the widespread occurrence of leaf
roll in Germany and the UK led to poor yields, and in the mid-nineteenth
century late blight devastated crops in the New York and New England
states of the USA. A little later, the catastrophic blight epidemics ravaged
the Irish crop, resulting in famine and mass emigration of the population.
Towards the end of the nineteenth century, wart disease, found first in
Czechoslovakia and later in other parts of Europe, threatened the survival
of the cultivars of the day. When the causes of these diseases were
discovered, early investigators formulated what have become established
as the basic methods of disease control. For example, studies of viruses and
wart led to the establishment of seed tuber production in areas less
favourable to the spread of disease, isolated from the main potato growing
regions and where close control by inspection for freedom from diseases as
well as for purity of cultivar could be maintained. More recently the occur-
rence of rots and blemishes on freshly harvested or stored potatoes has
stimulated investigations into ways of conrolling or at least avoiding them.
The potato is prone to more than a hundred diseases caused either by
bacteria, fungi, viruses or mycoplasms but fortunately relatively few reach
serious proportions in anyone growing area. Whereas late blight is
generally the most important disease wherever potatoes are grown, some
diseases are serious only in cool temperate regions (e.g. skin spot and
gangrene) and others in warmer climates (e.g. early blight, bacterial wilt or
brown rot), perhaps related to their region of origin.
It seems probable that most potato pathogens have been transported
from country to country in the normal course of international trade in
404 Disease aspects of potato production
potatoes over the centuries. Speedier transport may well have increased
their chances of survival on tubers; certainly expanding trade has provided
greater opportunities in this century .. The absence of a disease from a
particular country may result from failure of introductions to become
established because of unfavourable climate, but in more recent times
import restrictions could have prevented the initial introduction of some
diseases into countries whose climate is favourable for their development.
For example, at present importation of tubers affected with bacterial ring
rot is prohibited into the UK, where the disease is unknown, and strict
quarantine precautions are taken with tubers imported as germplasm from
S. America to avoid the potentially very serious smut disease caused by
Angiosorus (Thecaphora) solani. Also the incidence of some diseases on
table potatoes (ware) in England may be related to their prevalence in
Scotland and Ireland where many seed stocks are produced.
Pathogens may attack foliage, root systems and tubers, so disease can be
important throughout the crop cycle. Seed tubers may deteriorate before
or soon after planting especially when seed is cut; the growing plant may be
weakened by direct attack or systemic infection and thereby less productive,
or the roots or foliage may be attacked directly causing their premature
death. Tubers may become infected during growth or, often more import-
antly, through wounds made at harvest or subsequent handling. Diseases
therefore are capable not only of decreasing production but also of blem-
ishing and rotting tubers so that the quality of the crop is adversely affected.
Since potatoes are usually propagated vegetatively, the seed tuber often
provides the major source of inoculum, ensuring the continued survival of
some diseases (e.g. late blight), but symptoms may not always be present,
for example, when infection is 'latent' (e.g. blackleg) or when inoculum is
carried as a surface 'contamination' as often occurs with dry rot and
gangrene, or as oospores with pink rot. Therefore the disease status of a
crop often reflects the spectrum of diseases on the seed tubers. Ground-
keeper or 'volunteer' plants surviving from previous potato crops or
diseased tubers removed from store and dumped on cull piles may also
help perpetuate disease (e.g. late blight, viruses) as do some solanaceous
and other weed hosts. Sometimes susceptible agricultural crops provide a
reservoir of inoculum (e.g. bacterial brown rot) whereas the pathogens
causing powdery and common scab, pink rot and wart can survive
considerable periods in soil in the absence of known hosts.
Worldwide losses of potato yield through disease have been estimated at
30% (Cramer, 1967). In the USA, average losses for the period 1951-60
were 19%, with late blight, leaf roll, X and Y viruses and verticillium wilt
accounting for at least 2% each. However, these figures almost certainly
exclude losses occurring after lifting from damage and disease which in the
UK can account for 30% of the harvested yield (Twiss and Jones, 1965).
By contrast, it is more difficult to estimate costs to the potato industry of
preventing disease. In any calculation, the cost of producing healthy
Effects of diseases 405
seed tubers, manufacturing, developing and applying plant protective
chemicals, developing specialized machinery for potato handling and
storage, breeding disease-resistant cultivars and of the necessary research,
advisory and disease forecasting services are among many factors to be
considered.
Expenditure on disease control will differ in different regions, depending
on the value of the crop and the number of diseases of economic
importance. In the UK, growers acknowledge the need to buy certified
seed to ensure freedom from virus diseases and wart, to apply fungicides to
control late blight and insecticides to prevent spread of virus diseases. Only
recently, however, have diseases such as gangrene, blackleg, skin spot and
silver scurf received the attention they warrant as major causes of losses in
production, storage, export potential especially of seed tubers or in
presentation quality. In the latter respect the sale of washed, pre-packed
potatoes has elevated some hitherto unimportant blemishing diseases such
as silver scurf and black dot to a completely new status.
This chapter describes how the main potato diseases listed in Table 10.1
affect production, how they are spread and perpetuated, the methods
developed for their control and the possibilities for this to be improved.
10.2 EFFECTS OF DISEASES
10.2.1 Damage to seed tubers and early plant growth

Seed crops can carry a variety of diseases, and in many countries are
inspected during growth and certified for freedom from virus diseases
within pre-determined limits. Such certification rarely guarantees freedom
from tuber-borne bacterial and fungal diseases because the failure to
develop above-ground symptoms (for example, blackleg) is no guarantee
of freedom from pathogens (Graham and Harrison, 1986). Also some
disease symptoms develop only after the tubers have been stored.
Similarly, visual pre-sale examination cannot distinguish the presence of
microscopic infections or inoculum in adhering soil.
Perombelon (1972) found that all Scottish seed stocks examined in
1966-{)8 were contaminated with blackleg bacteria and when Hide (1981)
surveyed stocks of certified Scottish seed of cv. King Edward received by
farmers in England and Wales in 1963-76, 54% of tubers had skin spot,
16% powdery scab, 15% black scurf, 8% gangrene and 2% dry rot. With
some diseases affected tubers were found in all stocks, whereas with
others, most stocks had few or no apparently diseased tubers.
The main causes of seed tuber decay before planting are gangrene
(Phoma foveata, sometimes P. exigua var. exigua) , dry rot (mainly
Fusarium solani var. coeruleum) , late blight (Phytophthora infestans)
and soft rot (Erwinia carotovora ssp. carotovora and E. carotovora
Table 10.1 Diseases of the potato
Symptoms
Causal agent and common disease name Distribution Foliage Tubers
VIRUSES AND OTHER AGENTS
Virus X, mild mosaic Worldwide Mottling
Leaf roll virus Worldwide Rolling
Virus Y, severe mosaic Worldwide Necrosis, stunting, mottling
Virus A, mild mosaic Worldwide Mosaic
Paracrinkle virus Europe, N. America Crinkling
Virus S Worldwide Mosaic
Tobacco rattle virus Europe, N. America, Brazil, Japan Stunting, distortion Internal necrosis
Mop top virus W. Europe, Peru Chlorotic markings Internal necrosis
Yellow dwarf virus N. America Chlorosis, stunting Internal necrosis
Spindle tuber 'viroid' N. America, USSR, S. Africa Stunting Malformation
Witches' broom (mycoplasma-like Europe, N. America, Australasia, Chlorosis, stunting
bodies) China
BACTERIA
Clavibacter michiganensis var. Europe, N. America Wilting Rot in vascular
sepedonicus, Ring rot region
Erwinia carotovora (Jones) Bergey Europe, N. America Wilting Soft rot
et al. ssp. atroseptica (van Hall)
Dye, Blackleg, Soft rot
ssp. carotovora, Soft rot Europe, N. America Wilting (Arizona, USA) Soft rot
E. chrysanthemi Burkholder et al. Tropics, sub-tropics Wilting Soft rot
Blackleg, Soft rot
Pseudomonas solanacearum (E.F.Sm.) Tropics, Warm-temperate zones Wilting Soft rot
E.F.Sm., Brown rot
Table 10.1 Continued
Symptoms
FUNGI
Alternaria solani (Ell. and Mart.) Worldwide Progressive necrosis Rot
Sol., Early blight defoliation
Angiosorus solani Thirum. & O'Brien, Central and S. America Malformation,
Potato smut internal necrosis
Botrytis cinerea Fr., Grey mould Worldwide Decay of senescing stems Rot
Collectotrichum coccodes (Wallr.) Worldwide Premature senescence Superficial
Hughes, Black dot lesions
Fusarium spp., Fusarium wilt N. America Wilting Rot
Fusarium solani var. coeruleum Europe Rot
(Sacc.) Booth, Dry rot
Helicobasidium purpureum (Tul.) Pat., Worldwide Premature senescence Superficial
Violet root rot lesions and rot
Helminthosporium solani Dur. and Europe, N. America Blemish
Mont., Silver scurf
Macrophomina phaseolina (Tassi) N. America, India Stem rot Rot
Goid., Charcoal rot
Polyscytalum pustulans (Owen and N. Europe, N. America, Australasia Non-emergence Blemish, kills eyes
Wakef.) Ellis, Skin spot
Phoma exigua Desm. } { Europe, N. America, AustralaSia}
var. exigua Gangrene Non-emergence Rot
Phomafoveata Foister Europe
Phytophthora erythroseptica Pethybr., Europe, N. America, Australasia Wilting, premature Rot
Pink rot senescence
Phytophthora infestans (Mont.) de Worldwide Progressive necrosis, Rot
Bary., Late blight defoliation
Table 10.1 Continued
Symptoms
Pythium ultimum Trow., Watery wound Worldwide Wet rot
rot
Rhizoctonia solani Kiihn., Black scurf Worldwide Delayed emergence, rolling Superficial
and stem canker sclerotia
Sclerotinia sclerotiorum (Lib.) de Europe, N. America Stem rot
Bary, Stalk break
Spongospora subterranea (Wallr.) Worldwide Galls on roots Superficial
Lagerh., Powdery scab pustules, cankers
Streptomyces scabies (Thaxt.) Waksman Worldwide Superficial
and Henrici., Common scab lesions
Synchytrium endobioticum (Schilb.) Europe, N. and S. America, Deformation of petioles and Deformation
Perc., Wart S. Africa, Asia leaves
Verticillium albo-atrUm) Verticilium Worldwide Chlorosis, premature Vascular browning
R &B wilt, Early senescence
Verticillium dahliae death
Kleb.
subsp. atroseptica) either alone or in combination (Bokx and Mooi,
1974).
Cutting seed, as widely practised in the USA, increases the likelihood of
Fusarium rots, Erwinia soft rots, ring rot (Clavibacter michiganensis
ssp. sepedonicus) and, in the warmer states of the USA, brown rot
(Pseudomonas solanacearum). Sprout growth from seed tubers during
storage and after planting can be adversely affected by diseases. Black
scurf (Rhizoctonia solani) can kill tips of developing sprouts and roots
when seed is sprouted in high humidity and, in cooler potato growing
regions, skin spot (Polyscytalum pustulans) kills buds in the eyes of
un sprouted seed as also can wart (Synchytrium endobioticum) at high
storage humidity. Silver scurf (Helminthosporium solani) and common
scab (Streptomyces scabies) can delay early growth of plants (Adams and
Hide, 1981; Read and Hide, 1984) but usually, despite their prevalence,
tuber yields at harvest are not decreased.
Soil conditions after planting should favour rapid growth so that shoots
emerge quickly and give full plant populations. Gaps (misses) and delay in
emergence are usually caused by rotting of the seed tubers or by failure or
delay in sprouting in cold and dry or waterlogged soils, sometimes by
planting errors and inadvertent use of sprout inhibitors or by bird or animal
damage. Rapid and even emergence is of general importance in potato
production and it is essential for growers of early maturing cultivars to
ensure the early yields that command premium prices. Therefore, in the
UK, the physiological disorders 'little potato', which may be associated
with physiologically old seed tubers or with Rhizoctonia infection, and
'coiled sprout', sometimes associated with Verticillium nubilum (Ali et al.,
1970) or with increased ethylene production (Catchpole and Hillman,
1976) by shoots in compacted soil or after deep planting, can delay
emergence of shoots and formation of tubers.
Most of the diseases of stored seed also affect the establishment and
growth of plants. After planting, whole or cut seed tubers may decay so
rapidly with rots caused by bacteria or Fusarium spp. that no plant is
produced. A less rapid rot may result in death of shoots soon after
emergence. These and other pathogens may also be associated with the
rotting of seed tubers several weeks after plant establishment (Lapwood,
unpublished) although its effect on crop growth is not known. However,
even if shoots are not killed, infection can modify the subsequent growth of
shoots and stolons and the number and size of tubers produced. A severe
attack of skin spot can result in few stems and few but large tubers per
plant (Hide et al., 1973) whereas gangrene stimulates sprouting and
increases stem and tuber numbers (Griffith et al., 1974).
Infection of shoots by Rhizoctonia solani (stem canker) soon after
planting delays stem emergence and development of foliage (Hide et al.,
1985). Later, pruning of stolons removes sinks for photosynthates which
then temporarily accumulate in stems and leaves and in some cultivars
growth of axillary shoots is encouraged, so supplementing leaf area;
sometimes aerial tubers develop in leafaxils. In consequence further
stolons develop and more tubers are initiated.
Compared with many annual crops, potatoes are grown wide apart
within and between rows to give a relatively low plant population,
so failure of plants to emerge causes large and easily observed gaps.
However, losses in yield due to gaps are not proportional to the number of
missing plants because plants bordering gaps are able to benefit from
decreased competition and can compensate to some extent for the 'miss'.
With a single gap, the yield of each of the two adjacent plants within the
row and in neighbouring rows may be increased respectively by 30% and
5% so that overall, the compensatory growth from these four plants can
account for 70% of the potential yield of the missing plant. With larger
gaps (two or more adjacent plants missing) production by neighbouring
plants may be larger than for single plant gaps but the amount of
compensation will be proportionally less.
Compensation or yield loss will be influenced by plant spacing, the time
when gaps occur and by the vigour of surviving plants. Thus, significant
yield losses were recorded only when at least 15% plants were missing at
emergence, or when at least 6% plants were removed at flowering time to
simulate plant death caused by blackleg (Hirst et al., 1973). But disease
also weakens plants and such plants are then overgrown by healthy plants
which suppress their growth and yield (Table 10.2).
Table 10.2 Effect on plant yield (kg) when neighbours were

healthy, diseased or missing
Neighbouring plants
Healthy Diseased Missing SED
Blackleg field trials
Healthy plants 1.59 1.84 2.41 0.099
Diseased plants 0.78 1.06 1.42 (50 DF)
Stem canker field trials
Healthy plants 1.59 1.80 2.55 0.021
Gangrene field trials
Healthy plants 1.63 1.90 2.80 0.016
Data from Adams and Lapwood (1983) and Hide and Read (unpublished).
10.2.2 Stunting and premature death of foliage
(a) Infection with viruses

Plants can become systemically infected with viruses following transmission
either mechanically or through vectors and, unlike diseases caused by
bacteria and fungi, once infection has become established development of
disease within the plant is not greatly affected by weather conditions.
Usually viruses decrease plant vigour by shortening internodes, and cause
colour changes in foliage which becomes mottled or chlorotic. Yields are
decreased and some viruses cause internal tuber symptoms which cannot
be seen until the affected tuber is peeled. Crops affected in this way may be
rendered unsaleable even though a relatively small proportion of tubers
are affected. Some viruses may rapidly kill plants whereas others cause
mild or no symptoms and the reaction of different cultivars to the same
virus can vary so that symptoms alone may not be a reliable guide to the
virus involved.
With some virus diseases, symptoms can be quite different in the year
when plants become infected (primary symptoms) from those showing in
plants derived from infected seed (secondary symptoms). For example
virus Y, transmitted by Myzus persicae and other aphids, may cause near
complete defoliation (leaf drop streak) within 3-4 weeks of infection and
yields are decreased according to the time and rate of defoliation. In the
following season plants from infected seed are severely stunted, mottled
(severe or rugose mosaic) and die early often giving only half the yield of
uninfected plants and this composed of a high proportion of small tubers.
The primary symptoms of some virus infections are mild or, if infection
occurs late in the season, plants show no symptoms, but even so a
proportion of tubers from them when planted as seed will produce plants
with severe symptoms in the following season. Plants infected during the
growing season with leaf roll virus transmitted by aphids, and potato
yellow dwarf virus transmitted by leaf hoppers, may show merely slight
rolling of upper leaves, often only on one stem. In the next season,
however, leaf roll infected tubers produce stunted plants with rolled lower
leaves. These are thickened and brittle with accumulation of carbohydrates
and are characteristically harsh and rattle if shaken. Aerial tubers may
form in leafaxils and yield is halved. Secondary symptoms of leaf roll
and virus Y also develop on plants following previous infestation by
viruliferous aphids on their sprouts before planting. Seed infected with
yellow dwarf virus produces chlorotic and stunted plants and sometimes
the growing point is killed; in warm soil tubers may fail to sprout or plants
die soon after emergence. Any tubers that are produced are small and mis-
shapen and, because rust coloured lesions develop in the pith, quality is
also affected.
Two other diseases that cause severe secondary symptoms are witches'
broom and spindle tuber. Plants produced from tubers infected with
witches' broom, caused by mycoplasma-like organisms, are slow to emerge
and have many weak stems bearing simplified pale-edged leaves. Spindle
tuber is caused by a virus-like particle comprising only RNA ('viroid') and
infected plants remain erect but with rugose leaves and the few tubers
produced are cylindrical or irregular in shape. The viroid can be trans-
mitted in true seed (Fernow et al., 1970) and therefore can be a problem to
the potato breeder.
Some virus diseases produce only mild symptoms or none at all in both
seasons. Virus X, transmitted mechanically by leaf contact, machinery or
on clothing, may produce no symptoms in some cultivars whereas in others
it causes an interveinal mottle but without visibly affecting plant vigour,
although yield may be decreased. Plants infected with virus A, transmitted
by aphids, show mild symptoms but in association with virus X or Y cause
crinkle symptoms. Paracrinkle and virus S, both transmitted by aphids,
produce such slight symptoms in plants that their prevalence in some
cultivars remained unrecognized until plants were freed from them
(Bawden and Kassanis, 1965).
Virus diseases decrease yield but as infected plants are usually scattered
through the crop, neighbouring healthy plants can compensate in growth,
as discussed in Section 10.2.1. For example, in three crops with 5-15,15-25
and 25-50% of plants affected with secondary symptoms of leaf roll and
virus Y, yields were decreased respectively by 2, 7.5 and 16% (Schick,
1952). However, the amount of compensation may be less than expected if
infection spreads from diseased to neighbouring healthy plants especially if
the spread occurs early in the season. Also, when several diseased plants
are together, they too can benefit from decreased competition, so calcula-
tion of losses in yield from crops with known proportions of diseased plants
is complex (Reestman, 1970).
(b) Destruction of foliage

Early blight (Alternaria solani) and late blight (Phytophthora infestans) are
probably the most important fungal foliage diseases of potatoes. Both are
dependent on weather conditions for sporulation and spread and the
amount of yield loss they cause depends on how early and quickly they
destroy the foliage. Late blight can attack and destroy foliage of develop-
ing plants at any stage of growth given suitable weather. In Mexico
where crops are planted in anticipation of the rainy season, unprotected
susceptible European cultivars are attacked soon after emergence, yielding
little or nothing (Lapwood, 1971). At the other extreme, as in much of the
UK, blight does not become epidemic until the canopy closes across rows
allowing the development of a suitable micro-climate for the disease to
spread. The fungus sporulates mainly on the undersides of leaves in a band
around the edge of the necrotic lesions and the sporangia may be released
into the air or splashed in water droplets during rain to other leaves.
Persistent leaf wetness is needed for successful infection and epidemics
develop only in wet weather or following heavy dew or mist and progress
most rapidly when rain is frequent. Because it is a disease which mostly
develops from discrete lesions, the rate of foliage destruction depends on
how soon every leaf becomes infected, although in some cultivars and in
some seasons stem infections can be common, resulting in death of the
stems distal to the girdling lesions.
Young foliage seems temporarily resistant to early blight. In Colorado,
USA, first (primary) infections produced from spores in soil are few and
develop on the lower leaves in June and provide the spores for secondary
spread which may be delayed, however, for as long as 2 months. Spore
production is increased by prolonged periods of high humidity and
maximum numbers develop at 20--25°C. In India the disease is favoured by
short photoperiods (Rambawale and Bedi, 1982), and when infection does
occur it can destroy the foliage very rapidly (Harrison et at., 1965)
especially in crops given low or moderate amounts of nitrogen fertilizer
(Mackenzie, 1981). Yields may be decreased by up to 40% (Harrison and
Venette, 1970), although in areas where the disease is severe, crops may
further be affected by wilting and early death caused by Verticillium spp.
The growth of tubers is dependent on the functioning leaf area, so
diseases that defoliate plants have serious effects on yield. The progress
curve for a late blight epidemic, expressed as the proportion of leaf area
destroyed, is essentially of the sigmoid type, and Large (1952) found in
England and Wales that tuber growth stops when 75% of the leaf area is
destroyed. From data on the progress of late blight in fungicide-sprayed
and unsprayed crops, reasonably reliable estimates of the percentage loss
in yield can be made by reference to their bulking curves.
In Canada where the growing season is 40 days shorter than in the UK,
this method seriously under-estimated losses (James et at., 1971), and a
method based on weekly increments of foliage destruction, with epidemics
classified as 'early' or 'late' depending on when they occurred in relation to
crop development, was found more reliable (James et at., 1972). More
recently, computer programs, e.g. Blitecast, in the USA (MacHardy,
1979) have been developed as an aid to forecasting the progress of
epidemics.
(c) Wilting and premature death of plants

Several pathogens may cause wilting symptoms because they infect roots
and invade the vascular system or produce toxins. Water supply to the
foliage is restricted and typically leaves become chlorotic and sometimes
roll before dying. If infection occurs late, symptoms may be confused with
and by senescence.
Of the bacteria attacking potato plants, three cause diseases which are of
major economic importance: Erwinia carotovora ssp. atroseptica the
common cause of blackleg, Clavibacter michiganensis ssp. sepedonicus
causing ring rot and Pseudomonas solanacearum causing brown rot. All are
tuber-borne and some affected tubers give rise to diseased plants. E.
carotovora ssp. atroseptica produces a necrotic rot on stems extending from
the seed tuber up to and, in very wet weather, above soil level. Leaves turn
yellow and leaflets roll upwards but not all stems on a plant may be
affected. In Arizona E. carotovora ssp. carotovora has been found to
cause blackleg symptoms (Stanghellini and Meneley, 1975), as does E.
chrysanthemi in warmer climates. Infection with Clavibacter also causes
yellowing and rolling of leaves early in the season in warm areas, although
in Europe symptoms develop later and can be confused with late blight or
senescence. Similarly infection with Pseudomonas results in wilting of
plants but like Clavibacter does not cause a 'blackleg' symptom.
Infection by fungi, including species of Verticillium and Fusarium and
Phytophthora erythroseptica can also cause wilting and early death.
Fusarium spp. mostly cause wilting in warm climates following infection of
roots or decay of seed tubers before plants are fully established. Verticillium
spp. infecting through roots grow into the vascular system and decrease
leaf area (Harrison and Isaac, 1969). Later, lower leaves become unilater-
ally chlorotic and symptoms progress up the stem until the plant becomes
desiccated and dies. Crop yields can be halved. Severity of the disease is
increased by infestation of roots with nematodes including Pratylenchus
penetrans, Meloidogyne hapla, Globodera rostochiensis and G. pallida
which seem to aid infection of roots (Morsink and Rich, 1968; Hide et al.,
1984; Evans, 1987). In N. America the disease can be caused by V.
alboatrum and V. dahliae (Busch, 1967) whereas in the UK, V. dahliae is
the predominant species (Isaac and Harrison, 1968).
Several other pathogens can cause wilting and death of foliage. In wet
areas, stems may be infected by Sclerotinia sclerotiorum (stalk break)
causing leaf chlorosis and death of stems, which bear sclerotia in the stem
cavity. Botrytis cinerea (grey mould) usually infects leaves but may
progress along petioles to infect stems late in the growing season, causing a
soft wet rot. Occasionally injured tubers are attacked. Colletotrichum
coccodes, a weak parasite, can infect roots and stems causing early death of
foliage with development of sclerotia on dead tissues (black dot) and plants
are most susceptible to infection when pre-disposed by factors such as
moisture stress. The fungus may also prune stolons and blemish tubers
(Thirumalachar, 1967; Read and Hide, 1988).
10.2.3 Infection of tubers during growth

Progeny tubers may be attacked during growth by bacteria and fungi which
gain entry through lenticels and growth cracks, unsuberized tissue around
eyes, insect wounds, intact skin or via the stolons. After infection, tubers
may rot in soil, become deformed or blemished or the pathogens may
remain latent to cause disease later. Alternatively, inoculum present at the
tuber surface or in the surrounding soil infects only when tubers are stored
in conditions favourable for infection or are damaged at or after lifting.
Amounts of infection are often related to the prevalence of disease on the
seed tubers.
(a) Rots
Tubers that are rotted by fungi usually remain intact unless they are
invaded by secondary organisms. Sporangia of Phytophthora infestans
washed down from diseased foliage can infect tubers through lenticels,
eyes or growth cracks to produce the typically hard brown marbled
progressive rot. Macrophomina phaseolina infects through lenticels, eyes,
stolons and through wounds made by larvae of the tuber moth (Phthorimaea
operculella) to cause black sunken lesions and later blackening of internal
tissues. Phytophthora erythroseptica gains entry through stolons causing a
wet but firm rot progressing from the heel towards the rose end of the
tuber. Infection can occur at the 'eyes' and oospores may be found in
surface tissues of symptomless tubers (Lonsdale et at., 1980). When lifted,
infected tubers frequently have small clumps of soil adhering to the skin
where moisture has exuded from lenticels. The cut surface characteris-
tically turns a bright salmon pink when first exposed to the air, giving the
disease its common name, pink rot. Occasionally the reddish brown
mycelium of Helicobasidium purpureum (violet root rot) is found on
tubers. The fungus penetrates the skin and causes a soft rot.
Unlike fungi, bacteria infecting tubers through lenticels and stolons
generally cause complete disintegration of affected tissues resulting in soft
mushy rots. All Erwinia spp. can cause a general decay of tissues.
Clavibacter attacks tissues of the vascular region, in which cavities develop
causing typical 'ring rot' symptoms. These may later be masked by
secondary bacteria invading other tissues and causing complete disintegra-
tion of tubers. Pseudomonas solanacearum also infects the vascular
elements which are stained brown (brown rot) and liquid can be squeezed
from infected tubers.
(b) Blemishes
Several diseases deform or blemish the tuber skin or flesh and so decrease
quality and saleability of crops. In wet soil, Synchytrium endobioticum
penetrates eyes to form warty outgrowths that disfigure tubers. Spongospora
subterranea infects lenticels in similar conditions and development of
powdery scabs containing resting spores (spore balls) occurs when soil
gradually dries. In continuing wet conditions resting spores may germinate;
the fungus penetrates deeper into tuber tissue resulting in the development
of tumour-like outgrowths which also form if eyes become infected. The
actinomycete Streptomyces scabies (common scab) causes little malforma-
tion. Infection of the lenticels of developing tubers gives rise to surface
scabs which enlarge so long as the tubers swell, but cause increasingly deep
and disfiguring lesions if successive attempts by the tuber to form a wound
barrier fail. Dry weather favours infection and, when prolonged during
early tuber formation, encourages severe attacks that can affect a large
proportion of the tuber surface area (Lapwood and Hering, 1970) and so
decrease saleability. Early infection of tubers by Rhizoctonia solani
sometimes causes blackening of the skin; these lesions break up as tubers
swell but tubers are often malformed. Most black scurf (sclerotia) forms as
the foliage senesces or following mechanical or chemical destruction, and
its severity increases as long as tubers remain in soil.
Tubers may become systemically infected with the viruses that infect
foliage but usually they show no symptoms. A few viruses, however, do
cause internal tuber symptoms which can be particularly insidious because
they are not seen until the tubers are prepared for cooking. Consequently a
small proportion of affected tubers can render a whole crop unacceptable.
The viruses capable of tuber damage include potato yellow dwarf (Section
10.2.2), potato mop top transmitted by the zoospores of the powdery scab
fungus Spongospora subterranea, and tobacco rattle transmitted by ecto-
parasitic nematodes of the genus Trichodorus. The last two viruses cause
arc-shaped rusty-brown necroses of the tuber flesh (spraing) which are
occasionally visible on the surface of the tuber; yields can be decreased by
25% or more.
(c) 'Latent diseases'

During growth, tubers can be infected or contaminated by bacteria and
fungi that may not cause disease until after crops are lifted. Bacteria may
penetrate lenticels or through stolons but cause rots only when tubers are
stored or planted. Spores of Helminthosporium solani (silver scurf)
produced on seed tubers cause symptomless infection of young tubers,
firstly at the stolon end; and Polyscytalum pustulans (skin spot) from
lesions on seed tubers, roots, stolons and stem bases infects scale leaves
and buds of tuber initials and young tubers. Sometimes silver scurf and skin
spot symptoms are seen on tubers at lifting but mostly these diseases
develop in store. Similarly the intensity of black dot can increase after
harvest. Phoma Joveata (gangrene) from soil, decaying seed tubers or from
pycnidia on dead stems may occasionally invade tubers before lifting but
more often, like Fusarium spp., spores contaminate the tuber surface and
surrounding soil and do not infect until tubers are injured at or after lifting.
Irrigation increases tuber infection by Erwinia, Polyscytalum, Phoma and
Colletotrichum, probably by increasing dispersal of the pathogen, but tends
to decrease infection by Helminthosporium (Adams et al., 1987).
10.2.4 Infection of tubers at lifting
When potato crops are lifted, tubers are particularly at risk from the
bacteria and fungi present at the tuber surface, in the surrounding soil, or
harboured in the decaying foliage and decaying seed tubers. Wounds
inflicted by stones and machinery may make tubers unsaleable but more
importantly may provide sites for infection by pathogens. In the UK almost
one-quarter of all tubers from 600 farms surveyed in 1965 and 1966 had
wounds penetrating several mm into the flesh (Church et al., 1970). Some
wounds, such as cuts, heal quickly and completely with the formation of a
cork barrier, but crush wounds are more slowly and often less completely
healed and are therefore more likely to become infected. Wounds are most
severe when lifting from dry stony soils and in dry seasons, especially if
tubers are immature, whereas moist soil cushions tubers and prevents
severe mechanical damage on the harvester web. Much fertilizer can delay
maturity and increase the incidence of mechanical damage, and so increase
the incidence of diseases that result from infection of wounds.
Immature tubers are prone to infection by Fusarium spp., Phoma
foveata, Pythium ultimum (watery wound rot) and Alternaria solani (Boyd,
1972; Venette and Harrison, 1973) but their resistance later increases and,
in Scotland, wounded tubers are most resistant to infection by F. solani
var. coeruleum when fully mature; tuber resistance is readily induced
by destroying foliage. Phoma foveata and Macrophomina phaseolina
(charcoal rot) can also infect wounds made during lifting, and tubers can be
infected by Phytophthora infestans from inoculum in soil and on foliage
but, because sporangia are short-lived, delaying harvest for 2 weeks after
foliage destruction can minimize this risk.
10.2.5 Post-lifting disease

Deterioration may start as soon as crops are lifted, during transport or at
the retail outlets. Early marketing can avoid the risk of loss in store but
some crops must be stored to ensure continuity of supply for the fresh and
pre-pack trade and for processing.
(a) During transport and at retail outlets

Most loss follows warm and humid conditions at or soon after lifting,
because they favour rapid rotting by the soft rot bacteria and fungi Pythium
ultimum and Phytophthora erythroseptica. These problems are usually
more acute in immature crops and are often associated with poor ventila-
tion in transit (Cromarty and Easton, 1973). Secondary rotting by bacteria
can also affect tubers infected with late blight. At retail outlets tubers
offered for sale are also at risk especially if they have been cleaned by
brushing or washing and packed in poorly ventilated plastic bags. These
conditions encourage bacterial rotting especially of immature tubers.
Susceptibility is decreased by storage before washing.
(b) In store
Most mature crops are kept in store initially in conditions that encourage
wound healing and thereafter in an environment that minimizes water loss
and sprouting. The method of storage varies, according to the available
facilities and to the degree of environmental control required (this volume,
Chapter 14), from clamps in the field insulated with straw and soil, to
improvised buildings or purpose-built stores with facilities to monitor and
control the environment. Disease development is greatly influenced by
storage conditions (Boyd, 1972) and, because crops for processing or for
sale as table potatoes or as seed may require different environments, so
different diseases may become predominant.
In stores kept cool (2-5°C) a major rotting disease is gangrene (Phoma
foveata) which develops on wounds that have not healed (Adams and
Griffith, 1978). Externally, lesions show as dark, dry spreading depressions
and cavities may form within tubers often much larger than the visible
symptoms would suggest. Dry rot (Fusarium spp.) is favoured by higher
temperatures and can be arrested in cool (5°C) stores but re-activated on
return to warm conditions. Under drying conditions rots gradually lose
moisture and become shrunken, developing concentric wrinkles round the
site of infection on which cushions of spore-bearing mycelium develop,
while internally the affected tissue is usually light brown. In damp
conditions the rots advance more rapidly and, as with tubers affected by
late blight, secondary organisms can hasten the decay and sometimes
initiate massive soft rotting. Susceptibility of tubers to infection increases
during storage and, like gangrene, much dry rot can develop from wounds
inflicted when tubers are sorted after storage.
Bacteria, either in wounds or lenticels or following fungal infections, are
the main cause of rapid deterioration in warm storage and especially if
crops go into store wet. Rotting of individual tubers may begin soon after
loading and, if conditions are favourable, moisture exuded from these will
affect neighbouring tubers leading to local pockets of rotting tubers in
which temperatures rise, the environment becomes increasingly anaerobic
favouring Clostridium spp. (Campos et al., 1982) and accelerating the rate
of spread through the bulk. If conditions become cooler and drier,
infections developing at lenticels may be arrested and affected tissues
collapse causing spots, a form of 'pit rot', or larger superficial 'hard rot'
lesions.
Of the blemishing diseases, skin spot can become particularly severe in
cool (2°C) humid stores. Symptoms usually develop 2-3 months after
lifting and subsequently the number of spots increases and buds in the
'eyes' of affected tubers may be killed. Infection occurs through 'eyes',
Survival and spread of pathogens 419
lenticels or skin abrasions and the disease is often prevalent in crops lifted
in wet cool conditions. The number and depth of skin spots can be
increased by the sprout suppressant chlorpropham, especially when
applied too soon after store loading. Humid storage encourages the
formation of wart, the development of tumours on tubers with powdery
scab and the superficial tuber to tuber spread of Phytophthora erythrosep-
tica mycelium. Similarly, warm and damp storage encourages the develop-
ment of Rhizoctonia solani sclerotia (black scurf). Silver scurf lesions
spread over the surface of tubers and conidia are released into the store
environment; these and the conidia of Polyscytalum pustulans (Carnegie et
al., 1978) spread the disease to other tubers.
Amounts of disease can increase after tubers are removed from store.
Tubers with soft rot disintegrate on the grader and can spread inoculum to
other tubers and similarly fungal inoculum in adhering soil may spread
from tuber to tuber. Gangrene and dry rot develop at fresh wounds which
on stored tubers are especially susceptible.
10.3 SURVIVAL AND SPREAD OF PATHOGENS
There are a number of ways in which potato pathogens survive but all
require dispersal to infect succeeding crops. Some are associated with soil
and can be regarded as soil-borne. Usually these spread only slowly, unless
introduced with infected seed tubers. However, they may survive for long
periods by forming thick-walled resting structures, e.g. Spongospora
subterranea whose spore balls may remain viable in soil for more than
10 years, and Synchytrium endobioticum resting sporangia for at least 30
years. Also found in soil are the oospores of Phytophthora erythroseptica
and P. infestans, the latter originally thought to occur only in Mexico but
now found in Europe (Tantius et al., 1986), the chlamydospores of
Fusarium spp., and sclerotia of Rhizoctonia solani, Sclerotinia sclerotiorum
and Verticillium dahliae. A few pathogens survive saprophytically in soil
in the absence of potatoes or potato crop residues. For example,
Streptomyces scabies is commonly found in soils under grass while
Rhizoctonia solani and Helicobasidium purpureum are also soil inhabit-
ants, although they can infect crops other than potatoes. Colletotrichum
coccodes can survive in soil for at least a year after decomposition of crop
residues and Phoma foveata for at least 7 years after potato crops but in
what form is not known. Phytopathogenic bacteria do not form resting
spores and seldom survive for long in soil. An exception is Pseudomonas
solanacearum which can survive in the deep soil layer (Graham and Lloyd,
1979) for 4-6 years in bare fallow and for up to 10 years in soils in non-
susceptible plants (Schuster and Coyne, 1975). The viruses causing spraing
can survive in the soil in their vectors, the fungus S. subterranea and
nematodes of the genus Trichodorus.
Other pathogens survive only in infected plant remains, and so the
length of survival of Alternaria solani, Polyscytalum pustulans and Erwinia
carotovora ssp. depends on the rate of decomposition of the debris.
Many of the above pathogens and also Clavibacter michiganensis ssp.
sepedonicus (Nelson, 1984) and Phoma foveata (Carnegie et al., 1978) can
be found contaminating planting and harvesting implements and in the
dust of potato stores.
Many pathogens survive from season to season in infected tubers, which
may be lifted and used as seed, or remain in the soil unharvested and
perpetuate infection on volunteer plants. Most viruses survive in tubers
which produce systemically infected foliage; an exception is tobacco rattle
virus which is seldom transmitted from infected seed tubers. Lesions on
seed tubers or soil adhering to them can be sources of bacterial and fungal
inoculum for infecting the resulting crop and for introducing diseases into
new potato growing areas.
Most bacterial diseases originate from seed tubers as do skin spot and
silver scurf, and discarded tubers affected with late blight may produce
infected stems and foliage which constitute primary sources of infection
each year. Plants other than potatoes may also become infected with
potato pathogens but, although it is not known how widespread these are
as sources of inoculum or as means of survival, they are likely to be most
important in seed-producing areas.
For the perpetuation of diseases, pathogens require not only means of
survival but also effective methods of spread. Pathogens on seed tubers
may be released into soil as infected tubers rot (Phoma foveata, Fusarium
spp. and the bacterial pathogens), as spores produced on tuber lesions
(Helminthosporium solani, Polyscytalum pustulans) , or mycelium may
grow out into the soil, e.g. from sclerotia (Rhizoctonia solani). Dissemina-
tion may be assisted by water, by soil fauna or the movement of soil by
implements or wind. The presence of growing roots or tubers can stimulate
the release of motile zoospores which are able to move in soil water films
from the resting spores of S. subterranea and probably of S. endobioticum.
Above ground, pathogens are spread by airborne spores, in or on insect
vectors or by implements. Spread of airborne fungal spores and subsequent
infection may require specific weather conditions: for example, sporangia
of Phytophthora infestans can be washed or splashed in rain on to
neighbouring plants or down to the tubers where they require a water film
in which to germinate. Spores of Alternaria solani in soil are blown or
splashed in heavy rain on to leaves to initiate primary lesions, which
produce spores in dew or following rain or irrigation. The spores are
released and spread by wind when leaves dry. Rain releases spores from
pycnidia of Phoma foveata on dead stems and washes Erwinia bacteria
from blackleg stems to tubers in soil. Both pathogens can be spread by
wind within and between crops in rain-generated aerosols (Quinn et al.,
1980) and also aerosols of sap may be produced when haulm is pulverized
Control of diseases 421
prior to harvest (Perombelon et al., 1979; Carnegie et al., 1987). Erwinia
spp. are also spread by insects attracted to blackleg stems (Kloepper et al.,
1979) and by tractors and implements contaminated by passing through
infected crops.
The amount and rate of spread of aphid-transmitted viruses depends on
the build-up of the vector populations, the stage of development of the
crop when first infested and vector movement within and between crops
(Broadbent, 1953). For example, in the south of England early infestations
of Myzus persicae carrying virus Y can almost destroy crops, and early
infection with leaf roll may result in the appearance of secondary
symptoms in the same season. There can be a 50-fold incre.ase in the
number of infected plants in one season. Older plants resist infection so
that the later the aphids arrive in crops, the less damaging are these
infestations. Usually the number of plants infected with leaf roll and virus
Y increases by up to lO-fold in one season: for example, in a stock of Ulster
Prince grown for three successive seasons, the incidence of leaf roll was
respectively 0.2,1.1,7.4% and of virus Y, 0.2, 0.3 and 8.8% (Broadbent et
al., 1960). The amount of spread may also be affected by the behaviour of
virus in aphids. For example, virus Y is readily transmitted soon after
acquisition but is non-persistent, the ability to transmit being restricted to a
small number of feeding probes, and therefore spread is limited. However,
leaf roll virus requires an incubation period in the aphid before transmis-
sion can be effected, but is then persistent so that it can be spread widely
following flights between different crops.
Seed tubers infected with viruses usually produce diseased plants, but
not all the progeny tubers of newly infected plants are necessarily diseased
because, perhaps, only one stem of the several independent stems com-
prising a plant, and the tubers attached to it, are infected. Usually infected
plants produce a larger proportion of small tubers. Consequently, infected
tubers are concentrated in the seed-size fraction, so if a stock is grown in
successive seasons the proportion of infected plants can increase in the
absence of vectors. Crops become less vigorous and yields decline progres-
sively (previously termed 'degeneration' or 'running out'). However, some
viruses infecting very susceptible cultivars kill plants before tubers form
and therefore tend to be self-eliminating (e.g. potato yellow dwarf).
10.4 CONTROL OF DISEASES
A knowledge of the etiology of a disease and of the methods of crop

production are essential if the disease is to be prevented or its severity and
effects minimized. The control measures employed will often depend on
eliminating, or severely restricting, the most important sources of inocu-
lum. Some diseases can be avoided by planting disease-free seed tubers,
planting into pathogen-free soil or by legislation preventing introduction of
new diseases. Eradication of pathogens may be possible by treating seed
tubers or soil, by removing alternate hosts and infected host tissues, or by a
rotation of crops. Crops may be protected by modifying their environment,
e.g. with irrigation, or by the application of chemicals that prevent
infection or control vectors. Development of disease in tubers may be
prevented or restricted by modifying the storage environment. Probably
the easiest method in practice is to use resistant cultivars and thereby avoid
the necessity of other measures.
10.4.1 Cultural (non-chemical) control

Non-chemical control usually means modifying cultural practices to make
the whole environment as unsuitable as possible for the disease in all its
phases, by decreasing inoculum available to the crop, e.g. by cultivations
to eliminate volunteer plants, as well as restricting spread during growth or
storage by various means. Diseases can sometimes be avoided by wise
choice of site, by the rotation of crops or by improved soil drainage, or
their effects ameliorated by planting a mixture of local cultivars, as
practised by Andean farmers to counter the range of indigenous diseases.
Severity may be lessened by changing planting or lifting dates or depth of
planting and adjusting temperature and humidity in store to encourage the
healing of wounds and thus decreasing the rate of disease development.
Attempts have been made to forecast the risk of disease development in
store from assessments of infection or inoculum on crops during growth
(Adams et al., 1980), and to provide an early warning of rotting in store by
detecting changes in volatile compounds in the storage atmosphere
(Waterer and Pritchard, 1984).
Much mechanical damage on tubers is caused by lifting from cold, dry
and stony soils, providing opportunity for wound infections to occur.
Tubers may also be damaged during their removal from store and this may
be diminished by correct adjustment and protection of machinery and by
allowing tubers to warm up before mechanical handling.
(a) Management of the storage environment

Modifying the conditions in which tubers are stored can prevent diseases
developing, although with some diseases, control by this method is
only palliative, the pathogen subsequently resuming its progress if the
environment becomes suitable. Minimizing mechanical injury, apart from
decreasing direct losses, is an important measure of control for some
diseases but, since injuries can never be avoided entirely, the provision of
conditions that favour rapid wound healing offers a useful additional
means of control. In the UK, 'curing' for 10--14 days at 15°C immediately
after loading into store decreases the amount of gangrene that develops
later in cooler conditions. Cut wounds heal more quickly than more
complex injuries (e.g. crush wounds), but disease control is less effective if
the start of curing is delayed for several days.
The development of bacterial rotting soon after tubers are put into store
may be favoured by warm 'curing' conditions, especially if tubers carry
surface moisture from soil, or from exposure to rain during lifting or
transport to the store. Condensation on tubers, which occurs when
ambient temperatures fall below that of the stack, can be avoided by a
thick straw covering or quilt by way of insulation; any condensation then
forms in this layer, keeping the tubers dry. But curing is unlikely to affect
the survival of some pathogens present on the tuber surface so that tubers
injured later, as during quality or size grading before sale, may become
infected unless they are cured again. However, if humidity is low (c. 80%
RH) during curing, superficial infections can be eradicated and sub-
sequently much less disease develops in the humid conditions of long-term
storage. The incidence of diseases such as skin spot (Hide and Cayley,
1987), silver scurf and black dot may be decreased in this way.
(b) Growing conditions

Diseases resulting from soil-borne inoculum are among the most difficult to
control and complete prevention is seldom achieved. Rotation with non-
susceptible crops can decrease early death of plants caused by Verticillium
albo-atrum (Emmond and Ledingham, 1972) but not when it is caused by
V. dahliae (Powelson, 1964). Soil-inhabiting Rhizoctonia solani, including
strains non-pathogenic to potatoes but able to form sclerotia on the tuber
surface, increases with the frequency of potato crops. Soil infested with
oospores of Phytophthora erythroseptica requires a rest from susceptible
crops of 4 years to ensure comparative freedom from pink rot, whereas
with powdery scab and wart, effective rotations are unlikely to be
practicable.
Flaming infected stems before lifting decreases Verticillium inoculum in
soil and increases yield of subsequent crops (Easton et al., 1972) but, like
rotations, the method is unlikely to be successful when the soil population
is already large, even if it proves practicable. Improving drainage promotes
rapid growth after planting and thereby decreases the chance of pink rot,
powdery scab and the bacterial diseases that are encouraged by water-
logged soil, and eliminating potato volunteers could decrease inoculum of
viruses, bacteria and fungi that are harboured in tubers.
Liberal applications of inorganic manures, especially N, delay the
development of symptoms of Verticillium spp. and Alternaria solani so that
growth of the crop is prolonged and yields are improved (Easton, 1970;
Barclay et al., 1973). However, an imbalance in fertilizer can affect the
severity of foliar diseases and render the tubers more liable to mechanical
injury and rotting by Phoma and Fusarium spp. Excessive N increases the
speed of foliage destruction by late blight although this can be partly offset
by increasing phosphate. With early blight both the efficiency of control by
chemicals applied to foliage and yields are decreased when excess N is
given (Soltanpour and Harrison, 1974). Organic manuring can decrease
pink rot and also common scab which can be severe in the first crop after
long-term grass and especially on light sandy or gravelly soils and on land
recently limed.
Common scab is also effectively controlled by irrigation provided this
achieves a soil water potential of not less than -0.4 bar (pF 2.6) for at least
6 weeks after tuber initiation (Lapwood et al., 1973). However, over-
watering or starting applications early relative to crop development, may
increase the incidence of powdery scab (Taylor et al., 1986). When soil
populations of S. scabies are large or watering impracticable, the disease
may be controlled by chemicals or combination of both treatments (Davis
et al., 1974). Irrigation decreases charcoal rot indirectly by lowering soil
temperature and Verticillium by encouraging anaerobic decomposition of
organic residues containing the pathogen (Watson and Huber, 1971). It
may, however, increase the incidence of late blight, blackleg and other
tuber diseases.
Spread of aphid-borne virus diseases can be decreased by roguing out
diseased plants and, as young plants are more susceptible than mature
plants, this must be done as soon as they are recognizable. Also, spread
may be prevented by covering growing crops with plastic sheet or coarse
nets (Cohen, 1981; Gibson and Gunenc, 1981).
Delaying planting until the soil becomes warm and planting shallowly
with sprouted seed tubers decreases the risk of seed tuber decay by
bacteria and Fusarium spp., and encourages development of shoots from
tubers affected with skin spot. Also, damage by Rhizoctonia solani on
stems (stem canker) is largely prevented and coiled sprout decreased.
Deep planting may increase stem canker, but a tall pointed ridge is best for
preventing greening of tubers and infection by late blight. Coiled sprout is
associated with the planting of seed with over-long sprouts and 'little
potato' with planting into cold soil immediately after desprouting and so
avoiding such practices will largely overcome these problems.
The amount of tuber infection is sometimes affected by lifting date, but
different diseases are affected in different ways. For example, R. solani
and Polyscytalum pustulans infect tubers increasingly during the growing
season and the diseases they cause, black scurf and skin spot, are decreased
by early lifting. Lifting soon after foliage destruction can decrease the
prevalence of gangrene (Logan et al., 1975) probably by avoiding the
accumulation of P. Joveata in soil. In India, lifting early before the ambient
temperature rises at the end of the growing season decreases charcoal rot.
By contrast, tubers may have little silver scurf when lifted early but high
ambient temperature in the first weeks of storage encourages its spread so
that the disease can be more severe than on tubers lifted later. With
Alternaria solani, although the amount of inoculum on foliage increases
towards the end of the growing season, this is accompanied by increasing
tuber resistance, so that crops lifted late may develop least disease.
Similarly, as tubers mature, their resistance to infection by Fusarium solani
var. coeruleum increases and this can be induced by destroying foliage
prematurely. Provided lifting is delayed to allow tuber skin to 'set', this
practice offers a means of control, but sometimes at the expense of yield.
Low storage temperatures, imposed to avoid sprouting and water loss,
encourage the development of gangrene and skin spot. Development of
severe silver scurf, charcoal rot and Fusarium rots is largely arrested by
these conditions but may recommence when tubers are removed to warmer
conditions; consequently in India charcoal rot is encouraged to develop by
storing seed crops warm for 2-3 weeks so that affected tubers can be seen
and removed when the crop is size graded.
When seed tubers are dried after lifting and stored in boxes or trays,
sporulation and spread of pathogens is prevented and if the temperature is
raised sprouting of the seed is encouraged. With tubers removed from bulk
stores and stored dry and warm, skin spot and damage to eyes from P.
pustulans will be minimized. Dry conditions may also decrease infection of
wounds but are unlikely to prevent gangrene once established. Heating can
rid tubers of late blight, skin spot, gangrene and silver scurf (Fox et al.,
1970; Hide, 1975), and recently techniques have been developed for
controlling Erwinia infection by hot water treatment (Wale et al., 1986).
Similarly, heating tubers will eradicate viruses X, Y, M, S, A and leaf roll
but this method is only likely to be used to control viruses in nuclear seed
stocks.
(c) Other cultural methods

Some viruses including X, Y, A, Sand paracrinkle can be eliminated by
growing plantlets from excised apical meristems on nutrient agar (Kassanis
and Varma, 1967) and also spindle tuber viroid can be eradicated provided
mother plants are grown in cool conditions (5-8°C) (Lizarraga et al., 1980).
These methods are useful when all tubers of a stock are infected and in the
UK, the cultivar King Edward, totally infected with paracrinkle, was
completely replaced with virus-free seed stocks in Scotland 11 years after
the first meristem culture (Bawden and Kassanis, 1965).
Larger portions of stems (stem cuttings) or nodal segments (micro-
propagation) can be rooted to produce plants free from some bacteria and
fungi. The method is useful for producing nuclear clones free from diseases
that are tuber-borne but strict hygiene is necessary to maintain this level of
health since pathogens surviving in soil or plant debris are quickly re-
acquired when stock are multiplied in farm conditions. Similarly, produc-
tion of plants from true seed could lead to much healthier crops.
10.4.2 Chemical control
(a) Soil
Some diseases could be controlled by application of chemicals to soil,
although development of the technique awaits production of effective
materials at economic cost and methods for thorough incorporation into
soil. Systemically-acting soil-applied materials and phloem-translocated
chemicals effective against root and tuber diseases could lead to a rapid
change in disease control practice. Currently, however, control by soil
applied chemicals finds but limited application for potato diseases.
Several materials have been used to prolong growth and increase yields
of crops infected with Verticillium spp., including nematicides which are
effective when the disease is associated with nematodes in soiL Fumigation
with methyl bromide, a very toxic material with a wide spectrum of activity
against arthropods, nematodes, fungi and weed seeds, decreases the
severity of disease but involves the use of gas-tight sheets making it
cumbersome and costly for large scale application. Pre-planting treatments
with dazomet or chloropicrin are also effective as are the nematicides
aldicarb, DD, and telone, even when nematodes known to increase the
disease are apparently not present in soil (Easton, 1970), although some
nematicides can increase the amount of infection by Rhizoctonia (Scholte,
1987). Benomyl incorporated in soil delays the appearance of Verticillium
symptoms, and is beneficial to subsequent potato crops, although the
major effect seems to be by controlling nematodes (Hide et al., 1984).
Quintozene (PCNB) incorporated in soil decreases common scab, stem
canker and black scurf so that saleable tuber yields are improved, but the
possibility of phytotoxicity and taint in tubers limits its general acceptance.
Common scab may also be decreased by soil applications of a range of
chemicals, including urea formaldehyde, sulphur, gypsum, copper salts
and magnesium sulphate, but whether through effects on the pathogen or
on the host is not understood. Sulphur has also been reported to decrease
pink rot, powdery scab and blight.
(b) Seed tubers

Treating seed tubers with chemicals offers a method of eliminating the
relatively small but important amounts of inoculum they carry. The most
common fungicides previously used on seed tubers contained mercury and
acted as eradicants. Bacterial and Fusarium rots, gangrene, skin spot,
silver scurf and black scurf are controlled by washing and dipping in
solutions or suspensions of organic mercury compounds. The treatment
encourages sprouting and early even emergence and increases the propor-
tion of small tubers in the crop, although it may increase the incidence of
blackleg. However, awareness of the toxic hazards of mercury compounds
and associated problems in the disposal of effluent stimulated a search for
alternative and less noxious materials.
In Scotland, fumigating potatoes with 2-aminobutane in gas-tight
chambers decreases gangrene and skin spot in stored tubers (Graham et
al., 1973) and, like treatment with mercury compounds, best control is
achieved by fumigating soon after lifting. Other fungicides including
thiabendazole, carbendazim, imazalil and prochloraz applied to tubers at
harvest, control gangrene, dry rot, skin spot and silver scurf during
storage. Also, when applied to diseased seed tubers they prevent escape of
inoculum after planting and so produce crops that remain healthy during
storage (Hide et al., 1987). Unfortunately, resistance to thiabendazole has
been found in isolates of Fusarium, Helminthosporium and Polyscytalum
and so the fungicide may fail to control dry rot, silver scurf and skin spot
(Langerfeld, 1986; Hide et al., 1988), but cross resistance to imazalil has
not yet been detected. Treatment of seed tubers with tolclofos-methyl,
pencycuron or iprodione prevents infection from tuber-borne Rhizoctonia
inoculum (Hide et al., 1987). However, the degree of control with all
materials will be related to the efficiency of the application method and will
be greatest when they are deposited evenly over the whole of the tuber
surface (Cayley et al., 1987).
(c) Foliage
Chemicals may be dusted, or more commonly sprayed, on to foliage either
to kill insects which transmit virus diseases (see Section 10.2.2 and Chapter
11), or as protectants against early or late blight. Systemically-acting
chemicals which can be applied to foliage to protect the whole plant,
including the underground parts, have been found and for late blight are
now widely used. Other materials that control common scab when applied
to foliage (McIntosh et al., 1982) have not yet been developed for
commercial use.
Insecticides sprayed on foliage can control the insect vectors of potato
viruses. Spread of leaf roll is prevented by conventional organophosphate
or carbamate insecticides applied as sprays or as granules during planting
but these materials have little effect on spread of virus Y because aphids
are killed too slowly. Pyrethroids have rapid 'knockdown' and are more
effective. Sprays of mineral oil or whitewash have also been found to
decrease spread of virus Y (Gibson and Cayley, 1984; Marco, 1986).
Both early and late blight can become epidemic but the timely
application of fungicide slows or prevents their progress and so prolongs
the life of the foliage, allowing the bulking of the crops to continue. For
late blight, the timing and frequency of spray applications, fungicides and
formulations and the weather conditions favouring infection, spread and
epidemic development have been much studied. Such information has
enabled many countries to operate a forecasting service to warn farmers of
outbreaks of this disease and when conditions are suitable for its spread.
Inorganic chemicals based on copper (which can be phytotoxic) were
used for many years but have been largely replaced by organic dithio-
carbamates (mancozeb, zineb, maneb, etc.), organo-tin compounds (fentin
hydroxide, fentin acetate) and, more recently, by systemically acting
phenylamides. High volume sprays applied to run-off, or medium volume
sprays (200-350 I ha- l ) designed to wet the foliage short of run-off, are
applied by tractor-mounted sprayers. Dusting is sometimes preferred as a
quicker method using light-weight equipment, especially in poor ground
conditions. More recently, highly concentrated formulations suitable for
low or ultra-low volume application from aircraft have been introduced.
After evaporation, the concentrated droplets leave 'spots' of chemical
which is then released in rain at rates determined by the retention qualities
of the formulation. Strains of P. infestans tolerant to phenylamides are now
appearing (Davidse et at., 1983) and this has led to spray programmes that
include surface acting dithiocarbamates, systemics and late-season organo-
tin compounds, the latter to decrease the incidence of tuber infection.
Early blight is becoming increasingly important in the USA, especially in
areas where overhead irrigation is more widely practised. First applications
are usually of dithiocarbamates or captafol, timed according to plant size
and maturity or when primary infection of lower leaves has occurred. The
number and timing of spray applications vary widely in practice, although
research has shown that three well-timed sprays can be very effective
(Harrison and Venette, 1970; Douglas and Groskopp, 1974).
Tractors used for spraying crops can damage foliage and compact soil so
that yields from damaged rows are decreased by up to 30%. Loss of crop
has been estimated at 5-7%, but will depend on the proportion of rows
damaged and varies greatly with soil type and season.
10.4.3 Legislation and certification of seed

The purpose of legislation and certification schemes is to prevent introduc-
tion of alien diseases and decrease the spread of endemic ones (Ebbles,
1979). Before it was realized that introduction of new diseases could be
prevented or initial introductions eradicated, diseases became established
in new areas that provided a favourable climate. Nowadays each country
has regulations prohibiting import of tubers carrying certain diseases, or of
all tubers from areas where certain diseases are known to be endemic. Such
procedures are more likely to be effective against pathogens that have no
airborne spores, for example Synchytrium endobioticum and Clavibacter
michiganensis ssp. sependonicus, than those having airborne spores or
vectors which can travel long distances (for example Alternaria solani,
Phytophthora infestans, insect-transmitted viruses). In the past, quarantine
restrictions have usually only delayed the introduction of diseases but this
delay could provide a valuable opportunity to develop control methods, or
perhaps breed resistant cultivars in anticipation of eventual introduction.
However, to embark on such a programme would demand the knowledge,
or at least the conviction, that the disease would, if introduced, pose a
serious threat, and this is not always so. Moreover, it would involve the
introduction of the pathogen concerned for experimental purposes, which
is always attended by some risk even with the constraints of rigid
quarantine regulations.
Spread of endemic diseases can be restricted by legislation prohibiting
production in, or movement away from, areas known to be infested with
serious pathogens, and by voluntary certification schemes controlling
health of seed tubers. Seed crops are often produced in areas isolated from
the main growing regions where aphids are scarce and where crops can be
maintained free of virus diseases. In the UK the importance of wart disease
and the persistence of the pathogen in soil were realized in the early 1900s,
but no method of control was known, although some cultivars did not
become infected ('immune'); seed stocks often contained rogue tubers of
non-immune cultivars, so schemes were introduced to ensure trueness to
type. Roguing (negative selection) and later multiplication from disease-
free plants (positive selection) were used to improve the health of seed,
and tolerances for diseases and abnormalities in field inspections were
gradually decreased. In many countries stem cuttings or micropropagated
plants are now used to produce nuclear stocks free from pathogenic
bacteria and fungi.
Certification of seed can control only those diseases that are tuber-borne
and, even if stocks are multiplied from healthy seed, they are not protected
from diseases that are acquired from soil. Healthy seed can be successfully
produced if multiplication is done hygienically and supplemented with
chemicals that can keep inoculum at a very low level, and the regular and
continual replacement of diseased seed with recently propagated healthy
stocks should itself help in this direction.
10.4.4 Resistant cultivars

Replacing susceptible cultivars with resistant ones is the most convenient
and, for the grower, the most economical and practical method of
controlling diseases, and even in the early nineteenth century differences in
susceptibility to disease were noted between cultivars. Genetic sources of
resistance, as well as of many other desirable characters, found in existing
cultivars or in wild or cultivated Solanum spp., have to be incorporated
into new cultivars. This is traditionally achieved by sexual crossing but
during the past decade other methods including genetic transformation and
protoplast fusion have been developed to bring about specific and pre-
determined changes in the genome.
Immunity to disease is seldom achieved and may not be long-lasting
because uncommon pathogenic races can be selected, or the pathogens
themselves adapt and become able to infect previously immune plants.
Cultivars exhibiting tolerance were once thought valuable for controlling
virus diseases. Now they are considered unsatisfactory as their use restricts
the growing of non-tolerant cultivars, which may be preferred in other
important respects, by maintaining perpetual reservoirs of inoculum. At
present, two types of resistance are exploited. Hypersensitivity relies on
single genes and with late blight, infected cells die rapidly and prevent
further spread of the disease. With hypersensitivity to viruses, normal
inoculation fails because infected cells die and therefore plants are 'field-
immune'. This hypersensitivity can be demonstrated by grafting with an
infected tolerant cultivar; the whole plant dies as a result of mass death of
infected cells. However, reliance on single major genes conferring hyper-
sensitivity is now generally recognized as unsatisfactory at least for
airborne pathogens capable of epidemic disease (e.g. P. infestans, causing
late blight). Experience suggests that such pathogens exist as mixed
populations from which new races are likely to emerge able to overcome
this type of resistance in response to the extreme selection pressure it
imposes. Multigenic (field) resistance is more broadly based and seldom
eliminates pathogens but allows a low level of disease to develop; less
selection pressure is imposed and the resistance is likely to be more
durable. In the past, most breeding of potato cultivars has concentrated on
resistance to wart, late blight and virus diseases, but now the testing of
agronomically acceptable selections for resistance to a range of trouble-
some diseases is more commonly practised.
Immunity to wart has long been recognized, although improved methods
of testing showed that 'immune' cultivars gave differing amounts of
hypersensitive reaction (field immunity). Characters governing resistance
are dominant and readily inherited and variability of the pathogen is low,
so the chances of selecting new virulent strains (or races) seem remote,
although races with the ability to infect cultivars immune to UK strains of
the pathogen, have been found in N. America, central Europe and India.
Hypersensitivity and multigenic resistance have been used in breeding
cultivars resistant to late blight. Hybrids derived from the Mexican
Solanum demissum showed hypersensitivity but later succumbed to new
races of P. infestans. Variability in the pathogen has shown that cultivars
depending on hypersensitivity for their resistance to late blight are unlikely
to maintain their resistance for very long and this approach has been
abandoned in favour of multi genic resistance. This is likely to prove more
durable and seems equally effective against all known races of the fungus.
Progress of the disease is restricted by effects on different components of
its development, induding resistance to infection, slow growth of the
pathogen through tissues, an extended incubation period and a reduction
in the number of spores produced, resulting overall in a prolongation of the
life of the foliage.
Many genes govern the reaction of Solanum spp. to viruses and a
Future prospects 431
dominant gene for hypersensitivity to potato virus X confers field immunity
to several established cultivars. Single gene hypersensitivity to strains of
virus X occurs in S. acaule and to viruses Y and A in S. stoloniferum.
Multigenic resistance to Y and leaf roll viruses is known in S. phureja, S.
acaule and S. demissum and for leaf roll this is being used because there are
no alternatives. Using protoplast fusion, somatic hybrids have been
produced between S. tuberosum and non-tuberizing S. brevidens and these
show resistance to leaf roll and Y and to Erwinia spp. (Austin et al., 1988;
Gibson et al., 1988).
10.5 FUTURE PROSPECTS
Potato production in a disease-free environment is as yet unknown and will

be difficult to achieve, but must be the aim if the costs of preventing
diseases and the losses they cause are to be avoided. At present, the use of
resistant cultivars is probably the easiest and cheapest method of disease
control for the farmer and it is hoped that current investigations with novel
genetic techniques will enable a range of beneficial characters to be rapidly
incorporated into new or existing cultivars.
Late blight has remained the most important disease in world potato
production for more than 100 years and any major improvement in control
must come both from resistant cultivars and from efficient fungicide usage.
The risk from oospores in soil will need to be assessed.
Chemicals applied to soil may help to control diseases that originate
from soil-borne inoculum if materials can be found that do not persist in
soil or in crops destined for human consumption. Alternatively, for
diseases such as powdery scab, common scab, Verticillium and black dot,
assay methods could be useful in helping determine the risk of infection
from inoculum in soil.
The prospects seem brighter for diseases that are mostly tuber-borne,
provided there are no major alternative sources of infection; although
healthy nuclear stocks can be produced readily, it is difficult to maintain
high standards of hygiene during multiplication to commercial quantities.
With micropropagation techniques it should be possible to decrease the
number of multiplication years. These diseases could also become less
frequent if the crop is produced from true seed.
Chemical tuber treatments are now widely used but, although their
application may be improved by the use of electrostatic sprayers, adhering
soil often prevents deposition of chemical on to the tuber skin or into
wounds. Also as chemicals are often only included as an additional factor
in crop production, their effectiveness should be improved by integrating
their use with agronomic practices that also discourage disease, such as
early harvesting and drying tubers before storage. With such strategies it
might be possible to decrease the amount of chemical used, and this would
be useful if fungicide resistance becomes widespread.
Storing potatoes always involves risks and improvement in storage
technology is needed to avoid infrequent catastrophes as well as the
smaller chronic losses which amount annually to a significant proportion of
world production. However, high costs will prohibit the installation of fully
controlled stores for as long as the potato remains a cheap staple food. The
major problems are the presence of pathogens and mechanical damage on
tubers as they are loaded into store. If crops likely to deteriorate in quality
could be distinguished from those which have a potential for long storage,
decisions on which crops to store and subsequent management of stores
could be modified to allow appropriate orderly marketing with minimal
losses. This might be done by simple assessments of disease potential
during growth, assessments of injuries at lifting, and by monitoring the
store environment for volatile chemicals that indicate the onset of trouble
(Varns and Glynn, 1979). Mechanical damage is one of the most important
causes of direct and indirect loss of crop and its avoidance would
undoubtedly make the most significant single contribution to production. It
is a problem worthy of considerable effort since the benefits could be very
great indeed; it has been estimated that on average more than 20% of the
crop in the UK sustains largely avoidable damage at harvest.
Besides agronomic aspects of diseases, there are many facets of the
biology of pathogens that need to be investigated before effective and
continuing control can be assured. Much is still to be learned about the
relative importance of different sources of inoculum, its spread and the
time and methods of infection. The role of volunteer plants and of perhaps
hitherto unknown sexual stages of fungi need to be evaluated. Resistant
cultivars may be the best method of decreasing effects of disease; mean-
while, other control measures can be made more effective if sources of
inoculum can be decreased.
REFERENCES
Adams, M.J. and Griffith, R.L. (1978) The effect of harvest date and duration of
wound healing conditions on the susceptibility of damaged potato tubers to
infection by Phoma exigua (gangrene). Ann. Appl. BioI., 88, 51-5.
Adams, M.J. and Hide, G.A. (1981) Effects of common scab (Streptomyces
scabies) on potatoes. Ann. Appl. Bioi., 98, 211-16.
Adams, M.J. and Lapwod, D.H. (1983) The effect of Erwinia carotovora subsp.
atroseptica (blackleg) on potato plants. II. Compensatory growth. Ann. Appl.
Bioi., 103, 79-85.
Adams, M.J., Hide, G.A. and Lapwood, D.H. (1980) Relationships between
disease levels on seed tubers, on crops during growth and in stored potatoes. I.
Introduction and black scurf. Potato Res., 23, 201-14.
Adams, M.J., Read, P.J., Lapwood, D.H., Cayley, G.R. and Hide, G.A. (1987)
References 433
The effect of irrigation on powdery scab and other tuber diseases of potatoes.
Ann. Appl. BioI. 110, 287-94.
Ali, M.A., Lennard, J.H. and Boyd, AE.W. (1970) Potato coiled sprout in
relation to seed tuber storage treatment and to infection by Verticillium nubilum
Pethybr. Ann. Appl. BioI., 66, 407-15.
Austin, S., Lojkowska, E., Ehlenfeldt, M.K., Kelman, A and Helgeson, J.P.
(1988) Fertile interspecific somatic hybrids of Solanum: a novel source of
resistance to Erwinia soft rot. Phytopathology, 78, 1216-20.
Barclay, G.M., Murphy, H.J., Manzer, F.E. and Hutchinson, F.E. (1973) Effects
of differential rates of nitrogen and phosphorus on early blight in potatoes. Am.
Potato 1., 50, 42-8.
Bawden, F.C. and Kassanis, B. (1965) The potato variety King Edward VII and
paracrinkle virus. Rothamsted Report for 1964, pp. 282-90.
Bokx, J.A. de and Mooi, J.C. (1974) Methods of quality assessment of seed
potatoes. Potato Res. 17, 410--33.
Boyd, AE. W. (1972) Potato storage diseases. Rev. Pl. Path., 51, 297-321.
Broadbent, L. (1953) Aphids and virus diseases in potato crops. Biological
Reviews, 28, 350--80.
Broadbent, L., Heathcote, G.D. and Burt, P.E. (1960) Field trials on the retention
of potato stocks in England. Eur. Potato 1., 3, 251--62.
Busch, L.V. (1967) Distribution in Ontario of Verticillium strains causing wilt of
potatoes. Can. PI. Disease Survey, 47, 76-8-
Campos, E., Maher, E.A. and Kelman, A. (1982) Relationship of pectolytic
clostridia and Erwinia carotovora strains to decay of potato tubers in storage.
Plant Disease, 66, 543--6.
Carnegie, S.F., Adam, J.W. and Symonds, C. (1978) Persistence of Phoma exigua
var. foveata and Polyscytalum pustulans in dry soil from potato stores im relation
to reinfection of stocks derived from stem cuttings. Ann. Appl. Bioi., 90,
179-86.
Carnegie, S.F., Adam, J.W. and Cameron, A.M. (1987) Spread of Phoma exigua
var. foveata to healthy potato plants. Plant Path., 36, 398-406.
Catchpole, A.H. and Hillman, J.R. (1976) The involvement of ethylene in the
coiled sprout disorder of potato. Ann. Appl. BioI., 83, 413-23.
Cayley, G.R., Hide, G.A, Lewthwaite, R.J., Pye, B.J. and Vojvodic, P.J. (1987)
Methods of applying fungicide sprays to potato tubers and description and use
of a prototype electrostatic sprayer. Potato Res., 30, 301-17.
Church, B.M., Hampson, C.P. and Fox, W.R. (1970) The quality of stored main
crop potatoes in Great Britain. Potato Res. 13, 41-58.
Cohen, S. (1981) Reducing the spread of aphid-transmitted viruses in peppers by
coarse-net cover. Phytoparasitica, 9, 69-76.
Cramer, H.H. (1967) Plant Protection and World Crop Production, Farbenfabriken
Bayer, AG, Levenkusen.
Cromarty, R.W. and Easton, G.D. (1973) The incidence of decay and factors
affecting bacterial soft rot of potatoes. Am. Potato 1., 50, 398-407.
Davidse, L.c., Danial, D.L. and van Westen, c.J. (1983) Resistance to metalaxyl in
Phytophthora infestans in the Netherlands. Netherlands 1. Plant Pathot., 89, 1-20.
Davis, J.R., McMaster, G.M., Callihan, R.H., Garner, J.G. and McDole, R.E.
(1974) The relationship of irrigation timing and soil treatments to control potato
scab. Phytopathology, 64, 1404-10.
Douglas, D.R. and Groskopp, M.D. (1974) Control of early blight in Eastern and
South central Idaho. Am. Potato J., 51, 361-8.
Easton, G.D. (1970) Systemic insecticides, soil fumigation, and nitrogen fertiliza-
tion for Verticillium wilt control. Am. Potato J., 47, 419-26.
Easton, G.D., Nagle, M.E. and Bailey, D.L. (1972) Effect of annual soil fumiga-
tion and pre-halVest vine burning on Verticillium wiU of potato. Phytopathology,
62,520-4.
Ebbles, D.L. (1979) Principles and problems of certification schemes for vegeta-
tively propagated crops, with special reference to potatoes, in Plant Health, (eds
D.L. Ebbles and J.E. King), Blackwell, Oxford, pp. 113-20.
Emmond, G.S. and Ledingham, R.J. (1972) Effects of crop rotation on some soil-
borne pathogens of potato. Can. J. PI. Sci., 52, 605-11.
Evans, K. (1987) The interactions of potato cyst nematodes and Verticillium
dahliae on early and maincrop potato cultivars. Ann. Appl. Bioi., 110, 329-39.
Fernow, K.H., Peterson, L.C. and Plaisted, R.L. (1970) Spindle tuber virus in
seeds and pollen of infected potato plants. Am. Potato J., 47, 75-80.
Fox, R.A., Dashwood, E.P. and Wilson, H.M. (1970) Gangrene of potato. Scottish
Horticultural Research Institute Report for 1969, pp. 30-1.
Gibson, R.W. and Cayley, G.R. (1984) Improved control of potato virus Y by
mineral oil plus the pyrethroid cypermethrin applied electrostatically. Crop
Protection, 3, 469-78.
Gibson, R.W. and Gunenc, Y. (1981) Effect of covering potato crops with clear
polyethylene film on spread of potato virus Y. Plant Path. 30, 233-5.
Gibson, R.W., Jones, M.G.K. and Fish, N. (1988) Resistance to potato leaf roll
virus and potato virus Y in somatic hybrids between dihaploid Solanum
tuberosum and S. brevidens. Theor. Appl. Genet., 76, 113-17.
Graham, D.C. and Harrison, M.D. (1986) (eds) Report of the International
Conference on Potato Blackleg Disease, Potato Marketing Board, London,
96 pp.
Graham, D.C., Hamilton, G.A., Quinn, C.E. and Ruthven, A.D. (1973) Use of
2-aminobutane as a fumigant for control of gangrene, skin spot and silver scurf
diseases of potato tubers. Potato Res., 16, 109-25.
Graham, J. and Lloyd, A.B. (1979) Survival of potato strain (race 3) of
Pseudomonas solanacearum in the deeper soil layers. Australian J. Agric. Res.,
30,489-96.
Griffith, R.L., Hide, G.A., Hirst, J.M. and Stedman, O.J. (1974) Effects of
gangrene (Phoma exigua) on potatoes. Ann. Appl. Bioi., 77, 237-50.
Harrison, J.A.C. and Isaac, I. (1969) Host/parasite relations up to the time oftuber
initiation in potato plants infected with Verticillium spp. Ann. Appl. Bioi., 64,
469-82.
Harrison, M.D. and Venette, J.R. (1970) Chemical control of potato early blight
and its effect on potato yield. Am. Potato J. 47, 81-6.
Harrison, M.D., Livingston, C.H. and Oshima, N. (1965) Epidemiology of potato
early blight in Colorado. I. Initial infection, disease development and the
influence of environmental factors. Am. Potato J., 42, 279-91.
Hide, G.A. (1975) Effect of heat treatment of potato tubers on Oospora pustulans.
Plant Path., 24, 233-6.
Hide, G.A. (1981) Fungus diseases on potato seed tubers planted in England and
Wales, 1963-76. Ann. Appl. Bioi., 98, 377-93.
References 435
Hide, G.A. and Cayley, G.R. (1987) Effects of delaying fungicide treatment and of
curing and chlorpropham on the incidence of skin spot on stored potato tubers.
Ann. Appl. BioI., 110, 617-27.
Hide, G.A., Hirst, J.M. and Stedman, O.J. (1973) Effects of skin spot (Oospora
pustulans) on potatoes. Ann. Appl. Bioi., 73, 151-{j2.
Hide, G.A., Corbett, D.C.M. and Evans, K. (1984) Effects of soil treatments and
cultivars on 'early dying' disease of potatoes caused by Globodera rostochiensis
and Verticillium dahliae. Ann. Appl. BioI., 104,277-89.
Hide, G.A., Read, P.J. and Sandison, J.P. (1985) Stem canker (Rhizoctonia
solani) of maincrop potatoes. II. Effects on growth and yield. Ann. Appl. Bioi.
106,423-37.
Hide, G.A., Read, P.J., Sandison, J.P. and Hall, S.M. (1987) Control of potato
diseases with fungicides applied to seed tubers. Tests of Agrochemicals and
Cultivars, 8, Ann. Appl. Bioi., 110, (Supplement), 72-3.
Hide, G.A., Hall, S.M. and Boorer, K.J. (1988) Resistance to thiabendazole in
isolates of Helminthosporium solani, the cause of silver scurf disease of
potatoes. Plant Path., 37, 377-80.
Hirst, J. M., Hide, G.A., Stedman, O.J. and Griffith, R.L. (1973) Yield
compensation in gappy potato crops and methods to measure effects of fungi
pathogenic on seed tubers. Ann. Appl. Bioi., 73, 143-50.
Isaac, I. and Harrison, J.A.e. (1968) The symptoms and causal agents of early-
dying disease (Verticillium wilt) of potatoes. Ann. Appl. BioI., 61, 231--44.
James, W.C., Callbeck, L.C., Hodgson, W.A. and Shih, C.S. (1971) Evaluation of
a method used to estimate loss in yield of potatoes caused by late blight.
Phytopathology, 61, 1471-{j.
James, W.e., Shih, C.S., Hodgson, W.A. and Callbeck, L.C. (1972) The quanti-
tative relationship between late blight of potato and loss in tuber yield.
Phytopathology, 62, 92-{j.
Kassanis, B. and Varma, A. (1967) The production of virus-free clones of some
British potato varieties. Ann. Appl. Bioi., 59, 447-50.
Kloepper, J.W., Harrison, M.D. and Brewer, J. (1979) The association of Erwinia
carotovora var. atroseptica and Erwinia carotovora var. carotovora with insects
in Colorado. Am. Potato J., 56, 351-{j1.
Langerfeld, E. (1986) Thiabendazole resistance in Fusarium sulphureum. Nachr.
Deutsch Pflanzenschutzdientes, 38, 165-8.
Lapwood, D.H. (1971) Observations on blight (Phytophthora infestans) and
resistant potatoes at Toluca, Mexico. Ann. Appl. BioI., 68, 41-53.
Lapwood, D.H. and Hering, T.F. (1970) Soil moisture and the infection of young
potato tubers by Streptomyces scabies (common scab). Potato Res., 13, 296-304.
Lapwood, D.H., Wellings, L.W. and Hawkins, J.H. (1973) Irrigation as a practical
means to control potato common scab (Streptomyces scabies): Final experiment
and conclusions. Plant Path., 22, 35-41.
Large, E.C. (1952) The interpretation of progress curves for potato blight and
other plant diseases. Plant Path., 1, 109-17.
Lizarraga, R.E., Salazar, L.F., Roca, W.M. and Schilde-Rentschler, L. (1980)
Elimination of potato spindle tuber viroid by low temperature and meristem
culture. Phytopathology, 70, 754--5.
Logan, C., Copeland, R.B. and Little, G. (1975) Potato gangrene control by ultra
low volume sprays of thiabendazole. Ann. Appl. BioI., 80, 199-204.
Lonsdale, D., Cunliffe, C. and Epton, H.A.S. (1980) Possible routes of entry of
Phytophthora erythroseptica Pethyb. and its growth within potato plants.
Phytopath. Z. 97, 107-17.
MacHardy, W.E. (1979) A simplified, non-computerized program for forecasting
potato late blight. Pl. Dis. Reptr., 63, 21-5.
MacKenzie, D .R. (1981) Association of potato early blight, nitrogen fertilizer rate,
and potato yield. Plant Disease, 65, 575-7.
Marco, S. (1986) Incidence of aphid-transmitted virus infections reduced by
whitewash spray on plants. Phytopathology, 76, 1344-8.
McIntosh, A.H., Burrell, M.M. and Hawkins, J.H. (1982) Field trials of foliar
sprays of 3,5-dichlorophenoxyacetic acid (3,5-D) against common scab on
potatoes. Potato Res., 25, 347-50.
Morsink, F. and Rich, A.E. (1968) Interactions between Verticillium albo-atrum
and Pratylenchus penetrans in the Verticillium wilt of potatoes. Phytopathology,
58,401.
Nelson, G.A. (1984) Survival of Corynebacterium sepedonicum in potato stems and
on surfaces held at freezing and above-freezing temperatures. Am. Potato J.,
62,23-8.
Perombelon, M.C.M. (1972) The extent and survival of contamination of potato
stocks in Scotland by Erwinia carotovora var. carotovora and E. carotovora var.
atroseptica. Ann. Appl. Bioi., 71, 111-17.
Perombelon, M.C.M., Fox, RA. and Lowe, R (1979) Dispersion of Erwinia
carotovora in aerosols produced by the pulverization of potato haulm prior to
harvest. Phytopath. Z., 94, 24~0.
Powelson, R.L. (1964) Studies on Verticillium wilt of potatoes in Oregon. Am.
Potato J., 41, 303.
Quinn, C.E., Sells, I.A. and Graham, D.C. (1980) Soft rot Erwinia bacteria in the
atmospheric bacterial aerosol. J. Appl. Bacteriol., 49,175-81.
Rambawale, O.M. and Bedi, P.S. (1982) Epidemiology of early blight of potato in
the Punjab. Indian Phytopathology, 35, 574-82.
Read, P.J. and Hide, G.A. (1984) Effects of silver scurf (Helminthosporium solani)
on seed potatoes. Potato Res., 27, 145-54.
Read, P.J. and Hide, G.A. (1988) Effects of inoculum source and irrigation on
black dot disease of potatoes (Collectotrichum coccodes (Wallr.) Hughes) and
its development during storage. Potato Res., 31, 493--500.
Reestman, A.J. (1970) Importance of the degree of virus infection for tb.e
production of ware potatoes. Potato Res., 13, 248--68.
Schick, R. (1952) Problems of potato multiplication. Deutsch. Landw., 3, 618-27.
Scholte, K. (1987) The effect of crop rotation and granular nematicides on the
incidence of Rhizoctonia solani in potato. Potato Res., 30, 187-99.
Schuster, M.L. and Coyne, D.P. (1975) Survival factors of plant pathogenic
bacteria. Nebraska Agric. Exp. Station Res. Bulletin, 268, 533 pp.
Soltanpour, P.N. and Harrison, M.D. (1974) Interrelations between nitrogen and
phosphorus fertilization and early blight control of potatoes. Am. Potato 1.,51, 1-7.
Stanghellini, M.E. and Meneley, J.C. (1975) Identification of soft-rot Erwinia
associated with blackleg of potato in Arizona. Phytopathology, 65, 86-7.
Tantius, P.H., Fyfe, A.M., Shaw, D.S. and Shattock, RC. (1986) Occurrence of
the A2 mating type and self-fertile isolates of Phytophthora infestans in England
and Wales. Plant Path., 35, 578-81.
References 437
Taylor, P.A., Flett, S.P., de Boer, RF. and Marshall, D. (1986) Effect of
irrigation regimes on powdery scab disease and yield of potatoes. Australian J.
Exp. Agric., 26, 745-50.
Thirumalachar, M.J. (1967) Pathogenicity of Colletotrichum atramentarium Oil
some potato varieties. Am. Potato J., 44, 241-4.
Twiss, P.T.G. and Jones, M.P. (1965) A survey of wastage in bulk-stored maincrop
potatoes in Great Britain. Eur. Potato J., 8, 154-72.
Yarns, J.L. and Glynn, M.T. (1979) Detection of disease in stored potatoes by
volatile monitoring. Am. Potato J., 56, 185-97.
Venette, J.R. and Harrison, M.D. (1973) Factors affecting infection of potato
tubers by Alternaria solani in Colorado. Am. Potato J., 50, 283-92.
Wale, S.J., Robertson, K., Robinson, K. and Foster, G. (1986) Studies on the
relationship of contamination of seed potato tubers with Erwinia spp. to
blackleg incidence and large scale hot water dipping to reduce contamination.
Aspects of Applied Biology, 13, 285-91.
Waterer, D.R. and Pritchard, M.K. (1984) Monitoring volatiles: A technique for
detection of soft rot (Erwinia carotovora) in potato tubers. Can. 1. Pl. Path., 6,
165-71.
Watson, R.D. and Huber, D.M. (1971) Effects of nitrogen and cultural practices
on the severity of Verticillium wilt of potato. Proc. Int. Verticillium Symp., Wye
College, p. 32.
CHAPTER 11
Pest aspects of potato production

Part 1. The nematode pests of
potatoes
K. Evans and D.L. Trudgill
11.1 INTRODUCTION TO THEIR BIOLOGY
Plant-parasitic nematodes are typically small (0.2-10 mm long) worm-like

animals, able to move between soil particles, between folded leaves of
plant buds, in the air spaces of leaves and stems or in plant tissues
themselves. Their movement in soil is influenced by the thickness of
the water films surrounding soil particles and they are inactive in dry
conditions. Agricultural soils usually contain many species of plant-
parasitic nematodes, as well as predatory and microphagous types, and the
species composition is affected by many factors including climate, soil type
and cropping patterns. Species parasitic on plants all possess a mouth stylet
which they use to puncture plant cells and, in all except the Trichodoridae,
this stylet is hollow and used to extract cell contents. Some feed on the
cytoplasm which accumulates in the region of the stylet following the
injection of saliva. Others modify the cells they feed upon so that the
supply of nutrients is increased and they are able to become sedentary
and lose their vermiform shape (Giobodera, Heterodera, Meioidogyne,
Nacobbus). Such nematodes usually produce large numbers of eggs and
include the most harmful species. The damage may be direct, due to their
feeding and penetration of roots, but some species transmit harmful viruses
and damage by other species may be made worse by interaction with
pathogenic fungi and bacteria.
Nematodes attacking potatoes can be grouped according to whether they
attack stems and leaves, tubers or roots (see Table 11.1 for the most
important species). Jensen et ai. (1979) list 67 species from 24 genera
reported to be associated with potatoes but many of these are of little
importance to crop production. The most damaging are the potato cyst
Published in 1992 by Chapman & Hall, London. ISBN 0412296403
Table 11.1 The main nematode pests of potatoes grown in temperate soils
Latin name Common name Part of plant attacked Known geographical
distribution
Globodera rostochiensis Golden nematode of potato Roots Worldwide
Globodera pallida White potato cyst nematode Roots Worldwide
Meloidogyne hapla Northern root-knot nematode Roots N. America and temperate
areas in general
Meloidogyne chitwoodi Columbia root-knot nematode Roots North-west USA and
The Netherlands
Nacobbus aberrans False root-knot nematode Roots Peru and Bolivia
Pratylenchus penetrans Root lesion nematode Roots N. America and Europe
Ditylenchus destructor Tuber-rot nematode Tubers N. America, W. Europe
and USSR
Ditylenchus dipsaci Stem nematode Stems, leaves, tubers W. Europe
Trichodorus and Stubby root nematodes Transmit tobacco rattle N. America and Europe
Paratrichodorus spp. virus (sprain g) to tubers
440 The nematode pest of potatoes
nematDdes (Globodera rostochiensis, and G. pal/ida), which cause tre-
mendDus IDsses in many cDuntries. Other species distributed wDrldwide
which cause significant problems include roDt knDt (Meloidogyne
spp.), stubby rDDt (Trichodorus and Paratrichodorus spp.), root lesiDn
(Pratylenchus spp.) and pDtatD rDt (Ditylenchus spp.) nematDdes. SDme
species, such as the false roDt knDt nematDde (Nacobbus aberrans) and the
sting nematDde (Belonolaimus longicaudatus) , are IDcally impDrtant
problems.
11.1.1 Nematodes which attack stems and leaves

The stem nematDde, Ditylenchus dipsaci, produces pectDlytic enzymes
which dissDlve the middle lamella between adjacent cells, so. creating
spaces and enabling the nematDde to. attack stem tissues (Riedel and Mai,
1971). Ditylenchus dipsaci has been repDrted causing distDrtiDn Df pDtatD
haulms in Germany and The Netherlands (SeinhDrst, 1957) and in England
(NewtDn and DuthDit, 1954). The aerial parts Df plants are invaded from
the sDil Dnly during wet weather bl!lt, Dnce within the prDtected environ-
ment Df the plant, multiplicatiDn is rapid and haulms may be severely
distDrted and stunted. The pseudo. stem nematDde (Neotylenchus vigissi)
has also. been repDrted as a parasite Df leaves, stems and tubers and, in
Russia, causes IDsses Df up to. 50% (BrDdie, 1984).
11.1.2 Nematodes which attack tuhers

Ditylenchus dipsaci may also. attack tubers, eventually causing tuber rot,
but dDes so. less frequently than the tuber-rot nematDde Ditylenchus
destructor. BDth species invade the tubers via eyes Dr lenticels, but D.
dipsaci lesiDns may extend throughDut the tuber whereas D. destructor
lesiDns are mDre superficial. In advanced stages Df infectiDn diseased tubers
may be readily recDgnized by sunken, dark cDIDured pits and skin cracks.
Damage by bDth species is accentuated by secDndary fungal Dr bacterial
infectiDns and tends to. becDme mDre apparent during stDrage. Weeds provide
alternative hDStS fDr D. destructor but damage to. tubers DCCurs Dnly rarely
in western Europe because Df the relatively high standards Df weed cDntrol,
the use Df gDDd quality seed and crDp rotatiDn. Severe damage has been
repDrted frDm Canada and the USA, where infested land is nDW aVDided,
and also. frDm eastern EurDpe. The eCDIDgy, biDIDgy and contrDI Df these
nematDdes are reviewed by WinslDw and Willis (1972) and BrDdie (1984).
Tuber quality may be spDilt by 'spraing' disease Dr 'cDrky ringspDt'
caused by infectiDn with tDbaccD rattle virus (TRV). The virus is trans-
mitted to' tubers when they are fed upDn by viruliferDus Trichodorus and
Paratrichodorus spp. There are many serolDgically identifiable strains Df
TRV, to' which SDme pDtatD cultivars are resistant and Dthers susceptible.
Typical tuber symptDms are irregular shape, skin cracking and internal
necrotic arcs, thDugh these may also. be caused by pDtatD mDp tDP virus
Introduction to their biology 441
transmitted by the fungus Spongospora subterranea. Nine species of
Trichodorus and Paratrichodorus are vectors of TRV (Brown et al., 1989).
The Trichodoridae are all ectoparasitic nematodes with wide host ranges
and recent research indicates that each species is the vector of specific
serotypes of the virus. Most species are confined to lighter sandy soils
(Winfield and Cook, 1975) so that, although spraing is widespread in
western Europe, significant amounts of infection occur only if the soil is
moist enough for nematode activity in the period following tuber initiation
when tubers are most susceptible (van Hoof, 1964). When they are
abundant, Trichodorus and Paratrichodorus spp. sometimes cause stunting
of roots, and therefore tops, and abrasion of the surface of tubers due to
mass feeding.
The root knot nematodes (Melodiogyne spp.) are endoparasitic with
sedentary females and, because they occur mainly in warmer soils, have
several generations per year. They produce galls on roots and heavy
infestations may also result in attack of tubers, producing warty,
gall-like protuberances which render affected tubers unsaleable. The soil
temperature requirements of Melodiogyne spp. vary, M. hapla and M.
chitwoodi being adapted to lower temperatures than M. javanica, M.
incognita and M. arenaria (Santo and Q'Banon, 1981). Meloidogyne hapla
is the dominant species attacking potatoes in Europe, North America and
Japan; M. chitwoodi is locally important in north-western USA and has
also been found in The Netherlands. It poses more of a threat than M.
hapla because it can also parasitize cereals and therefore is less readily
controlled by crop rotation. Meloidogyne javanica and M. incognita are the
species which cause most damage in Australia, Africa, South America and
parts of Asia.
11.1.3 Nematodes which attack roots

Nematodes parasitic on potato roots exhibit a range of specialization. At
one end of the spectrum are migratory, ectoparasitic polyphagous genera
(e.g. Trichodorus, Tylenchorhynchus) which browse on epidermal cells
close to the root tip. These nematodes generally cause little damage except
when very abundant or when they interact with other pathogens or
environmental stress. More damaging, but still very polyphagous, are the
migratory endoparasites (e.g. Pratylenchus) which migrate within the
cortex of attacked roots. These nematodes, living in a protected environ-
ment within the roots, have high reproductive rates and may cause damage
in their own right. However, this damage is made worse by interaction with
other pathogens (e.g. with fungi, Verticillium spp.). The most specialized
nematodes are those with a sedentary, endoparasitic habit (e.g. Meloidogyne,
Nacobbus, Globodera). Invading juveniles of these genera induce the
formation of enlarged, cytoplasm-enriched feeding cells which enable them
to mature and produce many eggs without moving from cell to cell. This
sedentary habit is frequently accompanied by gall formation. The success
of these parasites leads to high densities in the soil which are often
associated with severe crop damage. Interaction with other pathogens is
frequent.
The degree of specialized adaptation to the host of these three main
types of root parasites determines the likelihood of evolution of host
resistance: the more specialized the nematode's relationship with the host,
the more likely the development of resistance. Thus, very effective
resistance to potato cyst nematodes (Globodera spp.) is found in a
number of Solanum species but no resistance to ectoparasites such as
Tylenchorhynchus has been reported (Roberts,1982).
(a) Migratory ectoparasites

Genera of ectoparasitic nematodes occasionally causing poor growth
include Belonolaimus, Criconemoides, Helicotylenchus, Longidorus,
Paralongidorus, Paratrichodorus, Paratylenchus, Rotylenchus, Trichodorus,
Tylenchorhynchus and Xiphinema (Winslow and Willis, 1972; Jensen et al.,
1979; Brodie, 1984). Most have wide host ranges and can be injurious to
potatoes when large populations are left by preceding crops. Damage by
Longidorus spp., which induce root-tip galls, has been reported in several
countries. Brown and Sykes (1975) reported that Longidorus elongatus
caused yield losses in potatoes, with yield related linearly to nematode
population density. In a sandy soil, populations of L. leptocephalus below
cultivation depth may have reduced yields by preventing roots entering the
sub-soil, so depriving the crop of moisture in mid-season (Evans, 1979).
Paralongidorus maximus caused typical galling of root tips and depressed
growth in patches in potato crops in Germany (Sprau, 1960). Large
numbers of Trichodorus spp. feeding on root tips may also stunt root
growth.
(b) Migratory endoparasites

Root lesion nematodes (Pratylenchus spp.) are the only migratory endo-
parasites which cause serious damage to potato roots. All stages are
vermiform and move freely into and out of roots. They have a powerful
stylet which they use to penetrate cell walls as they move within the roots.
Their host ranges are wide, although some hosts enable them to multiply
much more than others (Oostenbrink et al., 1957). Above ground
symptoms caused by high population densities of root lesion nematodes
appear as areas of poorly growing plants which turn yellow and die
prematurely (Dickerson et al., 1964). Pratylenchus penetrans is one of the
most harmful species in temperate regions and, in association with
Verticillium dahliae, causes an early wilt disease (Martin et al., 1982). It is,
however, uncommon in warmer climates where P. brachyurus (USA and
Africa), P. coffeae (Japan) and P. pratensis (tropical and subtropical areas
generally) replace it (Brodie, 1984).
(c) Sedentary endoparasites

Nacobbus aberrans (false root knot nematode)
The false root knot nematode which attacks potatoes appears to be native
to Andean regions of Peru and Bolivia. It was known as Nacobbus
serendipiticus bolivianus until Sher (1970) synonymized it with N.
batatiformis from sugar beet and N. serendipiticus from tomato as N.
aberrans. However, N. aberrans may be composed of several distinct races
(Jatala and Golden, 1977) and it is only those in South America which
attack potatoes (Mai et al., 1981). The restricted distribution of races
attacking potatoes may be due to poor persistence in the absence of a host
crop so that only in the high Andes in the region of the Bolivian altiplano,
where almost all the commonly grown crops (potatoes, quinoa, mashuar)
are hosts for this nematode, does it survive. Potatoes in this area are
frequently severely damaged and N. aberrans is often the dominant
nematode pest (Franco and de Scurrah, 1972), out-competing cyst nema-
todes which also occur. In nearby areas where a greater range of crops
including non-hosts (barley, beans, onions) are grown, cyst nematodes,
which are able to persist in the absence of a host crop, are the dominant
nematode pests of potatoes.
Nacobbus aberrans overwinters mainly as eggs. Second stage juveniles
hatch from these eggs and invade roots where they settle and feed in the
vascular tissues. After two moults some juveniles leave the roots and
undergo the final moult to adult males or females in the soil. Other
juveniles remain in the roots and moult to adult females. Those individuals
which develop into females in the soil re-invade larger roots and commence
feeding near the stele; the cells surrounding their heads enlarge and galls
form and envelop the entire females. The posteriors of the females extend
into the cortex, an opening is formed at the root surface and the eggs are
discharged into a gelatinous matrix. Tubers may also be invaded but the
nematodes rarely develop to maturity. However, this provides an ideal
means for dissemination of the nematodes over long distances (Mai et al.,
1981).
Infected plants are stunted and show signs of chlorosis and other
symptoms of root damage. Examination of roots shows spherical galls
occurring in a beadlike manner, which differ from those induced by
Meloidogyne spp. in their more regular shape.
Meloidogyne spp. (root knot nematodes)

Root knot nematodes occur more commonly in warmer, lighter soils: they
are not presently a major worldwide problem on potatoes because potatoes
are mainly grown in temperate regions. However, certain species cause
severe damage locally. Six species are pests of potatoes (Brodie, 1984) and
all are parthenogenetic, have wide host ranges and can have several
generations per year. Second stage juveniles invade roots and settle to feed
within the stele, where they induce the cells around their head to form
large, multinucleate 'giant' cells, which supply the developing juveniles
with an enriched food source, thus enabling them to remain sedentary. The
globe shaped females produce many eggs (c. 1000) and secrete a gelatinous
matrix into which the eggs are laid.
The roots of infected plants generally become heavily galled. Gall shape
ranges from almost spherical (M. arenaria) to a very rough and irregular
appearance (M. hapla) but M. chitwoodi, and to a lesser extent the other
species, does not always induce gall formation. Meloidogyne hapla causes
extensive lateral root proliferation in addition to gall formation. The tops
of infected plants are often stunted, wilt readily and their yields are
reduced. Tubers may also be infected and disfigured (see Section 11.1.2).
Rotylenchulus reniformis (reniform nematode)

The reniform nematode is closely related to Nacobbus. It is a sedentary
parasite which is usually not completely embedded within the root. It has
been reported on potatoes in Hawaii, India and Egypt (Brodie, 1984) and
shown to be pathogenic in glasshouse tests (Rebois et al., 1978).
Globodera rostochiensis and Globodera pallida (potato cyst nematodes)

The two species of potato cyst nematode originate from the Andean
regions of South America, Globodera pallida being commoner north of
Lake Titicaca and G. rostochiensis commoner to the south of this lake
(Evans et al., 1975a). From these regions both species have spread to many
of the potato growing areas of the world. Figure 11.1 shows the current
world distribution of the two species and indicates the probable routes of
their dissemination, with Europe acting as a secondary distribution centre.
Consequently, only a restricted part of the South American gene pool has
been introduced to many countries. For example, virulence genes enabling
G. rostochiensis to mature on potatoes with resistance gene HI. derived
from Solanum tuberosum ssp. andigena epc 1673, are absent from
populations in Britain and many other countries but present in some of
those in mainland Europe.
Potato cyst nematodes are highly specialized and have few hosts outside
the genus Solanum. Most cyst nematodes are adapted to hosts which grow
seasonally; they persist during the winter, when few or no roots are avail-
able for invasion, as dormant eggs contained within a hardened cyst formed
from the dead body of the female. Eggs of potato cyst nematodes contained
within their cysts can withstand drying, unlike the eggs of most other nema-
todes. Consequently, potato cyst nematodes can be transported over long
distances in contaminated soil adhering to tubers or any other root crops.
In years when non-host crops are grown only a small proportion of
.... _- ----
~- Globodera pal/ida
Figure 11.1 The known world distribution of the two species of potato cyst
nematode; the arrows indicate their probable movements.
potato cyst nematode eggs hatch, but when a host crop is grown most o{ the
dormant second-stage juveniles are stimulated to hatch by a specific
chemical 'hatching {actor' diffusing from the host roots. The juveniles are
attracted to and invade the roots, cutting a path through the cortical cells
until they reach the stele where they induce formation of specialized
transfer cells (syncytia) on which the now sedentary juveniles feed. The
syncytia are formed mainly from phloem parenchyma cells by the break-
down of cell walls and enlargement of the cells. Special wall ingrowths de-
velop on the walls adjacent to vessels to increase the surface area of the plas-
malemma (Fig. 11.2) and so increase the rate of transport of solutes (Jones
and Northcote, 1972). During feeding, the nematodes produce coiled,
tube-like structures from their stylet tips. These 'feeding tubes' are thought
to enable the nematode to extract only the cytosol (Rumpenhorst, 1984).
Provided a juvenile induces a syncytium of sufficient size it receives all
the food necessary to develop into an adult female, but when the syncytium
is inadequate for this purpose the juvenile either dies or becomes a male.
Experimental evidence suggests that food availability may affect the sex of
the developing juveniles, poorly nourished ones becoming males and well-
nourished ones becoming females (Trudgill, 1967; Mugniery and Fayet,
1981). After fertilization the female deposits eggs in her own body, her
cuticle tans and when she dies the dead 'cyst' containing hundreds of eggs is
left in the soil. In the absence of host crops soil infestations may persist for
20-30 years, and it is in this dormant condition that potato cyst nematodes
have been spread, either in infested soil or adhering to tubers harvested for
Effects of root-feeding nematodes on growth and yield 447
use elsewhere as seed. The large reproductive capacity of these nematodes
(up to 70-fold and more) and their slow rate of decline in the absence of a
host crop (about 30% per annum) make them a persistent and serious pest
of potatoes.
Living females of G. pallida and G. rostochiensis on potato roots can be dis-
tinguished by their colour; those of G. pallida remain white throughout their
lives whereas those of G. rostochiensis become yellow (Guile, 1966). Other
differences used to discriminate the species are juvenile stylet length and
shape (Webley, 1970) and the proteins separated by electrophoresis (Trudgill
and Carpenter, 1971; Bakker and Gommers, 1982; Fleming and Marks,
1983). There are also small differences in their biology which suggest that
G. pallida is adapted to slightly cooler conditions than G. rostochiensis
(Foot, 1978; Mugniery, 1978a; Franco, 1979; Webley and Jones, 1981).
However, Hominick (1979, 1982) has shown that, in an area where early
maturing potatoes are grown each year, G. rostochiensis has adapted to
hatch and develop at lower temperatures than other populations of this
species. In soils in which the temperature during the growing season exceeds
30°C, neither species survives (Trudgill, 1970; Evans and Stone, 1977).
The damage caused by both species is similar: affected crops are stunted,
have a tendency to wilt and die early (Grainger, 1951). Typically, field
damage occurs in patches, the cause of which can be readily confirmed by
examining the roots for developing females.
11.2 EFFECTS OF ROOT -FEEDING NEMATODES ON GROWTH AND

YIELD
The classical proof of the role of pathogens in causing disease is that they
satisfy Koch's postulates. To do this they must be isolated from a diseased
plant and, when introduced to a healthy plant, re-establish the disease. The
causal organisms must then be re-isolated from the newly infected plant.
Because most root-feeding nematodes multiply very s,lowly compared to
Figure 11.2 (A) Transverse section of potato root showing giant cell complex
around head of female potato cyst nematode. x 240. (B) Whole mount of female
potato cyst nematode in potato root showing giant cell complex associated with the
stele. x 200. (C) Scanning electron micrograph of a longitudinal section of root
infected by soybean cyst nematode with cell contents digested away to show
dissolution of cell walls and formation of a syncytium. x 200. (D) Scanning electron
micrograph of a longitudinal section of a root infested by tobacco cyst nematode
showing dissolved cell walls and wall ingrowths. x 1000. (E) Wall ingrowths in
giant cells induced by root knot nematodes in roots of balsam. x 3000. n,
nematode; gc, giant cells; wi, wall ingrowths.
(A) Preparation by R.A. Rohde, photograph by c.c. Doncaster.
(B) Preparation by D.L. Trudgill, photograph by Rothamsted Experimental
Station (C-E) by M.G.K. Jones.
fungi and bacteria, it may not be possible to establish, in simple test
conditions, population densities which exceed the threshold for plant
damage and so allow the diseased condition to be reproduced. Repeated
cropping with a host may be necessary before the population density
becomes damaging, so a rigorous satisfaction of Koch's postulates is
difficult and the establishment of the role of nematodes in causing plant
damage is frequently based upon less direct evidence.
The similarity of above-ground symptoms caused by all types of root-
feeding nematodes suggests that models developed from work on relatively
few species may be applied to infestations by most root-pathogenic species.
Seinhorst (1965, 1979) proposed that the relationship between total plant
weight (Y) and initial density of nematodes (P) can be expressed by the
equation
Y = m + (1 - m) ZP-T
in which m is the minimum yield, T is the nematode population density
corresponding to the crop's tolerance limit (below which yield is unaffected
by change in nematode density) and z is a constant. Yields are expressed as
a proportion of the nematode-free yield and, when the initial nematode
population density is expressed on a logarithmic scale, the relationship is
sigmoidal. Seinhorst and den Ouden (1971), Seinhorst (1982a) and Greco
et al. (1982) have shown experimentally that this model can be applied to
the relationship between initial density of potato cyst nematodes and yield
of potato tubers.
However, a major limitation of this and other models relating pre-
planting numbers of nematodes to crop yield is their failure to take account
of known interactions with various environmental factors (e.g. soil type -
damage is usually greater on light than heavy soils) or differences in yield
potential between sites and between cultivars. Groups in The Netherlands
(Ward et al., 1985) and the UK are seeking to overcome these limitations.
Seinhorst's model has too many unknowns to be used for predictive
purposes and Brown (1969) used untransformed linear regression analysis
to compare yield losses and nematode population density at many sites
infested with potato cyst nematodes. He conluded that average losses were
2.13 t ha- 1 for each increase of 20 eggs g-l soil. In a further series of
experiments Brown and Sykes (1983) found that damage differed markedly
between sites and that average losses had risen to 6.2 t ha- 1 for each
increase of 20 eggs g-l soil.
For most other nematode species attacking potatoes, information on the
relationship between pre-planting nematode population density and yield
loss is scarce. Brown and Sykes (1975) estimated that losses due to
Longidorus elongatus were 3 t ha- 1 for every 100 nematodes per 200 g soil;
Olthof and Potter (1972) reported that potato yields were decreased
significantly by 666 M. hapla per kg soil; Oostenbrink (1958, 1961) and
Dickerson et al. (1964) indicate that there is a relationship between initial
population density of P. penetrans and potato yield losses.
The mechanisms of damage by potato cyst nematodes have been
Effects of root-feeding nematodes on growth and yield 449
extensively investigated. Much of the damage is caused by the invading
juveniles which destroy root cells and inhibit root growth (Evans, 1982a;
Storey, 1982) with lateral roots more affected than primary roots (Evans et
al., 1977; Trudgill and Cotes, 1983b). For most and probably all of the
growing season, infected plants, especially those of intolerant cultivars,
have smaller root systems (Trudgill, 1980; Evans, 1982a; Trudgill and
Cotes, 1983b) that explore a smaller volume of soil (Table 11.2) than do
those of uninfected plants. The damage decreases the uptake of most
minerals, N, P and K being particularly affected (Trudgill et al., 1975a,b),
leading to chronic nutrient deficiency (Trudgill, 1980, 1987) and a reduced
rate of top growth (Fig. 11.3). Uptake of Ca, however, is much increased
and this may be due to endodermis damage (Price and Sanderson, 1984).
The number of leaves per stem is not affected but leaf size, internode
length and numbers of stems are all decreased (Table 11.3). The decrease
Table 11.2 The effect of different nematode population densities on the

estimated volumes of soil (m 3 X lO-2) exploited by potato plants* (from
Evans et ai., 1977)
Pentland Dell Maris Piper
Nematode population density Small Large Small Large Larget
Depth
Ridge soil 1.8 0.7 2.7 2.7 1.1
Ploughed layer 2.1 0.1 3.0 1.5 1.3
Sub-soil 0.6 0.0 1.0 0.1 0..3
Total 4.5 0.8 6.7 4.3 2.7
Yield of tubers per plant (g) 1071 200 1424 977 815
Total K in tops and tubers (g) 7.8 1.7 9.4 5.8 2.9
* NB It is assumed that nutrients move to roots over distances of up to 0.5 em.
t G. pallida, all other populations G. rostochiensis.
100
§ ~:::=:=:=.~
1 J//~~ \~O~
60
~ 20 f-/ \ .,.\ \Few

f-I> '" Many.,. Intermediate
oL-~~--~--~---~~L-~~--~--
8 12 16 20
Weeks after planting
Figure 11.3 Changes, with time, in ground cover of Pentland Dell potato plants
infested with few, intermediate numbers or many potato cyst nematodes. (From
Evans et al., 1975b).
Table 11.3 Growth analysis and Nand K contents of Pentland
Dell potato plants heavily or lightly infested with G.
rostoehiensis, 10 weeks after planting (from Trudgill et a!.
J975a)
Heavily infested Lightly infested
(unfumigated) (fumigated)
Haulm
Fresh weight (g) 199 538
Dry matter (%) 9.2 8.1
Height (em) 34.2 43.4
Number of stems/plants 4.2 8.6
Number of nodes/stems 13.0 11.2
Number of leaves/stem 11.6 10.6
Leaf area/plant (em 2) 1598 5882
Area each leaf (em 2) 32.8 64.5
N in dry matter (%) 4.5 4.8
K in dry matter (%) 4.0 6.8
Tubers
Fresh weight (g) 49 77
Number of tubers 15.4 19.4
Dry matter (%) 14.6 13.6
K in dry matter (%) 2.3 2.5
Roots
Fresh weight (g) 48.6 104.2
in top fresh weight is proportionally greater than that of the root system
(Evans, 1982a).
Because potato cyst nematodes decrease the rate of top growth they also
decrease the amount of solar radiation intercepted by the leaves and
thereby total assimilation. Water uptake does not appear to be greatly
affected in the early stages of growth hut, as plant size and temperatures
increase and water availability decreases, infected plants often become
more water stressed than uninfected ones (Evans et al., 1975b; Evans,
1982a,b). Infected plants therefore tend to wilt and to lose their lower
leaves. The efficiency of water use, as measured by the transpiration ratio,
is also decreased (Evans, 1982b).
Assimilation efficiency (increase in dry matter per unit of light inter-
cepted) may also be decreased for some cultivars. A comparison of the
effect of heavy infestations of potato cyst nematodes on light interception
and total tuber dry matter at harvest, using data from Evans (1982a),
showed (Table 11.4) that both were decreased by a similar percentage for
cultivars Cara, Maris Peer and Pentland Crown, but for cv. Pentland Dell
the reduction in tuber dry matter (64%) was much greater than that for
light intercepted (42%). These differences were paralleled by a difference
Interactions with other organisms 451
Table 11.4 Tuber dry matter yields and amounts of light intercepted by four
cultivars grown in plots lightly or heavily infested with G. rostochiensis (data from
Evans, 1982a)
Maris Peer Pentland Dell Pentland Cara
Crown
Initial nematode 8 105 8 105 8 105 8 105
population density
Pi (eggs got)
Fresh tuber yields 24.8 11.0 46.3 13.4 55.9 31.3 59.4 54.6
(t ha- t )
Estimated dry tuber 4.2 2.3 7.9 2.8 9.5 6.6 10.1 9.3
yields (t ha· t)*
Reduction in dry 45.3 64.3 30.8 8.1
matter yield (%)
Total light 667 366 837 484 991 690 1209 1031
intercepted
(MJ m-2 )
Reduction in light 45.1 42.2 30.4 14.7
interception (%)
• Percentage dry matter estimated at 17% in tubers from four plots with Pi = 8 eggs g-! and
21% from those with Pi = 105 eggs g.! (except for Cara where 17% was used). No estimate
was made of the amounts of dry matter remaining in the tops.
in stomatal resistance: that of heavily infested Pentland Dell was increased

13 weeks after planting whereas that of Cara was almost unchanged. The
effects of potato cyst nematodes on crop growth are discussed in greater
detail by Trudgill (1986).
The effects on plant growth of other root-feeding nematodes are
essentially similar to those of potato cyst nematodes. Meloidogyne spp.
seem to have a particularly marked effect on plant water relations.
O'Bannon and Reynolds (1965) found that cotton plants infected by
Meloidogyne spp. grew as well as uninfected plants when the soil was kept
at field capacity, but that they used more water per unit weight. When the
soil was allowed to dry to 50% of field capacity between waterings infected
plants grew more slowly than uninfected ones. Odihirin (1971) found that
at first M. incognita greatly increased the rate of transpiration of infected
tobacco plants but that eventually the plants wilted even when abundant
water was available, apparently because the roots became too severely
damaged to meet the demand for water. Wallace (1974) evaluated the
effects of root knot nematodes on photosynthesis and nutrient demand by
roots of tomato plants, and concluded that the nematode's most important
effect was probably on water uptake and thereby on photosynthesis.
Other ways in which nematodes may affect plant growth include
disturbing the balance of plant growth substances. Setty and Wheeler
(1968) found that roots of tomato plants infected with Meloidogyne spp.
contained more auxin than those of uninfected plants.
11.3 INTERACTIONS WITH OTHER ORGANISMS
When plants are infected by a combination of nematodes and pathogenic

fungi or bacteria the damage they suffer is often greater than the sum of
that which could be caused by the organisms singly. Morgan (1926)
suggested that 'potato sickness' was caused by a combination of
Rhizoctonia solani and potato cyst nematodes, a view supported by
Grainger and Clark (1963) and by Dunn and Hughes (1967) who showed
that there was also an interaction with Oospora pustulans. Corbett and
Hide (1971) showed that yield losses were increased when plants were infec-
ted with a combination of G. rostochiensis and Verticillium dahliae. Evans
(1987) reported an interaction between G. pallida and V. dahliae to which
early maturing cultivars were usually (not always) more susceptible than later
maturing cultivars and Storey and Evans (1987) were able to relate the
variation amongst early cultivars to differences in root anatomy. Trudgill et al.
(1975b) observed that treatment with dazomet, a partial soil sterilant,
improved potato growth in field trials more than could be accounted for by
the control of cyst nematodes alone. They suggested, as did Brown (1983),
that fungi might be interacting with the nematodes to increase the damage
caused. Jatala et ai. (1976) investigated the interaction between G. pallida
and Pseudomonas solanacearum and found that their effects on growth
were synergistic, even on potatoes that were resistant to the nematode.
In North America there is considerable evidence that 'early dying'
disease of potatoes is caused by a combination of P. penetrans and V. albo-
atrum. Cetas and Harrison (1963) suggested that Verticillium wilt was more
severe in soils heavily infested with P. penetrans and Biehn et al. (1971)
that control of P. penetrans delayed infection by V. albo-atrum and the
expression of wilt symptoms. Morsink and Rich (1968) also found inter-
actions between P. penetrans and V. albo-atrum on potatoes. However,
Burpee and Bloom (1978) and Gould (1974) indicated that the effects of
the two pathogens might simply be additive rather than interactive. The
severity of wilt in cultivars tolerant of V. dahliae was increased by P. thornei
(Krikun and Orion, 1977) leading to 30-40% yield losses and Martin et ai.
(1982) found an interaction between V. dahliae and P. penetrans.
Root knot nematodes are noted for their role in pre-disposing many
species of plant to infection by other pathogens, so resulting in disease
complexes. In potatoes the severity of Verticillium wilt is increased by M.
hapla (Jacobsen et al., 1979) and damage by R. so/ani may also be
increased (Joubert and Dalmasso, 1972). Infection by M. incognita has
been reported to increase the severity of bacterial wilt (caused by P.
solanacearum) and to advance the expression of symptoms (Feldmesser
and Goth, 1970).
Infection by Spongospora subterranea of the galls produced by N.
aberrans is thought to cause extra yield loss (Brodie, 1984) but there are
many difficulties in the study of interactions and it seems likely that there
are many more interactions which have yet to be identified.
Methods of decreasing nematode damage 453
11.4 METHODS OF DECREASING NEMATODE DAMAGE
Damage due to nematodes is avoided if they are absent from the

environment in which a crop is grown. Hence, considerable effort is
devoted to statutory regulations which aim to limit the further spread
of serious pests such as potato cyst nematodes. If a pest is present,
however, its numbers can be decreased to levels where yield losses will be
insignificant, by:
1. chemicals;
2. a variety of biological means such as growing resistant or non-host
cultivars or crops;
3. planting the crop in cool soils where its roots grow but the nematodes
are not very active;
4. harvesting the crop before the pest has completed its life cycle;
5. possibly the use of natural pathogens or predators for biological control.
Damage may also be decreased by growing cultivars which are particularly
tolerant of nematode attack or by modifying crop husbandry. In heavily
infested soils an integration of several methods of decreasing damage is
usually necessary.
11.4.1 Legislative control

Nematodes which are endoparasitic or which, like potato cyst nematodes,
can withstand drying, are readily spread by infested seed tubers.
Certification schemes which aim, amongst other things, to produce
nematode-free seed are therefore a foundation of legislative controL In
many countries it is a requirement that seed potatoes are produced on land
free of pathogenic nematodes, particularly potato cyst nematodes, so
that the seed tubers are free of infection. In the European Community,
seed-producing land must be sampled and if potato cyst nematodes are
found that land is scheduled: the growing of seed potatoes is then
prohibited. Also, even on clean land the frequency with which seed
potatoes may be grown is restricted. In practice, this legislation is only as
effective as the sampling and detection methods employed; and the
difficulties of detecting small infestations become clear when it is realized
that one cyst in 500 g soil (the usual sample size) may represent more than
10 million cysts per hectare.
Potato cyst nematodes are recognized as a serious threat to potato
production and, as they are amongst the most difficult of all crop pests to
control, most countries from which they are thought to be absent have
strict regulations governing the importation of any produce that might
carry them. This includes seed potatoes, transplants, nursery stock, bulbs
and corms. Quarantine regulations may also be imposed within a country
to prevent further spread once the pest is known to have entered. For
example, in New York State on mainland USA and on Long Island large
areas are quarantined and, in addition to stringent regulations concerning
movement from these areas of hay, straw, propagating material and farm
machinery, all potatoes grown must be packaged in non-reusable paper
bags. Potatoes may not be grown on or near land known to be infested
(Spears, 1968). However, as indicated above, it is difficult to detect light
infestations of potato cyst nematodes and they may be spread over a wide
area before their presence is noticed (Jones, 1969).
11.4.2 Chemical control

Treatment of soil on a field scale with chemicals to control nematodes is
costly but many thousands of hectares are now treated annually. The
chemicals used may be fumigants or non-volatile organophosphates and
oxime carbamates. Fumigant nematicides are usually biocides and there-
fore kill nematodes. Organophosphate and oxime carbamate nematicides
are nerve poisons and seem to disorientate nematode juveniles and so
prevent them from invading plant roots (Boparai and Hague, 1974). All
are generally expensive (so good yield responses are necessary to justify
their use) and special precautions are required during the application of
non-volatile nematicides because of their extremely poisonous nature.
Farmers apply nematicides firstly to improve the growth and yield of the
current crop and, secondly, to prevent nematode multiplication, so that
following host crops are less severely damaged. Fumigant nematicides
rarely kill sufficient nematodes to prevent the population increasing
(Whitehead, 1986). However, good yield responses are generally obtained
because the young plants are protected from severe damage, other
pathogens may be killed, and the availability of nitrogen may be increased.
When properly applied, non-volatile nematicides can prevent nematode
multiplication and are effective at doses as low as 3 kg ha- 1 . Of these, the
oxime carbamates aldicarb and oxamyl are particularly effective in
virtually all soil types (Whitehead, 1973) but, like fumigant nematicides,
the organophosphate types seem to work less well in organic soils
(Whitehead, 1975).
Early research on the chemical control of potato cyst nematodes
concentrated on soil fumigation and double treatments were used in the
USA to decrease population densities to undetectable levels (Spears,
1968). Fumigant nematicides have now been largely replaced by non-
volatile types and the effectiveness of these has been improved by better
methods of incorporation. New application methods may lead to decreased
application rates and, in view of the recent implication of aldicarb in
contamination of ground water on Long Island, USA (Brodie, 1984), this
is an important goal.
Some success in the control of Ditylenchus destructor by fumigation
using ethylene dibromide has been achieved in the USA (Thorne, 1961).
This method of control has also been effective against root knot nematodes
and, when nematode densities are great, fumigation can decrease them by
up to 99% with increases in yield of up to 55% (Winslow and Willis, 1972).
Non-volatile nematicides have shown promise for control of root knot
nematodes on potatoes (Abdel-Rahman et al., 1974) and as a seed-piece
dip to reduce infection of seed (Rodriguez-Kabana and Ingram, 1977).
Infection of seed can also be reduced by heat treatment (Nirula and Bassi,
1965; Martin, 1968). Centre pivot irrigation systems provide a convenient
method for applying chemicals directly to crops and Santo and Qualls
(1984) describe control of M. chitwoodi and M. hapla damage to potatoes
by injecting metham sodium into irrigation lines.
Fumigants have been used to control Pratylenchus spp. on potatoes but
the yield increases obtained may not always be economic. When Verticillium
wilt fungus was also present, Hawkins and Miller (1971) reported yield
increases of 10--14% using Vorlex. Non-volatile nematicides are more
practical for controlling Pratylenchus spp. on potatoes and oxamyl has
been used as a seed-piece dip to control P. brachyurus (Rodriguez-Kabana
and Ingram, 1977).
Brown and Sykes (1973) found non-volatile nematicides to be effective
in decreasing transmission of tobacco rattle viruses to potato tubers by
Trichodorus spp. Trichodorid nematodes have been more difficult to
control by fumigation, apparently because they reside deeper in the soil
(Brodie, 1984). The same was true for control of Longidorus leptocephalus
by Telone, which only killed nematodes in the top 30 cm of a sandy soil;
chloropicrin, however, killed nematodes to a depth of 60 cm (Evans,
1979).
11.4.3 Non-chemical control

Crop rotation is one of the oldest and most important means of controlling
indigenous nematodes. It is particularly effective with those that have
narrow host ranges (e.g. potato cyst nematodes) or those which persist
poorly in the absence of their host (e.g. Nacobbus and Meloidogyne
spp.). However, some nematodes (e.g. Meloidogyne spp.) which persist
poorly have wide host ranges and this greatly increases the difficulties of
induding acceptable non-hosts within the rotation. In Zimbabwe M.
javanica is effectively controlled by growing certain grasses before potatoes
(Jensen et al., 1979) and in the north-western USA the population density
of M. hapla is greatly decreased by cereals; it is, however, essential to
identify M. hapla infestations accurately because M. chitwoodi occurs in
the same area and is not controlled by growing cereals (O'Bannon and
Santo, 1984).
Crop rotation is widely used as a means of reducing potato cyst
nematode population densities below the threshold for economic damage.
In Britain, populations decrease typically by about 30% per annum in the
absence of a host crop and, in practice, potatoes can be grown once every 4
or 5 years in heavy soils, but only once every 6 or 7 years in light and peat
soils (Cooper, 1953; Winfield, 1965). Potato cyst nematodes may adapt to
longer rotations by becoming more persistent but they persist less well in
warm than in cold regions (Schluter, 1976). In very warm soils potato cyst
nematodes are unable to establish, e.g. irrigated coastal areas of Peru,
where the nematodes are able to reproduce if introduced to a potato crop
but are unable to survive in the hot dry soil between growing seasons
(Simon, 1955).
The effectiveness of rotation can be increased by eliminating alternative
hosts. Certain weeds are important hosts for D. destructor and must be
eliminated if the nematodes are to be kept under control (Gratwick, 1989).
Other nematodes, such as Pratylenchus penetrans, have such wide host
ranges that they cannot be controlled by crop rotation. With these, the
need is to avoid planting tolerant crops which are good hosts immediately
before intolerant crops, e.g. rye before potatoes on land infested with P.
penetrans (Brodie, 1984). Some husbandry practices can be used to
decrease nematode population densities. A good example is early lifting of
potatoes before potato cyst nematode females have completed their
development (van den Brande and d'Herde, 1964; Mugniery, 1978a,
Webley and Jones, 1981).
Plant-parasitic nematodes are prey for many other organisms but there
have been no reports of nematode parasites of potato being naturally
controlled, although the population densities of the cereal cyst nematode,
Heterodera avenae, are frequently held under natural control by fungi
(Kerry, 1974). For nematodes of potato the most promising work is that of
Jatala et al. (1979) who found that the fungus Paecilomyces lilacinus
infected 70-90% of G. pal/ida eggs and 80-90% of M. incognita acrita eggs.
The bacterium Pasteuria penetrans is also considered a potentially effective
bio-control agent (Kerry, 1987).
Resistant cultivars (those that are non-hosts or only very poor hosts) also
control nematodes in an environmentally safe and biologically sound
manner. Provided they are agronomically satisfactory, resistant cultivars
have several advantages over chemicals, a major one being that their use
involves the farmer in little additional expense. Cook and Evans (1987)
make a comprehensive review of breeding for resistance to plant parasitic
nematodes. Sources of resistance to most of the important nematode pests
of potato seem to be available but breeding for resistance to potato cyst
nematodes is the most advanced. Ellenby (1954) identified resistance to
potato cyst nematodes in several clones of S. tuberosum ssp. andigena and
this resistance, which is conferred by a single major gene (HI)' has been
incorporated in many excellent cultivars. Gene HI confers resistance to all
British (and those derived from Britain) populations of G. rostochiensis,
many in Europe, and others around the world (e.g. those in the eastern
United States) where G. rostochiensis Rol is the only pathotype present.
The Hl gene may also confer partial resistance to P. penetrans (Brodie,
1984), preventing populations of P. penetrans from increasing on the
potato crop and conferring a measure of tolerance. Parrott (1981) showed
that virulence in G. rostochiensis against the Hl resistance gene is
conferred in the nematode by a complementary recessive gene which
occurs widely in South American and some European populations but not
in Britain. Thus, again it seems likely that only a part of the nematode gene
pool has been introduced into Britain. Bakker (1987), using 2-dimensional
electrophoresis of proteins from a large number of European populations
of both G. rostochiensis and G. pallida, has shown that many alleles
present in founder populations have been progressively lost as these
nematodes were spread within Europe.
Resistance to virulent pathotypes of G. rostochiensis and to G. pallida
has been identified in the wild species S. vernei and this source forms the
basis of breeding programmes in several countries. Considerable back-
crossing to S. tuberosum has been required to eliminate undesirable
characteristics but resistant cultivars for starch production containing ex-So
vernei resistance have been available for several years in The Netherlands
and 'table' cultivars are now becoming available. A second important
source of resistance, effective against G. pallida but not G. rostochiensis,
was identified in the clone S. tuberosum ssp. andigena CPC 2803 (Howard
et al., 1970) and table cultivars with this resistance should soon be
available. A further difficulty in breeding for resistance to G. pallida is that
resistance from both these sources is inherited quantitatively and crosses
with S. tuberosum ssp. tuberosum produce progenies with variable degrees
of resistance. The problems of assessing and quantifying this resistance and
its effect on potato cyst nematode population densities in the field are the
subject of much research and discussion (Phillips and Trudgill, 1983;
Phillips, 1984; Dellaert and Vinke, 1987). The most consistent method of
assessing this resistance is to relate that of the test clones to that of a series
of internationally agreed standard test cultivars (Phillips et al., 1989). In
collaborative experiments in several European countries it was found that,
despite variations in the absolute multiplication rates achieved by several
nematode populations on a range of partially resistant clones, the resis-
tances were ranked in a similar order by each population. This has led to a
recommendation to the European Plant Protection Organization for
standard resistance testing procedures (Mugniery et al., 1989). However,
there is some concern that this resistance is unlikely to prove completely
durable (Stone and Turner, 1982). Other sources currently being used to
breed for resistance to potato cyst nematodes include S. multidissectum
and S. sanctae rosae (both of which confer the major gene H 2 ), S.
speggazzinii and S. gourlayi (Uhrig and Wenzel, 1981).
Cultivars resistant to several other nematodes attacking potatoes have
been bred. Resistance to D. destructor has been introduced in several
cultivars (Jensen et al., 1979) and sources of resistance to D. dipsaci have
been identified (Olefir, 1972). Several sources of resistance to Meloidogyne
spp. have been found in wild and cultivated species of Solanum and good
resistance to M. incognita, M. javanica, and M. arenaria and to the false
root knot nematode (Nacobbus) has been identified in S. sparsipilum
(Jatala and Rowe, 1976; Mai et al., 1981) and to M. hapla in S. tuberosum
ssp. andigena (Brodie and Plaisted, 1977). Breeding for resistance to
Nacobbus and to Meloidogyne is, however, often complicated by dif-
ferences in virulence between populations (Canto Saenz, 1984) and the
expression of resistance is sometimes affected by temperature.
Frequently, new sources of resistance reveal populations of nematodes
able to overcome the resistance, leading to the recognition of pathotypes
or races. Several schemes have been proposed for identifying pathotypes
of potato cyst nematodes, the most recent being the related schemes
proposed by Kort et al. (1977) and Canto Saenz and de Scurrah (1977).
These schemes used an absolute multiplication rate of greater or less than
one and clones with quantitatively inherited resistance to differentiate
pathotypes and have proved unworkable (Stone and Parrott, 1985, Trudgill,
1985). Using clones with major gene resistance two pathotypes of G.
rostochiensis and two of G. pallida can be distinguished in Europe on the
basis of their ability to overcome the Hl (ex-S. tuberosum ssp. andigena)
and H2 (ex-S. multidissectum) genes respectively. Populations of G. pal/ida
and G. rostochiensis do differ in their rates of multiplication on cultivars
with quantitative resistance (Kort et al., 1977; Phillips and Trudgill, 1983)
but such population differences should not be classified as distinct
pathotypes.
11.4.4 Damage avoidance

There is a well established relationship between soil population densities of
potato cyst nematodes and potato yield (see Section 11.2), so estimates of
the population density can be used to decide whether to grow potatoes or
not, whether to use a nematicide and which cultivar to choose (resistant,
tolerant, etc.). Careful monitoring of the nematode population density
helps decrease losses and services to farmers such as that provided by the
Agricultural Development and Advisory Service (AD AS) in England and
Wales are valuable in this respect.
Early maturing potato cultivars are frequently grown in relatively frost-
free areas of the UK (e.g. the Ayrshire coast of Scotland; Pembrokeshire;
Cornwall) in order to capture the early potato market. The absence of
frosts allows early planting at a time when potatoes will grow but soil
temperatures are too low for much nematode activity (see Mugniery,
1978a, for base temperatures for nematode activity). By the time soil
temperatures rise to levels which favour nematode activity, the plants
are well established and they suffer less damage. Also, the early harvest
means that fewer females have matured and started to produce eggs, so
post-harvest population densities are low and the challenge to subsequent
potato crops is not severe. Because populations are held in check by early
lifting, repeated annual cropping with potatoes is possible and some
growers are even exploring the possibility of two potato crops in one year,
the second being protected with a nematicide.
Late planting can also be of benefit for certain problems. Kirjanova and
Krall (1971) found that soil moisture was inadequate for optimum invasion
of tubers by D. destructor in late planted crops and that early harvest was
helpful in providing healthy seedstock.
Because the primary effect of root-feeding nematodes is on root growth,
attacked plants show symptoms of water and nutrient stress. These
symptoms can be partially alleviated by applying irrigation or additional
fertilizers (Jones, 1977; Jensen et al., 1979; Trudgill, 1987). Extensive use
of irrigation might increase problems caused by those nematodes that
require moisture films in light soils for activity, viz. activity of trichodorid
nematodes may be increased during and after tuber initiation so that they
are able to transmit TRV to the tubers.
Huijsman et a1. (1969) tested 118 potato cultivars and found that Multa
was significantly more able to tolerate potato cyst nematode attack than all
others. Evans and Franco (1979) showed that such differences in tolerance
could lead to large differences in yield and their results were confirmed by
Evans (1982a), Trudgill and Cotes (1983a) and Whitehead et al., (1980).
Haydock (1989) attempted to increase tolerance of G. pal/ida attack in
several cultivars by physiological ageing of the seed tubers prior to
planting. It was hoped that faster initial growth rates from physiologically
'older' seed would enable the plants to withstand nematode attack better;
when this occurred it was offset by earlier senescence of the crop and yields
were not significantly improved by physiological ageing.
At infested sites tolerant cultivars will grow better and yield more than
intolerant ones but they will also support greater nematode multiplication.
This would constitute a serious threat to subsequent crops and even
tolerant cultivars might eventually yield poorly. Although cultivars that are
resistant to potato cyst nematodes prevent nematode reproduction, they
are still invaded by the juveniles and may suffer considerable damage.
Combining resistance and tolerance would therefore have the dual
advantages of preventing the numbers of nematodes from increasing to
unacceptable levels on tolerant cultivars and minimizing the damage
suffered by resistant cultivars as a consequence of invasion. Tolerance has
proved to be an especially important goal· in the breeding of potato
cultivars with quantitatively inherited resistance to G. pal/ida, where
intolerance is very common (Trudgill and Cotes, 1983a). Tolerance alone
would be useful where the nematode is endemic and suitable resistant
cultivars are not available; such a situation is found in relation to pota,to
cyst nematodes in certain parts of South America.
Screening plants for tolerance in a breeding programme is very difficult
because tolerance can ony be identified easily in field grown plants. Evans
(1982a) reviews field trial designs which are useful for assessing tolerance
and Trudgill and Cotes (1983a) suggest the use of nematicide trials
followed by genotype x environment analysis of the results to identify
tolerance. Trudgill (1986) reviews the importance of tolerance in
minimizing yield losses in potatoes due to potato cyst nematodes.
The multiplication rates of nematodes are density dependent and several
models describing the reduction in multiplication with increasing pre-
planting population densities have been formulated. Seinhorst (1967)
produced an equation which links the effect of nematode density on
reducing host growth with its effect on overall nematode multiplication
rate. Such information is important and this approach has much to
commend it. Jones and Perry (1978) formulated equations relating to the
population dynamics of cyst nematodes which allowed for the use of
resistant cultivars and nematicides and should therefore be helpful in
predicting their effects.
11.4.5 Integrated control

No presently available method of controlling the nematode pests of
potatoes is ideal: control solely by rotating potatoes with non-host crops
restricts the frequency of cropping with potatoes to less than many farmers
would wish; cultivars resistant to all species and pathotypes are not yet
available and the use of those that are encourages multiplication of virulent
portions of the population; chemical control is expensive, sometimes of
uncertain outcome and involves the handling of highly toxic materials.
Jones (1970) outlines various rotations which incorporate the use of
resistant cultivars with and without nematicides and shows that very large
reductions in the population density of potato cyst nematodes may be
obtained, even with quite short rotations, so that a non-resistant cultivar
might then be grown. Including a non-resistant cultivar may have the
advantage of slowing selective multiplication of virulent pathotypes, so
prolonging the useful life of resistant cultivars (Trudgill et al., 1987). The
use of a nematicide may have additional benefits on land where harmful
nematodes such as Longidorus and Trichodorus spp. are troublesome.
Other nematodes may also be killed and in The Netherlands, where the use
of nematicide has been compulsory for a number of years when potatoes
are grown frequently on potato cyst nematode infested land, considerable
additional benefits have been obtained through the control of H. avenae
and M. hapla.
11.4.6 Cost effectiveness of control methods

In order to assess the cost effectiveness of a nematode control measure we
need to know the potential losses with no control and the likely benefits
and cost of the control measure. The overall losses caused by potato cyst
nematodes are difficult to assess but there can be little doubt that they
cause sufficient damage to justify considerable efforts to control them. If a
nematode pest is thought to be absent from an ar~a--it is worth trying to
avoid or delay its introduction, but once fields are infested the farmer is
faced with control using rotation, nematicides or resistant cultivars. High
yielding, good quality resistant cultivars are much the best method of
control as there is little extra cost to the farmer. Resistant cultivars take
time and are costly to develop, but Bottrell (1979) estimated that in the
USA pest and disease resistant cultivars earn $300 for each $1 spent on
research. Lawes (1988) estimated the cost of producing new cereal
cultivars in Britain (1982 prices) at £0.5-O.75M each, and Cook and Evans
(1987) give figures of £1.5m per new potato cultivar in the UK and $lm in
the USA.
Appropriate resistant cultivars are not always available and the only
alternative open to many farmers is to employ a rotation designed to keep
the numbers of nematodes below damaging levels. Empsom and James
(1966, 1967) examined the economics of enforced changes in rotation and
found that lengthening the period between potato crops from 3 to 5 years
was profitable on farms where all fields were infested, provided the yield
potential was 10 ton (10.16 tonnes) acre-I, but less so when it was 15 ton
(15.24 tonnes) acre-I. In order to establish suitable rotations, the distri-
bution and numbers of nematodes have to be determined. However, the
value of such information is limited by the inaccuracies associated with
using relatively small samples to detect or estimate nematode populations
(Seinhorst, 1982b). Even so, Adamson (1972) estimated that the value of
the advice given after processing each soil sample might be as much as £87
(at 1970 prices and assuming each sample covered 5 acres. Based on advice
given to potato growers in S.W. Lancashire). The cost of maintaining an
appropriate service to provide this advice should be considered when costs
of control are assessed on a national scale. More obscure costs are also
involved such as not growing potatoes on the best potato land as often as
would be desirable, the cost of nematode-free certification schemes for
seed potatoes and propagating material such as bulbs, and the potential
harm to a country's export trade in seed potatoes and other planting
material. If a pest can be totally excluded from a country many of these
costs are avoided, but they are replaced by those of monitoring land for
appearance of the pest and maintaining quarantine measures.
Despite the availability of cultivars with good resistance to G. rostochiensis
in the UK, chemical control is a vital part of the management used to
minimize yield losses due to potato cyst nematodes. This is because G.
paUida has become increasingly important and potato production has
tended to become concentrated on the best potato land. Agronomically
desirable cultivars with full resistance to G. paUida are not yet available, so
a considerable proportion of the potato crop is treated with nematicides.
According to Trudgill (1986) more than 20000 ha were treated in 1982.
With a current price for the materials of c. £250 ha- 1 , the annual cost of the
chemicals alone exceeds £5m, but the improved yields obtained following
nematicide use are likely to be worth as much as £25m.
11.4.7 Conclusions
The extensive movement of machinery and planting material means that
nematodes are continually being introduced to new areas, and that
eventually all species will have access to all areas which provide a suitable
environment and suitable hosts. However, quarantine controls, such as the
United States Department of Agriculture's campaign against the golden
nematode (G. rostochiensis), can be effective in delaying introduction of
pests. The golden potato cyst nematode was first introduced on Long
Island in 1941, and in 1944 New York State instituted quarantine pro-
cedures. Systematic surveys showed that many properties on the island
were infested but it was not until 1967 that the nematode was discovered on
the mainland in Steuben County (Spears, 1968). Further discoveries in
additional counties in western New York State during the 1970s and 1980s
suggest that potato cyst nematodes are spreading inexorably in the USA,
but that the rate of spread is probably slower than occurred in Europe 100
years earlier because of the quarantine measures which have been enforced
(Brodie, 1984).
If a nematode pest is very widespread strict legislative control cannot be
effectively applied but if it is relatively sparsely distributed legislation can
delay further spread and minimize losses. Perhaps more importantly it
means that seed may be produced on uncontaminated land so that spread
via seed is eliminated. Operation of reliable clean seed schemes is a
prerequisite for any control programme. Pure G. rostochiensis pathotype
R01 populations might be eliminated by repeated growing of HI ex-
andigena cultivars but cultivars with high enough levels of resistance to
eliminate other pathotypes are not available. Thus, where potato cyst
nematodes are endemic farmers will have to learn to live with these by
using rotations, nematicides and resistant cultivars in an integrated policy
designed to keep populations below the threshold for damage.
11.5 FUTURE DEVELOPMENTS IN NEMATODE RESEARCH
Because many plant-parasitic nematodes live in the soil, and are only one
of several agents that might impair root function, it is often quite difficult
to determine the amount of damage they cause. There are usually obvious
associations between the symptoms in the plant and attack by Globodera
spp., Meloidogyne spp. or Nacobbus spp., but the effects of ectoparasites
such as Longidorus spp. and endoparasites such as Pratylenchus spp. are
Future developments in nematode research 463
more insidious and difficult to diagnose. A good example of the difficulties
of diagnosis is the story of the so-called 'Docking Disorder' of sugar beet,
which was variously ascribed to fungal pathogens and unusual chemical
and physical conditions of the soil before Gibbs and Harrison (1963)
associated the damage with Longidorus spp. and Whitehead et at. (1966)
associated it with Trichodorus spp. More work is required to assess the
effects of these nematodes on potatoes.
Increased knowledge of the mechanisms of nematode damage to plants
may enable us to apply remedial treatments. Research into the optimum
rates of fertilizer for plants infested with small to moderate numbers of
potato cyst nematodes may provide a cheaper method of increasing yield
than application of nematicide (Trudgill, 1986) but could lead to much
higher post-harvest nematode population densities.
Increased knowledge of mechanisms of nematode damage to plants
might also provide plant breeders with more rapid and rational methods
of testing progeny for resistance. Giebel (1970) attempted to develop
a chemical test to determine the degree of plant resistance to G. rostochiensis
and found that resistant and susceptible cultivars differed in the ratio of
mono- to polyphenols found in their roots. Rice et at. (1985) described a
hypersensitive response in the root cells surrounding the invading juveniles
in resistant clones and Robinson et at. (1988) used fluorescence microscopy
to define differences in the chronology and location of this hypersensitive
response between resistant and susceptible hosts. They suggested that it
may be possible to use quantified fluorescence to determine the degree of
compatibility of the host-parasite relationship. Saunders (1989) examined
changes in concentrations of a range of metabolites associated with the
hypersensitive response and showed that some changes were associated
with the expression of resistance but was unable to identify the agent
directly responsible.
Current potato breeding programmes seek to produce cultivars with
reasonable tolerance to potato cyst nematodes, largely because of recent
experiences with several prospective cultivars which, although highly
resistant to G. pallida, were extremely intolerant of nematode attack
(Whitehead, 1986). Incorporating tolerance in new cultivars requires a
suitable assay for tolerance, and Huijsman et at. (1969) described different
anatomical reactions in the roots of susceptible, resistant and tolerant
cultivars but the techniques involved in assessing the reactions of root cells
to nematode feeding were too difficult to apply on a routine basis. Because
a primary effect of potato cyst nematodes is the slowing of root growth,
measurements of the effect of invading juveniles on root growth may
provide one measure of tolerance. However, many factors interact in
determining crop performance in the field and the only reliable estimates
of nematode tolerance come from field trials. Such trials are usually
expensive in terms of planting material and other resources but a system
described by Phillips et at. (1988) and Evans and Russell (1990) uses single
plants as plots. Good estimates of tolerance levels are obtained and the
assays can be made at an earlier stage of the breeding programme than was
previously possible and this should enable breeders to ensure that new
cultivars with G. pallida resistance are also tolerant. Even so, we need to
know more about the mechanisms involved in tolerance/intolerance and
their inheritance. Tolerance and/or resistance to secondary root pathogens
may also be worth seeking but as yet there is insufficient information about
their role in hastening senescence of nematode-infested plants, except
possibly that of Verticillium dahliae (Corbett and Hide, 1971; Storey and
Evans, 1987).
There is also a need to understand better the ways in which nematodes
interact with the plant environment to cause damage (Trudgill, 1986).
Small numbers of potato cyst nematodes can cause severe damage to
potatoes grown in soil poor in Mg (Trudgill et al., 1975c) and Evans (1971)
suggested that L. leptocephalus caused most damage to potatoes when
water was scarce.
There is still scope for improvement in control of nematodes and the
losses they cause in potatoes. Biological control of potato cyst nematodes
seems unlikely as they are devastating in the area from which they
originated as they are in areas to which they have been introduced.
However, Nishizawa (1986) used an isolate of Pasteuria penetrans from
Heterodera glycines in a pot test and reduced the rate of multiplication of
G. rostochiensis. Because biological control agents establish a dynamic
equilibrium with their hosts rather than eliminating them, it may become
necessary to produce potato cultivars with tolerance of, but no resistance
to, potato cyst nematodes.
The use in the UK of potato cultivars resistant to G. rostochiensis
pathotype R01 has diminished its occurrence but has increased the
occurrence of G. pallida (Hancock, 1988) making the breeding of cultivars
with resistance to this species apriority. In order to provide cultivars
resistant to all pathotypes, such as occur in South America, a wider
spectrum of resistance is required. This is probably to be found in wild
potatoes (Chavez et al., 1988) and Jackson et al. (1988) point out that a
wide genetic base should be used in order to decrease the likelihood of
selection for virulence in the nematode populations, perhaps using species
such as S. brevicaule and S. leptophyes in which they found G. pallida
resistance.
The production of new cultivars with a wide spectrum of resistance will
be made easier if nematode populations with clearly defined levels of
virulence can be selected, and some preliminary attempts are being made
with G. rostochiensis (Janssen et al. 1989). This work is paralleled by
attempts to define virulence levels in populations of the soybean cyst
nematode (Luedders, 1990). More understanding of the interactions
between potato cyst nematode populations and potato clones carrying
minor genes for resistance is required before the pathotype situation can be
Future developments in nematode research 465
properly resolved (Trudgill, 1985; Cook and Evans, 1987). At present, two
pathotypes of G. rostochiensis (separated by the major resistance gene H 1 )
and two pathotypes of G. pallida (separated by the major resistance gene
H 2 ) can be recognized. Classifying populations of the nematodes even
according to this simple scheme is time-consuming when traditional pot-
testing methods are used. Much more rapid systems of analysis will
undoubtedly be developed soon, based on the use of monoclonal anti-
bodies (Jones et al., 1988) and/or nucleic acid probes (Burrows, 1988).
Similarly, improvements in techniques for gene transfer between plants
may soon make it possible to isolate resistance genes and transfer them to
potato clones which provide a particularly suitable genetic background.
Also, an increased understanding of mechanisms involved in the elicitation
of resistance and of host-parasitic relationships should make it possible to
identify genes which will provide resistance by novel methods, perhaps
using completely unrelated species of plants as gene donors.
Although much attention is focused on mechanisms of resistance to cyst
and root knot nematodes, the susceptible response is probably even more
interesting. The syncytia induced by cyst nematodes are formed by cell wall
breakdown and cell enlargement accompanied by increases in DNA, RNA
and cytoplasm content. Giant cells induced by root knot nematodes
become much enlarged, multinucleate and the nuclei become polyploid.
These changes are thought to be induced by saliva coming from the dorsal
pharyngeal gland cell in the nematode. In addition, secretions from this
gland cell produce the spiral 'feeding tube' through which the nematode
probably withdraws cytosol without damaging the cytoplasmic structure of
the syncytium or giant cell. Advances in our understanding of these
processes may have far-reaching consequences.
Improvements in the availability of nematicides may also occur..
Spraying cabbage plants with oxamyl decreased invasion of the roots by
Heterodera schachtii juveniles (Potter and Marks, 1971) but Whitehead et
al. (1973) reported that there was no evidence to suggest that potato cyst
nematodes could be controlled in this way. Oxamyl is only translocated in
phloem to a small extent (Bromilow et al., 1987) whereas other compounds
(e.g. herbicides such as glyphosate and growth regulators such as maleic
hydrazide) are better transported in phloem due to their more appropriate
physico-chemical properties, the requirements for which have been
reviewed by Bromilow and Chamberlain (1989). Using these principles, it
may be possible to produce nematicides which can be sprayed onto foliage
and will be transported to roots to give effective control of nematodes,
although potatoes present a particular problem in that a large root system
is produced and invaded by cyst nematodes before the haulm is large
enough to spray. The use of phloem-translocated nematicides would
therefore have to be supplemented by the use of a soil-applied nematicide
for early protection of the root system. However, the use of foliar
applications only of nematicide may be effective in protecting potatoes
from ectoparasitic nematodes such as Longidorus spp. and preventing
infection of tubers with tobacco rattle virus transmitted by Trichodorus
spp. It is also possible that new nematicides of lower mammalian toxicity
may be developed, perhaps by targeting vital life processes in the
nematode (e.g. moulting) for inhibition.
For the present then, integration of control measures as suggested by
Jones (1970) and Mugniery (1978b) and applied to whole farms based on
estimates of soil nematode population densities by extension services,
seems likely to provide the best control of cyst and other nematode pests of
potatoes.
Some nematode problems of potato growing may be solved by
agronomic changes. For instance, it may be possible to eliminate spraing
in potatoes by growing barley, which is a non-host for most strains of TRV,
prior to growing potatoes, although good weed control would be essential.
However, the spread of potato cyst nematodes (especially G. pal/ida) and
an increased awareness of the effects of other nematodes will mean that
nematode problems in potatoes in temperate regions will continue to occur
for many years to come. And in the same way that cyst nematodes have
spread N. aberrans and M. chitwoodi may spread beyond their present
fairly limited known distribution ranges. More research is required to
understand what factors control the distribution of such potentially serious
pests, especially as N. aberrans has been reported in a glasshouse in
England (Graham, 1959). It seems likely too that potatoes will be grown
more in lowland tropical areas and root knot nematodes will undoubtedly
prove troublesome in these localities, making research into root knot
nematode resistant lines another priority. However, the rapid progress
made in recent years in understanding and controlling nematode problems
of potatoes suggests that control measures will usually keep abreast of
them.
REFERENCES
Abdel-Rahman, T.B., Elgindi, D.M. and Oteifa, B.A. (1974) Efficacy of certain
systemic pesticides in the control of root-knot nematodes of potato. Plant Dis.
Rep., 58, 517-20.
Adamson, A. (1972) The value of advice on potato cyst eelworm in south west
Lancashire. ADAS Q. Rev. 6, 1-11.
Bakker, J. (1987) Protein variation in cyst nematodes, Ph.D. Thesis, Agricultural
University, Wageningen, The Netherlands, 159 pp.
Bakker, J. and Gommers, F.J. (1982) Differentiation of the potato cyst nematodes
Globodera rostochiensis and G. pallida and of two G. rostochiensis pathotypes.
Proc. van de Koninklijke Academie van Wetenschappen, C85, 309-14.
Biehn, W.L., Hawkins, A. and Miller, P.M. (1971) Effect of 1,3-dichloropropene
and a combination of 1,3-dichloropropene + chloropicrin on infection of
potatoes by Verticillium albo-atrum. Plant Dis. Rep., 55, 968-71.
References 467
Boparai, J.S. and Hague, N.G.M. (1974) The effect of thioxamyl nematocide on
the development and reproduction of the potato cyst nematode Heterodera
rostochiensis Woll. Meded. Fac. Landbouwwet. Rijksuniv. Gent, 39, 719-25.
Bottrell, D.R. (1979) Integrated Pest Management, United States Government
Printing Office, Washington, USA.
Brodie, B.B. (1984) Nematode parasites of potato, in Plant and Insect Nematodes
(ed. W.R. Nickle), Marcel Dekker Inc., New York and Basel, pp. 167-212.
Brodie, B.B. and Plaisted, R.L. (1977) Breeding for resistance to root-knot
nematodes in potatoes. Nematropica, 7, 2.
Bromilow, R.H. and Chamerberlain, K. (1989) Designing for systemicity, in
Mechanisms and Regulations of Transport Processes (ed. R.K. Atkin), British
Plant Growth Regulator Group Monograph 18, pp. 113-28.
Bromilow, R.H., Rigitano, R.L.O., Briggs, G.G. and Chamberlain, K. (1987)
Phloem translocation of non-ionised chemicals in Ricinus communis. Pestic. Sci.
19,85-99.
Brown, D.J.F., Ploeg, A.T. and Robinson, D.J. (1989) A review of reported
associations between Trichodorus and Paratrichodorus (Nematoda:
Trichodoridae) and tobraviruses with a description of laboratory methods for
examining virus transmission by trichodorids. Revue Nematol., 12, 235-4l.
Brown, E.B. (1969) Assessment of the damage caused to potatoes by potato cyst
eelworm, Heterodera rostochiensis Woll. Ann. Appl. Bioi., 63, 493-502.
Brown, E.B. (1983) The relationship of potato yield with and without nematicide
to density of potato cyst nematodes, Globodera rostochiensis and G. pallida.
Ann. Appl. Bioi., 103,471--6.
Brown, E.B. and Sykes, G.B. (1973) Control of tobacco rattle virus (spraing) in
potatoes. Ann. Appl. Bioi., 75, 462-4.
Brown, E.B. and Sykes, G.B. (1975) Studies on the relation between density of
Longidorus elongatus and yield of barley and potatoes. PI. Path., 24, 221-3.
Brown, E.B. and Sykes, G.B. (1983) Assessment of the losses caused to potatoes
by the potato cyst nematodes, Globodera rostochiensis and G. pallida. Ann.
Appl. Bioi., 103, 271--6.
Burpee, L.L. and Bloom, J.R. (1978) The influence of Pratylenchus penetrans on
the incidence and severity of Verticillium wilt of potato. 1. Nematology, 10,
95-9.
Burrows, P.R. (1988) The use of nucleic acid probes to identify plant parasitic
nematodes. Proceedings, Brighton Crop Protection Conference, Pests and
Diseases, pp. 811-20.
Canto Saenz, M.A. (1984) Races of Meloidogyne incognita. OTAN Newsletter,
16(1), 34-5.
Canto Saenz, M.A. and de Scurrah, M.M. (1977) Races of the potato cyst
nematode in the Andean region and a new system of classification.
Nematologica, 23, 340--9.
Cetas, R.C. and Harrison, M.B. (1963) Evaluation of fumigants for control of early
maturity wilt of potatoes on Long Island. Phytopathology, 53, 347-8.
Chavez, R., Jackson, M.T., Schmiediche, P.E. and Franco, J. (1988) The
importance of wild potato species resistant to the potato cyst nematode,
Globodera pallida, pathotypes P4 A and PsA, in potato breeding. I. Resistance
studies. Euphytica, 27, 9-14.
Cook, R. and Evans, K. (1987) Resistance and tolerance, in Principles and Practice
of Nematode Control in Crops (eds R.H. Brown and B.R. Kerry) Academic
Press, Sydney, pp. 179-231.
Cooper, B.A. (1953) Eelworm problems in north fenland with special reference to
crop rotation. Hort. Educ. Ass. Ann. Report, pp. 106-15.
Corbett, D.C.M. and Hide, G.A. (1971) Interactions between Heterodera
rostochiensis Woll. and Verticillium dahliae Kleb. on potatoes and the effect of
CCC on both. Ann. Appl. Bioi., 68, 71-80.
DelJaert, L.M.W. and Vinke, J.H. (1987) Testing potatoes for resistance to
Globodera pallida pathotype Pa-3; resistance spectra of plant genotypes and
virulence spectra of Pa-3 isolates. Rev. Nematol., 10,445-53.
Dickerson, O.J., Darling, H.M. and Griffin, G.D. (1964) Pathogenicity
and population trends of Pratylenchus penetrans on potato and corn.
Phytopathology, 54, 317-22.
Dunn, E. and Hughes, W.A. (1967) Interactions of Oospora pustulans,
Rhizoctonia solani and Heterodera rostochiensis on potato. Eur. Potato 1., 10,
327-8.
Ellenby, C. (1954) Tuber forming species and varieties of the genus Solanum
tested for resistance to the potato root eelworm Heterodera rostochiensis
Wollenweber. Euphytica, 3, 195-202.
Empson, D.W. and James, P.J. (1966) An economic approach to the potato root
eelworm problem. NAAS Q. Rev. 73, 22-9.
Empson, D. W. and James, P.J. (1967) A further note on the economics of potato
root eelworm. NAAS Q. Rev., 76,160-5.
Evans, K. (1971) Potatoes in the Woburn ley-arable experiment. Rep. Rothamsted
Exp. Stn for 1970, Part 1, pp. 155-6.
Evans, K. (1979) Nematode problems in the Woburn ley-arable experiment, and
changes in Longidorus leptocephalus population density associated with time,
depth, cropping and soil type. Rep. Rothamsted Exp. Stn for 1978, Part ll, pp.
27--45.
Evans, K. (1982a) Effects of infestation with Globodera rostochiensis (Wollenweber)
Behrens Rolon the growth of four potato cultivars. Crop Protection, 1, 169-79.
Evans, K. (1982b) Water use, calcium uptake and tolerance of cyst-nematode
attack in potatoes. Potato Research, 25, 71-88.
Evans, K. (1987) The interactions of potato-cyst nematodes and Verticillium
dahliae on early and maincrop potato cuItivars. Ann. Appl. Bioi., 110,329-39.
Evans, K. and Franco, J. (1979) Tolerance to cyst-nematode attack in commercial
potato cultivars and some possible mechanisms for its operation. Nematologica,
25, 153-62.
Evans, K. and Russell, M.D. (1990) The use of single potato plants as plots in field
trials to assess tolerance of potato cyst nematode attack. Ann. Appl. Bioi., 117,
595-610.
Evans, K. and Stone, A.R. (1977) A review of the distribution and biology of the
potato cyst-nematodes Globodera rostochiensis and G. pallida. PANS, 23,
178-89.
Evans, K., Franco, J. and de Scurrah, M.M. (1975a) Distribution of species of
potato cyst-nematodes in South America. Nematologica, 21, 365-9.
Evans, K., Parkinson, K.J. and Trudgill, D.L. (1975b) Effects of potato cyst
nematodes on potato plants III. Effects on the water relations and growth of a
resistant and susceptible variety. Nematologica, 21, 273-80.
References 469
Evans, K., Trudgill, D.L. and Brown, N.J. (1977) Effects of potato cyst nematodes
on potato plants V. Root system development in lightly- and heavily-infested
susceptible and resistant varieties, and its importance in nutrient and water
uptake. Nematologica, 23, 153-64.
Feldmesser, J. and Goth, R.W. (1970) Association of a root knot with bacterial wilt
of potato. Phytopathology, 60, 1014.
Fleming, C.C. and Marks, R.J. (1983) The identification of the potato cyst
nematodes Globodera rostochiensis and G. pallida by isoelectric focusing of
proteins on polyacrylamide gels. Ann. Appl. Bioi., 103, 277-81.
Foot, M.A. (1978) Temperature responses of three potato-cyst nematode popu-
lations from New Zealand. Nematologica, 24, 412-17.
Franco, J. (1979) Effect of temperature on hatching and multiplication of potato
cyst nematodes. Nematologica, 25, 237-44.
Franco, J. and de Scurrah, Maria M. (1972) Nematodes, in Enfermedades de la
Papa en el Peru, Boletin Tecnico (77) Estacion Experimental Agricola La
Molina, Ch. VI, DGIA, Lima.
Gibbs, A.J. and Harrison, B.D. (1963) Docking disorder of sugar beet. Rep.
Rothamsted Exp. Stnfor 1962, pp. 113-14.
Giebel, J. (1970) Phenolic content in roots of some Solanaceae and its influence on
IAA-oxidase activity as an indicator of resistance to Heterodera rostochiensis.
Nematologica, 16, 22-32.
Gould, M.D. (1974) The root lesion nematode - Verticillium wilt disease complex
of potatoes and pathogenicity studies of the lesion nematode on selected
ornamental and cover crops. Diss. Abs. Int., 35B(2), 783.
Graham, C.W. (1959) A nematode genus new to Europe. Pl. Path., 7,114.
Grainger, J. (1951) The Golden Eelworm. Res. Bull. 10 West of Scotland Agric.
Call. Auchincruive, Ayr.
Grainger, J. and Clark, M.R.M. (1963) Interactions of Rhizoctonia and potato root
eelworm. Eur. Potato J., 6, 131-2.
Gratwick, M. (ed.) (1989) Potato Pests, Reference Book No. 187, HMSO,
London, 104 pp.
Greco, N., di Vito, M., Brandonisio, A., Giordano, I. and de Marins, G. (1982)
The effect of Globodera pallida and G. rostochiensis on potato yield. Nemato-
logica, 28, 379-86.
Guile, C.T. (1966) Cyst chromogenesis in potato cyst-eelworm pathotypes. Pl.
Path., 16, 125-8.
Hancock, M. (1988) The management of potato cyst nematodes in UK potato
crops. Aspects of Applied Biology, 17,29-36.
Hawkins, A. and Miller, P.M. (1971) Row treatments of potatoes with systemics
for meadow nematode (P. penetrans) control. Am. Potato J., 48, 21-5.
Haydock, P.P.J. (1989) Seed tuber physiological age and the growth of potato
cultivars with partial resistance to the potato cyst nematode Globodera pallida.
Tests of Agrochemicals and Cultivars, Ann. Appl. BioI., 114, (suppl), 158-9.
Hominick, W.M. (1979) Selection for hatching at low temperatures in Globodera
rostochiensis by continuous cultivation of early potatoes. Nematologica, 25,
322-32.
Hominick, W.M. (1982) Selection of a rapidly maturing population of Globodera
rostochiensis by continuous cultivation of early potatoes in Ayrshire, Scotland.
Ann. Appl. BioI., 100, 345-51.
Howard, H.W., Cole, C.S. and Fuller, J.M. (1970) Further sources of resistance to
Heterodera rostochiensis Woll. in the Andigena potatoes. Euphytica, 19,210-16.
Huijsman, C.A., Klinkenberg, C.H. and Duden, H. den (1969) Tolerance to
Heterodera rostochiensis Woll. among potato varieties and its relation to certain
characteristics of root anatomy. Eur. Potato 1., 12, 134-47.
Jackson, M.T., Hawkes, J.C., Male-Kayiwa, B.S. and Wanyera, N.W.M. (1988)
The importance of the Bolivian wild potato species in breeding for Globodera
pallida resistance. Plant Breeding, 101, 261-68.
Jacobsen, B.J., MacDonald, D.H. and Bissonette, H.L. (1979) Interaction
between Meloidogyne hapla and Verticillium wilt disease of potato.
Phytopathology, 69, 288-92.
Janssen, R., Bakker, J. and Gommers, F.J. (1989) Selection of virulent and
avirulent lines of Globodera rostochiensis for the Hl resistance gene in Solanum
tuberosum ssp. andigena CPC 1673. Rev. Nematol., 13, 265-8.
Jatala, P. and Golden, A.M. (1977) Taxonomic status of Nacobbus species
attacking potatoes in South America. Nematropica, 7, 9-10.
Jatala, P. and Rowe, P.R. (1976) Reaction of 62 tuber-bearing Solanum species to
the root-knot nematode, Meloidogyne incognita acrita. J. Nematology, 8, 290.
Jatala, P., Gutarra, L., French, E.R. and Arango, J. (1976) Interaction
of Heterodera pallida and Pseudomonas solanacearum on potatoes. 1.
Nematology, 8, 289-90.
Jatala, P. Kaltenbach, R. and Bocangel, M. (1979) Biological control of
Meloidogyne incognita acrita and Globodera pallida on potatoes. 1. Nematology,
11, 303.
Jensen, H. J., Armstrong, J. and Jatala, P. (1979) Annotated Bibliography of
Nematode Pests of Potato, International Potato Center, Lima, Peru, 315 pp.
Jones, F.G.W. (1969) Some reflections on quarantine, distribution and control, in
Nematodes of Tropical Crops (ed. J.A. Peachey), Commonwealth Agricultural
Bureaux, Farnham Royal, pp. 67-80.
Jones, F.G.W. (1970) The control of the potato cyst nematode, Heterodera
rostochiensis. 1. R. Soc. Arts, 118, 179-99.
Jones, F.G.W. (1977) Pests, resistance and fertilizers. Proceedings 12th
Colloquium of the International Potash Institute, Izmir, pp. 233-58.
Jones, F.G.W. and Perry, J.N. (1978) Modelling populations of cyst-nematodes
(Nematoda: Heteroderidae). 1. App/. Eco/., 15,349-71.
Jones, M.G.K. and Northcote, D.H. (1972) Nematode-induced syncytium: a
multinucleate transfer cell. 1. Cell Sci., 10, 789-809.
Jones, P., Ambler, D.J. and Robinson, M.P. (1988) The application of monoclonal
antibodies to the diagnosis of plant pathogens and pests. Proceedings Brighton
Crop Protection Conference, Pests and Diseases, pp. 767-76.
Joubert, J. and Dalmasso, I. T.P. (1972) PossibiJites d'interaction entre Rhizoctonia
solani et Meloidogyne incognita sur pommes de terre. Ann. de Phytopathol., 4,
409.
Kerry, B.R. (1974) A fungus associated with young females of the cereal cyst-
nematode, Heterodera avenae. Nemat%gica, 20, 259-60.
Kerry, B.R. (1986) Biological control, in Principles and Practice of Nematode
Control in Crops (eds R.H. Brown and B.R. Kerry), Academic Press, Sydney,
pp.233-63.
Kirjanova, E.S. and Krall, E.L. (1971) Plant Parasitic Nematodes and Their
References 471
Control, Vol. II, Nauka Publishers, Leningrad Section, Leningrad (English
translation, Amerind Publishing co. Pvt. Ltd, New Delhi).
Kort, J., Ross, H., Rumpenhorst, H.J. and Stone, A.R. (1977) An international
scheme for identifying and classifying pathotypes of potato cyst nematodes
Globodera rostochiensis and G. pallida. Nematologica, 23, 333-9.
Krikun, J. and Orion, D. (1977) Studies on the interaction of Verticillium dahliae
and Pratylenchus thornei on potato. Phytoparasitica, 5, 67.
Lawes, D.A. (1988) Resistant varieties, in Control of Plant Diseases: Costs and
benefits (eds B.C. Clifford and E. Lester), Blackwell, Oxford, 263pp.
Luedders, V.D. (1990) A recessive soybean cyst nematode allele for incom-
patibility with soybean PI 88287. Ann. Appl. Bioi., 116,313-19.
Mai, W.F., Brodie, B.B., Harrison, M.B. and Jatala, P. (1981) Nematodes,
in Compendium of Potato Diseases (ed. W.J. Hooker), American
Phytopathological Society, St Paul, pp. 93-101.
Martin, G.c. (1968) Control of Meloidogyne javanica in potato tubers and M.
hapla in roots of young roses by means of heated water. Nematologica, 14,
441--6.
Martin, M.J., Riedel, R.M. and Rowe, R.C. (1982) Verticillium dahliae and
Pratylenchus penetrans: Interactions in the early dying complex of potato in
Ohio. Phytopathology, 723, 640-4.
Morgan, D.O. (1926) Some remarks on the etiology of potato disease in
Lincolnshire. J. Helminthology, 4, 49-52.
Morsink, F. and Rich, A.E. (1968) Interactions between Verticillium albo-atrum
and Pratylenchus penetrans in the Verticillium wilt of potatoes. Phytopathology,
58,401.
Mugniery, D. (1978a) Vitesse de developpement, en fonction de la temperature, de
Globodera rostochiensis et G. pallida (Nematoda: Hetero:deridae). Rev.
Nematol., 1,3-12.
Mugniery, D. (1978b) Lutte culturale contre les nematodes a kystes de La pomme
de terre Globodera rostochiensis (WoH.) et G. pal/ida (St.) et perspectives de
lutte integree. Ann. Zool. Ecol. Anim., 10, 187-203.
Mugniery, D. and Fayet, G. (1981) Determination du sexe chez Globodera pal/ida
Stone. Rev. Nematol., 4, 41-5.
Mugniery, D., Phillips, M.S., Rumpenhorst, H.J., Stone, A.R., Truer, A. and
Trudgill, D.L. (1989) Assessment of partial resistance of potato to, and pathotype
and virulence differences in, potato cyst nematodes. EPPO Bull., 19, 7-25.
Newton, H.C.F. and Duthoit, C.M.G. (1954) Stem and bulb eelworm on potatoes.
PI. Path., 3, 139-40.
Nirula, K.K. and Bassi, K.K. (1965) Thermotherapy for root-knot nematode
(Meloidogyne incognita) in potato tubers. Indian Potato J., 7, 9-11.
Nishizawa, T. (1986) On a strain of Pasteuria penetrans parasitic to cyst nematodes.
Rev. Nematol., 9, 303-4.
O'Bannon, J.H. and Reynolds, H.W. (1965) Water consumption and growth of
root-knot nematode-infected and uninfected cotton plants. Soil Sci.. , 99, 251-5.
O'Bannon, J.H. and Santo, G.S. (1984) Effect of soil temperature on reproduction
of Meloidogyne chitwoodi and M. hapla alone and in combination on potato and
M. chitwoodi on rotation plants. J. Nematology, 16, 309-12.
Odihirin, R.A. (1971) Effects of root-knot and lesion nematodes on transpiration
and water utilization by tobacco plants. J. Nematology, 3, 321-2.
Olefir, V.V. (1972) [Breeding of potato for resistance to potato tuber nematode].
Tezisy soveshchaniya. Moskova dekabr 1972. Moscow, USSR VASHNIL,
101-3.
Olthof, T.H.A. and Potter, J.W. (1972) Relationship between population densities
of Meloidogyne hapla and crop losses in summer-maturing vegetables in
Ontario. Phytopathology, 62, 981-6.
Oostenbrink, M. (1958) An inoculation trial with Pratylenchus penetrans in
potatoes. Nematologica, 3, 30-3.
Oostenbrink, M. (1961) Nematodes in relation to plant growth. III Pratylenchus
penetrans (Cobb) in tree crops, potatoes and red clover, Neth. J. Agric. Sci., 9,
188-208.
Oostenbrink, M., s'Jacob, J.J. and Kuiper, K. (1957) Over de Waardplanten van
Pratylenchus penetrans. Tijdschr. PIZiekt., 63, 345-60.
Parrott, Diana M. (1981) Evidence for gene-for-gene relationships between
resistance gene Hl from Solanum tuberosum ssp. andigena and a gene in
Globodera rostochiensis, and between H2 from S. multidissectum and a gene in
G. pallida. Nematologica, 27, 372-84.
Phillips, M.S. (1984) The effects of different levels of inoculum of Globodera
pallida on the assessment of nematode reproduction on partially resistant clones
derived from Solanum vernei. Nematologica, 30, 57-65.
Phillips, M.S. and Trudgill, D.L. (1983) Variations in the ability of Globodera
pallida to produce females on potato clones bred from Solanum vernei or S.
tuberosum ssp. andigena CPC 2802. Nematologica, 29, 217-26.
Phillips, M.S., Trudgill, D.L. and Evans, K. (1988) The use of single, spaced
potato plants to assess their tolerance of damage by potato cyst nematodes.
Potato Research, 31, 469...,.75.
Phillips, M.S., Rumpenhorst, H.J., Trudgill, D.L., Evans, K., Gurr, G., Heinicke,
D., Mackenzie, M. and Turner, S.J. (1989) Environmental interactions in the
assessment of partial resistance to potato cyst nematodes. I. Interactions with
centres. Nematologica, 35, 187-96.
Potter, J.W. and Marks, C.F. (1971) Effect of DuPont (R) 141O-X on rate of
development of Heterodera schachtii on cabbage. J. Nematology, 3, 325.
Price, N.S. and Sanderson, J. (1984) The translocation of calcium from oat roots
infected by the cereal cyst nematode Heterodera avenae Woll. Rev. Nematol., 7,
239-43.
Rebois, R.V., Eldridge, B.J. and Webb, R.E. (1978) Rotylenchulus reniformis
parasitism of potatoes and its effect on yields. Plant Dis. Rep., 62, 520-3.
Rice, S.L., Leadbeater, B.S.C. and Stone, A.R. (1985) Changes in cell structure in
roots of resistant potatoes parasitized by potato cyst-nematodes. I. Potatoes
with resistance gene Hl derived from Solanum tuberosum ssp. andigena.
Physiol. Pl. Pathol., 27, 219-34.
Riedel, R.M. and Mai, W.F. (1971) Pectinases in aqueous extracts of Ditylenchus
dipsaci. J. Nematology, 3, 28-38.
Roberts, P.R. (1982) Plant resistance in nematode pest management. J.
Nematology, 14, 24-33.
Robinson, M.P., Atkinson, H.J. and Perry, R.N. (1988) The association and
partial characterisation of a fluorescent hypersensitive response of potato roots
to the potato cyst nematodes Globodera rostochiensis and G. pallida. Rev.
Nematol., 11, 99-107.
References 473
Rodriguez-Kabana, R. and Ingram, E.G. (1977) Treatment of potato seed-pieces
with oxamyl for control of plant-parasitic nematodes. Plant Dis. Rep., 61, 29-3l.
Rumpenhorst, H.J. (1984) Intracellular feeding tubes associated with sedentary
plant parasitic nematodes. Nematologica, 30, 77-85.
Santo, G.S. and D'Bannon, J.H. (1981) Effect of soil temperature on the
pathogenicity and reproduction of Meloidogyne chitwoodi and M. hapla on
Russet Burbank potato. J. Nematology, 13,483--6.
Santo, G .S. and Qualls, M. (1984) Control of Meloidogyne spp. on Russet Burbank
potato by applying metham sodium through centre pivot irrigation systems. J.
Nematology, 16, 159--6l.
Saunders, K.S. (1989) The biochemical basis for major gene resistance to potato
cyst nematodes in Solanum tuberosum, Ph.D. Thesis, University of Birmingham.
Schluter, K. (1976) The potato cyst eelworm Heterodera rostochiensis Woll. in
Morocco: its distribution and economic importance. Z. PJlanzenkrank.
P[lanzensch., 83, 401--6.
Seinhorst, J.W. (1957) Phytonematology in Western Europe. Technical
Committee, Southern Regional nematology Project (S-19), Auburn, Alabama.
Seinhorst, J.W. (1965) The relation between nematode density and damage to
plants. Nematologica, 11, 137-54.
Seinhorst, J.W. (1967) The relationships between population increase and popu-
lation density in plant parasitic nematodes. V. Influence of damage to the host
on multiplication. Nematologica, 13, 481-92.
Seinhorst, J.W. (1979) Nematodes and growth of plants: formalization of the
nematode-plant system, in Root-Knot Nematodes (Meloidogyne species) (eds F.
Lamberti and C.E. Taylor), Academic Press, London and New York, pp.
231-55.
Seinhorst, J.W. (1982a) The relationship in field experiments between populatioo
density of Globodera rostochiensis before planting potatoes and yield of potato
tubers. Nematologica, 28, 277-84.
Seinhorst, J.W. (1982b) The distribution of cysts of Globodera rostochiensis in
small plots and the resulting sampling errors. Nematologica, 28, 285-97.
Seinhorst, J.W. and den Duden, H. (1971) The relation between density of
Heterodera rostochiensis and growth and yield of two potato varieties.
Nematologica, 17,347--69.
Setty, K.G.H. and Wheeler, A.W. (1968) Growth substances in roots of
tomato (Lycopersicon esculentum Mill.) infected with root-knot nematodes
(Meloidogyne spp.). Ann. Appl. Bioi., 61, 495-50l.
Sher, S.A. (1970) Revision of the genus Nacobbus Thorne and Allen, 1944
(Nematoda: Tylenchoidea). J. Nematology, 2, 228--35.
Simon, J.E. (1955) Contribuciones al estudio del nematodo dorado de la papa en el
Peru. Informe mensual Estacion experimental agricola La Molina, 29, 8--14.
Spears, J.F. (1968) The Golden Nematode Handbook, US Dept Agric., Agric.
Handbook 353.
Sprau, F. (1960) Uber ein vermutlich Pflanzenschadigendes auftreten eines
freilebenden Nematoden, Longidorus maximus (Butschli) an einer Reihe von
Kulturpflanzen. Nematologica Suppl., 2, 49-55.
Stone, A.R. and Parrott, D.M. (1985) Pathotypes. Rep. Rothamsted Exp. Stn for
1984, p. 116.
Stone, A.R. and Turner, S.J. (1982) The nature of resistance to potato
cyst-nematodes, in Research for the Potato in the Year 2000, International
Potato Center, Lima, Peru, pp. 178-9.
Storey, G.W. (1982) The relationship between potato root growth and reproduc-
tion of Globodera rostochiensis (Woll.). Nematologica, 28, 210-18.
Storey, G.W. and Evans, K. (1987) Interactions between Globodera pallida
juveniles, Verticillium dahliae and three potato cultivars, with descriptions of
associated histopathologies. Pl. Path., 36, 192-200.
Thorne, G. (1961) Principles of Nematology, McGraw-Hill, New York, 553 pp.
Trudgill, D.L. (1967) The effect of environment on sex determination in
Heterodera rostochiensis. Nematologica, 13, 263-72.
Trudgill, D.L. (1970) Survival of different stages of Heteroda rostochiensis at high
temperatures. Nematologica, 16, 94-8.
Trudgill, D.L. (1980) Effects of Globodera rostochiensis and fertilisers on the
mineral nutrient content and yield of potato plants. Nematologica, 26, 243-54.
Trudgill, D.L. (1985) Potato cyst nematodes: a critical review of the current
pathotyping scheme. EPPO Bull., 15, 273-9.
Trudgill, D.L. (1986) Yield losses caused by potato cyst nematodes: a review of the
current position in Britain and prospects for improvements. Ann. Appl. BioI.,
108, 181-98.
Trudgill, D.L. (1987) Effects of rates of nematicide and of fertiliser on the growth
and yield of cultivars of potato which differ in their tolerance of damage by
potato cyst nematodes (Globodera rostochiensis and G. pallida). Plant and Soil,
104, 235-43.
Trudgill, D.L. and Carpenter, J.M. (1971) Disk electrophoresis of proteins of
Heterodera species and pathotypes of Heterodera rostochiensis. Ann. Appl.
BioI., 69, 35-41.
Trudgill, D.L. and Cotes, L.M. (1983a) Differences in the tolerance of potato
cultivars to potato cyst nematodes (Globodera rostochiensis and G. pallida) in
field trials with and without nematicides. Ann. Appl. Biol., 102, 373-84.
Trudgill, D.L. and Cotes, L.M (1983b) Tolerance of potato to potato cyst
nematodes (Globodera rostochiensis and G. pallida) in relation to the growth
and efficiency of the root system. Ann. Appl. BioI., 102, 385-97.
Trudgill, D.L., Evans, K. and Parrott, D.M. (1975a) Effects of potato cyst-
nematodes on potato plants. 1. Effects in a trial with irrigation and fumigation
on the growth and nitrogen and potassium contents of a resistant and a
susceptible variety. Nematologica, 21, 169-82.
Trudgill, D.L., Evans, K. and Parrott, D.M. (1975b) Effects of potato cyst
nematodes on potato plants. II. Effects on haulm size, concentration of
nutrients in haulm tissue and tuber yield of a nematode resistant and a
nematode susceptible potato variety. Nematologica, 21, 183-91.
Trudgill, D.L., Parrott, D.M., Evans, K. and Widdowson, F.V. (1975c) Effects of
potato cyst nematodes on potato plants. IV. Effects of fertilisers and Heterodera
rostochiensis on the yield of two susceptible varieties. Nematologica, 21,
281-6.
Trudgill, D.L., Phillips, M.S. and Alphey, T.J.W. (1987) Integrated control of
potato cyst nematode. Outlook on Agriculture, 16, 167-72.
Uhrig, H. and Wenzel, G. (1981) Solanum gourlayi Hawkes as a source of
resistance against the white potato cyst nematode Globodera pallida Stone. Z.
Pjlanzenzuchtung, 86, 148-57.
References 475
van den Brande, J. and d'Herde, J. (1964) Phenological control of the potato root
eelworm (Heterodera rostochiensis Woll.). Nematologica, 10, 25-8.
van Hoof, H.A. (1964) Het tijdstip van infectie en veranderingen in de
concentratie van ratelvirus (Kringerigheid) in der aardappelknol. Meded.
LandbHoogesch. opzoekingsstations van de staat te Gent, Deel XXIX, 3,
944--55.
Wallace H.R. (1974) The influence of root knot nematode, Meloidogyne javanica
on photosynthesis and on nutrient demand by roots of tomato plants.
Ward, S.A., Rabbinge, R. and den Ouden, H.(1985) Construction and preliminary
evaluation of a simulation model of the population dynamics of the potato cyst-
nematode Globodera pallida. Neth. J. PI. Path., 91,27-44.
Webley, D.P. (1970) A morphometric study of the three pathotypes of the potato
cyst eel worm (Heterodera rostochiensis) recognised in Great Britain.
Webley, D.P. and Jones, F.G.W. (1981) Observations on Globodera pallida and
G. rostochiensis on early potatoes. Pl. Path., 30, 217-24.
Whitehead, A.G. (1973) Control of cyst-nematodes (Heterodera spp.) by organo-
phosphates, oximecarbamates and soil fumigants. Ann. Appl. Bioi., 75, 439-53.
Whitehead, A.G. (1975) Chemical control of potato cyst-nematode. ARC Res.
Rev., 1, 17-25.
Whitehead, A.G. (1986) Problems in the integrated control of potato cyst-
nematodes, Globodera rostochiensis and G. paUida, and their solution. Aspects
of Applied Biology, 13, 363-72.
Whitehead, A.G., Greet, D.N. and Fraser, Janet E. (1966) Plant parasitic
nematodes and their control in relation to stunting of sugar beet in England.
Proc. Third Br. Insecticide and Fungicide Conf., Brighton, 1965, pp. 100--5.
Whitehead, A.G., Tite, D.J. and Fraser, Janet E. (1973) Control of potato cyst
nematode, Heterodera rostochiensis, in sandy loam, by Du Pont 1410 (S-methyl
1-( dimethylcarbamoyl)-N-[ (methylcarbamoyl)oxy1thioformimidate) applied to
the soil at planting time. Ann. Appl. Bioi., 73, 325-8.
Whitehead, A.G., Tite, D.J., Fraser, J.E., French, E.M. and Short, L. (1980)
Effects of aldicarb and ox amyl in peaty loam soils on potato cyst nematode,
Globodera rostochiensis, and on resistant and susceptible potatoes. J. Agric.
Sci. Camb., 95, 213~17.
Winfield, A.L. (1965) Potato root eelworm in Holland, Lincolnshire. NAAS Q.
Rev., 67,110--17.
Winfield, A.L. and Cooke, D.A. (1975) The ecology of Trichodorus, in Nematode
Vectors of Plant Viruses (eds F. Lamberti, C.E. Taylor and J.W. Seinhorst),
Plenum, London, pp. 309-41.
Winslow, R.D. and Willis, R.J. (1972) Nematode diseases of potatoes, in
Economic Nematology (ed. J.M. Webster), Academic Press, London, pp.
17-48.
CHAPTER 11
Pest aspects of potato production

Part 2. Insect pests
K. V. Raman and E.B. Radcliffe
11.6 INTRODUCTION
The potato ecosystem is inhabited by many different species of insects and

mites. Boiteau (1983) collected 565 taxa (species or species groups) in a
3-year study of potato fields in New Brunswick, Canada. Each continent,
growing region, and production system has its own pest complex (Fig.
11.4). Radcliffe et al. (1991) listed 170 arthropod species as potato pests in
North America alone. But the greatest diversity of arthropods associated
with potatoes occurs in South and Central America, where the crop and its
many related wild species originated.
In each region and production system, however, there are only certain
key pests that must be routinely or specifically targeted for control. In seed
tuber production, the pests of greatest concern are usually the aphid
vectors of potato virus diseases, especially green peach aphid, Myzus
persicae. In ware production, the key pests may be insects which attack
tubers, such as potato tuber moth (= potato tuberworm), Phthorimaea
aperculella, Andean potato weevils, Premnotrypes spp., or wireworms. In
other situations, the key pests may be foliage feeders such as Colorado
potato beetle, Leptinotarsa decemlineata, leafhoppers, or leaf-feeding
coccinellids.
In high input potato production systems, acceptable control of these key
pests usually can be obtained only with use of insecticides. But, once
insecticides have been introduced into the system, relative pest status of
the various species may be greatly altered. Certain potentially injurious
species will be incidentally suppressed to noneconomic status, while
others, perhaps because of elimination of their natural enemies or develop-
ment of insecticide resistance, may assume greater importance.
CENTRAL ANI) ......
NORTH AMERICA SOUTH AMERICA EUROPE AFRICA ASIA AUSTRALIA ::I
......
<:> "'1
z Myzus persicae 0
Myzus persicae Myzus persicae Myzus persicae Myzus persicae Myzus persicae
!:l Macrosiphum euphorbiae 0..
<:>z Macrosiphum euphorbioe Macrosiphum euphorbiae A ulacorthum .mlan; Macrosiphum euphorbiae Macrosiphum euphorbiae
ZO Aphis nasturti; ~
a:tu Aphis gossypii Aphis nasturtii Uriomyza triIm;; Aphis Jabal! Phthorimaea apercu/ella ()
Aulacorthum solani Liriomyza spp. Aulacorthum mlani Phthorimaea Aphis gossypi; Henosepilachna .\parsa ......
~irl
en a: " Empoasca Jahal! I:..pitrix spp. i.eptinotarsa decem/ineata apercu/ella Empoasca devostans vigintb.expunclata O·
ween ParatriollJ cockerell; Diabrotica spp. Austroasca viridigrisea
>Z . Phthorimaea aperculella Skis/oeerea gregoria Eplilachna spp. ::I
~C~ Leptinotarsa decemlineata Epicaula spp. Epilachna spp. Heliolhis armigera Listroderel' obliquus
-'<:>- Epitrix spp. Premnolrypes spp. Polyphagolarsonemus Spodoplera exigua
Z~
S2~ Phthorimaea operculella latus Plusia orichalcea
gci Scrobipalpula absoluta Thrips palmi Sihenaridea pusilla
en~ Po/yphagotarsonemus latus Psylloides plana
Epicaula hirticornis
I::::>
U
Phthorimaea operculella
Polyphagolarsonemu,~ lalus
Leptinolarsa decem/ineala
Thrips palmi
Phlh'orimaea Phthorimaea opercu/ella I Phlhorimaea opereu/el/a

Symmetrischema
I"'-"- .~~. operculella Agrotis ipsilon Agrotis spp.
plaeseosema Agrotis segetum Heteronychus aralor
Agrotil' segetum
Scrobipalpopsis solanivora Agroti,{ ipsi/on Ustroderes obliquu.{
Liriomyza palagonica
{~ I Feltia expena
Stenotycha sp.
Umonius sp. Premnotrypes spp. Limonius sp. Phthorimaea Phthorimaea operculella Phrhorimaea operculella
Me/anolu, ,po
I Phthorimaea operculella Conoderus sp. operculella Anomala dimidiata Cheiroplaly.~lalipes
Clenicera sp. Symmetrischema Ctenicera sp. Gryl/ola/po africana Phyllognalhus dionysiu.{ Agrotis spp.
Agriotes sp. p/aeseosema Agrioles spp. .tlelolontha spp. Graphognathul' leucoloma
l-{I Agrotis spp.
Hypolithus sp. Scrobipalpopsis mlanivora Phthorimaea operculella Odontoterme.\' obel'us Heteronychus arator
Conodorus sp. Diabrotica spp. Eremolerme,\' spp.
/:.pitrix spp. Epilrix spp. Agrotis spp.
Phthorimaea operculella Phyllophaga sp. Gryllotalpa africana
Copitarsia turbata Alcidodel' wel'lermanni
Feltia experla Myllocerus .\'ubfal'ciatus
Rothinus maimon Pyralis farinalil'
Phenacoccus grenadensis Nipaecoau.\ va,~tator
Figure 11.4 Major insect and mite pests of the potato crop, worldwide. Information summarized from Shands and Landis (1964),
Radcliffe et al. (1991) for North America; from Delgado and Aguilar (1980), King and Saunders (1984) for South and Central +:--
America; from Gibson (1978) for Europe; from Raman (1987), Kibatta (1982) for Africa; from Misra and Agrawal (1988), Eveleens ::j
and Woodford (1982) for Asia; and from Hamilton (1983) for Australia. Original figure, International Potato Center (CIP).
478 Insect pests
Yield losses resulting from arthropod pests are not documented as well
as might be expected. For example, research on pesticides has often
measured only reductions in pest numbers and not effects on tuber yield or
quality. Potatoes are a relatively high value crop and yields are difficult to
measure with precision. Economic thresholds are usually computed by
regression techniques in experiments with a single stress variable. In
temperate countries, economic thresholds have been published for most
key species, but relatively little research has been done to quantify
interacting effects of multiple biotic and abiotic stress factors. Even
differences in cultivar response to pests are not well documented. In the
tropics, especially in low input production systems, crop/pest interactions
are even less well known. Additional research is needed not only on the
present key pests, but also on potential pests, as the status of these could
change as pest management systems evolve.
When pest pressure is severe, many of the major individual pest species
can routinely cause tuber yield reductions of 30-70%. Losses of this
magnitude have been shown in untreated plots for aphid transmitted
viruses [potato leafroll virus (PLRV) and potato virus Y (PVY)], potato
tuber moth, leafminer flies, Liriomyza spp., Andean potato weevils,
potato leafhopper, Empoasca fabae, and Colorado potato beetle.
Growers tend to have a lower tolerance threshold for insect inflicted
defoliation than does the potato, whereas they often overlook, at least
initially, the less conspicuous damage caused by sucking insects or tuber
pests. Pests that attack the plant over a prolonged period of time, such as
potato tuber moth, leafminer flies, Andean potato weevils, thrips, mites
and aphid transmitted viruses cause the greatest yield losses. Lesser yield
reductions result from pests that defoliate or damage plants only briefly, as
do cutworms and leaf feeding beetles. Potato has considerable ability to com-
pensate for early season loss of foliage (Midmore, 1986), but there is little
compensation for adverse effects on plant health, as caused by virus infec-
tion, disruption of nutrient transport or reduced photosynthetic efficiency.
Despite considerable recent efforts to implement integrated pest
management (IPM) programmes for potato, use of chemical pesticides
remains the pre-eminent tactic of pest management in most technologically
advanced potato production systems. In the developing world, most potato
growers are small landholders who generally do not use great quantities of
insecticides on their crops. However, as farm sizes increase and farmers are
educated to the benefits, insecticide use increases greatly (Fano and Ewell,
1986). This is particularly true when production is intensified or growing
the crop is extended into high-risk agroecological zones. After cotton,
potato is the crop most treated with pesticides (Horton, 1987).
The benefits of insecticide use are undeniable. Between 1945 and 1950,
average US potato yields increased from 10.5 (10.67 tonnes) to 17.1 tons
(17.37 tonnes) ha- 1 (Potato Association of America, 1980). This dramatic
improvement has been attributed in large measure to the introduction of
Introduction 479
DDT and more effective fungicides (McNew 1963). Pimentel et al. (1977)
estimated that losses in yield and quality of potatoes in the US due to
insects were 22% in 1910-35, 16% in 1942-51, and 10-12% in 1975.
Without insecticides it is estimated that potato yield losses would increase
30% (Pimentel et al., 1979).
Much of the insecticide used on potato is applied not to remedy pest
outbreaks, but as an insurance against the uncertain risk of unacceptable
loss. Therefore, insecticides are commonly applied to potatoes on a fixed
schedule, without regard to actual pest pressure. While this has often
proved very effective, at least initially, the adverse consequences of
excessive reliance on pesticides have been repeatedly demonstrated.
Adoption of pest management strategies that rely largely on pesticides is
always risky. This is especially so in developing countries where there may
be inconsistent supplies of pesticides, costs of these materials may be
determined externally and unpredictably, and lack of proper application
equipment or applicator training may present safety hazards (Brader, 1988).
However, for the foreseeable future, insecticides will continue to be a
necessary ingredient in potato production. It is imperative that insecticides
be used as proficiently and as responsibly as possible. One concern is that
the rate at which new pesticides are reaching the market has slowed
dramatically (Larson et al., 1985). Moreover, nearly all the candidate
insecticides presently in development are pyrethroids, as are most cur-
rently important potato insecticides. It is critically important that insecti-
cide resistance be effectively managed to preserve the efficacy of our
present insecticides. This will require a co.mmitment to insect co.ntrol
strategies that fully inco.rpo.rate IPM precepts.
Problems asso.ciated with an excessive reliance o.n insecticides include
develo.pment of insecticide resistance, pest resurgence, secondary pests,
adverse environmental effects, and human expo.sure to. pesticides. In
certain instances, po.tato. growers have fo.und themselves trapped on a
pesticide treadmill that ultimately caused productio.n to. be unprofitable; a
classic example is from Lo.ng Island, New Yo.rk (Fo.rgash, 1985).
On Lo.ng Island, Co.lorado. po.tato beetle has been a target fo.r insectici-
dal control fo.r mo.re than 100 years. Resistance to. DDT was first o.bserved
in 1952, after just seven seasons (14 beetle generatio.ns) use. Each new
insecticide introduced o.n the Island failed, with the useful life o.f each
successive introductio.n o.f a chemical class being progressively sho.rter. For
a time, high rates o.f aldicarb (Temik) were applied in-furrow at planting
fo.r first generation beetle co.ntrol, but the insecticide was withdrawn in
1980 because o.f its appearance as a ground water contaminant. The
pyrethroids, fenvalerate (Pydrin) and permethrin (Ambush, Po.unce),
were granted emergency registrations in 1979, but by 1981 no. lo.nger
provided adequate co.ntro.l. Growers got through the 1982 and 1983
seasons using fenvalerate in co.mbinatio.n with the synergist piperonyl
buto.xide. By 1984, growers fo.und that no.ne o.f the synthetic insecticides
480 Insect pests
available to them performed adequately and they had to resort to rotenone

with piperonyl butoxide or cryolite (Kryocide). Contol costs eventually
reached levels of $(US)300-700 ha- 1 . In the period 1978-88, potato
hectarage declined by 65% on the Island.
11.7 PEST BIOLOGY, DAMAGE AND DISTRIBUTION
11.7.1 Aphids
Worldwide, aphids (Homoptera: Aphididae) are the insects of greatest
economic importance on potato. When abundant, aphids can cause direct
plant injury (Adams and Kelley, 1950; Carden, 1965; Kolbe, 1970; French,
1983), but the primary importance of aphids is as vectors of potato viruses
(see Chapter 10). At least ten potato viruses are aphid transmitted
(Bagnall, 1977; International Potato Center, 1988): PLRV, PVY, potato
virus A (PVA) and the closely allied SB29 virus (SB29V), potato virus S
(PVS) and the closely related potato virus M (PVM) , potato aucuba
mosaic virus (PAMV = potato virus F and including potato virus G),
alfalfa mosaic virus (AMV = potato calico virus), and the closely allied
SB22 virus (SB22V), and cucumber mosaic virus (CMV).
All potato-colonizing aphids have four nymphal instars. Their partheno-
genetic life style coupled with a rapid turnover of generations (typically
7-10 days) allows for spectacular population increases. Under ideal field
conditions, green peach aphid populations can double in as little as
1.7 days. A single female can give birth to 100 or more progeny. In
temperate regions, most aphid species show a seasonal alternation of hosts
(heteroecy) and overwinter as fertilized eggs (holocycly) (Fig. 11.5). The
overwintering (primary) host is often a woody plant, whereas the summer
~ ..ad"Hs
parthenogenetic
9W-
reproduction
adult
Figure 11.5 Life cycle of typical heteroecious, holocyclic aphid. (From Raman,
1982).
Pest biology, damage and distribution 481
(secondary) hosts are usually short-lived herbaceous species. The prima,ry
host range is usually very specific, whereas the secondary host range may
be broad. Alternation of hosts is an evolutionary adaptation that has
permitted aphids to exploit agricultural crops perhaps more successfully
than any other insect group.
The potato-colonizing aphid species that is most widely distributed,
commonly abundant, and efficient as a virus vector is green peach aphid
(van Emden et al., 1969, Mackauer and Way, 1976). Like most aphids of
agricultural importance, green peach aphid originated in the Palearctic
region (Dixon, 1987). Green peach aphid is a remarkably polyphagous
species having more than 875 secondary hosts (Tamaki, 1981).
Primary hosts of the green peach aphid are certain Prunus spp. including
peach and nectarine, P. persicae. In Europe, black cherry, P. serotina is an
important overwintering host (Hille Ris Lambers, 1972), although in
North America, where the tree is native, it is not considered a host.
Overwintering eggs hatch in spring giving rise to apterous, partheno-
genetic, viviparous stem mothers (fundatrices) which feed and reproduce
on the primary host (Miyazaki, 1987). Fundatrices that drop from the tree
may survive on secondary hosts. Beginning with the second generation,
alate morphs (spring migrants) appear. Peak alate production and migra-
tion occur in the third generation. Migrants invading potatoes usually are
both true spring migrants from primary host( s) and alatae produced on
secondary hosts, often weed species. Progeny of spring migrants are
invariably apterous, but in succeeding generations a small proportion are
alate. On potato, apterae are almost always the predominant morpho As
host plant quality declines, more summer migrants are produced and
these disperse to other plants. As day lengths shorten, fall migrants are
produced; these are both male and female (sexuales). Fall migrants return
to the primary host where the females (gynoparae) give birth to an
apterous female morph (oviparae). Oviparae mate with male fall migrants
and lay a fertilized overwintering egg. These eggs are usually laid near buds
of the primary host. In mild climates, continuous asexual reproduction
(anholocycly) can occur. However, under short photoperiods these popu-
lations may give rise to an increased proportion of alate virginoparae and
some males (androcycly).
Other potato-infesting aphid species of worldwide importance include
potato aphid, Macrosiphum euphorbiae; buckthorn aphid, Aphis nasturtii;
and the stolon-infesting Rhopalosiphoninus latysiphon. Among species
of regional importance are the Aphis gossypii complex (including A.
frangulae) , Aphis fabae, Aphis craccivora, Myzus (= Nectarosiphon)
ascalonicus, Aulacorthum (= Neomyzus) circumflexum, Aulacorthum
solani, Rhopalosiphum padi, Lipaphis erysimi, and Brachycaudus heli-
chrysi. Aphids differ from most other major taxa in that the greatest
diversity of species associated with a given plant species occurs in the
temperate regions (Dixon, 1987).
482 Insect pests
Potato viruses can be classified as either circulative (persistent) or
noncirculative (nonpersistent or semipersistent) with respect to their mode
of transmission by aphid vectors (Kennedy et al., 1962). All potato viruses
except PLRV are noncirculative. Noncirculative viruses are borne on the
stylets or are egested via the food canal of the maxillary stylets. Such
viruses can be acquired or transmitted in probes of only a few seconds
duration. In contrast, PLRV can only be acquired in extended phloem
feeding. After ingestion, PLRV passes through the midgut wall into the
haemolymph and other tissues. The insect is not competent to transmit
until the virus reaches the salivary glands, typically 8-24 h after acquisition.
Once acquired, PLRV-vector competence is usually retained for life.
Noncirculative viruses are quickly lost and never retained through a moult.
The primary vector of PLRV is green peach aphid. Other known vector
species include M. euphorbiae, M. ascalonicus, A. circumflexum, A.
solani, and A. nasturtii. In contrast, most noncirculative viruses can be
transmitted by many aphid species, including some that do not colonize
potato.
Circulative and noncirculative viruses present very different problems
with respect to vector control. Insecticides are generally much more
effective in preventing the spread of PLRV than in preventing the spread
of noncirculative viruses (Broadbent, 1957; Shanks and Chapman, 1965).
Even persistent toxic residues may not kill quickly enough to prevent the
transmission of noncirculative viruses by alate aphids that invade the field.
The principal benefit of insecticides used against noncirculative viruses is in
preventing within field (secondary) transmission.
Insecticide resistance and pesticide induced outbreaks are common
phenomena in aphid popUlations. Green peach aphid, like Colorado beetle
and Liriomyza spp., has demonstrated an ability to develop resistance
to all major insecticide classes: DDT and methoxychlor, BHC and
cyclodienes, organophosphates, carbamates and pyrethroids (Metcalf,
1980). Insecticide resistance greatly limits the insecticidal options that can
be used to control green peach aphid and accordingly is a significant
obstacle to developing potato IPM. Insecticide resistance in green peach
aphid has been attributed to the duplication of the gene sequence which
codes for carboxyesterase production. In highly resistant variants, carboxy-
esterase may account for 3% of total body protein (Devonshire and
Moores, 1982). When insecticidal treatments are discontinued against
green peach aphid, resistance levels remain stable for many generations,
but eventually collapse and revert to complete susceptibility (Bauernfeind
and Chapman, 1985).
Potato plants are most susceptible to virus infection when young
(Knutson and Bishop, 1964). Movement of PLRV and presumably other
viruses from the stem first inoculated to other stems decreases with plant
age at time of inoculation. (Flanders et al., 1990). Enzyme-linked immuno-
sorbent assays (ELISA) are useful in detecting specific potato viruses and
are commonly used to monitor the health of seed potatoes. However, reli-
ability of ELISA is dependent upon virus titre and testing of foliage may
not give a true indication of tuber infection (Tamada and Harrison, 1980a,
b; Hill and Jackson, 1984).
European seed producers have long relied on monitoring aphid flights
for making decisions on when to top kill potatoes. Suction traps, yellow
sticky traps, and yellow pan traps have all been used with success
(Heathcote, 1972; Taylor and Palmer, 1972). Yellow pan traps of the
M6ericke design have been credited with greatly benefiting the seed
industry in The Netherlands because each day harvest can be delayed
increases tuber yields (Hille Ris Lambers, 1972). Monitoring of alatae is
also used in parts of North America (Singh and Boiteau, 1987). The
rationale for monitoring aphid flights is the assumption that some
unacceptable proportion of the influx will be viruliferous. While traps are
useful for identifying periods of aphid flight activity on an area wide basis,
they do not provide a reliable basis for scheduling insecticide applications.
By the time alatae are captured in traps, apterae are usually already
present on potatoes. If the alatae are coming off virus-free hosts then pan
captures are of no significance other than indicating increasing aphid
numbers in the vicinity, which could be more readily determined by foliage
sampling. Leaf counts are considered the most reliable practical means of
monitoring within field populations.
Most European workers contend that virus spread is correlated with
captures of alatae and not with within field populations of apterae
(Broadbent, 1950, 1965). However, Ribbands (1965) offered contrary
evidence with respect to PLRV and showed that even the spread of PVY
sometimes correlated better with apterae than alatae. Hanafi et al. (1989)
found that the spread of PLRV from within field sources was correlated
with numbers of apterae and that within row spread was greater than
across row spread.
Various workers have proposed action thresholds for within field popu-
lations of green peach aphid apterae (Bacon et al., 1978; Byrne and
Bishop, 1979; Radcliffe et al., 1981a; Shields et al., 1984; Misra and
Agrawal, 1987; Parry, 1987). All suggested thresholds that differed little
from the 30 green peach aphid apterae per 100 leaves proposed more than
55 years ago by Davies (1934). For ware potatoes, these thresholds are
probably low, unless the cultivar is susceptible to net necrosis. For seed
potatoes, the concept of vector thresholds is somewhat questionable since
there is no practical way of knowing what proportion of the population is
viruliferous. For Minnesota seed potatoes tentative thresholds of 3-10
green peach aphid apterae per 100 leaves have been proposed (Flanders et
al., 1991).
484 Insect pests
11.7.2 Leafhoppers (Jassids)
Leafhoppers (Homoptera: Cicadellidae (= J assidae» are common pests in
the tropics and subtropics. In India, Amrasca biguttala biguttala and
Empoasca devastans are major potato pests (Saxena et al., 1974). However,
it is in North America that leafhoppers are of greatest importance as potato
pests. In the eastern half of the US and Canada, potato leafhopper must be
routinely controlled to prevent unacceptable yield losses (Granovsky,
1944; Johnson et al., 1986, 1987).
Potato leafhopper overwinters only in a permanent breeding area along
the Gulf Coast. In spring, flying adults are caught in updraughts and
transported north on upper level airstreams. Influxes of leafhoppers into
the North Central States are associated with southerly winds of 36 h or
more duration and precipitation in the fallout area (Huff, 1963). These
long-distance migrants often arrive before the potatoes have emerged.
Significant populations seldom appear on potato until a generation of new
adults has been produced on alternative hosts (Flanders and Radcliffe,
1989).
In the literature, potato leafhopper has been described as a stylet-sheath
feeder, but recent research (Kabrick and Backus, 1990) has shown that
sheath formation does not occur in most feeding probes. More typical of
potato leafhopper is lacerate and flush feeding. Hypertrophy of tissues
distal to the feeding site results in a physiological injury of potato termed
'hopperburn'. Hopperburn is associated with accumulation of photosyn-
thate due to occlusion of the phloem. Evidence suggests that it is the
insect's saliva that induces plant cells to enlarge blocking the phloem.
The first symptom of leafhopper feeding is subtle paling of the veins and
curling of the leaflet. Hopperburn begins as a triangular lesion at the tip of
infested leaflets. Lesions spread progressively back and inwards from the
margins finally destroying the leaves; plants senesce and die prematurely.
Feeding injury by nymphs and adult leafhoppers increases plant respira-
tion, depleting reserves available for growth and tuber development (Ladd
and Rawlins, 1965). Close linear relationships exist between increasing
leafhopper numbers, percent hopperburn and decreasing yield.
Potato leafhopper was originally thought to be the only empoascan
attacking potato in North America. However, there are at least four
closely related species: E. solani in humid southern areas; E. filamenta in
arid, high altitude western intermountain regions; and E. abrupta and E.
arida in arid, low altitude Pacific coast regions (Delong, 1931).
Certain other leafhopper species cause little direct feeding injury, but
are important on potato because they are vectors of potato diseases. In
North America, aster leafhopper, Macrosteles fascifrons transmits the aster
yellows mycoplasma-like organism (AY-MLO) which causes purple top. A
variety of other mycoplasma-like and viral pathogens of potato are
tranmsitted to potato by various leafhopper species.
11.7.3 Potato psyllid
Potato psyllid, Paratrioza cockerelli (Homoptera: Psyllidae), occurs
through the western US and is of particular importance in the Mountain
States. Uncontrolled, the insect can cause total crop loss. The saliva of the
nymph is toxicogenic and induces a condition called psyllid yellows. When
infestation occurs early, tuber set is increased, but the tubers fail to bulk.
The economic threshold for potato psyllid is said to be one nymph per 100
sweeps.
11.7.4 Colorado potato beetle

No insect defoliator is more destructive than the Colorado potato beetle,
Leptinotarsa decemlineata (Coleoptera: Chrysomelidae). It is also a pest on
tomato and eggplant. Colorado potato beetle is an oligophagous species
that feeds exclusively on Solanaceae and primarily on Solanum spp.
Prior to 1859, Colorado potato beetle was an uncommon species
occurring in the arid eastern foothills of the Rocky Mountains where it fed
on buffalo bur, S. rostra tum and S. angustifolium. Adaptation to potato
may have taken as long as 40 years (Casagrande, 1987). By the 1860s,
devastating losses were occurring throughout the midwestern states. During
the next 15 years Colorado potato beetle extended its range north into
Canada and east to the Atlantic. Colorado potato beetle became per-
manently established in Europe prior to 1922. Its spread east was rapid.,
reaching Germany in 1936, Poland in the mid-1950s, the western borders
of the USSR by 1959, and the Caucasus and Turkey by 1976.
Colorado potato beetle has two or three generations per year throughout
most of its range. In northern and high altitude areas there may be only
one generation; in southern regions four may be possible. The beetle
overwinters as an adult in the soil. Emergence in spring usually occurs
before potatoes have emerged. The insect is remarkably fecund, with
females typically laying 300-500 eggs. Eggs are laid in masses of 20 or
more. There are four larval instars, with 75% of total foliage consumption
by immatures occurring in the final instar. The insects are voracious
feeders and complete defoliation and even haulm destruction can result.
Where defoliation is continuous, yield loss can be total.
Colorado potato beetle has had a prominent place in the development of
crop protection entomology (Gauthier et al., 1981). Use of Paris green,
cuperous acetoarsenite, for beetle control was the first successful example
of insecticidal control of a crop pest. Many significant advances in
application equipment were in response to this pest. Examples include: the
first hand-operated compression sprayers, the first wheel-drawn sprayers,
the first traction-operated dusters, the first engine operated sprayer and the
first air-blast sprayer. Potatoes were also one of the first crops to be treated
by airplane.
486 Insect pests
Colorado potato beetle is an insect renowned for its ability to develop
resistance to insecticides. Resistance developed first and has been most
severe along Virginia's eastern shore and Long Island. However, high
levels of pyrethroid resistance now exist in beetle populations throughout
the northeastern and midwestern states. In the western US, resistance has
developed slowly, but has increased each year with seeming inevitability.
Insecticide resistance has, in recent years, greatly enhanced the pest status
of Colorado potato beetle over much of North America.
11.7.5 Flea beetles

Flea beetles (Coleoptera: Chrysomelidae) are important defoliating pests
in the Americas and Asia. In Peru, there are at least five species: Epitrix
parvula; E. subcrinita; E. ubaquensis; E. yanazara; E. harilana rubia. In
North America, the most important species are E. cucumeris, common
throughout the eastern half of the continent; western potato flea beetle, E.
subcrinita, widespread in the West and Mountain States; tobacco flea
beetle, E. hirtipennis, common in the South and Mountain States; and
tuber flea beetle, E. tuberis, which occurs in the Northwest. Adults are
1-2 mm long and feed on foliage, producing numerous minute circular
holes (1 mm dia.). Damaged leaves may desiccate. The feeding wounds
can serve as entry point for potato pathogens including bacterial diseases
and early blight (Hodgson et al., 1974). Larvae of all species feed on roots
and tubers, but most species cause only superficial injury. Tuber flea beetle
makes deep tunnels which render the potatoes unmarketable. These
tunnels are often invaded by fungi. Common scab infection is often
associated with flea beetle injury.
11.7.6 Leaf-feeding coccinellids

In India, Epilachna beetles, Henosepilachna (= Epilachna) VlgmtlOcto-
punctata and H. ocellata (Coleoptera: Coccinellidae) are important
defoliators which often limit successful potato cultivation (Misra and
Agrawal, 1988). Both adults and grubs skeletonize the leaves.
11.7.7 Wireworms
Many different species of wireworms (Coleoptera: Elateridae) are pests of
potatoes. In most regions, no one species can be regarded as a key pest,
but collectively wireworms represent a serious problem (Thompson, 1987;
Radcliffe et al., 1991). In most localities, the relative importance of the
various species is not well known. Wireworms have long life cycles (one to
several years) so knowledge of the previous cropping history is important.
Wireworms are usually worst following cereals. Damage is done by the
larvae tunnelling into the tubers. Tuber damage is usually characterized by
a clean, round entry hole (2-3 mm dia.) often entering straight into the
tuber. Injury does not reduce yields, but can make the crop unmarketable.
Important genera in North America include Limonius, Melanotus,
Ctenicera, Agriotes, Hypolithus, and Conoderus.
11.7.8 White grubs

During the last three decades, white grubs (Coleoptera: Scarabaeidae)
have become major potato pests in certain countries of Asia and Central
America. In India, at least eight species have been reported. Major pest
species include: Anomala spp. and Melolontha spp. (Singh and Verma,
1982). Larvae are the damaging stage and cause severe injury to seed
tubers and roots of new seedlings. In North America, Phyllophaga spp. are
occasional pests when potatoes are planted on land previously in sad or
pasture.
11. 7.9 Andean potato weevil complex

The Andean potato weevil complex, Premnotrypes spp. (Coleoptera:
Curculionidae), and a few other closely allied weevils, are potato pests of
exceptional destructiveness in high altitude (3000 m) tropics from Chile
and Argentina to Venezuela (European Plant Protection Organization,
1984). There are at least 15 species which attack potatoes, the most
important of which are: P. latithorax, P. pusillus, P. solani, P. suturicallus,
P. vorax, P. fractirostris, P. sanfordi, P. piercei, Amitrus (= Canephoro-
tomus) jelskysi and Rhigopsidius tucumanus (Munro, 1968). The adult
weevils feed on young potato plants, and severe infestations may com-
pletely denude the field. Life-cycles of the species differ only in detail.
There is usually one generation per year. The dry winter is passed as an
adult within the tuber or the soil. When the rains start, adults emerge and
feed mainly on leaves and stems of potato plants. Eggs are laid on potato,
weeds, or in the soil. The larvae tunnel in tubers, and in fields not
protected by insecticides, loss of marketable yield can be total (Munro,
1954; Alcala and Alcazar, 1976; Calvache, 1986). Pupation is either within
the tuber or in an earthen cell of surrounding soil.
Cultural control measures are effective against Andean potato weevils.
Recommended measures include selection of clean seed, deep planting,
hilling, careful soil cultivation, removal of weed hosts, early harvest, and
ditches to protect fields from invasion by migrating adults (Yabar, 1988a).
Adult weevils can be monitored with pit fall traps (Rojas, 1978). The traps
are checked weekly and insecticides are applied upon first capture of
adult weevils. Use of this method can greatly reduce insecticide use over
what would be applied on a prophylactic basis. In Colombia, more than
$(US) 22 million is spent annually on insecticides for control of P. vorax
(International Potato Center, 1984).
488 Insect pests
11.7.10 (Potato) tuber moths
Lepidopterous species in several families cause serious damage to
potatoes. Most devastating are four species of tuber moths (Lepidoptera:
Gelechiidae): potato tuber moth, Scrobipalpula absoluta, Symmetrischema
plaesiosema and Scrobipalpopsis solanivora. Tuber moths are pests in
warm temperate and tropical areas. Potato tuber moth is a cosmopoli-
tan pest now of worldwide distribution (Commonwealth Institute of
Entomology, 1968; Izhevsky, 1985). S. absoluta, S. plaesiosema and S.
solanivora are pests of the Neotropics.
Although quarantine is commonly practised against infested tubers,
tuber moths continue to spread. Potato tuber moth is assumed to have
originated in the humid subtropics of South America. Evidence for this
origin is that its principal host plants, potato and tobacco, Nicotiana
tabacum, are indigenous to that area and there the pest is associated with a
rich parasite complex and is of minor importance (Lloyd, 1972).
Potato tuber moth damages both foliage and tubers. On foliage, eggs are
deposited on dorsal leaf surfaces and emerging larvae mine leaflets. As
leaflets are consumed, larvae may leave their tunnels and web adjacent
leaflets together. Larvae sometimes mine stems causing entire terminals to
die. When tubers are exposed moths may oviposit directly on them.
Partially grown larvae sometimes descend from the plant and invade
tubers, particularly when the soil is dry and deeply cracked. When tuber
infestations are high, the crop may not be worth harvesting as it is
prohibitively time consuming to sort out infested tubers. When infested
tubers enter storage, larvae can continue to develop, tunnelling exten-
sively, filling the tubers with frass, and permitting entry of decay
organisms. In storage, losses can be almost total. Potato appears to be the
preferred host, but many plant species, cultivated and wild, are attacked.
Potato tuber moth is the most important potato pest throughout the
Middle East and North Africa (von Arx et al., 1987,1988). In primitive
storage systems, losses can be total even when insecticides are used. In San
Ramon, Peru, tuber infestation of over 90% has been recorded following
four months of storage (Raman et at., 1987).
Sex pheromones provide a useful tool for monitoring populations of
potato tuber moths under both field and storage conditions. Potato tuber
moth pheromone consists of two components, trans-4, cis-7-tridecatrien-
1-01 acetate (PTM-1) (Roelofs et al., 1975) and trans-4, cis-7, cis-10-
tridecatrien-1-01 acetate (PTM-2) (Persoons et al., 1976). Blends of PTM-1
and PTM-2 are more effective than either alone, but the ratio is not
important (Voerman and Rothschild, 1978). Various trap designs have
been used, but combination pheromone/water traps work best (Bacon et
al.,1976).
For potato tuber moth, mass trapping using sex pheromones has
demonstrated the feasibility of direct control (Raman, 1988a). In field
studies, trap densities of one funnel trap per 25m 2 reduced tuber damage
by 45% compared to plots receiving no pheromones. In storage, both
water-pan and funnel traps were effective in reducing tuber moth damage.
In the western US, pheromone traps are used to schedule insecticide
applications (Flint, 1986). A threshold of 40 moths in anyone night or a
mean of 15-20 per night is used for fresh market potatoes. In Costa Rica, a
threshold of 100 moths per trap per week is used to determine the need to
treat with insecticide (Rodriguez and Lepiz, 1988).
Symmetrischema plaesiosema now occurs in Peru, Colombia, the US and
Australia. In the field, larvae mine foliage and bore into stems making
chemical control extremely difficult (Sanchez and Aquino, 1986). In
storage, the tubers are destroyed by larval mining. Infested tubers have a
foul odour and are unfit for consumption or seed. Tuber damage in 51 farm
stores surveyed in Peru varied between 2% and 78% (Raman, 1988b).
Scrobipalpula absoluta damages only above ground portions of the
plant. It is a pest of both potatoes and tomatoes in high altitude, temperate
locations in Peru, Chile, Argentina, Colombia and Brazil (Delgado and
Aguilar, 1980; Moore, 1983). Susceptible potato genotypes can be com-
pletely destroyed by this pest.
Scrobipalpopsis solanivora is a major pest in Central America. Larvae of
this species primarily attack tubers. It is believed that this species origin-
ated in Guatemala (Povolny, 1973). S. solanivora was discovered in Costa
Rica in 1970, apparently having been introduced with a shipment of
potatoes. In 1972, it caused yield losses of 20-40% over a 2000 hectare area
of Costa Rica. Recently, this species has been reported in Venezuela and
Colombia. It is imperative that strict quarantines be established to prevent
further spread of this pest. A sex pheromone, (E)-3-dodecenyl acetate, has
been identified for S. solanivora (Nesbitt et al., 1985).
11.7.11 Cutworms
Numerous species of cutworm (Lepidoptera: Noctuidae) attack potato.
Among the most cosmopolitan and destructive are Agrotis spp. In India,
A. ipsilon and A. segetum are major pests on potato (Misra and Agrawal,
1988). Cutworm larvae are nocturnal, living just below the ground surface.
Larvae can cut the stalks of young growing plants, with severely infested
fields appearing grazed. On more mature plants, cutworms can damage
tender stems. After tuberization, cutworm damage is largely confined to
the tubers (Chandla et al., 1977).
11.7.12 European corn borer

Potato is a preferred host of European corn borer, Ostrinia nubialis
(Lepidoptera: Pyralidae). Plant damage is usually minor, but in some
locations, e.g. North Carolina, European corn borer is regarded as a
490 Insect pests
serious pest. Injury is most likely when oviposition occurs before maize has
emerged or is still very young. Tunnelled haulms wilt readily and are prone
to twist and break in the wind. Larval tunnelling may predispose the plant
to Erwinia spp. Cultivars differ greatly in susceptibility.
11.7.13 Leafminer flies

Leafminer flies, Liriomyza spp. (Diptera: Agromyzidae), are the most
important dipterans attacking potato. Leafminer flies are pests in warm
temperate and tropical regions. In South America, L. huidobrensis can
cause yield reductions of 35% or more (Chavez and Rama, 1987; Yabar,
1988b). Plants are attacked shortly after emergence. Adult females
puncture the leaves to feed or lay eggs. Eggs are inserted individually on
lower leaf surfaces and hatch within 3---4 days. Larvae tunnel into the
leaflets making characteristic serpentine mines. L. trifolii is the most
important species in Africa, from Senegal to Kenya and Mauritius
(Bourdouxhe, 1982). Outbreaks of leafminer flies are often induced by
intensive use of insecticides. In the Canete Valley of Peru, insecticide-
resistant L. huidobrensis has, over the past 10 years, emerged as a major
pest. Sticky traps have been used by Chavez and Raman (1987) to reduce
leafminer L. huidobrensis damage.
11.7.14 Thrips and spider mites

Thrips (Thysanoptera: Thripidae) and mites (Acarina: Tetranychidae) are
becoming increasingly important in many tropical countries. In Indonesia
and the Philippines, severe thrips and mite infestations have been
reported in lowland potato production. Two thrips species, Thrips palmi,
Megaulothrips usitatus and the broad mite, Polyphagotarsonemus latus, are
abundant in these countries (Eveleens and Woodford, 1982; Misra and
Agrawal, 1988). The biology of these thrips is still not fully known. An
international effort has been initiated by CIP to develop IPM programmes
for these pests.
11.8 PEST CONTROL MEASURES
11.8.1 Host plant resistance (HPR)

Host plant resistance has not been exploited against insects to nearly the
extent that it has against plant pathogens and nematodes. Ideally, host
plant resistance should be a primary defence against insect pests. Unfortu-
nately, cultivar differences alone usually do not provide adequate control.
Many wild species have characteristics potentially useful in breeding for
resistance to insects (Radcliffe et al., 1981b; Radcliffe, 1982). Present
Pest control measures 491
European and North American cultivars represent a decidedly narrow

sampling of the genetic diversity existing in the primitive cultivated
potatoes of South America and owe even less to the approximately 160
known wild potato species.
Resistance to Colorado potato beetle has long been known to be
associated with the presence of foliar glycoalkaloids, as in S. chacoense
(Kuhn and Low, 1955), but breeding efforts to exploit this resistance have
met little success. Two major problems have been encountered: levels of
tuber glycoalkaloids tend to be highly correlated with levels in the foliage,
and resistance always declines in backcrosses. Now that resistance has been
associated with specific glycoalkaloids, especially leptines, commersonine
and dehydrocommersonine (Sinden et ai., 1980), it may be possible to
make greater progress.
In other wild potato species, insect resistance is associated with glandular
pubescence as in S. berthaultii (Gibson, 1971; Tingey, 1984). Gibson
(1971) described two types of glandular trichomes (Fig. 11.6): 'Type A'
which are short-stalked with a four-lobed, membrane-enclosed head that
ruptures on contact releasing a sticky exudate; and 'Type B' which are
longer, tapering, and terminate in a simple sticky globule which is
continuously exuded. Small insects and mites are entrapped by the sticky
droplets of the type B hairs. Larger insects are entrapped by the combined
action of both trichome types (Gibson and Turner, 1977). Body parts that
Figure 11.6 Glandular trichomes of Solanum berthaultii. 'Type A' , short-stalked,

four-lobed; 'Type B', long-stalked, tapering, with a simple droplet. x 126.
(Original SEM, International Potato Center (CIP).)
492 Insect pests
become coated with Type B exudate tend to adhere to the membrane head
of Type A trichomes which rupture as the insect struggles to free itself.
Type A exudate, which contains polyphenol oxidase, hardens on the feet
and mouthparts, preventing feeding and often trapping the insect (Gregory
et al., 1986). Contact with Type B exudate appears to agitate green peach
aphid causing it to break the membranes of the Type A glands and become
entrapped. Type B exudate contains a sesquiterpenoid fraction similar or
identical to aphid alarm pheromone (Gibson and Pickett, 1983).
Breeding for trichome-conferred resistance began at Cornell University
in 1977 when accessions of S. berthaultii were crossed with the cultivar
'Hudson'. From this population, Tingey et al. (1982) selected three F3
hybrid clones, with aphid resistance equal to the S. berthaultii parent.
Mehlenbacher et al. (1983) concluded that this resistance behaved as a
quantitatively inherited trait. S. tuberosum x S. berthaultii hybrids have
been selected with high levels of resistance to Colorado potato beetle
(Wright et al., 1985). Larval populations were reduced by as much as 90%
on the most resistant hybrids. Resistance appears to involve deterrence of
oviposition and a slowing of larval growth.
Potato breeders at CIP are endeavouring to introduce glandular
trichomes into material adapted to the lowland tropics. For developing
countries, glandular trichome mediated resistance could be a great benefit
in the suppression of potato tuber moth, aphids, leafminer flies, thrips and
mites. CIP has screened its germplasm collection for resistance to potato
tuber moth, L. huidobrensis, and P. suturicallus (Raman, 1989). P.
operculella-resistant clones have been distributed to national programmes
in several developing countries to confirm resistance. The US Department
of Agriculture has funded cooperative agreements with various universities
to screen the US potato germplasm collection for resistance to Colorado
potato beetle, green peach aphid, and potato leafhopper.
11.8.2 Cultural control

Cultural practices have proved effective against many potato pests. For
potato tuber moth, infestation is minimized by hilling, timely irrigation,
early harvest, rapid handling, and storage in well designed stores. In the
tropics, potatoes are commonly intercropped with maize, beans or other
crops. By the choice of intercropped species and appropriate agronomic
practices it is possible to reduce populations of various potato pests (Potts,
1988). Potato intercropped with wheat is less damaged by L. huidobrensis
because of the attraction of wheat to parasitoids of the leafminer
(Raymundo and Alcazar, 1983). Peruvian farmers sometimes intercrop
potato with maize, Oxalis tuberosa, and other tuberous crops to reduce
pests. The same approach is used by Asian farmers who intercrop potato
with onion or garlic to reduce aphids and possibly other pests. Crop
rotation effectively delays the establishment of Colorado potato beetle
Pest control measures 493
(Wright, 1984). For long-lived insects such as wireworms and white grubs,
previous cropping history determines whether there will be a problem or
not.
11.8.3 Repellent plants and botanical insecticides

For centuries, farmers in many developing countries have protected their
crops with natural insect repellents found in leaves and fruits of certain
plants (Grainge and Ahmed, 1988). Andean farmers burn dried capsicum
fruits to fumigate potato stores against potato tuber moth. The Incas used
dried leaves of Muiia, Minthostachys spp., in their potato stores to obtain
clean healthy potatoes. Unfortunately, this effective, age-old practice has
largely disappeared with the introduction of modern pesticides. Several
local Peruvian plants, including Lantana camara, Eucalyptus globulus and
Minthostachys spp. can be used to protect stored tubers against potato
tuber moth and aphids (Raman et al., 1987). L. camara also proved
effective in rustic stores in Nepal (Pradhan, 1987). In India, L. aculeata has
been used (Lal, 1987). P. vorax has been controlled in Colombia (Oilvache
and Posada, 1987) by planting borders of O. tuberosa around potato fields.
In collaboration with CIP, many national programmes are evaluating local
plants as protectants against potato tuber moth.
Much recent attention has focused on the botanical insecticide, neem,
extracted from the kernel of the neem tree, Azadirachta indica. Neem is
highly effective in controlling the Colorado potato beetle. Against the
beetle, neem is both an antifeedant and a disruptant of metamorphosis.
Beetles exposed to low dosages have reduced fecundity and generally are
sterile (Schmutterer, 1987).
11.8.4 Biological and behavioural control

Despite the vast amount of research done on potato pests, little attention
has been given to natural enemies and microbial agents. What information
does exist has only in a few instances been integrated into successful pest
management strategies.
Aphid populations are often severely depleted by parasitoids, predators
and entomopathogenic fungi, but there is usually a time lag between aphid
colonization of the crop and establishment of effective control by biotic
agents. In Maine, parasitoids were usually not effective in controlling
the aphids that colonize potato (Shands et al., 1965). That insecticides
are routinely applied to potato obviously does not favour parasitoids.
Predators appear to be considerably more effective than parasitoids
(Shands et al., 1972) and are especially valuable when they attack the
aphids on the primary hosts (Tamaki et al., 1967). Coccinellids, syrphids,
chrysopids, anthocorids, nabids and lygaeids are among the most impor-
tant taxa of aphid predators.
494 Insect pests
A complex of entomopathogenic fungi also attack potato-infesting
aphids. Fungi can be exploited by cultural manipulations to enhance
infection or as mycoinsecticides. The most important aphid pathogen in
North America is Conidiobolus obscurus. This fungus has been used as a
mycoinsecticide in the USSR. Another fungus, Verticillium lecanii, has
been developed for aphid control in greenhouses (Bennett, 1984) and
should be tested as a control for aphids on potato in open stores.
Colorado potato beetle has been controlled experimentally with Bacillus
thuringiensis var. thuringiensis ~-exotoxin (Cantwell and Cantelo, 1984),
entomopathogenic fungi, particularly white muscardine disease, Beauveria
bassiana (Roberts et al., 1981) and Paecilomyces farinosus (Bajan, 1973) and
the nematodes Neoaplectana spp. (Bennett, 1984). Exploration for natural
enemies of other species of Leptinotarsa in South and Central America has
led to the discovery in Colombia of Edovum puttleri (Hymenoptera:
Eulophidae) (Schroder and Athanas, 1985). This parasitoid is seasonally
effective, but cannot overwinter in temperate regions. The potential of
strains from colder areas should be assessed. In more valuable crops such
as tomato or egg plant, annual inoculative releases may be useful. European
efforts to establish North American parasitoids and predators against
Colorado potato beetle were unsuccessful (Szmidt and Wegorek, 1967).
Bacillus thuringiensis var. san diego and B. thuringiensis var. tenebrionis
produce a polypeptide toxin effective against young Colorado potato
beetle larvae (Herrnstadt et al., 1986). These have been commercialized as
microbial insecticides for use against insecticide resistant Colorado potato
beetles. The products are relatively expensive and have a short persistence
on the foliage, but it is possible to insert the gene sequence into a different
bacterium which can then be killed with the cell remaining intact,
encapsulating the toxin. This approach could increase toxin yields and
produce an environmentally more stable product.
Recently, several private companies have inserted the gene for produc-
tion of Bacillus thuringiensis {i-endotoxin into potato plants. Transformed
potato plants are now being tested for resistance to Colorado potato beetle
and potato tuber moth. Availability of such cultivars might greatly reduce
the need for insecticides, but there is concern that this approach would
pose considerable selection for B. thuringiensis resistance in the pests.
The potential for control of Andean potato weevils by manipulation of
parasitoids and predators appears to be limited, but prospects for microbial
control are more promising. B. bassiana was highly effective in field trials
in Colombia and Peru (Bennett, 1984).
Eleven parasitoids of L. huidobrensis have been reported from Peru;
the predominant species was Halticoptera patellana (Hymenoptera:
Pteromalidae) (Raman and Redolfi, 1984). The Commonwealth Institute
of Biological Control (CIBC) has conducted surveys for natural enemies of
L. trifolii and has arranged the introduction of several North American,
Caribbean and South American parasitoids into Senegal (Murphy, 1984).
Integrated pest management (IPM) 495
Earlier introduction of several of these parasitoids into Hawaii reduced the
need for chemical control.
Potato tuber moth parasitoids from South America have been intro-
duced in countries around the world by the CIBC and other organisations.
Parasitoids successfully established in other countries include: Campoplex
haywardi (Hymenoptera: Ichneumonidae); Apanteles subandinus and
Orgilus lepidus (Hymenoptera: Braconidae); and Copidosoma kohleri
(= uruguayensis) , and C. desantisi (Hymenoptera: Encrytidae). In Cyprus,
releases initiated in 1965 resulted in the establishment of at least
three parasitoid species and substantial potato tuber moth control
(Commonwealth Institute of Entomology, 1968). Following the establish-
ment of A. subandinus and C. kohleri in Zimbabwe in 1965 (Mitchell,
1978), potato tuber moth became so scarce that regular surveys for it were
discontinued in 1969 (Sankaran and Greathead, 1980). Similar success was
achieved in Zambia (Cruickshank and Ahmed, 1973). Partial control has
also been reported in Australia, Mauritius, Madagascar, St Helena, New
Zealand, South Africa and the US. Unfortunately, biological control has
since been largely negated in east Africa by increased use of insecticides
targeted against other potato pests. In other countries, failure to establish
potato tuber moth parasitoids or the ineffectiveness of those that are
established may be a consequence of heavy use of insecticides on the crop.
Future success of biological control will depend on finding ways that
insecticides can be used selectively.
A highly pathogenic granulosis virus specific to potato tuber moth larvae
was identified in Australia (Reed and Springett, 1971). Applied as a spray,
the virus gave control equivalent to that obtained using insecticides with
the added benefit of persistence from year to year. A granulosis virus has
also been observed in Peru. A simple procedure has been devised for mass
rearing potato tuber moth (Raman, 1988b) permitting production of large
quantities of this virus for use in developing countries. The virus has
proved effective in preventing potato tuber moth development and
reducing tuber damage under both field and storage conditions (Raman
and Alcazar, 1988). This virus has proved effective in Colombia,
Guatemala, Bangladesh, Yemen, Egypt and Tunisia. B. thuringiensis var.
thuringiensis is also effective in control of potato tuber moth in storage.
11.9 INTEGRATED PEST MANAGEMENT (IPM)
In recent years, considerable attention has been given to the development

of potato IPM. IPM has been defined variously, but the common theme is
that our reliance on pesticides should be reduced by substituting the
coordinated use of alternative control measures. US President Carter, in
his 1979 Environmental Message, expressed this concept in these words:
'IPM uses a systems approach to reduce pest damage to tolerable levels
496 Insect pests
through a variety of techniques, including natural predators and parasites,
genetically resistant hosts, environmental modifications and, when necessary
and appropriate, chemical pesticides'. A major impetus for development
of IPM has been recognition of the adverse consequences of excessive
reliance on pesticides.
Many national and international organizations have committed resources
to IPM. For example, CIP has increased its participation in IPM activities
by establishing collaborative arrangements with several national agricul-
tural research programmes. CIP presently participates in five collaborative
research networks, three in Latin America, one in east Africa and one in
south-east Asia. These networks are dedicated to the improvement of the
potato crop research and transfer of technology (Ezeta, 1988).
IPM strategies generally rely first upon biological defences against pests
before chemically altering the environment. For potato, these control
measures may include use of resistant or tolerant varieties, introduction or
enhancement of natural enemies, cultural manipulations of the crop and
pest environment, and careful selection and timing of chemical applica-
tions (Cisneros, 1984). All pest management decisions are, in the final
analysis, economic decisions so the ultimate goal must be optimization of
return. External and long-term costs should be included in this analysis. In
view of such considerations, successful implementation of IPM often
necessitates inputs from both biological and social scientists to facilitate
development of appropriate component technologies and assure their
successful adaptation to user and society needs.
In North America, efforts are being made to implement potato IPM
programmes on a regional scale (Smilowitz, 1981; Bishop et al., 1982;
Shields et al., 1984; Parry, 1987). The focus of each of these programmes
varies somewhat, but the goal of each is comprehensively to apply
knowledge from various subdisciplines to optimize potato pest manage-
ment. Implementation efforts often reveal knowledge gaps which in turn
serve to focus research where most needed. For example, in Maine and
New Brunswick, implementation of potato seed IPM programmes have
necessitated reassessment of virus/aphid vector ecology (Storch, 1981;
Singh and Boiteau, 1987). An important recent advance has been the use
of microcomputers for data input and processing, pest and yield loss
prediction, and for disseminating information. Many growers have their
own microcomputers and either use software directly or subscribe to
centralized programs that can be accessed by telephone (Slack, 1987).
The US National IPM Program (Miller and Griffitts Clifford, 1985) has
targeted potatoes for competitive funding of IPM research in two of the four
USDA regions, North Central and North East. Funding priority is given to
multidisciplinary, multi state projects that address research topics and pests
as identified by the Regional IPM Committees. In the North Central
States, a funded project (NC-166) was established to apply a systematic,
systems approach to the allocation of research resources (Johnson and
References (IPM) 497
Mather, 1988). This analysis was subsequently embodied in the proposal
guidelines for North Central Potato IPM proposals. Topics identified in
these guidelines include epidemiology and ecology of insects and weeds,
pest-crop interactions, development and validation of biological monitor-
ing for IPM, development of innovative control tactics, and pest manage-
ment science. Within each of these topics, specific areas of emphasis were
identified. A primary objective of the National IPM Program has been to
focus research resources on objectives of national priority.
Much discourse exists on the status of IPM in developing countries, but
little has been done to implement the ideas that are often expressed. The
argument is often advanced that in developing countries there usually is
not enough information available on the various components to formulate
IPM programmes. However, it is clear that considerable knowledge already
exists, and that in many or possibly most situations, practical IPM pro-
grammes could be formulated. Implementation may in fact be the best way
of determining where scarce research resources should be allocated. Modi-
fications to fine tune the programmes can proceed as our knowledge base
expands. If we are to wait until we have all the answers we will never start.
Some initial efforts have been made to formulate IPM programmes for
the control of potato tuber moth in developing countries. Several combina-
tions of control components have been recommended for testing locally to
identify the most effective method (Raman and Booth, 1983). For the
tropics, IPM must be tailored to fit the specific conditions of an area. Once
the programme is available, the widespread adoption depends largely on
the success of demonstrating to farmers that the system works advan-
tageously. Best results occur in areas where growers have some kind of
organization and there is a system to provide information and technical
assistance.
IPM in the tropics will become reality only when the component
technologies are taken by farmers and adapted to their conditions.
Sponsored pest surveys and on-farm research can play important roles in
facilitating this process. National and international programmes should
encourage such activities. Farmers' perceptions and knowledge of pests are
an important source of information and can be very useful in formulating
IPM programmes. There remains, however, pressing need for much
additional research.
REFERENCES
Adams, J.B. and Kelley, R.A. (1950) Potato aphid control studies, 1946-1949, at
Woodstock, N.B., Canada. Am. Potato J., 27,175-82..
Alcal<l, c.P. and Alcazar, S.J. (1976) Biologia y comportamiento de Premnotrypes
suturicallis Kuschel (Col.: Curculionidae). Rev. Peruana Entomol., 19(1),49-52.
Bacon, D.G., Seiber, J.N. and Kennedy, G.G. (1976) Evaluation of survey
498 Insect pests
trapping techniques for potato tubeworm moth with chemical baited traps. J.
Econ. Entomol., 69, 569-72.
Bacon, D.G., Burton, V.E. and Wyman, J.A. (1978) Management of insect pests
on potatoes. Calif. Agric., 32(2), 2&-7.
Bagnall, R.H. (1977) Resistance to the aphid-born viruses in the potato, in Aphids
as Virus Vectors (eds K.F. Harris and K. Maramorosch), Academic Press, New
York, NY, pp. 501-26.
Bajan, C. (1973) Paecilomyces fumoso-roseus (Wize)-pathogenic agent of the
Colorado beetle (Leptinotarsa decemlineata Say). Ekol. Pol., 21(45), 705-13.
Bauernfeind, RJ. and Chapman, R.K. (1985) Nonstable parathion and endosulfan
resistance in green peach aphids (Homoptera: Aphididae). J. Econ. Entomol.,
78,51&-22.
Bennett, F.D. (1984) Biological control in IPM, in Report of XXII Planning
Conference on Integrated Pest Management, 4-8 June 1984, Int Potato Cent.
(CIP), Lima, Peru, pp. 189-97.
Bishop, G.W., Homan, H.W., Sandvol, L.E. and Stoltz, RL. (1982), Management
of Potato Insects in the Western States. A Western Regional Extension Publica-
tion, WREP 64, pp. 1-31.
Boiteau, G. (1983) The Arthropod Fauna of Potato Fields: Composition and
abundance, Agric. Canada Res. Br. Conltrib. 1983-16E, pp. 1-57.
Bourdouxhe, L. (1982) La mineuse nord-americaine des feuilles (Liriomyza trifolii),
diptere, Agromyzidae) sur cultures maraicheres au Senegal. Bull. Phytosan#'Iilire
FAO, 30, 81-2.
Brader, L. (1988) Needs and directions for plant protection in developing coun-
tries. The FAO view. FAO Plant Prot. Bull., 36(1), 2-8.
Broadbent, L. (1950) The correlation of aphid numbers with spread of leafroll and
rugose mosaic in potato crops. Ann. Appl. Bioi., 37, 58-65.
Broadbent, L. (1957) Insecticidal control of the spread of plant viruses. Ann. Rev.
Entomol., 2, 339-54.
Broadbent, L. (1965) The importance of alate aphids in virus spread within crops.
Proc. XIlth Int. Congr. Entomol., London, 1964, pp. 523-4.
Byrne, D.N. and Bishop, G.W. (1979) Relationship of green peach aphid numbers
to spread of potato leaf roll virus in southern Idaho. J. Econ. Entomol., 72,
809-11.
CHvache, H.G. (1986) Aspectos biol6gicos y ecol6gicos del gusano blanco de la
papa Premnotrypes vorax (Hustache). Memorias del curso sobre 'Control
Integrado de Plagas de Papa' (ed L. Valencia), Int. Potato Cent. (CIP)-Inst.
Colombiano Agropecuario (ICA), Bogota, Colombia, pp. 18-24.
CaIvache, H.G. and Posada, L. (1987) Efecto de barreras vegetales en el control
del gusano blanco de la papa. XIII Reunion (ALAP) Asoc. Latinoamericana de
la Papa. Memoria, Mar. 9-13, 1987, Panama, Panama, pp. 355-62.
Cantwell, G.E. and Cantelo, W.W. (1984) Control of the Colorado potato beetle
with Bacillus thuringiensis var. thuringiensis. Am. Potato J., 61, 451-9.
Carden, P.W. (1965) Economics of aphid control on ware potatoes. Proc. Br.
Insectic. Fungic. Conf., 3rd. Brighton, 1965, pp. 308-16.
Casagrande, RA. (1987) The Colorado potato beetle: 125 years of mismanage-
ment. Bull. Entomol. Soc. Am., 33 (3), 142-50.
Chandla, V.K., Misra, S.S., Verma, K.D. and Bist, B.S. (1977) Insecticidal control
of Agrotis spp. infesting potato crop. Pesticides, 11(2), 29-30.
References 499
Chavez, G.L. and Raman, K.Y. (1987) Evaluation of trapping and trap types to
reduce damage to potatos by the leafminer Liriomyza huidobrensis (Diptera:
Agromyzidae). Insect Sci. Applic., 8, 369-72.
Cisneros, F.M. (1984) The need for integrated pest management in developing
countries, in Report of the XXII Planning Conference on Integrated Pest
Management, 4--8 June, 1984, Int. Potato Cent. (CIP), Lima, Peru, pp. 19-30.
Commonwealth Institute of Entomology (1968) Distribution maps of insect pests.
Phthorimaea operculella (Lepidoptera: Gelechiidae). Commonw. Inst. Entomol.
Publ., Ser. A (Agric) Map 10 (revised), p. 1.
Cruickshank, S. and Ahmed, F. (1973) Biological control of potato tuber moth
Phthorimaea operculella (Zell.) (Lep.: Gelechiidae) in Zambia. Commonw.
Inst. Bioi. Control (CIBC) Tech. Bull., 16, pp. 147--62.
Davies, W.M. (1934) Studies on aphids infesting the potato crop. II. Aphis survey:
its bearing upon the selection of districts for seed potato production. Ann. Appl.
Bioi., 21, 283-99.
Delgado, J.J. and Aguilar, P.G. (1980) Control integrado en papa.. Rev. Peru.
Entomol., 23, 102-4.
DeLong, D.M. (1931) Distribution of the potato leafhopper (Empoasca fabae
Harris) and its close relatives of Empoasca. f. Entomol., 24, 475-9.
Devonshire, A.L. and Moores, G.D. (1982) A carboxylesterase with broad substrate
specificity causes organophosphorus, carbamate, and pyrethroid resistance in
peach-potato aphids (Myzus persicae). Pestic. Biochem. Physiol., 18, 235-46.
Dixon, A.F.G. (1987) The way of life of aphids: host specificity, speciation and
distribution, in Aphids: Their biology, natural enemies, and control (World Crop
Pests; 2A) (eds A.K. Minks and P. Harrewijn), Elsevier, Amsterdam, The
Netherlands, pp. 197-207.
European Plant Protection Organisation (1984) EPPO data sheets on quarantine
organisms, Premnotrypes spp. (Andean species). EPPO Bull., 14(1), 55--60.
Eveleens, K.G. and Woodford, J.A.T. (1982) Management of insect pests of
potato in Indonesia, in Potato Production in the Humid Tropics (eds L.J.
Harmsworth, J.A.T. Woodford and M.E. Marvel). Proc. of Third Int. Sympo-
sium on Potato Production for the Southeast Asian and Pacific regions, 12-17
October, 1980, Bandung, Indonesia. Int. Potato Cent., Far East and Southeast
Asia Regional Office, c/o Philippine Counc. Agric. Resour. Res. Dev.
(PCARRD), Los Banos, Laguna, Philippines, pp. 289-305.
Ezeta, F.N. (1988) Collaborative country research networks. Reg. Res., Int. Potato
Cent. (CIP), Lima, Peru, pp. 1-12.
Fano, M. and Ewell, P.T. (1986) EI control de plagas en papa en la sierra del Peru
Sur. Bo!. Inf. Agrario (Cuzco, Peru) ano 9, 96, 40--6.
Flanders, K.L. and Radcliffe, E.B. (1989) Origins of potato leafhoppers (Homop-
tera: Cicadellidae) invading potato and snap bean in Minnesota. Environ.
Entomo!., 18, 1015-24.
Flanders, K.L., Ragsdale, D.W. and Radcliffe, E.B. (1990) Use of enzyme-linked
immunosorbent assay to detect potato leaf roll virus in filed grown potato, cv
Russet Burbank. Am. Potato f., 67, 589--602 and 67, 811.
Flanders, K.L., Radcliffe, E.B. and Ragsdale D.W. (1991) Potato leaf roll virus
spread in relation to densities of green peach aphid (Homoptera: Aphididae):
Implications for management for Minnesota seed potatoes. f. Econ. Entomol ..
83, 1028-36.
500 Insect pests
Flint, M.L. (1986) Integrated Pest Management for Potatoes in the Western United
States. Publ. 3316, Univ. of Calif. pp. 1-146.
Forgash, A.J. (1985) Insecticide resistance in the Colorado potato beetle, in
Proceedings of the Symposium on the Colorado Potato Beetle, XVIIth Inter-
national Congress of Entomology (eds D.N. Ferro and RH. Voss). Mass.
Agric. Exp. Stn, Res. Bull. 704, pp. 35-52.
French, H.J. (1983) Macrosiphum euphorbiae (Thos.) injury to potato plants,
Ph.D. Thesis, University of Cambridge, UK.
Gauthier, N.L., Hofmaster, RN. and Semel, M. (1981) History of Colorado potato
beetle control, in Advances in Potato Pest Management (eds J.H. Lashomb and
R Casagrande), Hutchinson Ross Pub. Co., Stroudsburg, Penn. pp. 13-33.
Gibson, R.W. (1971) Glandular hairs providing resistance in certain wild speci,es.
Ann. Appl. BioI., 68, 113-9.
Gibson, RW. (1978) Pest aspects of potato production, in The Potato Crop: The
scientific basis for improvement (ed. P.M. Harris), Chapman and Hall, New
York, NY, pp. 470-503.
Gibson, R.W. and Pickett, J.A. (1983) Wild potato repels aphid by release of aphid
alarm pheromone. Nature, 302, 608-9.
Gibson, R.W. and Turner, R.H. (1977) Insect-trapping hairs on potato plant.
PANS (Pest. Artic. News Summ.), 22, 272-7.
Grainge, M. and Ahmed, S. (1988) Handbook of Plants with Pest-Control
Properties, John Wiley & Sons, New York, NY pp. 1-470.
Granovsky, A.A. (1944) Tests of DDT for the control of potato insects. J. Econ.
Entomol., 37, 493-99.
Gregory, P., Tingey, W.M., Ave, D.A. and Bouthyette, P.Y. (1986) Potato
glandular trichomes: a physicochemical defense mechanism against insects, in
Natural Resistance of Plants to Pests - Roles of allelochemicals (eds M.B. Green
and P.A. Hedin), Am. Chern. Soc., Washington, DC, pp. 160-7.
Hamilton, J.T. (1983) Insect pests of potatoes. Agfact H8 AEI, first edition 1983.
Agdex 262/620, Dept. Agric., New South Wales, pp. 1-7.
Hanafi, A., Radcliffe, E.B. and Ragsdale, D.W. (1989) Spread and control of
potato leafroll virus in Minnesota. J. Econ. Entomol., 82, 1201-6.
Heathcote, G.D. (1972) Evaluating aphid populations on plants, in Aphid Tech-
nology with Special Reference to the Study of Aphids in the Field (ed. H.F. van
Emden), Academic Press, London, New York, NY, pp. 105-45.
Herrnstadt, c., Soares, G.G., Wilcox, E.R and Edwards, D.L. (1986) A new
strain of Bacillus thuringiensis with activity against coleopteran insects. Bio-
techno!., 4, 305-8.
Hill, S.A. and Jackson, E.A. (1984) An investigation of the reliability of ELISA as
a practical test for detecting potato leaf roll virus and potato virus Y in tubers.
Plant Pathol. (London), 33, 21-6.
Hille Ris Lambers, D. (1972) Aphids: their life cycles and their role as virus
vectors, in Viruses of Potatoes and Seed-Potato Production (ed. J.A. de Bokx),
Wageningen Cent. Agric. Publ. Doc., Netherlands, pp. 36-56.
Hodgson, W.A., Pond, D.D. and Munro, J. (1974) Diseases and Pests of Potatoes,
Canada Dep. Agr. 1492, pp. 1-69.
Horton, D. (1987) Potatoes in the Third World. The Courier, No. 101, 82-4.
Huff, F.A. (1963) Relation between leafhopper influxes and synoptic weather
conditions. J. App!. Meterol., 2, 39-43.
References 501
International Potato Center (1984) Potatoes for the Developing World: a collabora-
tive experience, Int. Potato Cent. (CIP), Lima, Peru, pp. 1-150.
International Potato Center (1988) Control of Virus and Virus-like Diseases, Annu.
Rpt, Int. Potato Cent. (CIP), Lima, Peru, pp. 79-91.
Izhevsky, S.S. (1985) Review of the parasites of the tubermoth, Phthrorimaea
operculella (Zeller) (Lepidoptera: Gelechiidae). Entomol. Rev., 64, 148-60.
Johnson, K. and Mather, B. (1988) An analysis of potato production systems and
identification of IPM research needs in the North Central Region. Rept. NC-166
Potato 1PM Res. Priorities Subcomm., NCS-3 Proj., Mar., 1988, pp. 1-39.
Johnson, K.B., Radcliffe, E.B. and Teng, P.S. (1986) Effects of interacting
populations of Alternaria solani, Verticillium dahliae, and the potato leafhopper,
(Empoasca fabae) on potato yield. Phytopath., 76, 1046-52.
Johnson, K.B., Teng, P.S. and Radcliffe, E.B. (1987) Analysis of potato foliage
losses caused by interacting infestations of early blight, Verticillium wilt, and
potato leafhopper; and the relationship to yield. Z. Pjlanzenkr. Pjlanzenschutz,
94,22-33.
Kabrick, L.R. and Backus, E.A. (1990) Salivary deposits and plant damage
associated with specific probing behaviors of the potato leafhopper, Empoasca
fabae, on alfalfa stems. Entomol. Exp. Appl., 56, 287-304.
Kennedy, J.S., Day, M.F. and Eastop, V.F.A. (1962) A Conspectus of Aphids as
Vectors of Plant Viruses, Commonw. Inst. Entomol., London, UK, pp. 69-75.
Kibatta, G.N. (1982) Important insect pests of the potato, in Potato Seed
Production for Tropical Africa (eds S. Nganga and F. Shideler), Int. Potato
Cent. (CIP), Lima, Peru, pp. 70-3.
King, A.B.S. and Saunders, J.L. (1984) Las plagas invertebradas de cultivos
alimenticios anuales en America Central. Trop. Dev. Res. Inst. (TDRI),
Overseas Dev. Admin. (aDA), London, UK, pp. 1-182.
Knutson, K.W. and Bishop G.W. (1964) Potato leaf roll virus: effect of date of
inoculation on percent infection and symptom expression. Am. Potato J., 41,
227-38.
Kolbe, W. (1970) Influence of direct feeding damage on yields of heavily aphid-
infested potato crops. Pjlanzenschutz-Nachr., 23, 273-82.
Kuhn, R. and Low, 1. (1955) Resistance factors against Leptinotarsa decemlineata
Say, isolated from the leaves of wild Solanum species, in Origins of Resistance to
Toxic Agents (eds M.G. Sevag, R.D. Reid and O.E. Reynolds), Academic
Press, New York, NY, pp. 122-32.
Ladd, T.L., Jr and Rawlins, W.A. (1965) The effects of the feeding of the potato
leafhopper on photosynthesis and respiration in the potato plant. J. Econ.
Entomol., 58, 623-8.
Lal, L. (1987) Studies on natural repellents against potato tuber moth, Phthorimaea
operculella (Zeller) in country stores. Potato Res., 30, 329-34.
Larson, L.L. Kenaga, E.E. and Morgan, R.W. (1985) Commercial and Experi-
mental Organic Insecticides (1985 Revision), Entomol. Soc. Amer., pp. 1-105.
Lloyd, D.C. (1972) Some South American parasites of the potato tuber moth
Phthorimaea operculella (Zeller) and remarks on those in other continents.
Commonw. 1nst. BioI. Control Tech. Bull., 15, pp. 35-49.
Mackauer, M. and Way, M.J. (1976) Myzus persicae Sulz. an aphid of world
importance, in Int. BioI. Programme 9. Studies in Biological Control (ed. V.L.
Delucchi), Cambridge Univ. Press, London, UK, pp. 51-119.
502 Insect pests
McNew, G.L. (1963) Pest control in relation to human society, in New Develop-
ments and Problems in the Use of Pesticides. Proc. Symp. 12th Ann. Meeting
Liaison Panel Food Prot. Comm. Publ. 1082, Nat. Acad. Sci.-Nat. Res. Coun.
pp. 1-22.
Mehlenhacher, S.A., Plaisted, R.L. and Tingey, W.M. (1983) Inheritance of
glandular trichomes in crosses with S. berthaultii. Am. Potato 1., 60, 699-708.
Metcalf, R.L. (1980) Changing role of insecticides in crop protection. Ann. Rev.
Entomol., 25, 219-56.
Midmore, D.J. (1986) Respuesta de la planta de papa (Solanum spp.) al dano de
insectos: algunos efectos de compensaci6n. Memorias del curso sobre 'Control
Integrado de la Papa' (ed. L. Valencia), Int. Potato Cent. (CIP)-Inst. Colom-
biano Agropecuario (ICA), Bogota, Colombia, pp. 176-200.
Miller, R.J. and Griffitts Clifford, D. (1985) The National IPM Program, in CIPM
Integrated Pest Management on Major Agricultural Systems (eds R.E. Frisbie
and P.L. Adkisson), Texas Agric. Exp. Stn, MP-1616, pp. 729-39.
Misra, S.S. and Agrawal, O. (1987) Potato aphids: a review of the species, their
identification, importance, control and pesticide residues in potatoes in India.
Trop. Pest Manage., 33(1), 39-43.
Misra, S.S. and Agrawal, O. (1988) Potato pests in India and their control. Trop.
Pest Manage., 34(2),199-209.
Mitchell, B.L. (1978) The biological control of potato tuber moth Phthorimaea
operculella (Zeller) in Rhodesia. Rhodesia Agric. 1., 75, 55-58.
Miyazaki, M. (1987) Forms and morphs of aphids, in Aphids: Their biology, natural
enemies, and control (World Crop Pests; 2A) (eds A.K. Minks and P.
Harrewijn), Elsevier, Amsterdam, The Netherlands, pp. 27-50.
Moore, J.E. (1983) Control of tomato Ie.afminer (Scrobipalpula absoluta), in
Bolivia. Trop. Pest Manage., 29, 231-8.
Munro, J.A. (1954) Entomology problems in Bolivia. FAG Plant Prot. Bull., 2(7),
97-101.
Munro, J.A. (1968) Insects affecting potatoes in Bolivia. 1. Econ. EntomoL, 61(3),
882.
Murphy, S.T. (1984) Final report on surveys in the NeotropicallNearctic region for
natural enemies of Liriomyza trifolii and L. sativae (Diptera, Agromyzidae),
Commonw. Inst. Bioi. Control (FAOICIBC) Contract no: TCP/SEN/2202-1
AGOA, pp. 1-28.
Nesbitt, B.F., Beevor, P.S., Cork, A., Hall, D.R., Murillo, R.M. and Leal, H.R.
(1985) Identification of components of the female sex pheromone of potato
tuber moth, Scrobipalpopsis solanivora. Entomol. Exp. Appl., 38, 81-5.
Parry, R.H. (1987) Aphid and virus management in potatoes in Eastern Canada, in
Potato Pest Management in Canada (eds G. Boiteau, R.P. Singh and R.H.
Parry), Proc. Symp. on Improving Potato Pest Prot. Fredericton, N.B.,
Canada, pp. 9-22.
Persoons, c.J., Voerman, S., Verwiel, P.E.J., Ritter, F.J., Nooyen, W.J. and
Minks, A.K. Sex pheromone of the potato tuberworm moth, Phthorimaea
operculella: isolation, identification and field evaluation. Entomol. Exp. Appl. ,
20, 289-300.
Pimentel, D., Terhune, E.C., Dritschillo, W., Gallahan, D., Kinner, N., Nafus,
D., Peterson, R., Zareh, N., Misiti, J. and Haber-Schaim, O. (1977) Pesticides,
insects, in foods and cosmetic standards. BioScience, 27, 178-85.
References 503
Pimentel, D., Krummel, J., Gallahan, D., Hough, J., Merrill, A., Schreiner, 1.,
Vittum, P., Koziol, F., Back, E., Yen, D. and Fiance, S. (1979) A cost-bene.fit
analysis of pesticide use in U.S. food production, in Pesticides: Contemporary
roles in agriculture, health and the environment (eds T.J. Sheets and D.
Pimentel), Humana Press, Clifton, NJ, pp. 97-149.
Potato Assocation of America (1980) Commercial Potato Production in North
America (eds R.E. Thornton and J.B. Sieczka); Am. Potato f. Suppl., 57, 1-36.
Potts, M.J. (1988) The influence of intercropping on pests and diseases of potato
with special references to their control. Asian Potato Assoc. Proc., Kunming,
China, 12-26 June 1988, pp. 116-19.
Povolny, D. (1973) Scrobipalpopsis solanivora sp. n.-a new pest of potato
(Solanum tuberosum) from Central America. Acta. Univ. Agric. Bmo. Fae
Agron., 21, 133-46.
Pradhan, R.B. (1987) Control of potato tuber moth by weeds.. Rep. Nat. Potato
Dev. Prog. (His Majesties Govt./Swiss Assoc. Tech. Assist.), Lalitpur, Nepal,
pp. 1-18.
Radcliffe, E.B. (1982) Insect pests of potato. Ann. Rev. Entomol., 27,173-204.
Radcliffe, E.B., Cancelado, R.E. and Cranshaw, W.S. (1981a) Estabishing action
thresholds for insect pests, in Integrated Plant Protection for Agricultural Crops
and Forest Trees Vol. II (ed. T. Kommedahl), Proc. Symp. IXth Int. Congr.
Plant Prot. Washington, DC, 5-11 August 1979, pp. 477-80.
Radcliffe, E.B., Lauer, F.I., Lee, M. and Robinson, D.P. (1981b) Evaluation of
the United States potato collection for resistance to green peach aphid and
potato aphid. Univ. of Minn. Agric. Exp. Stn. Tech. Bull., 331, pp. 1-41.
Radcliffe, E.B., Flanders, K.L., Ragsdale, D.W. and Noetzel, D.M. (1991) Potato
insects-pest management systems for potato insects, in CRC Handbook of Pest
Management in Agriculture, Vol III, 2nd edn (ed. D. Pimentel), CRC Press
Inc., Boca Raton, FL, pp. 587-621.
Raman, K.V. (1982) Transmission of potato viruses by aphids. Tech. Inf. Bull. 2,
3rd edn (rev.), Int. Potato Cent. (CIP), Lima, Peru, pp. 1-23.
Raman, K.V. (1987) Survey of diseases and pests in Africa: Pests. Acta Horti-
culturae, 213, 143-50.
Raman, K.V. (1988a) Control of potato tuber moth Phthorimaea operculella with
sex pheromones in Peru. Agric. Ecosys. Environ., 21, 85-99.
Raman, K.V. (1988b) Integrated insect pest management for potatoes in develop~
ing countries, CIP Circular,16, No.1, pp. 1-8. Int. Potato Cent. (CIP), Lima,
Peru.
Raman, K.V. (1989) Host plant resistance of potato to arthropod and mite pests.
Proc. II th Int. Congr. Plant Prot., 5-9 October 1987, Manila, The Philippines
(in press).
Raman, K.V. and Alcazar, J. (1988) Biological control of potato tuber moth,
Phthorimaea operculella (Zeller) using a granulosis virus in Peru. Asian Potato
Assoc. Proc. Second Triennial Conf. Kunming, China, 12-26 June 1988, pp. 173-4.
Raman, K.V. and Booth, R.M. (1983) Evaluation of technology for integrated
control of potato tuber moth in field and storage. Tech. Eva!. Ser. 10. Int.
Potato Cent. (CIP), Lima, Peru, pp. 1-18.
Raman, K.V. and Redolfi, I. (1984) Progress on biological control of major potato
pests, in Report of XXll Planning Conference on Integrated Pest Management,
4-8 June 1984, Int. Potato Cent. (CIP), Lima, Peru, pp. 199-208.
504 Insect pests
Raman, K.V., Booth, R.H. and Palacios, M. (1987) Control of potato tuber
moth, Phthorimaea operculella (Zeller) in rustic potato stores. Trop. Sci., 27,
175-94.
Raymundo, S.A. and Alcazar, J. (1983) Effects of polyculture (mix cropping) on
the incidence and severity of potato pests and diseases, in Research for the
Potato in the Year 2000 (ed. W,J. Hooker), CIP, Lima, Peru, pp. 159-60.
Reed, E.M. and Springett, B.P. (1971) Large-scale field testing of a granulosis
virus for the control of the potato moth (Phthorimaea operculella (Zeller)
(Lep., Gelechiidae». Bull. Entomol. Res., 61, pp. 223-33.
Ribbands, C.R. (1965) The significance of apterous aphids in the sprcad of viruses.
Proc. XXIlth. Int. Congo Ent., London, UK, 1964, pp. 525-6.
Roberts, D.W., LeBrun, R.A. and Semel, M. (1981) Control of the Colorado
potato beetle with fungi, in Advances in Potato Pest Management (eds J.H.
Lashomb and R. Casagrande), Hutchinson Ross Publ. Co., Stroudsberg, Penn.
pp. 119-37.
Rodriguez, c.L. and Lepiz, C.S. (1988) Manejo adecuado de las feromonas de la
polilla de la papa. Bol. Divulgatorio 90, Min. Agric. Ganaderja, Dpto.
Entomol., San Jose, Costa Rica, pp. 1-13.
Roelofs, W.L., Kochansky, J.P., Carde, R.T., Kennedy, G.G., Henrick, C.A.,
Labovitz, J.N. and Corbin, V.L. (1975). Sex pheromone of the potato tuberworm
moth, Phthorimaea operculella. Life Sci., 17,699-706.
Rojas, D.A.M. (1978) Evaluaci6n de plagas insectiles en variedades comerciales de
papa. Tesis Ing. Agronomo, Univ. Nac. Cent. Peru, Huancayo, Peru, pp. 1-51.
Sanchez, G. and Aquino, C. (1986) La polilla de la papa, Symmetrischema
plaesiosema (Turner, 1919). Bol. Tecnico, 5(1), Lima, Peru: Inst. Nac. Invest.
Promocion Agropecuaria (INIPA), pp. 1-21.
Sankaran, T. and Greathead, D.J. (1980) The current status of biological control of
the potato tuber moth. Biocontrol News and Inf., 1, 207-1 t..
Saxena, K.N., Ghandi, J.R. and Saxena, R.c. (1974) Patterns of relationship
between certain leafhoppers and plants.!' Responses to plants. Entomol. Exp.
Appl., 17,303-18.
Schmutterer, H. (1987) Fecundity-reducing and sterilizing effe.cts of neem seed
kernel extracts in the Colorado potato beetle, Leptinotarsa decemlineata in
Natural Pesticides from the Neem Tree and Other Tropical Plants (eds H.
Schmutterer and K.R.S. Ascher), Proc. 3rd. Int. Neem Conf., Nairobi, Kenya,
10--15 July 1986, pp. 351-60.
Schroder, R.F.W. and Athanas, M.M. (1985) Review of research on Edovum
puttleri Grissell, egg parasite of the Colorado potato beetle, in Proceedings of
the Symposium on the Colorado Potato Beetle, XVIlth International Congress of
Entomology (eds D.N, Ferro and R.H. Voss), Mass. Agric. Expt. Stn Res.
Bull. 704, pp. 29-32.
Shands, W.A. and Landis, B.J. (1964) Potato Insects: Their biology and biological
and cultural control, US Dep. Agric. Res. Servo Agric. Handb. 264, pp. 1-61.
Shands, W.A., Simpson, G.W., Muesebeck, C.F.W. and Wave, H.E. (1965).
Parasites of potato-infesting aphids in northeastern Maine. Univ. Maine Agric.
Exp. Stn Tech. Bull. Tl9, pp. 1-77.
Shands, W.A., Simpson, G.W., Wave, H.E. and Gordon, C.C. (1972) Importance
of arthropod predators in controlling aphids on potatoes in northeastern Maine.
Life Sci. Agric. Univ. Maine Tech. Bull. 54, pp. 1--49.
References 505
Shanks, C.H., lr and Chapman, R.K. (1965) The effects of insecticides on the
behavior of the green peach aphid and its transmission of potato virus Y.
1. Econ. Entomol., 58, 79-83.
Shields, E.l., Hygnstrom, 1.R., Curwen, D., Stevenson, W.R., Wyman, 1.A. and
Binning, L.K. (1984) Pest management for potatoes in Wisconsin - A pilot
program. Am. Potato 1., 61, 508--16.
Sinden, S.L., Sanford, L.L. and Osman, S.F. (1980) Glycoalkaloids and resistance
to the Colorado potato beetle in Solanum chacoense Bitter. Am. Potato 1.,57,
331-43.
Singh, R.P. and Boiteau, G. (1987) Control of aphid borne diseases: nonpersistent
viruses, in Potato Pest Management in Canada (eds, G. Boiteau, R.P. Singh and
R.H. Parry), Proc. Symp. Improv. Potato Pest Prot. Fredericton, N.B.,
Canada, pp. 30--53.
Singh, S.P. and Verma, K.D. (1982) A note on white grubs and wireworms
infesting potatoes in Simla, in Potato in Developing Countries (eds B.B.
Nagaich, G.S. Shekhawat, P.c. Gaur and S.c. Verma), Centr. Potato Res.
Inst. (CPRI), Simla, India, pp. 379-80.
Slack, S.A. (1987) Some interesting aspects of recent and expected developments
in the control of diseases and pests in North America. Acta Hortic., 213, 161-7.
Smilowitz, Z. (1981) GPA-CAST: a computerized model for green peach aphid
management on potatoes, in Advances in Potato Pest Management (eds 1.H.
Lashomb and R. Casagrande), Hutchinson Ross Pub. Co., Stroudsburg, Penn.,
pp. 193-203.
Storch, R.H. (1981) Insects and diseases in the production of seed potatoes, in
Advances in Potato Pest Management (eds 1.M. Lashomb and R. Casagrande),
Hutchinson Ross Pub. Co., Stroudsburg, Penn. pp. 138--52.
Szmidt, A. and Wegorek, W. (1967) Populationsdynamische wirkung von Perillus
bioculatus (Fabr.) (Het. Pentatomidae) auf der kartoffelkafer. Entomophaga,
12,403-8.
Tamaki, G. (1981) Exploiting the ecological interaction of green peach aphid on
peach trees. US Dep. Agric. Tech. Bull. 1640, pp. 1-6.
Tamaki, G., Landis, B.l. and Weeks, R.E. (1967) Autumn populations of green
peach aphid on peach trees and the role of syrphid flies in their control. J. Econ.
Entomol., 60, 433--6.
Tamada, T. and Harrison, B.D. (1980a) Factors affecting the detection of potato
leaf roll virus in potato foliage by enzyme-linked immunosorbent assay. Ann.
Appl. BioI., 95, 209-19 .
. Tamada, T. and Harrison, B.D. (1980b) Application of enzyme-linked immuno-
sorbent assay to the detection of potato leaf roll virus in potato tubers. Ann.
Appl. BioI., 96, 67-78.
Taylor, L.R. and Palmer, 1.M.P. (1972) Aerial sampling, in Aphid Technology
with Special Reference to the Study of Aphids in the Field (ed. H.F. van Emden),
Academic Press, New York, NY, pp. 189-234.
Thompson, L.S. (1987) The control of potato flea beetles, leafhoppers, wireworms,
and white grubs, in Potato Pest Managetnent in Canada (eds G. Boiteau, R.P.
Singh and R.H Parry), Proc. Symp. Improving Potato Pest Prot. Fredericton,
N.B., Canada, pp. 99-111.
Tingey, W.M. (1984) Glycoalkaloids as pest resistance factors. Am. Potato J., 61,
157--67.
506 Insect pests
Tingey, W.M., Plaisted, R.L., Laubengayer, J.E. and Mehlenbacher, S.A. (1982)
Green peach aphid resistance by glandular trichomes in Solanum tuberosum and
S. berthaultii hybrids. Am. Potato J. 59, 241-51.
van Emden. H.F., Eastop, V.F., Hughes, R.D., and Way, M.J. (1969) The
ecology of Myzus persicae. Ann. Rev. Entomol., 14, 197-270.
von Arx, R., Goueder, J. Cheikh, M. and Ben Temine, A. (1987) Integrated
control of potato tubermoth Phthorimaea operculella (Zeller) in Tunisia. Insect
Sci. Appl., 8, 989-94.
von Arx, R., Ewell, P.T., Goueder, J. et al. (1988) Management of the Potato
Tuber Moth by Tunisian Farmers: A Report of On-Farm Monitoring and a
Socioeconomic Survey. Int. Potato Cent. (CIP) , Lima, Peru, and Inst. Nat.
Rech. Agron. Tunisie (INRAT), Tunis, Tunisia, pp. 1-30.
Voerman, S. and Rothschild, G.H.L. (1978) Synthesis of the two components of
the sex pheromone systef~ of the potato tuberworm moth, Phthorimaea
operculella (Zeller) (Lepidoptera: Gelechiidae) and field experience with them.
J. Chem. Ecol., 4, 531-42.
Wright, R.J. (1984) Evaluation of crop rotation for control of Colorado potato
beetles (Coleoptera: Chrysomelidae) in commercial potato fields on Long
Island. J. Econ. Entomol., 77, 1254-9.
Wright, R.J., Dimock, M.B., Tingey, W.M. and Plaisted, R.L. (1985) Colorado
potato beetle (Coleoptera: Chrysomelidae). Expression of resistance in Solanum
berthaultii and interspecific potato hybrids. J. Econ. Entomol. , 78,
576--82.
Yabar, E.L. (1988a) Integraci6n de pn'icticas culturales para el control del Gorgojo
de los Andes (Premnotrypes spp.). Rev. Latinoamericana de la Papa, 1, 120-31.
Yabar, E.L. (1988b) La mosca minadora de la papa en el Peru. Informe Especial,
Sector Agrario, Inst. Nac. Invest. Agraria Agroindustrial (INIAA) , Lima, Peru,
1(2), 1-37.
CHAPTER 12
Tuber quality
R.M.]. Storey and H. V. Davies
12.1 INTRODUCTION
The potato is one of the world's major staple food crops and produces
more dry matter and protein per hectare than the major cereal crops
(Burton, 1989). The main production areas are in Europe and the USSR,
which account for nearly 56% of the world acreage and 58% of output.
Poland has the greatest production in Europe (1981/82: 33.6 million
tonnes) and although a large proportion is used for stockfeed and industrial
starch, consumption is estimated to be 121 kg head- 1 year- 1 (Horton and
Fano, 1985). In western Europe, Great Britain, with an annual per capita
consumption in 1988 of 115 kg, is the only country with increasing
consumption (Potato Marketing Board, 1989). In contrast, North America
accounts for about 3% of the world acreage but 6% of production, most
being consumed as processed product. China produces 55.8 million tonnes
from 5.65 million hectares and the International Potato Center (CIP) has
programmes to improve potato production in the sub-tropics and tropics.
In countries with developed economies, there is an increasingly sophisti-
cated market for fresh potatoes, e.g. prepacked potatoes and 'bakers',
and for the increasing amounts of processed potato products such as
chips (crisps in the United Kingdom) and French fries which are available.
The consumers' requirements for these markets are often associated with
the aesthetic qualities of the potato and may have a marked effect on their
acceptability as food. These include, for example, the appearance of the
tuber and its freedom from defects and disorders including damage and
bruising and diseases. Some of the diseases may give superficial surface
blemishes but are still considered to detract from the quality of the potato.
The structural and biochemical factors which contribute to the culinary
quality and suitability for processing purposes are also considered to be
very important in this respect: for example, those affecting the freedom
from discolouration after cooking and influencing the texture of the cooked
potato or product. In contrast, in other countries with less well developed
508 Tuber quality
market economies, the nutritional value of the potato has been recognized
as a major criterion. This aspect of quality has been the impetus for the
major development programmes by ClP and for breeding projects to
improve the protein quality and content of the potato.
All elements which contribute to potato quality, be they nutritional or
aesthetic, are influenced by various factors related to the morphology,
structure and chemical composition of the tuber. All can be affected by the
environment during growth and development and by storage conditions
(see Chapter 14). As a result, it is often difficult to produce, consistently,
potatoes for commerce of the required quality. This chapter therefore
considers not only the morphological and biochemical differences which
are the basis for quality but also the part played by genotype, cultural
methods and the environment. Nutritional quality is considered initially
and subsequent sections examine other quality aspects associated with the
fresh, and processed potato.
12.1.1 Nutritional value

The potato provides significant amounts of protein, vitamin C, carbo-
hydrate and iron. The carbohydrates, which constitute about 75% of the
total dry matter, are the main energy source. Elton (1978) using an average
consumption figure of 240 g potato day-l (88 kg year- 1) for European
Community countries, estimated that potatoes provided about 5.3% of
daily energy requirements. As the energy value of 290 kJ per 100 g is only
half that of rice or pasta and about a quarter of that of bread; this means
that a considerable amount (about 4.5 kg of cooked potatoes) would have
to be eaten to meet total daily energy requirements. However, the potato's
contribution to the diet will vary depending upon the cooking method, and
when cooked with oil or fat, potatoes have a secondary function and act as
a carrier for absorbed and surface fat (Table 12.1).
The potato is also an important source of dietary fibre and in Great
Table 12.1 Proximate composition of raw and cooked maincrop potatoes (g per
100 g) (from Finglas and Faulks, 1984)
Dry Protein Non-protein Starch Sugars Fat Dietary
matter (N x 6.25) nitrogen fibre
Uncooked* 21.2 2.1 0.17 17.3 1.3 0.2 1.7
Boiledt 19.9 1.8 0.14 16.7 1.2 0.1 1.6
Bakedt
(without skin) 21.1 2.2 0.18 16.9 1.5 0.2 1.9
Roastt 34.1 2.8 0.23 25.1 1.7 4.3 2.7
Chippedt 44.0 4.6 0.29 30.5 1.7 6.8 3.3
* Mean value for four main crop varieties purchased quarterly (1981/82 season) at three
centres.
t For cooked potatoes mean values per 100 g as consumed.
Introduction 509
Britain it contributes about 15% of the intake (Finglas and Faulks, 1984).
This factor and the insignificant fat content when boiled or baked has
meant that potatoes have been identified as an important dietary item in
the NACNE and COMA reports (Anon., 1983, 1984) to the United
Kingdom (UK) government. The consumers' perception of the potato's
importance in a balanced diet is also increasing (Potato Marketing Board,
1988).
12.1.2 Protein
Recent estimates of the protein content of UK potatoes were 1.6 and 2.1 g
per 100 g fresh weight (FW) for new season and maincrop potatoes,
respectively (Finglas and Faulks, 1984). Although potatoes are not usually
regarded as a protein source because of the low protein content on a fresh
weight basis compared to other foods, they can make a significant
nutritional contribution to the diet in countries with a high per capita
potato consumption (Woolfe, 1986a). In Great Britain in 1983, potatoes
were estimated to contribute 3.4% of total protein intake, compared with
4.6%, 4.8% and 5.8% contributed by eggs, fish and cheese (Woolfe,
1986b). Moreover, the high quality dietary nitrogen in potatoes means that
a small quantity (100 g) of boiled potato supplies 8-13% of the FAO-
WHO recommended daily allowance (RDA) of protein for children and
6-7% of the adult RDA (Horton and Sawyer, 1985). Amino acid analyses
have shown that potatoes are a very good source of lysine. However,
they have lower concentrations of the sulphur-containing amino acids,
methionine and eystine (Woolfe, 1986b). In mixed diets, an important role
for potatoes is supplementing foods low in lysine, e.g. rice or pasta.
The nutritive value of potato protein varies between lots of the same
variety and is affected by preparation, cooking and storage (Chick and
Slack, 1951; Finglas and Faulks, 1984).
The soluble and insoluble protein fractions are two components of the
total N content of the tuber. The remainder of the nitrogen content is
present as non-protein N, which contains both organic and inorganic
nitrogen. The organic nitrogen fraction contains free amino acids and the
ami des glutamine and asparagine.
Total N content ranges, in general, from 0.11 to 0.58 g per 100 g fresh
weight (from Gray and Hughes, 1978); of this the protein-N component
has been estimated at between 51 and 63% (Pol and Labib, 1963) and
between 38 and 64% (Burton, 1966). The soluble protein fraction aceounts
for 28-51 % of the total N and is fairly evenly distributed throughout the
tuber, whereas another 8-10% of the total N fraction is insoluble protein
and is concentrated largely in the skin (Neuberger and Sanger, 1942; Burton,
1966). Munshi and Mondy (1989) reported lower concentrations of protein
(and non-protein N) in the pith region than in the cortex. However, as the
pith comprises the greater part of the potato it contains 71 % of total tuber
protein. Peeling removes 2% of the total protein content.
510 Tuber quality
The protein content can vary for different cultivars and between crops of
the same variety depending on location and fertilizer usage (Mulder and
Bakema, 1956; Talley et aI., 1961, 1970; Pol and Labib, 1963; Desborough
and Weiser, 1974; Eppendorfer et aL, 1979). Differences which occur have
been related to tuber dry matter (DM) content (Talley et aI., 1961; Li and
Sayre, 1975), N availability (Talley, 1983) and crop maturity (Mazza et al.,
1983). Rosenstock and Zimmerman (1976) observed that young tubers of
cv. Saskia had twice the protein content of older tubers (>3 cm diameter).
Early studies (Mulder and Bakema, 1956) identified two main com-
ponents of the soluble protein fraction as globulin (tuberin) 70%, and
albumin (tuberinin) 30%. These two fractions had similar amino acid
composition but albumin was distinguished by its lower isoelectric point
and more hydrophilic nature (Groot et aI., 1947). Kapoor et al. (1975) also
identified albumin (49% of total protein-N) and globulin (26%) as the
main protein fractions and also smaller amounts of glutelin (8.8%) and
prolamin (4.3%). However, these fractions are not chemically distinctive
and subsequent studies have revealed up to 40 proteins (Stegmann, 1975).
A major soluble glycoprotein was isolated by Racusen and Foote (1980)
and named patatin. Park (1983) found it could be up to 40% of the total
soluble protein and it has been characterized as an esterase enzyme
complex. The soluble proteins have characteristic electrophoretic patterns
which are genotype specific and they have been used in conjunction with
isozyme patterns for routine cultivar identification (Stegemann and Schnick,
1982), detection of differences between species, and as biochemical gene
markers in hybrid progenies (see Desborough, 1983). Because of the
importance of potato protein, several breeding programmes have been
developed and several hybrids have been shown to have improved protein
quantity and quality.
The free amino acid pool contains 40--60% of the total N. However, this
nitrogen contribution to the diet is proportionally less important than the
essential amino acids contributed by the potato proteins (Desborough,
1985). The factors which affect protein levels also influence the free amino
acids and the amides glutamine and asparagine (Talley et al., 1970, 1984;
Davies, 1977). These two amides account for about half of the soluble non-
protein N (Neuberger and Sanger, 1942), and are generally increased by
increasing N fertilizer (Mulder and Bakema, 1956; Hippe, 1988). In three
cultivars examined by Mazza et al. (1983), non-protein N was between 25
and 40% of total N, with the lowest level in Russet Burbank, the cultivar
with the highest DM content. Li and Sayre (1975) similarly found that clones
with a high DM content contained lower proportions of non-protein N.
12.1.3 Vitamins
Vitamin C is the main vitamin in potatoes and Burton (1974) suggested
that they were the most important single source in the UK diet and possibly
Introduction 511
supplied over 30% of the intake. A lower figure (19.4%) was estimated by
Finglas and Faulks (1984). In freshly harvested tubers the usual range is
15-25 mg per 100 g FW. It is present in both the reduced state (ascorbic
acid) which predominates in the tuber (c. 85-100%) and the oxidized state
(dehydroascorbic acid) the two forms being readily interchangeable. The
concentration of vitamin e increases during growth and development of
the tuber (Lampitt et at., 1945a; Mazza et at., 1983) but immature potatoes
may have higher levels (Shekhar et at., 1978) than mature ones. The
vitamin e concentration is not influenced by tuber DM content or size
(Noonan et at., 1951) and Lampitt et at. (1945a) found it evenly distributed
throughout the tuber. However, other early reports (Rolf, 1940; Smith and
Gillies, 1940) suggested a higher concentration in the skin than the cortex
and at the bud end of the tuber. Shekhar et at. (1978) and Klein et at.
(1980) confirmed higher concentrations at the bud end, but the latter
authors and Munshi and Mondy (1989) found the concentration to be
slightly higher in the pith (25-29 mg per 100 g) than in the outer cortex (22-
26 mg per 100 g).
Differences in vitamin e concentration between cultivars have been
recorded (Lampitt et at., 1945b; Shekhar et at., 1978; Finglas and Faulks,
1984) and conflicting results are reported on the effect of N application;
Augustin (1975) with cv. Russet Burbank found a fall with increasing N,
whereas Mondy et at. (1979) found the reverse. With the former author,
tuber ascorbic acid was influenced by soil type with higher levels in crops
from a sandy soil than from a loam. The reported effects of these factors on
vitamin e in the harvested crop however are small compared to the length
and temperature of storage (Barker and Mapson, 1950; Augustin et at.,
1978) and cooking or processing of the tubers (see Burton, 1989).
During storage there appears to be a sharp initial decline in vitamin e
levels (Augustin et at., 1978; Keijbets, 1984) with those tubers having a
high concentration, for example some immature tubers, showing the most
marked decrease. Such losses amount to 33% over 3-4 weeks at lO o e for
immature King Edward, compared to 10% loss for mature crops (Burton,
1989). This initial decrease is reflected in ascorbate levels found in retail
potatoes in the UK (Finglas and Faulks, 1984). Thereafter there is a period
of little change, but this appears to depend in part on cultivar and storage
temperature. Augustin et at. (1978) found little difference for five cultivars
held between 4 and 8 months at 7.2°e, whereas Keijbets (1984) stored five
different cultivars at 5-6°e and found only a slight decrease in ascorbic acid
between December and March, but in the period to July the fall was much
more rapid. Other authors have reported a gradual decline in ascorbic acid
content throughout storage (Mazza et at., 1983). Barker and Mapson
(1950) also found less ascorbic acid in tubers stored at low than high
temperatures.
During preparation for cooking, ascorbic acid may be oxidized on the
cut surface of the tubers, and although this does not markedly affect the
512 Tuber quality
level of antiascorbutic activity it can lead to higher vitaminC losses during
cooking as the dehydroascorbic acid is converted to diketogulonic acid
which represents a net loss of vitamin C. Vitamin C losses which occur
during cooking and processing can be considerable and are dealt with in
detail by Leichsenring (1957a, b) and Burton (1989). Burton (1989)
estimated average percentage losses for various methods of preparation:
unpeeled-steamed, 10-15%; boiled and baked, 20%; peeled-steamed,
10-30%; pressure cooked, 15-25%; boiled, 25%; microwaved, 25%; roast,
20-45%; chipped, 35-50%; reconstituted instant powder or flake, 70%.
Although the losses from processing can be large, addition of ascorbic
acid by manufacturers can give high vitamin C contents in the final product
(Paul and Southgate, 1978).
The other main vitamins in the tuber are those of the B group (Table
12.2). Although they occur at much lower concentrations than vitamin C
they can make a significant contribution to the daily requirement (Burton,
1989). Levels are reported to be influenced by a range of cultural
treatments (Augustin, 1975; Augustin et al., 1978) and they are reduced
during cooking (Finglas and Faulks, 1984).
Table 12.2 Vitamin composition of raW potatoes (mg per 100 g FW)
Reference Ascorbic Thiamin Riboflavin Niacin Total Pyridoxin
acid BJ Bz Bz folate (J.lg) B6
8*-30t 0.11 0.04 1.2 14 0.25
2 8-30 0.03-0.15 0.01-0.05 0.5-3.1 5-10* 0.13-0.42
3 21*-36t 0.12-0.14 0.06-0.07 2.2-2.5 19-24 0.26-0.39
4 1-54 0.02-0.18 0.01-0.02 0.6-2.0* 0.19
5 16*-19t 0.2*,t 0.02*'t 0.4*-O.6t 25*-35t
1 Paul and Southgate (1978), * stored 9 months, t freshly harvested.
2 Augustin (1975) (* Folic acid).
3 Augustin et al. (1978), * stored, t freshl~· Irlarvested.
4 Adler (1971) (* nicotinamide).
5 ·Finglas and Faulks (1984), * first early, t maincrop.
12.2 QUALITY ASSOCIATED WITH THE FRESH TUBER
The morphological characteristics of the tuber, such as its size and shape or
skin colour, its defects, for example greening or growth cracking and
damage and the incidence of diseases and internal disorders all contribute,
to a greater or lesser extent, in determining the initial acceptability of the
fresh potato to the consumer and its suitability for the processor. The
external characteristics and internal disorders are examined in this section
but mechanical damage and bruising are considered separately as these, in
part, depend upon the structural characteristics and biochemical properties
of the tuber. Tuber diseases are dealt with in detail elsewhere with
Quality associated with the fresh tuber 513
particular emphasis on surface blemishing diseases, such as silver scurf,
which are a major concern for those producing crops for the washed
prepacked and baker markets (see Chapter 10). Many of the factors which
have a bearing on these elements of quality are interrelated and account
needs to be taken of varietal differences and the cultural and environ-
mental factors which affect plant growth and tuber development.
12.2.1 Morphological characteristics
(a) Tuber size and shape

The size of tubers required by consumers varies considerably and will also
depend upon the acceptable level of peeling loss as this is proportionally
greater for small than large tubers. With tubers of 200 g a peeling loss of
23% was found; this decreased to 16% for 500 g tubers (Weaver et al.,
1979). However, when small new potatoes are scraped, rather than peeled
prior to cooking, weight loss is reduced to 4-5% (Burton, 1989). In
selecting potatoes for different cooking methods size was considered more
important than cultivar for the domestic consumer (Potato Marketing
Board, 1988). Although there were indications that a range of sizes was
desirable, a large tuber was preferred for chips and jacket potatoes and a
small size for new potatoes. Generally, for new potatoes those sized 35-55
mm are used; jacket potatoes are usually sized 65-80 mm, although
because of price premiums available selection on an individual tuber
weight basis has been introduced with bands of 175-225 g, 225-275 g, 275-
325 g and 325-400 g widely adopted. Interest in developing more general
weight grading schemes has also increased because better electronic
systems offer the opportunity for high speed weighing of individual tubers.
Glasbey et al. (1988) have suggested parallel weight grading and size
criteria following analysis of weight, length, breadth and depth of the ten
most commonly grown varieties in the UK.
Potatoes used by the processing sector have stringent size and shape
requirements and these are usually associated with particular varieties
suitable for the different products (Kirkman, 1986). Tuber size is deter-
mined in part by variety and the number of tubers set per stem, but a large
measure of control over size is achieved in commerce by varying plant
population and crop husbandry (see Chapter 7).
For most commercial purposes, in addition to an even-sized sample, a
uniform tuber shape is preferred, although the desired shape may vary
depending upon the end use or regional preference (Cowan, 1985). Deep
eyes, protuberances on the tuber, secondary growth defects and growth
cracks reduce the acceptability of the potato (see Section 12.2.2). Tuber
shape is largely a varietal characteristic and Burton (1989, referring to
Huaman et al., 1977) describes eight shapes: compressed, round, ovate,
obovate, elliptic, oval, oblong and long. A number of less usual tuber
514 Tuber quality
shapes also occur. However, in developed countries the shapes of varieties
in present commercial use are very limited in comparison to the diversity
found in Andean cultivars. In general, round-oval are preferred for
prepacking, oval to long-oval for baking, long-oblong for French fries and
round for canned new potatoes and chipping (crisping). The preferred
shape can markedly improve out-turn. For example with French fries, use
of long-oval Pentland Dell results in only 10% loss from short strips «5
cm) after slicing, whereas with a short-oval variety, Pentland Squire, losses
are about 17% (Harkett, pers. comm.). Shape is also influenced in some
cultivars by cultural and environmental conditions but more in those with
long rather than round tubers (Reust and Munster, 1978).
(b) Skin texture

A clean, bright, blemish free skin is required for the most demanding
outlets of washed prepacks and bakers as potatoes are selected initially by
consumers on the appearance of the sample (Potato Marketing Board,
1983). Increasing importance is being attached to the reduction of mech-
anical damage, bruising and the control of field and storage diseases
which cause surface blemishing diseases, e.g. common scab or silver
scurf.
The periderm thickness is partially dependent on cultivar and skin
texture and can be influenced by soil type, growing conditions and crop
maturity (Vander Zaag, 1970). More smooth skinned samples occur in
seasons with high rainfall and increased netting was found in cultivars
Wilja and Estima with late harvests (Kerr, 1986). With high soil tempera-
tures the number of cell layers and periderm thickness increase (Cutter,
1978) and more russetting occurs (Yamaguchi et aI., 1964).
For new season early potatoes an immature skin is necessary for
marketing purposes but immaturity is a disadvantage later in the season
because it increases susceptibility to scuffing at harvest and potential for
increased weight loss in store.
(c) Flesh and skin colour

A preference for a particular flesh colour in the raw potato is often
expressed by consumers but, irrespective of the household cooking method
adopted, it is usually of secondary importance to the flavour and cooking
quality of the variety (Potato Marketing Board, 1988). In Britain, although
the consumers' stated preference is for white-cream flesh, a number of
varieties with yellow flesh (e.g. Desiree and Estima) are widely accepted
and in total yellow fleshed varieties (including those for processing)
accounted for about 45% of the area planted in GB in 1988. This stated
preference is in contrast to most of continental Europe where, with
perhaps the exception of Scandinavia, the preferred flesh colour is yellow.
For certain processing purposes a yellow flesh colour is required and
cultivar Record is grown for chip (crisp) manufacture in the UK as, when
processed, it produces an attractive, bright coloured product.
Carotenoids
The degree of yellow colouration in the flesh and skin is due to the
concentration of carotenoids in the tuber (Iwanzik et al., 1983) and is a
heritable characteristic (Howard, 1978). The carotenoids are loosely
associated with cell membrane lipids. The mean whole tuber contents
range from 14 to 343 mg per 100 g fresh weight and violaxanthin (38%),
lutein (37.7%) and lutein 5, 6-epoxide (22.2%) are the main constituents.
These are not distributed uniformly throughout the tuber and in cv. Assia
higher concentrations of total carotenoids are found in the skin (499 mg per
100 g) than in the outer storage parenchyma (323 mg per 100 g) or the rest
of the tuber (255 mg per 100 g) (Iwanzik et al., 1983).
Anthocyanins
Where there is a national or regional preference for potatoes with either
fully or parti-coloured skins, a range of cultivars are exploited for
marketing purposes. The red, blue or purple colour in potato skin depends
upon the range and concentration of anthocyanins dissolved in the cell sap
of the periderm and/or peripheral cortical cells. The most common
anthocyan ins are pelargonidin, delphinidin, cyanidin, petonidin, peunidin
and malvidin (Harborne, 1960) and the colour of their pigment depends on
the presence and number of hydroxyl and methyl groups in the molecule.
A limited number of cultivars normally have anthocyanin pigmentation
either wholly or partially distributed through their flesh e.g. Purple Congo
or Red Salad.
Flavones and flavines are also present in the tuber in small or trace
amounts but have no effect on flesh or skin colour (Lampitt and Goldenberg,
1940).
12.2.2 Tuber defects
(a) Greening
Many crops contain green tubers at harvest, often resulting in high losses
when tubers are dressed for sale. Indeed, greening may be the major cause
for rejection by processors, prepackers and merchants. In the field the
major causes of greening are either insufficient cover over the tubers at
planting (Lewis and Rowberry, 1973) or the exposure of tubers following
heavy rainfall on light soils. While greening at harvest may be a problem,
chlorophyll production in tubers stored and displayed in retail outlets is
more so. Ways of reducing or preventing the greening process have been
investigated over many years. Light intensities as low as 3-11 W m- 2 for
516 Tuber quality
periods as short as 24 h induce greening (Larsen, 1949). Light quality is
also important, studies showing that daylight-type fluorescent tubes are
most effective in inducing the development of chlorophyll (Liljemark and
Widoff, 1960). Short, repeated daily doses of light have an effect several
times greater than single exposures to much larger duration (Akeley et aI.,
1962; Brown and Riley, 1976). The temperature to which tubers are
exposed also has a profound effect on the rate of greening, lower
temperatures increasing the lag phase between exposure to light and
chlorophyll accumulation by several days (Harkett, 1975). No greening is
found at 5°C, but it is extensive at 20°C (Larsen, 1949). Tubers tend to
remain green even when stored in the dark for 30 days or more. An
absolute minimum exposure to light remains one answer to the problem;
this includes displaying prepacks outside lit cabinets. The greening process
is also affected by the prevailing gaseous environment (Forsyth and Eaves,
1968). Atmospheres containing 15% or more CO 2 are particularly effective
in preventing greening, as is the coating of tubers with surfactants such as
Tween 81 or 85. Effective concentrations of surfactant vary with genotype
(Thomas, 1984). The infiltration of tubers with Ca is also reported to
restrict the greening response (Arteca, 1982). Genotypic variation in the
extent of greening has been noted by Akeley et al. (1962), offering the
possibility of modifying the greening response through breeding pro-
grammes. However, progeny testing indicates that multiple factors are
involved in inheritance and that dominance is incomplete.
(b) Growth cracking

Although the phenomenon of growth cracking is known to be associated
with certain cultivars, the reasons for the phenomenon are not well
understood. It is generally thought to occur in response to fluctuating water
stress (Robins and Domingo, 1956; Iritani, 1981; van Loon, 1981) but such
changes do not reliably induce the disorder. It is also suggested that
cracking is associated with conditions which give rise to rapid changes in
growth rate (Gray and Hughes, 1978). However, both suggestions of
the conditions leading to growth cracking may be over-simplifications
(MacKerron and Jefferies, 1985). For example, cracking in cv. Record
occurred when there was relief of soil moisture deficits of greater than 70
mm. The amount of cracking was greater when re-wetting was greater and
when re-wetting occurred from greater soil moisture deficits (Jefferies and
MacKerron, 1987). In contrast, tubers of cv. Guardian did not crack when
severe soil moisture deficits were relieved, but did when a deficit of only 20
mm was relieved and when the soil moisture content remained close to
field capacity. Conditions which produce very rapid growth of tubers,
which mayor may not accompany the relief of water stress, can produce
the large cleavage cracks observed in cv. Guardian if growth rates are high
enough. The relief of water stress can also induce the jagged tearing cracks
observed in cv. Record where the preceding stress has been sufficient,
along with other conditions, to cause cessation of tuber growth. Fracture
cracking of tubers, a form of mechanical damage at harvest, has been
considered to be a separate phenomenon from growth cracking. However,
Schoorl and Holt (1983) concluded from compression studies that cracking
occurred when the potential energy within a tuber exceeded a critical
value. Susceptibility to mechanical damage, including fracture damage,
varies with cultivar and is influenced by environment. Lampe (1960)
showed that tubers from high rainfall areas were damaged by the lowest
rupture force. Similarly, Finney and Findlen (1967) observed that tubers
from irrigated plots damaged more easily than those from unirrigated
plots, and that high turgor increased damage susceptibility. These are the
conditions associated with growth cracking in cv. Guardian but not in cv.
Record which is resistant to mechanical damage (Grant and Hughes,
1985). If the static forces within a tuber that is turgid and expanding rapidly
can be equated with the quasi-static forces from compression tests then it
may be that fracture cracking and growth cracking of the type in cv.
Guardian are related phenomena. Growth cracking in cv. Record may be
the result of uncoordinated changes in cell sizes in the several tissues of the
tuber causing stresses in cortex and periderm. This would be consistent
with the observation that growth cracks in cv. Record are, initially at least,
shallow and extensive (Jefferies and MacKerron, 1987).
12.2.3 Internal disorders

Hiller et af. (1985) classified internal physiological disorders as either major
or minor. Major disorders are considered to be those occurring most
frequently or those which affect a large percentage of tuber production.
Major internal disorders include hollow heart, brown centre, internal rust
spot (also commonly known as internal brown spot and chocolate spot),
vascular discolouration and bruising. Minor disorders include translucent
end, jelly end, blackheart and low temperature necrosis. The following
sections deal only with the major internal physiological disorders. These
disorders cannot be ascribed to pathogens.
(a) Hollow heart

This is normally manifested by a hole of varying dimensions towards the
tuber centre. There are no external symptoms but acoustic and X-ray
detection of hollows are possible (Watts and Russell, 1985). Hollow heart
may often be preceded by the onset of brown centre, i.e. necrosis of pith
cells resulting in the weakening of tissue and the splitting apart of cells as
the tubers grow. Whether preceded by brown centre or not, the consensus
of opinion is that hollow heart is the result of tissue tension associated with
rapid tuber enlargement. Nelson and Thoreson (1986) showed that the
518 Tuber quality
frequency of hollow heart was considerably higher in large compared with
small tubers. This is not always the case, however, and the rate of tuber
growth at the time of imposition of stress may be equally important.
Several researchers have reported the development of hollow heart within
a few days of tuber initiation (Rex and Mazza, 1989 and references
therein). Certain environmental and cultural practices have been impli-
cated in the induction of hollow heart, but the literature is often conflict-
ing. However, high soil temperatures and fluctuations in water supply to
developing tubers appear to be important and there is some evidence that
modifications in nutrient supply also playa role (Kallio, 1960; Rex and
Mazza, 1989). High N applications, particularly around initiation, can
induce higher incidences of hollow heart, whereas K appears to lower the
incidence (Jackson et al., 1984). There is also evidence for an inverse
relationship between the severity of hollow heart and soil Ca status (van
der Zaag and Ffrench, 1987) although work by Silva et al. (1988) conflicts
with this finding. Factors giving rise to slow, steady growth without
excessive stress will reduce hollow heart (Watts and Russell, 1985) but, at
present, it seems impossible to guarantee absolute freedom from the
disorder through manipulations of cultural practices only. As with many
physiological disorders genotypic variation exists in the degree of suscept-
ibility (Rex and Mazza, 1989). Despite good evidence that fluctuations in
tuber growth rate are involved there is no evidence that inter-cultivar
variations in growth rates are indicative of susceptibility. Mogen and
Nelson (1986) showed that there was no difference in water potential
between pith cells from various parts of the tuber or between cultivars.
Also pith cell size was not related to hollow heart susceptibility. Detailed
analysis of mineral ion distribution within tubers during the onset of hollow
heart would appear to implicate either Mg toxicity, or Ca deficiency (due
to high Mg: Ca ratios) as causative factors (Arteca et al., 1980; Mohr et at.,
1984). Nevertheless, the physiological basis for genotypic variation in
susceptibility remains poorly understood.
(b) Brown centre

Brown centre is generally considered to be a precursor to hollow heart
although hollow heart does not always arise (Hiller et at., 1985 and
references therein). It is characterized by the appearance of small groups
of necrotic cells in the central pith region, by the depletion of starch in
surrounding cells and eventually by the production of wound cambium
around the necroses. Cool temperatures around the time of tuber initiation
are reported to induce brown centre e.g. between 10-15°C (Hiller et al.,
1979; Van Denburgh et al., 1980) and the severity is increased by high soil
moisture.
As with hollow heart, tuber growth rate plays an important role in the
initiation of brown centre and again the disorder can manifest itself in small
as well as large tubers. It also appears that some crops have a high
incidence of brown centre early in the season but show only a low incidence
at harvest. This is almost certainly due to the dissipation of dead cells as the
tuber grows and not to some reversible phenomenon. Brown centre has
been reduced by applIcations of ethephon (Hiller and Koller, 1984). Other
than growing resistant cultivars, however, avoidance of the disorder is
difficult.
(c) Internal rust spot

Internal rust spot (IRS) is characterized by rust coloured lesions, of various
size, in the medullary tissues internal to the vascular ring. Lesions are
generally more pronounced towards the apical end of the tuber. The
disorder is reported to be induced by several environmental factors
including restricted water supply and high temperatures (O'Brien and
Rich, 1976; Hooker, 1981) and by conditions unfavourable to even rates of
tuber growth. As to the basic underlying cause of IRS, there is a strong
body of evidence implicating a role for Ca. Although many soils have a
high Ca content, Ca ions are transported within plants primarily in the
xylem stream, a high phosphate concentration precluding substantial
transport within the phloem network. Water potential gradients within the
potato plant favour xylem transport into the foliage, which has a high Ca
content. Tubers, however, are likely to receive most of their water along
with sucrose in the phloem mass flow stream. Tubers therefore have a very
low Ca content (0.1-0.4 mg g-J FW), a content which may be considered
marginal for maintaining cellular integrity. Decreasing the supply of Ca to
developing tubers increases the incidence of IRS (Collier et al., 1978;
Davies and Ross, 1986a). There are also cases where soil applications of Ca
reduces the severity of the disorder (Tzeng et al., 1986).
Environmental/cultural conditions giving rise to IRS would also be
expected to modify Ca transport into and out of tubers. In hot, dry
weather, for example, tubers lose water and probably Ca to the haulms.
The absolute concentration of tuber Ca required to induce IRS is un-
certain, but it is clearly genotype-dependent (Collier et al., 1980; Davies
and Ross, 1986a; Monk and Davies, 1989). Factors other than overall
tissue Ca content must be considered and genotypic extremes in suscept-
ibility exploited to examine the relative roles of tuber water relations and
Ca regulated metabolism in modifying responses to restricted Ca supplies.
Current suggestions for controlling IRS include growing less susceptible
cultivars, delayed planting (and other treatments reducing tuber size) and
modifying cultural practices to prevent rapid fluctuations in tuber growth
rate (Hiller et al., 1985).
i
520 Tuber quality
100
Slight
80 Moderate
en Severe
~
Q) 60
0
c:
Q)
"0
·0
..5
40
<fl.
20
o
~ ~ \ ~ ~ ~ '0 '0 1"~ ~ '0 ~
~~f$>~~Q~~~<JI..-:~~
~ ~ '~ 0. ~ ~ ~ ~ ~ ~ ~ IS'
~ ~ o~'~ ~ ~ ~ ~ ~ ~ ~ '.-(\
0 '% <0 "0 "0'
<0
~':)~O'i:."''i:.'' <0 o~ "0, 0'£9""- 0
~ Of$>~Oo- OO~. 'i:. f$> ~
,... f$>~ . f$>~ f$> ~ 1!3< 0 ~"o~
" ~.,. 0- f$>-'"
9..- '" 0f$>~
Figure 12.1 Percentage incidence (slight, moderate and severe) of internal rust
spot in tubers of a range of genotypes grown either in the field or in vermulite
under low (1 mol m· 3 ) or high (10 mol m- 3 , control) calcium regimes. 15075 and
10337 de 40 are clones obtained from the breeding programme at the Scottish Crop
Research Institute (Talbot and Davies, unpublished data).
(d) Vascular discolouration

Under certain stress conditions aberrant synthesis of anthocyan ins can
occur in the flesh of varieties where it is not normally present (Cromack,
1981). Problems have arisen with cvs Record and Ulster Sceptre in Great
Britain and with cv. Roseval in France. In Canada and the USA the
occurrence appears to be commercially rare. Precursors of anthocyanins
are present in all tubers but so are inhibitors of synthesis and the purple/red
pigmentation occurs when the balance of these components is affected.
The pigmentation is largely confined to the centre of the tuber and it is
believed to result from one or two causes. It can develop under conditions
which check tuber growth and in some, but not all instances, the disorder is
associated with greening of the tubers. The disorder therefore seems to be
most prevalent when tubers have formed near the surface of the ridge or
when earthing up has been inadequate. Irrigation may reduce the number
of tubers affected, whereas temperatures less than lOoC, particularly at
night, have an adverse effect (Kirkman, pers. comm.). High levels of
organic or mineral N may also favour pigment synthesis as may several
Quality associated with internal factors 521
diseases, including leaf roll virus. More detailed studies are needed to
provide quantitative data on effects of climatic and cultural variables on
biosynthesis in the tuber. However, for a degree of control, a switch to
wider rows, deeper planting or bed systems has proved beneficial.
12.3 QUALITY ASSOCIATED WITH INTERNAL FACTORS
The internal composition of the tuber, together with the structural and
biochemical properties of the tissue, affect diverse aspects of quality in
both the raw and cooked potato. For example the dry matter content of the
tuber is associated with the susceptibility to damage and bruising and also
with the texture of the cooked potato and yield of the processed product.
The range of intrinsic factors involved with anyone of these aspects of
quality is large. Inevitably variation with and between crops occurs as
individual elements, for example the DM content, starch composition
within cells and cell size all vary to differing degrees from tissue to tissue
and tuber to tuber. This section deals with elements such as DM content
and damage and bruise susceptibility which are associated, initially, with
the raw tuber, and considers environmental, cultural and developmental
factors which affect them. Subsequent sections examine the influence of
internal factors on the cooked tubers.
12.3.1 Dry matter content

The DM content of tubers is an important measure of quality and it is used
to assess suitability for processing purposes as it influences process
efficiency, product yield and oil absorption. Tubers with a high DM
content require less energy input during frying or dehydration to remove
water; they have a greater product yield per unit fresh weight than tubers
with lower solids content, and absorb less oil during frying. The DM
content is also used to provide a measure of dryness or mealiness in boiled,
chipped and baked potatoes and an indication of tuber susceptibility to
internal bruising. However, the relationship between DM content and
these quality attributes is often imprecise and some of the effects are
indirectly associated with other factors.
The specific gravity of tubers determined from the underwater weight of
a sample, or by hydrometer readings, is extensively used as a measure of
the potato's DM content. (See Chapter 14.)
The relationship between specific gravity and percentage DM has been
reported by a number of workers (von Scheele et al., 1937; Porter et al.,
1964; Houghland, 1966; Verma et at., 1971; Schippers, 1976; van Es and
Hartmans, 1987a) but slight differences are found because evaluations
were made with different cultivars and on crops grown under a range of
environmental conditions. The varietal differences are, in part, a result of
522 Tuber quality
differences in intercellular space in the tuber (Kushman and Hayes, 1971)
and differences in the composition of the DM. Internal disorders, such as
hollow heart (Drew and Dreasy, 1941) and to a lesser extent the tempera-
ture of the tubers and water when carrying out the assessment also affect
the estimation of DM. However, correction factors can be used (Gould
and Plimpton, 1985). The specific gravity also provides an estimate of the
starch content of the tuber as starch is the major component of the DM,
usually compromising 65-75% of the total solids. The relationship between
the starch content and total DM is reasonably constant and is little affected
by environmental conditions. Consequently, as a general indicator of DM
content, specific gravity determination has much merit because of its ease
of measurement and it is widely used for determining final product quality.
The selection of suitable cultivars is probably the most usual way of
achieving the desired tuber DM for commercial purposes and levels of
between 18-26% occur for most European and North American cultivars
(Burton, 1989). The tuber DM is influenced by a wide range of factors
affecting the growth and development of the crop including, most impor-
tantly, environmental factors such as intercepted solar radiation, soil
temperatures, available soil moisture and cultural treatments.
The DM content of early maturing cultivars is reported to be usually
lower than that of later maturing varieties (Burton, 1966) but this may be
due on occasions to harvesting immature crops (Nelson et al., 1988).
Nevertheless, this does result in the supply of high DM tubers preferred for
processing being limited during the earlier part of the growing season in
temperate regions.
Genotypic control of tuber DM is modified by environmental and
cultural factors although interactions between these and variety are
considered small (Killick and Simmonds, 1974; Jefferies and MacKerron,
1986). In general, cool dull years and short growing seasons reduce DM
production and percentage DM content whilst the reverse occurs in warm
sunny years and long growing seasons.
Scott and Wilcockson (1978) showed that for potatoes the amount of
intercepted solar radiation was correlated with the total DM yield. A linear
relationship between total and tuber dry weights for a range of agronomic
treatments was demonstrated by Allen and Scott (1980). They also showed
that tuber dry weight was directly related to the intercepted radiation.
Hogge (1989) has established that the linear relationship between tuber
DM and light interception per unit area is not changed for a wide range of
individual husbandry treatments for cv. Pentland Dell at different sites.
Applied Nand K usually reduce tuber DM (see Chapter 4), but the
effects are not always consistent (Birch et al., 1967; Schippers, 1968).
However, the DM changes are usually small over the range of fertilizer
levels used to give optimum yield. Nitrogen applications tend to delay early
tuber growth rates and crop maturity and may also indirectly influence
percentage DM if foliage growth is curtailed. It is suggested that the effect
of K is on tuber hydration (van der Zaag and Meijers, 1970; Kunkel et al.,
1973) and this may be an indirect effect through the foliage (Hiller et al. ,
1985) as the mineral element composition of the tuber is reported to be
relatively constant regardless of the amount of fertilizer applied (Kunkel et
al., 1973; Harris, 1978). With K, those fertilizers applied as KCI reduce
DM levels slightly more than when K is applied as K2 S0 4 (Dickens et al.,
1962; Birch et al., 1967). These effects may also depend on soil moisture
(Harward et aI., 1956) and plant age (Forster and Beringer, 1983). Applied
P does not appear to have either a consistent or a marked effect on tuber
percentage DM, some authors reporting a decrease, others an increase for
higher application rates (see Kunkel and Holstad, 1972).
In general, dry seasons have been associated with high percentage DM
(Burton, 1966). However, depending upon the stage of growth and develop-
ment of the plant, the effects of soil moisture on tuber percentage DM are
more complex; authors report different responses depending on the periods
of rainfall, irrigation or water stress (Haddock, 1961; Carr, 1985; Shimshi and
Susnoschi, 1985). Irrigation during the early stages of growth increases DM
content because starch production is increased, but continuous or late season
irrigation can depress DM. The rate of water uptake following a period of
moisture stress may also reduce percentage DM for a period of time and
conversely severe water stress during tuber bulking and up to the time of crop
defoliation can increase the tuber percentage DM. As well as the direct
effects of soil moisture on percentage DM, water stress during the period
of tuber initiation will reduce the number of tubers set (van Loon, 1981;
MacKerron, 1985; MacKerron and Jefferies, 1986) and so indirectly affects
the final percentage DM of those tubers which remain to develop.
The effect of defoliation on tuber percentage DM will depend upon the
timing of the treatment in relation to the stage of growth and development
of the crop and weather conditions until harvest. Defoliation occuring
during the main phase of bulking will almost invariably result in tubers
with a lower percentage DM than those allowed to grow until haulm
senescence, from which time there is a reported slight decrease in
percentage DM due perhaps to respiration losses and a change in the
balance between water uptake and DM accumulation (see Gray and
Hughes, 1978). Where there is a requirement for a specific percentage DM
in a defined size grade for processing purposes, the mean tuber percentage
DM for total yield is not the primary consideration. The tuber size distribu-
tion of the crop and differences in the percentage DM of the size grades
have to be considered in relation to the timing of the defoliation treatment.
Water uptake by roots and an influx into the tuber following defoliation
increases fresh weight but there is a corresponding reduction in percentage
DM, as it is usually assumed that there is little change from respiration in
absolute levels of tuber DM during the interval from defoliation to harvest.
Whilst desirable to increase fresh weight yield there may be disadvantages,
particularly for short season processing crops (Hogge, 1989).
524 Tuber quality
As tubers increase in size during the season, the DM content increases,
but this relationship is not linear (Ifenkwe et al., 1974) and a maximum
percentage DM occurs in medium sized tubers. Wurr and Allen (1974)
showed in two maincrop cultivars maximum DM was reached at a tuber size
of about 50 mm. The tuber percentage DM showed a decline for larger size
grades and other studies (Cole, 1975; Wurr et al., 1978; Nelson et al., 1988)
demonstrated a similar relationship in a wider range of cultivars. For a
given number of tubers, the size distribution alters with increasing yield
and at early harvests Nelson et al. (1988) showed that differences in
percentage DM between size grades were smaller than at later harvests and
that the size grade with the highest percentage DM increased over time.
These changes in tuber DM reflect changes in the rate of increase in starch
and cell wall material compared to water uptake. In the early stages of
growth, starch deposition is rapid but the rate declines as growth proceeds
and there is continued uptake of water (Gray and Hughes, 1978).
The DM differences between tubers of different sizes will also be related
to the relative growth of the cortex and pith since these tissues differ in
their DM content (Glynne and Jackson, 1919). In general the percentage
DM is highest in the storage parenchyma between the cortex and vascular
ring, is lower for the periderm and cortex and decreases inwards towards
the medullary tissue of the pith (Johnston et al., 1968). The DM content
also decreases from the stem to the bud end of a tuber (Glynne and
Jackson, 1919).
With an understanding being developed of the intrinsic factors influenc-
ing crop growth and tuber DM accumulation, manipulation of husbandry
practices can be used to manage size distribution and indirectly influence
percentage DM. This would include, for example, the use of physio-
logically aged seed and modified planting populations to affect stem and
tuber numbers. By such means tubers of the desired size and chemical
composition may be produced for specific markets.
12.3.2 Mechanical damage and bruising

Mechanical damage, including bruising, is a major problem for the potato
industry worldwide. For example, a 1981 survey in Great Britain revealed
that after harvest 9% of tubers were severely damaged and 26% were
bruised (Balls et al., 1982). Handling during store loading, unloading and
grading can add considerably to the damage incurred at harvest.
Damage which occurs can be divided into two groups, external and
internal, and classified according to the different forms of damage and its
severity. External damage includes scuffing of the skin, cuts or gouges,
crushing and splits or cracking. It is apparent on inspection and leads to
direct losses at grading or preparation for consumption, increases weight
loss during storage and allows for the ingress of disease pathogens.
Internal damage is usually of two types, internal shattering or cracking
and blackspot. The internal shattering is characterized by cracking of the
tuber tissue which may extend to the outer surface. The fractured tissue
may dehydrate, with some shrinkage and discolouration occurring in the
damaged cells around the margins of the shattered tissue. Blackspot is an
internal discolouration of the tissue which develops some time after the
impact. The bruised area is typically blue-grey with diffused edges,
although in some tissue it may have well defined margins. The discoloura-
tion in both instances is due to the formation of melanin within the
damaged cells.
The internal damage may be visible under the skin of the tuber, but in
most instances it is not apparent until tubers are cut or peeled. Compres-
sion bruising can also occur where potatoes impinge on one another during
storage and where prolonged pressure results in tuber tissue deformation.
All types of internal injury result in increased losses during preparation for
cooking or processing.
The extent and type of damage which occurs to potatoes is influenced by
the type of impact and the energy absorbed and by the interactions with the
physical, and in the case of blackspot, the biochemical properties of the
tuber. The impacts may arise from tubers colliding with moving or stationary
surfaces, stones, clods or other tubers, and the resulting damage and/or
internal bruising is markedly increased by handling at low temperatures
(Ophuis et al., 1958; Smittle et at., 1974).
0.69
R2 = 0.9460
0.67
2- 0.65
"0
Q)
.7,6.5
-e0
~.9~/·8
·7
(/)
.0 0.63
d··
I1l
>.
E' 13.11
Q)
c: 12
w 0.61 ~/ • •15 Y = 0.29068 + 0.02441x
19 .17
• .18
0.59
.20
Damage rating = 0.04591 + 0.92348 ext. score
.21 + 0.14573 width + 0.04401 depth
0.57
12 13 14 15 16 17 18
Damage rating
Figure 12.2 Relationship between energy absorbed and damage rating for a range
of commercial varieties and breeding line accessions. (From Grant and Hughes,
1985.)
526 Tuber quality
Improved mechanization methods throughout the harvesting and hand-
ling operation can be adopted to reduce damage. Where land has a high
stone and clod content, separation and windrowing of the unwanted
material may significantly reduce severe damage (Witney, 1984). On
harvesters where damage is cumulative, improvements in share and web
rod design (McRae et al., 1982; Hutchison and McRae, 1987) and use of
horizontal web agitation, rather than vertical agitation (McRae et al., 1986)
can each make a significant contribution to reducing tuber damage; all
have been incorporated into a harvester which has low damage characteris-
tics. During subsequent handling, store loading and grading, damage can
be reduced by minimizing drop heights, use of cushioning material and
plastic link grading screens (McRae, 1986). Further improvements in
damage reduction may be achieved by the development of electronic,
rather than mechanical, sizing and grading systems and through greater
care in packaging and handling potatoes during transportation (McRae et
al., 1984; Turczyn et al., 1986).
Despite better handling, damage levels are often unacceptable and the
rheological properties of the tuber have been investigated to assess the
effects of abiotic and biotic factors on the type and degree of damage. This
approach helps to identify genotypes less susceptible to damage for use in
breeding programmes, enables more effective evaluations to be made on
the performance of harvesting and handling equipment and allows assess-
ment of the effects of different store management regimes.
The range of tests used to evaluate aspects of damage include simple
tuber drop tests (Ophuis et al., 1958; McRae et al., 1976), impact by falling
bolt (Kunkel and Gardner, 1959; Schippers, 1971) or from pendulums with
either restrained or suspended tubers (Hughes et al., 1985; Noble, 1985;
Skrobacki et at., 1989), vibrating trays, penetrometers and crush tests
(Blight and Hamilton, 1974; De Maine, 1986). The simplest do not provide
standardized impact conditions but can give practical information on the
size of drop which tubers can tolerate and gross varietal differences in
damage susceptibility. The more sophisticated pendulum tests, with appro-
priate instrumentation provide information on impact parameters such as
energy absorbed, tuber tissue deformation and impact duration which
enable more precise relationships to be established between the tuber
characteristics and type and extent of damage. (Fig. 12.2)
(a) Tuber characteristics

To cause damage an impact must occur, and the relative velocity and
masses of the colliding objects have a marked effect on the extent of
damage, with the type of damage being associated with the inherent
rheological and biochemical properties of the tuber interrelated with the
modifying environmental influences. In this context the size and shape of
the tuber influence the amount of damage occurring. A large tuber has
more kinetic energy and will be damaged to a greater extent than a small
tuber dropped from the same height. Hughes et al. (1985) demonstrated
that a very good correlation (r = 0.913) existed between the amount of
energy absorbed from a pendulum impact by tubers held under a static
load with a score for external damage. This measure of the dynamic tissue
strength reflected the tuber susceptibility to cell wall fracture, and the
correlation was improved slightly when a measure of internal cell wall
damage was included. With regard to shape, tubers having a small radius of
curvature, for example in long tubers, are more prone to damage, perhaps
through a more limited contact area and conversely impacting bodies with
a large radius produce less damage (Carruthers, 1982). It has also been
suggested (Murfitt, pers. comm.) that several small impacts, each of which
was insufficient to cause damage, but occurring at the same place on the
tuber may produce bruising.
Although tuber size and shape are varietal characteristics and are aspects
which should be considered in selecting breeding lines for damage resis-
tance, tuber size, in particular, changes during growth and development of
the crop. Husbandry and environmental factors which increase tuber
weights are associated with higher damage levels (Parke, 1963; Hunnius et
al., 1972). As with the susceptibility to internal bruising the precise reasons
why certain varieties are more predisposed to external damage are unclear.
The cultivar responses can be modified by growing conditions and mineral
nutrition; both affect specific gravity and perhaps cell wall structure and
cell size and so influence the rheological properties of the tuber.
The extent of external damage which occurs on impact and the tuber
susceptibility to crushing and splitting have usually been associated with
tissue firmness and the strength of the skin. Cultivars with weak skins,
determined by penetrometer readings (Lampe, 1960) or impact deforma-
tion tests (Hunnius and Munzert, 1972) are often, but not invariably, more
susceptible to cracking. Maintaining an intact skin has a marked effect on
the tuber by limiting the degree of deformation of the underlying tissue
and dissipating absorbed energy. The skin strength is also associated with
the maturity of the tuber, and where immature tubers are harvested or the
skin has not set following defoliation, abrasion during harvesting and
handling operations can result in scuffing of the skin. This is unsightly and
may lead to enzymic browning of the damaged surface tissue which affects
tuber marketability. Increased water loss occurs over the scuffed area
(Burton, 1989), leading to changes in cell turgor and bruise susceptibility.
Tissue firmness has been associated with cell wall strength and internal
turgor. Direct and indirect evidence has established that within a cultivar
excessively firm tubers are more prone to external cracking and shatter
bruise than flaccid tubers, which suffer more from blackspot (Sawyer and
Collin, 1960; Kunkel and Gardner, 1965). Generally, growing conditions
which increase turgor increase susceptibility to cell wall fracture and
external cracking. However, the susceptibility of cultivars to suffer
528 Tuber quality
external damage does differ and turgid potatoes of certain cultivars may be
more prone to external splitting whilst others may sustain internal tissue
damage - shatter bruising, with skin breakage. Both types of damage were
associated by Hughes (1980) with ceU wall fracture and both are generally
increased by high levels of tuber hydration.
(b) Internal bruising

The role of tissue composition and the modifying influences of environ-
mental conditions are more complex in the control of blackspot as both
structural and biochemical components of the tuber are involved. Two
independent conditions are necessary for blackspot development - a
damaging impact on susceptible tuber tissue and tissue that has the
potential to produce the coloured oxidation products (Section 12.3.3).
The effect of turgor pressure on the degree of internal shatter bruising
and blackspot is modified by tuber temperature which affects the rheo-
logical properties of the tissue. As with external damage, there is a
decrease in total internal bruising (shatter cracking plus blackspot) with
increasing temperature (Smittle et aI., 1974). However, as temperature
increases the level of hydration must also increase to minimize the
occurrence of blackspot (see Hiner et al., 1985).
The level of tuber firmness cannot be used to predict blackspot as
different samples produce different levels of blackspot for the same
amount of tuber deformation in a pendulum test (Hughes et aI., 1975a).
Furthermore, very low or high turgor levels can have inconsistent effects.
For example, when potatoes were flaccid due to sprouting the tubers were
less prone to blackspot (Kunkel and Gardner, 1959). Noble (1985) was
able to predict occurrence of shatter bruising using an accelerometer
mounted to the impacting head of a pendulum but was unable to
distinguish between undamaged tubers and those which would develop
blackspot (Fig. 12.3). The accelerometer was perhaps sensitive enough to
detect the cell wall fracture but not the damage to cell contents which has
been suggested as an important element in blackspot susceptibility.
Gray and Hughes (1978) suggested that damage to membranes within
the tuber cell by starch grains initiated blackspot development. This was
related to the greater degree of deformation which occurred in a flaccid cell
under a given pressure than occurred in cells with a high turgor pressure
(Nilsson et aI., 1958). The differences between varieties it was suggested
would be a result of differences in the rheological properties associated
with the cell wall composition and cell wall elasticity for a given turgor
pressure. Cell size, which has been shown to affect susceptibility to internal
damage (Olsson and Fridell, 1975; van Es and Hartmans, 1975) and
bruising (Hudson, 1975) would also be important.
Potassium has the most consistent effect on blackspot and tubers grown
with low K levels are more susceptible to blackspot (Vander Zaag and
(a) (b)
c;-
CI) 200
:sz
0
i= 400
«
0:
w
....J
W
()
w 600
Cl
800 , , ,
0 2 4 6 0 2 4 6
TIME (x1 0-3 5- 1)
Figure 12.3 Acceleration - time curves produced by a pendulum - mounted

accelerometer showing (a) single peak associated with undamaged tubers and
(b) multiple peaks related to internal crushing or shatter bruising; cv. King
Edward. (From Noble, 1985.)
Meijers, 1969). Increased K levels often reduce specific gravity and it was
supposed the effect in reducing blackspot was on tuber hydration (Kunkel
and Gardner, 1965; van der Zaag and Meijers, 1970). An indirect effect of
high K levels has been related to lower levels of tuber phenols and
phenolase activity.
Increasing N may also reduce specific gravity of tubers, but the effects on
blackspot can be quite different to that of increased K and may be due to
an interaction of a number of factors, including increased tuber size and
possibly differences in turgor. It appears that the specific gravity of the
tubers is not a major factor determining blackspot susceptibility between
cultivars, although within a cultivar it is an important consideration, with
increasing specific gravity generally giving higher bruising.
In stored potatoes a good correlation exists between increased blackspot
and increased specific gravity, but this may reflect changes in turgor due to
water loss. Noble (1985) found that the impacts which resulted in the most
tissue deformation produced the most blackspot in the susceptible cultivar
Record. This was associated with a low loading velocity and long impact
duration. For the same energy absorbed with a high loading velocity and
relatively short impact more shatter cracking occurred. The long duration
and low loading in the viscoelastic potato tissue where the strain is time
530 Tuber quality
dependent may be reflected in the higher levels of blackspot rather than
internal cracking which occurs in tubers subjected to pressure deformation
and suffering from dehydration as a result of prolonged storage or low
humidity.
The interactions causing external and internal damage are complex and
the reasons why particular tubers are more susceptible than others is not
fully understood. Although several components have been studied in
detail, results reported have been apparently conflicting. However, in most
of these instances not all factors which may have had a bearing on damage
incidence, e.g. tuber specific gravity, soil nutrient status or temperature,
had been considered. Nevertheless, of the factors influencing damage,
most in some way seem to be associated with tuber hydration. Although
little can be done which affects tuber tissue structure, the range of cultural
practices adopted during growth and subsequent storage can have a
significant effect on damage levels. The maintainance of optimum tuber
turgor, avoidance of unfavourable temperatures and reducing impacts can
produce marked benefits.
12.3.3 Enzymic browning

The discolouration of potato flesh from external and internal bruising
caused by mechanical damage is a major cause of loss to producers and
processors. Discolouration through enzymic browning is well documented
(Walker, 1977; Rhodes and Wooltorton, 1978). The reasons for this
enzymic browning of cut surfaces, external bruising and blackspot caused
by internal bruising are well understood. The principal cause is the
oxidation of tyrosine to 3, 4-dihydroxyphenylalanine (DOPA) catalysed by
the enzyme polyphenol oxidase (PPO). DOPA is then oxidized to dop-
aquinone. The latter cyclizes to 5,6-dihydroxyindole derivatives which are
oxidized to reddish orange dopachrome pigment. Once dopachrome has
been formed a series of non-enzymic polymerizations, oxidations and
reactions with proteins occur to produce, finally, a black melanin pigment
(Joslyn and Ponting, 1951). Enzymic browning can also involve oxidation
of o-hydroxyphenols e.g. chlorogenic acid, producing lighter brown 0-
quinones (Pierpoint, 19uO). Chemical control of browning involves inhibit-
ing PPO activity by adjusting pH (Amla and Francis, 1961), adding
bisulphite or sulphydryl compounds (Lerner, 1953), chelating the copper
from the enzyme (Matthew and Parpia, 1971), the use of reducing
compounds which convert o-quinones to the o-hydroxyphenol state, or
chemicals which react with o-quinones to give colourless addition products
(Muneta, 1981 and references therein). Muneta (1981) has shown that
sodium sulphite, cysteine, dithiothreitol and sodium diethyldithiocarbamic
acid (DIECA) inhibit browning by restricting the oxidation of tyrosine.
Sodium bisulphite and DIECA also cause enzyme inactivation. Cysteine
and dithiothreitol inhibit tyrosine oxidation but do not inactivate PPO.
Ascorbate and dihydroxyfumarate also inhibit blackening by reducing
dopaquinone to DOPA. There is also evidence (Burrell, 1984) that
phenoxyacetic acids such as 3,5-dichlorophenoxyacetic acid inhibit brown-
ing, probably by delaying an increase in phenylalanine ammonia lyase
(P AL) and cinnamate-4-hydroxylase activities. This would reduce the
synthesis of phenols such as chlorogenic acid.
The incidence of enzymic browning is affected by climatic and edaphic
factors. In certain cases a direct relationship between rainfall at a given
centre. and the amount of tyrosine in tubers has been reported (Mapson et
al., 1963). In other instances the levels of browning, tyrosine and some-
times phenolase are reduced when higher levels of K are fed to plants
(Welte and Miiller, 1966). Nitrogen rarely increases the severity of
discolouration. Whilst a direct relationship has been observed between
tyrosine and enzymic browning the relationship varies depending on
variety and environmental conditions (Umaerus and Olsson, 1974). This
suggests that factors other than tyrosine content playa role.
E 0.5
E
0
0
If)
0.4
co
c:i
Q.
Ol
0.3
L.S.D. I
c
·c 0.2
~
e
.0
~ 0.1
E
*
0..
0.0
a b d e 9 h GW PJ PD MP D
Solanum hjertingii accessions Solanum tuberosum varieties
Figure 12.4 Comparison of the potential browning of freeze-dried potato

macerates of Solanum hjertinjii accessions and commercial Solanum tuberosum
varieties. (After Oubb et al., 1989b.) OW = Golden Wonder; PJ = Pentland
Javelin; PD = Pentland Dell; MP = Maris Piper; D ;; Desiree.
Genotypic variation exists in the extent of enzymic browning (Fig. 12.4)·

and the work of Firbas (1961) and Woodwards and Jackson (1985) suggests
that Solanum hjertingii and other wild species from the series Longipedi-
cellata possess a non-browning character. Substrate analysis of some of the
accessions revealed lower chi orogenic acid contents in the wild type
compared with commercial varieties, while tyrosine levels varied by up to
an order of magnitude between different accessions (Gubb et al., 1989a).
Gubb et al. (1989b) evaluated enzymic browning quantitatively for fifty
genotypes, representing seven accessions of S. hjertingii together with five
commercial cultivars of S. tuberosum with a known range of enzymic
browning. Of S. hjertingii clones, 94% were less severely affected by
532 Tuber quality
browning than commercial varieties. Eighteen clones showed no visible
discolouration when sliced. Furthermore, both mono- and diphenolase
(PPO) activities were reduced in non-browning clones with the hydroxyla-
tion of monophenol being the most important. These data implicate
reduced PPO activity as the factor limiting browning, while tyrosine
content determines the final degree of discolouration.
12.3.4 Glycoalkaloids
The major glycoalkaloids in potatoes are a-solanine and a-chaconine, both
of which are derived from solanidine (Maga, 1980). They have been
associated with a bitter taste (Sinden et aI., 1976) and are reported not to
be destroyed by cooking (Sizer et al., 1980; ladhev et al., 1981), although
Ponnampalam and Mondy (1983) showed some decrease by baking and
frying. The glycoalkaloids are not uniformly distributed throughout the
tuber, 30-80% being associated with the peel (Maga, 1980). High con-
centrations (200-400 mg per 100 g) are also found in potato sprouts (Wood
and Young, 1974; Fitzpatrick et al., 1977). Because most glycoalkaloids are
in the skin, concentrations are usually low in prepared potatoes and the
bitter taste means that only in exceptional circumstances has it been
responsible for poisoning (see Maga, 1980; Burton, 1989).
Potatoes with mean levels above 20 mg per 100 g FW are considered
undesirable for human consumption (Jadhev and Salunkhe, 1975) because
of their bitterness and toxicity, although Lepper (1949) considered 15 mg
per 100 g to be too high. Some individuals are also able to detect a bitter
taste in tubers with a glycoalkaloid content as low as 10 mg per 100 g (Ross
et al., 1978). Most common European and American varieties have
glycoalkaloid contents below 20 mg per 100 g level and Burton (1989)
quoting several authors reports typical levels of 2-10 mg per 100 g FW. An
extensive survey of retail potatoes in the UK (Davies and Blincow, 1984)
found mean glycoalkaloid levels of 10.4 mg per 100 g for maincrop
varieties, 11.3 mg per 100 g for UK grown earlies and 12.3 mg per 100 g for
imported earlies. Only two samples out of 133 in the survey exceeded the
20 mg per 100 g level. Other authors (Wo,]f and Duggar, 1946; Verbist and
Monnet, 1979) have similarly reported higher concentrations in immature
tubers than in mature tubers; this may be due to the larger surface area to
volume ratio for the small tubers. Verbist and Monnet (1979) found 44.8
mg per 100 g in one immature sample of cv. Sirtema and an average level of
21.8 ± 3.6 mg per 100 g for immature tubers of eight cultivars. Certain
Scandinavian cultivars also have high glycoalkaloid contents (Baerug,
1962) and in the United States in 1970 the cultivar Lenape was withdrawn
from commerce after its glycoalkaloid content was found to range from
18.6 to 35.4 mg per 100 g FW. The high levels in some cultivars may also be
influenced by the growing season as glycoalkaloid contents are significantly
influenced by environmental factors (Sinden and Webb, 1972). Those
Aspects affecting quality of the cooked potato 533
cultivars with genetically high levels are apparently more susceptible to
excessive glycoalkaloid production when stressed,
Although differences have been found amongst the same cultivar grown
at different locations these have been attributed to factors such as frost or
hail damage to foliage (Hutchinson and Hilton, 1955) or crop maturity and
tuber damage (Sinden and Webb, 1972),
Exposure to light increases the levels of glycoalkaloids in the tuber,
However, glycoalkaloid formation is independent of chlorophyll formation
and the associated greening of tubers (Gull and Isenbreg, 1960), The
increase in glycoalkaloid levels depend upon the duration of exposure,
wavelength and intensity of light (Conners, 1937; Wolf and Duggar, 1946;
Gull and Isenberg, 1960; Liljemark and Widoff, 1960), with green light
most effective in minimizing glycoalkaloid formation. It is influenced by
temperature (Salunkhe et al., 1972) and high levels are associated with
damage and bruising (Fitzpatrick et al., 1978). Some workers have
attempted to minimize light-dependent glycoalkaloid synthesis with chem-
icals and there has been some success following treatment of tubers with
mineral oil and lecithin (Jadhev and Salunkhe, 1974; Wu and Salunkhe,
1977).
Glycoalkaloids increase during storage (Cronk et al., 1974) and Hilton
(1951) has attributed a more bitter taste in tubers stored at a low compared
with a high temperature stored potatoes to a greater accumulation of
solanine. Similarly, Zitnak (1953) has shown a marked increase in glycoal-
kaloid levels in cv. Netted Gem stored at 4-8°C compared with 12-15°C.
Glycoalkaloid content increased from 7.9 to 15.4 mg per 100 g over 6
weeks at the low temperature whereas at the higher temperature only a
slight increase occurred, from 7.5 to 8.7 mg per 100 g.
Although the presence of glycoalkaloids is undesirable for consumption
purposes it has been suggested that they playa role in protecting the plant
against pathogens. Allen and Kuc (1968) have shown glycoalkaloids to
possess fungitoxic properties and McKee (1959) reported toxicity to spores
of Fusarium caerulum. However, Deahl et al. (1973) found no correlation
between blight resistance and glycoalkaloid levels.
12.4 ASPECTS AFFECTING QUALITY OF THE COOKED POTATO
12.4.1 After-cooking blackening

After-cooking blackening is a non-enzymic discolouration of the tuber
which is also referred to as stem-end blackening as it is usually more
obvious at the stolon end of the tuber. It occurs to some extent in most
cultivars and to potatoes grown under a wide range of cultural conditions,
with maximum intensity of colour usually developing when potatoes stand
for a period of time after cooking. Its absence was considered to be the
534 Tuber quality
most important quality attribute for boiled new and maincrop potatoes to
consumers in Great Britain (Potato Marketing Board, 1988). Although the
defect is at its worst when susceptible varieties are boiled or steamed, the
discolouration may also be evident following cooking of baked potatoes,
French fries, canned and dehydrated potato products.
Hughes and Swain (1962a, b) found that during cooking chlorogenic acid
combined with iron to forma complex which oxidized on cooling to give
the coloured ferri-dichlorogenic acid responsible for the blue-grey dis-
colouration of tubers at the normal pH (6.0-6.5) of cooked potatoes. At
pH 5.5 a grey-green colour occurred due to the formation of the mono-
phenolate complex, whereas the triphenolate gave a dark grey-reddish
pigment at pH 8.0. The formation of the pigment was also found to be
affected by the presence of chelating agents, most importantly citric acid
(Mulder, 1949) but also malic acid and phosphate which competed with the
chlorogenic acid for the iron to form a colourless complex. The citric acid
sequesters iron more efficiently than chlorogenic acid (Mulder, 1949) and
although the level of iron in the tuber may vary (Lampitt and Goldenberg,
1940; Wurster and Smith, 1963) the amount available to complex with the
chiorogenic acid can be less dependent on this than on the citric acid
content of the tuber. Consequently, the degree of after-cooking blackening
is mainly dependent upon the amount of the two acids present and Hughes
and Swain (1962a) showed that the distribution of after-cooking blackening
within the tuber was dependent upon the ratio of chlorogenic acid to citric
acid. The level of citric acid was lower at the stem end of the tuber
(Shekhar and Iritani, 1979) where more after-cooking blackening tends to
take place.
As well as the difference between the stem and bud ends of the tuber
with regard to chlorogenic acid levels, Brandl and Herrmann (1984)
reported that concentrations of the caffeic acid compounds increased
greatly from the inner to the outer sections of the tuber with about 50%
located in the peel and adjoining tissue. These workers identified the main
chlorogenic acid as 3-0-caffeoylquinic acid (n-chlorogenic acid) and total
chlorogenic acid levels for ten varieties ranged from 33 to 104 mg g.t FW.
A similar range of 34-144 mg g-t FW was reported for ten cultivars by van
Es and Hartmans (1987a). Hughes :and Evans (1967) demonstrated that
such differences between cultivars in the amounts of chlorogenic acid and,
to a lesser extent, citric acid, were largely responsible for the different
levels of after-cooking blackening which occurred. The ranking of the
degree of blackening was usually similar in any given year or soil for those
varieties which had either very good or very poor freedom from discoloura-
tion. In general, those with high chlorogenic acid levels blackened the
most. Muneta and Kaisaki (1985) have also identified the purple coloured
complex from ascorbic acid as another possible source of after-cooking
blackening.
The cultivar differences in susceptibility and the importance of growing
conditions and season in modifying the incidences of after-cooking
blackening have long been recognized and have been extensively reviewed
(see Gray and Hughes, 1978).
Certain factors such as a high ratio of N to K and cool wet seasons were
usually found to make tubers more susceptible to after-cooking blackening.
These environmental factors were considered to affect, primarily, citric
acid levels (Hughes and Evans, 1967). However, in some situations both
citric acid and chi orogenic acid levels are reported to be affected. Lynch
and Kaldy (1985) found consistently higher levels of citric acid and K and
also lower levels of chlorogenic acid in Russet Burbank grown in southern
as compared with central Alberta. The lower chlorogenic acid levels were
associated with cooler temperatures and soils with a higher organic matter
content (Kaldy and Lynch, 1983). Hughes and Evans (1967) showed that
potatoes blackened more and had a low level of citric acid and were only
sometimes higher in chlorogenic acid when grown on a fen soil (high
organic matter and low K) than when grown on a clay soil (low organic
matter with high K). Experiments by Hughes and Mapson (1966) on plants
grown under controlled nutrient regimes substantiated the role that Nand
K play in controlling the discolouration of potatoes. Very good correla-
tions were generally found between the amount of K in the tuber and
citrate levels, except when chloride was present in the nutrient solution.
When ammonium ions or urea were present the uptake of K and level of
citric acid was reduced. Chlorogenic acid levels were not greatly affected
by mineral nutrition except for the effect of high levels of nitrate and urea;
the high N, it was considered, might have explained why higher amounts of
chlorogenic acid were sometimes found in tubers grown on fen soils. Very
high levels of chloride affected the level of water soluble iron and in certain
soils uptake of iron may also be affected by pH and high calcium levels.
Under such circumstances iron may have a greater involvement in control-
ling blackening (Heisler et al., 1963).
An understanding of varietal differences and the role of Nand K has
provided the advisory base for minimizing the incidence of severe after-
cooking blackening in the harvested crop. However, despite the large
proportion of the crop which is stored, little attention has been given to the
effects of the storage environment which appears to influence the degree of
after-cooking blackening in some cultivars. There is evidence of consistent
changes in the levels of after-cooking blackening and the ranking for eight
cultivars following storage at 7°C (Potato Marketing Board, unpublished).
Changes in the levels of chemicals contributing to after-cooking blackening
occur during storage, but their effects on the cooking quality of the samples
have not been established. Rogozinska et al. (1986) reported cultivar
differences in, and seasonal effects on, chlorogenic acid during storage at
4°C and 95% RH. Most cultivars in a 2-year study showed a decrease in
chlorogenic acid during 4 months' storage. However, in one year the level
rose in two of the nine varieties. Rumpf (1972) reported a decline of over
536 Tuber quality
90% in the citric acid content of tubers during 8 months' storage at lOoC
and 85-90% RH. In contrast, Muller (1975) (reported by Burton, 1989)
examining 28 cultivars stored at 8°C for 10 weeks showed in some
instances, e.g. cv. Tasso, a steady increase in citric acid content but in
general there was no consistent change.
12.4.2 Texture
The texture of the potato has been related to many structural and chemical
properties of the tubers and to environmental factors that influence them.
However, differences in texture from material with, for example, similar
dry matter contents do occur and these are influenced by the cooking
conditions. The tendency to breakdown on cooking is a primary considera-
tion and this has been established to be due to cell separation and factors
influencing elements of this relationship have been examined in detail.
However, how these physico-chemical differences are reflected in the
consumer's perception of a particular texture and their preferences are
even less well understood and remain to be developed further.
Although environmental factors affecting the growth and development
of the plant can influence, to some extent, the texture of the potato,
particularly in relation to its dry matter accumulation, it is well established
that genotype plays a most important role (see Howard, 1978). Descriptions
of cultivars usually include an evaluation of their dry matter content and
less frequently an assessment of their texture, e.g. mealiness (see National
Institute of Agricultural Botany, 1990).
Salaman (1926) used the terms floury, close, waxy and soapy to describe
the texture of steamed potatoes. The emphasis in these categories is on
disintegration of the tubers. However, in an evaluation of texture in
cooked potatoes it is important to encompass aspects of palatability and
other terms, often not well defined, have been used in the literature.
Mealiness, which is synonymous with floury, appears to have three
components - disintegration, mouth feel and dryness (from appearance)
(Gray and Hughes, 1978). The European Association for Potato Research
Table 12.3 Characters and degrees of expression used to describe aspects of

cooked potato texture (after Lugt, /961)
Texture Expression of characteristic
Disintegration None Slight Moderate Complete

Consistency Firm Fairly firm Soft Soft, uneven
Mealiness Not mealy Slightly mealy Mealy Very mealy
Dryness Humid Slightly humid Slightly dry Dry
Structure Fine Fairly fine Slightly coarse Coarse
has established a widely used system (Lugt, 1961) which provides a basis
for comparison between subjective assessments.
Because of its ease of measurement the specific gravity or dry matter
content of tubers is frequently used to give a general indication of the
degree of disintegration which can be expected but, although these
parameters have often been correlated, particularly within a variety, there
are instances where the relationship is poor.
The relationship between high specific gravity and increased disinteg-
ration has been attributed to the level of starch in the tuber; the swelling
during gelatinization exerts greater internal pressure in tubers of a high
starch content leading to greater cell distention and more effective
separation of adjacent cells (see Burton, 1966). However, there appears to
be no experimental evidence of tuber starch levels directly increasing cell
separation by a swelling pressure (see Hoff, 1972) and in a study where
intercellular adhesion was measured objectively a direct relationship was
found to exist with starch content of the tuber (Sharma et at., 1959).
An indication of the role of cell separation, rather than the starch
content, in influencing the degree of disintegration was also provided by
studies which showed that tubers with high starch and amylose levels are
associated with increased measurements of tuber firmness (Linehan and
Hughes, 1969a; Iritani et al., 1977). These results suggested that the effect
of starch levels on cell separation were probably indirect and possibly
reflected differences in cell wall composition and cell size (Gray and
Hughes, 1978). Increased firmness associated with high starch levels has
also been attributed to increased viscosity (Warren and Woodman, 1974).
An increase in cell size has been related to a reduction in cell adhesion
during cooking (Barrios et at., 1963; Linehan et al., 1968; Gray, 1972). The
cell size is affected by variety (Linehan et at., 1968) and increases with
tuber development (Plaisted, 1957; Gray, 1972; Reeve et al., 1973).
However, cell size alone does not always explain differences in texture with
maturity (Gray, 1972). In developing tubers Geddes et al. (1965) have
shown an increase in starch grain size and an increasing proportion of
amylose in the starch content which affects gelatinization temperature.
Such differences may, together with structural or chemical changes affect-
ing cell adhesion (Hughes et al., 1975b), be also associated with texture
changes during maturity.
The softening of the potato and decreased cell adhesion during cooking
are generally attributed to solubilization of the pectic substances in the
middle lamella. The cell walls of potato tubers are largely hemicellulose
and cellulose within a pectic matrix and permeated by water (Selvendran et
al., 1987) with components held by covalent and hydrogen bonds. The
pectic substances in the cell wall and linking adjacent cells at the middle
lamella are largely polymers of esterified galacturonic acid and polysac-
charides (Hoff and Castro, 1969; Jarvis et al., 1981; Ring and Selvendran,
1981) with polygalacturonic acid chains joined by Ca and Mg bridges.
538 Tuber quality
Most evidence suggesting that solubilization of the middle lamella was
associated with decreased cell adhesion was largely circumstantial (see
review by Linehan and Hughes, 1969b). However, direct relationships
were established by Hughes and co-workers who demonstrated that
objectively measured changes in cell firmness were closely related with the
release of pectic polyuronide into the cooking medium when potatoes discs
of a uniform starch content and cell size had been cooked for varying times
(Hughes et al., 1975c,d).
The loss of cell adhesion and pectin degradation during cooking is
associated with leaching of depolymerized pectic substances into the water
(Bettelheim and Sterling, 1955) and the removal of Ca and Mg bridges
between the galacturonic acid chains by ion exchange and hydrogen bond
breakage (Hughes and Faulks, 1973; Hughes et al., 1975b), or chelation by
naturally occurring phytin (Wager, 1963). Keijbets (1974) has shown that
solubilization of pectic substances can also occur by transelimination.
The role of these different mechanisms in affecting cell cohesion will be
influenced by the prevailing cooking conditions and the composition of the
tissue. For example, addition of monovalent ions in the cooking medium or
pretreatment with a chelating agent (e.g. sodium hexametaphosphate)
reduce intercellular adhesion (Hughes et al., 1975b), whereas soaking
potatoes in CaCl2 prior to cooking is an effective firming agent. The latter
effect has been attributed to formation of Ca bridges between the
galacturonic acid chains. However, the effect may also be partly the
consequence of a reduction in transelimination due to a loss of methoxyl
groups, as the pH of the cooking medium is lowered (see Gray and
Hughes, 1978).
The structural and chemical differences of cell walls also interact with
the cooking medium to affect the rate and degree to which the progressive
changes in solubilization of the middle lamella occur. During cooking the
composition of polymers released from the potato tissue is altered (Hughes
and Faulks, 1973) and changes in cell adhesion are gradual until the pectins
are completely degraded (Nonaka, 1980). It has also been suggested that
extruded starch gel between cells and strands of coagulated protein linking
cells (Roberts and Procter, 1955; Reeve, 1967) are involved in cell
adhesion.
With the range of factors influencing cell adhesion it is not surprising that
the cooking quality of individual tubers can reflect differences in structure
and composition of the potato when non-uniform behaviour is observed.
Sloughing of the outer layer of the tuber as a result of disintegration in the
region of the vascular ring has been attributed to observed differences in
the DM content of the tissue. However, elements such as heat penetration,
the resulting degree of pectin degradation and leaching from different
depths of the tissue will all play a role in affecting cellular adhesion in a
particular tuber (see review in Burton, 1989). Differences in firmness
.between bud and item end of tubers are recorded after cooking (Iritani et
al., 1977; Nonaka, 1980) and where growing conditions affect DM
distribution, e.g. second growth or chain tuberization, texture differences
are apparent.
The changes which occur during cooking affecting the middle lamella
also affect the cell walls and weakening of the pectic structure can lead
to cell wall disintegration. This breakage may be exacerbated with
mechanical handling of the cooked potato, by mashing or mechanical
processing, e.g. extrusion cooking. In these circumstances, the starch
content and its composition may playa role in determining the degree of
stickiness which results.
12.4.3 Flavour
Flavour of the cooked potato is one of the attributes most frequently cited
by consumers when evaluating a variety's acceptability. From a sensory
viewpoint it is usually considered as a composite of the tuber's aroma and
taste, both of which are dependent upon its constituents and the changes
occurring to them during cooking.
The particular aroma of boiled potatoes is produced by reactions
between amino acids, sugars and pectins (Self et al., 1963) and is likely to
be affected by the relative contribution of these compounds (Casey et aT.,
1963) and will therefore be influenced by the growth and storage of the
crop. Although a great many compounds contribute to the aroma of the
cooked potato the carbonyl- and sulphur-containing volatiles have been
identified as making the most important contribution (Swain and Self,
1964) despite occurring in much smaller quantities than methanol which
was the major volatile produced during cooking. An 'earthy' aroma is
often described for potatoes and Buttery and Ling (1973) ascribed this to
2-methoxy-3 isopropylpyrazine. Pareles and Chang (1974) have identified
various methylpyrazine compounds as contributing to the distinctive aroma
of baked potatoes.
The aroma and more particularly the taste of the potato are also
associated by the consumer with its 'mouth-feel' or texture and whether it
is perceived to be waxy or mealy, dry or moist and with its consistency and
structure after cooking (see Section 12.4.2).
A mild, but not insipid, taste is preferred by consumers in Great Britain
for maincrop potatoes and new season potatoes are sometimes considered
to have a slightly sweet taste. This sweetness is often associated with a firm,
waxy texture of immature harvested potatoes. Although the tuber total
sugar levels usually show a decline towards maturity, the relative dif-
ferences during crop development are less than can occur during storage.
Sweetness in cooked potatoes is more frequently described from stored
potatoes which have either been subjected to low temperatures or have
been held for an extended period and are exhibiting senescent sweetening.
540 Tuber quality
12.5 PROCESSING QUALITY
The proportion of the crop utilized in a processed form is increasing and

represented 28% of consumption in Great Britain in 1988-89. French
fries and chips (crisps in the UK) are the major products for direct
consumption. On an international basis considerable tonnages are also
used to produce a range of dehydrated products and a proportion of the
crop is also canned. Starch and alcohol production from potatoes are
important elements in certain countries. The production of these are
considered by Burton (1989). Aspects of the quality requirements for
processing into French fries and chips, e.g. tuber size, shape and dry
matter content have been examined. However, a most important quality
criterion for these products relates to the fry colour of the fresh and stored
potato.
12.5.1 Non-enzymic browning

The browning of chips (crisps) and French fries at high processing
temperatures is due to a typical Maillard reaction between reducing sugars
and the a-amino groups of nitrogenous compounds (Schallenberger et al. ,
1959). In practice, the quantities of amino-N are rarely limiting and the
extent of browning is better correlated with the concentration of the
principal reducing sugars (hexoses) glucose and fructose (Gray and
Hughes, 1978). The ideal reducing sugar content for processing into chips
(crisps) is generally accepted to be 0.1 % of tuber FW with 0.33% as the
upper limit. For French fries the upper limit may be as high as 0.5%
(Burton and Wilson, 1970). Nevertheless, published correlations between
reducing sugar concentrations and browning are variable, as are the slopes
of the regressions (Schwimmer et al., 1954; Schallenberger et at., 1959;
Wunsch and Schaller, 1972), possibly reflecting variation in amino acid
content. Recent findings suggest a correlation between soluble amino
acids, particularly asparagine, and reducing sugar levels in cv. Pentland
Dell during the early weeks of storage at 5°C (Hart and Cobb, pers.
comm.). Other factors affecting correlations and regressions include
differences between experimenters in procedures of sampling and frying
and in methods of estimating sugars and fry colour. Tuber pH and organic
acid content may also be important (Fuller and Hughes, 1984).
Four types of sweetening can be defined. The first results from tuber
immaturity, the second from rapid sprout growth, the third from senescent
sweetening and the fourth from exposure of tubers to low temperatures.
All categories undoubtedly share some common underlying biochemical
mechanisms. Tuber maturity is extremely important in determining reduc-
ing sugar content at the beginning of storage (van Es and Hartmans,
1987b). Immature tubers can accumulate substantial quantities of glucose
and fructose within a few days or weeks of transfer to lOoC storage
Processing quality 541
(Richardson et al., 1990). In many cases this may be attributed to high
sucrose levels at harvest (Burton, 1965; Sowokinos, 1978). However,
sucrose losses may not totally account for reducing sugar gains, particularly
in mature tubers (Burton and Wilson, 1978). To maintain desirable levels
of reducing sugars in stored tubers, rapid sprout growth and starch
breakdown at storage temperatures above 5°C are prevented through the
use of chemicals, principally propham and chlorpropham. Even at high
temperatures, however, manufacturers may be obliged to remove excess
sugars from the raw product, by costly blanching procedures for example.
Furthermore, with increasing concern over the use of chemicals on
foodstuffs alternatives to the use of chemical sprout suppressants are
sought. Low temperature storage (below SOC) inhibits sprout growth and
has the added advantage of reducing the incidence of bacterial soft-rots.
However, low temperatures induce a substantial increase in reducing sugar
content with a concomitant increase in the extent of browning (Isherwood,
1973). This deterioration in fry colour may be reversed by reconditioning
tubers at an elevated temperature (Iritani and Weller, 1978; Storey and
Shackley, 1987a).
Low temperature sweetening is a complex phenomenon involving the
interaction of several pathways of carbohydrate metabolism. The factors
which regulate starch, sucrose and hexose turnover rates are all important
in establishing the sugar balance in storage. Isherwood (1973, 1976)
provided convincing evidence that starch degradation can provide the
carbon skeletons for reducing sugar accumulation. The regulation of starch
mobilization in potato tubers is not well understood, in partkular the
relative roles of amylases and phosphorylases. Morrell and ap Rees (1986)
suggested that starch breakdown was phosphorylytic but recent evidence
(Brisson et al., 1989) has shown that starch phosphorylase is located
outside the starch granule in mature tubers, casting doubt upon the
importance of this enzyme. Other studies have also found no direct
relationship between phosphorylase activity and starch breakdown in
sprouting tubers (Davies and Ross, 1986b; Davies and Viola, 1988). Some
workers have reported that substantial increases in amylase activity are
not associated with either low temperature sweetening or rapid starch
breakdown in sprouting tubers (Davies and Ross, 1986b; Morrell and ap
Rees, 1986). However, recent evidence indicates enhanced exo-amylase
and endo-amylase activities in tubers held at 4°C compared with lOoC
(Cochrane and Duffus, 1989). Genotypes accumulating less reducing
sugars than others at low temperature also showed the least difference in·
activities between temperature extremes.
Sucrose accumulation is known to precede or accompany the rise in
reducing sugars at low temperatures. Since sucrose is a precursor of
glucose and fructose (via invertase action), the regulation of sucrose
formation has received some attention. Sucrose phosphate synthase cata-
lyses the synthesis of sucrose at low temperature (Pressey, 1970) and the
542 Tuber quality
amylases
glucosidase phosphoryl"se
G-1-P
i
In~vrtase
Invertase
inhibitor
:::
'--------G-6-P+---------'
phospho
glucomutase
I SUCROSE IIFRUCTOSE I phosphohexose

isomerase
1
~M"T ~
UTP
00_ .."","
SUCROSE synthase UDP
PHOSPHATE "'(;--A.,.----.--'---'-------F-6-P
pyrophosphorylase
1
UDP A T P - - -__ ___---~PPi
PPi: PFK
ATP:PFK
ADP Pi
F-1,6-P 2
GLYCILYSIS
CYANIDE INSENSITIVE UDPG

RESPIRATION?
Figure 12.5 Possible metabolic pathways regulating hexose accumulation in

tubers.
activity of the enzyme is considered to be in excess of that required to
account for the rate of sugar accumulation (Pollock and ap Rees, 1975).
Indeed, there is little evidence that coarse control mechanisms regulate
sucrose accumulation in the cold (ap Rees et al., 1988). The present
evidence favours an accumulation of sucrose following the diversion of
hexose monophosphates away from glycolysis and into sucrose biosyn-
thesis (Dixon and ap Rees, 1980). In tubers the key glycolytic enzymes
ATP-dependent phosphofructokinase (ATP-PFK), and pyruvate kinase
are cold labile (Pollock and ap Rees, 1975; Dixon and ap Rees, 1980;
Dixon et al., 1981). This is believed to reduce carbon flow into respiration
and increase the flux into sucrose. Four forms of ATP-PFK exist in potato
tubers (Kruger et al., 1988) and in the standard UK crisping cv. Record
only one form does not exhibit cold-lability. Cultivar Brodick, bred at the
Scottish Crop Research Institute, accumulates much lower levels of
reducing sugars at low temperature than Record and none of the four
forms of ATP-PFK is cold labile (ap Rees et al., 1988). The picture is
made more complex by the existence of a far more active pyrophosphate-
dependent phosphofructokinase (PPj-PFK) in tubers (van Schaftingen et
al., 1982). This raises the possibility of by-passing the cold labile ATP-
PFK. Although some genotypes which show restricted accumulation of
sugars in the cold may also have high PPj-PFK activity (van Es
and Hartmans, 1987d), there is, as yet, no convincing evidence that
pyrophosphate-PFK acts as a glycolytic enzyme in plants (ap Rees et aI.,
1988). Furthermore, PPj-PFK is itself cold labile (ap Rees et al., 1988) and
the Ka for its activator fructose-2, 6-bisphosphate increases by an order of
magnitude when the temperature is reduced from 25°C to 2°C (Trevanion
and Kruger, 1989). Irrespective of which enzyme plays the major role it
appears that the potential for converting fructose-6-phosphate to fructose-
1, 6-bisphosphate is reduced substantially at low temperatures.
The importance of the reaction catalysed by the phosphofructokinases
hinges upon the nature of the compound crossing the amyloplast
membrane (ap Rees et al., 1988). Evidence obtained from recent nuclear
magnetic resOnance studies on potato (Viola and Davies, unpublished)
confirms similar studies on wheat (Keeling et al., 1988) i.e. six-carbon
molecules eulter the amyloplast to support starch biosynthesis. If C6
compounds are transported in the opposite direction then a primary role
for the phosphofructokinases in regulating the supply of carbon for sucrose
synthesis at low temperature is envisaged. If C3 compounds are trans-
ported then the cold lability of pyruvatekinase could be the key regulatory
step. However, several questions remain to be answered about the
importance of the cold lability of these enzymes in regulating cold induced
sweetening. Why, for example, does tuber respiration rate increase on
transfer to low temperature if glycolysis is impaired (Burton, 1966 [cited by
van Es and Hartmans 1987c]; Isherwood, 1973)? Also, if pyruvate kinase is
the key regulatory enzyme how do hexose monophosphates accumulate via
544 Tuber quality
reverse glycolysis if PPi-PFK activity is also cold labile? There is little
evidence that substantial reverse glycolysis in tubers is mediated by
fructose-I, 6-bisphosphatase (ap Rees et al., 1988).
While considering these factors it is worth commenting further on the
relationship between tuber respiration and sugar accumulation in the cold.
Ehlenfeldt et al. (1989) measured CO2 output from intact tubers of a range
of genotypes and concluded that, in general, good chipping cultivars had
higher rates of respiration than poor ones. However, the relationship was
not clear cut and the high CO 2 output characteristic is probably not
primarily responsible for low sugar accumulation. Amir et al. (1977)
demonstrated an increase in tuber A TP content and CO 2 output in tubers
stored at 4°C together with the expected rise in sugar content. The burst in
respiration at low temperature appears to result from the induction of the
cyanide resistant pathway (Sherman and Ewing, 1982). Solomos and Laties
(1975) proposed that the activation of cyanide insensitive respiration in
tubers de-regulates aerobic respiration, with sucrose synthesis acting as a
sink for the excess ATP formed. Furthermore, low oxygen levels, which
suppress the cyanide insensitive pathway, are effective in suppressing sugar
accumulation at 1°C (Sherman and Ewing, 1982 and references therein).
Further research is required to establish any causal relationship between
cyanide insensitive respiration and sugar accumulation in cold-stored
tubers.
In many potato varieties the ratio of glucose to fructose in storage
approximates unity (Schwimmer et aI., 1954; Davies, et aI., 1989b) and
sucrose hydrolysis often accompanies reducing sugar accumulation in
storage (Richardson et aI., 1990). The evidence therefore favours sucrose
hydrolysis via the action of invertase rather than sucrose synthase. Sasaki et
al. (1971) demonstrated five forms of invertase in cold-stored tubers, three
forms showing lower activity in tubers stored at 20°C. In vitro the activity
of invertase is affected by the presence of a proteinaceous invertase
inhibitor (Pressey, 1966, 1969; Anderson and Ewing, 1978) which forms a
normally undissociable complex with the enzyme (Pressey, 1967). Pressey
and Shaw (1966) showed that during the initial period of cold treatment,
when reducing sugars were increasing rapidly, invertase formation pro-
ceeded until the level of the enzyme exceeded that of the inhibitor.
Transfer of tubers to higher temperature caused invertase activity to
decline and an excess of inhibitor to develop. The relationship between
invertase, invertase inhibitor and reducing sugar accumulation is not clear
cut, however. An analysis of 37 genotypes after 3 months' cold storage
revealed that reducing sugar content was not proportional to invertase
activity (Pressey, 1969). High sugar contents were associated with low
levels of inhibitor but low sugars not necessarily with high inhibitor levels.
Richardson et al. (1990) monitored changes in invertase activity in storage
for three genotypes known to differ in their susceptibility to low temperature
sweetening. The patterns of change in reducing sugar content during
storage at lOoC were, in general, related to changes in acid invertase
activity. Total invertase activity, i.e. measured after destroying the
endogenous inhibitor, reflected sugar changes more closely than did basal
activity (see also Davies et al., 1989a,b). However, genotypic variation in
the extent of hexose accumulation was not always paralleled by significant
differences in total invertase activity. Furthermore, transfer of tubers from
lOoC to 3°C had little effect on total activity whilst hexose content
increased substantially. What the study did reveal, however, was that both
total invertase activity and tuber hexose content increased markedly
following harvest and transfer to storage at lOoC. A similar phenomenon
occurs in rapidly growing, immature tubers, following excision from the
mother plant. Whether or not substantial increases in reducing sugars
during storage would occur if this initial increase in invertase activity were
suppressed remains to be established.
Since 1982 the Institute for Storage and Processing of Agricultural
Produce (IBVL), Wageningen, has been conducting trials on new potential
processing cultivars (Meijers, 1983). In the UK breeders have so far been
unsuccessful in producing an alternative to the cv. Record as have their
contemporaries in Europe and North America where cvs Bintje and
Russett Burbank still predominate (Beukema and van der Zaag, 1979), In
attempts to breed new improved processing cultivars, breeders have now
begun to examine the possibility of introducing low sweetening and long
dormancy characters from wild species into the cultivated form Solanum
tuberosum (Anon., 1987; Thomson et al., 1987). It is the case, however,
that variation in the low sweetening characteristic already exists within S.
tuberosum and may already be available to exploit in advanced breeders'
clones as potential cultivars or parental material in conventional breeding
programmes (Mackay et al., 1990). From the Scottish Crop Research
Institute breeding programme at least one genotype superior to cv. Record
has emerged. This genotype, previously known as clone 13737 1 and now
named Brodick, shows sufficient yield potential, disease resistance and
generally good agronomic characteristics to be considered as a possible
successor to cv. Record (Fig. 12.6).
Increased accumulation of hexoses during rapid sprout growth is well
documented (Hughes and Fuller, 1984 and references therein). However,
an irreversible accumulation of reducing sugars can occur in unsprouted
tubers following prolonged storage at high temperatures. This is referred
to as 'senescent sweetening' (Burton, 1966) and is characterized by changes
in the amyloplast membrane resulting in a breakdown of cellular
compartmentation and metabolic control (Isherwood and Burton, 1975;
Isherwood, 1976). The rate and extent of senescent sweetening depends on
variety, together with the history of the crop during growth and at the time
of lifting (Dwelle and Stallknecht, 1978). There is also a connection
between this type of sweetening and sprouting potential (Burton, 1966). If
546 Tuber quality
cv. Record
2.0
~\
J\ ,
1.5
,i "\"\ .........._...........1I--.... _-.... __ .... l
i
• /
Is.........
oIJ
.......
"
,l ..... 6
I
: ......"j'''
/.¥'/
0.5 ).---/------
i i i
o 20 40 60 80 100 120 140
Days in storage
cv. Brodic1<
2.0
1.5
I!?
IU_
g>"i
"'.<::
",,,,
<=" 1.0
·O.t::
"01<
al-
po..........••••••a •••······.a····-..
/ "":.;::~...:·. ..... . ..,.
II:
1li
0.5
/ /
,IS ./
~ /..-1
I .B a .,./. •
iii
o 20 40 60 80 100 120 140
Days in storage
Figure 12.6 Genotypic variation in tuber reducing sugar content during storage at
lOoC ( 0 ) and 3°C ( 0 ) and following transfer from lOoC to 3°C ( b. ). (From
Richardson et aI., 199O.)
sprouting is inhibited by chemical or physical means sugar accumulation is
accentuated, possibly because of reduced demand from growing sprouts
(Burton, 1966). This may also explain accelerated senescent sweetening in
irradiated tubers.
12.5.2 Dehydration
Dehydrated granules are used to prepare a reconstituted mash on the
addition of water. Manufacturers require tubers with above 20% DM to
minimize their energy costs for dehydration to granules with moisture
levels of 6-7% (Talburt et al., 1987.) Greened potatoes and the presence of
bacterial rots which may taint the product are the least desirable defects in
the raw material. Granule production has traditionally utilized outgrade
material, but with the development of more sophisticated markets a wider
range of processed food and snack products are being produced from
potato granules. Where a cooking stage is involved and product colour is
an important consideration, low levels of reducing sugars are specified in
the raw potato «0.25%) and the granule.
Dried potato flakes are produced from a precooked mash which is dried
on steam-heated rollers (Willard et al., 1987). With the dehydrated
products maintenance of cell wall integrity is a primary consideration as
cell wall fracture leads to starch leakage and an unacceptably sticky mash
after reconstitution. Primary causes for deterioration of dehydrated pro-
ducts are the non-enzymic browning reaction between amino acids and
reducing sugars (Burton et al., 1962a,b, 1963a,b; McWeeny and Burton,
1962) and oxidation of lipids, particularly linoleic and linolenic acids to
give rancid off-flavours, which are associated with the level of hexanal
produced (Buttery, 1961; Buttery et al., 1961). Sodium metabisulphite is
usually used to prevent browning during drying and storage and the
possible mechanisms for inhibition are reviewed by Wedzicha (1984).
Antioxidants such as butylated hydroxyanisole (BHA) and butylated
hydroxy toluene (BHT) are also added to retard rancidity.
12.5.3' Canning
Canned potatoes are prepared in several forms, including whole, diced and
sliced, but by far the largest proportion is of whole new and small potatoes.
However, the total tonnage canned in the UK is limited (c. 9000 t in 1988)
and market developments appear to be linked to finding improved cultivars
which will give more presentable and palatable products.
There are three main types of usage (Potato Marketing Board, 1988): diced
used as an ingredient in soups, salads, stews, etc.; small whole potatoes
graded from suitable maincrop cultivars canned for the catering trade, and
canned new potatoes. Several speciality potato products e.g. rosti, julienne
or shoestring potatoes (Smith and Davis, 1968) may also be canned.
548 Tuber quality
For all purposes, low incidence of damage, bruising, greening and
blemishes are desirable and tubers should have shallow eyes to minimize
peeling, trimming and rejection losses. However, the shape and size of
tubers required vary. Oblong tubers of 50 mm are preferred for diced,
whereas for canned small and new potatoes, round tubers of 20-40 mm are
preferred, with cvs Maris Peer and Arran Comet the most often used in
the UK for the latter purpose (Potato Marketing Board, 1989). The dry
matter content should be below 20% and be evenly distributed in the
tuber. Following cooking the potatoes should be free from after-cooking
blackening and disintegration, with a firm waxy texture. To minimize
sloughing CaCh is often used during processing as a firming agent. In the
finished product total calcium should not exceed 0.05%. Details of the
canning process are described by Talburt (1987).
12.6 IRRADIATION
12.6.1 Effects on sugar metabolism

The literature is often conflicting on the effects of irradiation on the sugar
content of tubers, possibly reflecting genotypic variation in response,
length of time between harvest and treatment and subsequent storage
regimes imposed (Thomas, 1984). y-irradiation is reported to increase
sucrose content (Becker and Somogyi, 1977), the level remaining high with
high irradiation doses. Doses in the region of 2-3 kGy are reported to
produce maximum accumulation (Hayashi and Aoki, 1985 and references
therein). Burton et al. (1959) observed a marked increase in sucrose with
exposures to 10 krad and storage at 10°C. Maximum accumulation
occurred 5 days after treatment but by day 26 sucrose levels had returned
to normal. There was also a temporary rise in reducing sugars, the increase
being greater the later the date of irradiation.
Jaarma (1958) reported that in tubers irradiated with 4-10 krad sucrose
and glucose increased while fructose decreased. These changes were again
reversible. There are also observations of a reduction in sucrose content
following irradiation (Filep and Koposztassy, 1971).
There have been several comparative studies on the processing qualities
of tubers following irradiation treatment under actual commercial storage
and processing conditions. In a large scale operation using a 60Co
irradiation 400 t potatoes were treated and successfully processed into
chips, mashed flakes, French fries and prepeeled fresh boilers (Thomas,
1984). In other studies Kennebec tubers iradiated with 8 krad and stored
for 8 months gave no undesirable effects on chipping quality (Gardner
and MacQueen 1965). However, Storey and Shackley (1987b) reported
that processed products from Pentland Dell, Record and Maris Piper
irradiated with 6 or 10 krad were not as acceptable as from non irradiated
Irradiation 549
tubers and a decline in the overall suitability for processing occurred
with both irradiated and non-irradiated samples stored at lOoC over a 6
month period. Freund (1965) examined the suitability of irradiated and
chemically sprout-inhibited Russet Burbank potatoes and found that both
treatments produced generally comparable processed products. However,
evaluation of these products showed that after 6 months' storage of the
final product only French fries made from irradiated tubers stored for 8
months after harvest were still commercially acceptable.
In an attempt to understand some of the changes induced by irradiation
many researchers have investigated the effects of irradiation on carbo-
hydrate metabolizing enzymes (Thomas, 1984 and references therein).
Ussuf and Nair (1972) showed that phosphorylase activity was increased
within 2 h of irradiation (10 krad) , but doses as high as 500 krad decreased
activity. Becker and Somogyi (1977) demonstrated that the increase in
sucrose after irradiation was related to a rise in phosphorylase activity
ranging from 147 to 355%, depending on variety. Increases in phospho-
glucomutase, UDP-glucose-pyrophosphorylase, sucrose synthase and
sucrose phosphate synthase have also been reported (Rubin and Metlisky,
1958; Hayashi and Aoki, 1985 and references therein).
12.6.2 Other effects
(a) Inhibition of sprout growth

Irradiation with doses in the range 7-lO krad inhibits sprouting irreversibly,
regardless of cultivar and storage temperature (Thomas, 1984). The best
results are obtained with good quality tubers, sufficiently cured to heal
bruises and other wounds. Handling should be minimized because wound
periderm formation is impaired after irradiation. Optimal storage tempera-
ture is in the region of 7-lO°C. Long term storage of irradiated tubers at
high temperatures characteristic of the tropics is not possible as spoilage
due to soft rots occurs very quickly (Thomas et al., 1978).
(b) Greening and glycoalkaloids

There are several reports on the effects of y-irradiation on greening and
glycoalkaloid production in tubt!rs exposed to light (Thomas, 1984). In
general, tubers exposed to y-rays are more resistant to light-induced
greening. It appears that with irradiation greening can be delayed for 9
days (Jandrell, 1979). Since, under normal conditions, greening starts to be
a problem after 3 days in the supermarket, this treatment prolongs shelf
life considerably. Doses of 5, 15, 50 and 250 krad inhibited chlorophyll
synthesis by 62, 67, 75 and 80%, respectively in tubers of Russett Burbank
illuminated for 3 days after irradiation (Schwimmer and Weston, 1958).
Others (Patil et al., 1975) have confirmed significant inhibition with lO krad
550 Tuber quality
but failed to show any significant effect on glycoalkaloid production.
Similarly, Wu and Salunkhe (1971) reported that 200 krad of y-irradiation
failed to inhibit light-induced glycoalkaloid production. However,
synthesis of glycoalkaloids induced by wounding or mechanical damage
was inhibited by 25-200 krad.
(c) Respiration
An immediate effect of ionizing radiation is an apparently transient
increase in respiration rate (Ojima et al., 1970; Ussuf and Nair, 1972). This
increase has been attributed to an increased oxidation of substrate coupled
to the phosphorylation of ADP (Jaarma, 1967). There is also evidence that
increased respiration immediately after irradiation is due to the cyanide
insensitive pathway (Chachin and Iwata, 1981).
(d) Nutritional quality

A study of the nutritional quality of tuber proteins has shown no significant
effects on digestibility or protein biological value due to irradiation with 8
krad (Varela and Gurbano, 1971). However, proline, glutamate and y-
aminobutyric acid (GABA) levels are increased (Thomas, 1984). In
contrast, Ussuf and Nair (1972), analysing the free amino acid pool, found
a decrease in glutamate and proline, together with methionine and
phenylalanine 24 h after exposure to 10 krad. Aspartate, asparagine,
threonine, serine, alanine, isoleucine, leucine, lysine and arginine all
increased. Lysine increased six-fold after 1 week and remained high even
after 1 month. Some attention has been paid to the effects of irradiation on
GABA levels as this is a well known inhibitory neurotransmitter in the
human brain. Results are again conflicting, Jaarma (1969) reporting
increases in GABA following irradiation and Satyanarayan and Nair
(1986) reporting no significant changes (despite a three-fold increase in the
operation of the 4-aminobutyrate shunt 2 days after irradiation). As far as
vitamins are concerned a reduction in ascorbic acid content has been
observed during the early storage period following irradiation. However,
the content after prolonged storage appears to be comparable to, or even
higher than, that in non-irradiated tubers stored under the same conditions
(Salkova, 1957). Levels of vitamins Bl and B2 also appear to remain
unaffected (Thomas, 1984).
(e) Wound healing

Increased susceptibility of irradiated tubers to rotting has been related to
the inability to produce wound periderm (Thomas, 1984). The capacity for
normal periderm development and suberization following exposure to low
doses, although initially retarded, appears to recover within a few weeks
Concluding comments 551
(Brownell et al., 1957). Doses as high as 8-10 krad completely destroy the
capacity to produce wound periderm and, while suberization is not
completely inhibited, it is retarded (Thomas, 1984). In the absence of
wound periderm pathogens still enter the tuber inducing a faster develop-
ment of rot. Gamma irradiation, up to 10 krad, does not appear to affect
the first important event in the development of a wound reaction i.e.
formation of quinones from phenolic acids, but does reduce the capacity
of tubers to synthesize chlorogenic acid (Pendharkar and Nair, 1987).
Treatment impairs the activity of cinnamic acid-4 hydroxylase. It has been
suggested that after harvest tubers should be stored for 2 weeks at is-20°C
with high relative humidity and good ventilation to induce rapid wound
healing before irradiation is used to prevent sprouting (Thomas, 1984).
12.7 CONCLUDING COMMENTS
The quality of the potato is associated with its Itutritional value and its
suitability for a wide range of cooking purposes. The factors affecting the
characteristics contributing to quality are various and influenced to differ-
ing degrees by the genotype and by environmental and cultural conditions
and the interactions between them. The extent to which different attributes
are affected depends upon the character being considered - for example
tuber shape and skin and flesh colour are largely determined by genotype -
and it is relatively easy to exercise a degree of control over them in a
breeding programme. Other quality characteristics such as dry matter
content, texture and enzymic and non-enzymic browning are affected by
both environmental and cultural factors and this is often complicated by
interactions between these and the cultivar. Consequently it is often
difficult to specify cultural practices which give consistent quality.
However, as much as anyone quality characteristic is the composite of
different factors, elements from different scientific disciplines are able to
contribute to the greater understanding of the underlying structural,
physiological and biochemical principles which affect it. For example the
interactions between growth and development of the tuber, cultural and
environmental constraints on dry matter accumulation and the tubers'
carbohydrate metabolism are all important, particularly the latter, in
relation to fry colour of the processed product. This better understanding
of the complex interactions has coincided with the advent of genetic
manipulation and the identification of putative targets within the potato for
improvement of nutritional and other quality aspects, particularly relating
to sugar metabolism and processing quality.
The possible nutritional and economic benefits arising from the
successful adoption of this technology are enormous. This opportunity for
improvements in quality also coincides with another pressing challenge;
that of increasing environmental concern, most notably over use of N
552 Tuber quality
fertilizers and application of pesticides and sprout suppressants. These
have been a mainstay of economic production and maintenance of various
quality attributes for many years. If limitations are imposed on the use of
these substances, the understanding of the intrinsic factors affecting
quality, together with their interactions with cultivation methods and the
environment, become of much greater significance to the industry and the
consumer.
REFERENCES
Adler, G. (1971) Kartoffeln und Karoffelerseugnisse. Grundlagen und Fortschritte

der Lebensmitteluntersuchung, 13, Parey, Berlin.
Akeley, RV., Houghland, G.V.C. and Schark, A.E. (1962) Genetic differences in
potato tuber greening. Am. Potato J., 39, 409.
Allen, E.H. and Kuc, J. (1968) a-Solanine and a-chaconine as fungitoxic com-
pounds in extracts of Irish potato tubers. Phytopath., 58, 776.
Agric. Sci., Camb., 94, 583-606.
Amir, J., Kahn, V. and Unterman, M. (1977) Respiration; ATP level and sugar
accumulation in potato tubers during storage at 4°C. Phytochem., 16, 1495.
Amla, B.L. and Francis, F.J. (1961) Effect of dipping solution on the quality of
pre-peeled potatoes. Am. Potato J., 38, 121.
Anderson, R.S. and Ewing, E.E. (1978) Partial purification of tuber invertase and
its proteinaceous inhibitor. Phytochem., 17,1077.
Anon. (1983) National Advisory Committee on Nutritional Education (NACNE),
HMSO, London.
Anon. (1984) Committee on Medical Aspects of Food Policy (COMA), Diet and
Cardiovascular Disease, HMSO, London, 32 pp.
Anon. (1987) Cultivating our wild relations. Potato World, 4, 22.
ap. Rees, T., Burrell, M.M., Entwistle, T.G., Hammond, J.B.W., Kirk, D. and
Kruger, N.J. (1988) Effects of low temperature on the respiratory metabolism
of carbohydrates by plants, in Plants and Temperature (eds S.P. Long and F.1.
Woodward), S.E.B. Symp. 42, 377-93.
Arteca, R.N. (1982) Calcium infiltration inhibits greening in Katahdin potatoes.
HortSci., 17,79.
Arteca, R.N., Poovaiah, B.W. and Hiller, L.K. (1980) Electron microprobe and
neutron activation analysis for the determination of elemental distribution in
hollow heart potato tubers. Am. Potato J., 57, 241.
Augustin, J. (1975) Variations in the nutritional composition of fresh potatoes. J.
Food Sci., 40, 1259-99.
Augustin, J., Johnson, S.R., Teitzel, C., Toma, R.B., Shaw, R.L., True, R.H.,
Hogan, J.M. and Deutsch, RM. (1978) Vitamin composition of freshly
harvested and stored potatoes. J. Food Sci., 43,1566--74.
Baerug, R (1962) Influence of different rates and intensities of light on
solanine content and cooking quality of potato tubers. Eur. Potato J., 5,
242-51.
Balls, RC., Gunn, J.S. and Starling, A.J. (1982) The National Potato Damage
References 553
Awareness Campaign, Potato Marketing Board and Agricultural Development
and Advisory Service, Oxford, 32pp.
Barker, J. and Mapson, L.W. (1950) The ascorbic acid content of potato tubers. II.
The influence of temperature of storage. New Phytol., 49, 283-303.
Barrios, E.P., Newsom, D.W. and Miller, J.C. (1963) Some factors influencing the
culinary quality of Irish potatoes II. Physical characters. Am. Potato J., 40, 200--8.
Becker, D.P. and Somogyi, LC. (1977) The action mechanism of the increase in
saccharose in irradiated potatoes 1. Inter-relation between the increase in
saccharose content and phosphorylase activity after gamma irradiation. Mitt.
Gebitete Lebensm. Hyg., 68, 409.
Bettelheim, F.A. and Sterling, C. (1955) Factors associated with potato texture. II.
Pectic substances. Food Res., 20, 118--29.
Beukema, H.P. and van der Zaag, D.E. (1979) Potato Improvement, Some Factors
and Facts, LA.C., Wageningen.
Birch, J.H., Devine, J.R., Holmes, M.R.J. and Whitear, J.D. (1967) Field
experiments on the fertilizer requirements of maincrop potatoes. J. Agric. Sci.,
Camb., 69, 13--24.
Blight, D.P. and Hamilton, A.J. (1974) Varietal susceptibility to damage in
potatoes. Potato Res., 17,261-70.
Brandl, W. and Herrmann, K. (1984) Uber das Vorkommen der chlorogensauren
in der Kartoffel. Z. Lebensm. Unters. Forsch., 178(3), 192--4.
Brisson, N., Giroux, H., Zollinger, M., Camirand, A. and Simard, C. (1989)
Maturation and subcellular compartmentation of potato starch phosphorylase.
The Plant Cell, 1, 559.
Brown, E. and Riley, W. (1976) Greening of potato tubers: varietal response to
controlled exposure to light. J. Nat. Inst. Agric. Bot., 14, 70.
Brownell, L.E., Gustafson, F.G., Nehemias, J.V., Isleib, D.R. and Hooker, W.J.
(1957) Storage properties of gamma-irradiated potatoes. Food Technol.,
11, 306.
Burrell, M.M. (1984) Inhibition of browning, phenoxyacetic acids and phenolic
metabolism in potato tuber discs: a model system to study chemicals that control
common scab. Plant Pathol., 33, 325.
Burton, H.S., McWeeney, D.J. and Bintcliffe, D.O. (1962a) Development of
chromophores in the carbonyl-amino system. Nature, London, 196,40-1.
Burton, H.S., McWeeney, D.J., Pandhi, P.N. and Bintcliffe, D.O. (1962b)
Fluorescent compounds and non-enzymatic browning. Nature, London, 196,
948-50.
Burton, H.S., McWeeney, D.J. and Bintclife, D.O. (1963a) Sulphur dioxide and
ketose-amino reactions. Chemy Ind., 1963, 693-5.
Burton, H.S., McWeeney, D.J. and Bintcliffc, D.O. (1963b) Non enzymic
browning, the role of unsaturated carbonyl compounds as intermediates and of
S02 as inhibitor of browning. J. Sci. Fd Agric., 14, 911-20.
Burton, W.G. (1965) The sugar balance in some British varieties during storage. 1.
Preliminary observations. Eur. Potato J., 8, 80.
Burton, W.G. (1966) The Potato: A survey of its history and of the factors
influencing its yield, nutritive value, quality and storage, H. Veenman and
Sonen, N.V., Wageningen.
Burton, W.G. (1989) The Potato, 3rd edn, Longman, London, 742 pp.
Burton, W.G. and Wilson, A.R. (1970) The apparent effect of the latitude of the
554 Tuber quality
place of cultivation upon the sugar content of potatoes grown in Great Britain.
Potato Res., 13,269.
Burton, W.G. and Wilson, A.R. (1978) The sugar content and sprout growth of
tubers of potato cultivar Record, grown in different localities, when stored at
10,2 and 20°e. Potato Res., 21,145.
Burton, W.G., Horne, T. and Powell, D.B. (1959) The effect of gamma radiation
upon the sugar content of potatoes. Eur. Potato J., 2, 105.
Buttery, R.G. (1961) Autoxidation of potato granules, II. Formation of carbonyls
and hydrocarbons. J. Agric. Fd Chem., 9, 248-52.
Buttery, R.G. and Ling, L.C. (1973) Earthy aroma of potatoes. J. Agric. Fd
Chem., 21, 745-6.
Buttery, R.G., Hendel, C.E. and Boggs, M.M. (1961) Autoxidation of potato
granules. I. Changes in fatty acids. J. Agric. Fd Chem., 9, 245-8.
Carr, M.K. V. (1985) The role of water in the productivity of potatoes, in Irrigating
Potatoes. UK Irrigation Association Technical Monograph 2 (eds M.K.V. Carr
and P.J.C. Hamer), Cranfield Press, Silsoe, pp. I-II.
Carruthers, J. (1982) A comparison of impact damage levels from two shapes of
potato harvester web rods. Scottish Inst. Agric. Engng. Dept. Note SIN/329.
Casey, J.C., Self, R. and Swain, T. (1963) Origin of methanol and dimethyl sulfide
from cooked foods. Nature, London, 200, 855.
Chachin, K. and Iwata, T. (1981) Respiratory metabolism and potassium release of
irradiated potatoes. Paper No. IAEA-SR-60/15. Seminar on Food Irradiation
for Developing Countries in Asia and the Pacific, Tokyo.
Chick, H. and Slack, E.B. (1951) Biochem. J., 45, 211-2I.
Cochrane, P.M., and Duffus, e.M. (1989) Amylolytic activity in stored potato
tubers. Am. Potato J., 66, 510.
Cole, e.S. (1975) Variation in dry matter between and within potato tubers. Potato
Res., 18,28-37.
Collier, G.F., Wurr, D.C.E. and Huntington, V.e. (1978) The effect of calcium
nutrition on the incidence of internal rust spot in the potato. J. Agric. Sci.
Camb., 91, 24I.
Collier, G.F., Wurr, D.C.E. and Huntington, V.C. (1980) The susceptibility of
potato varieties to internal rust spot. J. Agric. Sci. Camb., 94, 407.
Conners, H.W. (1937) Effect of light on solanine synthesis in the potato tuber.
Plant Physiol., 12, 79.
Cowan, e.A. (1985) Potatoes - influence of age and regional differences on
preference for size. Irish J. Ag. Econ. Rural Sociol., 10(2),119-27.
Cromack, H.T.H. (1981) Purple discoloration of the flesh of potato tubers. ADAS,
Trawscoed.
Cronk, T.e., Kuhn, G.O. and McArdle, F.J. (1974) The influence of stage of
maturity, level of nitrogen fertilization and storage on the concentration of
solanine in tubers of three potato cultivars. Bull. Environ. Contamin. Toxicol.,
11, 163.
Cutter, E.G. (1978) Structure and development of the potato plant, in The Potato
Crop (ed. P.M. Harris), Chapman & Hall, London, pp. 70-152.
Davies, A.M.C. (1977) The free amino acids of tubers of potato varieties in
England and Ireland. Potato Res., 20, 9-2I.
Davies, A.M.C. and Blincow, P.J. (1984) Glycoalkaloid content of potatoes and
potato products sold in the UK. J. Sci. Fd Agric., 35, 553-7.
References 555
Davies, H.V. and Ross, H.A. (1986a) The development of internal rust spot in
potato tubers. Aspects of Applied Biology, 13, 433.
Davies, H.V. and Ross, H.A. (1986b) Hydrolytic and phosphorolytic enzyme
activity and reserve mobilisation in sprouting tubers of potato (Solanum
tuberosum L.). 1. Plant Physiol., 126, 387.
Davies, H.V. and Viola, R. (1988) The effect of gibberellic acid on starch
breakdown in sprouting tubers of Solanum tuberosum L. Annals Bot., 61, 689.
Davies, H.V., Richardson, D.L. and Ross, H.A. (1989a) Invertase and hexose
accumulation in stored tubers. Am. Potato 1., 66, 514.
Davies, H. V., Jefferies, R.A. and Scobie, L. (1989b) Hexose accumulation in cold-
stored tubers of potato (Solanum tuberosum L.): The effects of water stress. 1.
Plant Physiol., 134, 471.
Deahl, K.L., Young, R.J. and Sinden, S.L. (1973) A study of the relationship of
late blight resistance to glycoalkaloid content in fifteen potato clones. Am.
Potato 1., 50, 248-53.
Desborough, S.L. (1983) Potato (Solanum tuberosum L.), in lsozymes in Plant
Genetics and Breeding (eds S.D. Tunksley and T.J. Orton), Elsevier,
Amsterdam, Part B, pp. 167-88.
Desborough, S.L. (1985) Potato proteins, in Potato Physiology (ed. P.H. Li),
Academic Press, London, pp. 329-51.
Desborough, S. and Weiser, e.J. (1974) Improving potato protein. I. Evaluation of
selection techniques. Am. Potato 1., 51, 185-96.
Dickens, J.e., Harrap, F.E.G. and Holmes, M.R.J. (1962)1. Agric. Sci., Camb.,
59,319-26.
Dixon, W.L. and ap Rees, T. (1980) Carbohydrate metabolism during cold-
induced sweetening of potato tubers. Phytochem., 19, 1653.
Dixon, W.L., Franks, F. and ap Rees, T. (1981) Cold-lability of phosphofructo-
kinase from potato tubers. Phytochem., 20, 969.
Drew, J.P. and Dreasy, D. (1941) Potato growing in Ireland with particular
reference to production for industrial purposes. 1. Dep. Agric. Repub. Ire., 38,
220-38.
Dwelle, R.B. and Stallknecht, G.F. (1978) Respiration and sugar content of potato
tubers as influenced by storage temperature. Am. Potato 1., 55, 561.
Ehlenfeldt, M.K. and Boe, A.A. (1989) Respiration and sugar accumulation of
cold chipping and conventional potato clones. Am. Potato 1., 66, 519.
Elton, G.A.H. (1978) European diets in relation to standards of need, in Diet of
Men: Needs and wants (ed. J. Yudkin), Applied Science Publishers, London,
pp.25-40.
Eppendorfer, W.H., Eggum, B.O. and Bille, S.W. (1979) Nutritive value of potato
crude protein as influenced by manuring and amino acid composition. 1. Sci. Fd
Agric., 30, 361-8.
Filep, G. and Koposztassy, E. (1971) Effect of temperature and gamma irradiation
on changes in carbohydrate composition of some potato varieties during
storage. Noventermeles, 20, 289.
Finglas, P.M. and Faulks, R.M. (1984) Nutritional composition of UK retail
potatoes, both raw and cooked. 1. Sci. Fd Agric., 35, 1347-56.
Finney, E.E. and Findlen, J. (1967) Influence of pre-harvest treatment upon turgor
of Katahdin potatoes. Am. Potato 1., 44, 383.
Firbas, H. (1961) Beitrag zur selektion von im rohzustand nichtdunkelnder
556 Tuber quality
kartoffeln. Aus dem Max-Planck-Institutfur Zuchtungsforschung, 90th Birthday
von metternich, Koln-Vogelsang.
Fitzpatrick, T.J., Herb, S.F., Osman, S.F. and McDermott, J.E. (1977) Potato
glycoalkaloids: increases and variations of ratios in aged slices over prolonged
storage. Am. Potato J., 54, 539-44.
Fitzpatrick, T.J., McDermott, J.A. and Osman, S.F. (1978) Evaluation of injured
commercial potato samples for total glycoalkaloid contents. J. Food Sci., 43,
1617.
Forster, H. and Beringer, H. (1983) Starkegehalte von Kartoffelknollen
in Abhangigheit von Kalium-Ernahrung und Knollenentwicklung.
Z. Pfanzenerniihr. Bodenk., 146, 572-82.
Forsyth, F.R. and Eaves, e.A. (1968) Greening of potatoes. Food Tech., 22, 48.
Freund, G.A. (1965) Suitability o.f potato products prepared from irradiated
and chemically inhibited potatoes. Report IDO-l0042, Western Nuclear
Corporation, US Atomic Energy Commission, Washington DC.
Fuller, T.J. and Hughes, J.C. (1984) Factors influencing the relationships between
reducing sugar and fry colour of potato tubers of cv. Record. J. Food Technol.,
19,455.
Gardner, D.S. and MacQueen, K.F. (1965) Effect of gamma rays Dn storage life
and chipping qualities of Ontario grown Kennebec potatoes.. Potato Chipper,
24,38.
Geddes, R., Greenwood, C.T. and MacKenzie, S. (1965) Studies on the bio-
synthesis of starch granules III. The properties of the components of starches
from the growing potato tuber. Carbohydrate Res., 1, 71.
Glasbey, C.A., McRae, D.C. and Flemi:ng, J. (1988) The size distribution of potato
tubers and its application to grading schemes. Ann. Appl. Bioi., 113, 579-87.
Glynne, M.D. and Jackson, V.G. (1919) The distribution of dry matter and
nitrogen in the potato tuber. Variety King Edward. J. Agric. Sci., Camb., 9,
237-58.
Gould, W.A. and Plimpton, S. (1985) Quality evaluation of potato cultivars ~or
processing. Ohio Agricultural and Research Centre, Research Bulletin 1172,
Ohio State Univ., Wooster, 25pp.
Grant, A. and Hughes, J.C. (1985) The relationship between physical properties of
tubers measured during pendulum impact tests and tuber fracture damage.
Potato Res., 28, 203-13.
Gray, D. (1972) Some effects of variety, harvest date and plant spacing on tuber
breakdown on canning, tuber dry matter content and cell surface area in the
potato. Potato Res., 15, 317-34.
Gray, D. and Hughes, J.C. (1978) Tuber quality, in The Potato Crop (ed. P.M.
Harris), Chapman & Hall, London, pp. 504-44.
Groot, E.H., Jansen, L.W., Kentie, A., Oosterhuis, H.K. and Trap, H.L.J. (1947)
A new protein in potatoes. Biochim. et Biophys. Acta, 1,410--14.
Gubb, I., Callow, J.A., Faulks, R.M. and Jackson, M.T. (1989a) The biochemical
basis for the lack of enzymic browning in the wild potato species Solanum
hjertingii Hawkes. Am. Potato J. , 66, 522.
Gubb, I.R., Hughes, J.e., Jackson, M.T. and Callow, J.A. (1989b) The lack of
enzymic browning in the wild potato species Solanum hjertingii Hawkes
compared with commercial Solanum tuberosum varieties. Ann. Appl. Bioi.,
114, 579.
References 557
Gull, D.D. and Isenberg, F.M. (1960) Chlorophyll and solanine content and
distribution in four varieties of potato tubers. Proc. Am. Soc. Hart. Sci., 75, 545.
Haddock, J.L. (1961) The influence of irrigation regime on yield and quality of
potato tubers and nutritional status of plants. Am. Potato J., 38, 423-34.
Harborne, J.B. (1960) Plant polyphenols. I. Anthocyanin production in the
cultivated potato. Biochem. J., 74, 262-9.
Harkett, P.J. (1975) The influence of temperature and skin colour on chlorophyll
synthesis in potato tubers exposed to light. Proc. 6th Trienn. Conf. Eur. Assoc.
Potato Res., Wageningen, p. 179.
Harris, P.M. (1978) Mineral nutrition, in The Potato Crop (ed. P.M. Harris),
Chapman & Hall, London, pp. 195-243.
Harward, M.E., Jackson, W.A., Nielson, L.W., Pilanol, J.R. and Mason, D.D.
(1956) Effects of soil moisture tensions and of chloride and sulphate treatments
on the yield, composition and bacterial soft-rot of Irish potatoes. Proc. Soil Sci.
Soc. Am., 20, 59--{)3.
Hayashi, T. and Aoki, S. (1985) Effect of irradiation on the carbohydrate
metabolism responsible for sucrose accumulation in potatoes. J. Agric. Fd
Chern., 33,14.
Heisler, E.G., Siciliano, J., Treadway, R.H. and Woodward, C.F. (1963) After-
cooking discolouration of potatoes. Iron content in relation to blackening
tendency of tissue. J. Food Sci., 28, 453-9.
Hiller, L.K. and Koller, D.C. (1984) Effect of early season soil moisture levels and
growth regulator applications on internal quality of Russett Burbank potatoes.
Proc. Wash. State Potato Conf., 23, 67.
Hiller, L.K., Koller, D.C. and van Denburgh, R.W. (1979) Brown center of
potatoes - what have we learned? Proc. Wash. State Potato Conf., 18, 21.
Hiller, L.K., Koller, D.C. and Thornton, R.E. (1985) Physiological disorders of
potato tubers, in Potato Physiology (ed. P.H. Li), Academic Press, London, pp.
389-455.
Hilton, R.J. (1951) Factors in relation to tuber quality in potatoes. II. Preliminary
trials on bitterness in Netted Gem potatoes. Can. J. Agric. Sci., 31, 61.
Hippe, J. (1988) HPLC-analysis of the concentrations of free asparagine and
glutamine in potatoes grown with varying amounts of nitrogen. Potato Res., 31,
535-40.
Hoff, J.E. (1972) Starch swelling pressure of cooked potatoes. J. Agric. Fd Chern.,
20, 1283-4.
Hoff, J.E. and Castro, M.D. (1969) Chemical composition of potato cell wall. J.
Agric. Fd Chern., 17, 1328--31.
Hogge, M.C. (1989) Effects of site, season and husbandry on yield and processing
quality of potato variety Pentland Dell, Ph.D. Thesis, University of Cambridge,
208 pp.
Hooker, W.J. (ed.) (1981) Compendium of Potato Diseases, Am. Phytopathol.
Soc., St. Paul, Minnesota.
Horton, D.E. and Fano, H. (1985) Potato Atlas, CIP, Lima. 135 pp.
Horton, D. and Sawyer, R.L. (1985) The potato as a world food crop, with special
reference to developing areas, in Potato Physiology (ed. P.H. Li), Academic
Press, London, pp. 1-34.
Houghland, G.V.c. (1966) New conversion table for specific gravity, dry matter
and starch in potatoes. Am. Potato J., 43, 138.
558 Tuber quality
Howard, H.W. (1978) The production of new varieties, in The Potato Crop (ed.
P.M. Harris), Chapman & Hall, London, pp. 607-46.
Huaman, Z., Williams, J.T., Salhluana, W. and Vincent, L (1977) A list of
descriptors for the cultivated potato. Report of Planning Conference on
Utilization of the Genetic Resources of the Potato II, CIP, Lima.
Hudson, D.E. (1975) The relationship of cell size, intercellular space, and specific
gravity to bruise depth in potatoes. Am. Potato 1., 52, 9-14.
Hughes, J.e. (1980) Role of tuber properties in determining susceptibility of
potatoes to damage. Ann. Appl. Bioi., 96, 344-5.
Hughes, J.e. and Evans, J.L (1967) Studies on after-cooking blackening in
potatoes. IV Field experiments. Eur. Potato 1., 10, 16-36.
Hughes, J.C. and Faulks, R.M. (1973) Texture of cooked potatoes of different
maturity in relation to pectic substances and cell size. Proc. 5th Triennial EAPR
Conference, 1972, p. 163. ,.
Hughes, J.e. and Fuller, T.J. (1984) Fluctuations in sugars in cv. Record during
extended storage at lO°e. Potato Res., 27, 229.
Hughes, J.C., Grant, A. and Faulks, R.M. (1975a) Susceptibility of tubers to
internal bruising, Potato Res., 18, 338-9.
Hughes, J.C. and Mapson, L.W. (1966) Potatoes for the food processing industry.
Biochemical studies. Food Trade Review, 34, 43-9.
Hughes, J.e. and Swain, T. (1962a) After-cooking blackening in potatoes. II. Core
experiments. 1. Sci. Fd Agric., 13, 229-36.
Hughes, J.C. and Swain, T. (1962b) After-cooking blackening in potatoes. III.
Examination of the interaction of factors by in vitro experiments. 1. Sci. Fd
Agric., 13, 358-63.
Hughes, J.e., Faulks, R.M. and Grant, A. (1975b) Texture of cooked potatoes.
Relationship between compressive strength, pectic substances and cell size of
Redskin tubers of different maturity. Potato Res., 18, 495-514.
Hughes, J.e., Faulks, R.M. and Grant, A. (1975c) 1. Sci. Fd Agric., 26, 731-8.
Texture of cooked potatoes: relationship between compressive strengths of
cooked potato discs and release of pectic substances.
Hughes, J.C., Grant, A. and Faulks, R.M. (1975d) Texture of cooked potatoes:
the effect of ions and pH on the compressive strength of cooked potatoes. 1. Sci.
Fd Agric., 26, 739-48.
Hughes, J.e., Grant, A., Prescott, E.H.A., Pennington, D.E. and Worts, W.H.
(1985) A portable pendulum for testing dynamic tissue failure susceptibility of
potatoes. 1. Agric. Engng. Res., 32, 269-77.
Hunnius, W. and Munzert, M. (1972) Z. Acker-u Pft Bau., 136, 245-60.
Hunnius, W., Butcher, G. and Munzert, M. (1972) Zum Einflus des Stickstoffs
auf die Vollerntevertraglich keit der Kartoffelknodle. Potato Res., 15,
54-66.
Hutchinson, A. and Hilton, R.J. (1955) The influence of certain cultural practices
on solanine content and tuber yields in Netted Gem potatoes. Can. 1. Agric.
Sci., 35, 485.
Hutchison, P.S. and McRae, D.C. (1987) The development of a hollow web rod for
potato harvesters. Scottish Inst. Agric. Engng., Dept Note SIN/483.
Ifenkwe, D.P., Allen, E.J. and Wurr, D.C.E. (1974) Factors affecting the
relationship between tuber size and dry-matter content. Am. Potato 1., 51,
233-42.
References 559
Iritani, W.M. (1981) Growth and preharvest stress and processing quality of
potatoes. Am. Potato J., 58, 71.
Iritani, W.M. and Weller, L.D. (1978) Factors influencing reconditioning of Russet
Burbank potatoes. Am. Potato J., 55, 425-30.
Iritani, W.M., Powers, M.J., Hudson, L. and Weller, L. (1977) Factors influencing
time to breakdown (TTB) of cooked potato tissues. Am. Potato J., 54, 23-32.
Isherwood, F.A. (1973) Starch-sugar interconversions in Solanum tuberosum.
Phytochem., 12, 2579.
Isherwood, F.A. (1976) Mechanisms of starch-sugar interconversioJil in Solanum
tuberosum. Phytochem., 15, 33.
Isherwood, F.A. and Burton, W.G. (1975) The effect of senescence, handling,
sprouting and chemical sprout suppression upon the respiratory quotient of
stored potato tubers. Potato Res., 18, 98.
Iwanzik, W., Tevini, M., Stute, R. and Hilbert, R. (1983) Carotinoidgehalt und-
zusammensetzung verschiedener deutscher Kartoffelsorten und deren
Bedautung fur die Gleischfarbe der Knolle. Potato Res., 26, 149-62.
Jaarma, M. (1958) Influence of ionizing radiation on potato tubers. Arkiv. Kemi.,
13,97.
Jaarma, M. (1967) Influence of ionizing radiation on oxidative phosphorylation,
adenosine triphosphatase, and apyrase in potato tubers. Acta Chem. Scand., 21,
2069.
Jaarma, M. (1969) Comparison of chemical changes in potato tubers induced by
gamma irradiation and by chemical treatment. Acta Chem. Scand., 23, 3435.
Jackson, T.L., McBride, R., Powelson, M.L. et al. (1984) Soil fertility, plant
nutrition, and plant disease interactions affecting potatoes. Oregon State Univ.
Agric. Exp. Stn Spec. Rept July, pp. 34-7.
Jadhev, S.J. and Salunkhe, D.K. (1974) Effects of certain chemicals on photo-
induction of chlorophyll and glycoalkaloid synthesis and on sprouting of potato
tubers. J. [nst. Can. Sci. Technol. Aliment., 7, 178.
Jadhev, S.J. and Salunkhe, D.K. (1975) Formation and control of chlorophyll and
glycoalkaloids in tubers of Solanum tuberosum and evaluation of glycoalkaloid
toxity. Adv. Food Res., 21, 307-54.
Jadhev, S.J., Sharma, R.P. and Salunkhe, D.K. (1981) Naturally oc.curring toxic
alkaloids in foods. CRC Crit. Rev. Toxico!., 9, 21-104.
Jandrell, A.c. (1979) Potato production in South Africa and the potential use of
irradiation in the potato industry. Proc. Nat! Symp. Food [rradiat., Pretoria, S.
Africa, p. 14.
Jarvis, M.C., Hall, M.A., Threlfall, D.R. and Friend, J. (1981) The polysaccharide
structure of potato cell walls: Chemical fractionation. Planta, 152, 93-100.
Jefferies, R.A. and MacKerron, D.K.L. (1986) Tuber dry matter concentration of
potato cultivars in relation to soil moisture. Aspects of Applied Biology, 13,
425-7.
Jefferies, R.A. and MacKerron, D.K.L. (1987) Observations on the incidence of
tuber growth cracking in relation to weather patterns. Potato Res., 30, 613.
Johnston, F.B., Hoffman, 1. and Petrasovits, A. (1968) Distribution of mineral
constituents and dry matter in the potato tuber. Am. Potato J., 45,
287-92.
Joslyn, M.A. and Ponting, J.D. (1951) Enzyme catalysed oxidative browning of
fruit products. Adv. Food Res., 3, 1.
560 Tuber quality
Kaldy, M.S. and Lynch, D.R. (1983) Chi orogenic acid content of Russet Burbank
potato in Alberta. Am. Potato 1., 60, 375-7.
Kallio, A. (1960) Effect of fertility level on the incidence of hollow heart. Am.
Potato 1., 37, 338.
Kapoor, A.C., Desborough, S.L. and Li, P.H. (1975) Potato tuber proteins and
their nutritional quality. Potato Res., 18, 469-78.
Keeling, P.L., Wood, J.R., Tyson, R.H. and Bridges, I.G. (1988) Starch bio-
synthesis in developing wheat grain. Evidence against the direct involvement of
triose phosphates in the metabolic pathway. Plant Physiol., 87, 311.
Keijbets, M.J.H. (1974) Pectic substances in the cell wall and the intercellular
cohesion of potato tissue during cooking. IBVL Publication 275, Wageningen.
Keijbets, M.J.H. (1984) Vitamin C during long-term storage of potatoes. Abst. 9th
Triennial EAPR Conference, 1984, pp. 390-1.
Kerr, S.P. (1986) Potato skin texture from NIAB trials. Aspects of Applied
Biology, 13, 409-12.
Killick, R.J. and Simmonds, N.W. (1974) Specific gravity of potato tubers as a
character showing small genotype-environment interactions. Heredity, 32,
109-12.
Kirkman, M.A. (1986) What the market wants: the processing sector, in Pottkto
Quality: The production of potatoes to meet market requirements, Potato
Marketing Board, Oxford, pp. 26-32.
Klein, L.B., Chandra, S. and Mondy, N.I. (1980) The effect of phosphorous
fertilization on the chemical quality of Katahdin potatoes. Am. Potato 1., 57,
259-66.
Kruger, N.J., Hammond, J.B.W. and Burrell, M.M. (1988) Molecular characteris-
ation of four forms of phosphofructokinase purified from potato tuber. Arch.
Biochem. Biophys., 267, 690.
Kunkel, R. and Gardner, W.H. (1959) Blackspot of Russet Burbank potatoes.
Proc. Am. Soc. Hart. Sci., 73, 436-44.
Kunkel, R. and Gardner, W.H. (1965) Potato tuber hydration and its effects on
blackspot of Russet Burbank in the Columbia Basin of Washington. Am. Potato
1., 42, 109-24.
Kunkel, R. and Holstad, N.M. (1972) Potato chip colour, specific gravity and
fertilization of potato with N-P-K. Am. Potato 1., 49, 43-62.
Kunkel, R., Holstad, N.M. and Russell, T.S. (1973) Mineral element content of
potato plants and tubers vs. yields. Am. Potato 1., 50, 275-82.
Kushman, J.J. and Hayes, F.L. (1971) Influence of intercellular space differences
due to variety and storage upon tuber specific gravity - dry matter relationships.
Am. Potato 1., 48, 173-81.
Lampe, K. (1960) Die Widerstandsfahigkeit von Kartoffelknollen gegan
beschadigungen. Eur. Potato 1.,3, 13-29.
Lampitt, L.H. and Goldenberg, N. (1940) The composition of the potato. Chemy
Ind., 1940, pp. 748-61.
Lampitt, L.H., Baker, L.c. and Parkinson, T.L. (1945a) Vitamin-C content of
potatoes I. Distribution in the potato plant. 1. Soc. Chem. Ind., 64, 18-22.
Lampitt, L.H., Baker, L.c. and Parkinson, T.L. (1945b) Vitamin-C content of
potatoes II. The effect of variety, soil and storage. 1. Soc. Chem. Ind., 64, 22-6.
Larsen, E.J. (1949) Investigations on cause and prevention of greening of potatoes.
Bull. Idaho Agric. Exp. Stn, No. 16.
References 561
Leichsenring, J.M., Norris, L.M. and Pilcher, H.L. (1957a) Ascorbic acid contents
of potatoes. I. Effect of storage and of boiling on the ascorbic, dehydroascorbic
and diketogulonic acid contents of potatoes. Food Res., 22, 37--43.
Leichsenring, J.M., Pilcher, H.L. and Norris, L.M. (1957b) Ascorbic acid contents
of potatoes. II. Effects of baking and pressure cooking on the ascorbic, dehydro-
ascorbic and diketogulonic and contents of potatoes. Food Res., 22, 44-50.
Lepper, W. (1949) Solaningehalte von 58 Kartoffelsorjten. Z. Lebensm. Unter. 89,
264-73.
Lerner, A.B. (1953) Metabolism of phenylalanine and tyrosine. Adv. in Enzymol.,
14,73.
Lewis, W.C. and Rowberry, R.G. (1973) Some effects of planting depth and time
and height of hilling on Kennebec and Sebago potatoes. Am. Potato J., 50, 301.
Li, P.H. and Sayre, K.D. (1975) The protein, non-protein and total nitrogen in
Solanum tuberosum ssp. andigena potatoes. Am. Potato J., 52, 341-50
Liljemark, A. and Widoff, E. (1960) Greening and solanine development of white
potato in fluorescent light. Am. Potato J., 37,379-89
Linehan, D.J. and Hughes, J.C. (1969a) Texture of cooked potato. II.
Relationship between intercellular adhesion and ch.emical composition of the
tuber. J. Sci. Fd Agric., 20,113-19.
Linehan, D.J. and Hughes, J.e. (1969b) Texture of cooked potato I - Introduction. J.
Sci. Fd Agric., 20, 11G-12.
Linehan, D.J., Stooke, e.E. and Hughes, J.C. (1968) The importance of cell size in
influencing the texture of the cooked potato. I. Preliminary observations. Eur.
Potato J., 11,221-5.
Lugt, e. (1961) Results of the assessment of the cooking quality of internationally
exchanged potato samples. Proc. 1st Triennial Conf. Eur. Assn Potato Res., pp.
321-3.
Lynch, D.R. and Kaldy, M.S. (1985) Citric acid and potassium contents of Russet
Burbank potato in Alberta. Can. J. Plant Sci., 65, 793-5.
MacKay, G.R., Brown, J. and Torrance, e.J.W. (1990) The processing potential
of tubers of the cultivated potato, Solanum tuberosum, after storage at low
temperature. 1. Fry colour. Potato Res., 33, 211-18.
MacKerron, D.K.L. (1985) Timing of irrigation in relation to yield and quality of
potatoes, in Irrigating Potatoes. UK Irrigation Association Technical
Monograph 2 (eds M.K.V. Carr and P.J.C. Ham,er), Cranfield Press, Silsoe, pp.
54-60.
MacKerron, D.K.L. and Jefferies, R.A. (1985) Observations on the relief of late
water stress in potato. Potato Res., 28, 349.
MacKerron, D.K.L. and Jefferies, R.A. (1986) The influence of early soil moisture
stress on tuber numbers in potato. Potato Res., 29, 299-312.
Maga, J .A. (1980) Potato glycoalkaloids. CRC Crit. Rev. Food Sci. Nutr., 12,
371--405.
Maine, M.J. De (1986) A method of testing the resistance of potato cultivars to
tuber damage caused by squeezing. J. Agric. Sci., Camb., 106, 255-8.
Mapson, L.W., Swain, T. and Tomalin, A.W. (1963) Influence of variety, cultural
conditions and temperature of storage on enzymic browning of potato tubers. J.
Sci. Fd Agric., 14, 673.
Matthew, A.G. and Parpia, H.A.B. (1971) Food browning as a polyphenol
reaction. Adv. in Enzymol., 19, 75.
562 Tuber quality
Mazza, G., Hung, J. and Dench, M.J. (1983) Processing/nutritional quality
changes in potato tubers during growth and long term storage. Can, lnst. Food
Sci. Technol. J., 16, 39--44.
McKee, R.K. (1959) Factors affecting the toxicity of solanine and related com-
pounds to Fusarium caeruleum. J. Gen. Microbial., 20, 686.
McRae, D.C. (1986) Mechanical damage to potatoes, causes, measurement and
remedies. Aspects of Applied Biology, 13, 383-92.
McRae, D.C., Carruthers, J. and Porteous, R.L. (1976) The effect of drop height
on damage sustained by some main-crop varieties. Scottish lnst. of Agric.
Engng, Dept. Note SlN1202.
McRae, D.C., Hutchison, P.S. and Fleming, J. (1982) The development of power
driven disc shares for two row high output potato harvesters. Am. Potato J., 59,
479.
McRae, D.C., Fleming, J. and Fisher, A. (1984) The development of a rigid carton
for retailing potatoes. The Agricultural Engineer, 39, 74-9.
McRae, D.C., Hutchison, P.S. and Carruthers, J. (1986) Sieving control and
horizontal agitation of potato harvester chains. Trans. Am. Soc. Agric. Engrs,
29,366-9.
McWeeny, D.J. and Burton, H.S. (1962) Sulphites and carbonyl-amino
chromophore development. Nature, London, 196,41-2.
Meijers, C.P. (1983) In search of the ideal chip variety. Rapport 497 B, IBVL,
Wageningen.
Mogen, K.L. and Nelson, D.C. (1986) Some anatomical physiological potato tuber
characteristics and their relationship to hollow heart. Am. Potato J., 63, 609.
Mohr, W.P., Spurr, A.R., Fenn, P. and Timm, H. (1984) X-ray microanalysis of
hollow heart potatoes. Food Microstructure, 3, 41.
Mondy, N.I., Koch, R.L. and Chandra, S. (1979) Influence of nitrogen fertilization
on potato discolouration in relation to chemical composition. 2. Phenols and
ascorbic acid. J. Agric. Fd Chem., 27,418-20.
Monk, L.S. and Davies, H.V. (1989) Antioxidant status of the potato tuber and
Ca2»fH deficiency as a physiological stress. Physiol. Plant., 75, 411.
Morrell, S. and ap Rees, T. (1986) Control of the hexose content of potato tubers.
Phytochem., 25, 1073.
Mulder, E.G. (1949) Mineral nutrition in relation to the biochemistry and
physiology of potatoes. Pl. Soil, 2, 59-121.
Mulder, E.G. and Bakema, K. (1956) Effect of the nitrogen, phosphorus,
potassium and magnesium nutrition of potato plants on the content of free
amino acids and on the amino acid composition of the protein of tubers. PI.
Soil, 7, 135-66.
Miiller, K. (1975) Veranderungen wertgebender Inhaltsstoffe in der Kartoffelpflanze
und-kolle im Verlauf von Vegetation und Lagerung, ihre Bedcutung fUr die
Qualitat der Knolle. Hefte filr den Kartoffelbau, 17 Schriftenreihe der
Forderungsgemeinschaft der kartoffelwirtschaft, Hamburg.
Muneta, P. (1981) Comparison of inhibitors of tyrosine oxidation in the enzymatic
blackening of potatoes. Am. Potato 1., 58, 85.
Muneta, P. and Kaisaki, F. (1985) Ascorbic acid-ferrous iron (Fe»fH»fH) complexes
and after cooking darkening of potatoes. Am. Potato J., 62, 531-6..
Munshi, c.B. and Mondy, N.I. (1989) Ascorbic acid and protein content of
potatoes in relation to tuber anatomy. J. Food Sci., 54, 220-1.
References 563
National Institute of Agricultural Botany (1990) Potato Variety Handbook 1991,
NIAB, Cambridge, 44 pp.
Nelson, D.C. and Thoreson, M.C. (1986) Relationships between tuber size and
time of harvest to hollow heart initiation in dryland Norgold Russet potatoes.
Am. Potato J., 63, 155.
Nelson, D.G., Jenkins, P.D. and Gillison, T.C. (1988) Processing potential of
potato cultivars at early harvest. Potato Res., 31, 633-42.
Neuberger, A. and Sanger, F. (1942) The nitrogen of the potato. Biochem. J., 36,
662-71.
Nilsson, S.B., Hertz, C.B. and Falk, S. (1958) Physiologica Pt., 11,818--37.
Noble, R. (1985) The relationship between impact bruising in potato tubers. J.
Agric. Engng. Res., 32, 111-2l.
Nonaka, M. (1980) The textural quality of cooked potatoes. I. The relationship of
cooking time to the separation and rupture of cells. Am. Potato J., 57, 141-9.
Noonan, J.C., Webb, R.E., Attaya, R.B. and Miller, J.C. (1951) Influence of
location on the ascorbic acid content of the Irish potato. Am. Potato 1., 28,
521--4.
O'Brien, M.J. and Rich, A.E. (1976) Potato diseases. US Dep. Agric., Agric.
Handbook, 474.
Ojima, T., Hori, S. and Torantini, H. (1970) Effects of ionizing radiation upon the
sprout inhibition, loss of weight and respiration in potato tubers. Ann. Rep.
Radiat. Center Osaka Prefect., 11, 143.
Olsson, K. and Fridell, J.E. (1975) Differences in blackspot susceptibility of two sib
varieties of potato in relation to cell volume and some chemical factors. Potato
Res., 18, 338.
Ophuis, B.G., Hesen, J.C. and Kroesbergen, E. (1958) The influence of
temperature during handling on the occurrence of blue discolourations inside
potato tubers. Eur. Potato J., 1(3),48-65.
Pareles, S.R. and Chang, S.S. (1974) Identification of compounds responsible for
baked potato flavour. J. Agric. Fd Chem., 22, 339--40.
Park, W.D. (1983) Tuber proteins of potato - a new and surprising molecular
system. Plant Msl. Bioi. Rep., 1, 61--6.
Parke, D. (1963) The resistance of potatoes to mechanical damage caused by
impact loading. J. Agric. Engng. Res., 8, 173-7.
Patil, B.C., Singh, B. and Salunkhe, D.K. (1975) Formation of chlorophyll and
solanine in Irish potato (Solanum tuberosum L.) tubers and their control by
gamma irradiation and CO 2 enriched packaging. Lebensm. Wiss. TechuoL, 4,
123.
Paul, A.A. and Southgate, D.A.T. (1978) The Composition of Foods, 4th edn,
HMSO, London, 418 pp.
Pendharkar, M.B. and Nair, P.M. (1987) Alteration in phenylpropanoid
metabolism in gamma irradiated potatoes. Potato Res., 30, 589.
Pierpoint, W.S. (1970) Formation and behaviour of o-quinones in some processes
of agricultural importance. Ann. Rep. of Rothamsted Expt. Stn, pp.
199-218.
Plaisted, P.H. (1957) Growth of the potato tuber. PI. Physiol., 32, 445-53.
Pol, G. and Labib, A.I. (1963) De voedingswaarde van aardappelen van
verschillende rassen en de invloed daarop van bemesting en bewaring. Voeding,
83,479-99.
564 Tuber quality
Pollock, C.l. and ap Rees, T. (1975) Activities of enzymes of sugar metabolism in
cold-stored tubers of Solanum tuberosum. Phytochem., 14, 613.
Ponnampalam, R. and Mondy, N.1. (1983) Effect of cooking on the total
glycoalkaloid content of potatoes. J. Agric. Fd Chem., 31, 493-5.
Porter, W.L., Fitzpatrick, T.l. and Talley, E.A. (1964) Studies on the relationship
of specific gravity and total solids of potatoes. Am. Potato J., 41, 329-36.
Potato Marketing Board (1983) Report on a Survey of Consumer Attitudes and
Behaviour 1981182, Potato Marketing Board, Oxford, 72 pp.
Potato Marketing Board (1988) Report on a Usage and Attitude Survey into the
British Potato Market, Potato Marketing Board, Oxford, 80 pp.
Potato Marketing Board (1989) Potato Processing in Great Britain 1988, Potato
Marketing Board, Oxford, 10 pp.
Pressey, R. (1966) Separation and properties of potato invertase and invertase
inhibitor. Arch. Biochem. Biophys., 113, 667.
Pressey, R. (1967) Invertase inhibitor from potatoes: purification characterization,
and reactivity with plant invertases. Plant Physiol., 42, 1780.
Pressey, R. (1969) Role of invertase in the accumulation of sugars in cold-stored
potatoes. Am. Potato 1., 46, 291.
Pressey, R. (1970) Changes in sucrose synthetase and sucrose phosphate synthetase
activities during storage of potatoes. Am. Potato J., 47, 245.
Pressey, R. and Shaw, R. (1966) Effect of temperature on invertase, invertase
inhibitor, and sugars in potato tubers. Plant Physiol., 41,1657.
Racusen, D. and Foote, M. (1980) A major soluble glycoprotein of potato tubers.
J. Food Biochem, 4, 43-52.
Reeve, R.M. (1967) Suggested improvements for microscopic measurement of cells
and starch granules in fresh potatoes. Am. Potato J., 44, 41-50.
Reeve, R.M., Timm, H. and Weaver, M.L. (1973) Parenchyma cell growth in
potato tubers. I. Different tuber regions. Am. Potato J., 50, 49-57.
Reust, W. and Munster, 1. (1978) Facteurs responsables de la variation de la forme
des tubercules de pommes de terre. Revue Swisse Agric., 10, 13-17.
Rex, B.L. and Mazza, G. (1989) Cause, control and detection of hollow heart in
potatoes: A review. Am. Potato J., 66, 165.
Rhodes, 1.M.C. and Wooltorton, L.S.C. (1978) The biosynthesis of phenolic com-
pounds in wounded plant storage tissues, in Biochemistry of Wounded Plant
Tissues. (ed. G. Kahl), Walter de Gruyer & Co., Berlin and New York. pp. 243-86.
Richardson, D.L., Davies, H.V., Ross, H.A. and Mackay, G.R. (1990) Invertase
activity and its relation to hexose accumulation in potato tubers. J. Exp. Bot.,
41,95-9.
Ring, S.G. and Selvendran, R.R. (1981) An arabinogalactoxyloglucan from the
cell wall of Solanum tuberosum. Phytochem., 20, 2511-19.
Roberts, E.A. and Procter, B.E. (1955) The comparative effect of ionising
radiations and heat upon the starch containing cells of potato tuber. Food Res.,
20,254-63.
Robins, 1.S. and Domingo, C.E. (1956) Potato yield and tuber shape as affected by
severe soil-moisture deficits and plant spacing. Agron. J., 48, 488.
Rogozinska, I., Hippe, 1. and Miiller, K. (1986) Einfluss der Langzeitlagerung
und einer kontrollierten Stossbeschadigung mit anschliessender Kurzzeitla-
gerung auf den Gehult an phenolischen sauren in Knollen verschiedener
Kartoffelsorten. Potato Res., 29, 239-43.
References 565
Rolf, L.A. (1940) The effect of cooking and storage on the ascorbic acid content of
potatoes, J. Agric. Res., 61, 381-95.
Rosenstock, V.G. and Zimmerman, H.J. (1976) Vergleichende Studien uber den
Protein-und Nucleinsaurestoffwechel beim Speicherparenchyn von Solanum
tuberosum L. mit primarer und sekundarer mitotischer Aktivitut. Beitr. BioI.
Pjlanz., 52, 413-26.
Ross, H., Pasemann, P. and Nitzsche, W. (1978) Der Glykoalkaloidgehalt von
Kartoffelsorten in seiner Abhangigkeit zum Geschmack. Z. Pjlanzenzucht., 80,
64-79.
Rubin, B.A. and Metlisky, L.V. (1958) A study of the action of ionizing radiation
on the metabolism of potato tuber in relation to the problem of their all year
round storage, in Proc. 2nd Int. Conf. Peaceful Uses of Atomic Energy, 27,437.
Rumpf, G. (1972) Gaschromatographische Bestimmung loslicher Inhaltsstoffe in
bestrahlten und mit chemischen Keimhemmungsmitteln behantdlten
Kartoffeln. Potato Res., 15, 236-45.
Salaman, R.N. (1926) Potato Varieties, Cambridge University Press, Cambridge,
378 pp.
Salkova, E.G. (1957) The influence of irradiation with Cobalt 60 on vitamin C
content in potatoes. Dokl. Akad. Nauk SSSR, 114, 757.
Salunkhe, D.K., Wu, M.T. and Jadhev, S.J. (1972) Effects of light and
temperature on the formation of solanine in potato slices. J. Food Sci., 37, 969.
Sasaki, T., Tadokoro, K. and Suzuki, S. (1971) Multiple forms of invertase in
potato tubers stored at low temperature. Phytochem., 10,2047.
Satyanarayan, V. and Nair, P.M. (1986) Enhanced operation of 4-aminobutyrate
shunt in gamma-irradiated potato tubers. Phytochem., 25, 1801.
Sawyer, R.L. and Collin, G.H. (1960) Blackspot of potatoes. Am. Potato J., 37,
115-26.
Schallenberger, R.S., Smith, o. and Treadaway, R.H. (1959) Role of sugars in the
browning reaction in potato chips. J. Agric. Fd Chem., 7, 274.
Schippers, P.A. (1968) The influence of rates of nitrogen and potassium application
on the yield and specific gravity of four potato varieties. Potato Res., 11, 23-33.
Schippers, P.A. (1971) Measurement of blackspot susceptibility of potatoes. Am.
Potato J., 48, 71-81.
Schippers, P.A. (1976) The relationship between specific gravity and percentage
dry matters in potato tubers. Am. Potato J., 53, 111-22.
Schoor!, D. and Holt, J.E. (1983) Cracking in potatoes. J. Texture Studies, 14,61.
Schwimmer, S. and Weston, W.J. (1958) Chlorophyll formation in potato tubers as
influenced by gamma irradiation and by chemicals. Am. Potato J., 35, 534.
Schwimmer, S., Benvenue, A., Weston, W.J. and Potter, A.L. (1954) Survey of
the major and minor sugar components of the white potato. J. Agric. Fd Chem. ,
2, 1284.
Scott, R.K. and Wilcock son , S.J. (1978) Application of physiological and
agronomic principles to the development of the potato industry, in The Potato
Crop (ed. P.M. Harris), Chapman & Hall, London, pp. 678-704.
Self, R., Rolley, H.L.J. and Joyce, A.E. (1963) Some volatile compounds from
cooked potatoes. J. Sci. Fd Agric., 14, 8-14.
Selvendran, R.R., Stevens, B.J.H. and DuPont, M.S. (1987) Dietary fibre:
chemistry, analysis and properties. Adv. Food Res., 31,117-209.
Sharma, M.K., Isleib, D.R. and Dexter, S.T. (1959) The influence of specific
566 Tuber quality
gravity and chemical composItIOn on the hardness of potato tubers after
cooking. Am. Potato J., 36, 105~12.
Shekhar, V.C. and Iritani, W.M. (1979) Changes in malic and citric acid contents
during growth and storage of Solanum tuberosum L. Am. Potato J. , 56, 87~94.
Shekhar, V.e., Iritani, W.M. and Arteca, R. (1978) Changes in ascorbic acid
content during growth and short-term storage of potato tubers (Solanum
tuberosum L.). Am. Potato 1., 55, 663·70.
Sherman, M. and Ewing, E.E. (1982) Temperature, cyanide and oxygen effects on
the respiration, chip color, sugars and organic acids of stored tubers. Am.
Potato J., 59,165.
Shimshi, D. and Susnoschi, M. (1985) Growth and yield studies of potato
development in a semi-arid region 3. Effect of water stress and amounts of
nitrogen top dressing on physiological indices and tuber yield and quality of
several cultivars. Potato Res., 28, 177~91.
Silva, G.H., Chase, R.W. and Kitchen, R.B. (1988) Influence of irrigation,
nitrogen and calcium levels on specific gravity and internal defects of potatoes.
Am. Potato J., 65, 498.
Sinden, S.L. and Webb, R.E. (1972) Effect of variety and location on the
glycoalkaloid content of potatoes. Am. Potato J., 49, 334-8.
Sinden, S.L., Deahl, K.L. and Aulenbach, B.B. (1976) Effects of glycoalkaloids
and phenolics on potato flavour. J. Food Sci., 41, 520.
Sizer, C.E., Maga, 1.A. and Craven, e.J. (1980) Total glycoalkaloids in potatoes
and potato chips. J. Agric. Fd Chem., 28, 578~9.
Skrobacki, A., Halderson, 1.L., Pavek, 1.1. and Corsini, D.L. (1989) Determining
potato tuber resistance to impact damage. Am. Potato 1., 61, 401~13.
Smith, A.M. and Gillies, 1. (1940) The distribution and concentration of absorbic
acid in the potato (Solanum tuberosum). Biochem. J., 34, 1312··20.
Smith, O. and Davis, e.o. (1968) Potato processing, in Potatoes: Production,
storing, processing (ed. O. Smith), A VI, Westport, pp. 558-602.
Smittle, D.A., Thornton, R.E., Peterson, e.L. and Dean, B.B. (1974) Harvesting
potato with minimum damage. Am. Potato J., 51, 152-64.
Solomos, T. and Laties, G.G. (1975) The mechanism of ethylene and cyanide
action in triggering the rise in respiration in potato tubers. Plant Physiol., 55, 73.
Sowokinos, J. R. (1978) Relationship of harvest sucrose content to processing
maturity and storage life of potatoes. Am. Potato J., 55, 333.
Stegmann, H. and Schnick, D. (1982) Index of European Potato Varieties,
Mitteilangen der Biologischen Bundesanstalt Heft 211, Berlin, pp. 1~127.
Storey, R.M.l. and Shackley, D.l. (1987a) The effect of reconditioning on the
processing quality of Pentland Dell, Maris Piper and Wilja. Abst. 10th Triennial
EAPR Conference, 1987, Aalborg pp. 61~2.
Storey, R.M.J. and Shackley, D.l. (1987b) The potential of Cohalt-60 gamma
irradiation for long-term storage and its effect on processing quality. Potato
Res., 30,137.
Swain, T. and Self, R. (1964) Studies on potato flavour. Eur. Potato J., 7,
228~32.
Talburt, W.F. (1987) Canned white potatoes, in Potato Processing, 4th edn (eds
W.F. Talburt and O. Smith), AVI, New York, pp. 683~95.
Talburt, W.F. and Smith, O. (1975) Potato Processing, 3rd edn, AVI Publishing
Co., Westport, 705pp.
References 567
Talburt, W.F., Boyle, F.P. and Hendel, C.E. (1987) Dehydrated mashed potatoes
- potato granules, in Potato Processing, 4th edn (eds W.F. Talburt and O.
Smith), AVI, New York, pp. 535-55.
Talley, E.A. (1983) Protein nutritive value of potatoes is improved by fertilization
with nitrogen. Am. Potato J., 60, 35-9.
Talley, E.A., Fitzpatrick, T.H., Porter, W.L. and Murphy, H.l. (1961) Chemical
composition of potatoes. I. Preliminary studies on the relationship between
specific gravity and nitrogenous constituents. J. Food Sci., 26, 351-5.
Talley, E.A., Fitzpatrick, T.l. and Porter, W.L. (1970) Chemical composition of
potatoes. VIII. Effect of variety, location and year of growth on the content of
nitrogen compounds. Am. Potato J., 47, 231-44.
Talley, E.A, Toma, R.B. and Orr, P.H. (1984) Amino acid composition offreshly
harvested and stored potatoes. Am. Potato J., 61, 267-79.
Thomas, P. (1984) Radiation preservation of foods of plant origin. Part 1. Potatoes
and other tuber crops. CRC Crit. Rev. in Food Sci. and Nutr., 19, 327.
Thomas, P., Srirangarajan, AN., Hoshi, A.N. and lanave, M.T. (1978) Storage
deterioration in gamma irradiated and unirradiated Indian potato cultivars
under refrigeration and tropical temperatures. Potato Res., 22, 261.
Thomson, A.l., Hughes, l.e. and Starr, C. (1987) Breeding for long dormancy and
low reducing sugar content following extended storage at low temperature, in
Abstracts of Conf. Papers and Posters, 10th Trienn. Conf. of EAPR, Aalborg,
pp.22-3.
Trevanion, S.l. and Kruger, N.l. (1989) Influence of temperature on the kinetic
properties of pyrophosphate: fructose-6-phosphate phosphotransferase from
potato tubers. Plant Physiol. (suppl.), 4, 56.
Turczyn, M.T., Grant, S.W., Ashby, B.H. and Wheaton, F.W. (1986) Potato
shatter bruising during laboratory handling and transport simulation. Trans.
Am. Soc. Agric. Engrs, 29, 1171-5.
Tzeng, K.e., Kelman, A., Simmons, K.E. and Kelling, K.A. (1986) Relationship
of calcium nutrition to internal brown spot of potato tubers and sub-apical tip
necrosis of sprouts. Am. Potato J., 63, 87.
Umaerus, M. and Olsson, K. (1974) Varietal differences in tyrosine and chlorogenic
acid in relation to enzymic discoloration of potato tubers. Potato Res., 17, 157.
Ussuf, K.K. and Nair, P.M. (1972) Metabolic changes induced by sprout inhibiting
dose of gamma irradiation in potatoes. J. Agric. Fd Chem., 20, 282.
van Denburgh, R.W., Hiller, L.K. and Koller, D.C. (1980) The effect of soil
temperatures on brown center development in potatoes. Am. Potato J., 57, 371.
van Es, A. and Hartmans, K.l. (1975) Some differences in susceptibility to internal
blackspot. Pot. Res., 18,338.
van Es, A. and Hartmans, K.l. (1987a) Structure and chemical composition of the
potato, in Storage of Potatoes (eds. A Rastovski, A. van Es et al.), Pudoc,
Wagenigen, pp. 15-78.
van Es, A. and Hartmans, K.l. (1987b) Sugars and starch during tuberisation,
storage and sprouting, in Storage of Potatoes (eds A. Rastovski, A. van Es et
al. ), Pudoc Centre for Agricultural Publishing and Documentation,
van Es, A and Hartmans, K.l. (1987c) Respiration, in Storage of Potatoes (eds A.
Rastovski, A. van Es et at.), Pudoc, Centre for Agricultural Publishing and
Documentation, Wageningen, pp. 133-9.
568 Tuber quality
van Es, A. and Hartmans, K.J. (1987d) Carbohydrate metabolism during
storage
of potatoes. Abstr. Conf. Papers and Posters of the 10th Trienn. Conf. of EAPR,
Aalborg, p. 63.
van Loon, C.D. (1981) Effect of water stress on potato growth, development and
yield. Am. Potato J., 58, 51-69.
van Schaftingen, E., Lederer, B., Bastrons, R. and Hers, H.G. (1982) A kinetic
study of pyrophosphate: fructose-6-phosphate phosphotransferase from potato
tubers. Eur. J. Biochem., 129, 191.
Vander Zaag, K. and Ffrench, S. (1987) Preliminary evaluations of foliar calcium
applications with respect to yields and processing quality of the potato cultivars
Atlantic and Norchip. Onto Potato Cultivar Eval. Assoc. Bull. No.2. pp. 3-5.
Vander Zaag, D.E. (1970) Groei, morfologie en anatomie van de aardappelknol
(vervolg). De Pootaardappelwereld, 1-5.
Vander Zaag, D.E. and Meijers, C.P. (1970) Blackspot - practical aspects. Proc.
Trienn Conf. Eur. Assoc. Potato Res., 1969,4, 93-103.
Varela, G. and Gurbano, G. (1971) Effect of irradiation on the Imtritional quality
of protein in potatoes. Agrochimica, 16, 171.
Verbist, J.F. and Monnet, R. (1979) A propos de la teneur en solanine des petite
tubercules nouveaux de pomme de terre (Solanum tuberosum L.). Potato Res.,
22,239-44.
Verma, S.C., Malhotra, V.P., Joshi, K.C. and Sharma, T.R. (1971) Specific
gravity and dry matter content of potato. Potato Res., 14, 94-5.
von Scheele, C., Svensson, G. and Rasmusson, J. (1937) Die Bestimmung des
Starkegehalts und der Trockensubstanz der Kurtoffel mit Hilfe des spezifischen
Gewichts. Landw. Vers Sta., 127, 67-96.
Wager, H.G. (1963) The role ofphytin in the texture of cooked potatoes. J. Sci. Fd
Agric., 14, 583-6.
Walker, J.R.L. (1977) Enzymic browning in foods: Its chemistry and control. Food
Tech. in New Zealand, 12, 19.
Warren, D.S. and Woodman, J.S. (1974) The texture of cooked potatoes: a review.
J. Sci. Fd Agric., 25, 129-38.
Watts, K.c. and Russell, L.T. (1985) A review of techniques for detecting hollow
heart in potatoes. Can. Agric. Engin., 27, 85.
Weaver, M.L., Ng, K.C. and Huxsoll, C.C. (1979) Sampling potato tubers to
determine peel loss. Am. Potato J., 56, 217-24.
Wedzicha, B.L. (1984) Chemistry of Sulphur Dioxide in Foods, Elsevier Applied
Science Publishers Ltd, London, pp. 183-207.
Welte, E. and Muller, K. (1966) Uber den einflus der Kalidungung auf die
dunkelung von rohem kartoffelbrei. Eur. Potato J., 9, 36.
Willard, M.J., Hix, V.M. and Kluge, G. (1987) Dehydrated mashed potatoes -
potato flakes, in Potato Processing, 4th edn (eds W.F. Talburt and O. Smith),
AVI, New York, pp. 557-612.
Witney, B.D. (1984) The investigation and promotion of stonc/dod windrowing for
potato production systems. Res. Dev. Agric., 1, 1-20.
Wolf, M.J. and Duggar, B.N. (1946) Estimation and physiological role of solanine
in the potato. J. Agric. Res., 73, 1-32.
Wood, F.A. and Young, D.A. (1974) TGA in Potatoes. Canada Dept. Agric.,
Publication no. 1533.
References 569
Woodwards, L. and Jackson, M.T. (1985) The lack of enzymic browning in wild
potato species, Series Longipedicellata, and their crossability with Solanum
tuberosum. Zeitsch. fur Pjlanzenzuchtung, 94, 278.
Woolfe, J.A. (1986a) The Potato in the Human Diet, Cambridge University Press,
Cambridge.
Woolfe, J.A. (1986b) Potato - a gift from the Andes. Brit. Nut. Fdn. Bull. 46, 11,
29-45.
Wu, M.T. and Salunkhe, D.K. (1971) Effect of gamma irradiation on wound
induced glycoalkaloid formation in potato tubers. Lebensm. Wiss. Technol., 4,
123.
Wu, M.T. and Salunkhe, D.K. (1977) Use of spray lecithin for control of greening
and glycoalkaloid formation in potato tubers. J. Food Sci., 42, 1413.
Wunsch, A. and Schaller, K. (1972) Uber wechselwirkungen zwischen zuckern und
aminosauren bei de ausbildung der chipsfarbe. Potato Res., 15, 12.
Wurr, D.C.E. and Allen, E.J. (1974) Some effect of planting density on the
relationship between tuber size and dry-matter percentage in potatoes. J. Agric.
Sci., Camb., 82, 277-82.
Wurr, D.C.E., Bean, J.N. and Allen, E.J. (1978) Effect of variety and date of
harvest on the tuber dry-matter percentage of potatoes. J. Agric. Sci., Camb.,
90, 597--604.
Wurster, R.T. and Smith, o. (1963) Potato quality XVIII. The distribution of
radioiron in the potato tuber and its significance in after cooking darkening.
Am. Potato J., 40, 415-20.
Yamaguchi, M., Timm, H. and Spurr, A.R. (1964) Effects of soil temperature on
growth and nutrition of potato plants aM tuberization, composition, and
periderm structure of potatoes. Proc. Am. Soc. Hort. Sci., 84. 412-23.
Zitnak, A. (1953) The influence of certain treatments upon solanine synthesis in
potatoes, M.S. Thesis, Univ. of Alberta, Edmonton.
CHAPTER 13
Mechanization of crop production

and handling operations
B.D. Witney and D.C. McRae
13.1 INTRODUCTION
Major improvements have taken place in the design of potato crop

establishment, harvesting and grading machinery over the past decade.
These changes have been consumer-driven with premium payments for
quality produce, particularly for pre-packs of uniformly sized tubers. The
psychological impact of freshness is fully recognized and large retail outlets
often display such produce to maximum advantage at the entrance to the
store. There is no place in this quality image for the bag trade - an
unknown quantity of dirty, damaged and even diseased tubers.
Attaining a high marketable yield requires attention to detail at all stages
of the crop production process in order to minimize mechanical damage
and bruising of the tuber. There is a wealth of evidence to show that clods
present in the seedbed will remain throughout the growing season and will
exacerbate harvesting problems. Consequently, the emphasis on providing
complex trash separation systems for harvesters in the 1960s has given way
to the more modern concept of simple, low damage, harvesters preceded
by stone and clod separation in the spring to allow the potato setts to
multiply in a fine tilth (Fig. 13.1)
13.2 SEEDBED PREPARATION
13.2.1 Tilth requirement

The majority of arable crops can be grown in soils with a moderate stone
content below 100 mm in nominal diameter. Indeed, a proportion of the
plough layer in the gravel fraction is considered to assist soil drainage by
keeping the structure more open, and to enhance soil warming because the
Published in 1992 by Chapman & Hall, London. ISBN 0 412296403
Ch. 13 © 1992 B.D. Witney and D.C. McRae.
Seedbed preparation 571
Figure 13.1 Windrows of stones and clods in the valleys between the potato
ridges.
thermal conductivity of stone is higher than well-structured, moist soil. For

the potato crop, however, any stones and clods which cannot pass through
the minimum web bar spacing on the harvester of between 25 and 31 mm
will decrease the commodity throughput capacity of the machine and will
increase crop damage (Witney, 1988). As potatoes are seldom grown on
the same land every year, this specific crop requirement for a stone and
clod free tilth may be accommodated by a springtime operation of stone
and clod windrowing which partially replaces alternative cultivation
operations to provide the necessary seedbed conditions.
13.2.2 Potato land preparation

It is recommended that deep ploughing, to 30 cm if possible, is completed
by Christmas on heavier soils to avoid smearing the furrow bottom and the
creation of clods. Tractor overloads which cause high wheelslip should be
avoided and grass swards should be rotary cultivated in the autumn to
destroy the turf.
The land should be cultivated to a depth of 24 cm. The degree of soil
comminution depends on the soil conditions and the energy applied (Fig.
13.2). Of the power-driven machines, the vertical axis rotary cultivator
(e.g. Lely Roterra), the reciprocating harrow (e.g. Vicon), and the
horizontal axis spiked rotary cultivator (e.g. Howard Rotavator) are all
effective in producing a good tilth in a single pass (Pascal et at., 1983). In
stony conditions, the horizontal axis rotary cultivator tends to bury stones
572 Mechanization of crop production
and unbroken clod; the other machines have the reverse effect, but are
more prone to blockages when large stones are present. Mounted spring
tine harrows have a high work rate but usually require two passes to
achieve a similar tilth to that produced by the power-driven machines.
Draught implements impose a higher traction requirement on the tractor
and therefore have the disadvantage of introducing more soil damage;
whereas power take-off (pto) driven machines such as the horizontal axis
spiked rotary cultivator tend to push the tractor forward so that the power
used for traction is a small proportion of the total.
Figure 13.2 Lely Roterra vertical axis rotary harrow preparing a seedbed.
13.2.3 Stone and clod windrowing

Stone and clod windrowers separate out the stones and clods from the top-
soil and concentrate them into windrows away from the vicinity of the crop
for the period from planting to harvesting (Witney, 1984). Even in some
stone-free soils, the separation and breakdown of clods may be beneficial.
The major advantages of the technique are:
l. reduction of 30-50% in severe damage to the tubers during harvesting;
2. improved harvesting spot rate of work of up to 40%;
3. unaffected yields of subsequent crops provided that stone windrows are
adequately redistributed by cross cultivation.
Extra management is required because planting may be delayed both
through waiting for suitable drier soil conditions and through planter work
rates being restricted by the slower windrowing operation. Soil conditions
Fertilizer application 573
are critical as too many clods leave insufficient tilth for crop production and
incur a greater risk of exposed tubers.
After cultivation, the land is set up into two-row beds with a bed-former
equipped with heavy duty furrow openers which loosen the valley bottoms
to the depth of the plough layer in preparation for burying the stones and
clod. The stone and clod windrower lifts the bed of soil, sifts out the fine
material and transfers the hard clod and stones into the valley between the
beds of untreated soil where it is pressed into the loose furrow by the
tractor wheels during the next pass of the machine.
Potato planting may follow as a separate operation or may be combined
with the stone and clod windrowing in a single machine (Fig. 13.3). The
combined operation has a lower rate of work than separate activities
because the slow progress of the windrower in the soil is combined with the
stoppages for filling the planter on the headland. A simple machine,
however, does reduce labour requirements and avoids the potential
weather risk of leaving cultivated land unplanted.
Figure 13.3 Reekie combined stone and clod windrower/potato planter transfer-
ring the separated material into the adjacent valley.
13.3 FERTILIZER APPLICATION
Fertilizer is used more efficiently, if placed rather than broadcast and,

therefore, less should be used if placement is adopted. Fertilizer close to
the potato setts, however, can damage young shoots. It should be placed
well to the side of, and slightly below, the setts. The risk of damage is
increased in light, dry soils and can be minimized by broadcasting half of
the fertilizer and placing half.
The number of machinery operations can be reduced by combining the
fertilizer placement and potato planting operations. A front mounted,
liquid fertilizer tank acts as a counter balance to the rear mounted potato
planter and the filling operations on the headland can be carried out
simultaneously with replenishing the planter. The complexities start to
accrue when stone windrowing and potato planting are undertaken as a
joint operation; in this event, fertilizer placement is transferred to the bed-
forming operation for which timeliness is less critical (Fig. 13.4). The bands
of placed fertilizer which are processed by the stone and clod windrowerl
potato planter remain in the same position relative to the setts but the
granules are more dispersed in the soil.
Figure 13.4 BW furrow opener equipped with a fertilizer placement unit

preparing beds for stone and clod windrowing.
13.4 POTATO PLANTING
13.4.1 Accuracy of planting

The market requirement for specific tuber size ranges - whether for a seed
crop, normal ware or baker size - means that the farmer has to attempt to
control the size spectrum of the crop by adjusting his seed size and spacing.
A number of complex interactions occur when the spatial arrangement of
seed tubers is altered.
Potato planting 575
There is ample evidence (Pascal and Robertson, 1977) that there is little
overall effect on yield from irregularly spacing seed tubers though uneven
spacing does influence the number and size of tubers. Palmer (1976)
showed that, although close grading of seed had little effect on total yield,
a large tuber planted too closely to another large tuber tends to encourage
the production of a relatively greater number of small tubers. Small tubers
planted too far apart will tend to produce small numbers of large tubers.
Depth of planting has an effect on yield. Lang et al. (1986) found that
deep planting can depress yield especially with smaller seed tubers. If
planted too shallow, large seed tubers may produce a higher than average
number of small tubers near the surface which may suffer from greening.
Considering the planting patterns for tubers, Travis (1987) suggests that
crop tuber size is influenced mainly by the number and weight of tubers
planted/unit area.
13.4.2 Planter design

The key elements in planter design are:
1. the hopper and hopper emptying system;
2. the feed arrangement;
3. the planting belt;
4. the opening and closing bodies.
The hopper of a two-row planter usually has a capacity of up to one
tonne of seed. Where chitted seed is handled in trays some planters may
have a tray platform which allows the seed to be emptied into the feed
system by hand.
Some larger planters (e.g. four row) have hoppers which can be tipped
to keep a steady flow to the feed system. Bridging may occur with the
longer varieties and with chitted seed it is likely to be a serious problem if
sprouts exceed about 30 mm length. Vibrating plates, tilt trays, or small
conveyor belts controlled by a trip switch may be used to improve
movement of seed to the feed system. It is common to have provision to
alter the degree of agitation.
From the hopper, potatoes move to the singulation sytem. In the most
common type of planter, this comprises a double run of staggered cups
(Fig. 13.5) mounted on a flat conveyor belt running vertically from the
hopper base to a delivery tube leading to the planter opening share. The
purpose of the double run of cups is to reduce the tuber pick-up speed as
the cups plough through a small reservoir of tubers usually situated outside
the main supply in the hopper. The double run cup arrangement can cope
with evenly sized seed, or split graded seed with the use of small cup inserts
to enable smaller seed to be used.
The main advantage of the cup planter is accuracy of tuber spacing,
especially at low speeds. Drawbacks are damage to chits, an inability to
Figure 13.5 Cup planter. (By courtesy of Reekie.)
perform so well with long cultivars such as Pentland Dell, or Shepody

which are favoured by chip producers and some gapping on side slopes
when tubers may fall out of cups.
Belt planters singulate tubers into a stream with consecutive tubers
touching. At this stage, there is usually a transfer system to a planting belt
leading to the ridge opener. Belt planters generally operate faster than cup
planters and for some markets the spacing accuracy is adequate. Chitted
seed suffers less damage. According to Meijer and Frederiks (1975)
describing tests on a belt type machine, broken sprouts and bruised
sprouts amounted to 10% and 12% respectively at forward speeds of
2.6-6.0 km h- 1 .
Apart from handling chitted seed well, some farmers favour belt fed
planters because the spacing of the tubers in the ground is related to the
contiguous length of tubers on the planter belt. Thus, a small tuber
followed by a large tuber will compensate in the crop to give a more
consistent yield. Against this, ground spacing variations can be consider-
able and there is no mechanism to compensate for gaps.
At the higher forward speeds there can be some rolling as tubers are
released by cups or belts with a forward trajectory relative to the ground.
Covering is generally carried out by discs and these tend to leave a peaked
and fragile ridge crest. On light soils which do not readily smear, a profile
Potato planting 577
former comprising a flat plate with notches for each drill can be mounted at
the rear of the machine. The profile plate can produce a ridge configuration
which may be less prone to slumping in heavy rain.
The depth of planting is an important consideration and some farmers
find that as the planter hopper empties, the opening shares may rise by
50-75 mm above the initial operating depth. Planters with floating planting
shares may partly overcome this drawback.
New planter designs are being considered. There is a need for planters
capable of handling well chitted seed, and handling a wider seed size range
with less need for split grading or multiple grading. The design of feed
systems which impart low or even zero velocity to seed relative to the
ground would also be advantageous.
From time to time, interest is expressed in using cut seed in the UK. The
pick machines favoured in USA and Canada do not necessarily achieve the
planting accuracy commonly expected mainly due to doubles. Entz and La
Croix (1983) cite the performance of one machine which gave a coefficient
of variation in spacing of 76.5% at 39.2 cm spacing, which fell to 50% at
47.5 cm spacing. This is similar to the performance of some cup planters
cited in a series of tests of planters carried out in Holland (IMAG, 1978).
During the same tests 4.8-9% wheelslip was noted.
13.4.3 Spacing control

Wheelslip in ground drive planters can vary due to changes in the drive
power requirement and ground conditions. Recently at the Scottish Centre
of Agricultural Engineering (SCAE), a planter controller has been
developed which avoids the wheelslip problem. This unit uses the land
wheels only, to drive a wheel speed sensor and drives the planting
mechanism by means of a hydraulic motor through a hydraulic proportion-
ing valve. A great advantage of this arrangement is that the tractor driver is
furnished with a digital display and spacing control in his cab. This shows
the actual tuber spacing and by adjusting a control knob the spacing can be
altered without leaving the cab.
A planter monitor is now being developed in the UK, which gives a
read-out of mean tuber spacing, area planted and plant popUlation (tubers
ha- 1). Such a device used in conjunction with a controller could ensure
much more accurate planting and control of the tuber population than has
been previously possible.
13.4.4 Bed planting

In regions where soil moisture conservation is important and irrigation
tends to produce slumping and greening of tubers near the surface, some
farmers prefer bed planting. Often beds comprise three rows which have
staggered spacing to even out competition between plants. Choices of
planter are limited for this operation though triple run cup planters are
available.
13.4.5 Mini- and micro-tubers

The development of mini- and micro-tubers in the seed industry has been
rapid and planters for these types of seed tend to be adaptations of maize
drills. There is a need for improved designs for handling this type of seed.
Though some cultivars are quite spherical in the small sizes, others still
display elongation. Some of the techniques used for drilling crops such as
sugar beet may be applied to these small tubers.
13.5 HARVESTING
13.5.1 Potato quality and damage

In the last decade, there has been a rising demand for quality potatoes.
Sales through traditional outlets such as the greengrocer and small corner
shop have declined as the supermarkets have increased their share of the
retail market. The great buying power of the supermarkets has provided a
strong incentive for the farmer to meet a tight quality specification.
Likewise, an expanding processing industry has developed a clear set of
specifications for quality, size and shape.
The wide size and shape spectrum and the basic vulnerability of potatoes
to damage, makes a potato crop a difficult one to harvest and handle. The
avoidance of damage throughout the harvesting and handling chain and the
final corrective by efficient inspection prior to sale, must remain a prime
objective of the farmer. Along with this goes the need to maximize
operating rates to ensure timeliness at harvest and in meeting target
outputs in post harvest handling operations.
In the UK, the Potato Marketing Board Ware Prescription is the
statutory standard for quality and size. The quality regulation does not
allow the aggregate of damage (both external mechanical damage over
3 mm deep and internal bruising), blemished, green and mis-shapen
potatoes and a range of other defects, to exceed 5% of the sample by
weight. This is one of the strictest standards in Europe. The processing
industry has additional requirements and there is particular concern
regarding internal bruising which must not exceed limits set by individual
processors - generally not more than 5% of tubers with one or more
bruises.
In addition to the UK Prescription for New Potatoes and Ware Potatoes
published annually by the Potato Marketing Board, an international
quality standard used in Europe is set out in the Rules of Usages of Intra
European Trade in Potatoes - known as the RUCIP Rules. These rules
Harvesting 579
cover the requirement for quality pota,toes being moved within and
between European countries. The rules include seed, new and ware
potatoes. For ware potatoes, the tolerance by weight of the defects listed in
the UK Ware Prescription is, in the RUCIP Rules, 6%. In addition, a
maximum of 2% waste, such as dirt tare and totally unusable potatoes is
allowed.
Whilst the above regulations set the standard for general quality of
potatoes in the UK and in Europe, some retail food chains operate a higher
standard for both external defects and bruising. It would appear likely that
in future, more attention will be focused on skin condition, which, though
it may not necessarily affect the internal quality of the potatoes, does affect
consumer choice in a very competitive market. Studies of skin condition
and skinning resistance are being carried out at SCAE (Muir et at., 1990).
13.5.2 Methods of assessing external damage and bruising
(a) Damage index

With a view to relating damage occurring at harvest time to actual loss
likely to be incurred by the consumer, Robertson (1970) developed a
damage index. The index was based on the weight of tissue which had to be
cut out as potatoes were peeled, in order to remove unsightly areas of rot,
corking and other tissue damage. A broad classification was applied and
the classes then weighted by a constant according to the typical quantities
of damaged tissue found in each category. The classification was as follows:
Damage Category Constant

Undamaged
Scuffing (surface scoring) 1
Peeler - damage to 1.5 mm depth 3
Severe - damage deeper than 1.5 mm 7
The damage index is calculated as follows:

Damage index = % scuffed +3 x % peeler + 7 x % severe damage
It can be derived on a number or weight basis. The damage index has wide
applications in the UK and similar indices are used in other European
countries. Over a period of years the maximum depth for the peeler
category has been increased from 1.5 mm to 3 mm. This really alters the
original basis for the index, i.e. that it is a measure of the likely loss
damaged tissue means for the consumer. The 3 mm depth now used for
peeler in effect reduces the loading factor of 7 for severe damage and raises
the loading factor of 3 for peeler. The whole concept of the damage index
is currently being re-examined at SCAE.
(b) Bruising
Various methods of assessing the severity of bruising in tubers have been
tried. Ideally the total volume of discoloured tissue should be removed
from the tuber and its weight expressed as a proportion of the tuber
weight. Alternatively, the product of length, breadth and depth of the
bruised tissue could be used to estimate bruise volume (McRae et aI.,
1976). Both methods are laborious and not really practicable for assessing
large samples.
The most common way of quantifying bruising is to cut each washed
tuber with a multiple knife cutter into lO-mm thick slices. Holding all the
slices in the palm of the hand, an inspector can quickly count the number of
slices showing one or more bruises. The percentage bruising is the number
of potatoes in a sample of 100 with at least one bruise.
Attempts have been made to combine external damage and bruising
(McGechan, 1980) but since external damage and bruising represent two
different phenomena, and are not part of a continuum in terms of the level
of abuse applied to the potatoes, it is probably best to have a separate
damage index for each type of damage.
(c) Mechanical aspects of damage

External damage can be classified as splits, gouges and crushing. Splitting
is usually the result of impact on a hard surface, after dropping for example
from a conveyor to a hopper floor, or on a harvester from a web to a
transfer conveyor. During impact on a flat surface, the tuber surface
deflects, the contact area enlarging as the tuber decelerates. The zone of
contact forms the base of a cone which then behaves like a wedge. In
extreme cases where splitting is severe, cracks radiate from the periphery
of the contact area, the potato tissues failing in tension along the crack
lines (McRae et aI., 1976).
Crushing may be caused when the tubers sustain heavy impacts or are
trapped against solid parts of handling machinery. Impact against a web
rod can cause a narrow band of crushed tissue to be formed. Gouging is
generally attributable to collision with sharp objects such as the edges of
metal sheets or protruding bolts.
Whilst crushing and gouging are little affected by variety or other
considerations such as temperature, splitting is more dependent on these
factors. Some older potato varieties now rarely grown, for instance Red
Craigs Royal, in some seasons could be very prone to severe splitting with
only minor impacts. Generally, as the temperature of tubers falls, splitting
becomes more likely (Johnston and Wilson, 1969).
Bruising is a complex phenomenon which, though attributable to impact
damage to tissues, also depends on physiological factors. During impact,
the tuber cell contents in the impact zone may be distributed and rupture
can occur. Enzyme reactions are then set in motion which lead to the
Harvesting 581
development of black pigmentation (melanin) over a period of two or more
days. At least three types of bruise have been identified. These are:
1. black-spot - a crescent or spherical blackened zone which occurs at
various depths below the tuber surface;
2. crushing bruise - a substantial area of very blackened tissue is evident,
resulting from a greater impact;
3. shatter bruise - a blue-black zone with striations of darker hue where the
tissues have shattered.
Bruising appears to occur more frequently in high dry matter varieties and
in both cold and over-warm conditions. A number of factors affect the
occurrence of bruises (Table 13.1). Sometimes where the development of
blackening is inhibited by the presence of ascorbic acid, white bruising may
occur where the damaged tissues dry out without blackening.
Table 13.1 Factors affecting likelihood of bruising

Likelihood of bruising
Less More References
More turgid Less turgid Sawyer and Collin (1960)
Smaller cells Larger cells van Es et al. (1975)
Lower specific gravity Higher specific gravity Ophius et al. (1958)
Higher soil potash Lower soil potash Hughes et al. (1975)
Higher temperature Lower temperature Johnston and Wilson (1969)
Clay soil or clay loam Dry sandy soil PMB (1974)
Small potatoes Large potatoes McRae et al. (1976)
Short storage period Long storage period Ophius et al. (1958)
In storage, pressure bruising is found especially if turgor falls late in the

storage season. Though it is usually associated with deep bulk stores
(Rastovski and van Es, 1981) it has been known to occur in one tonne
boxes.
(d) Equipment for measuring damage resistance

A range of instruments has been designed to measure varietal resistance
to damage (McRae, 1985). Drop tests can provide some guidance to
designers of potato handling equipment, but varietal resistances to damage
by dropping have been found to vary so greatly between seasons that no
single variety is consistently outstanding in terms of damage resistance.
Nor have there been very satisfactory correlations between harvesting
damage levels and varietal resistances as assessed by drop tests. Improved
methods of carrying out the drop tests could increase their usefulness.
Kershaw and McRae (1985) describe a method of increasing the quantity
of useful data from drop tests by minimizing the number of tubers dropped
well beyond the mean drop height likely to produce 10% severe damage.
Further improvements to the drop test method could be obtained if the
steel impact block used in the experiments were to be replaced by a slightly
resilient surface which is more likely to simulate harvester webs suspended
between support rollers.
(e) Pendulum tests

A pendulum tester was described by Gall et af. (1967) and later an
improved version was introduced by Hughes et af. (1975). Tubers to be
tested are usualy cut in half transversely and held with some pre-loading
against a stop with a hole in it. The pendulum is released from a suitable
starting position, allowing it to swing through a pre-determined angle. It
passes through the hole in the stop and impacts against the potato sample.
The deflection, rebound height, coefficient of restitution and absorbed
energy can all be measured. Improvements are now being made to
pendulum designs, for example to ensure freedom from torsional vibra-
tions and to eliminate the need to make measurements on cut potatoes. It
would appear possible that a closer relationship between the pendulum
results and damage resistance shown in passing through harvesting and
handling machinery may be obtained.
(f) Skin strength

Until recently, little attention has been paid to tuber skin strength. Now,
with growing interest in skin quality and harvesting methods such as the
two-stage system, a means of measuring skin strength is being examined.
Results obtained so far by Muir et af. (1990) indicate that there are changes
in skin strength after harvesting and during storage, but determining the
significance of the changes will await at least a further season's work. There
is some evidence that skin strength in storage falls for about 4 weeks after
harvesting - suggesting that moving potatoes at this time could increase the
risk of scuffing damage.
13.5.3 Preparation for harvest

Prior to the harvest, the potato haulm is usually treated to arrest growth
and hasten skin set. In the UK, the crop is either sprayed down with a
desiccant or a haulm pulverizer is used. Haulm pulverizers may be front or
rear mounted on the tractor and comprise a pto driven rotor running inside
a hood and fitted with a series of flails which conform to the ridge profile.
Front mounted pulverizers have the considerable advantage of running
ahead of the tractor wheels, avoiding crushing haulm and folding it parallel
to the ridge flanks, where flails generally do not pick it up. The operation
Harvesting 583
of a rear mounted pulverizer could be improved by fitting crop deflectors
on the tractor wheels such as the SCAE 'Golden Slippers'. Pulverizers
must be set high enough to avoid damaging any potatoes lying near the
ridge crests or flanks. Though quite effective in a good standing crop,
pulverizers are less effective where the haulm is straggly, or has been
flattened and broken by wind and rain.
A general tightening of the regulations relating to the use of crop
desiccants in Europe and banning of some crop sprays has led to an
upsurge of interest in alternative forms of haulm treatment. Techniques
currently in use or being investigated are:
1. flame desiccation;
2. haulm pulling;
3. lifting and re-burying.
In some areas, especially where early potatoes are grown, green top
harvesting is practised, but there can be problems when immature tubers
remain attached to the stolons and may be lost or cracked in haulm
stripping rollers.
(a) Flame desiccation

Experiments have been carried out in Holland (Philipsen et al., 1974) for a
number of years and more recently in Sweden (Larsson, 1990) to initiate
haulm desiccation by flame, using liquid petroleum gas (LPG) burners
mounted behind the tractor and spanning two or four rows. The intense
heat causes cell rupture in the lower stems. In order to minimize gas usage,
the haulm is generally pulverized prior to flame desiccation leaving stumps
of about 200 mm length. The cost of flame treatment after pulverizing is
about 30% more than the cost of pulverizing and spraying with a reduced
concentration of desiccant. Total costs tend to be twice the cost of spraying
which may, however, not be an option in future. Typically a flame unit uses
60 kg ha- 1 of LPG and operates at 1 ha h- i for four-row units. There are
some dangers with LPG operation due to the need for large burners to be
fitted with pre-heaters. The Swedish unit uses hot water for pre-heating
rather than a proportion of the direct heat from the flame units. If either
type of system is adopted in the UK, stringent safety measures would be
necessary.
(b) Haulm pulling

A proportion of the seed crop in Holland (last estimated at 60%) is treated
with haulm pullers. A typical haulm pulling operation is preceded by
pulverizing, usually by a pulverizer mounted on the front of the tractor
operating the haulm puller. Stumps of around 200 mm length are left for
the puller to grip, and in good conditions, over 95% of all stems are pulled.
Snapping of the haulm is minimal if it is gripped by the machine as near to
the ridge crest as possible.
Haulm pullers usually pull two rows simultaneously. Dutch haulm
pullers use one of the following mechanisms:
1. pairs of inflated rubber rollers which pull in a substantially vertical
plane;
2. pairs of pulling belts which pull in a substantially horizontal plane;
3. pairs of inflated rubber tyred wheels which pull at around 45° to the
horizontal.
Several critical relationships govern the success of haulm pullers. The
ratio of the pulling unit speed to ground speed is important, so that the
rootstocks clear the ground before the gripping unit releases the haulm.
The roller units rotate with a peripheral speed of around 6.7 m S·1 at a
forward speed of 1.4 m S·1. The belt units operate at 3.1 m S·1 for a forward
speed of 1.6 m S·1. The belt units tend to pull the rootstocks clear of the
ground more effectively than the roller units and are also regarded as being
superior to the wheel units which are considered to have inadequate
pulling contact area.
Depth control is particularly important with haulm pullers. The belt type
puller (Fig. 13.6) is best set so that the lower edges of the belts run at or
just 1 cm below the crest of the ridge. If the stumps are gripped more than
100 mm above the soil, they snap more easily, because they are succulent
Figure 13.6 Belt type haulm puller. (By courtesy of Nimos.)

Harvesting 585
rather than woody above this point. Losses through uprooting the tubers
are minimized by shoes running under the belts along the ridge flanks. The
roller and wheel types likewise use stripping bars.
Attempts to pull haulm in the UK with imported machines have had
limited success. A single-row unit capable of delivering the haulm two rows
to one side developed at SCAE also operated with variable efficiency
(Hutchison and McRae, 1985). From experience in operating this machine
it would appear that the requirements to enable pulling to be successful in
the UK - with efficiencies of over 95% - would be:
1. very straight drills and the haulm growing upwards from the centre of
the drills, with little spread;
2. few stones which very easily block the belt type pullers;
3. good depth control as variations of over 20 mm can mean the difference
between efficient and unsuccessful pulling.
Advantages and limitations

Haulm pulling offers an alternative to spraying if successfully carried out.
In Holland, it has proved useful in control of Rhizoctonia in seed crops.
There are undoubted advantages for subsequent harvesting, since removal
of the haulm and root minimizes the need for much trash removal
equipment on harvesters. In a field experiment at SCAE, it was observed
that the forward speed of the harvester was doubled after haulm pulling
compared with green top lifting.
Though when conditions are ideal, haulm pullers can be effective, the
limitations of both design and haulm condition combine to ensure that the
operation is never 100% efficient. Stems which have been left unpulled
may have to be lightly sprayed down to avoid re-growth. Pullers at present
leave the haulm, stems and roots lying on the ridge top rather than
delivering them between the ridges. Generally, haulm is regarded as an
obstacle to faster harvesting yet if it is not dealt with properly, there will be
problems during harvesting through crop loss, damage to sound tubers and
reduced harvesting rate. In seed crops there are risks from diseases. A
more detailed account of haulm pulling research in Holland is given by
Bouman and Bouma (1983).
13.5.4 The harvesting operation
(a) Stages
Harvesting potatoes involves removing some 40 tonnes of potatoes per
hectare from 500 to 1500 tonnes of soil, stone and clod. The stages in lifting
the crop are as follows:
1. digging the ridge or bed;

2. sieving and conveying the potatoes and remaining stones and clods,
together with root, haulm and any weeds;
3. separating haulm and roots;
4. performing a final cleaning/separation operation;
5. transferring the potatoes to trailers, boxes or a bunker on the harvester.
On most harvesters, digging is by means of a solid or three part passive

share usually preceded by a diabolo depth roller running on the ridge and
haulm cutting discs on either side of the share. Tuber damage can occur
during digging, particularly at the transfer point from the share to the front
of the web. At this point the web rods can be travelling vertically at over
2 m S-1 and may damage the lowest tubers in the ridge.
Alternatives to flat shares have been developed. Johnson (1974)
improved digging rates with a vibrating share and Carruthers (1980) found
that even small tubers were brought to the surface by a vibrating share and
parallel rod combination. Some blockages by weed did occur and the share
did not work so well on sloping ground when digging downhill., due to roll
back.
Power driven disc shares with static scrapers have proved effective on
some harvesters marketed in eastern Europe. A disadvantage was con-
sidered to be the rate of wear of the discs in abrasive soils. Some harvesters
are still marketed with powered side discs for cutting through trailing
Figure 13.7 Disc share with di-cone depth roller. (By courtesy of SCAE.)
Harvesting 587
haulm, rather than for the main digging function. An improved power
driven double disc share with inverted dished powered scraper discs (Fig.
13.7) has been found to have a number of clear advantages over flat shares
(Hutchison and Fleming, 1980).
Potato damage attributable to this share design is reduced by 23-29% in
comparison with the flat share, mainly due to delivering the ridge well clear
of the front of the primary web. Draught is greatly reduced (by 60-88%)
and due to better clod break-up, some 37% less clod passes over the rear of
the primary web. The only drawback is a tendency to lift 18-40% more
soil. This, however, can be sieved out readily on the webs and should not
prove a real problem with good depth control.
The use of powered disc shares enables the front of the primary web to
run above the soil level and this is likely to minimize wear and blockage by
small stones. Neither diabolo rollers nor side discs are required with disc
shares, a single roller running between ridges sufficing to control the
digging depth.
(b) Soil sieving

The primary and subsequent webs on modern harvesters perform the task
of sieving the soil and breaking up soft clod. Usually vertical agitation is
used to effect the sieving either by eccentric idlers, or rocking support
rollers. McGechan (1977), in experiments at SCAE, showed that horizon-
tal agitation gives superior soil sieving rates and much reduced tuber
damage. An agitation amplitude of 25 mm at a frequency of 3 Hz was
found to give good results. The sieving process takes twice as long with a
clay soil, compared with a light sandy soil and consequently on heavy soils,
maintenance of a soil cushion on the webs to reduce tuber damage is
seldom a problem. Trials of harvesters fitted with horizontally agitated
webs have shown that this form of agitation can reduce tuber damage by
from 15 to 39% and increase sieving rate by 15% or more.
Continental webs used on all European harvesters are constructed of
solid spring steel rods 9-10.3 mm in diameter, rivetted to carrier webs. By
substituting hollow rods 12.7 mm in diameter, the 40% lower mass has
been found to produce surprising reductions in potato damage. McRae and
Hutchison (1988) showed that with hollow web rods there was 16-33% less
external damage and up to 30% less bruising. Recent trials indicated that
up to 60% less external damage can occur. The rods are 30% stronger than
the slightly smaller diameter solid rods and their performance with respect
to potato damage is mainly due to the mass reduction and not to bending
during impact.
A harvester incorporating powered discs and horizontally agitated
hollow web rods has proved capable of causing 22% less potato damage
and operating at higher lifting rates, compared to conventional machines.
This harvester, called 'Pulsar' through the method of agitation, has been
Figure 13.8 'Pulsar' general view.
licenced for manufacture but is not yet commercially available (Figs. 13.8
and 13.9).
(c) Cleaning systems on harvesters

It is unfortunate that potato damage during harvesting is cumulative and all
efforts to remove fine clod root and haulm tend to produce slight increases
in damage, particularly in the scuffing and peeler categories.
Manned harvesters are usually provided with separation systems based
on sloping 'hedgehog' belts. The slope is adjustable and, when set
carefully, this separator can remove useful quantities of stones and clods,
but suffers from the fact that the shape characteristics of both stones and
clods overlap those for potatoes.
Star cleaners, which use curved fingered rollers in series can be useful for
removing fine clod, though in some conditions may cause damage. A
cleaning device recently developed in the USA comprises a series of pairs
of rollers driven by a hydraulic motor. If any of the rollers become jammed
by a stone, they reverse almost instantaneously. This idea has some useful
potential.
The development of stone windrowing systems which greatly reduce the
quantity of clod and stone which pass over the picking tables on harvesters,
Harvesting 589
Horizontally agitated
rear web
Powered scraper discs
Double disc share
Figure 13.9 Pulsar diagram. (By courtesy of Farmers' Weekly.)
or in the case of unmanned machines pass directly to the trailer, has

reduced the urgency of producing new forms of separator. The first
electronic separator used the principle of X-ray transmission through plant
material to differentiate between potatoes and stones. This device was
fitted for some years on harvesters but was more successful as a farm store
separator when unmanned harvesters became popular. Other types of
separator have been marketed, including an infrared scanning unit. The
widely successful 'Airvac' system used in eastern USA and Canada relies
on the ability of potatoes to float in an airstream velocity of around
40 m S-1. The system has been applied in limited areas of the UK. The
power consumption to drive the large paddle fan used to generate the
appropriate airflow is up to 40 kW and this is a considerable drawback.
(d) Types of harvester

The main types of harvester used in the UK and other European countries
are:
1. single-row manned;
2. two-row manned;
3. two-row unmanned (some with optional limited manning);
4. four-row self propelled.
Generally, the forward speed of each type is about the same. As a result,
two-row harvesters tend to harvest about twice as quickly as single-row
machines. The headland turning time which is unproductive takes slightly
longer per turn with two-row machines compared to single-row machines
but the number of turns is halved.
In some parts of Europe, bunker model harvesters are popular. These
are useful where the haulage distance to the farm store is considerable and
the bunker can then act as a buffer store should transport delays occur.
They are not suited to wet conditions because of the additional tonne or so
of potatoes they carryon board and may pose problems on sloping ground.
A recent upsurge in interest in self-propelled harvesters has occurred.
These machines have advantages in manoeuvrability, good driving position
to see the main conveyors, and good weight distribution. The four-row
versions are very large machines but have a high capacity and reduced
headland idle time per hectare compared with single- and two-row
machines.
(e) Two-stage harvesting

Two-stage harvesting is the term applied to the system of digging the crop,
removing some of the soil and clod and laying the potatoes back on the
ground in the form of a windrow, which is later picked up by a lifter.
The idea is not a new one. It had a brief start in England in the 1950s as the
Johnson six-row system. A windrow was created with two pairs of outside
rows being windrowed between two undug rows. The whole windrow was
then lifted by a second machine which delivered the potatoes into a trailer.
Scottish trials of two-stage harvesting were carried out at the then NIAE
Scottish Station, Mid Calder in 1959. A simple single-row digger was used,
followed two hours later by a lifter fitted with a round bar share. Potato
damage figures showed that there was an increase of 20--27% in damage
index after re-lifting the windrow.
In the eastern states of the USA, particularly Maine, and just over the
Canadian border in New Brunswick, windrowers have been in general use
for 20 years or more. Sometimes, two rows are deposited in between two
undug rows and after a variable time interval, a digger lifter completes the
lifting operation. Typical harvesting rates of about 1.25 ha h- 1 in a 17 t ha- 1
crop were recorded in the 1970s.
Since the early 1980s, there has been a revival of interest in two-
stage harvesting in parts of Europe. In the Luneberg Heath area of
Germany, the soils are particularly suited to the system. More recently in
Harvesting 591
this country, a UK manufacturer has introduced two-stage harvesting
machinery.
A number of advantages have been claimed for two-stage harvesting.
1. In the windrow, the potato skins dry and soil is knocked off easily
during re-lifting. This is a particular advantage in parts of Germany
where dark soils cause unsightly mottles on potato skins when the
potatoes are lifted in the conventional way. In the UK, less soil on the
skin prior to storage might improve the efficiency of fungicides.
2. Tubers are usually warmer when they are re-lifted and therefore less
damage prone. At SCAE, trials in autumn 1989 showed that there was a
rise of 4°C in tuber temperature over a two hour period after digging
(Muir et al., 1990).
3. The better skin condition and apparent improvement in skin strength
reduces ingress of disease through scuffing damage.
4. Faster lifting rates are possible especially with four- and six-row
windrows.
The equipment
Two main types of system are now being evaluated in the UK. One system
employes a four-row digger with two transverse conveyors putting the
potatoes into a single windrow. The soil on which the potatoes are laid is
firmed by a roller running just ahead of the potatoes and lateral spillage is
avoided by two flanking ridges produced by side discs. Lifting is achieved
by a modified harvester without diabolo rollers. The normal shares are
replaced by a full width onion share which is held just below the windrow
by two small depth wheels.
The other system used creates a windrow from two rows. Again a heavy
roller prepares a trough ahead of the potatoes to contain the windrow.
Lifting is carried out by a simple harvester with a small diameter powered
steel roller share and a light rubber roller above it, to transfer the windrow
smoothly to the first web.
Field performance
The Potato Marketing Board carried out damage tests with the two-row
windrow system in 1986 and 1989. Though low damage levels were
recorded in 1989, Statham and Cunnington (1990) found that there was no
significant difference between two-stage harvesting and direct harvesting.
An over-winter assessment of the storability of crop lifted by the two-stage
system showed little appreciable difference in crop quality compared with a
crop lifted in the conventional way.
Whether or not the two-stage harvesting system becomes more widely
accepted will depend on a number of factors. The low damage figures
obtained in UK trials were on land with an insignificant amount of stone
and clod. The presence of either, even in modest quantities, could lead to
additional damage during lifting. Climatic factors could also be a problem
with windrows being rained on, though local weather forecasts are now
much more dependable than in the past. The benefits of soil loss and skin
drying have yet to be fully evaluated in terms of storability. Skin strength
measurements at SCAE (Muir et al., 1990) up to two hours after digging
have not shown consistent improvement in the windrow but more work is
needed in this area. Impressive lifting rates have been claimed for both of
the systems used, but account should be taken of the additional labour and
machinery input required in preparing the windrow prior to the final lift.
If the four-row and two-row systems are compared, the four-row
windrower is likely to be considerably faster than the two-row machine. In
addition to lifting twice as many rows at once, headland turning time is
reduced. Windrowing four rows into one is likely adversely to affect the
drying out rate of potatoes in the deeper windrow. Some further evaluation
of this system will be necessary before it is likely to be widely adopted in
the UK.
13.5.5 Transport from field to store

Potatoes are transported to the store, or sometimes directly to processing
plants or pack houses, in a variety of trailers and containers. Selecting the
most suitable transport involves considering a number of factors including
soil conditions, haulage distance, unloading arrangements and the output
of the harvester or harvesters with which the transport is associated.
In the USA, it is common for trucks of 12 tonnes capacity or more to be
used to take potatoes directly from the harvester to the farm store, or some
other location. This type of transport has advantages where haulage
distances are considerable and capital is available to invest in the trucks. In
the UK and most parts of Europe, tractor drawn trailers with capacities of
a few tonnes up to 12 tonnes are preferred but, unless soil conditions are
good, even smaller trailers must be used.
A problem in filling large trailers from the harvester is the drop height
between the delivery conveyor and the trailer floor. Trailer floors can be
lined with foamed plastic sheet, but the cushioning effect tends to be
confined to the first two or three layers of potatoes. Automatic height
controls on the harvester conveyor are useful in minimizing drop height
when filling the trailers. Break fall chutes and fall arrester hands across
trailers can help to reduce damage (de Haan and Van Zwol, 1974). For
long hauls over rough roads, some springing in tandem axles may be an
advantage.
Care must be exercised when tipping at the store into bulk hoppers and
rising tail-gate trailers can help to reduce cascading at this stage. Some
farmers operate a fleet of self-emptying trailers with floor conveyors driven
by an electric hook-up at the store. Due to the great height variations in
different designs of trailer there is often an incompatibility between the
Handling and grading 593
trailer height and the reception hopper, or conveyor height, leading to
considerable risk of damage.
So far no integrated approach to transport and unloading seems to have
gained commercial acceptance. Experiments in the 1970s with such a
system combining a sprung clamshell trailer and associated unloading
hopper (McRae and Fleming, 1978) showed that transport and unloading
could be accomplished with a very low damage index in the range 4-13.
The rapid extension of box storage systems even for ware potatoes
means that in some cases boxes are loaded in the field. The boxes may be
carried in groups of four to six on a flat trailer. It is important to provide
cushioning at the junctions between boxes.
13.6 HANDLING AND GRADING
Though it is generally accepted that harvesting and transporting the crop to

the farm store is the most damaging of all the operations in potato
handling, the second most important is considered to be handling in and
out of store. In a questionnaire (Barr et al., 1987), sent to all UK potato
growers, 21% of those who replied considered this operation to be the
single most important cause of damage to potatoes.
13.6.1 Handling into and out of store

Apart from problems cited regarding heights of trailers in relation to
reception facilities, delivery of potatoes to the bulk heap in the store can
cause damage and if the crop has much adhering soil, soil coning which
may lead to local hot spots will occur unless the bulk loading conveyor has
a slewing head. In deep storage, an added problem is damage to potatoes
avalanching down the face, which can be combatted to an extent by
terraced filling. A number of bulk conveyors are now fitted with height
control sensors based on ultrasonic proximity switches and these help to
minimize drops. On some farms instead of bulk conveyors, high lift tipping
trailers are used for store filling.
Much of the UK crop is loaded out of store by root buckets, mounted on
tractors or rough terrain vehicles. Powered scoops and buckets mounted
on fork lift masts may also be used. Root buckets can cause up to 9%
severe potato damage (PMB, 1974) unless handled carefully. It is prefer-
able to use slightly under-powered tractor units to minimize shock loads on
the bulk heap.
Box handling requires special equipment. Mobile box tipplers are
required if the boxes are to be emptied in installations which cannot readily
be serviced by static box tipplers. Whether these are forward tipplers or
rotary tipplers there is poor control over discharge rate and drop height. A
control lid for rotary box tipplers has been developed at SCAE (McRae et
al., 1985) which reduces damage to the first 100 kg or more delivered from
the box to the reception hopper by 75%. A range of static box tipplers is
also fitted with this control lid.
13.6.2 Handling within stores and grading areas

In most grading and handling operations, the effect of drops must be
minimized to reduce cumulative damage to the potatoes. Drops may be
fixed, as when potatoes are transferred from one conveyor to another, or
they may vary depending on the level within a holding hopper for instance.
Several types of fall breaking device are available to help to reduce impact
velocity if the potential drops are substantial. De Koning and Lerink
(1981) cite an experiment to reduce damage using 12 mm diameter strings
35 mm apart spanning a trailer or hopper and connected in the middle in
groups of three. They found that only 32% of the incoming material
passing through the strings slowed down to an acceptable impact velocity.
Problems did arise with haulm and weed. More elaborate equipment for
minimizing damage through dropping, such as zig zag chutes, is described
by de Haan and Van Zwol (1974). In using these chutes, there is a risk that
mutual collisions between potatoes may cause some bruising. Nevertheless
if the recommended throughputs are not exceeded, such devices can make
a valuable contribution to damage redudion.
Where drops are unavoidable, careful regulation of levels, for instance
by using proximity switches in hoppers, can avoid impact of potatoes on
the hopper floor or elevator belt flight edges. A range of cushioning
materials is now marketed for use in areas where there is an impact risk.
Some are homogeneous expanded plastic foams based on polyurethane or
other resilient plastics. Sheets of these materials can be attached to hopper
sides with industrial adhesives. Wear resistance 'is an important considera-
tion and composites can give better service. This type of cushioning has a
resilient base material, bonded to a top layer of woven plastic fibres. Muir
(1990) showed that provided the thickness of the underlying material is at
least 12 mm, and 20 mm would be even better, cushioning is excellent.
Figure 13.10 compares the energy absorption capability of one homo-
geneous and one composite foam. Regular inspection of all cushioned
surfaces is needed to spot wear points, before the cushioning ceases to be
effective.
13.6.3 Grading
The term 'grading' is applied rather loosely and can mean either sizing or
inspection. These operations are at present separate, but rapid develop-
ments in electronics and machine vision suggest that it is becoming possible
to carry out both operations at once.
400
300
c:
:8200
~
Q)
"8
Q)
0 100
o Smm composite foam
" 5.5mm homogeneous
O+---,----r--~--~----r_--._--,_--.
10 20 30 40
Drop height (em)
Figure 13.10 Comparison of cushioning materials (Muir, unpublished).
13.6.4 Sizing
The aim of sizing is to present samples with sufficient uniformity to satisfy
the perceived market requirements. Examples are potatoes for baking,
chipping or seed. Uniform size simplifies packing and filling count boxes
and enhances eye appeal on the market stall or supermarket shelf. Though
visual uniformity is subjective, sizing by eye in comparison with a square
mesh hand riddle for citrus fruit has been shown to give an error of little
over 3% (Kaser, 1965). No comparable figure is available for potatoes but
it is likely that judgements of length and girth would give errors of under
10%.
In the UK, the part of the Potato Marketing Board Ware Potato
Prescription referring to size permits only 5% of the sample to consist of
outgrades when measured against a square mesh standard. It has been
recognized for some time that dimensions such as major and minor axes in
the case of tubers with an elliptical cross section which govern their passage
through a square mesh aperture, express size inadequately. For some
markets, length is also measured with a view to improving the grades.
Kolchin and Smekhunov (1975) confirmed the earlier work of Goryachkin
in the 1930s which suggested that weight should be the ultimate sizing
criterion. For many practical purposes this is correct. The use of weight
reduces anomalies with long varieties which can vary in weight by as much
as ±40% if sized by the traditional square mesh criterion.
McRae et at. (1986) carried out measurements on 6000 tubers of ten
leading UK maincrop varieties. In later experiments, a further 4000 tubers
were examined. The square mesh riddle size and single dimensions such as
length, breadth and thickness were all found to be poor predictors of
weight, with a range of up to ±41 %. Riddle size is slightly more effective
as a predictor of weight than thickness for instance, which is the criterion
for parallel roller sizers. Accurate weight grading can be related to volume
grading if the specific gravity is known. In practice, tests at SCAE have
shown that volume can be used as a predictor of weight with a range of only
±1.8%.
13.6.5 Types of sizer
For ware size potatoes agitated mesh screens are widely used in the UK.
The agitation encourages long tubers to orientate so that they pass through
the screen meshes if they are below the mesh size. A recently introduced
mesh screen uses horizontal rather than vertical agitation to reduce risk of
damaging the tubers. Accuracy of this type of grader ranges from 95 to
98%. Whilst most screens use 3 mm diameter wire mesh, a 12.5 mm plastic
link screen (developed at SCAE by Carruthers) has been introduced. This
type has several advantages. Farm trials have shown that damage attri-
butable to impacts when potatoes are transferred from a feed conveyor to
the screen and during passage over the screen, is reduced by 50%. In cold
conditions, the incidence of skin splits such as 'thumbnailing' can also be
considerably reduced. The plastic screen is much lighter than the wire
screen and one man can carry and change a 1.5 m wide plastic screen single
handed. Wear rates can also be very low.
Many seed growers who need to produce a range of sizes for different
customer requirements prefer oscillating screens based on the jog trough
principle. Some screens are fitted with recirculating clearing bars which
move back and forth under the riddles dislodging any trapped potatoes or
clods and rubbish.
For the high throughputs required if pre-storage grading is practised, a
number of farmers have turned to expanding roller sizers. The simplest of
these machines would have plain rollers. Hutchison and McRae (1980)
measured the efficiency of a plain roller expanding sizer and found that
even with a width of 0.6 m a throughput of 12 t h- 1 at 85-91% sizing
efficiency was possible. Larger machines are marketed which provide
greater sizing accuracy using staggered moulded rollers, or drop rollers to
improve performance (Fig. 13.11).
Alternatives to mechanical sizers are now being developed and a number
of machines using either optical or gravimetric sizing methods have
appeared on the market. Of the optical sorters one of the first types
developed in the USA uses a square aperture carrying two sets of infrared
emitters and receivers, one set mounted in a bank at 90° to the other. A
signal derived from a series of cross sections of the tuber passing the
emitters is integrated after passage of the potato through the aperture is
complete and allows a volume estimate to be made in time to route
potatoes by means of pneumatic fingers to an appropriate take-off point.
Figure 13.11 Expanding roller sizer. (By courtesy of Peal.)
The device requires a singulation system and there is a need to limit tuber
velocity to 1.5 m S-l to avoid tuber damage.
In the UK, a machine vision system for size grading has been developed
by the Silsoe Research Institute (Marchant et al., 1990) which uses a
camera to scan the tubers as they rotate. The signals from the camera are
analysed to estimate the volume of each tuber. Length can also be
measured if required (Fig. 13.12).
Weight graders are currently used for baker-sized tubers. Usually the
potatoes are singulated in channels, the singulating systems being derived
from fruit grading machines. Some hand assistance is often required and
smaller throughput weight graders depend on hand feeding and spacing to
obtain accurate results. The weigh head on these machines is typically a
slack belt weigher using a viscous damped load cell. Auto-taring is
provided to avoid errors through dirt and debris accumulation. So far, high
throughput weight graders, e.g. of the order 15-20 t h- 1 for the whole crop,
though technically feasible using multiple weigh heads are likely to be too
costly for general use for some time to come. Combined sizing and
inspection will be considered later in the chapter.
With the currently rapid development of electronic sizing methods, the
existing sizing standard based on the square mesh riddle requires to be
related to a weight standard. The results of tests on ten leading maincrop
cultivars have shown that two potatoes of the same cultivar can differ in
size by almost as much as two potatoes from different cultivars. Whilst the
Figure 13.12 Quality inspection and sizing machine. (By courtesy of Loctronic
International. )
two sizing criteria - square mesh and weight - are different and cannot be
entirely reconciled, an analysis of the test data by Glasbey et al. (1988) has
provided the nearest threshold weights which minimize misclassification by
the square mesh criterion (Table 13.2).
13.6.6 Inspection
Inspection is the second grading process which usually follows sizing,
although sometimes severely damaged or defective potatoes are removed
during pre-grading prior to storage. The operation is almost entirely
manual in the UK, but the application of electronics and machine
vision systems is likely to make inspection an automatic process in the
future.
Table 13.2 Square mesh and
weight sizing standards
(Glasbey et ai., 1988)
Mesh size (em) Weight (g)
3.5 40
4 55
4.5 75
5 100
5.5 130
6 170
6.5 210
7 260
7.5 320
8 380
8.5 450
Inspection tends to be under-rated, considering that it is a key operation

in the whole sequence of producing a marketable product. This may be due
to a conflict between reaching a standard and maximizing saleable yield.
Whilst it is important that the out-turn from inspection conforms to the
PMB Prescription, over-zealous inspection where the defect level is fairly
near 5% can lead to actual financial loss. It is therefore in the interest of
the farmer to regulate the inspection process carefully, to make sure that
the required standard is being maintained.
The most common methods of inspection are to use roller tables, or a flat
belt with a means of turning the potatoes over as they traverse the
inspection area.
There have been several studies of the effects of altering the operating
settings on roller tables. Some of the earliest research was carried out in
the USA by Malcolm and De Garmo (1953). Artificial potatoes (made
from wood) were used extensively and the quantity of field test data with
real potatoes was limited. Hunter and Yaeger (1970) considered feed rate,
percentage of defective tubers, translation speed and roller rotation rate
and direction. Artificial blemishes were used in the tests.
Tests using potatoes with naturally occurring blemishes were carried out
at PMB Sutton Bridge Experimental Station (PMB, 1972, 1973). A grading
efficiency (the percentage of defective material removed) of 33% at 2.2 t h- 1
for one operator was recorded and this rose to 57% at 0.7 t h- 1 . With two
operators an efficiency of 67-72% was achieved at the same throughputs.
A major difficulty with research on roller table operation has been the
amp ling process which is laborious and prone to error. A new technique
was developed at SCAE by Elstob et al. (1990) to enable more extensive
studies of the effect of altering each of a number of possible settings on the
efficiency of inspection.
Before a test run, a sample of potatoes was made up consisting of a
known proportion of defective potatoes. The defective potatoes (selected
by the inspector who would carry out the test) were marked with an
invisible ink which fluoresced in ultraviolet light. Code marks were used to
differentiate between mechanical damage, and greening. The sample was
then circulated round a low damage recirculating conveyor system and
over the roller table. After passing the inspector, who moved apparently
defective potatoes to one side of a partition on the table, the two s,treams of
potatoes were passed through a lighting box fitted with powerful ultraviolet
lamps, which caused the normally invisible markings to fluoresce. The
proportion of sound and defective potatoes in each stream was then
identified from a video recording made throughout each run.
In addition to altering throughput, defect proportion, roHer translation
speed, rotation rate and direction, tests were carried out to determine the
effect of no rotation, partial rotation (zero rotation followed by brief
rotation of one roller) forward presentation and the use of artificial
markings, on grading efficiency. The following conclusions were reached.
(a) Roller translation speed

There is a steady decline in sorting efficiency in the range 6-10 m min- l for
the roller translation speed.
(b) Roller rotation rate

The rate seems to have little effect on sorting efficiency with small tubers
but a greater effect with larger tubers. The direction of rotation seems to
matter more to individual operators than the rotation rate (Fig. 13.13).
There is a very serious problem in rejecting sound tubers especially if they
100
x Forward Rotation
o Reverse Rotation
~ 95
>-
u
c
OJ 90
'0
~
:E
OJ
OJ
c 85
t
0
~
en
Cii 80
Q;
>
0
75
3 4 5 6 7 8
Roiler rotation (revs m-1 )
Figure 13.13 Relationship between roller rotation direction and efficiency of

roller table,
are large ware or baker size (above 60 mm). This could be related to the
reduced numbers to be scanned per second even with a full table, when the
tubers are large. It has important economic consequences due to the high
value of baker sized potatoes.
(c) Throughput
The efficiency of the inspection operation falls as throughput increases from
12 000 to 28 000 tubers per hour, especially above 18 000 tubers per hour.
(d) Defect percentage

When the defect level exceeded 20%, the out-turn did not reach the PMB
Standard. A higher efficiency was measured when only one type of defect
only (e.g. greening or mechanical defects) was present: the greater the
number of different defects being searched for, the lower the overall
efficiency.
(e) Zero rotation

There is a considerably lower inspection efficiency with zero rotation
compared with normal roller rotation. A higher than normal rejection rate
for sound potatoes also occurs. This finding suggests that flat belts used for
inspection, even with a turning mechanism half way along their length,
could reduce inspection efficiency.
(f) Partial rotation with two operators

This arrangement again gave lower efficiency although the difference was
small at high translation speeds.
(g) Forward presentation

This method (contrary to the findings of Malcolm and De Garmo, 1953)
did not show an overall gain compared with side presentation. There were
practical problems of grasping tubers and visual difficulties. An exception-
ally high number of sound tubers was rejected.
(h) Artificial markings

The study using the video sampling technique clearly showed the unrelia-
bility of using artificial markings for such tests - the inspection efficiency
was very high, with no incorrectly rejected sound tubers.
At least one UK manufacturer supplies roller tables with drives which

100
-UNDAMAGED
- - - FRESH WOUND
----- OLD WOUND
~
Q)
\
u
c:
ro 50 \
\
13 "...... '\
' .... _-_ ..... -_.- .. , ..
Q)
'ii5 .......
a:
(a) 0
500 1000 1500 2000
Wavelength (nm)
100
-HEALTHY
---SOFT ROT
>R
o ----- DRY ROT
Q)
u
~ 50
~ '\
'i 5
a:
~ .......
(b) o+-------~------~------~
500 1000 1500 2000
Wavelength (nm)
Figure 13.14 Reflectance characteristic curves of healthy, damaged and diseased

tubers. (From Muir et at., 1982.) (a) Damaged and undamaged tubers and (b)
healthy and diseased tubers.
allow control over translation and roller rotation speed. There would be
additional benefit in allowing for the inspector's preferred direction of
roller rotation.
Apart from the operating parameters discussed, lighting has consider-
able effect on operator performance. There is quite a wide range of
tolerances to light intensity. Zegers and Van den Berg (1988) reported that
a range from 500 to 2000 lux does not appear significantly to affect
inspection performance, but that the quality of the light does have a
significant effect. A 'cool white' light with a colour rendering index of
85 Ra was found to give the best results with potatoes.
Whilst the range of lighting intensities tolerated can be considerable, the
lighting in some farm installations is clearly well below requirement and
often the inspection tables run in very dark sheds with poor background
lighting. The problem is sometimes exacerbated by heavy deposits of dust
on the fluorescent tubes.
Figure 13.15 Optical disease detection machine. (From Muir et aI., 1982.)
13.6.7 Machine vision

Though potato inspection is a much more difficult operation than most
industrial inspection processes where the objects being inspected are
relatively uniform in size and shape, there are now devices for inspecting
agricultural produce which can be adapted for potatoes. A typical system
comprises a scanning camera, a signal conditioner and a computer which is
instructed to make decisions to accept or reject the produce on the basis of
how nearly it conforms to the ideal standard. It is possible for instance to
select on the basis of surface area occupied by a blemish, or colour, such as
greening. The operation of scanning for blemishes can in certain situations
also perform a sizing operation where length and cross section are analysed
(incorporating a series of views) and a volume estimate made (Fig. 13.12).
The most difficult inspection operation using the machine vision
technique is detection of disease - sometimes at a sub-clinical stage. Muir
et al. (1982) found that the near infrared reflectance curve of diseased areas
in potatoes compared with healthy tubers was different for each of a range
of common diseases. Some 14 diseases have so far been identified with this
system. Figure 13.14a shows the reflectance characteristic curves for an
undamaged and a damaged tuber, and for a healthy compared with
diseased tuber (Fig. 13.14b). At present a scan of 1500 pixels (3 mm
diameter areas) on the surface of each potato takes about 4-5 s, but the use
of a parallel processing computer is likely to reduce the time taken, thus
enabling commercial throughputs to be possible. This equipment, which is
shown in prototype form in Fig. 13.15, is likely to have particular signifi-
cance for inspection in the seed potato industry.
In the near future it should be possible for large scale packing lines to be
fitted with automatic sizing and inspection systems to enable produce
quality to be much more closely controlled than it is at present.
13.7 CONCLUSIONS AND FUTURE PROSPECTS
Stone and clod windrowing is an essential technique to ensure minimum

crop damage at harvest. There is considerable market interest in the
development of a new planter which incorporates accurate and easily
altered spacing with high speed operation for a wide range of tuber sizes
and shapes. The commercial exploitation of horizontal agitation for sieving
soil on the potato harvester will further reduce tuber damage compared
with vertical agitation mechanisms. Without significant extra complexity, a
multi-purpose self-propelled unit could be designed for both crop estab-
lishment and crop harvesting. This would have the advantage of containing
fixed costs which currently represent about 40% of the total crop produc-
tion costs. Machine vision for disease and defect detection is an exciting
future prospect not only for potatoes but also for a wide range of other fruit
and vegetable crops.
REFERENCES
Barr, L.H., Wood, J., McRae, D.C. and Hutchison, P.S. (1987) A survey to
establish the current problems of potato crop mechanisation. Dept Note SIN/
482, Scot. Inst. Agric. Engng, Penicuik, (unpub!.).
Bouman, A. and Bouma, J. (1983) The effect of the potato haulm on the haulm
pulling efficiency. Agric. Engr, 38(1), 25-32.
Carruthers, J. (1980) Vibratory share for lifting twin rows or beds of potatoes and
vegetables. Dept Note SIN/294, Scot. Inst. Agric. Engng, Penicuik (unpub!.).
Elstob, R.H., Kirkland, J., McRae, D.C. and Fleming, J. (1990) Investigation of
the factors which limit the efficiency of operation of roller grading tables.
Abstracts of 11th Triennial Conference, Eur. Assoc. for Potato Res., Edinburgh,
pp. 239--40.
Entz, M.H. and La Croix, L.J. (1983) A survey of planting accuracy of commercial
potato planters. Am. Potato J., 60(8), 617-23.
Gall, H., Lamprecht, P. and Fechter, E. (1967) First results with a rebounding
pendulum to determine the susceptibility of potatoes to damage. Eur. Potato J.,
10,272-85.
Glasbey, C.A., McRae, D.C. and Fleming, J. (1988) The size distribution of potato
tubers and its application to grading schemes. Ann. Appl. Bioi., 113, 579-87.
References 605
Haan, P.H. de and Van Zwol, B.H. (1974) Check ways and chutes for potato
transport. Pub!. 279, IBVL, Wageningen.
Hughes, J.C., Grant, A. and Faulks, R.M. (1975) Susceptibility of tubers to
internal bruising. Potato Res., 18, 338-9.
Hunter, J.H. and Yaeger, E.C. (1970) Use of a Float Roll Table in Potato Grading
Operations. Bull. 690, University of Maine, Orono.
Hutchison, P.S. and Fleming, J. (1980) An investigation into the performance of a
twin disc share for potato harvesters. Dept Note SIN/280, Scot. Inst. Agric.
Engng, Penicuik (unpub!.).
Hutchison, P.S. and McRae, D.C. (1980) A variable grader for fruit and
vegetables. Dept Note SINJ293, Scot. Inst. Agric. Engng, Penicuik (unpub!.).
Hutchison, P.S. and McRae, D.C. (1985) An investigation into haulm removal in
the early potato crop. Dept Note SIN/445, Scot. Inst. Agric. Engng, Penicuik
(unpub!.).
IMAG (1978) Series of Tests of Potato Planters. IMAG publication 96, Wageningen,
Netherlands.
Johnson, L.F. (1974) Design and field testing of a low damage potato harvester.
Am. Soc. Agric. Engrs, Paper 74, 1509. St Joseph, Michigan 49085, USA.
Johnston, E.F. and Wilson, J.B. (1969) Effect of soil temperature at harvest on the
bruise resistance of potatoes. Am. Potato 1., 46, 75-82.
Kaser, L.O. (1965) Citrus sizing. Am. Soc. Agric. Engrs, Paper No. PC 65-13,
St Joseph, Michigan 49085, USA.
Kershaw, C.D. and McRae, D.C. (1985) Sequential design for estimating levels
with low response rates in potato drop tests and other binary response
experiments. 1. Agric. Sci., Camb., 105,51-7.
Kolchin, N.N. and Smekhunov, E.A. (1975) Promising principles of grading
potatoes and vegetables. Trakt Set" khozmash., 10, 19-2l.
Koning, K. de and Lerink, P. (1981) Fall breakers for potatoes and decelerating
devices to prevent damage. Landbouw Mechanisatie, 32(7), 663--6.
Lang, R.W., Bevis, J. and Franklin, M. (1986) The performance of seed of
different sizes under varying conditions. Proc. Eur. Assoc. for Potato Res.
Agron. Section, Northern Ireland, pp. 17-18.
Larsson, K. (1990) Destroying of potato haulm by flaming. Abstracts of 11th
Triennial Conference, Eur. Assoc. for Potato Res., Edinburgh, p. 497.
McGechan, M.B. (1977) An investigation into the relative effectiveness of various
riddling motions for removal of soil from potatoes. 1. Agric. Engng Res., 22,
229-45.
McGechan, M.B. (1980) An investigation into the damage sustained by different
varieties of pototoes during riddling to remove soi!. 1. Agric. Engng Res., 25(4)!
345-53.
McRae, D.C. (1985) A review of developments in potato handling and grading. 1.
Agric. Engng Res., 31, 115-30.
McRae, D.C. and Fleming, J. (1978) A cradle trailer for potatoes. Dept Note SIN/
255, Scot. Inst. Agric. Engng, Penicuik (unpub!.).
McRae, D.C. and Hutchison, P.S. (1988) Entwicklung eines hohlen siebstabes fur
kartoffelroder (Development of a hollow web rod for potato harvesters). Der
Kartoffelbau, 39 JG (7), 248-5l.
McRae, D.C., Carruthers, J. and Porteous, R.L. (1976) The effect of drop height on
potato damage. Dept Note SIN/202, Scot. Inst. Agric. Engng, Penicuik (unpub!.).
McRae, D.e., Statham, O.J.H. and Fleming, J. (1985) Retractable lid attachment
for a box tippler. Agric. Engr, 40, 12~3.
McRae, D.e., Glasbey, e.A., Melrose, H. and Fleming, J. (1986) Size grading
methods and their relationship to the dimensions, mass and volume char-
acteristics of potato cultivars. Potato Res., 29, 477-86.
Malcolm, D.G. and De Garmo, E.P. (1953) Visual inspection of products for
surface characteristics in grading operations. USDA Marketing Research Rep.
45, Washington, DC.
Marchant, J.A., Oriyango, C.M. and Street, M.J. (1990) Computer vision for
potato inspection without singulation. Computer and Electronics in Agriculture,
4,235--44.
Meijer, E.N.C. and Frederiks, J. (1975) Development of a new machine for
planting pre-chitted potatoes. IMAG Res. Report 75-3, Wageningen,
Netherlands.
Muir, A.Y. (1990) Reducing damage in handling systems. Abstracts of 11th
Triennial Conference of Eur. Assoc. for Potato Res., Edinburgh, p. 504.
Muir, A.Y., Porteous, R.L. and Wastie, R.L. (1982) Experiments in the detection
of incipient diseases in potato tubers by optical methods. J. Agric. Engng Res. ,
27, 131-8.
Muir, A.Y., Ostby, P.B. and Zender, F.N. (1990) The measurement of the
resistance of tuber skin to scuffing. Dept Note 30, Scot. Centre Agric. Engng,
Penicuik.
Ophius, B.G., Hesen, J.e. and Kroesbergen, E. (1958) The influence of the
temperature during handling on the occurrence of blue discolouration inside
potato tubers. Eur. Potato J., 1,48-65.
Palmer, G.M. (1976) Effect of close graded seed on the yield of main crop potatoes.
Expl. Husb., 30, 18-23.
Pascal, J.A. and Robertson, T.P. (1977) Yield effects of regularly and irregularly
spaced potato tubers. Expl Husb., 32, 25-33.
Pascal, J.A., Langley, A., Sheppard, B.W. and Hamilton, A.J. (1983) Energy
requirements for the reduction of clod populations during potato land
preparation. Agric. Engr, 38, 2-6.
Philipsen, P.J.J., Schlepers, H. and Hak, P.S. (1974) Thermische behandeling van
aardappelloof IBVL - mededelingen 433.
Potato Marketing Board (1974) Report on a National Damage Survey, 1973,
Potato Marketing, Board, London.
Potato Marketing Board Sutton Bridge Experimental Station, Annual Reviews
(1972) pp. 29-32; (1973) pp. 26-32.
Rastovski, A. and van Es, A. (1981) Storage of Potatoes, Pudoc, Wageningen, pp.
159-62.
Robertson, I.M. (1970) Assessment of damage in potato tubers. Dept Note SIN/60,
Scot. Inst. Agric. Engng, Penicuik.
Sawyer, R.L. and Collin, G.H. (1960) Black spot of potatoes. Am. Potato J., 49,
243-54.
Statham, O.J.H. and Cunnington, A.e. (1990) Two stage harvesting - an assess-
ment of systems and their impact on lifting rate and tuber quality. Abstracts of
11th Triennial Conference of Eur. Ass. for Potato Research, Edinburgh, pp.
388-9.
Travis, K.Z. (1987) Use of a simple model to study factors affecting the
References 607
size distribution of tubers in potato crops. J. Agric. Sci., Camb., 109,
563-71.
van Es, A. and Hartmans, K.l. (1975) Some differences in susceptibility to internal
blackspot. Potato Res., 18, 338.
Witney, B.D. (1984) The investigation and promotion of stone/clod windrowing for
potato production systems. Research and Development in Agriculture, 1, 1···20.
Witney, B.D. (1988) Choosing and Using Farm Machines, Longman Scientific and
Technical, London.
Zegers, D. and Van den Berg, V. (1988) The effect of lighting quality and intensity
on the efficiency of seed potato inspection. Agric. Engr, 43(1), 5-11.
CHAPTER 14
The physics and physiology of

storage
W.C. Burton~ A. van Es and K.J.Hartmans
14.1 INTRODUCTION
14.1.1 General
Whether or not to store potatoes and how to store them are commercial
decisions which must depend upon the circumstances .of the individual
case. The technicalities of storage should never be considered in isolation.
The influence of many storage variables upon the quality of stored potatoes
and upon storage losses can be found in the following pages. On the basis of
this information technically optimal methods of storage might be suggested.
Three variables determine storage losses: the potato, storage condi-
tions and storage duration. It must be realized that storage l.osses cannot be
avoided even by optimal storage. Good storage can merely limit storage
losses in good product over relatively long periods of storage. Bad storage
results in high storage losses, even in an originally good product.
Storage losses are often specified as weight losses and losses in the
quality of potatoes, although the two cannot always be distinguished.
Storage losses are mainly caused by the following processes: respiration,
sprouting, evaporation of water from the tubers, spread of diseases,
changes in the chemical composition and physical properties of the tuber
and damage by extreme temperatures. These processes are influenced by
storage conditions. All the losses mentioned above depend on the storage
conditions and therefore can be limited by maintaining favourable
conditions in the store. However, the storability of potatoes is already
determined before the beginning of storage, by such factors as cultivar;
growing techniques; type of soil; weather conditions during growth;
diseases before harvesting; maturity of potatoes at the time of harvesting;
damage to tubers during lifting, transport and filling of the store.
There are four main outlets for stored potatoes: seed potatoes, the
Introduction 609
ordinary ware market, the processing industry and potatoes as raw
material for the production of starch (mainly in The Netherlands) or
alcohol. Choice of storage method must be conditioned by the require-
ments for each purpose, but for all uses wound healing is essential
immediately after harvest or, where relevant, after grading, or, in the case
of consignments of seed arriving from elsewhere, immediately on receipt.
14.1.2 Requirements
The requirements for each purpose are briefly described in the following
sections.
(a) Wound healing

For all uses wound healing is essential immediately after harvest, or, where
relevant, after grading. This involves a period of about 2 weeks at lOoC or
above (Section 14.3). It is occasionally possible to take a calculated risk in
not allowing wound healing, where it is known that storage is to be so short
that wound pathogen development is unlikely to be serious - in buffer
stocks for canning for example. There is, however, some risk.
(b) Seed
Points to be considered are: method of planting and the importance placed
upon being able to plant rapidly when conditions are right; size of handling
and storage units to suit the method of planting and other requirements;
importance placed upon avoiding sprout damage - rapid mechanical
planting could damage sprouts which had grown and perhaps delay
emergence, but slower planting could lead to later planting and even
longer delay; length of sprout which can be tolerated in the light of the
foregoing; importance or otherwise of apical dominance.
Exposure to light is not deleterious, nor advantageous except for its
effect in giving stronger sprouts; sweetening is not a disadvantage; and it is
desirable for the dormant period to be over - even if the sprouts are not
growing they should be on the verge of so doing. Uniformity of behaviour
is important, which means that storage conditions, in particular temperature,
should be uniform.
In accordance with the above it could be suggested that the minimum
storage temperature should be well above any danger of low tempera-
ture injury, say 2°C; that the long-term maximum should not be
such that undesirable growth occurred in the cultivar stored i.e. usually
SoC or below; and that the temperature range throughout any batch
of seed should not exceed 2°C at the most. There could be a short
period at, say lOOC to encourage growth if and when required (see Chapter
6).
610 The physics and physiology of storage
The main requirement for seed storage therefore is close temperature
control within the range 2-5°e after wound healing.
(c) Ware for domestic and catering consumption

Potatoes for consumption must be protected from light because of the
undesirable formation of glycoalkaloids induced by exposure to it (Section
14.5.4). Wilting and sprout growth should be avoided as far as economically
possible. Apart from these, storage impinges on domestic and catering
requirements by affecting the degree of internal bruising during grading. If
it is feasible, susceptible stocks should be allowed to rise to 15°e or so
before grading. Sugar content, affected by storage temperature (Section
14.5.3) is not critical provided that potatoes which have sweetened at a low
storage temperature are allowed to de-sweeten, at woe or above, for
about 2 weeks during distribution and in the home. Senescent sweetening
is not normally a serious problem, although it can become objectionable in
very late stored potatoes.
From the above we could conclude that a temperature of 5°e or so, after
wound healing, could be optimal. This would avoid serious sprout growth
and senescent sweetening (Section 14.5.3). Low temperature sweetening
could be acceptably reversed during distribution. The temperature range
would need to be kept within narrow limits if we depended upon
temperature to avoid sprout growth - say 4-6 0 c. An alternative method
would be to hold the temperature roughly in the range 5-lOo e and use
chemical sprout suppressants for late storage - it is of course a waste of
money to use these on potatoes which are only to be stored for a period
during which sprouting could be prevented by a simple storage regime such
as night ventilation. Whatever method of storage is adopted the avoidance
of wilting is important - water loss should be kept, if possible, below 5%.
(d) Ware for processing

'Processing' usually means either crisp or chip manufacture or dehydra-
tion in the form of flakes or granules. The storage requirements for all
these are compatible and they can be considered together. A voidance of
internal bruising is important but is not really a storage problem except
for raising the temperature, if necessary, before grading. The main
requirement of the manufacturers is a very low and uniform content of
reducing sugars - preferably about 0.1 % of the fresh weight, with a
maximum of about 0.25% for crisp and 0.5% for chip or French fry
manufacture. For this, in the early part of the storage season, the higher
the temperature the better, up to about 20 0 e (Section 14.5.3). Long term
storage of potatoes in any quantity at a high temperature is hazardous,
however, and a temperature of about woe is the best commercial
compromise. Because of the requirement for uniformity a range of ± l°e is
Introduction 611
desirable. A manufacturer usually requires supplies which last until freshly
harvested potatoes are available at an economic price. This entails late
storage, and late storage at lOoC results in a degree of senescent sweeten-
ing which is unacceptable to the processor. It would be avoided by storage
at 5°C, but this gives low temperature sweetening in most cultivars, which,
although reversible, is incompletely so, and not uniform (Section 14.5.3).
A compromise solution is to store the potatoes required for late use at
about 5-7°C for chip manufacture and at about 7-lOoC for crisps manu-
facture, after wound healing.
Senescent sweetening is sufficiently delayed but there is some low
temperature sweetening (see Fig. 14.26). This is reversed by brief storage,
batch by batch at a high temperature, up to 20°C, and the residual rather
higher sugar and lack of uniformity accepted as the lesser of two evils.
Storage at the temperatures mentioned would result in vigorous sprout
growth, and chemical suppression is essential except for stocks stored
only briefly. Wilted potatoes present processing difficulties, and water loss
should, if possible, be kept below 5%.
(e) Canning potatoes

Canned potatoes are typically small immature tubers. These normally have
a high content of sugar and the sweetening which results from storage at
about 4-5°C, after wound healing, is apparently acceptable. This avoids
sprout growth. Immature potatoes are very susceptible to water loss
(Section 14.2.2) and have a high initial rate of respiration (Section 14.2.3)
and consequent heat production. Avoidance of drying conditions in store is
thus essential, and storage in bulk to depths greater than about 2 m is
inadvisable.
(f) Industrial potatoes

The main industrial use of potatoes is for starch manufacture. Sweetening
represents a loss of starch - the extra sugar accumulation at a temperature
of 4°C, above that at lOoC, illustrated in Fig. 14.26, represents a loss of
about 1.3% by weight of starch, probably about 8-9% of that originally
present. Sweetening must therefore be avoided. Unventilated storage in
bulk, although with ducts under the stack for the introduction of a sprout
inhibitor if desired, at a temperature within the approximate range 8-12°C,
is probably the best method. Sprouting represents a loss of starch, but 1%
by weight of sprouts only represents a loss of about 0.1 % of starch, which
together with the extra respiration accompanying sprouting may be about
1% of the original content. The cost of sprout suppression must thus be set
against the realizable advantage. Wilting is not important.
14.2 PHYSICS
14.2.1 Temperature
(a) Effects of temperature

Some important physiological mechanisms that occur in the potato tuber
during storage are briefly mentioned here.
Respiration
The short-term effect of temperature upon the rate of respiration is well
illustrated by the results of Miiller-Thurgau (1882), which showed a two-
fold increase for a lOoC rise in temperature over the range 0-20°C. This
short-term effect is not maintained during storage at different tempera-
tures and after the re-establishment of all the dynamic equilibria between
temperature-sensitive reactions which may influence the overt symptoms
of respiration, the QlO may be very different from 2. Burton (1974) found a
mid-storage temperature coefficient, 20/lOoC, of about 1.2-1.3, but
respiration at 2°C (sweetened tubers) could be higher than at 20°e. Later,
increased respiration accompanying sprouting - most vigorous at 20°C and
non-existent at 2°C - changed this pattern. For a more detailed description
see Section 14.2.3.
Cooling effect of evaporation

At a storage temperature of SoC, the latent heat of vaporization of water is
about 2.49 kJ g-l. The specific heat of potatoes at the same temperature is
about 3.6 Joel g-l. The evaporation of 1 mg of water will thus remove an
amount of heat sufficient to lower the temperature of 1 g of potato tissue by
O.69°C.
Water loss
For a more extended discussion on the water loss of potatoes see Section
14.2.2. Briefly, temperature has a considerable influence on water vapour
pressure deficit of the surrounding air in potato stores. Hence temperature
will influence the loss of water from the tubers.
Dormancy and sprout growth

There is a significant relationship between temperature, and dormancy and
sprout growth. The effects will be described more in detail in Section
14.4.1.
Wound healing
The higher the temperature the more rapid wound healing takes place. A
more detailed description is given in Section 14.3.4.
Physics 613
Metabolic changes
Of the numerous metabolic reactions the formation of the reducing
carbohydrates glucose and fructose as well as the sweetening carbohydrate
sucrose will get special attention with regard to temperature changes
during storage. See Section 14.5.3.
(b) Temperature of a single tuber

In a tuber which is in thermal equilibrium with its surroundings, metabolic
heat production raises the temperature slightly above the environment -
the specific heat of the tissue is about 3.6 Joel g-l (Mann, reported in
Burton, 1966) and a typical heat production of, say, 50 mJ g-l h- l (Section
14.2.3) could thus raise the temperature 0.014°C h- l were it not transferred
to the surrounding air. This transfer involves a temperature gradient of
about 0.2°C from the centre to the periphery, and about O.l°C from just
below the skin to the surrounding air (Burton, 1969). These are the
approximate equilibrium values.
A tuber placed in an environment differing in temperature, equilibrates
at a rate depending on the temperature difference, the thermal conductivity
of the tissue, the surface area and the rate of air movement. Re-
equilibration of ware tubers exposed to an 8°C temperature differential
takes about 12 h with free air circulation. (Burton, 1969). Farrimond
(1973) found a heat transfer of about 0.8 W from 200 g tubers in an air
current of 1 m S-l with a mean surface temperature differential of about
3°C. Theoretically, the rate of heat transfer from a spherical body to an air
stream is approximately proportional to 01. 6 , where V is the linear velocity
of the stream, but, as Mann (1963) pointed out, heat transfer from the
centre of a potato is limited by the thermal conductivity of the tissue. He
found the overall heat transfer from potatoes of about 7 cm diameter,
weighting about 215 g to be proportional to 01. 31 . The index may vary with
the size, shape and moisture content of the tuber, and in practice
proportionality to the cube root of the velocity is a convenient approximation.
The foregoing has been concerned with the response of a single tuber to
the environment immediately surrounding it. Before we can make use of
this information for anything other than single tubers, we need to have a
fairly precise idea of the environment in a stored bulk. For a single tuber
the whole of the metabolic heat production is then being transferred to the
surrounding air as fast as it is being produced. The rate of heat production
depends on the rate of respiration, but for a mature 200 g tuber at normal
storage temperatures, it could be expected to be within the range 8-16 J h- l
(Section 14.2.3). Transfer of this heat to the surrounding air raises the
temperature of the latter - the specific heat of air is about 1.0 J g-l °e l
and 10 J would thus raise the temperature of 1 g (i.e. about 800 cm 3) of
air by lO°C, or of 1 I of air by 8°C. In the case of a single tuber exposed to
free air circulation in a room, the ratio of the weight of air to tuber is so
great that the effect of the transfer of metabolic heat upon air temperature
is negligible.
(c) Temperature in a stack of potatoes

One tonne of fairly clean potatoes heaped together will, however, have
only about 0.5 m 3 of air in the spaces between them. The heat production
of the tonne of potatoes (40-80 kJ 11-1) would be sufficient to raise the
temperature of this amount of air by about 65-130°C in 1 h. The air is not
trapped, however, but can escape from the heap. Within the temperature
range encountered in a potato stack, every 1°C rise in air temperature leads
to a drop of about 0.36% in its density (there is also a small additional drop
due to humidification of the air in a stack), and this warmed lighter air will
rise out of the heap at a rate which would be proportional to the drop in
density (or the rise in temperature) were it not that the resistance to air
flow through heaped potatoes increases in proportion to V 1 .8 (Burton et al.,
1955). The net result is that the air rises out of the heap at a rate
proportional to (A1)°·55 where AT is the temperature difference activating
the movement (Burton et al., 1955). The more the temperature rises,
therefore, the faster the air rises out of the heap, cooler air being drawn in
from the surroundings to replace it and being in turn warmed and replaced
by cooler air. A convective movement of air into and out of the heap has
thus been activated. Every m 3 of air leaving the heap at a temperature 1°C
above that at which it entered would remove about 1.25 kJ.
Burton et al. (1955) derived a relationship between the approach velocity
of convective air movement and the temperature difference activating it.
They also found that in a stack of potatoes in equilibrium, the maximum
temperature was about 1.5 times the average temperature. The precise
temperature difference activating convection and over what distance may
be open to question.
Burton et al. (1955) used the average stack temperature, which gives a
value 20% less than that calculated using the maximum temperature. The
average will be used here, with the above caveat. On this basis, with a stack
of fairly clean potatoes 3 m high, comparable with commercial practice, it
can be calculated from the relationship mentioned above that 1°C
temperature difference would activate convection at a rate of 4.6 m 3 C1 h- 1 ,
capable of removing 5.75 kJ h- 1 plus about 12.6 kJ h- 1 by evaporation. A
rise of the temperature difference to 3°C would increase the convection to
8 m 3 t- 1 h- 1 (i.e. IX. (A 1)0.55), capable of removing 30 kJ h- 1 plus about 22 kJ
h- 1 by evaporation, about equivalent in total to the normal average heat
production.
In general terms, the effectiveness of convection in removing heat
increases in proportion to (A1)1.55 so far as direct cooling is concerned
(Burton et al., 1955), but in proportion to (A1)°·55 so far as evaporative
cooling is concerned. Respiration, and heat production, may also increase
Physics 615
with increasing temperature, but at normal storage temperatures, to a
much smaller extent than does the effectiveness of convection; and, except
under exceptional circumstances, which will be mentioned below, thermo-
dynamic equilibrium is always established. Burton et at. (1955) found it to
be reached after a period of about 2 weeks without marked fluctuation in
ambient temperature. In stacks in such a state the temperature is of course
not uniform - if the average temperature were 3.5°C above ambient there
would be gradation from little above ambient to perhaps 5°C above (see
'Dimensions of the stored mass' below), and the rate of convection, and its
effectiveness, would show local variation and local transfer of heat. Burton
et at. (1955), in bins of potatoes 1.9-3.7 m high, found average tempera-
tures to differ from ambient by 1.8 to 3.0°C and maximum temperatures by
2.8 to 5.8°C.
They derived a general equation for the calculation of average
temperatures in stacks in thermodynamic equilibrium with the ambient air,
based on continuous records of temperature measurements, at up to 73
points in anyone stack, on 12 stacks ranging from about 10 to 100 t.
Modified to SI units and with a correction to allow for evaporative heat loss
(Hylmo et at., 1975a), disregarded in the original equation, it is:
K Qm - C - E 1.8
(Tp - T a )2.8 ( ) (14.1)
= 1.17 X 10-2 A
where Tp = average temperature of the potatoes (OC)

Ta = average temperature of the ambient air CC)
Q = metabolic heat production (kJ t- 1 h- 1)
m = weight of potatoes (t)
C = heat loss by conduction (kJ)
E = heat loss by evaporation (kJ)
A = area of top surface of stack (m 2 ).
The value for K varies with the sample of potatoes. Burton et at. derived
a value of 7.7 x 10-5 for a commercial sample with little adherent earth.
The data of Ophuis (1957) give a value of 5.3 x 10-5 for cleaned potatoes.
With 20% adherent earth, Wager et at. (1952) found 2 X 10-4 and the same
potatoes, sprouted to the extent of 5% by weight gave 8 X 10-4 . There is
thus a 15-fold difference between extremes, but for good un sprouted
commercial samples a value of 8 x 10-5 may be a reasonable estimate. Q,
as we see from Section 14.2.3, is likely to be in the range 40--80 kJ r 1 h- 1 . C,
in the case of large commercial stacks, is small in comparison with
convective loss and may be disregarded.
Measurements of weight loss in practice suggest that water evaporates
during the season at an average rate of 10 g t- 1 h- 1 . This corresponds with a
heat loss of about 25 kJ r 1 h- 1 : less than half the normal total loss (Hylmo et
at., 1975a). Dry ambient air would give excessive evaporation and
corresponding cooling. On average in England (Q m - C - E) = 35 kJ rl
h- 1 could probably be used.
(d) Ambient temperature

The thermodynamic equilibrium values discussed above are all related to
ambient temperature. The temperature of a stack of potatoes, if it is out of
equilibrium with the surrounding air, will, as outlined above, always move
in the direction of that equilibrium. There will be no tendency to equate
with the actual ambient air temperature, but with a higher temperature
according to the height of the stack. Thus the average temperature of a
stack 3 m high will follow the general trend of a temperature about 3.5°C
higher than the average daily ambient, although with an amplitude of
fluctuation normally less than 1°C per day. The thermal inertia of a stack of
potatoes is too great for it to follow rapid ambient fluctuations. Similarly
the maximum temperature of the stack would follow the general trend of a
temperature about 5.5°C higher than average ambient, again with damped
15
-.. .Ie
nc1l ~.
•
.-
• • ex:! 0'
10 • . .wPl_
..~
~O.~ ; oio
• •
-
0
~
~ °eo ·.0 cPO

:::J • • • • • -0 p
~~ ·..,on ~
~
..... 5
••••
-
. ~~ .~ • "'tal
Q)
c.. • 1Sl~
E
•
• • ••• •• •
.:
Q)
f-
0 ••
•
•
-5
20 40 60 80 100
Days stored
Figure 14.1 Daily average temperatures, over a period of 80 days, of potatoes (cv.
Majestic) stored in an unventilated bin to a height of 2.5 m related to a temperature
3°C higher than daily average stare air temperature (Data from Wager et at., 1952).
0: potato temperature; e: store air temperature plus 3°C.
Note: (1) Approximate thermodynamic equilibrium during periods of minor
fluctuation in store temperature (days 20-43).
(2) Damped response to major consistent trends in store temperature, tending to
re-establishment of thermodynamic equilibrium with temperature 3°C above
ambient (days 44--100).
Physics 617
fluctuation. It was stated above that the minimum temperature was usually
that of the potatoes in the most exposed top corner. This is true of the state
of thermodynamic equilibrium; and true also of the long-term average.
These potatoes however follow a temperature about 1°e above average
daily ambient, with only such damping of fluctuation as results from their
limited heat conductivity (Section 14.2.1). If there was a sudden marked
rise in ambient temperature they could, transiently, be the warmest
potatoes in the stack - see, for example the temperatures given by Wager et
al. (1952).
The response to ambient temperature, described above, is illustrated in
Fig. 14.1, in which the average temperature of potatoes stpred to a depth
of 2.5 m is shown in relation to a temperature 3°e above daily average
ambient, over a period of 80 days.
(e) Dimensions of the stored mass

Effect of lateral dimensions of stack on temperature
Burton et al. (1955) found the temperature in an unventilated mass of
potatoes, averaged over the whole height, to increase from the periphery
inwards and to reach a maximum value at a distance from the sides of the
mass which was about equal to its height (Fig. 14.2). Thereafter there was
no further increase. The cooler periphery is due to cool air being drawn in
by convection at the sides of the stack, where resistance to air flow is least,
.----
(j)
.---- -----------
Q)
~
_ 4
o
a.
.!..
.ctj
~7mhigh
[)3
o
/" 0_ 0-------------
'-'
Q)
g 2
"/.,...-----1.9 m high
....
Q)
Q)
:::::
'6 o
1
....
Q)
:::J
10
....
~ 0 L -________~__________~__________~~
E
Q)
0 1 2 3
I- Distance from bin wall (m)
Figure 14.2 Temperatures of potatoes (stored in bulk in bins) at different
distances from the bin wall. Each temperature is an average of those at several
heights from the floor (0.23, 1.0, 1.75,2.5 and 3.5 m in the 3.7 m bin; at the first
three in the 1.9 m bin). (From Burton et at., 1955.)
the effect being noticeable over the distance given and being most marked
at the corners. Here the temperature of the top potatoes differs little
« 1°C) from ambient. Over the rest of the stack - and this would
represent the greater part of a large commercial stack - it would appear
that local convective currents are drawn in over the top surface. It follows
from the foregoing that the lateral dimensions of potato stacks of com-
mercial size have little effect upon the temperature of the potatoes,
although in crates and small bins peripheral cooling would have an
appreciable effect. This expected behaviour has been confirmed in com-
mercial stacks of 500 tonnes and upwards.
Temperature in relation to the height of the stack

In contrast to the effect of lateral dimensions the height of a stack can be
expected to influence its temperature markedly. Burton (1978a) carried
out some calculations on the effect of increased height of storage.
Observations in practice confirm this calculation of the magnitude of the
effect of increased height. Burton et al. (1955) concluded, on the basis of
their results over a number of heights from 1.8 to 3.7 m, that the
temperature difference !1T with ambient temperature was proportional to
ho. 64 , or approximately h 2!3, h being the height of the stack, and that in the
middle of the storage season !1T approximated to 1.8 h2J3 giving an average
temperature 2.8°C above ambient for a stack 2 m high, and 4.4°C above for
4 m high - temperatures in about the same ratio (1.6) as was calculated
theoretically above. The actual temperatures are of course very dependent
upon the heat production of the potatoes (ex: QO.64 where Q is the heat
production) and upon other characteristics of the stack, and individual
results of Burton et al. (1955) showed some variation, e.g. 1.8°C above
ambient for stacks 1.8 m high, and in one case only 2.2°C above ambient
for a stack 3.7 m high. In stacks of the heights they studied, Burton et al.
concluded that the maximum temperature approximated to 1.5 times
the average temperature. Taking the relationship between height and
temperature above, this suggests a maximum of 2.7 h 2/3 oC. These con-
clusions were based on continuous records of temperature measurements,
extending over a few years, at up to 73 points in anyone stack, on twelve
stacks ranging from about 10 to 100 tonnes. Table 14.1 gives the maximum
temperatures reached at different levels in two unventilated stacks of
potatoes. The positions of these maxima illustrate the fact that with
convective cooling the maximum temperature is not reached at the top of
the stack, where ambient air is being drawn in by convection, but usually
about 0.5-1 m below the top surface.
Subdivision of the mass

A small unit, such as a 0.5-1 t crate of potatoes, behaves as a miniature
unventilated bulk. The dimensions are such that the maximum tempera-
ture, controlled by convection approximates to 1°C above ambient, and
Physics 619
Table 14.1 Approximate maximum temperatures reached at different
levels in unventilated bins of fairly clean, mature unsprouted potatoes
(based on Burton et aI., 1955)
Total height Level at which temperature Approximate maximum
of potatoes was taken (m above floor) temperature (OC above
(m) average ambient)
1.85 0.25 1.5
1.0 3
1.7 2.8
3.7 0.25 2.5
1.0 4
1.75 5.5
2.5 5.7
3.55 5
the mlmmum temperature to ambient. We thus have fairly uniform

temperature conditions in each crate. It remains to ensure that the crates
do not differ from each other, which means that the air throughout the
store must be at about the same temperature. This is done by circulating
the store air. The circulating air, moving over the potatoes is warmed
by the metabolic heat; 30 m3 passing over a tonne, producing 35 kJ net
after evaporative cooling (Section 4.2.2), would be raised nearly loe in
temperature, which means that circulation at 30 m 3 t- l h- l would result in a
loe gradient in the store air. Allowing also for the gradient in each crate,
circulation at this rate is sufficient to hold the potatoes within a range of
about ± loe of mean store temperature. It remains to control the latter at
the required temperature - ideally about 3-4°e for seed; about lOoe for
short-term storage for processing; about 7°C for long-term storage for
processing, and about 4-5°e for canning. Potatoes for such uses can best
justify the expense of crate storage, for which the crates represent a
sizeable capital investment. In the main potato growing areas of the UK
it is possible, on average, to obtain sufficient control for every purpose,
until about March, by an adequately insulated store and judicious use of
ventilation with outside air during the night, or whenever the temperature
is suitable, using the same fans as for recirculation, i.e. with a capacity
of 30 m3 t- l h-l. This suffices for seed storage. For storage during the
succeeding months for other purposes, in some cases until July or August,
refrigeration is necessary.
Ideally, refrigeration would also be desirable for seed storage in years
when the winter was very warm, but the economic justification for it would
have to be based on the frequency of such years and the monetary loss
from not having such a facility.
Inclusion of earth and sprouting
The inclusion of earth may be expected to have a great influence. The
effect of earth included in the stack is two-fold. A coating of earth on the
tubers may influence the rate of transfer of heat from tubers to air, as
described earlier; and earth in the spaces between the tubers will increase
the resistance of the stack to air flow. Calculations from the data of Ophuis
(1957) for clean potatoes give for K, the coefficient of resistance to airflow
a value of 5.3 x 10-5 . Misener (1986) determined the pressure drop per unit
depth of product P for increasing amounts of adherent earth (Fig. 14.3).
100. 6%
4%
2%
E
-
ell
a..
-
..c
c-
Ol
CLEAN
-
1:l
'0 10 .0
'c
:::J
a.
0....
1:l
....
Ol
:::J
en
en
a:
Ol
1.0
0.5
0.03 0.10 1.00
Air flow (rrr m-2 5- 1 )
Figure 14.3 Airflow versus static pressure drop for potatoes with various soil
content. (From Misener, 1986.)
Burton et ai. assessed the effect of increased resistance to air flow from
Equation (14.1). As we have seen above, inclusion of excessive earth has
been observed in one case to increase the value of K in this equation by a
factor of about 3, which means that (Tp - T af· 8 would have been
increased three-fold and hence (Tp - Ta) by a factor of about 1.5. Potatoes
stored to a depth of 3 m would thus reach thermodynamic equilibrium at an
average temperature 5-5.5°C above ambient instead of 3.5°C above. The
magnitude of the effect obviously depends upon the amount and nature of
the earth included, and the consequent influence upon K, which will vary
from case to case. The above factor of 1.5 is quoted merely to give an idea
of the likely upper limit of the effect. Besides the effect of earth, sprouting,
if this occurs late in the storage season also reduces the air spaces available
Physics 621
for convective air movement and has similar effects on resistance of the
stack to air flow.
(f) Heat production

For every g of carbohydrate consumed in normal respiration about 16 kJ of
energy are released. In part this is transferred into energy-rich phosphate
molecules and utilized in metabolism; in part it is lost as heat. There have
not been many measurements of the proportion so lost. Green et al. (1941)
suggested it was as much as about 90%. If we accept this for tubers in
which no marked change is occurring, then for every g of CO 2 produced in
respiration about 10 kJ of energy are released in the form of heat. The
amount could differ during periods of rapid change but evidence for this is
lacking.
Heat balance in a potato store

According to Rastovski (1987) the heat balance in a potato store involves
the following factors. In a store filled with potatoes, a certain amount of
heat is continuously produced and heat evacuation also takes place. The
temperature at which the potatoes are placed in the store is usually higher
than the required storage temperature, so that the potatoes first have to be
cooled to that temperature. More heat than is being produced in the store
has to be evacuated during the cooling period. Once the storage tempera-
ture is reached, heat evacuation must be equal to the heat production in
the store if the required storage temperature is to be maintained. Fig. 14.4
shows the heat supply or heat production and heat evacuation in a store.
The heat supply thus stems from the heat sources described in the
/-_0----.
following sections.
Figure 14.4 Heat sources in the store. (From Rastovski, 1987.)

Heat content of the potatoes The amount of heat to be evacuated to cool a
given quantity of potatoes from the initial temperature to the storage
temperature can be calculated as follows:
Quantity of heat to be removed per day:
3600 m (T j - Ts)
ql = (J day-I)
n
in which:
n = time in days in which the potatoes are to be cooled from Tj to Ts
Tj = temperature of potatoes brought in the store (0C)
Ts = storage temperature COC)
m = weight of potatoes (kg)
Respiration of potatoes The heat of respiration of the potatoes depends

on the potato temperature and the factors mentioned in Section 14.2.3. In
healthy potatoes some time after harvesting, this heat may range from 0.01
J kg- l S-l at 5°C to 0.05 J kg- l S-l at 30°C (Burton, 1966). Three to six times
as much heat may, however, be developed in immature and damaged
potatoes. The heat produced by respiration is calculated as follows:
q2 = 86400 m qr (J day-I)
in which:
m = weight of potatoes (kg)
qr = heat of respiration (J kg- l S-l)
Heat flow through walls, roof and floor The heat flow through the walls,
roof and floor of the store arises owing to the difference between the
ambient and storage temperatures. If the former is higher than the latter,
there will be a heat flow from the outside to the inside. If the outside
temperature is lower, heat will flow in the opposite direction - i.e. heat will
be removed from the store. The amount of heat exchanged depends on the
temperature difference between the outside and the storage temperatures,
the area of walls, roof and floor and on their resistance to the heat flow.
The rate of heat flow through the store is:
fl.T
q3 = A -3600 x 24 (J day-I)
R
where:
A = surface area of wall, roof or floor (m 2 )
fl.T = Ts - TooC - difference between ambient temperatures on inner and
outer side of walls, roof and floor
R = (km2 W- l ) resistance to heat transfer of wall, roof or floor structure
respectively
Physics 623
Heat production by fans A fan produces heat while operating. The total
power of the electric motor which drives the fan is converted into heat.
Hence the heat supplied by the fan is:
q4 (J day-1)
Heat supplied by air renewal Fresh air is required to provide the potatoes
with the necessary oxygen and to remove the CO 2 produced in the store.
Air renewal supplies heat to the store if the fresh (external) air is warmer
than that in the store, but removes heat from the store if the outside air is
colder. A volume of air equal to the fresh air supplied to the store is also
removed from it. The temperature and relative humidity of this discharg-
ing air are those prevailing in the store. Hence V m3 day-I of fresh air with a
heat content io is supplied to the store while the same amount of air but
with a heat content is is removed from it. The amount of heat brought into
the store is accordingly:
qs = V!1i e
in which:
V = amount of fresh air (m 3 day-1)
f..i = difference in enthalpy between storage air is and outside air io (J kg-I)
e = air density of fresh air (kg m- 3 )
The enthalpy and air density can be obtained from the Mollier diagram
at known air temperature and relative humidities.
Heat produced by lighting, machinery, etc. The heat produced in a store

by leaks in the structure, lighting, machinery, people and open doors is not
easy to determine. This amount of heat is usually added as a percentage to
the sum of the components of the total heat supplied in accordance with
the items mentioned previously. This is usually taken as being 5% for large
stores and 10% for small ones; hence.:
q6 = 0.05 to 0.1
Total heat The total heat supply or heat production in the store is then:
This amount of heat has to be evacuate9 from the store if the required
storage conditions are to be reached and maintained. This heat is therefore
equivalent to the cooling load in the store. For many practical reasons the
installed cooling capacity must have higher (20%) capacity:
q = 1.2 X qtot (J day-I)
Overheating
We have discussed above the establishment of thermodynamic equilibrium.
With normal potatoes in stacks of normal height and under normal
ambient conditions this is established with no very great temperature rise.
On the other hand, we have seen that, with considerably sprouted potatoes
the rise necessary to activate sufficient convection is quite considerable. As
the temperature of the potatoes rises in the move towards the establish-
ment of equilibrium it would be possible in such potatoes to reach a
situation in which the extra heat generated as a result of a further rise
in temperature, with increased respiration, was greater than could be
removed by the increase in convection brought about by that rise in
temperature. The production of heat would thus outstrip its removal and
overheating would occur. This is imminent (Burton elal., 1955) when:
(
Tp - Ta + dT 1.55 Q + dQ
(14.2)
Tp - Ta Q
where Tp, Ta and Q are as defined for equation (14.1) and:
dT = a small rise in temperature, say O.S-l°C
dQ = increase in production of metabolic heat (kJ C1 hoi) caused by an
increase of temperature d T
Table 14.2 Conditions under which uncontrollable overheating could occur in

unventilated stacks of potatoes (approximate: based on Burton et aI., 1955)
Mean potato Mean ambient
temperature (DC) at temperature ceC)
at
which uncontrollable which foregoing
overheating could dangerous potato
start temperature could be
reached
Height of storage (m)
1.85 3.7 1.85 3.7

Characteristics of potatoes
(a) Freshly harvested
Immature, fairly clean 27 22 13 6
Immature, much earth 23 18 8 o
Mature, fairly clean 30 27 19 13
Mature, much earth 27 23 14 8
(b) Stored, unsprouted
Fairly clean 36 30 24 19
Much earth 30 27 20 14
(c) Stored, considerably sprouted
Fairly clean 27 22 13 6
Much earth 23 18 8 o
Physics 625
It is possible to derive the conditions under which overheating could
occur from Table 14.2.
(g) Temperature control by ventilation response of stored potatoes

Instead of subdividing the crop into crates and circulating the store air to
reduce temperature scatter, the same result can be achieved by storing the
potatoes in bulk and continuously ventilating them by means of ducts in or
on the floor. Continuous ventilation at 30 m3 t- 1 h- 1 would give a difference
of about 1°C between the minimum and maximum temperature in the
stack, assuming a net heat production, after evaporative cooling, of 35 kJ t- 1
h- 1 • Ventilation ensures a reasonably uniform temperature but does not
control the level of that temperature, and this must be done by controlling
the temperature of the ventilating air by recirculation, the air intake being
above the stack, coupled with admixture with outside air of suitable
temperature, or with refrigeration.
Two rates of ventilation have been mentioned above. A rate of 30 m3 t- 1
h- 1 was suggested for air circulation, and also for ventilation with suitable
outside air, in cases such as crate stores and bulk stores with recirculation
where air movement is virtually continuous and there is no build-up of
temperature among the potatoes. A rate of 70 m3 t- 1 h- 1 was mentioned in
connection with the intermittent ventilation of stacks through which air
was not continuously circulated. These stacks of potatoes, for domestic or
catering use, may not usually justify expensive temperature control, as by
refrigeration. Warm spells of weather can lead to undesirably high
temperatures. As we have seen earlier in this section an ambient average of
lOoC would result in an average temperature of 13-14°C, and a maximum
of 15-16°C, in a 3 m high stack. It is therefore necessary to take full
advantage of cooler spells of weather to cool the stack by ventilation.
For this purpose the faster the rate of ventilation the better, up to the
point where frictional heating of the ventilating air, and also increased
power requirements, render further increase in rate either undesirable or
uneconomic. The fact that cooling is proportional only to about the cube
root of the rate of ventilation (p. 613) is relevant here. Recommended
rates vary from 70 (Mann, 1963) to 160 m3 t- 1 h- 1 (Ophuis, 1957). These
higher rates of ventilation would also be desirable for crate stores and bulk
stores with recirculation if it were attempted to store potatoes late without
provision for refrigeration. By ventilating whenever the air was cooler than
the potatoes, but above freezing point, and using 160 m3 c 1 h- 1 , Ophuis
calculated that it was possible on average in The Netherlands, to hold
potatoes at mean temperatures of about lOoC in September, 7°C in
October, 4°C from November to April, 8°C in May, 11°C in June and 13°C
in July and August. In south-east England., Burton and Mann (1954)
calculated that potatoes could on the average be held, by a similar rate of
ventilation, at 5-6°C until the end of April and perhaps at 4°C until the end
of March. In Edinburgh in the 1968/9 season, Nash and Lennard (1973)
using a rate of about 100 m 3 t- 1 h- 1 , held the maximum temperature of 30-
40 t lots within a range of about 4-6°C until the end of April. Thereafter
ventilation when the air was cool became increasingly ineffective, and
towards the end of May the maximum temperature in the stacks had risen
to about 13°C. The cooling achieved by intermitent ventilation has been
considered theoretically by Ophuis (1957), Burton (1966) and, in more
detail, with experimental confirmation, by Hylmo et al. (1957b).
(h) Sprout growth

The effect Of sprout growth is also two-fold. It is accompanied by increased
respiration (Section 14.2.3) and consequent heat output; and the sprouts
growing in the spaces between the tubers increase the resistance of the
stack to air flow. Again we can make use of equation (14.1). Sprout growth
to an average extent greater than 5% by weight is rarely encountered in
practice. Tubers sprouted to this considerable extent could be respiring at
four to five times the rate of unsprouted tubers, which means that Q m in
the equation could be five times greater and Q m - C - E possibly as much
as eight times greater; although E is potentially much greater in sprouted
than in un sprouted potatoes it is, in unventilated storage, limited by the
water-holding capacity of the convective air, and thus by the rate of
convection, and could be perhaps doubled in the case under consideration.
(T~ - Taf8 could thus be increased by a factor of at least 51.8 and possibly
81. and hence Tp - Ta by a factor of 2.8--4.2. In addition there is the effect
upon resistance to flow, i.e. upon K, which could be increased by a factor
of 4, as we have seen above (admittedly in the presence of considerable
earth). This would increase Tp - Ta by a further factor of 1.6. The overall
effect could well be a four-fold increase in Tp - Ta. The average
temperature of a 3 m high stack, instead of being 3-5°C above ambient
would tend to be 17-18°C above. Again, as in the case of inclusion of earth,
this figure is not precise, but it gives an idea of the potentially serious effect
of considerable sprout growth in unventilated stacks.
(i) Refrigeration
The previous sections have shown that for most purposes the temperature
of stored potatoes can be controlled acceptably until March or April by
using intermittent ventilation with cool air, coupled, where uniformity of
temperature is important, with recirculation of the store air. It has also
been shown that this method cannot be relied upon later in the storage
season, and the temperature can then only be controlled by refrigeration.
This can be justified for uses for which temperature control is essential, and
it is usually applied by passing the circulating air over cooling surfaces,
such that the inlet temperature of the air is about 0.5°C below the desired
Physics 627
mean potato temperature. Such a method of temperature control is
efficient. Burton and Mann (1954), for example, stored SO t of potatoes at a
mean temperature of 5.2°C with a difference, between the minimum and
maximum temperatures in the mass, of about O.S°c.
There is one essential proviso, however, relating to the size of cooling
surfaces. Ventilating air removes water from the potatoes at a rate
proportional to the water vapour pressure deficit (WVPD) of the air
(Section 14.2.2). In the case of refrigeration the air passing over the
cooling surface deposits moisture or ice on it, if the surface temperature is
below the dew-point of the air, until the moisture content of the air falls to
that equivalent to saturation at the cooler temperature. The lower this is
the less the moisture retained in the air leaving the coolers, consequently
the higher its WVPD, and the greater the evaporation from the potatoes;
the resultant increased moisture content of the air leaving the potatoes
being again reduced on its next passage over the cooling surface. Efficient
transfer of heat, and hence temperature control, can be achieved either by
a large cooling surface at a temperature not much below the desired air
temperature, or by a smaller surface operating at a lower temperature. The
latter, however, could cause serious wilting of the potatoes. Burton and
Mann (1954) obtained satisfactory results using finned pipe coolers capable
of removing approximately 26 kJ h- l (= 7.2 W), per °C temperature
differential, for every tonne of potatoes they served. The whole of the
normal heat production could thus be removed by a 2°C differential, even
disregarding evaporative cooling, the WVPD of the air entering the
potatoes being therefore only about 1 mbar in the absence of heat leakage
into the store - the latter would necessitate an increased differential.
14.2.2 Water loss
(a) Water transport and water vapour pressure deficit

Water loss exemplifies a move of a dynamic system towards equilibrium -
in this case between a tuber which is in equilibrium with an approximately
water saturated atmosphere, and a surrounding atmosphere which usually
is not saturated. The rate at which water will evaporate from a tuber in
response to the above tendency is directly proportional to the difference
between the equilibrium water vapour pressure of the tuber (i.e. of a
saturated atmosphere at tuber temperature) and the water vapour pressure
of the air in contact with it. If the tuber and the air are at the same
tempeTature this difference approximates closely to the water vapour
pressure deficit of the air. The WVPD of the ambient air varies. Burton
(1966) gave rough average values for eastern England ranging from a
minimum of O.S mbar in the winter months to a maximum of 2.7 mbar in
May and June, the average over the whole storage season, October to
May, being about 1.4 mbar.
If we use this value, with the realization that it is a local overall average,
and not a precise figure, then every m 3 of convective air will remove about
2.75 kJ by evaporation, averaged over the season. If the temperature of the
tuber is changed, then the water vapour pressure in air saturated at tuber
temperature is changed, to an extent which is shown in Table 14.3. If the
temperature of the air is changed, over the range encountered in potato
storage, the vapour pressure of the water in it, if the quantity remains
unchanged, is unaltered. For example, air saturated with water vapour at
lO°C contains 9.33 g of water vapour m- 3 , exerting a pressure of 12.16
mbar. If we raise the temperature to 20°C, at constant atmospheric
pressure, the cubic metre will expand to 1.035 m3 , still containing 9.33 g
water vapour - i.e. 9.01 g m- 3 , exerting a similar pressure (Table 14.3). By
contrast, the effective vapour pressure of a tuber raised from 10 to 20°C
will be increased from 12.16 to 23.09 mbar. The quantity of water does not
always remain unchanged, however, and a fall in air temperature can result
Table 14.3 The content and partial pressure of the water vapour in 1 m 3 of
saturated air at a total barometric pressure of a standard atmosphere (=
1013.25 mbar; 101.325 kPa) (from Burton, 1973)
Temperature Weight of Volume of Partial Grams of
("C) water vapour water vapour pressure of water vapour
(g) (I) water vapour per mbar
(mbar)
0 4.84 6.00 6.08 0.80
1 5.18 6.44 6.53 0.81
2 5.54 6.92 7.01 0.80
3 5.92 7.43 7.53 0.79
4 6.33 7.96 8.07 0.79
5 6.76 8.54 8.65 0.78
6 7.22 9.15 9.27 0.78
7 7.70 9.80 9.93 0.78
8 8.21 10.49 10.63 0.77
9 8.76 11.22 11.37 0.77
10 9.33 12.00 12.16 0.77
11 9.93 12.82 12.99 0.77
12 10.57 13.69 13.87 0.76
13 11.25 14.62 14.81 0.76
14 11.96 15.60 15.81 0.76
15 12.71 16.64 16.86 0.76
16 13.50 17.74 17.97 0.75
17 14.34 18.90 19.15 0.75
18 15.22 20.12 20.39 0.75
19 16.14 21.42 21.70 0.74
20 17.12 22.79 23.09 0.74
25 22.80 30.87 31.28 0.73
30 30.04 41.88 42.43 0.71
Physics 629
in a lower water vapour pressure if the temperature drops to below dew-
point. If we lowered the temperature of the m3 of air, saturated at 10°C, to
5°C, then its volume would contract to 0.98 m3 , which if it still held the
original 9.33 g of water vapour would contain 9.50 g m- 3 . Air at 5°C
however can only hold 6.76 g m- 3 , exerting a pressure of 8.65 mbar, and the
excess water therefore condenses out. The initial effects of any changes in
air or tuber temperature upon the rate of water loss can now be calculated.
These initial effects are transitory, and alter in predictable fashion as the
temperatures of the tuber, or of the air, or of both, change in the direction
of thermodynamic equilibrium between tuber and air; and also as, if the
volume of air is limited, the water evaporated into it, from the tuber,
increases its water vapour pressure (see below).
(b) Rate of evaporation from tuber

Evaporation - limitation by the periderm - effect of damaging
The periderm limits the rate of evaporation from the tuber to much below
that from a free water surface, exposed to the same WVPD. The dead cork
cells are not freely permeable to water which migrates slowly through them
to the surface where it evaporates in response to the WVPD. About 98%
of the water loss appears to occur in this way and only about 2% by direct
diffusion in the gas phase through the lenticels (Burton, 1973). Limitation
by the periderm results in rates of water loss, from mature undamaged
tubers exposed to free air circulation, of the order of 6-10 Ilg cm- 2 h- 1
mbar- 1 (Burton, 1973) whereas in the absence of the skin, Burton (1955)
found a rate prior to wound healing, of about 3.5 mg cm- 2 h- 1 mbar- 1 ,
several hundred times as great.
Damaging and bruising due to lifting and grading are significant causes
of high moisture loss. Burton and Hannan (1957) suggest that the resulting
moisture loss can easily become three to five times as great. Also the
degree of sprouting is important; moisture loss is much greater in potatoes
that have sprouted (Burton and Hannan, 1957) (Fig. 14.5). Burton (1955)
calculated that the epidermis of the sprouts is about 100 times as permeable
to water as the surface of the tuber. Sprout growth corresponding to 1%
increase in tuber area causes an increase of 100% in moisture loss.
Influence of ventilation
Evaporation from a tuber into the air raises the amount of water in the
latter and hence its water vapour pressure, thus reducing the WVPD. If the
amount of air concerned is small the proportionate effect of evaporation of
a given amount of water into it is great. Increasing the amount of air by
increased ventilation can considerably reduce this effect. Conversely, if the
amount of air is great, the effect is small and still further increasing the
amount of air can have little influence. This is illustrated in Table 14.4
using typical values for the characteristics of the tubers. Changing the rate
x10-2
1.0
o
:,.,
3:
~ 0.5
'"'"
.Q
:E
OJ
~ o
2 4 6 8 10 12
Sprout growth (% by weight)
Figure 14.5 Rate of evaporation from tubers of cv. Dr McIntosh in relation to

sprout growth. Temperature: lO o e; water vapour pressure deficit of store air: 2.7 x
102 Pa. (From Burton and Hannan, 1957.)
Table 14.4 Calculated rates of evaporation from potato tubers at different

rates of ventilation
Rate of
ventilation Evaporation from tuber
Mature (1) Immature (2)
I h- 1 mg h- 1 % of mg h- 1 % of
potential\ potential
1 0.58 58 0.77* 2
2 0.75 75 1.54* 4
4 0.87 87 3.08* 8
10 0.94 94 7.69 19
20 0.97 97 14.6 36
100 0.99 99 31.9 80
200 1.00 100 35.6 89
1000 1.00 100 39.0 98
Characteristics of air: 10°C; water holding capacity, 9.33 mg 1-1; saturated water vapour
pressure, 12.16 mbar; WVPD, 1 mbar (= a deficit of 0.77 mg water 1-1)
Characteristics of tuber: (1) Mature stored tuber; 10°C; surface area 100 cm 2 ; wt, 100
g; evaporative potential, lO!1g cm- 2 h- 1 mbar- 1 . (2) Immature tuber 1 day after harvest;
10°C; surface area, 100 cm 2 ; wt, 100 g; evaporative potential, 400 !1g cm- 2 h- 1 mbar- 1
* Loss limited by water-holding capacity of air.
Physics 631
of ventilation is not marked except at very low rates or except in the case of
tubers - for example immature tubers - which can lose water readily.
Hunter (1985) showed the relationship between the percentage of weight
loss per week and the air velocity for different relative humidities (Fig.
14.6). Villa and Bakker-Arkema (1974) developed a model for moisture
loss from potatoes during storage.
0.18
0.16 90% RH
"""OJ
~ 0.14
Q;
c.
(j)
.Q 0.12
~
c
~ 0.10
OJ
.~ _________9_5_%_R_H
__
c...
--------------------
97.8% RH
0.08 ~
99% RH
0.06
0.00 50 100 150 200 250 300 350 400
Air velocity (m h- 1 )
Figure 14.6 Weight loss vs air velocity and relative humidity. (From Hunter,
1985.)
(c) Evaporative capacity of convection within a stored bulk of potatoes

Upper limit to the evaporative capacity
The air which enters a potato stack by convection is usually not fully
saturated with water, and thus has a capacity to take up more, and a
consequent WVPD in response to which evaporation occurs from the
potatoes. As the air moves through the stack it is warmed and can thus take
up still more water (Table 14.3), but when the warm air rises out of the
stack it is cooled by admixture with incoming air, and by the potatoes in
equilibrium with this mixture. Ultimately the air is cooled to ambient
temperature and cannot then hold any more water vapour than the water-
holding capacity of air at ambient temperature. If in fact it has taken up
more than this in the warm state then the excess will condense out.
In a stack with no such covering as a layer of straw, this condensation
will occur on the top potatoes. If there is a sufficiently thick straw covering,
the top potatoes will be warmer and the air will have moved above them
before being cooled down to dewpoint, and condensation will occur in the
straw. In either case, the air which leaves the stack, plus its covering,
cannot on average be more than saturated at store air temperature. The
upper limit to the evaporative capacity of convection is thus provided by
the quantity of water which the limited volume of convective air can take
up. Any excess above this which condensed on the top potatoes or the
straw would humidify subsequent convective air entering through the top
of the stack and reduce subsequent evaporative loss. The long-term
average would be as above. If we take the normal rate of convection to be
about 10 m3 t- 1 h- 1 , then for every mbar WVPD in the ambient aIr the
upper limit of evaporative loss would be about 7.8 g t- 1 h- I (Table 14.3)
(about 0.13% week- 1 mbar- I).
Evaporation from an unventilated bulk of potatoes

We can compare the above estimate of the maximum possible loss from
unventilated potatoes with some results of Wager et al. (1952) relating to a
small bin (9.5 t) of Majestic potatoes. These were weighed in and out at the
beginning and end of storage. The amount of earth was excessive (about
18%). From the average temperature, 6.6°C, 3.8°C above average
ambient, and the measured resistance to air flow, we can calculate an
average rate of convection of about 7.7 m 3 t- I h-l, less than would be
expected with cleaner potatoes. The average temperature of the store air
was 2.8°C and its average WVPD 1.5 mbar. It thus held 4.66 g of water m- 3
and had a saturated capacity of 5.84 g m- 3 (Table 14.3). Hence it was
capable of removing 9.1 g of water t- 1 h- 1 (about 0.15% week-I). The
potatoes actually lost 3.8% by respiration and evaporation over a period of
23 weeks. Of this, about 0.4-0.8% might have been due to respiration
(Section 14.3), leaving an evaporative loss of 3-3.4%, an average of 0.13-
0.15% week-t, approximating to the suggested possible maximum. We can
thus probably regard the actual evaporation from unventilated potatoes as
being equal to the capacity of the convective air on entry i.e. for fairly
clean potatoes, about 0.13% per week per mbar ambient WVPD.
(d) Modification of the evaporative capacity according to requirements

and response of stored potatoes
Ventilation
Convection at a rate of 10 m- 3 t- I h- I is equivalent to the lowest rate of
ventilation given in Table 14.4, at which humidification of the air is
such that evaporation from a tuber is only about 60% of the potential
evaporation into air of any given inlet WVPD. Evaporation from typical
mature stored tubers, capable of losing 0.17% week- 1 mbar- 1 would thus be
restricted to about 0.1 % week- 1 mbar- 1 were it not that, as we have seen
(Section 14.2.1) the convective air is warmed as it passes through the stack,
its evaporative capacity being thus enhanced.
Physics 633
Replacement of convective air movement by continuous positive ventila-
tion at a higher rate, through ducts under the potatoes, affects both the
temperature and the humidification of the air. The humidification is less
because of the greater volume of air, and the consequent restriction on
evaporative loss is progressively removed, as illustrated in Table 14.4, as
the rate of ventilation is increased. Metabolic warming of the air is
decreased, again because of its greater volume. There is also cooling of the
air because of evaporation from the potatoes. In the case of convection,
metabolic warming exceeds evaporative cooling and there is net warming,
but in the case of positive ventilation it would be theoretically possible for
cooling to exceed warming. This has in fact been observed by Hylmo et al.
(1975a). With a heat production of 50 kJ t- 1 h- 1 and a potential for water
loss of 10 g t- 1 h- 1 mbar- 1 it can be calculated that cooling and warming are
in balance with an average WVPD in the stack ranging from about 2.1
mbar (ventilation rate 200 m 3 t- 1 h- 1 ) to 3.5 mbar (ventilation rate 10 m3 t- 1
h- 1). With increase of WVPD above the balance point, cooling progres-
sively exceeds warming; with decrease below the balance point, warming
progressively exceeds cooling. The resultant temperature changes affect
the operative WVPD in the stack, cooling leading to a decrease, over and
above that due to humidification; and warming leading to an increase
which can more than offset humidifcation.
Calculated values for the average WVPD which is operative in potato
stacks ventilated at different rates with air of stated WVPD are given in
Table 14.5. Rather different values would be obtained with potatoes
differing in heat production and water loss, and with different storage
temperatures - this last because the increase in WVPD resulting from a 1°C
Table 14.5 Calculated average WPVD operative in a continuously

ventilated potato stack in a state of balance with air of constant
temperature and humidity*
Rate of ventilation (m 3 t- 1 h- 1)
10 20 40 60 SO 100 200
Inlet WVPD of Average WVPD operative in stack (mbar)
ventilating air (mbar)
0 1.1 0.6 0.4 0.3 0.2 0.2 0.1
0.5 1.5 1.0 O.S 0.7 0.6 0.6 0.5
1.0 1.S 1.4 1.2 1.2 1.1 1.1 1.0
2.0 2.5 2.2 2.0 2.0 2.0 2.0 2.0
3.0 3.2 2.9 2.9 2.9 2.9 2.9 2.9
4.0 3.S 3.S 3.S 3.S 3.S 3.S 3.9
• The following assumptions are made: heat output, 50 kJ rl h- ' ; potential
water loss, 10 g t- I h- 1 mbar- I (= 0.17% week- I mbar- ' ); inlet temperature of
ventilating air, 10°C; relationship between rate of ventilation and water loss
as in Table 14.4.
rise in temperature rises progressively from 0.45 mbar at 0-1°C to 1.39
mbar at 19-20°C (Table 14.3).
The main point of interest which emerges from Table 14.5 is that with
humid air, the faster the rate of continuous ventilation the less the WVPD
operative in the stack, and the lower would be the evaporation. This is to
be expected, in that, with humid inlet air, the main factor influencing
WVPD is temperature rise, and this is less the higher the rate of
ventilation.
With drier inlet air, on the other hand, the rate of ventilation, at least
within the range of normal commercial rates, has little or no effect upon
water loss. The foregoing applies only to continuous ventilation when a
state of heat balance has been reached and the temperature gradient is
solely due to the interaction between heat production, water loss and the
rate of air flow. The evaporative capacity of ventilating air during out-of-
balance periods, as for example when potatoes are being cooled by
ventilation, is altogether different and is discussed below. It will have been
noticed that the conclusions above, on the effect of the rate of ventilation
upon evaporative capacity and water loss, appear at first sight to contradict
the previous conclusions reached when discussing convection. This is only
because, as mentioned earlier in this Section condensation at the cool top
of a convectively cooled stack humidifies the ingoing air. Replacement of
convection by positive ventilation from below at a similar rate removes this
source of humidification and can, as shown in the first column in Table
14.5, appreciably increase water loss if the ambient air is humid. If
ventilation is used for cooling a stack of potatoes it is frequently in the form
of intermittent ventilation when suitable cool air is available. The effect of
this upon water loss can be illustrated by an example based upon
temperature measurements on a 4000 t stack, by Hylmo etal. (1975b). The
stack, 4.5 m high, was continuously ventilated with humid air to maintain
uniformity of temperature and minimal evaporation. The fans were then
switched off for 3 days, thus allowing the stack to rise to a temperature
ranging from 7.8 to 8.7°C. It was then cooled by ventilation at 30 m3 t- 1 h- 1
with air averaging 6.1°C and 97% relative humidity (RH). From the
humidity at entry and the figures in Table 14.4, it can be estimated that the
air would probably leave the stack at about 99.5% RH. The calculated
water loss during the period of cooling is given in Table 14.6, from which it
will be seen that the rate of water loss at the beginning of the cooling
period would have been about six times as great as given in Table 14.5 for
continuous ventilation at the same rate with air of similar humidity, and
that the weighted mean rate of loss over a 50 h cooling period would have
been over three times as great (27 g t- 1 h- 1), which, on the assumption of a
water loss of 10 g C 1 h- 1 mbar- 1 would be equivalent to a mean operative
WVPD in the stack of 2.7 mbar. Losses from stacks differing more from
the temperature of the ventilating air would be even greater during cooling
by ventilation.
Physics 635
Table 14.6 Calculated water loss, and equivalent average operative WVPD, in a potato stack during cooling
by ventilation (temperatures during cooling from Hylmo et ai., 1975a, b)
Time after start Air entering stack Air leaving stack Warer loss Operative
of ventilation (h) (g t- I h-I) WVPD
Temperature ("C) Water Temperature (0C) Water equivalent
content content to water loss·
(g per 30 m3) (g per 30 m3) (m bar)
0-1 6.1 210 8.8 260 50 5

10-11 6.1 210 8.2 250 40 4
20-21 6.3 215 7.8 240 25 2.5
50-51 6.1 210 6.6 225 15 1.5
See Table 14.3. RH at entry 97%; on leaving 99.5%.

• Assuming a water loss of 10 g t- I h- I mbar- I .
Recirculation
Recirculation is the term applied to contioous ventilation with store air, in
the case of bulk storage being introduced through ducts under the potatoes
and the air intake usually in the roof space above the potatoes. If the store
is sealed, depending for temperature control upon heat leakage through
the structure, the air is rapidly humidified by its passage through the
potatoes - the volume of air in a filled bulk store could be some 2 m3 t- 1 ,
capable of holding at a temperature of, say 7°C, only about 15 g of water
vapour per tonne stored (Table 14.3). There will be some loss of this water
by condensation on cold parts of the structure and by air leakage from the
store; there may also be dilution when it is necessary to introduce outside
air, for cooling purposes, but nevertheless the store air can often be
maintained with a very low WVPD, of the order of 0.5 mbar. In such a
case, recirculation is equivalent to continous ventilation with air with an
inlet WVPD of 0.5 mbar. At the rate of 30 m 3 t- 1 h- 1 this would give an
operative WVPD in the stack of about 0.9 mbar (Table 14.5) and an
evaporative loss of perhaps 0.15% week-I.
Refrigeration
The effect upon humidity of the temperature differential between coolers
and air has already been discussed (Section 14.2.1). An efficiently designed
refrigeration plant could ensure that the ventilating air entering the
potatoes has a WVPD not exceeding about 1 mbar, the WVPD in the
stack, again with ventilation at a rate of 30 m 3 t- 1 h- 1 , being about 1.3 mbar
(Table 14.5) and evaporative loss perhaps rather more than 0.2% week-I.
If refrigeration is employed, it is often intermittent and coupled with
continuous recirculation, the overall average loss being then somewhat
less. It would be pointless to have the expense of refrigeration without
virtually continuous ventilation, as temperature gradients in the stack
would override any benefit of a low inlet WVPD (Table 14.6).
Humidification
We have seen that ventilation takes two forms: continuous, which may be
in the form of recirculation, to maintain a uniform temperature; and
intermittent, often with outside air, to cool a stack which, between periods
of ventilation, is unventilated except by convection and thus develops a
temperature gradient (Section 14.2.1). In the case of continuous ventila-
tion the WVPD which is operative in the stack is governed by the inlet
humidity and the rate of ventilation, particularly the former (Table 14.5).
In the case of intermittent ventilation, the most important factor influenc-
ing the operative WVPD is the temperature gradient which has developed
in the stack (Table 14.6). As in the case of refrigeration, humidification of
the inlet air using, for example, a water spray in the air stream, can only
really be effective in the absence of a l.arge temperature gradient - that is
with continuous ventilation. With this, humidification, as used by Hylmo in
Sweden, will give an entry WVPD of about 0.3 mbar and an operative
WVPD, in a stack ventilated at 30 m 3 t- I h-l, of perhaps 0.7 mbar.
Evaporative losses can thus be kept down to about 0.12% week-I,
14.2.3 Respiration
(a) General
Respiration (biological oxidation) is the oxidative breakdown of the more
complex substrates normally present in the cells, such as starch, sugars,
and organic acids, to simpler molecules (C0 2 and H 2 0), with the con-
current production of energy and other molecules, which can be used by
the cell for synthetic reactions. Such metabolic reactions are essential for
maintenance of cellular organization and membrane integrity in living
cells. Maintaining the supply of adenosine triphosphate (ATP) is the
primary purpose of respiration.
The overall process of aerobic respiration involves the regeneration of
ATP from ADP (adenosine disphosphate) and Pi (inorganic phosphate)
with the release of CO 2 and H 2 0. If hexose sugar is used as the substrate,
the overall equation can be written as follows:
C 6 H 12 0 6 + 602 + 38 ADP + 38 Pi ~ 6C0 2 + 44 H 2 0 + 38 ATP
About 42% of the total energy produced (2786.22 kJ) is biologically useful
energy, and the remainder is dissipated as heat.
Aerobic respiration involves a series of reactions, each of which is
catalysed by a specific enzyme and breaks down a complex molecule to a
simpler one (Kader, 1987). It involves the following three metabolic
pathways.
1. glycolysis, i.e. the breakdown of glucose into pyruvate, which takes

place in the cytoplasm;
Physics 637
2. tricarboxylic acid (TCA) cycle, i.e. the breakdown of pyruvate into
CO 2 , which occurs in the mitochondrial matrix;
3. electron transport system, where low-energy nicotinamide adenine
dinucleotide (NAD) is reduced to the high-energy reduced form,
NADH.
With a good oxygen supply, the volumes of the amount of oxygen taken
up and the amount of carbon dioxide released per unit time are virtually
the same. The respiration quotient:
volume absorbed (0 2 h-I)
RQ= =1
volume formed (C0 2 h-I)
Burton (1974), however, found values of RQ = 0.8-0.9 early in the
storage period, during dormancy, while RQ later rose to 1.3 during
sprouting.
In senescent potatoes, CO 2 production is permanently greater than
oxygen consumption (RQ less than 1) (Isherwood and Burton, 1975).
Isherwood (1973) gives for basal respiration at 10°C a formation value of 2
ml CO 2 kg- I h- l , which is equivalent to the formation of 1.4 mmol ATP 100
g-I 24 h-l. This also entails the breakdown of starch and the formation of
anhydroglucose units of 0.5 x 10-2 mmol g-l 24 h- I or dry matter conversion
of 0.855 mg g-I 24 h- l . This corresponds to a loss of dry matter of about 0.85
mg g-l fresh weight per 24-h period. Part of this loss is compensated by the
water released. The above mentioned values for the respiration rate during
storage are influenced by a number of factors. Firstly, they are influenced
by factors associated with the potato itself as presented for storage, e.g.
morphological characteristics. The skin and lenticels are very important for
the oxygen permeability. Other determinants are also important, e.g.
maturity, dormancy, physiological age, cultivar, inhibitory or stimulatory
chemical compounds present and handling followed by healing of wounds.
Secondly, of course, the conditions prevailing in the store are very
important, e.g. the storage temperature and the oxygen and carbon
dioxide concentrations in the storage atmosphere. Respiration rates
and evaporative loss rates are important in considering the design and
management of potato stores.
Besides the normal respiration pathway, in potato tubers the cyanide-
resistant respiration occurs (Solomos, 1977). The latter is a type of cellular
respiration insensitive to terminal inhibitors such as cyanide, azide and
carbon monoxide, and to inhibitors that act between b- and c-type
cytochromes (Meeuwse, 1975). Temporary replacement of a functioning
classical electron transport chain by the alternate is energetically inefficient
as two of the three 'sites' for A TP-synthesis are bypassed. Electron transfer
to oxygen via the alternative pathway only occurs when the cytochrome
pathway is fully saturated (Theologis and Laties, 1978). For that reason it
is doubtful whether cyanide-resistant respiration is even engaged in storage
organs like the potato, although the alternative path might be involved in
these tissues when substrate mobilization is enhanced to such an extent
that the cytochrome path is overwhelmed (Solomos, 1977). Indeed, van
der PI as and Wagner (1983) showed that in potato tuber callus during some
stages there is a relation between endogenous sugar concentration and the
activity of the alternative pathway, pointing to an 'energy overflow
function'. There is also a general tendency for cyanide-resistant respiration
to survive in ageing and senescence as the cytochrome path disintegrates
(Laties, 1982).
(b) Gas exchange

The ability of a tuber to respire aerobically must depend upon the supply of
oxygen to the respiring tissue. This is limited by the periderm of the tuber,
which presents a barrier to gaseous diffusion. The bulk of this diffusion
occurs through the lenticels, of which there are some 0.5-3 per cm 2 of
surface of the fully grown tuber (Burton, 1966; Wigginton, 1973). Their
effective area can only be the area of the exposed intercellular spaces of the
underlying tissue, and has been calculated to be of the order of 10'6 times
the surface area of the tuber, or 1-2% of its volume (Burton, 1950).
The number of lenticels was observed to be about 100 per tuber. This
number was influenced by the size of the tuber, climatic conditions and soil
type; higher phosphate fertilization gave less lenticels (Meinl, 1966). Rates
of oxygen diffusion at 10°C varied from 0.024 to 0.296 cm 3 h'l atm,l with a
mean of 0.1105 ± 0.013 cm 3 h'l atm,l in King Edward tubers (Wiggington,
1973).
The intercellular space offers little resistance to diffusion through most
of the space at the rates encountered in plant respiration. Even in the
potato, with its very small space, the resistance is so slight that it is difficult
to detect any consistent oxygen gradient from periphery to centre (Burton,
Table 14.7 Rates of diffusion through the integument and internal atmosphere
immediately under the integument, of some samples of potato
Internal Diffusion Reference
atmosphere (%) through skin
Commodity immediately (ml kg'1 h'l per
under skin 1 % gradient)
CO 2 O2 CO 2 O2
Potato, stored
Arran Consul (lO°e) 1.8 18.2 1.0 0.7* Burton (1950)
Arran Consul (25°e) 7.0 13.0 0.8 0.7* Burton (1950)
Majestic (l0°e) 2.2 18.3 1.1 0.9 Burton (1974)
* Derived from CO 2 output on the assumption that RQ = 1. (From Burton, 1978b.)
Physics 639
1950). In senescent tissues the intercellular spaces may become filled with
cellular solution, which impedes O 2 movement and results in anaerobic
conditions within the tissue. The resistance to gas diffusion in a potato
tuber is not limited by the lenticels but by the intercellular spaces
immediately underlying them (Burton, 1978b), which means that the O 2
and CO 2 status of the tissue is little different from that of the ambient
atmosphere, as can be seen in Table 14.7.
Effect of a water film on the internal atmosphere

Because O 2 enters the tuber, and CO 2 leaves, by gaseous diffusion through
the lenticels, the internal atmosphere responds to conditions which inter-
fere with this. For example, if the surface of a tuber is completely covered
by a film of water, diffusion must occur, much more slowly, in solution. As
a result of this, Burton and Wiggington (1970) found a persistent film of
water to lead to tubers becoming anaerobic in about 6.5 h at lOoC and in
about 2.5 h at 21°C (Fig. 14.7). Ringel (1976) confirmed these results at
21.5°C and measured that after drying within a few hours the oxygen
concentration reached the original value. Tubers partly immersed in water
did not become anaerobic although the oxygen tension, determined in the
centre of the immersed part, was reduced to an extent related to the area of
periderm covered by water (Burton and Wigginton, 1970).
w5
:::l
en
~
E<..l 4
Q;
C.
'"b 3
x
100 200 300 400

Length of time (min)
Figure 14.7 The oxygen content of the centre of tubers which had been covered
by a water film (average thickness c. 3 x 10- 2 mm) for different lengths of time.
(0), 10°C; (e), 21°e. (From Burton and Wiggington, 1970.)
Oxygen concentration
According to Bahr and Bonner (1973a, b) the total respiratory rate (Vtot ) is
composed as follows:
V tot = pValt + V cyt + V res
In this equation V cyt represents the capacity of the cytochrome pathway
under the given experimental conditions, Valt is the capacity of the
alternative pathway and p is the fraction of the alternative pathway
capacity which is operative in uninhibited respiration. V res is the residual
respiration, which is insensitive to inhibitors of both cytochrome and
alternative pathways.
Since potato tubers are bulky organs with a rather impermeable
periderm, the question arises as to whether the availability of oxygen to the
cell is an explanation for the low rate of respiration. However, taking into
account the low O 2 demand of the whole organ, the periderm of freshly
harvested tubers appears rather permeable, allowing a sufficient rate of O 2
diffusion of 20 to 50 III cm- 2 S-l atm- 1 (Burton, 1950; Mapson and Burton,
1962). Under ordinary conditions, air is adequate to sustain respiration
fully, since elevation of external O 2 concentration has no effect on organ
respiration rate (C0 2 production).
The major part of respiration of intact tubers is mediated by a high
oxygen affinity terminal oxidase, the cytochrome-c oxidase, which is
unaffected by O 2 concentration except at low levels. Craft (1963) studied
1-6% O 2 ; Mapson and Burton (1962), 1.4-100% O 2 ; and Burton (1974)
air, 5% and 100% O 2 , Table 14.8 shows results of this, typical of tubers
during the first weeks after harvest. Mapson and Burton (1962) found in
pure O 2 however an increased O 2 uptake in comparison with the O 2 uptake
in air, which in view of the concentrations of O 2 found in the centre of the
tuber in air was explained as an increased uptake of low-affinity oxidase
(e.g. polyphenoloxidase). Burton (1974) found low-affinity uptake to be up
to about 30% of the whole at the time of harvest, but to fall almost to zero
over a period of about 1.5-3 months. Thereafter, it reappeared in the
spouting tuber, although it is possible that this reappearance was in the
respiration of the sprouts and not of the tubers.
When mature potato tubers (cv. Bintje) were stored in normal air at 4°C
and the O 2 uptake of tuber discs measured at regular intervals at 25°C, then
Table 14.8 Typical relation, within a few weeks of harvest, between ambient O2
concentration and O2 uptake by potato tubers at 20°C (cv. King Edward, O2 uptake
in air, 5-8 ml kg- 1 h- J ) (based on Mapson and Burton, 1962)
O 2 (%) O 2 uptake as %
uptake in air
14.7 92
9.8 81
7.0 73
5.0 70
2.5 59
1.4 55
Physics 641
the average O 2 uptake during a storage period of 200 to 350 days of V CYI
was about 17 nmol O 2 min- 1 g-1 fresh weight, of Vall about 3.5 nmol O 2 min- l
g-1 fresh weight. The V res between 200 and 300 days was about 8.5 nmol O 2
min- 1 g-l fresh weight and between 300 and 350 storage days decreased to
about 4.5 nmol O 2 min- 1 g-1 fresh weight. Lowering the O 2 concentration of
the storage atmosphere resulted in significant lower values of V cyl , but not
of Vall' When potatoes were stored at 5%, O 2 there was no significant
change in V cyl or Vall (Michielsen and Hartmans, 1988, unpublished).
Consideration of the effects of O 2 concentration on CO 2 output is
complicated by a delayed response of decarboxylation (Burton, 1974).
Despite this, exposure for only a few hours to 5% O 2 leads to some
reduction in CO 2 production compared with production in air (Burton,
1974) and the effect becomes more marked as the O 2 concentration is
further reduced (Craft, 1963). Even in N 2 , however, Craft found the
production of CO 2 over 24 h to be about 60% of that in air. Sherman and
Ewing (1983) measured the CO 2 production in N2 at 10 and lac. Storage at
laC reduced CO 2 production and storage in N2 at WaC depressed CO 2
output for 8-10 days; after that time, CO 2 production increased until the
experiments were terminated (4-5 weeks).
Exposure for some hours to pure O 2 has little or no effect on CO 2 output
(Chin and Frenkel, 1977; Theologis and Laties, 1982). Prolonged exposure
leads at first to an increase in output, over a period of 2 weeks or so, but
thereafter the effects of '0 2 poisoning' become apparent and after 5--6
weeks CO 2 output may be negligible (Barker and Mapson, 1955).
Carbon dioxide concentration

When the ventilation in a potato store is in any way insufficient, the CO 2
concentrations may increase by virtue of the gas exchange: O 2 uptake and
CO 2 release. Vendelbo (1980) found in commercial potato stores a
maximum of 3.5% CO 2 and up to 7.5% in pilot stores. Yarns (1986)
measured, in an unventilated commercial potato store during the suberiza-
tion period just after harvest, CO 2 concentrations between 4 and 6%.
These high CO 2 concentrations were rapidly reduced to < 2% if wind
speeds outside averaged 4.5 m S-1.
The influence of increased CO 2 concentrations on the respiration rate
has been studied by several authors. Burton (1952) measured O 2 uptake
and CO 2 output in the presence of 5-7% CO 2 over periods of 11-14 weeks
at wac. The increased CO 2 reduced both O 2 uptake and CO 2 output by
about 25-30%, the respiratory quotient being practically unaffected.
Perez-Trejo et ai. (1981) found however that application of high CO 2
concentrations (3-30%) to whole potato tubers at room temperature
gave a rapid and pronounced respiratory gas exchange, which persisted
for a prolonged time (at least 48 hr). COz-stimulated respiration was
accompanied by a pronounced decline in the content of starch and glucose-
6-phosphate, suggesting an active utilization of respiratory substrates.
When potaotes at 20 C were exposed to high levels of CO 2 (5-25%) for a
D
few days only and returned to air afterwards, the respiration was reduced,
with the higher concentrations resulting in a greater retardation. The
reduced respiration rates were maintained for at least 2 weeks after
treatment while the produce was ventilated with air (Wills et ai. 1979).
Increased CO 2 concentrations probably occur very often in many potato
stores, due to insufficient ventilation. The influence of CO 2 on changes in
respiratory metabolism of whole potatoes may be one of the most
important processes associated with the metabolic effects of CO 2 • It might
be that the changes are especially dependent on the concentration,
exposure time, temperature and physiological age of the tubers.
(c) Temperature and heat production

Temperature
The direct effect of temperature on respiration in potatoes is of relatively
short duration (7-10 days) (Hunter, 1986). The short-term effect of
temperature upon the rate of respiration is well illustrated by the results of
Miiller-Thurgau (1882), showing a two-fold increase for a lODC rise
in temperature over the range 0-20 C. This short-term effect is not
D
maintained during storage at different temperatures and the QlO may be

very different from 2. Burton (1974) found a mid-storage temperature
coefficient, 20/lO C, of about 1.2-1.3, but respiration at 2 C (sweetened
D D
tubers) could be higher than at 20 DC. Later, increased respiration accom-

panying sprouting - most vigorous at 20 DC and non-existent at 2DC -
changed this pattern. An example of the relationship between rate of
respiration and temperature at various stages during the storage period is
presented in Fig. 14.8. The increase in rate of respiration in this case
noted at higher storage temperatures after December was undoubtedly
due, at least in part, to sprouting (Schippers, 1977). Even when sprouting
is inhibited there is however a constant change in temperature coefficient
during storage (Fig. 14.9), as shown by Hunter (1986). It is therefore
difficult to establish a fixed relationship between the rate of respiration and
the storage temperature since at some temperatures the rate of respiration
is fairly constant, whereas at others it is not.
Heat production
For every g of carbohydrate consumed in normal respiration about 16 kJ of
energy are released. In part this is transferred into energy-rich phsphate
molecules and utilized in metabolism; in part it is lost as heat. There have
not been many measureme'nts of the proportion so lost. Green et ai. (1941)
suggested it was as much as about 90%. If we accept this for tubers in
which no marked change is occurring, then for every g of CO 2 produced in
respiration about 10 kJ of energy are released in the form of heat. The
amount could differ during periods of rapid change but evidence for this is
Physics 643
8
l Mar 1974
7 /
/
/
6
/
/
5 / P Feb 1974
:c:
'0)
.x: / . . l/
4
/ ./
C\j
0
()
0) /..... ,e Jan 1974
E /Y"'" /'
3 _ _x / l/'
~.......... ~ / ....d/ ___x Nov 1973
/ ...... ;.--
2 ~~~ ~
......... Y •••••••• ~/~ _._._.0 Dec 1973
D"·····..~~···.. .......... ~~ ·0'
---.-...:~-- ::::.:~~-.: ~;;."..,. .........
2.5 5.0 7.5 10.0 12.5 15.0

Temperature (0C)
Figure 14.8 Relationship between storage temperature and rate of respiration at

various dates during the storage period 1973174 (average of six cultivars). (From
Schippers, 1977,)
lacking. The heat produced must be considered in selecting proper

methods for cooling, package design, method of stacking packages, and
refrigerated storage facilities.
(d) Period of storage and reconditioning

Period of storage
Mature tubers show a fall in respiration rate after harvest and an increase
after the end of dormancy (Schippers, 1977; Frydecka-Mazurczyk, 1978;
Hunter, 1986). In general, respiration rate decline is more rapid at higher
temperatures during the falling rate period and rate increase more
rapid during the rising rate period (Schippers, 1977; Hunter, 1986). The
respiratory rate of dormant potato tubers is extremely low. Many available
results are in the range of 2-8 J.lI O2 g-l fresh weight h- 1 (Dizengremel,
1985). Several authors have found a reasonably constant level in rate of
respiration during part of the main storage period. At normal storage
6 (a)
r
5 l
:c l
x
~4 /
OJ l
X
E.
c
3 X
/
o .x' .
~
'5. 2
/'
(/)
Q)
II:
0
0 20 58 84 112 140 188 196 224 252
Days in storage
6 (b)
:c
';"
OJ 4
::t:
OJ
E. 3
c
0
~
'5. 2
(/)
Q)
II:
o
o 20 58 84 112 140 188 196 224 252
Figure 14.9 Influence of maleic hydrazide (MH) treatment of the plants (cv.
Kennebec) on the respiration of the mature tubers during storage at three different
temperatures: (a) untreated 1974-76 and (b) treated MH 1974-78. D, 3.3°C; 6,
7.22°C; +, 12.8°C. (From Hunter, 1986)
temperatures, the rate of respiration changes slowly over the course of the
storage period (Schippers, 1977).
Reconditioning
Many authors found that fluctuating temperatures in general lead to a
'respiration burst'. Burton (1974) showed that it was necessary neither to
Physics 645
make temperature changes extreme in order to obtain the 'respiratory
burst', nor to change from a low to a high temperature. He changed the
storage temperature of tubers of the cultivar of King Edward from 10 to
2°C and found a rapid increase in rate of respiration in about 2 weeks. The
rate then dropped in another 2 weeks and remained there until the tubers
were transferred back to 10°C about a month later. In this case the rate of
respiration increased in 2 days. The following decrease to a constant level
took about 2 weeks. (see Fig. 14.10).
10
o
n
8 1\
I'B
1\ B..-
:c I \ ...-
~ 6 •
I \0 . . - ...-"'-
I 1\ I '0---0"'-
Q)
-'" ~ \ I
R4 /\ ~.\ I
I·'
,
o
:0
:c \ I
1 ' . - -/- - - ::..... ~.-~_-- .. ---- __ ~'"
oL-________L-____
1 December 1 January
•
~ __
Sample B
~ ____
1 February
•
2_~
Sample C
____
1 March
•
~ _ _ _ _L __ _ _ _ _ _
Sample B
1 April
~L_ _ __ _
1 May
placed at 2°C placed at 2°C replaced

at 10°C
Figure 14.10 Oxygen uptake of potatoes, cv. King Edward, at 10°C and during
sweetening at 2°C and subsequent de-sweetening at lO°C. (0), lOoe; (e), 2°C;
---, at lOoe throughout; ----, subjected to temperature change on the dates
specified. (From Burton, 1974.)
(e) Influence of handling, wounding and irradiation

Handling
Barker (1935) found that turgid tubers could be handled without serious
disturbance of their respiration, but that if the tubers were soft, gentle
squeezing could increase the rate of respiration by about 30%. More
vigorous treatment, such as that involved in washing and drying the tubers,
was found by Burton (1974) to more than double the O 2 uptake by slightly
wilted tubers and to cause a 30% increase in that by freshly harvested
tubers. In both cases the effect was temporary and disappeared in the
course of a day. Isherwood and Burton (1975) found handling to lead
to variable respiratory quotients, an effect which could last for several
weeks.
Aeppli and Keller (1980) subjected mature harvested potato tubers to a
standardized shaking treatment and measured the respiration at lOoC.
Respiration after treatment increased with violence of the shaking as did
also the severity of blue spot. It was observed that increasing susceptibility
of the tubers to blue spot was accompanied by a proportional increase in
respiration rate (Fig. 14.11). During the storage season 1977178 the
respiration rate was found to increase with approximately equal sharpness
following three shaking treatments carried out at intervals of 6 weeks.
During 14 weeks subsequent to treatment it never fell to the level of that of
14 •
• •
12
••
•
•
· ,,
."
••
::-- 10 o ,
/ /
:c , /
~ o
N
o
r
<.> o
8
c
o •
~
'0.
~
.S
6 .• . ..
Q)
• y = 2.06 + 0.092 x
••
~
~ r = 0.83
<.>
E 4 •
o
o 20 40 60 80 100
Blue index
Figure 14.11 Relationship between blue spot and increase in the respiration rate
up to 48 h after a shaking treatment, based on 144 tests of tubers from field
experiments in 1976 (0) and 1977 (e). (From Aeppli and Keller, 1980).
Physics 647
untreated tubers. The maximum respiration rate was reached 43 h after
shaking at the end of October and 96 h afterwards in the middle of
January. Later treatment resulted in a moderate increase in the tendency
to develop blue spot.
Wounding
When potatoes are wounded there is also an increase in respiration. This
increase in respiration develops during the dedifferentiation of the
cells beneath the wound surface and depends on the synthesis of RNA
protein. This dedifferentiation is followed by a redifferentiation phase
(suberization, periderm formation, callus outgrowth). When periderm is
formed the induced respiratory rise is generally transient (Fig. 14.12).
After the 'closing of the wound' the cells beneath the wound surface
become metabolically less active again and the level of respiration returns
to that of the intact organ. In particular the respiratory rise seen during
the first days after wounding the tissue (induced respiration) has been
extensively studied (see reviews of Kahl, 1973, 1974; van Steveninck, 1975;
Laties, 1978).
;:-
5-L:
~'7C)_
.!:N;: Induced
5}O {j5 respiration
&"3.~
+- Freshly sliced tissue

Initial
increment
+- Intact organ
Time
Figure 14.12 Respiratory activity of potato tuber tissue after wounding and during
differentiation in different directions. (Data derived from Lange, 1970; Rosenstock
et at., 1971; after van der Plas, 1985.)
Irradiation
y-irradiation causes a marked temporary increase in respiration, rising to a peak
in a week or so and then falling to about the control level (Sussman, 1953).
(f) Influence of chemicals

'Metabolic' volatiles
Hanes and Barker (1931), who investigated the influence of hydrogen
cyanide because of its known effect as a depressant of cell respiration,
found that it increased the rate of respiration of potato tubers three-fold
when the tubers were kept in an atmosphere containing low dosages. After
~ 10 days of treatment the rate of respiration dropped even if the tubers
remained in the same atmosphere. At a higher dosage the increase was as
high as six-fold, but in that case a contributing factor may have been injury
to the tubers which occurred in the form of brown patches and sunken
areas. Craft (1963) in short-term experiments, found stimulation of O 2
uptake and CO 2 output at 25°C to be caused by a much higher (1 %)
concentration of HCN.
One class of chemicals which might be expected to have an influence on
the rate of respiration consists of those which are able to break the
dormant period of the tubers. Huelin and Barker (1939) investigated the
influence of ethylene shown by Rosa (1923) to have a restbreaking effect,
and found that a concentration of 0.1 % of ethylene in the air was effective
in increasing the rate of respiration at 15°C, particularly later in the storage
season. Ethylene treatment of whole potato tubers leads to an increase in
respiration of the intact organs (Reid and Pratt, 1972; Solomos and Laties,
1975) and induces the operation of the alternative pathway in freshly-cut
slices (Chin and Frenkel, 1977; Rychter et al., 1978) and in mitochondria
isolated therefrom (Rychter et al., 1979a). The response to ethylene in
pure O 2 is much greater than in air (Rychter et at., 1978; Theologis and
Laties, 1982).
The ability for ethylene to break dormancy in dormant tubers can
be obtained easily with Rindite, a mixture of anhydrous ethylene chloro-
hydrin, ethylene dichloride and carbon tetrachloride. The treatment of
whole tubers by Rindite induces dramatic changes in the activities of
subcellular fractions.
Rindite appears to be a more effective treatment than ethylene for
inducing both a general increase in oxidative activities and the functioning
of an important cyanide-resistant electron transfer (Bonnerot et al., 1985).
Other naturally occurring volatiles may act similarly to ethylene. Propylene
(Chin and Frenkel, 1977), ethanol, acetaldehyde and acetic acid, when
applied in a volatile state in air to potato tubers, led to a climateric-like
upsurge in respiration (Rychter et al., 1979b). Again the respiratory
upsurge was markedly enhanced when the volatiles were applied in 100%
O2 ,
Most attempts, other than the above, to elucidate the pathways of
respiration by using metabolic inhibitors and stimulants have employed
discs of tuber tissue, because only by such methods can inhibitors or
stimulants, other than gaseous ones, be introduced. The pathways so
elucidated, however, are those occurring in discs of tuber tissue, and not
necessarily in intact tubers. The literature on this subject was extensively
reviewed by Dizengremel (1985).
Physics 649
Sprout inhibitors
Sprout inhibitors are another group of chemicals which may be expected to
have an influence on the rate of respiration, if not during the dormant
period, then at least after the termination of the formal rest period. Only a
few authors have investigated the influence of commercial sprout inhibitors
like isopropyl N-phenyl carbamate/isopropyl N-(3-chlorophenyl) carbamate
(IPc/CIPC) and maleic hydrazide (MH) on respiration. Boe et al. (1974)
studied the influence of CIPC, Lougheed et al. (1975) CIPC and MH, and
Hunter (1986) MH. Boe et al. and Lougheed et al. concluded that the
respiration rate of the tubers which had been treated was less than that of
untreated tubers. However it can probably be assumed that sprouting
played a significant role in the rate of respiration of untreated tubers,
although the degree of sprouting was not mentioned in the two papers.
Hunter found, during a long storage period of about 250 days at 3.3 or
7.2°C, no difference in respiration rate between MH-treated and
untreated tubers of the cvs Kennebec and Russet Burbank. At a storage
temperature of 12.8°C for both cultivars, there was an increase in respira-
tion rate of untreated tubers, when dormancy ended and these tubers
started to sprout.
When potatoes were hand-desprouted the respiration of stored tubers
was not markedly different from CIPC-treated tubers (Isherwood and
Burton, 1975); however, respiration increased considerably, by about four
to five times, when tubers were allowed to sprout normally (Table 14.9).
At mitochondrial level there was no effect of CIPC treatment on
respiratory activity or behaviour of mitochondrial respiration during the
early stage of storage (Chung, 1985). There were also similar patterns in
Table 14.9 Rates of respiration, cv. Home Guard; January-June 1973

(monthly average values, ml kg- 1 h- 1 at NTP). Storage temperature till
January lOoe and then transferred to 200 e
Month Untreated sprouting Hand desprouted CIPC-treated
tubers
CO 2 O2 CO 2 O2 CO 2 O2
Jan. (2 weeks) 8.2* 7.0
Feb. 9.5 8.7 3.8 3.6
March 12.3 11.2 3.9 3.6
April 14.0 12.6 4.8 3.7 4.5 3.8
May 15.8 14.9 4.7 3.8 4.9 4.3
June 18.1 16.6 3.7 4.2 4.7 4.8
(1 week) (1 week) (2 w·ceks)
* The samples were hand de sprouted on 12 January, before the start of
measurements on 16 January, and thereafter left undisturbed. Sprout regrowth
started immediately and the rate of respiration during January would be enhanced
because of this. (From Isherwood and Burton, 1975.)
mitochondrial respiratory activity between CIPC-treated and untreated
tubers during the early stage of reconditioning.
(g) The effect of tuber characteristics

Maturity
The rate of respiration of potato tubers, measured after removal from the
plant, is high soon after tuber initiation and decreases throughout the
growing season, reaching a fairly steady, though slowly declining, value at
maturity (Appleman and Miller, 1926; Singh and Mathur, 1937; Burton,
1964, 1974; Peterson et al., 1981). Burton (1974) and Peterson et al. (1981)
measured respiration rates continuously at lOoC over a short period after
harvest. As an example the values after 1 and 3 days are shown in Fig.
14.13 (Burton, 1974).
The high rate of respiration of immature tubers was not maintained after
harvest but falls off rapidly, approximating to that characteristic of mature
tubers after 2-4 weeks (Burton, 1963, 1974). Peterson et al. (1981)
however found that respiration rates were high immediately after harvest
and then declined to an equilibrimn level after approximately 6 days.
Throughout the season, the respiration rate after 1 day was always higher
than at 6 days after harvest. However differences declined as the tubers
matured. This would suggest that harvesting trauma was more significant
on the more immature tubers. Both authors showed a close correlation
between respiration and maturity of tubers, this correlation being
independent of the physical characteristics of the tuber, namely, surface
area or tuber size (Peterson et al., 1981). Respiration may be a good index
of potato maturity.
24
';-
Cl
~ 16
I ...
~ 12
til
a.
~ ... ... ... •
8
0'" ... ... ...
4
20 40 60 80
Days after 27-06
Figure 14.13 Oxygen uptake of tubers, cv. Majestic, during the first 3 days after
harvest. (e), day 1; (A), day 3.(Redrawn from Burton, 1974.)
Wound healing and curing 651
Cultivar and tuber size
The rate of respiration is a clear characteristic of the cultivar (Schippers,
1977; Frydecka-Mazurczyk, 1978; Hunter, 1986) and independent of both
earliness in the field and length of the dormant period (Schippers, 1977).
The question of whether there is an influence of tuber size on respiration
has to be considered in sampling procedures. Several authors mentioned a
relationship between tuber size and respiration rate. Michaels (1932) and
Weber-Dahlmann (1957) found that the rate of respiration of smaller
tubers was higher than of larger tubers. Meinl (1967) confirmed the role of
tuber size, but obtained different results. Schippers (1977) found no
significant differences in respiration rate at 7.5°C between small (70-100 g)
and large (160-250 g) tubers when studied in 11 cultivars. Also Peterson et
al. (1981) concluded that no correlation existed between tuber size (90-400
g) and respiration rate.
Sprout growth
Sprout growth, which involves mobilization and transfer of material from
the tuber to the sprout and its synthesis into the sprout structure, is
accompanied by increased respiration. The respiration of a sprouting tuber
includes both that of the tuber and of the sprouts, and Burton et al. (1955)
found an approximately 50% increase in respiration to have occurred by
the time sprout growth had reached about 1%, by weight. The increase
continues with increasing growth, and Isherwood and Burton (1975)
recorded a four to five-fold increase in respiration by the beginning of June
in considerably sprouted tubers. Much of this may be accounted for by the
respiration of the sprouts themselves. In desprouted tubers Isherwood and
Burton found the increase to be not more than 30% above the basal level,
when comparing the respiration of mature tubers during storage from MH-
treated and untreated plants. Hunter (1986) showed that the increase due
to sprouting depended on the storage period and temperature (Fig. 14.9).
14.3 WOUND HEALING AND CURING
14.3.1 Changes in wounded tissue

Potato tubers are subject to various environmental conditions or factors,
the action of which may entail a partial or even complete destruction of the
tuber. If tubers are not lethally damaged, however, they are able to
regenerate destroyed tissue, to protect themselves against parasitic attack
and loss of water through a process of wound healing.
When potato tissue is damaged a variety of physiological, biochemical
and anatomical changes occurs in the cells surrounding the wound. The
metabolic changes include rapid synthesis of RNA (Tripathi and Kahl,
1982) and the de novo synthesis of RNase (Isola and Franzoni, 1986).
Anatomically, wounding of the tuber tissue initially gives rise to suberizing
of the cells on the surface of the cut. This is called primary suberization
(Rosenstock, 1963). The wound periderm is formed after the onset of
primary suberization. A few cell layers away from the wound surface, new
cell walls are formed parallel to the wound surface (Fig. 14.14). This gives
rise to a phellogen (cork cambium), which serves as the basis for wound
periderm formation: the phellogen cells form a number of phellem cells
parallel to the wound surface. These phellem cells subsequently become
suberized, this process being known as secondary suberization (Rosenstock,
1963).
Wounded surface
Suberized parenchyma
t-+----'~--::::r- ~=1;;~~==t::==t=~ Wound periderm
L..--'-_-L-'>.._---'--'--------<~'---_.L-__'__~"'__ ___"J. Storage parenchyma

Figure 14.14 Wound periderm formation. (Redrawn from Lange and
Rosenstock, 1963.)
Suberization of cells involves the deposition of suberin within the cell

wall and outside the plasma membrane (Kolattukudy, 1984). Suberin
(cork) formation is an important process, as suberization protects under-
ground organs and periderms from diffusion of moisture and microbial
attack. Suberization also occurs at a variety of other anatomical regions in
plants. In the case of the disorder 'hollow heart' suberin can also develop in
the tissue surrounding the hollow regions (Dean et al., 1977). The structure
and composition of suberin are poorly understood. Suberin is insoluble
polymeric material composed of aliphatic domains which are held together
by ester bonds and phenolic domains in which the aromatic components
are held together by linkages that might be similar to those found in lignin
(Kolattukudy, 1984). Suberization in wound-healing potato tuber tissue is
associated with a variety of biochemical changes like the appearance of an
anionic peroxidase (Espelie et al., 1986), the rise in phenylalanine-
ammonia-lyase (PAL) (Smith and Rubery, 1981), and increase in
chorismate mutase (Kuroki and Conn, 1988) and the rise in phenolase
(Rhodes and Wooltorton, 1978). The wound healing process is influenced
by a number of factors discussed in the following sections.
14.3.2 Type of wound

According to Artschwager (1927), abrasion results in a deep seated
periderm, whereas the wound periderm which forms after a smooth clean
cut may be within one layer of cells of the surface. Infection of crush
wounds, as commonly encountered in practice, is usually more severe than
infection of sliced surfaces like those of tuber discs (Adams and Griffith,
1978; Adams, 1980). Healing is more rapid over cut surfaces on tubers than
over bruises on whole tubers (Bland et al., 1987). Vapour loss from cut
surfaces on whole tubers decreased more rapidly than that from cores of
tissue (Bland et al., 1987). Wound healing rates in cortical and medullary
tissue were generally equal (McGee et al., 1985b).
14.3.3 The effect of tuber characteristics
(a) CuItivar
Many workers have reported potato cultivar differences in the rate of
wound healing (Priestley and Woffenden, 1923; Artschwager, 1927; Smith
and Smart, 1959; Nielsen, 1968; Wigginton, 1974; McGee et al., 1985b)
although Wigginton pointed out that the cuItivar differences he observed
may not have been significant because of the variation between individual
tubers. Varietal rankings of the wound healing potential of 15 UK
maincrop cultivars were moderately consistent over four experiments in 2
years, with cvs Desiree, Bintje and Pentland Hawk showing rapid healing
and Majestic, Redskin and Pentland Crown rather slow healing (McGee et
al., 1985b). Correlations with established ratings for resistance to in-
dividual diseases were not found, but some protection against a broad
spectrum of storage diseases is likely. Wound healing ran kings correlated
with varietal resistance to mechanical damage (McGee et al., 1985b).
(b) Maturity
McGee et al. (1985a) showed with the cv. Record a more rapid wound
healing with increasing maturity; the cv. Red Craigs Royal however,
showed little variation with increasing maturity. After storage until March
in the following year, the effect of maturity at lifting on wound healing rate
was no longer evident, but the difference between cultivars still was.
(c) Physiological state of the tuber

Lange and Rosenstock (1964) reported a continuous reduction in the
rapidity of wound periderm formation throughout the development and
storage from young tubers to mature tubers stored for 12 months after
harvest, the reduction being more marked the higher the storage tempera-
ture (Fig. 14.15). Related to this decrease in the number of periderm cell
layers, the suberization decreases and the closing of the wound becomes
more and more incomplete. This decrease in suberization might be
correlated with the decrease in the content of abscisic acid (ABA) during
prolonged storage (Kolattukudy, 1984).
t)
Q)
11 ~
ctI
10 I
!!l 9
-
Qi
(.)
8
0
7
Q;
.0 6
E
z 5
:::J
4
3
2
IX X XI XII I II III IV V VI VII VIII IX X XI XII

Months
Figure 14.15 Number of periderm cells formed after wounding and subsequent
incubation in moist air of potato tubers (cv. Olympia) stored for various times at
7°C (after Lange and Rosenstock 1964). (From van der Plas, 1987.)
During storage of 15 UK main crop cultivars at lOoC their mean wound

healing rate decreased in one year but increased in the next year (McGee et
al., 1985b). The authors suggested a possible influence of sprouting
superimposed on any general trend in that case. Bland et al. (1987) found
no significant change in wound healing of cores or cut tubers as time in
storage varied from 2 to 27 weeks at 6°C before injury.
The conflicting data from changes in wound healing rates during storage
are probably caused by the fact that wound healing has been measured by
decreased conductance using tissue cores. Healing of cores perhaps is
unrepresentative of that which occurs in whole tubers, as Bland et al.
(1987) found less variability of conductance measurements and a more
rapid decrease of vapour loss from cut surfaces on whole tubers than that
from cores of tissue.
14.3.4 The effect of storage conditions
(a)Temperature
Wound healing is more rapid the higher the temperature over the range 2.S
to 20°C (Appel, 1906; Priestley and Woffenden, 1923; Artschwager, 1927;
Radatz, 1967; Wigginton, 1974). The rate of healing was shown to increase
approximately three-fold between S and 10°C, and three-fold between 10
and 20°C (Wigginton, 1974).
Complete suberization of the superficial layer of cells could take 3-6
weeks at SoC, 1-2 weeks at 10°C and 3-6 days at 20°C. Wigginton (1974)
could detect no formation of wound periderm until 3-S days after
wounding at 20°C, 1-2 weeks at 10°C, and up to 4 weeks at SoC.
Experience suggests that the timescale of wound healing in freshly
harvested tubers at 10°C is adequate to prevent serious invasion by wound
parasites, indicating that suberization of the surfaces within 4 days, with
subsequent wound cork formation, may normally be sufficiently rapid.
Thomas (1982) showed that the wound-induced suberization and
periderm formation processes occurred most rapidly at 2SoC. A tempera-
ture of 3SoC prevented periderm formation and retarded suberization.
Impairment of the wound periderm formation therefore probably is a
major cause of the bacterial soft rot occurring in tubers when stored under
high tropical ambient temperatures.
(b) Humidity
With a high water vapour pressure deficit drying occurs at the surface of
the wound and the resulting crust inhibits or greatly delays suberization.
On the other hand, a water-saturated atmosphere hinders wound healing
by causing proliferation of cells at the wounded surface (Olufsen, 1903;
Lange et al., 1970).
At 10°C healing was most rapid above 80% RH and was shown to be
inhibited at 70 and 60% RH. At 20°C little difference was found in the rate
of healing at humidities of 70-100% RH. At S2% the rate of drying of the
surface cells was great enough to inhibit suberization and periderm
formation. At 30% RH the drying of the surface cells was so rapid and
deep that suberization was totally prevented (Wigginton, 1974).
A high humidity appears not quite so important at high temperatures
(20°C) where the effects of surface drying are counterbalanced by the rapid
rate of healing. The effect was not strongly marked, however, and relative
humidities above 80% are to be recommended.
The results are in agreement with the results of Nnodu et al. (1982), who
concluded also that the effect of temperature is more critical than the effect
of relative humidity in the wound healing process.
(c) Sprout suppressants, protective chemicals, radiation
Several chemical treatments and irradiation have been found to inhibit
sprouting effectively, but because they also inhibit wound periderm
initiation, it has been recommended that their application be delayed after
harvest to ensure formation of a continuous wound periderm (Sawyer,
1959). Audia et al. (1962) and Reeve et al. (1963) have investigated the
inhibiting influence of isopropyl N-(3-chlorophenyl) carbamate (CIPe) on
wound healing and Ives (1955) the same phenomenon of isopropyl N-
pheny1carbamate (IPe).
Dimethylnaphthalene (DMN) was shown to be an effective sprout
suppressant (Beveridge et al., 1981b), but also inhibited wound healing
(Hartmans and van Es, unpublished). y-irradiation which can be used to
suppress sprouting (Section 14.4.2) will also suppress the formation of
wound periderm (Waggoner, 1955; Sawyer, 1956). As a general rule sprout
suppressants should be applied only after wound healing is complete,
although tetrachlornitrobenzene (TCNB) appears to provide an exception
to this principle.
In many countries (for instance the United States and Canada) it is
common planting practice to use tubers cut into seed pieces, which exposes
tuber tissues to desiccation and to bacterial and/or fungal seed-piece decay.
Erwinia and Fusarium can become established more quickly than the
natural barriers are formed in response to wounding. For this reason
chemical seed piece treatments are often employed to slow infection, but
seed treatments must not interfere with wound healing. The wound healing
process was only minimally affected by chemical treatment of seed pieces
by mancozeb, zineb-streptomycin, zineb-fir bark, captan, captan-mertect,
captan-fir bark, and sodium hypochlorite (Vander Zaag et al., 1985; Nolte
et al., 1987).
(d) Oxygen and carbon dioxide concentration

The necessity of available oxygen was already known for a long time. As
yet it is unclear, which mechanism regulates this increase in wound-
induced respiration rate.
The, in general, low metabolic activity of tuber tissue during storage can
be activated immediately by wounding. Cutting leads to a four to five times
stimulation of respiration, while during subsequent days of incubation an
extra respiratory rise is induced, which amounts to three to four times that
of freshly sliced tissue (van der Plas, 1987). Lipton (1967) and Wigginton
(1974) found anaerobic conditions prevented both suberization and
periderm formation. Slight suberization occurred in 1 % O 2 , and increas-
ingly in higher concentrations up to 10% and sometimes 21 %. Periderm
formation did not occur below 3-5% and also increased with higher
concentrations up to 21%. Lange et al. (1970) stated that in cv. Saskia
however some periderm formation occurred even in 1% Oz, and was as
Dormancy, sprout growth and ageing 657
effective in air as in 5% O 2 . Wigginton (1974) found 5, 10 and 15% CO 2 , in
the presence of 21 % O 2 , prevented periderm formation and progressively
reduced suberization at 10°C.
14.4 DORMANCY, SPROUT GROWTH AND AGEING
14.4.1 Dormancy
(a) Dormancy (general)

From a point of view of keeping quality and storage, dormancy in potatoes
is very important. The term 'dormancy' has been used to describe different
states of growth arrest (Emilsson, 1949; Schippers, 1956b; Burton, 1963,
1978a; Goodwin, 1966; Davidson, 1958; Rappaport and Wolf, 1969;
Hemberg, 1970; Taylorson and Hendricks, 1977; Saunders, 1978; Harkett,
1981), and it is important to distinguish clearly the two main reasons for a
dormant seed or bud failing to grow. During the last decade there have
been different opinions about the definition of dormancy in potaotes.
Emilsson (1949) distinguishes two separate states of dormancy:
1. the time after harvesting when tubers will not sprout under favourable
conditions (he defines this as the 'rest period');
2. the time after harvesting when the buds are not growing, for whatever
reason.
Burton (1963) refers to a bud as dormant if it is not growing for any reason,
including an adverse environment, provided the bud has retained the
potential for future growth. This definition is the same, as far as the period
after harvesting is concerned; see (2) above, distinguished by Emilsson
(1949). In his review (Burton, 1978a), a bud is referred to as 'dormant' if it
does not grow at a favourable temperature; the period over which this holds
true is referred to as the 'dormant' period. The latter definition is the same as
that used by Goodwin (1966) and also that used for 'rest period' by Emilsson
(1949) (see (1) above). Burton (1978a) concludes that his first definition
(Burton, 1963) may be too broad, and states that it serves no useful
purpose to restrict the term 'dormancy' to a condition of no change. A con-
dition in which no changes at all take place, does not in fact exist; such a
bud would be dead, not dormant. Burton (1978a) also states that it is not
useful to contrast a 'dormant' bud with a resting bud (Emilsson, 1949).
In 1982 however Burton proposed referring to a bud which is not
growing for any reason as 'dormant', and describing the state of a still
viable bud, which will not grow under optimum environmental conditions
as 'endo-dormant'.
Many authors will agree with the concept of 'dormancy' as described in
the foregoing. It will, however, be necessary to indicate clearly the defini-
tion of dormancy and the precise conditions under which experimental
data are collected. This is to avoid confusion, because most of the
experimental data obtained so far in the past decade deal with the old
concept (Emilsson, 1949; Schippers, 1956b; Burton 1963).
In order to avoid differences in opinion a group of physiologists of the
European Association for Potato Research (EAPR) working on definitions
of terms (Reust, 1974) decided to propose the following:
Dormancy. The physiological state of the tuber in which autonomous
sprout growth will not occur, even when it is placed under ideal conditions
for sprouting (darkness, temperature 15-20oe, relative humidity c. 90%).
Dormancy period. In the literature, the following distinctions were often
made:
Rest period: period during which no measurable extension growth occurs
in any bud on the tuber when stored after harvest under ideal conditions
for sprouting (darkness, temperature IS-20oe, relative humidity c. 90%).
Dormant period: time following the rest period when environmental
conditions can inhibit extension growth in any bud; the dormant period
is defined as ended when 80% of the tubers h,ave sprouts of at least 3 mm
length.
The length of the dormancy period depends on cultivar, maturity of the
tuber and soil and weather conditions during growth and even in the period
just before harvesting if the foliage is still intact. Extreme weather
conditions affect the length of the dormancy period of the tubers (Krijthe,
1962; Burton, 1963). Extremely cold, wet weather increased the dormancy
period by about 4 weeks. Extremely dry, warm weather reduced it by some
9 weeks. Under more normal conditions, however, this effect is relatively
slight (Burton, 1966). Krijthe (1962) considered the tubers no longer
dormant as soon as 90% of the tubers in a sample put at 200 e for a period
of 3 weeks showed sprouts of at least 3 mm. She used 20 tubers per sample.
This procedure was shown to be useful in practice.
Besides cultivar, maturity and weather conditions and whether or not
the tuber is injured, the storage regime is an important factor influencing
the length of the dormant period.
There is also a lack of agreement among potato physiologists as to the
time when measurement of the dormant period should be initiated. Burton
(1957,1961,1963) considered that it was incorrect to measure the dormant
period from the date of harvest, as was usual in published work, in that the
date of harvest had no physiological significance - for example the foliage
at the time could be still functional, or senescent, or could even have been
dead for some weeks. He suggested that the most logical physiological or
biochemical period from which to measure the beginning of dormancy, was
the period during which tuber initiation was occurring: in other words, a
period during which the biochemical balance in the stolon tip was swinging
away from a state permissive of extension growth towards that favouring
lateral expansion, cell enlargement, and ultimately the practical cessation
of cell division. He later developed this theme further (Burton, 1972).
Cho et al. (1983) made comparisons of different methods of measuring
length of the dormant period. The study resulted in the following conclusions.
1. When measured from planting or tuber initiation to sprouting, the
dormant period of tubers harvested on the same date was longer from
an early planted crop than from a late planted crop.
2. When measured from harvest to sprouting, the dormant period was
shorter for tubers from an early compared to a late planting.
3. With the same planting date and different harvest dates, when
measured from planting or tuber initiation to sprouting, the dormant
period was shorter for tubers from an early compared to a late harvest.
4. On the other hand, with the same planting date, when measured from
time of harvest to sprouting, the dormant period was longer for tubers
from an early compared to a late harvest.
Planting date to sprouting was considered the best practical field measure
of dormancy since it closely correlated with tuber initiation to sprouting, a
method which was more accurate but difficult to determine. Both methods
were much better than harvest date to sprouting which is commonly used.
Van der Plas (1987) demonstrated after data from Burton (1978a) the
difference in length of the dormancy period between the 'tuber initiation'
and 'harvest time' method (Fig. 14.16).
(b) Cultivar
Extensive information on the dormant or resting periods of different
varieties after harvest was published by Emilssotl (1949; 51 varieties) and
33
~
30
27
24
-0
21
C/)
..l<:
Q)
18
Q)
15
3:
12
9
6
3
2 4 6 8 10 12 14 16 18 20 22 24
Temperature (0C)
Figure 14.16 Length of the dormancy period calculated from tuber initiation (0)
and from harvest (6) for nine different British cultivars stored at various
temperatures (after Burton, 1978). (From van der Plas, 1987.)
by Schippers (1956a; 41 varieties). Burton (1963) gave durations of the
dormant periods of 11 varieties, measured both from the time of tuber
initiation and from the time of harvest. The average duration after harvest
found by Schippers at lOoC, for tubers harvested immature in mid-July,
was 19.5 weeks and the range 17-27 weeks. Durations after harvest found
by Burton at lOoC for tubers harvested mature in September averaged 8
weeks, range 5-14 weeks, and after tuber initiation averaged 24 weeks,
range 20-33 weeks. There is no evidence in any of the above work that the
duration of the dormant period bears any relation to the maturity class of
the variety. Thomson et al. (1987) succeeded in breeding for long
dormancy. They crossed clones from Solanum berthaultii with commercial
cultivars and got tubers with a dormancy considerably longer than control
cultivars.
There does not appear to be any necessary connection between the
length of the dormant period and the subsequent rate of growth (Burton,
1963) but, like the dormant period, the rate of sprout growth shows
marked, and in general consistent, varietal differences. These are illust-
rated by some of the results of Emilsson (1949), although in considering
these we must remember to compare only cultivars with the same dormant
period after harvest and which therefore commenced growth at the same
time. Burton (1966) quantified the differences between the rates of growth
exhibited by four varieties by comparing the constants in the differentiated
form of the sigmoid growth curves, demonstrating a five-fold difference
between Craig's Defiance and Home Guard at one extreme and Arran
Consul at the other. There is no obligatory relationship between maturity
class and the rate of sprout growth, although in fact the maincrop varieties
which have survived in commerce tend to exhibit a slower rate of growth
than do commercial early varieties.
(c) Effect of storage conditions

Storage temperature
The obvious environmental factor which influences the rate of sprout
growth is temperature. The pattern normally observed is that growth is
very slow up to a temperature of 5°C. Some varieties show no sprout
growth below 5°C; other varieties which show rapid sprout growth under
favourable conditions may show slight growth at temperatures as low as
2°C (Krijthe, 1946). Above 5°C an increase in storage temperature causes
an increase in growth (Barker, 1937; Krijthe, 1946; Burton, 1948).
Barker (1937) stored samples of cv. King Edward 3 weeks after harvest.
Sprout weight expressed as a percentage of tuber weight was measured
after periods of 3 and 6 months. Figure 14.17 shows the difference in
growth at both chronological and hence physiological ages as a function of
storage temperature. This has been confirmed by Sadler (1961).
In Fig. 14.17 a shift in temperature optimum at the 3 and 6 months
storage periods may be observed. McGee et al. (1986) studied the influence
at greater intervals of higher temperatures for 11 European cultivars stored
10,----------------------------------------,
9
.l!l 8
e
:::J
0..
Ul 7
'0 6
.E
OJ
'a:; 5
3:
Q)
OJ 4
co
C
Q) 3
~
a... 2
Q)
O~~.,~~~_,--,__,_,--,__,_,--,__,_,~
o 4 8 12 16 20 24 28
Temperature (0C)
Figure 14.17 Effect of storage temperature on sprout growth of potatoes (cv.

King Edward). (0), sprouting after 3 months; (+), sprouting after 6 months. (Data
from Barker, 1937.)
in darkness. Short and Shotton (1970) measured the length of sprouts at

various temperatures. Krijthe (1962) investigated the influence of storage
temperature on sprout growth with and without desprouting.
In addition to the above direct effects of temperature on sprout growth,
Burton (1966) discussed the possibility that storage at a temperature
unfavourable to growth (say 2°C) may so alter the composition of the tuber
that, on subsequent transfer to a temperature favourable to growth, the
rate of sprout growth is greater than on tubers stored continuously under
favourable conditions. Certainly it may be expected that the rate of growth
can be influenced by previous storage history, whether favourably or not,
and Burton cited the results of Barker (1937) and, more particularly, of
Schippers (1956b) in evidence. The form of the sprouting may also be
influenced. For example storage, after the end of the dormant period at
lOoC, for some weeks at 2°C, with associated changes (including sweeten-
ing), may result in the sprouts which grow on return to lOoC being thicker,
and with much more growth of adventitious roots, than those on tubers
stored the whole time at lOoC.
Wurr and Allen (1976) found that a storage period of 14 days at 2-3°C,
followed by storage at 15.6°C, was favourable for sprouting. The sprouts
developed earlier and sprout length increased. Allen et al. (1978) dis-
covered this effect for a number of cultivars (including Desiree). In other
cultivars, however, this low-temperature treatment increased the number
of sprouts developing per tuber. Some of the cultivars (British and
American) examined by these authors exhibited no reaction to the low-
temperature treatment.
Humidity effect on sprout growth
Davidson (1958) studied the effects of storage conditions on initial
sprouting and subsequent sprout growth. He concluded that compared
with the influence of temperature and light, humidity for all practical
purposes showed only a slight effect. The influence of humidity and light
was judged where high humidity and dark conditions had promoted growth
in the latter part of the storage period. Compared to the influence of
humidity on dormancy-breaking there is only a minor effect on the
subsequent sprout growth.
The effect of light on sprout growth

Sprout growth during storage is inhibited by light. Seed tubers have been
successfully stored in diffuse daylight at higher temperatures (Potts et al.,
1983). McGee et al. (1987) investigated the relationship between wave-
length of light and growth inhibition, using a very long irradiation time of
23 days with a 12-h photoperiod. The inhibition showed a narrow peak of
far-red activity centered on 707 nm, with a shoulder in the red whose size
depended on the degree of inhibition chosen as standard, because the log
photon-fluence rate/response lines were not parallel. There was also
inhibitory activity in the blue «500 nm).
Blue light caused a positive phototropic response at similar or lower
fluence rates. Greening became visible only at the highest fluence rates,
between the two spectral regions inhibitory to growth. Broadband sources
~rJl 80
'E
(5
E
::l..c
;~ 60
._ 0
rJl'-
COl
Q)-
-00
Q)C
()o
c:;:::;
~;Q 40
:;:::-§
C·-
0;:,R
00
-all)
'5 20
Cii
e
()
Q.
'(3
Q)
II:
400 500 600 700
Wavelength (nm)
Figure 14.18 Wavelength relationship for sprout growth inhibition: reciprocal of
photon fluence rates giving 50% inhibition of growth at each wavelength. (From
McGee et al., 1983.)
had much less inhibitory activity in the 650-750 nm region (see Fig. 14.18).
The results confirmed the early photomorphogenetic research of Wassink
et al. (1950). They showed that red light above 650 nm and blue light below
500 nm both contributed to the inhibition of sprout growth.
(d) Break of dormancy

Dormancy release appears to be a gradual process with continuing bud
growth and development during the dormant phase (Emilsson, 1949;
Davidson, 1958). It is apparent that timing of dormancy begins during the
initial stage of tuber formation. Microscopic investigations have shown that
cell enlargement in the bud continues throughout dormancy. In a review
about biochemistry and physiology of dormancy release in potatoes,
Coleman (1987) evaluated published work over the past 20 years. A
complex picture of dormancy release emerges with the assumption of
multiple roles of most major plant growth regulator groups, including the
inhibitor-~-complex, which occur within the tuber. Current literature
continues to support the 'inhibitor/promotor' hypothesis. As already has
been stated by different authors (physiologists) slow changes in the buds
always take place during the period of endodormancy (Burton, 1982; see
also p. 657). Growth or lack of growth are only visible expressions of the
effects of some components of the biochemical balance in the tuber, the
identity and mode of action of which are still to a large degree unknown.
Rappaport and Wolf (1969) concluded that the substances most involved
were gibberellins (promoters) and abscisic acid (inhibitor). Both occur in
potato tubers. (Okazawa, 1959 - gibberellins; Cornforth et ai., 1966;
Milborrow, 1967 - abscisic acid). Of the gibberellic acids, GA3 or mixtures
have been shown to stimulate the break of dormancy of potato buds
(Rappaport et al., 1957; Bruinsma, 1966; Bruinsma and Swart, 1970).
Abscisic acid will inhibit potato bud growth (Blumenthal-Goldschmidt and
Rappaport, 1965; van Es and Hartmans, 1969), gibberellin synthesis
(Wareing et al., 1966) and the promoting effect of gibbereUins (van Es and
Hartmans, 1969). A balance of gibberellins and antigibberellins, such as
abscisic acid, has also been suggested as controlling tuber initiation (Tizio,
1966; EI-Antably et al., 1967; Racca and Tizio, 1968; Paupardin and Tizio,
1970; Holst, 1971; Hammes and Nel, 1975; van der Plas, 1987).
The concentration of ABA is high during dormancy; the concentration
drops 10-100 times during breaking of dormancy and the outgrowth of
sprouts (Fig. 14.19: Bielinska-Czarnecka and Bialek, 1976; Korableva et
al., 1977, 1980). When tubers of various cultivars were removed from 10°C
and placed (in the dark) at 20°C, initial ABA concentrations were
positively correlated with the duration of dormancy and negatively cor-
related with subsequent rates of sprout elongation at 20°C (Coleman and
King, 1984). Other growth inhibiting substances that show a comparable
decrease in concentration upon the end of dormancy are some phenolics
like caffeic acid and scopoletin (Korableva et al., 1977).
'Dormancy'
lr'----<\,
,,
,,
,,
VI VII VIII IX X XI XII

Months
Figure 14.19 Course of the GA and ABA contents during storage of potato tubers
(cv. Pierwiosnek) at 8°C (after Bialek and Bielinska-Czarnecka, 1978; Bielinska-
Czarnecka and Bialek, 1976.) (From van der Plas, 1987.)
14.4.2 Sprout growth
(a) Sprouting control

Sprouting control is very imortant when potato tubers must be stored for a
long period. Good storage practice is in particular important, when tubers
have to be stored for processing into crisps and French fries later in the
season.
Principally sprouting can be controlled by a good temperature regime.
When after the curing period the temperature is lowered to 2-4°C,
followed by storage at constant temperature, sprouting will be kept within
acceptable limits. However, in this temperature range the so-called low
temperature sweetening will develop, leading to a high reducing sugar
content which, depending on cultivar and growth circumstances will in turn
cause a brownish, bitter product when processed.
In this case storage at higher temperatures (7-8°C) combined with a
sprout inhibiting treatment is necessary.
The following general comments have to be made about sprouting
control treatments. Most of the treatments, having sprout-inhibiting effects
- i.e. which inhibit 'growth' - will also suppress suberization and the
formation of wound periderm, thus favouring infection from outside and
consequent rotting. High moisture losses may also occur as a result of
insufficient suberization. The potatoes must therefore be treated only after
the wounds have healed and wound periderm has formed (see Section
14.3.2). Only tecnazene treatments give no delay in wound healing (Dent,
1985). Internal sprout formation, in which sprouts grow inwards into the
tuber can be caused by insufficient sprout suppressant concentrations or too
late application. It is very important that low doses are required and that
very low residues in the tubers are present. Teratogenic effects must be
absent. The effects of secondary metabolites are currently under discussion.
(b) Sprout inhibition
Dimethylnaphthalenes (DMN)
In 1971 Burton and Meigh reported the results of laboratory observations
on potatoes stored in sealed containers in which respiratory CO 2 was
absorbed on solid NaOH and consumed oxygen continuously replenished.
Under these conditions sprouting was suppressed and some 20--30 com-
pounds were found to accumulate in low concentrations in the atmosphere
of the container. Meigh et al. (1973) identified several active chemicals in
the volatile fraction which included the dimethylnaphthalenes (DMN) ,
mainly the 1:4 and the 1:6 isomers. Investigations at Glasgow examined
these chemicals in detail (including DMN) in an attempt to identify which
of them if any would work successfully under commercial conditions
(Beveridge et al., 1981b). The suitability of DMN for this purpose was
confirmed. Duncan and van Es (1988) reported on DMNs and the results
of storage experiments on a practical scale.
The chemical structure of the compound is given in Fig. 14.20. The
different isomers are formed by substitution of two methyl groups at the
different corresponding carbon atoms.
Figure 14.20 Structure of 1, 4 DMN.
Supporting studies on the suitability or otherwise of different analogues

of naphthalene have been carried out, which have shown (1) a lack of
structure specificity i.e. most isomers of DMN are active (Table 14.10),
and (2) others including the ethylnaphthalenes, the isopropylnaphthalenes
(Stephen and Duncan, 1984) and the trimethylnaphthalenes (Filmer and
Rhodes, 1984) are particularly so.
Work at both the Chemistry Department of the University of Glasgow
and the Department of Physiology and Biochemistry of the IBVL at
Wageningen studied and confirmed the sprout suppressing properties
Table 14.10 Composition of DMN mixture and sprout
suppressant efficiency of DMN-isomers
DMN-isomers DMN concentration % inhibition
in synthetic compared to effect of
mixture control (Meigh et ai.,
1973)
1.2 11 70
1.3 33 95
1.4 8 100
1.5 4 90
1.6 12 100
2.3 6 0
2.6 22 0
Unknown 4
From Hartmans and van Es, unpublished.
and showed the commercial potential of the different isomers (Duncan and
van Es, 1988).
For long term storage, reapplication is necessary (see Table 14.11). To
minimize the initial rapid drop in concentration immediately after treat-
ment slow release formulations including specific adsorbents and porous
polymers are being studied. Experiments carried out by Duncan (1984)
indicated that sucrose levels as well as reducing sugar levels remained low
in all treatments including controls and hand desprouted controls.
Wound healing Since the effects of 100 mg kg- 1 DMN applied on a carrier
in large-scale trials were small, it seems probable that smaller doses applied
to stored tubers in the vapour phase as part of a programme of repeated
applications would have little or no effect on wound healing. At least it
would be preferable to allow a short 'curing' period before the first
application as is normal practice with CIPC.
Baking quality When such parameters as baking quality, taste, olfactory

properties, after-cooking blackening and susceptibility to damage were
assessed at IBVL, no deleterious effects were noted compared with
controls.
Propham-chlorpropham (IPC-CIPC)
Isopropyl N-phenylcarbamate ('propham') and isopropyl N-(3-chlorophenyl)
carbamate ('chlorpropham') are normally applied together as a mixture at
a rate of about 10 g of the chemicals per tonne. It is possible to apply them
as a dust (2% active ingredient in an inert filler) but because of adverse
effects upon wound healing (see e.g. Ives, 1955) this must be distributed
by blowing it through the main ventilating system after wound healing is
Table 14.11 Comparison between DMN (Synth. mixt.) and /PC/C/PC on tuber quality. Storage season /985/86
(cv. Bintje) (from Hartmans and van Es, /986)
Storage Sprout Doses (g) of active Application No Difference in taste mg kg- 1
temperature inhibitor susbtance per 1000 sprouting of cooked or fried products residue
eC) kg potatoes until compared to IPc/ levels
CIPC-treated potatoes
7-10 IPc/CIPC 3x 6i Swingfog June (end)
DMN 1x 100 Swingfog May None 1.3
DMN 2x 50 Swingfog June None 1.6
DMN 3x 33 Swingfog June None 1.7
DMN 1 x 100 Powder April None None
5-7 DMN 1x 100 Powder April None None
DMN 1x 100 Swingfog June (end) None 1.1
IPc/CIPC 3 x 6i Swingfog June (end)
Experimental store 'De Eest', Nagele (North-East Polder) - bins of 15 tonne each (IBVL Wageningen).
complete. Such a method does not give adequate distribution in box stores.
In these the chemical can be applied at the time of harvest in the form of
'delayed release granules' from which an inhibitory concentration does not
build up around the tubers until after the wounds have healed. If this
method is adopted it is of course essential that the tubers are stored for the
first 2-3 weeks under conditions favourable for wound healing, otherwise a
damaging concentration can build up before healing is complete.
An alternative method of distribution in box stores is to vaporize the
chemicals at about 200°C (pulse-jet system). Under normal circumstances
temperatures do not rise above the evaporation temperature of the solvent
in which the chemicals have been solved. However as soon as the solvent
fraction has been evaporated temperatures can rise considerably above the
decomposition temperature. The same rise in temperature can occur
when too high resistances in the ventilation ducts prevent the so-called
'Swingfog' equipment from mixing the jet stream with the necessary
volume of cold fresh air. During the treatment the vapour is introduced
into the recirculating system. The store is then closed for 24--48 h with the
circulating fans running continuously, after which there is a fairly uniformly
distributed deposit on the tubers. In bulk stores a similar method may be
used, or application may be in the form of an aerosol of an organic solvent
solution of the chemicals introduced into the main ventilating system with
the fan running. The aerosol is deposited on the tubers and the organic
solvent evaporates leaving a deposit of the sprout suppressant. A dose rate
of 10 g t- 1 is inadequate for prolonged storage, but a higher rate could give
residue problems, and it is usual to make three applications during a long
period of storage.
In several areas of Europe there is a tendency to prolong storage even
until June-July. In those cases repeated treatment is necessary to prevent
sprouting.
Corsini et al. (1979) determined chlorpropham concentrations in peel
samples of tubers taken from large commercial potato stores after aerosol
application. The minimum concentration of CIPC in the peel layer for
complete inhibition of sprouting was estimated to be 20 ppm (at 7.2°C).
Immediately after commercial aerosol application of CIPC, 99% of the
sample sites had concentrations high enough completely to inhibit sprout-
ing, and residues in the peel were generally highest in samples obtained
from the surface and from the bottom of the pile. CIPC concentrations
decreased during the storage season in nearly all sites tested. Storages
differed in the rate at which CIPC was lost. Mean peel residues fell below
the level necessary for sprout inhibition in some storages but not in others.
The rate of CIPC loss increased at the same time that air movement to
maintain the desired temperature increased. Retreatment before peel
residues drop below 20 ppm can extend sprout inhibition in storage.
Residues After treatment a gradual decrease in content of propham and
chlorpropham was observed. The decrease of chlorpropham was slower
than that of propham (Hajslova and Davidek, 1986). The presence of small
amounts of both N-phenylcarbamates was proved even in peeled tubers.
The penetration of chlorpropham into the flesh of the tuber was faster in
comparison with propham. From a point of view of health hazards the
residues of propham and chlorpropham in skin and tuber flesh are very
important. Also residues after cooking and frying become of great interest
to the health authorities. The maximum residue levels allowed in western
European countries, the USA, Canada and Scandinavia vary widely from
o to 50 mg kg- 1 for unpeeled potatoes. Besides the normal residues, today
attention is also focused on the presence of metabolites of propham and
chlorpropham (Heikes, 1985; Worobey and Sun, 1987; Worobey et al.,
1987). The tubers were found to contain isopropyl N-(3-chlor-4-methoxy-
phenyl) carbamate, 3-chloraniline and 3,3'-dichloroazobenzene (3,3'-DCAB)
in very low amounts.
Boyd and Duncan (1986a) determined headspace concentrations of
chlorpropham in a box potato store by adsorption onto a porous polymer
adsorbent followed by thermal desorption. Headspace concentrations of
0.24-1.13 f.lg per 10 I for chlorpropham were found throughout the store.
The same authors investigated the residue of chlorpropham in store
fabric (Boyd and Duncan, 1986b). Residue analyses showed that the fabric
of the store was heavily contaminated with chlorpropham. The floor
contained the highest concentrations varying from 2.05 to 9.47 mg g-l. The
wall samples showed chlorpropham concentrations which were not as high
as in the floor. Although the stored tubers had not been treated with
chlorpropham before the first sampling date, there was a readily detectable
concentration of chlorpropham above the tubers and it was assumed that
its source was the residue chlorpropham in the store. Subsequent treat-
ment of the tubers did not increase the concentrations of headspace
chlorpropham and the concentrations appeared to be influenced more by
the store temperature. Variations in temperature could be an important
factor that might account for variable sprout growth in commercial potato
stores where gradations in temperature have been regularly recorded.
Storage using propham and chlorpropham at high temperatures In

untreated potatoes, increasing storage temperature initially increases
sprouting (Booth and Proctor, 1972; Krijthe, 1977), up to an optimum
around 20°C (Burton, 1966). Above this temperature, sprouting vigour
declines, particularly at 25°C and higher. After 4 months' storage at 28°C,
rot began to appear; this would spread rapidly at this temperature. Hence
the sprout inhibitors CIPC and IPC seem to become ineffective at storage
temperatures above 18°C and if the storage period exceeds 3-3.5 months.
If the produce is mature and its skin has properly set, the sprout inhibitor
can be applied immediately after the harvest, on loading into the store.
With immature, insufficiently set potatoes, a curing period of about 10 days
must be allowed before the sprout inhibitor is administered. If this period
is not observed, the effect of the sprout inhibitor is negative. The sprout
inhibitor must be applied in such a case with a Swingfog unit, and the
method of storage must be appropriate for the Swingfog method. There is
no objection to the application of thiabendazole in powder form im-
mediately when the potatoes are placed in store (Hak and Vermeer, 1979).
A combination of the sprout inhibitors CIPC and IPC with the fungistat
thiabendazole (TBZ) seems to be promising for the purpose of limiting
quantitative and qualitative losses in the storage of potatoes at tempera-
tures of up to about 20°C. At storage temperatures of 25°C and above, the
use of CIPC stimulates sprouting, probably owing to a reduction of apical
dominance (Adlan, 1969).
Internal sprouting The growth of sprouts into the tuber instead of normal
growth was described many years ago (Miiller, 1846; Gager, 1912). It
results from the growth of axillary buds at the base of external sprouts of
which the growing point has been destroyed. Davis (1961) described the
inducing factors as (a) injury and loss of dominance of the apical meristem
without either inhibition of the development of unexposed buds or
complete inhibition of basal portions of the sprout, followed by (b)
circumstances which allow the full force of the expanding sprout to be
exerted on the periderm - by, for example, the pressure of another tuber
against the eye of origin, or by the mechanical components of a basipetally
expanding cluster of buds. Both predisposing factors can follow the use of
inadequate concentrations of sprout inhibitors giving initial suppression
but later a rosette of buds (see e.g. Twiss, 1963). It is however the
rosette following apical damage, not the subsequent presence of the low
concentration of inhibitor, which induces the complaint (Sawyer, 1961;
Sawyer and Dallyn, 1964).
Other sprout inhibitors
Maleic hydrazide (MH) Maleic hydrazide is not a storage treatment,

being applied as a foliar spray in the form of an aqueous solution of the
diethanolamine (sodium or potassium salt), 3-6 weeks before the death or
destruction of the foliage. As a sprout suppressant it is very effective
provided application can be made at the right time, but there is the
possibility of reduction in yield and of mis-shapen tubers, particularly with
early application. Applied too late it is ineffective, and successful use of
maleic hydrazide requires long periods of predictable weather.
Weis et al. (1980) carried out a 3-year study in Central Wisconsin, USA
with timed foliar applications of maleic hydrazide at 3lb acre- 1 (3.36 kg ha- 1).
The treatment resulted in an improvement in quality and a favourable
alteration in the shape of the tubers of cv. Russet Burbank. Applications
were made at intervals from mid-July till early August. Only application
in early or mid-June resulted in severely reduced yield and increased
incidence of mis-shapen tubers.
Struckmeyer et al. (1981) determined the increase in cell size in the
different tissues of the tuber. The cortical and peri medullary cells of the
potato tuber increased in size when treated with the diethanolamine and the
potassium salt of maleic hydrazide. The best time for maleic hydrazide
application appeared to be from mid to late July. There was a greater increase
in cell size in the cortex than the perimedullary region and treatment with the
amine salt was more effective than the potassium salt. The cells were smallest
at the bud end; however, with maleic hydrazide treatment, increase in cell
size was greater in this region compared to the midsection and stem end.
From late August to late September the cell size of untreated tubers
remained relatively unchanged, whereas tubers treated with the amine salt
showed a 16-46% increase with a lesser increase with the potassium salt.
The rounder form of the treated tubers can be explained by the shape and
increase in cell size in the cortical and perimedullary region. Increase in
cell size may also help prevent the formation of mis-shapen tubers. Maleic
hydrazide functioned as an effective sprout inhibitor on tubers from all
treatment dates except when applied in early June.
Ponnampalam and Mondy (1985) showed that foliar application of
maleic hydrazide significantly (p<0.01) increased NOTN content of the
tubers. Seed tubers cannot be taken from a maleic hydrazide-treated crop.
Tecnazene 2,3,5 6-Tetrachloronitrobenzene (Tecnazene, TCNB) has for

many years been used in the UK to control sprouting in potatoes and has
the advantage of having little effect on wound healing (Dalziel and
Duncan, 1980). Tecnazene has sprout-suppressive properties, as well as
fungicidal properties. For this reason its use increased during the early
1970s in the UK (Dent, 1985).
Tecnazene is volatile and active in the vapour phase and has been used
as a dust formulation (Fusarex, 3% or 6%) or as granules (5% or 10%).
Tecnazene will not effectively control sprouting if: dormancy has already
broken or is about to break; the store is excessively ventilated; or storage
temperature is kept too high (above lO°C). The rate of treatment at a
storage temperature of lOoC recommended by Dalziel and Duncan (1980)
was 135 mg a.i. kg-I.
According to Brown and Reavill (1954), free aeration for several weeks,
as occurs if seed tubers are trayed up, removes all trace of tetrachloro-
nitrobenzene from the tubers, and subsequent growth is normal. Seed
tubers may thus be taken from a treated store, provided they are graded
out several weeks before they are required and trayed up or otherwise
exposed to free air circulation. Tetrachloronitrobenzene is the only sprout
suppressant, applied post-harvest, which does not lead to enhanced rotting
by wound pathogens if applied before wounds have healed.
Nonanol 3-5-5-Trimethylhexan-l-01 (nonanol) is a liquid and is intro-
duced as a vapour, at a concentration of 0.1 mg 1-1 of air, into an airstream
blown through the main ventilating system of bulk stores at a rate of 10 m3
C1 h- 1. In box stores the vapour is introduced into the recirculating system.
The simplest method of introducing the vapour into the air is to blow the
latter over towelling dipping into a container of the liquid, the area of
towelling being adjusted to give roughly the correct concentration. This is
determined with sufficient accuracy by measuring the quantity of nonanol
required to replenish the container, the rate of ventilating being known.
Applicators incorporating a drip feed on to a hot plate have been used, but
under commercial conditions have suffered from considerable variation in
rate of feed. Nonanol, used as above, gives complete sprout suppression,
but it is not persistent and growth can occur within 2-3 weeks of stopping
the ventilation. The consumption of the chemical can be halved by
alternating 2-week periods of nonanol application with 2-week periods of
convection or ordinary ventilation, consumption over a 2-week period thus
being about 35 kg per 100 t. Nonanol provides the cheapest method of
sprout suppression but involves continuous supervision over the 2-week
period and is now little used.
Drauschke et al (1981) carried out experiments with 14C-Iabelled
nonanol to establish the penetration and residues in storage experiments at
different temperatures. Autoradiographic assays showed the following
results: a 48 hours experiment resulted in a homogeneous distribution over
the total surface area; the highest concentration of nonanol occurred in the
skin; a steady increase in radioactivity in the control showed evaporation
from the treated tuber surfaces; calculations based on autoradiography of
the penetrated 14C in the different layers were in good agreement.
Camptothecin Wall et al. (1966) isolated an alkaloid from Camptotheca

acuminata (Nyssaceae). The alkaloid was intensively studied as an anti-
tumour reagent. The effect of the plant growth regulating component was
tested on several species of mono- and dicotyledonous plants. Selective
growth inhibition was found. Wang et al. (1980) reported the action as a
growth inhibitor on potato sprouts. When camptothecia was applied as
lanoline suspensions at concentrations of 1.5, 3, 9 and 15 mM or to the
whole tubers as aqueous sprays or dips, potato sprouting at 15°C was
completely inhibited for 4 weeks. Inhibition was greater than 95% when a
solution of 5 X 10-1 mM was applied to the tubers as a spray or dip (three
times). The alkaloid treatment inhibited sprouting, reduced weight loss
and showed no symptoms of internal sprouting.
Alternative sprout inhibitors There are several essential plant oils which
are now applied in the food and cosmetics industry as fragrancing and
flavouring substances, which have promising sprout inhibiting properties
(Beveridge et al., 1981a).
Small-scale experiments have shown that the pulegone contammg
essential oils of Minthostachys glabrescens and Mentha pulegium, the
carvone containing essential oils of Anethum graveolens and Carum carvi
and the cuminaldehyde containing essential oils of Cuminum cyminum are
efficient sprout inhibitors. Apart from these three substances, perillaldehyde
and menthylacetate also have sprout inhibiting properties. From the results
of experiments on a small scale and semi-practical scale, it appears that a
number of these essential oils are efficient sprout inhibitors. Apart from an
optimum sprout inhibiting effect in prolonged storage the influence of
these substances on the quality of the potatoes is important. Therefore, a
number of quality aspects have been investigated, i.e. the effect on the
cooking and frying quality, the residue content, wound healing and attack
by fungi (Hartmans and van Es, 1988).
Evaluation of the cooking and frying quality after treatment did not give
any problems in potatoes treated with caraway oil or cumin oil. Potatoes
treated with pennyroyal oil have a deviant taste and the sprout inhibiting
effect of this compound was less satisfactory. The residue contents were
low when the bins were unloaded « 0.5 ppm).
The doses of essential oils applied were rather high: due to their
volatility a large part of the substances escapes from the bins rapidly after
application. This may probably be avoided by application with a slow-
release agent.
The preliminary results of experiments with caraway oil show that at
high doses a slight inhibiting effect on wound healing occurs, so that in this
case it seems advisable not to apply this compound to the potatoes directly
when loading the store.
Some essential oils have a fungistatic effect. Preliminary experiments
with caraway oil have shown that Fusarium sulphureum and silver scurf can
be suppressed.
An advantage of these oils is that no toxicological investigation is
necessary for application as a sprout inhibitor on potatoes. The dis-
advantages are: great volatility, which can possibly be solved with slow-
release dosing; possible odour and taste deviations, although application of
low dose rates, combined with strong ventilation before using the potatoes,
can avoid these problems; and as yet these oils are not commercially
available at acceptable costs.
Radioactive irradiation
Sprout growth may be prevented for a considerable period, in some cases
indefinitely, by irradiating the tubers with y-rays at doses of 50 to 200 Gy or
even lower (Brownell et al., 1954; Hagberg and Nylsom, 1954; Hannan,
1954; Sparrow and Christensen, 1954; Burton and Hannan, 1957; Patzold
and Kolb, 1957; Patzold and Weiss, 1957; Sparenberg and Vlmann, 1973).
Gamma irradiation with a cobalt source (60CO) has been found to be
particularly effective. A number of commercial-scale installations have
been in operation in various countries including Japan, Canada and The
Netherlands. Besides prevention of sprout growth it has however other
effects: there is marked temporary sweetening, and senescent sweetening
is hastened, so that very prolonged storage of irradiated tubers is not
possible (Burton et ai., 1959). For processing use the content of reducing
sugars may be too high after 3 months or so, but for domestic consumption
an acceptable sugar content could be retained for perhaps 6 months or
longer (Burton et ai., 1959). With irradiation at say 10 krad, we can thus
obtain complete sprout suppression for medium-term storage. The method
involves the provision of a facility for exposing a shallow layer of the tubers
to a fully shielded radiation source, usually cobalt-60, for a time sufficient
for adequate dosage. The provision of such a facility could scarcely be
envisaged on the farm. If it is provided at a central storage depot it could
not irradiate the whole tonnage of the catchment area at the time of
harvest unless the weather were so uniformly good that a steady intake was
possible over a fairly long predetermined period. In other cases double
handling of a considerable tonnage might be necessary.
Besides those disadvantages other objections are as follows. A grey
discoloration occurs when the potatoes are cooked although this has not be
observed in different laboratories to the same extent. It is assumed that
irradiated potatoes do not constitute a health hazard, but it has not been
established that irradiation does not give rise to the formation in the tuber
of compounds which may be harmful to health. The biggest disadvantage
however is the high cost of irradiation.
In the 1974-75 season a commercial irradiation plant was established in
Shihoro, Japan. It can irradiate 15-,17 t h- 1 and treat about 30000 t over a 3
month period of continuous operation. It employs a 300 kCi cobalt-60
source to give a mean dose of 10 krad. The capital cost of this facility to
irradiate 15-17 tonne h- 1 was $1.5 million US (then £0.7 million). Part of
this can be offset against alternative uses in the remaining 9 months of the
year - irradiation of onions, packed meals, etc. - and allowing for this the
cost of treating the potatoes has been estimated at about 10-11 SUS t- 1
(then £4.5-5).
(c) Sprout stimulation

Sprouting can be stimulated by chemicals, by damaging the tuber, by
increasing the humidity of the storage atmosphere and by temperature
variation. Although this subject is not directly relevant to potato storage, it
seems appropriate in the context of potato physiology to refer briefly to
sprout stimulation by means of chemical compounds and wounding of the
tuber tissue. The main chemical agents used in practice are Rindite, carbon
disulphide and gibberellic acid and to a lesser extent thiourea.
Rindite
The substance known by this brand name is a mixture of ethylene
chlorhydrin, ethylene dichloride and carbon tetrachloride (7:3:1, respec-
tively, by volume); the mixture was introduced by Denny (1945). The
method of application is typically as described by Keller and Berces (1966)
for tubers from 4 weeks after haulm killing onwards. A container, which is
capable of being tightly closed, is filled with potatoes surrounding a tube,
made of wire netting, running from bottom to top, which contains a coil or
strips of filter paper. The Rindite is poured on the filter paper, to the
amount of 10 ml per 10 I of empty container volume, and the container
closed and left at 24°C for 48 h. The tubers are then trayed up and kept
under diffuse lighting at 24°C to sprout.
Alternatives to Rindite have not so far proved as effective, but an
alternative is desirable because of the toxicity of Rindite vapour. Because
of the toxicity of the vapour strict safety precautions are necessary. Sikka
and Bryan (1980) reported an application method based on a dip treatment
of a range of genotypes, particularly Solanum andigenum, characterized by
long dormancy.
All concentrations of ethylene chlorhydrin tested accelerated sprouting;
3-5 ml of chemical to a litre of water were found to be optimum.
Compared with vapour treatment with ethylene chlorhydrin or Rindite,
sprouting response was not only faster but also there was uniformity in
sprouting behaviour unlike the erratic sprouting met with vapour treat-
ment. Significant rotting of tubers was observed with vapour treatment.
This was similar in single treatment (dip or vapour treatment) and triple
treatment consisting of dip or vapour treatment of ethylene chlorhydrin or
Rindite followed by treatment of GA: hence no additive effect of combined
treatment was discernible. There were, however, clear indications that
GA enhanced sprout growth significantly, the mean sprout length being
0.82 cm in the combined treatment and 0.30 cm in the single treatment.
Contrary to the use of Rindite, treatment with ethylene at low concentra-
tions, 1 III 1- 1 , at 20°C showed a decline in sprout growth (Timm et al., 1986).
Carbon disulphide (CS2 )

The use of sulphur compounds such as mercaptoethanol, thiourea and
CS 2 for dormancy breaking has long been familiar (Denny, 1926) and is
common, for example, in India and Brazil. Under Dutch conditions, CS 2
was tested by Meijers (1972). Treatment for about 3 days at 20°C using
12.5-25 ml CS 2 per m3 containing an average of 500 kg of potatoes is
sufficient to obtain optimum sprouting. CS 2 , however, also has the
disadvantage of being toxic.
Gibberellic acid (GA)

Immersion of tubers in a solution of GA (van Hiele, 1961) or spraying
them with this solution (Holmes et al., 1970; Marinus and Bodlaender,
1978) stimulates sprout growth and increases the number of stems per plant
and the number of smaller tubers (seed potatoes). The total yield is
unaffected by treatment of the tubers with GA (Holmes et aI., 1970).
Excessive concentrations of GA (over 25 ppm) can cause defects in the
plant, such as thin stems and small, rapidly yellowing leaves (Timm et al.,
1962).
GA is usually absorbed more efficiently by the tubers after wounding.
For this reason, application directly after harvesting or cutting of seed
potatoes is the most effective. The 'wound effect' caused by damage and
cutting of the tissue is well known. This gives rise, for example, to the
synthesis of gibberellins (Rappaport and Sachs, 1967), thus stimulating
sprout growth.
(d) Consequences of sprout growth

With regard to storage technology the effect of sprout growth is two-fold.
It is accompanied by increased respiration (see Section 14.2.3) and
consequent heat output; and the sprouts growing in the spaces between the
tubers increase the resistance of the stack to airflow. Sprout growth to an
average extent greater than 5% by weight is rarely encountered in practice.
Tubers sprouted to this considerable extent could be respiring at 4-5 times
the rate of un sprouted tubers, which means that the metabolic heat
production Q m in the equation on p. 615 could be five times greater,
and {Q m - C (heat loss by conduction) - E (heat loss by evaporation)}
possibly as much as eight times greater. Although E is potentially much
greater in sprouted than in unsprouted potatoes (Section 14.2.1), it is,
in unventilated storage, limited by the water-holding capacity of the
convective air, and thus by the rate of convection, and could be perhaps
doubled in the case under consideration. The difference in average
temperature of the potatoes and the average temperature of the ambient
air (Tp - T af· 8 could thus be increased by a factor of at least 51.8 and
possibly 81.8 and hence Tp - Ta by a factor 2.~.2. In addition there is the
effect upon resistance to flow, i.e. upon K, which could be increased by a
factor of 4, as we have seen earlier (admittedly in the presence of
considerable earth). This would increase Tp - Ta by a further factor of 1.6.
The overall effect could well be a five-fold increase in Tp - Ta. The
average temperature of a 3 m high stack, instead of being 3.5°C above
ambient would tend to be 17-18°C above. Again, as in the case of inclusion
of earth, this figure is not precise, but it gives an idea of the potentially
serious effect of considerable sprout growth in unventilated stacks.
14.4.3 Physiological ageing

At any given time from initial dormancy until incubation, the stage of
development of a seed tuber can be described as its physiological age
Changes in composition during storage 677
(Kawakami, 1952), a condition that is modified by, for example, increasing
chronological age. The significance of physiological age as a determinant of
crop yield was first recognized by Kawakami who found that the rapidity of
sprout growth increased with physiological age.
A seed tuber passes through a number of a developmental stages after its
formation. First, there is a period of dormancy from the time of initiation
until some months thereafter, during which, even if placed under condi-
tions favourable for sprouting, a tuber will not sprout except as a result of
the occurrence of second growth. Second, after the dormant period there
will be sprout growth (Fig. 14.21). Initially, usually only one sprout
develops; later there is a period of multiple sprout development after
which sprouts tend to branch and small tubers may develop on them.
Max. sprout capacity
,Physiological
old'
Max. dormancy
Days
Figure 14.21 Schematic representation of sprouting capacity vs time.
The effects of sprout development at the time of planting on subsequent

growth and yield are dealt with in detail in Chapter 6.
14.5 CHANGES IN COMPOSITION DURING STORAGE
14.5.1 Chemical composition and changes during storage

The chemical composition of the tuber at harvest is very important. It
determines keepability during storage and is, apart from the cell structure
of the tissue, the main factor determining cooking quality and quality for
processing into crisps, French fries and granules. The potato still occupies
an important position in the daily diet of large sections of the population in
many countries. It is important to know the composition of this diet in
terms of nutritional value.
The potato tuber contains very many potentially reactive systems of
2.4
2.2
A
2.0
j 1.8
.<:
rJl
1.6
~ 1.4
~ 1.2
C
2 1.0
c
8 0.8
~
::l
0.6
(/) 0.4
0.2
o ~--,---,---.---,---,---.---.---.---,----
Dec. '69, Jan. Feb: March Apr. May June July Aug. '70
06
1
~:1 ~~-,-==
Dec. '69 Jan. Feb. March Apr. May June July
I
Aug. '70
Figure 14.22 Changes in sugar content during storage. Storage conditions, lOoC
and 85-90% relative humidity. /:,. = sucrose; • = glucose and 0 = fructose
content in % of fresh weight. (From Rumpf, 1972.)
substrates and enzymes - in fact the presence of complex molecules which

have been elaborated in the tuber during growth suggests a potential
for continued reactions, and for reverse reactions. When a mature tuber
is harvested its content of the major known constituents has reached a
fairly steady level. Change is still occurring - we know that there is a
perpetual cycling of some constituents and may suppose that there is
cycling of others - but the production and breakdown of many of them are
nearly in balance: not quite, as some change can be detected, and of course
there is a perpetual loss of carbon from the system in the CO 2 evolved in
respiration.
Every reaction is influenced by its immediate physical and chemical
environment. The latter is affected by the products and demands of every
neighbouring reaction; which reactions are themselves subject to similar
influences. The whole tuber complex is thus very labile, and if the storage
conditions, or the stimuli to which the tuber is exposed, are very differ-
ent from those under which it has been growing, then the state of near-
balance is lost, and readjustments, which may be considerable, occur. If
the fresh stimuli include mechanical damage or infection by pathogens,
then the complexity of the response can be much increased and may
involve reactions, following membrane disruption or the introduction of
different reactants or enzymes, for which there is normally little or no
opportuni ty .
The complete response of the tuber to the time and conditions of
storage, and to other stimuli, as shown by changes in composition, is
unknown. We do, however, have some knowledge of a number of changes,
mainly those in the content of constituents of known or suggested
importance to the quality or dietary value of the tuber. For example much
research has been carried out in the field of dry matter content, metabolism
of reducing and non-reducing carbohydrates, proteins and other
nitrogenous compounds, although much work has still to be done.
14.5.2 Dry matter content
(a) Chemical composition of the dry matter

The composition of the dry matter in potatoes can vary substantially
according to cultivar, conditions during growth - e.g. type of soil, fertilizer
application, temperature, moisture supply and light - and degree of
maturity. Changes may occur during storage, as Fig. 14.22 shows for sugars
and Fig. 14.23 for organic acids (Rumpf, 1972). Levels and ranges for
a number of components are given in Table 14.12. See Table 14.13
(Augustin, 1975) for values of vitamins.
~0.6
-£i 0.5
!!?
- 0.4
~
:; 0.3
"C
.~ 0.2
.~ 0.1
«l
~ 0 L-~r-~--~--~--~--'r---~--~--r---
o Dec. '69 Jan. Feb. March Apr. May June July Aug. 70
Figure 14.23 Citric acid (e) and malic acid (0) content during storage. Storage
conditions: lOoe and 85-90% relative humidity. (From Rumpf, 1972.)
As with the dry matter content, the chemical compounds are not
distributed homogeneously over the tuber. Here again, there are dear
differences in concentration for a number of components. For instance,
chi orogenic acid is found in the outermost part of the tuber, and so is
solanine (see Tables 14.14 and 14.15). Solanine is concentrated mainly
around the eyes, while, for instance, the enzyme phenolase is distri-
buted throughout the tuber. Fig. 14.24 shows the distribution of the
components over the tuber in the radial direction relative to the apex-stem
axis.
Table 14.12 Approximate composition of the dry matter of the potato tuber
Chemical composition From Painter and From Burton (1966)
Augustin (1976)
Range (approx.) Normal value
(approx.)
Dry matter (%) 22.48
Alcohol insoluble solids (%) 92.65
Crude fibre (%) 1.08 1-10 2-4
Starch (%) 74.24 60-80 70
Reducing sugars (%) 0.55 0.25-3* 0.5-2*
Total sugars (%) 1.28
Total nitrogen (%) 1.16 1-2 1-2
Amino nitrogen (%) 4.09
Ascorbic acid (mg per 100 g) 92.08
Thiamin (mg per 100 g) 0.73
Niacin (mg per 100 g) 10.08
Riboflavin (mg per 100 g) 0.118
Ash (%) 4.20 4-6
Calcium (%) 0.019
Magnesium (%) 0.084
Potassium (%) 1.47
Sodium (%) 0.022
Iron (ppm) 15.70
Copper (ppm) 1.30
Citric acid (%) 0.5-7 2
Protein-N (%) 0.5-1 0.5-1
Fat (%) 0.1-1 0.3-().5
Sucrose (%) 0.25-1.5* 0.5-1.0*
All data except for dry matter are recorded on a dry weight basis. The dry matter m>rmally
comprises about 18-28% of the fresh weight, dependent upon factors discussed in the text.
Average value about 23%.
• These figures are representative of mature unstored tubers. The content of sugar is vexy
markedly affected by the stage of maturity and by temperature of storage.
Table 14.13 Effect of nitrogen fertilization on the contents of some water soluble vitamins in Russet
Burbank potatoes (in mg per 100 g dry matter) (from Augustin, 1975)
Nitrogen Ascorbic acid Thiamine Riboflav,m Niacin

(lb acre .1)
Mean CV(%) Mean CV(%) Mean CV(%) Mean CV(%)
Loamy soil 0 125.50 1.32 0.299 25.91 0.080 10.45 8.09 3.2
300 106.50 6.06 0.579 15.45 0.090 6.08 8.22 6.9
600 100.29 4.73 0.572 13.54 0.097 12.20 9.10 6.6
Sandy soil 0 150.50 1.25 0.252 30.70 0.102 9.80 8.22 7.9
300 137.82 5.76 0.360 24.84 0.103 13.73 8.48 6.4
600 127.81 4.70 0.457 6.91 0.097 6.52 9.19 6.3
Starch Protein Crude fibre Fat
(
0"',
9.0-14.0-25.0% 1.2-2.0-2.5% Bud end 0.4-0.7-1.0% 0.05-0.1-0.2%
o o
Sugars Organic acids
.:1-,;,-..:.;"
0.1-0.9-5.0% 1.0-2.0-3.0%
Stem end
® 0
Vitamins Minerals Phenol compounds Alkaloids
0.01-0.03-0.05% 0.3-1.0-1.2% 0.05-0.1-0.4% 0.003-0.006-0.011%

Figure 14.24 Chemical composition (% of fresh wt) and distribution within the
tuber. (From van Loon and Muller, 1984.)
Table 14.14 Comparative summary of relative concentrations of several

minor components within specific tuber tissues* (Reeve et al., 1969)
Tissue ChIorogenic Tyrosine Iron Solanin-
acid solanidine
Skin
Periderm
Phellogen + +
Phellem + ++
Cortex
Parenchyma ++ + + ++
Outer phloem ++ + + +
Phloem parenchyma ++ + + +
Vascular 'ring'
Xylem ++
Xylem parenchyma + ++ ++ +
Perimedullary zone
Storage parenchyma + ++ +
Internal phloem ++ ++ ++
Phloem parenchyma ++ ++ ++
Pith branches + ++ +
Pith
Pith parenchyma + + +
* Based on a general evaluation of analytical reports and histochemical evidence.
+= present, ++ = greater concentration. Lack of symbol does not indicate absence.
Table 14.15 Summary of bud end to stem end gradients of
minor components within the tuber (Reeve et aI., 1969)
Component Tuber zone*
'Eye' areas Bud end Middle Stem end

ChIorogenic acid + + ++ ++
Tyrosine + ++ +++
Ascorbic acid + + ++ +++
Citric acid +++ ++ +
Acidity + ++ +++
Iron ++ + ++ +++
Phosphorus +++ ++ +
Potassium +++ ++ +
Solanin-solanidine +++ ++ + +
* + = present, ++ = concentrated, +++ = more concentrated.
Lack of symbol does not indicate absence.
(b) Percentage of dry matter

The dry matter (DM) content is a very important factor in potatoes. It is
mainly determined genetically and thus depends on cultivar. Since 60-80%
of the DM consists of starch, there is a major correlation between DM
content and the starch content of the tuber. The DM content is important
for the quality of both boiled and fried potato products such as chips and
French fries. It is of direct importance in chips and French fries, as the
weight of the processed product depends directly on the amount of DM
present per quantitative unit of fresh potatoes. The same applies to dried
potato products. The DM content is also important with boiled potatoes,
as it may be a measure of the mealiness of the boiled products (Warren and
Woodman, 1974) and of mushiness on cooking (Talburt and Smith, 1967).
(The canning industry demands potatoes with a low DM content.) There is
also a relation between the dry content and susceptibility to black spot
caused by impact (Jacob, 1959). The DM content is influenced by a large
number of factors (Fig. 14.25), chief among which are: cultivar, maturity,
growth pattern as influenced by nitrogen fertilizer application, and climate,
soil and potassium fertilizer application. These factors have been reviewed
by Beukema and van der Zaag (1979).
Relationship between underwater weight, dry matter content, starch content

and specific gravity
It may be necessary to determine the DM content of batches of potatoes
not only for the marketing and processing of potatoes but also when they
are stored. Within limits this can be done very simply by weighing under
water. The underwater weight (UWW) is the weight under water in grams
Maturity of tubers I Variety

Date of harvest
Growth type
Variety
Physiological age of tuber
and sprout
Day length
Content dry matter in tuber Growth type Temperature
Light intensity
Water supply
Soil condition
Nsupply
Variety
Soil condition
Weather condition
Water and mineral N supply
uptake by the crop K supply
Cl supply
P2 0 5 supply
Figure 14.25 Simplified version of a survey of factors influencing the percentage

of dry matter in tubers. (From Beukema and van der Zaag, 1979.)
of 5000 g of potatoes. The specific gravity can be obtained directly from

this:
5000
Specific gravity (SG)
5000 - UWW
The relation between specific gravity (SG) and percentage DM (% DM) is
less simple; it has been determined empirically by a number of workers,
and varies to some extent as between different batches of potatoes. A very
well known relationship is that established by von Scheele et at. (1935):
%DM = 24.182 + (211.04 x SG) - (211.04 x 1.0988)
If the dry matter content is expressed in UWW then:
%DM = 0.0493 x UWW + 1.95

Other authors have given somewhat different regression lines (Porter et
at., 1964; Houghland, 1966; Carlsson, 1967; Verma et at., 1971; Ludwig,
1972; Schippers, 1976). One of the reasons is the difference in intercellular
space in the tuber tissue (Burton, 1966; Kushman and Haynes, 1971). The
temperatures of the potatoes and of the water in which they are weighed
are also relevant (Adler, 1971; Basker, 1975).
The relationship between underwater weight, DM content and starch
content for a large number of batch,es of potatoes used for starch, from
sandy and peat soils in the north of The Netherlands was found to be:
% DM = 0.05 x UWW + 1.0
The DM content of tubers is also an important measure of quality used
extensively by processors to assess suitability for the production of French
fries, chips and dehydrated products; this is because the yield of product is
greater per unit fresh weight from tuber with a high content of solids. To
avoid errors arising during conversion of underwater weight to dry matter
and starch content of tubers, Putz (1983) suggested using the underwater
weight expressed in grams as a basis for calculating payment according to
the quality of potatoes.
(c) Loss of dry matter during storage

Taking starch to be the primary respiratory substrate and assuming an
average evolution of CO 2 during storage of rather more than 3 ml kg- J h- 1
(see Section 14.2.3), then in 1 month a tuber will have lost about 0.3% of
its weight in the form of dry matter and gained about 0.15% of its weight in
water produced during respiration. If respiration were the only source of
net weight loss, a tuber with an initial percentage dry matter of 22.0%
would, after 1 month, have a dry matter of 21.7% and after 6 months,
20.4%. Water loss is also occurring during storage however (Section
14.2.2) and the percentage of dry matter need not therefore decrease,
although it does unless the potatoes are stored under conditions such that
water loss is excessive. Under good storage conditions a drop of about
0.1 % a month might be expected.
14.5.3 Carbohydrates
(a) General
Potatoes are fried to produce crisps and French fries by immersing them in
cooking oil. After the frying process the surface of the crisps and French
fries should be at the desired colour according to local preference. In
general a yellowish slight brown colour is preferred.
It is widely accepted that the Maillard reaction between the reducing
sugars and amino acids plays a major role in the process of the brown
colour development and decrease of nutritional value. Research has shown
that 2.5-3 mg reducing sugars per gram fresh weight must be regarded as
the maximum permissible level for crisping. For french fries the limit is
about 5 mg per gram fresh weight. Sometimes a high reducing sugar
content can be found already immediately after harvest and curing period.
In many cases however it develops during the storage period. The most
important reducing carbohydrates are glucose and fructose. Sucrose does
not contribute to the Maillard reaction but can increase to an undesirable
level giving a sweet taste of the products. Arbitrarily divided into major
and minor effects, a number of factors affecting reducing sugar content are
given below.
MAJOR EFFECTS MINOR EFFECTS
Temperature prior to cold storage Soil composition
Storage temperature Fertilization
Reconditioning Environment
Cultivar Water supply
Maturity
Storage conditions
It is well known that the formation of reducing sugars is strongly affected

by the storage temperature. Generally it can be stated that for a given
cultivar, storage at higher temperatures (7-100C) causes a lower reducing
sugar accumulation than storage at lower temperatures (2-4°C).
However, storage at the higher temperatures causes considerable
sprouting. Sprout prevention at those temperatures can only be achieved
by artificial sprout inhibitors. The most widely used inhibitors in this field
are still propham and chloropropham (IPC and CIPe). From a point of
view of research three different mechanisms, that take place simultaneously,
have to be distinguished: sugar accumulation at low temperature, sugar
formation accompanying the sprouting process and sugar formation due to
senescent sweetening.
(b) Influence of temperature

Storage at low temperatures - low temperature sweetening
The marked increase in sugars, particularly in the reducing sugars glucose
and fructose when stored at temperatures below 6°C, was recorded
and studied as long ago as 1882 by Miiller-Thurgau. Since then 'low
temperature sweetening' has been studied throughout the years by many
research workers. Many data and details are already known, although the
underlying mechanism has not yet been clarified.
In short-term storage experiments with mature tubers (cv. Majestic)
Burton (1965) plotted sugar formation against various temperatures, as
shown in Fig. 14.26. This figure clearly shows that below lOOC both
reducing sugars (glucose and fructose) and non-reducing sugar (sucrose)
are formed to an increasing degree. van Es and Hartmans (1986) showed
the varietal difference in behaviour during prolonged storage. Bintje
showed an increase while Saturna showed a decrease at 2°C (Fig. 14.27).
Work by van Es and Hartmans shows that potatoes transferred from 2°C
to 8°C show a marked decrease in reducing sugar content while the same
potatoes transferred from 8°C to 2°C showed a marked increase. The
a ~--------------------------------
a 4 8 12 16
Temperature (Oe)
Figure 14.26 Low temperature sweetening: sugar contents of potato tube.rs (cv.
Majestic) after storage for 4 weeks (17 December-14 January) at different
temperatures. (From Burton, 1965.) e, total sugar; D, glucose and fructose; .. ,
sucrose.
experiments were carried out with cvs. Bintje and Saturna (Fig. 14.28). In
fact the transfer from 2°C to 8°C followed by a decrease in reducing sugar
content provides an example of reconditioning (see below).
Storage at higher temperatures

Recently, more emphasis has been placed on storage at high temperature.
This is often necessary in the warmer developing countries where
mechanically cooled stores are not usually available. Verma et al. (1974a,
1974b) found that potaotes can be stored just as well, as regards sugar
content, at temperatures ranging from 24.7 to 36.2°C. Total sugar content
increases at these temperatures just as in the case of cold storage, but the
increase is smaller. On transfer to higher temperatures, the total sugar
content initially increased sharply, but then fell back to virtually the
original level. Just as with cold storage, the reducing sugar content also
increased, although only slightly at first, compared with cold storage.
Towards the end of the period of cold storage, however, the reducing
sugar content increased to 1.2%. Loss of dry matter and also rotting
increased sharply at high storage temperatures. Hak and Vermeer (1979)
found that even a product which was not fully mature and sufficiently
2.0
1.8
j 1.6
J::.
CJ)
~
-1.4
~
OJ
~ 1.2
ti
:::J
~ 1.0
c
til
~ 0.8 _---~ Saturna, sucrose

Cl
:::J
CJ) I
r "- , .......
g> 0.6 / .... -*_ 'X- ---x
·0 /,// ...........
/ x ...... x
ascr: 0.4
:::J
I \ / / " Bintje sucro;e---x
r' \ / /
\, '-x-'''/
0.2 'x_-x"
42 46 50: 2 6 10 14 18 22
1985-7:<- 1986 Week no.
Figure 14.27 Contents of reducing sugars and sucrose during storage at 2°C of cvs
Bintje and Satuma. (From van Es and Hartmans, 1986.)
hardened could be stored at up to 20°C for about 4 months if CIPC was

applied in combination with the fungcide thiabendazole at the right time -
i.e. after suberization and wound healing. The quantitative and qualitative
losses could then be limited.
Linnemann et al. (1985) investigated the difference in 'storage
behaviour' of cv. Bintje stored at 7, 16 and 28°C. The tubers were treated
with chlorpropham as a sprout inhibitor and stored for three months, with
the following results:
Sprouting. After 12 weeks' storage at 28°C the tubers had become soft and
showed 2-4 cm long, sturdy sprouts; sprouting started 2 weeks after the
beginning of storage. Tubers stored at 16°C were less soft and showed only
a few small sprouts of 0.5 cm length or less. The tubers stored at 7°C had no
sprouts at all and were still firm. The dry matter content of the tubers was
21.0, 21.7 and 24.1 % respectively.
Reducing sugars. Fig. 14.29 shows that during storage at 7 and 28°C the
glucose and fructose content decreased steadily. The potato tubers stored
at 16°C showed an initial decrease, followed by an increase.
Sucrose. At the storage temperature of 7°C the sucrose content in the
tubers remained constant. At 16°C the sucrose content showed a very slight
2.0 2.0
1.8 1.8
~ 0, Sir,t,.
.c 1.6 ~r;eT 1.6
-
<J)
'8ir,f <"'03
~ 1.4 ... 7e 1?
-;(?
' , , ...... 1.4
~ ...... , ' •......Cit.
<J) 1.2 ,00Cit.'(;/"I7Ci 1.2
<a "'~/"I,>', <0
OJ -..:.Ci l: ........... ~< 1 0
::::l
<J)
1.0 '~-::o. . o .
"-(f
OJ
"''0 ~'?>
c 0.8 0.8 \\
'u '0
::::l
"0
0.6 0.6 <?p"~
<D
a:
0.4 0.4 Bintie 16-12
""'
"' .....
.....
0.2 0.2 t
".,. """ <*'
..... ~..- Saturna 19/12 and 13/03

0 1 2 3 0 1 2 3
(a) Weeks (b) Weeks
Figure 14.28 Effect of transfer from (a) 2°e to 8°e and (b) 8°e to 2°e on the
reducing sugar content of cvs Bintje and Saturna, after two different periods of
storage. (From van Es and Hartmans, 1986.)
decrease followed by an increase. Storage at 28°C, however, showed a

sharp increase in sucrose content, probably because of the sprouting of the
tubers (final total sugar content 0.85 g per 100 g fresh weight). Total sugar
contents below 1.25 g per 100 g fresh weight do not cause a sweet taste,
when consumed (Ludwig, 1970).
Reconditioning
In potato reconditioning, the temperature is increased to 1(}....20o e in an
attempt to decrease the reducing sugar content, which has become too high
as a result of cold storage. During this process, a high proportion (80%) of
the reducing sugars is converted into starch; the remaining 20% is lost in
respiration. For several reasons, however, the sugars present are in-
completely eliminated by reconditioning. Burton found in 1975 that
potatoes which had first been stored cold for 4 weeks and then kept for 90
days at 10°C had a higher sugar content than potatoes stored constantly at
10°C.
Isherwood (1976) and Iritani and Weller (1978) showed that in general
only those sugars formed by cold storage could be reconditioned. Sugars
arising from senescence of the potato - i.e. later in the storage season -
could not be reconditioned. It was found that the factors involved in the
development of a high sugar content are also partly responsible for
.E 0.25l
OJ·f 0.20~
Cfl.s:: 0-••
.8
g.!=. rc0 - - 0
~ 0 15 " ~'0.,. ~-o ... --"o ___
Ltc) ,
g 0.10 'K,.. 16°C//
,.... ""X.---)t---x'
:9 0.05 28°C
o 2 4 6 8 10 12 o 2 4 6 8 10 12
Storage period (weeks) Storage period (weeks)
l
~ 0.80 ~ 1.0
1
OJ! 0.60 /28 C 0 0.8
§~ 0.40 ..,,4 x 0.6 X
If
/
~~ 16°C,.x, ...... '-0- ,/16°C
o 0.20
o
~ I --0---0----- -0-
"X _ _ ~--w--
)(" rc
""'t)---O
0.4
_x~''''x'';C
~-o--.o
- !! I ,!, !
o
!
2 4 6 8 10 12 o 2 4 6 8 10 12
Storage period (weeks) Storage period (weeks)
Figure 14.29 Changes in content of glucose, fructose and sucrose during storage
at 7, 16 and 28°C, of cv. Bintje (treated with CIPC). (From Linnemann et at.,
1985).
difficulties with reconditioning. On the one hand, reconditioning is easier

the later in the storage season it is carried out, probably by virtue of
sprouting and the associated increase in respiration. On the othe.r hand,
Burton and Wilson (1978) point out that the required decrease may not
then occur, because of senescence. The problem is to find the exact time at
which sprouting, respiration and senescence are in optimum balance for
good reconditioning.
The foregoing applies to potatoes lifted when mature. Yamaguchi (1959)
found that relatively immature potatoes were harder to recondition,
although there are exceptions. Hak (1980) showed that cultivars differed in
their response to reconditioning for 14 days at 20°C. Cv. Saturna appeared
to be more susceptible to reconditioning than cv. Bintje.
Probably because of these differences, the rate and effect of recondition-
ing may be influenced by the degree of maturity at harvest. This is
confirmed by Yamaguchi et al. (1966). They indicated that immaturity of
the tubers at harvest could slow reconditioning after cold storage. The
results reported above do not always agree with the findings of other
research workers. Hair and Gould (1979) state that the influence of
maturity, as measured by harvest date, on reconditioning is not very
clear. They found that some cultivars became more prediposed to rapid
conditioning with advancing maturity, whereas others were apparently not
affected by maturity in the same way. Besides the influence of maturity,
variations in cultivars are involved in the process of reconditioning.
(c) Formation of carbohydrates accompanying sprouting

Sprouting is associated with the mobilization of carbohydrates, mainly by
starch being hydrolysed from the tuber. Starch is the source of carbon and
energy for the developing sprouts. Microscopic investigations show that
the breakdown of the amyloplasts generally begins at the basal end of the
potato tuber. From there the sugars are translocated to the top region of
the tuber and the emerging sprout in the form of sucrose, and are
hydrolysed, at least partly, into glucose and fructose. Bailey et al. (1978)
showed that the levels of reducing sugar in potato tubers (cv. Majestic)
increased up to eight-fold during storage from October to May, with
almost all the rise occurring after emergence from dormancy in December.
Removal of all buds from the tubers within a month of harvest prevented
this rise in soluble carbohydrates. These experiments clearly show the
association between the onset and development of sprouts, and the
increase in reducing sugars in the tissue.
Sugar development during 'normal sprouting' at 4 and loDe

In experiments with cv. Bintje, stored at 4 and lODe, van Es and Hartmans
(1986) show the fluctuations in sucrose and in the reducing sugars glucose
and fructose. Over the period during which sprouting vigour developed
from dormancy to a maximum value, sprouting vigour was measured
according to a modification of the method described by Krijthe (1962).
At lODe the reducing sugars showed only a slight increase. At 4°e the
increase was much more pronounced, probably due (at least partly) to low-
temperature sweetening. The sucrose content at 4°e remained almost
constant whereas at lODe the increase was only moderate. Thus under
conditions of unlimited sprout growth, at a storage temperature of lODe,
the influence of sprout growth (even over a longer period) was not very
pronounced. In this case of immature tubers the picture was similar except
that the development of the reducing sugars at 4°e was even more
pronounced than in mature tubers, possibly due to low-temperature
sweetening (Figure 14.30).
13 4°C
(a)
Bintje 'mature' .g, 12
I
.::: 11
Ol
010
8,....
6 4°C 9 Bintje
Q)~
~ 'immature'
Q)
(fl-
o..c 5 (fl 8
nOl .9
E~ 4
.E
u 7
+-;;, 3 + 6
Q)O
(flO
10°C Q)
00 (fl
u,....
~~
2 8 5
a
-
C)~
Ol ~
4 ~100e
01..-----'-----'----1.- 3
o 100 200 300 days 0~--:1:-:!0""0---::~--"-:::::-
~~ 0.3 we 4°C sE 0.3
~I
.:.::: 0.2
i~02
>...: .
e~ 0.1
o.Ltl
~;
'- 0.1
~~ ~
~ 0 ~.9 0 Lc:..eo.-;:L.---~-----&.-
o 100 200 300 days 0 100 200 300 days
Storage period Storage period
(b) Bintje 'immature'

Bintje 'mature' 6
5
P
5
E E014
Ol 4
Q)I Q)I
(fl"': 4°C (fl"':
0-
tOl
3 §-;;' 3
~o
~o
0
2 ~8
0
2
,....
~ ~1
0 00
0 100 200 300 days 100 200 300 days
4°C
SE 0.3 sE
00l
0.3
00l
.2' I 02 .2' I 02
> to..: • > ~ .
"5;'
~-;;, 0.1 e Ltl
o.~
0.1
Ci~ ~Ol
~9 0 ~ 0
0 100 200 300 days 0 100 200 300 days
Storage period Storage period
Figure 14.30 Fluctuations in content of reducing sugars (a) and sucrose (b) in
relation to sprouting vigour, during storage of cv. Bintje (mature and immature) at
4 and lOoC. (From van Es and Hartmans, 1986.)
Manual desprouting
Hughes and Fuller (1984) investigated the relationship between sugar
development, dormancy and sprouting of tubers (cv. Record) s:tored at
lOoe. When tubers were in the stage of apical dominancy, they were
divided into two groups: a 'grow on' group and a 'desprouting' group.
Throughout storage the sprouts in the latter group were removed at weekly
intervals. Both the apical tissue and the basal tissue were analysed. The
difference in sucrose content was not very marked. The levels of reducing
sugar in the tissue surrounding the basal eye, however, were considerably
higher than those round the apical eye (Fig. 14.31). Regular removal of
sprouts caused levels of reducing sugar to continue to fall after the initial
rise, whereas the 'grow on' tubers showed a marked increase, in levels of
reducing sugar, particularly at the basal end. No marked increase in
sucrose occurred at the end of the storage season in the de sprouted tubers.
Weekly removal of the sprouts caused a loss of mobilized and transported
sugars. This will be at least one of the reasons why the sugar concentrations
in the tissue were kept at a low level by the manual desprouting.
eft.
-;;: 0.6
eel
(/)
C
~ 0.4
eel
OJ
m eyes 100%
g> 0.2 open sprouting
'0 grown on
~ __ ~~_----_o desprouted
a:: 0 0 - 50 100 150 200 250 300
(a) Days in store at 10°C
eyes
eft. open
~ 0.60
c. )\100%
eel
(/) / xsprouting
C
.;;; 0.40 x "\ / x grown on
(ij
OJ x X-0 x/
:::J '?--
(/) ....
OJ 0.20 ...... 0 desprouted
c " " _ 0 ____ --0
'0
:::J
'0
Q)
a:: 50 100 150 200 250 300
(b) Days in store at 10°C
Figure 14.31 Fluctuation in reducing sugars of cv. Record. (a) Apical eyes and (b)
basal eye. (From Hughes and Fuller, 1984.)
(d) Increase in sugar content due to senescence
The increase in sugar content due to senescence is a separate phenomenon.
In conditions of constant temperature, the sugar content remains relatively
constant for a certain time, and then increases, at first slowly and finally
very sharply. This is clearly illustrated in Fig. 14.32 (Barker, 1938).
Barker's experiment shows that the higher the storage temperature, the
earlier senescent sweetening starts. Burton (1965), investigating sweeten-
ing after prolonged storage at lOoe, found that the magnitude of this
sweetening shows considerable cultivar differences.
3.0..--------------------,
~
.£:
(/)
2.5
! • 10°C
ic: 2 . 0 J - - - - - - - - - - - - - - - - - - r - - - - - - - i
.0,
.§ 1.5
-#-en 1.0 o
OJ
::J
(/)
! O.:~~~~~~~~~~~~--~~~~~~~~~
N DI J F M A M J J A SON DIJ F M A
1935 1936 1937
Storage period
Figure 14.32 Changes in the sugar content of mature tubers of cv. King Edward
VII during prolonged storage at 15, 10, 7.5 and 5°C. Tubers stored at the three
higher temperatures exhibit senescent sweetening, those at 5°C, low-temperature
sweetening, with possibly some senescent sweetening later, in addition. (From
Barker, 1938.)
A characteristic of sweetening after prolonged storage is that it is not

reversible. Reconditioning measures are ineffective (Isherwood, 1976).
This sugar formation, called 'senescent sweetening' by Burton (1966), is
characterized by changes in the membranes surrounding the starch grains
(Isherwood and Burton, 1975). In popular terms, the membranes may be
said to have become 'leaky' (Fig. 14.33). Glucose and fructose originating
from the breakdown of starch accumulate. The rate and extent of this
sweetening depends on the cultivar. The history of the potato during
growth and at the time of lifting is also relevant (Dwelle and Stallknecht,
1978). There seems to be some connection with the sprouting potential:
sugar formation is stated to be higher the greater this potential (Burton,
1965). The only effect of sprouting control in this case is to increase the
'accumulation' of the sugars.
Proteins Lipids
Starch = n x Glucose
G GIJose -@--@-®-
F
.J.
Fructose
II1 Fructose
G Glucose
Figure 14.33 Schematic drawing of 'leakage' of sugar through starch grain

membranes due to senescence. (From van Es and Hartmans, 1987.)
(e) Effect of tuber characteristics

Cultivar
Of all the factors that determine the amount of reducing sugars during
storage and hence the suitability for processing, cultivar is probably the
most important.
van Vliet and Schriemer (1960), using 12 different cultivars grown under
identical conditions, showed that there were differences in maximum
reducing sugar levels, ranging from 5.9 to 26.5 mg per gram fresh weight
when the potatoes were stored at 2°C. The sucrose content varied from 4.3
to 14.9 mg per gram fresh weight. Approximately the same values were
measured in a different year. The order of the cultivars as regards sugar
content was in good agreement in the two years. Samotus et al. (1974) also
reported that sugar formation in a number of Polish cultivars stored at 1, 2
and 6°C for 18 to 22 weeks was correlated with cultivar.
Sowokinos (1973) states that the sugar content of potatoes during
tuberization and at the time of lifting is largely dependent on cultivar even
under virtually identical growing conditions from year to year at specific
locations in the USA. He also found that the response of potatoes to
storage at low temperatures is a typical cultivar characteristic and is
connected with the sugar content at the beginning of storage. It was found
that for a given degree of maturity, at the time when 97% of minimum
tuber size and maximum starch content were attained, there were
differences in the sucrose content ranging from 1.9 to 4.5 mg per gram of
tuber fresh weight. The main differences between cultivars in terms of
suitability for processing into potato products at full maturity were found in
the final concentrations of sucrose.
Experiments carried out by van Es and Hartmans (1986) with two
cultivars (Bintje and Saturna) clearly show cultivar differences in the
development of reducing sugars. Stored at 8-10°C cv. Bintje showed an
increase of about 0.2% (% fresh weight) in glucose + fructose. Under the
same conditions cv. Saturna showed only a very slight increase in reducing
sugar content.
Storage at low temperature (4°C), however, produced a great difference
between the two cultivars in the accumulation of reducing sugars. After a
storage period of 200 days at 4°C the reducing sugar content was about
0.9% for cv. Bintje and 0.2% for cv. Saturna. Compared with cv. Bintje,
cv. Saturna shows only a very limited reaction to the low-temperature
storage regime.
Maturity
Burton (1966) states that apart from cultivar differences, maturity may be
one of the principal factors affecting sugar content. Tuber maturity is very
important for the sugar content at the beginning of the storage period and
the variation in that content during storage, and determines the potato
quality for processing. The initial level of sugars - reducing sugars as well
as sucrose - at the moment of harvesting is affected by maturity. The
immature tubers contain more reducing sugar than the mature tubers.
Physiologically younger tubers show a faster response to an increase in
levels of reducing sugars as well as of sucrose (Samotus et al., 1974). A
clear relationship between harvest time, tuber size, dry matter content,
starch, sucrose and reducing sugar content of the tuber was demonstrated
by Wiese et al. (1975). The reducing sugar content decreased with
increasing size, starch and sucrose content.
Miller et al. (1975) showed the existence of an optimum harvest date at
which cultivars produced chips lightest in colour. Conversely, the darkest
coloured chips were obtained from each cultivar 15 days after the optimum
harvest date: this seemed to be attributable to cool, wet weather. There is
considerable variation among cultivars in their susceptibility to the degree
of maturation and related physiological processes. van Es and Hartmans
(1986; see Figs 14.30 and 14.34) show the cultivar influence of maturity on
the formation of reducing sugars during storage at 4 and 8°C; cv. Bintje
shows a distinct difference in sugar formation between 'mature' and
'immature'. In cv. Saturna, however, the difference with regard to
maturity is negligible.
Wiese et al. (1975) concluded that at the time of harvest, coinciding with
the end of flowering, net assimilation rate clearly correlates with falling
reducing sugar content. As the haulm begins to die, this relationship tends
to reverse. The starch content of the tubers then reaches its highest level.
1.4 1.4
1.2 1.2
f
-£ 1.0 !
r··· . •·· ..... 4 C immature
i
~ 1.0
g:
Q
! ..., ................................... .. g:
;f.
! ...... ~ 4°C mature
-; 0.8 ! ~ 08
~2
~ 0.6 i
:
:
i
i
~ 0.6
fii J fii
4°C immature
~
g 0.4 ~
g 0.4
a Q •••• d1"... ····.;j). •.• oQ. ••• "\I)o •••• o•••• ¢lo •••• o-···~ BOC immature a 't·""·······\
.....=.:::::t.. ....:
0.2
-oll-o--¢.-a_ll>-o_otl BOC mature
0.2 ! ~4ocma1ure
o~---- ____________________
i..a. ..... ~~~88°Cmature BOC immature
o
o 50 100 150 200 250 300 350 o 50 100 150 200 250 300 350
(a) Days after the first harvest (b) Days after the first harvest
Figure 14.34 Cultivar influence of maturity on the formation of reducing sugars

during storage at 4 and 8°C, of cvs (a) Bintje and (b) Saturna. (From van Es and
Hartmans, 1986.)
This is in agreement with Sowokinos (1973) who defined the state of

physiological maturity as the moment that: '97% of the final tuber size and
starch content has been attained'.
Sowokinos (1977), dealing with the quality of potatoes to be stored for
chipping and crisping, indirectly related the reducing sugar content to the
sucrose at the time of lifting.
14.5.4 N-fraction
The term 'crude proteins' is frequently used in the literature. To determine
this, the total weight of nitrogen is measured by the Kjeldahl method and
multiplied by a factor of 6.25. Desborough and Weiser (1974) showed that
for potato protein this factor should be 7.5, corresponding to the amino
acid composition. There are no major changes in the crude composition of
the major components of the N fraction during most of the storage period.
Consistent changes occur during sprout growth and development. These
are mainly in amide and free amino acid fractions (Tagawa and Okazawa,
1953; Szalai, 1957a, b, c; 1959a, b; Talley et al., 1964). The largest and
most consistent changes occur in the content of proline, which increases,
particularly towards the end of storage, and is translocated to the sprouts
(Breyhan et al., 1959; Heilinger and Breyhan, 1959; Heilinger, 1961;
Talley et al., 1964). Harvesting wounds or disease lesions could lead to a
local increase in the content of tyrosine, necessarily at the expense of some
other nitrogenous constituent. Johnson and Schaal (1957) found a doubling
in the content in cells immediately adjacent to the wound, but the effect on
the overall composition of the N-fraction of the whole tuber would be
slight. Important changes in protein composition take place during tuber
induction and subsequent development, although not during storage.
Nitrogen occurs in the form of inorganic nitrogen, in the form of proteins
and many different non-protein organic chemical compounds (such as free
amino acids).
(a) Protein N
Using differential solubility, proteins can be distinguished into albumin,
globulin, prolamin, glutelin and an insoluble residue. Using the extraction
method of Nagy et al. (1941), Kapoor et al. (1975) determined the levels of
these protein components (Table 14.16). The analysis also gave an
impression of the quality of potato protein in terms of nutritional value
(Table 14.17). Rexen (1976) determined the amino acid composition of
hydrolysed potato material from 20-33 cultivars grown with different doses
of nitrogen. Differences in amino acid content of hydrolysed potato
Table 14.16 Protein fractions as percentages of total protein

nitrogen expressed on a freeze-dry basis (from Kapoor et aI.,
1975)
Protein fraction mg of %of total

protein-N per g powder protein-N
Albumin 412 48.9
Globulin 218 25.9
Prolamin 36 4.3
Glutelin 74 8.8
Residue 75 8.9
Table 14.17 Chemical score, essential amino acid index (EAA)

and biological value determined in protein fractions (from
Kapoor et aI., 1975)
Protein fraction Chemical EAAt Biological
score* valuet
Albumin 61 82 77
Globulin 57 83 79
Prolamin 40 53 46
Glutelin 60 83 79
Residue 69 82 77
* Chemical score is the content of each essential amino acid in a protein
fraction expressed as a percentage of the content of the same amino acid
in the same quantity of egg (Anon., 1970).
t Essential amino acid index (EAA) is defined as the geometrical
means of the ratio of amino acids from potato and egg protein (Oser,
1959).
:j: Biological value was determined by an equation of Oscr (1959).
material among cultivars have been found to be roughly proportional to
differences in total nitrogen (Talley et al., 1984). The effect of 4 months
storage at two temperatures (3.3 and 7.2°C) on amino acids in the cvs
Russet Burbank and Katahidin was found to be minor (Talley et al., 1984).
Since the advent of gel electropheresis and chromatography a more
complex and realistic picture of the composition of potato protein has
emerged. Stegemann (1975) observed between 10 and 40 proteins by
electrophoresis. The electrophoretic protein patterns are characteristic and
have already been used for many years for identifying cultivars. Protein
crystals also occur naturally in potato tubers (Rodis and Hoff, 1984).
During senescence the molecular weight of some proteins falls sub-
stantially, owing mainly to dissociation of the larger protein complexes.
Much of the protein is made up of 'storage proteins', whose functions are
largely unknown; these storage proteins are highly ami dated (Mulder and
Bakema, 1956).
(b) Non-protein organic N

The non-protein nitrogen fraction contains both organic and inorganic
nitrogen. The organic nitrogen fraction contains 'free amino acids' and the
amides glutamine and asparagine. These compounds account for an
important part of the total fraction. The composition of the free amino acid
fraction is shown in Table 14.18 (Davies, 1977). Synge (1977) compares the
results of 13 cultivars in seven different countries.
The free amino acid content may vary from cultivar to cultivar and from
year to year, as Table 14.18 shows for four cultivars and several different
years (taken from Davies, 1977). The free amino acid content of potatoes
is affected not only by cultivar and year of cultivation but also, and just as
significantly, by factors such as soil type, fertilizer application, climate and
origin (Talley et al., 1970; Rexen, 1976; Davies, 1977).
As has already been noticed the most consistent changes occur in the
content of proline which increases particularly towards the end of storage
and is translocated to the sprouts.
There are many other organic nitrogen components in the potato tuber:
for instance, polyamines, purine derivatives, nitrogenous glycol derivatives
such as choline, and also the steroid alkaloids. The last group contains the
well-known glycoalkaloids, a-solanine and a-chaconine (Burton, 1966).
(c) Glycoalkaloids
All parts of the potato plant contain a toxic glycosidic steroidal alkaloid. A
normal potato tuber contains an insignificant amount of the glycoalkaloids.
However, certain environmental conditions, such as exposure to light, or
mechanical damage, will affect normal physiological sequences and induce
synthesis of the glycoalkaloids in the potato tuber. A bitter and/or musty
Table 14.18 Free amino acids of potato tubers (mg amino acid per 100 g tuber dry matter). Glutamine results for the
1969 and 1970 samples, including glutamic acid (from Davies, 1977)
Amino acid* Record Desiree Alpha Bintje
1969(2), 1970 1969, 1970, 1973 1969, 1970 1969, 1970
Aspartic acid 259 (199-376) 183 (162-212) 247, 233 197, 249
Asparagine 1937 (1200-2594) 2183 (1518-2756) 1159, 1947 1230, 2554
Threonine 69 (40-103) 62 (48-74) 39, 34 45, 63
Serine 63 (43-83) 66 (52-86) 47, 53 44, 47
Glutamine 1479 (973-2050) 1630 (822-2450) 871, 1260 984, 1861
Glutamic acid (-) (345-)
Proline 130 (42-303) 79 (48-137) 62, 201 49, 297
Glycine 11 (7-18) 13 (8-17) 9, 9, 20
Alanine 24 (23-25) 29 (19-45) 24, 30 36, 118
Valine 152 (74-281) 251 (179-312) 60, 182 110, 220
Methionine 48 (32-64) 95 (76-108) 55, 56 51, n
Isoleucine 58 (42-88) 131 (109-165) 45, 52 59, 67
Leucine 33 (25-38) 58 (43-71) 16, 31, 44
Tyrosine 66 (51-93) 229 (174-316) 36, 27 59, 33
Phenylalanine 81 (49-102) 149 (91-204) 56, 50 55, 27
Tryptophan (42-) 79 (67-91) , 145,
Lysine 52 (33-64) 112 (84-144) 39, 75 32, 45
Histidine 163 (85-247) 161 (71-239) 62, 107 91, 183
Arginine 434 (258-670) 326 (196-419) 181, 298 143, 280
4-Aminobutyric acid 171 (49-326) 206 (50-398) 66, 310 146, 442
Ornithine (26-) 75 (5-148) - , -,
• Where there were three or more samples of the same cultivar, the average and range are reported; otherwise the actual results are
given.
taste and off-flavour have been noticed in such tubers that have developed
excessive amounts of the glycoalkaloids. Potatoglycoalkaloids mainly
comprise two substances; a-solanine (solanidine-galactose-glucose-rhamnose)
and a-chaconine (solanidine-glucose-rhamnose-rhamnose). In addition to
the glycoalkaloids, the alkaloid solanidine occurs in the tuber. Other
glycoalkaloids also occur in potato tubers (Maga, 1980; Sinden and
Sanford,1981). The consumption of potatoes with high glycoalkaloid
contents may cause serious illness, sometimes leading to death (Jadhav et
at., 1981) in humans or in animals.
In their· review Jadhav et at. stated that the glycoalkaloids of potato are
not destroyed during boiling, baking, frying or drying at high tempera-
tures. However, Ponnampalam and Mondy (1983) showed that baking and
frying caused a significant decrease in total glycoalkaloid content. The
glycoalkaloid content in the tubers is concentrated in the peel and sprouts
and around the eyes, as shown in Table 14.19. The glycoalkaloids are
formed in the parenchyma cells of the periderm and cortex of the tubers
(Table 14.19). Hence about 40% is eliminated on peeling.
Table 14.19 Distribution of total glycoalkaloids in the potato

plant and various tuber tissues (after Wood and Young,
Agriculture Canada, Publication 1533, Canada Department of
Agriculture, 1974; from ladhav et aI., 1981)
Potato part Total glycoalkaloids
(mg per 100 g fresh weight)
Sprouts 200-400
Normal tuber tissue
Skin 2-3% of tuber 30-60
Peel, 10-15% of tuber 15-30
Peel and eye, liS-in. (3 mm) disk 30-50
Peels from bitter tubers 150-220
Flesh 1.2-5
Whole tuber 7.5
Bitter tubers 25-S0
The potato tuber normally contains small quantities (about 2-10 mg per
100 g) of solanidine and its glycoalkaloids. The total tuber glycoalkaloid
(TGA) content of cuitivars, whose tubers have not been exposed to light,
should not be greater than 20 mg per 100 g of fresh tissue weight and
preferably should be much less (Parnell et ai., 1984). It is assumed that a
level in excess of 20 mg per 100 g fresh weight constitutes a potential health
hazard (Jadhav and Salunkhe, 1975). A glycoalkaloid content exceeding 30
mg per 100 g fresh weight, corresponding to a level of 11-14 mg per 100 g
fresh weight in the peeled tuber, causes a bitter taste (Sinden et at., 1976).
Exposure to light
When potato tubers are exposed to light during harvesting, handling and
marketing, a green colour develops in the periderm and in the outer
parenchyma cells of the cortex. Also a bitter-tasting substance is formed.
The green coloration is a consequence of chlorophyll synthesis, while
the bitter taste is attributable to the formation of glycoalkaloids. The
formation of chlorophyll and glycoalkaloids on exposure to light are
independent processes (Gull, 1960). Light is the main cause of high
glycoalkaloid synthesis in the skin and adjacent layers, orange light being
the most effective while green light showed the least effects (Conner,
1937; Petermann and Morris, 1985). (Figure 14.35) The concentration
increases two to three times after 2 days of illumination (Baerug, 1962) .
• (J - Chac6nine
10 • ({ - Solanine
400 500 600 700

Wavelength (nm)
Figure 14.35 The spectral response of glycoalkaloid synthesis in potato tubers to

light of equal quanta (0.25 !lmol m- 2 S-I, 500 h). The k-ratio LSD is 21.0 for
a-solanine and 21.8 for a-chaconine. (From Petermann and Morris, 1985.)
The increase in glycoalkaloid and alkaloid levels depends on the

duration of exposure to light, the light intensity and the wavelength of the
light. Zitnak (1953) observed a rapid increase in glycoalkaloids when newly
lifted tubers were exposed to sunlight (Table 14.20).
Table 14.20 Increase in glycoalkaloid

content in mg per 100 g fresh weight of
newly lifted tubers of two potato cultivars
when exposed to sunlight (after Zitnak,
1953, from ladhav and Salunkhe, 1975)
Hours' light Netted Gem Katahdin
o 1.7 1.5
6 12.0 5.0
12 13.2 7.1
36 20.4 16.3
The property of forming high levels of glycoalkaloids is primarily

determined genetically, but it has been shown that a short cool growing
season may lead to the production of immature tubers, which have higher
levels. De Maine et al. (1988) showed that a cultivar which has a low
sensitivity to light, that is one in which the glycoalkaloids increase at a low
rate, can be more useful, for example, than one which, although it has a
low level, rapidly increases alkaloid content in the light.
Wounding
Damaging and bruising increase the glycoalkaloid content. Sinden
(1972) observed higher levels in immature tubers which had been graded
mechanically and then stored for two days in the dark than in tubers
exposed to sunlight in the field for two days. Fitzpatrick et al. (1978) and
Ahmed and Miiller (1978) observed an approximately 50% increase in
glycoalkaloid content due to mechanical damage. Olsson (1986) studied
the effects of impact damage on the accumulation of glycoalkaloids in a
wide range of genotypes and found that damage caused by drop test
resulted in increased glycoalkaloid levels which sometimes exceeded the
toxic level of 20 mg 100 g-! freshweight: there were large differences
between genotypes.
When potatoes are cut into chips for processing, the raw products exhibit
an increase in glycoalkaloid content when stored in the dark (Ahmed and
Miiller, 1978; Maga, 1980). This increase is extremely temperature-
dependent, being slight at low temperature (Salunkhe et al., 1972). It has
been found that the process of chipping is non-destructive to total
glycoalkaloids (Sizer et al., 1980). When the sprout suppressant CIPC is
used, the increase resulting from wounding is found to be inhibited (Wu
and Salunkhe, 1977).
Storage
The glycoalkaloid content of potatoes increases during storage (Cronk et
al., 1974; ladhav and Salunkhe, 1975). The storage temperature has a
marked effect (Zitnak, 1953; Salunkhe et al., 1972). At higher tempera-
tures there is an appreciably greater increase (Fig. 14.36). Storage at low
temperature (4-8°C) and high relative humidity doubled the glycoalkaloid
content of the cv. Netted Gem in 6 weeks, while storage for an equal
period in dry, warm conditions (12-15°C) had almost no effect on this
content (Zitnak, 1953). In a 3-month storage experiment at about 12°C
with 25 samples of different cultivars, Wilson et al. (1983) showed that the
highest glycoalkaloid content was the initial value before the tubers were
stored. Sprouting during storage increases the glycoalkaloid content in the
tissue surrounding the eyes. The sprouts themselves have a high alkaloid
content, which depends on cultivar and may amount to as much as 400 mg
of total glycoalkaloids and solanidine per 100 g fresh weight (Table 14.19).
a ~ooe
Oaoe
f]15°e
~
Itiiii24°e
11111
OJ
Dark Light
Figure 14.36 Effect of temperature on solanine formation in Russet Burbank
potato slices stored 48 h in dark or light (200 ft-c). Original concentration 0.3 mg
per 100 g fresh wt. (From Salunkhe et at., 1972.)
(d) Inorganic nitrate-nitrogen

Increasing interest is being focused on the nitrate content of various types
of agricultural produce for human consumption (Augustin et al., 1977;
McDole and McMaster, 1978).
Heisler et al. (1973) also give values for nitrite levels in potatoes: they
found an overall average of 0.44 ppm. Augustin et al. (1977) reported an
apparent slight incease in nitrate-N in the tubers with storage. Boosting the
nitrogen fertilization increases the nitrate content (Nitsch and Klein, 1983).
It has been found that high levels of nitrogen fertilizer application
combined with periods of moisture stress may give rise to very high nitrate
levels in potato tubers. McDole and McMaster (1978) found that with low
nitrogen fertilization and optimum, high irrigation, tubers contained 70-80
ppm N03-N. If there was a moisture deficiency during growth, the level
rose to 144 ppm. With excessive nitrogen fertilizer application, these
nitrate contents were 151-154 ppm and 370 ppm respectively.
Munzert and Lepschy (1983) determined nitrate content of different
cultivars grown at four different locations and found specific cultivar
differences. In 1982 the cv. ludika had the lowest value, its average nitrate
content being 53 ppm. The highest average content, 233 ppm, was
measured in the cv. Clivia.
Carter and Bosma (1974) determined the nitrate content over 3 years,
finding levels of 36-131,34-75 and 25-50 ppm respectively. A limit of 3.65
mg kg- 1 body weight day-l has been set as acceptable daily intake for the
nitrate ion; for nitrite this value is 0.13 mg (WHO, 1974).
14.5.5 Other constituents
(a) Vitamins
Vitamins are an important group of compounds for nutrition. The potato
makes an appreciable contribution to the dietary intake of vitamin C,
which is present in substantial quantities, and the vitamins of the B group.
The latter comprise Bl (thiamine), B z (riboflavin), Bs (probably nicotinic
acid = niacin) and B6 (pyridoxine) (Pol and Labib, 1963; Burton, 1966;
Adler, 1971; Muller, 1975). Niacin and folacin also belong to this group
(Augustin, 1975; Augustin et al., 1978). The mean vitamin composition of
freshly harvested and stored potatoes is given in Table 14.21.
Vitamin C is taken here to include both the reduced (ascorbic acid) and
oxidized (dehydroascorbic acid) form. There are several changes which can
occur during storage, including synthesis, interconversion of the two
forms, conversion of dehydroascorbic acid to diketo~gluconic acid (which
represents a loss of vitamin C), and other reactions leading to loss. The net
result which has usually been reported is marked loss of vitamin C during
storage: about 50%.
There was evidence of a temperature optimum in the work of Effmert et
al. (1961) but marked loss was found by Zilva and Barker (1939) at lOoC
not greatly different from 8°C, and the question of the extent of storage
losses of vitamin C appears to be unsettled. Barker and Mapson (1950)
found much evidence of effects of temperature, there being, in freshly
harvested tubers and after prolonged storage, an increase in ascorbic acid
during low temperature sweetening. Different results were observed in the
middle of the storage season, and clarification of the age-response and of
the reasons for this is needed. Barker and Mapson (1952) found anaerobic
conditions prevented change in ascorbic acid - either synthesis or loss - but
they stated this inhibition was not permanent, there being a slow decrease
in content after exposure to anaerobic conditions for 3-4 weeks. They
stated that there was no sign of tissue injury in the experimental tubers.
This is unusual after anaerobiosis for 3-4 weeks, however, and a more
detailed study of the effects of oxygen concentration, in relation to
temperature of storage, is required. Johnson and Schaal (1957), Jadhav et
al. (1984) and Mondy and Leja (1986) found an appreciable, although
temporary, increase in ascorbic acid in damaged tuber tissue. Muneta and
Kaisaki (1985) reported a complex of ascorbic acid plus ferrous iron
(Fe2+) forming a purple pigment similar to the after-cooking pigment that
Table 14.21 Mean vitamin composition of freshly harvested and stored potatoes (in mg per 100 g dry matter). Storage
conditions 7.2°C and 95% RH. Reconditioning at room temperature following cold storage (from Augustin et aI., 1978)
Vitamins Storage Russet Katahdin Norchip Kennebec Superior Pontiac
time Burbank
(months) 3* 1* 1(2)* 3* 1(2)* 1*
Ascorbic 1 82.~117.2 108.2 86.3 76.8-109.2 . 129.8

acid 4 48.9-57.8 52.0 44.8 49.3-51.8 52.8
8 42.0--53.4 54.2 43.4 46.5-50.4 48.1
Thiamine 1 0.3~.42 0.45 0.3~.37 0.37--0.45 0.31--0.35 0.44
4 0.30--0.48 0.45 0.30--0.47 0.34--0.48 0.44--0.46 0.57
8 0.43--0.45 0.54 0.39--0.40 0.41--0.48 0.37--0.39 0.50
Riboflavin 1 0.14--0.24 0.33 0.22--0.34 0.17--0.20 0.18--0.24 0.19
4 0.10--0.15 0.12 0.22--0.37 0.11--0.13 0.14--0.20 0.23
8 0.11--0.14 0.18 0.27--0.37 0.14--0.21 0.15--0.17 0.17
Niacin 1 5.1-9.4 8.4 5.5-6.4 6.3-10.3 7.1-9.6 8.9
4 5.5-7.4 7.4 5.5-9.1 7.9-9.1 4.1-13.5 8.9
8 4.7-6.3 5.5 4.5-6.3 3.5-7.6 3.2-11.9 7.2
Folic acid 54.2-97.4 84.2 59.3-73.6 81.4-81.5 74.9-77.7 87.9
4 41.1-49.6 48.9 53.4-55.0 45.5-58.0 48.0--57.7 65.0
8 44.3-64.1 50.7 51.3-52.5 44.0--71.4 46.2-62.5 72.8
Vito B6 1 0.95-1.01 0.92 0.49--0.69 0.60--0.90 0.71--0.89 0.69
(pyridoxin) 4 1.08-1.35 1.51 0.62--0.93 0.6~.99 1.05-1.12 0.79
8 1.41-1.72 2.08 1.1~1.35 1.15-1.57 1.50--1.85 1.62
* Number of locations sampled.
could be overcome with strong chelating or complexing agents such as
citric acid and sodium acid pyrophosphate.
The thiamine content fluctuates during storage, while the riboflavin and
niacin contents vary little (Augustin 1975). The vitamin C content, on the
other hand, falls appreciably as the N dose increases; it is also affected by
the type of soil. Ascorbic acid content increases with growth and maturity
of the tuber (Mazza et al., 1983). When analysing the major US potato
cultivars in various locations for their contents of water soluble vitamins,
both at harvest time and during subsequent storage, Augustin et al. (1978)
showed that the compositional ranges of each vitamin were quite large.
Only a few cultivar and location effects were detected. Prolonged storage
had little overall effect on thiamine and riboflavin. In all cases, however, it
resulted in a sharp initial decrease in ascorbic acid, significant decreases in
niacin and folic acid and a significant large increase in vitamin B 6 . A
storage temperature in the range of 3.3-7.2°C did not affect the vitamin
composition, nor did reconditioning of the tubers at room temperature
following cold storage (Augustin et al., 1978).
(b) Phenolic compounds - organic acids

Phenolic compounds of known importance to the cooking quality of the
tuber include tyrosine, already mentioned above; chI orogenic acid (Gray
and Hughes, 1978) and perhaps caffeic acid. Of the two last, chlorogenic
acid increases in concentration in the buds during storage (Amberger and
Schaller, 1973); is synthesized and increases very markedly in the
cells adjacent to wounds (Politis, 1948; Johnson and Schaal, 1957;
Ozeretskovskaya and Vasyukova, 1965), and shows a temporary rise,
followed by a fall, after y-irradiation (Penner and Fromm, 1972).
Malberg and Theander (1984) analysed free and conjugated phenolic
acids in the peel and the peeled tuber of six cultivars (Bintje, Magnum,
Bonum, Sabina, Ukama and Stina). Caffeic, protocatechuic, ferulic and
vanillic acids as free acids were found in all varieties in the peel. Brandl and
Herrmann (1984) investigated ten potato cultivars and determined the
amounts of the different isomers otchlorogenic acid and the n-, krypto-
and neo-chlorogenic acids. The predominant compound was shown to be
the n-chlorogenic acid (3-0-caffeoylquinic acid).
Malmberg et al. (1980) showed that a number of phenolic compounds
showed antifungal activity against Phoma exigua var. foveata (gangrene) in
potatoes.
The major organic acids in potatoes are citric, malic, oxalic and fumaric
acid. Bushway et al (1984) determined the organic acid content in several
potato cultivars (Table 14.22) and demonstrated a wide variation in acid
content among cultivars. The concentration of malic acid increases during
storage, while that of citric acid declines proportionately (Fig. 14.22)
(Adler, 1971; Rumpf, 1972).
Table 14.23 Organic acid content of several potato cultivars (from Bushway et at,
1984)
Cultivar No. mg acid per 100 g tuber wet weight
analysed
Oxalic Citric Malic Fumaric

Allagash Russet 6 37.S±1.0 414.2±33.2 68.4± 9.4 1.2±0.3
Atlantic 6 33.8±1.9 43S.9±24.9 86.1±1O.9 1.3±0.3
Monona 6 26.1±3.4 438.4±34.2 62.3± 9.4 1.2±0.3
Red Pontiac S 30.3±S.3 S03.7±S6.3 33.3±14.1 0.4±0.0
Russet Burbank 6 2S.7±2.0 374.S±SS.S 108.7±20.7 0.4±0.1
Norchip 2 3S.6±4.3 S70.S±16.8 107.8± 4.0 1.8±0.1
Superior 6 27.7±2.3 274.4±14.0 29.2± 4.4 0.8±0.2
Green Mountain 6 28.3±3.2 S06.6±SS.9 68.1±20.3 0.6±O.1
Cobbler S 22.9±1.4 348.6±33.8 2S.2± 2.6 O.S±O.l
(c) Pigments
The tissue of potatoes is coloured light yellow to a greater or lesser extent.
The degree of coloration is a cultivar characteristic (Saiaman, 1926; Burton,
1966). Caldwell et al. (1945) showed that potatoes with white flesh contain
0.014-0.054 mg of carotenoids per 100 g fresh material, while yellower
tubers contain 0.110--0.187 mg of carotenoids per 100 g fresh weight.
Respective average values are 0.021 and 0.138 mg per 100 g fresh weight.
Adler (1971) mentions values of 0.199 to 0.560 mg per 100 g fresh weight.
The carotenoids are highly fat-soluble, are modified by light and contain a
large number of different compounds, such as a-carotene. auroxanthine,
violaxanthine, s-carotene, ~-carotene, lutein, isolutein, a flavoxanthine
and neo-~-carotene. Smaller quantities of flavonols, flavones and flavin are
also found in the tuber (Lampitt and Goldenberg, 1940; Harborne, 1962).
Red-skinned potatoes contain anthocyanin dissolved in the cell sap of the
periderm cells and the outer cell layers of the cortical parenchyma.
(d) Chlorophyll
Chlorophyll is synthesized, particularly in the peripheral layers, if potato
tubers are exposed to light. After 1-2 days illumination there is already a
significant increase (Chaika et al., 1980) and after about 1 month the
chlorophyll concentration reaches a stable level (Mottley, 1980). Synthesis
is increasing with increasing light intensity over the range 140-3300 Ix
(Liljemark and Widoff, 1960). Liljemark and Widoff found daylight-type
fluorescent (350-750 nm) to be effective in promoting the synthesis. Within
this range of wavelength, green light, 500-600 nm, was least, and blue,
400-500 nm, and red, 600-700 nm, increasingly more effective. The degree
of chlorophyll synthesis which occurs under given conditions appears to be
an heritable varietal characteristic (Akeley et al., 1962). Harkett (1975)
found the rate of synthesis to be markedly temperature dependent, a
maximum rate being reached at 15°C and negligible synthesis occurring in
16 days under 100-1100 Ix at 2°C (Burton, 1978a). Once formed, chloro-
phyll is not lost, at least over a period of 33 days, during subsequent
storage in the dark (Harkett, 1975).
(e) Terpenoids
The production of sesquiterpenoid stress metabolites (SSM) by tuber tissue
is characteristic of the hypersensitive response of certain varieties of the
white potato (Solanum tuberosum L.) to strains of the late blight fungus
Phytophthora infestans (Mont.) de Bary. Four of the major SSMs produced
by S. tuberosum are katahdinone (solavetivone), lubimin, rishitin and
phytuberin. As well as an increase in these components there is an increase
in partly unidentified stress metabolites (Alves et al., 1984). They are men-
tioned here because of the widely publicized (Renwick, 1972) but unsub-
stantiated (Chaube et at., 1973; Elwood and MacKenzie, 1973; Emanuel
and Sever, 1973; Smith et al., 1973) suggestion of the teratogenic effects of
blighted potatoes. Increases have also been reported after inoculation with
various other fungi (Varns et al., 1971) and Erwinia caratovora var.
atroseptica (Lyon, 1972). The increases are local and marked accumulation
does not occur far beyond the edge of the rot (Lyon, 1972).
(f) Lipids, hormones and other components

There are many changes in quantitatively minor constituents of the tuber,
which have partly been studied because of their importance in tuber
metabolism, or relevance to the quality or nutritive value of the tuber. For
instance changes in organic acids (citric acid and malic acid) are of known
importance to the quality of cooked potatoes (Hughes, 1963; Bushway et
al., 1984). These acids affect potato quality directly and indirectly. For
example, they influence flavour directly because of their tartness and
colour by inhibiting after-cooking darkening and nonenzymatic browning.
Levels of lipids and their breakdown by the enzymes lipolytic acyl
hydrolase and lipoxygenase, present in the potato tuber, are important in
connection with off-flavours and rancidity problems when potatoes are cut
during processing, the fatty acids linoleic acid and linolenic acid quickly
being broken down. The lipid content of the tuber is generally low, about
0.5% of the dry weight (Galliard, 1973).
An increase in unsaturated fatty acids at low temperatures has been
described for potato tubers stored below 4°C (Cherif, 1973). This might be
important, as a changed membrane composition at low temperatures is
supposed to playa role in several physiological changes observed at low
temperatures.
References 709
Changes in hormones and inhibitors are related to dormancy and
sprouting. Bialek (1974) and Bialek and Bielinska-Czarnecka (1975)
showed the highest level of gibberellin-like substances at the end of
November, which is unlikely to be immediately before sprouting. Resting
potato tubers contain acidic growth inhibitors that disappear when rest
ceases naturally. They also disappear if the tubers are exposed to treat-
ments that break the rest (Hemberg, 1985).
REFERENCES
Adams, M.J. (1980) The significance of tuber damage and inoculum concentration
of Phoma exigua var. foveata in the development of gangrene in stored
potatoes. Ann. Appl. BioI., 95, 31-40.
Adams, M.J. and Griffith, R.L. (1978) The effect of harvest date and duration of
wound healing conditions on the susceptibility of damaged potato tubers to
infection by Phoma exigua (gangrene). Ann. Appl. BioI., 88, 51-5.
Adlan, A.M.B. (1969) The use of sprout inhibitors in simulated high-temperature
storage of potatoes in Sudan. IBVL-Publikatie 214, Wageningen, The Netherlands.
Adler, G. (1971) Kartoffeln und Kartoffelerzeugnisse. Grundlagen und Fortschritte
der Lebensmitteluntersuchung 13, Parey, Berlin, Hamburg.
Aeppli, A. and Keller, E.R (1980) Beziehung zwischen Respiration und Blau-
fleckigkeit von Kartoffeln nach einer mechanischen Behandlung der Knollen.
Potato Res., 23, 25-32.
Ahmed, S.S. and Muller, K. (1978) Effect of wound-damage on the glycoalkaloid
content in potato tubers and chips. Lebensm. Wiss. und Technol., 11, 144--6.
Akeley, R.V., Houghland, C.V.C. and Schark, A.E. (1962) Genetic differences in
potato tuber greening. Am. Potato J., 39, 409-17.
Allen, E.H., Beau, J.N. and Griffith, RL. (1978) Effects of low temperature on
sprout growth of several potato varieties. Potato Res. 21, 249-55.
Alves, L.M., Kirchner, RM., Lodato, D.T., Nee, P.B. (1984) Acetyldehydroris-
hitinol, a rishitinol-related potato stress metabolite. Phytochem, 23, 537-8.
Amberger, A. and Schaller, K. (1973) Wertgebende Inhaltsstoffe verschiedener
Kartoffelsorten in Hinblick auf ihre Verarbeitung zu Edelerzeugnissen.
Chemie, Mikrobiologie Technologie Lebensmittel, 2, 39-41.
Anon. (1970) Amino acid content of food and biological data on proteins. FAG.
Nutr. Stud., 24, FAO, Rome.
Appel, O. (1906) Zur Kenntnis des Wundverschlusses bei Kartoffeln. Ber. dt. bot.
Ges., 26, 118-22.
Appleman, C.O. and Miller, E.V. (1926) A chemical and physiological study of
maturity in potatoes. J. Agric. Res., 33, 569-77.
Artschwager, E.F. (1927) Wound periderm formation in the potato as affected by
temperature and humidity. J. Agric. Res., 35, 995-1000.
Audia, W.V., Smith, Jr, W.L. and Craft, c.c. (1962) Effects of isopropyl N-(3-
chlorophenyl) carbamate on suberin, periderm, and decay development by
Katahdin potato slices. Bot. Gaz., 123, 255.
Augustin, J. (1975) Variations in the nutritional composition of fresh potatoes. J.
Food Sci., 40, 1295-9.
Augustin, J, McDole, R.E. and Painter, e.G. (1977) Influence of fertilizers,
irrigation and storage treatments on nitrate-N content of potato tubers. Am.
Potato J., 54, 125-6.
Augustin, J., Johnson, S.R., Teitzel, e., Toma, R.B., Shaw, R.L., True, R.H.,
Hogan, J.M. and Deutsch, R.M. (1978) Vitamin composition of freshly
harvested and stored potatoes_ 1. Food Sci., 43, 1566-74.
Baerug, R. (1962) Influence of different rates and intensities of light on solanine
content and cooking quality of potato tubers. Eur. Potato J., 5, 242-51.
Bahr, J.T. and Bonner, W.D. (1973a) Cyanide-insensitive respiration. I. The
steady states of skunk cabbage spadix and bean hypocotyl mitochondria. J.
Bioi. Chem., 248, 3441-5.
Bahr, J.T. and Bonner, W.D. (1973b) Cyanide-insensitive respiration. II. Control
of the alternative pathway. J. Bioi. Chem., 248, 3446-50.
Bailey, K.M. Phillips, D.J. and Pitt, D. (1978) The roles of buds and gibberellin in
dormancy and the mobilization of reserve materials in potato tubers. Ann. Bot.,
42,649-57.
Barker, J. (1935) A note on the effect of handling on the respiration of potatoes.
New Phytol., 34, 407-8.
Barker, J. (1937) The effect of temperature of storage on the sprouting of potatoes.
Rep. Food Invest. Bd, London, 1936, pp 179-80.
Barker, J. (1938) Changes of sugar content and respiration in potatoes stored at
different temperatures. Rep. Food Invest. Bd, London 1937, pp 175-7.
Barker, J. and Mapson, L.W. (1950) The ascorbic acid content of potato tubers. II.
The influence of the temperature of storage. New Phytol., 49, 2.83-303.
Barker, J. and Mapson, L.W. (1952) The ascorbic acid content of potato tubers.
III. The influence of storage in nitrogen, air and pure oxygen. New Phytol., 51,
90--115.
Barker, J. and Mapson, L.W. (1955) Studies in the respiratory and carbohydrate
metabolism of plant tissues. VII. Experimental studies with potato tubers of an
inhibition of the respiration and of a 'block' in the tricarboxylic acid cycle
induced by 'oxygen poisoning'. Proc. Roy. Soc. B, 143,523-49.
Basker, D. (1975) Centigrade scale temperature corrections to the specific gravity
of potatoes. Potato Res., 18, 123-5.
Beukema, H.P. and Vander Zaag, D.E. (1979) Potato Improvement. Some factors
and facts, International Agricultural Centre (lAC), Wageningeh.
Beveridge, J.L., Dalziel, J. and Duncan, H.J. (1981a) The assessment of some
volatile organic compounds as sprout suppressants for ware or seed potatoes.
Potato Res., 24, 61-76.
Beveridge, J.L., Dalziel, J. and Duncan, H.J. (1981b) Dimethylnaphthalene as a
sprout suppressant for seed and ware potatoes. Potato Res., 24, 77-88.
Bialek, K. (1974) A preliminary study of activity of gibberellin-like substances in
potato tubers. Z. PJlanzenphysiol., 71, 370--2.
Bialek, K. and Bielinska-Czarnecka, M. (1975) Gibberellin-like substances in
potato tubers during their growth and dormancy. Bull. Acad. Pol. Sci., Ser.
Sci., Bioi., 23, 213--18 ..
Bialek, K. and Bielinska-Czarnecka, M. (1978) Gibberellin pattern in potato
tubers in relation to cultivar specificity and weather conditions. Bull. Acad. Pol.
Sci., Ser. Sci. BioI., 26, 505-12.
Bielinska-Czarnecka, M. and Bialek, K. (1976) Endogenous growth regulators in
References 711
potato dormancy and sprouting. Acta Univ. Nicolai Copernici, Nauki Mat. Prz.,
37,67-70.
Bland, W.L., Tanner, e.B. and Maher, E.A. (1987) Vapor conductance of
wounded potato tuber tissue. Am. Potato 1.,64, 197-204.
Blumenthal-Goldschmidt, S. and Rappaport, L. (1965) Regulation of bud rest in
tubers of potato (Solanum tuberosum L.) II. Inhibition of sprouting by
inhibitor-~ complex and reversal by gibberellin A. Plant Cell Physiol., 6, 601-8.
Boe, A.A., Woodbury, G.W. and Lee, T.S. (1974) Respiration studies on Russet
Burbank potato tubers; effects of storage temperature and chemical treatments ..
Am. Potato 1., 51, 355-61.
Bonnerot, C., Dizengremel, P. and Kader, J.C. (1985) Comparison between
ageing of slice and ethylene- or rindite-treatments on whole potato tubers:
studies on microsomal and mitochondrial enzymatic activities. 1. Exp. Bot., 36,
129-39.
Booth, R.H. and Proctor, F.J. (1972) Considerations relevant to the storage of
ware potatoes in the tropics. Pans, 18, 4.
Boyd, W.D. and Duncan, H.J. (1986a) Studies on potato sprout suppressants. 6.
Headspace analysis of tecnazene and chlorpropham in a commercial box potato
store. Potato Res., 29, 207-16.
Boyd, W.D. and Duncan, H.J. (1986b) Studies on potato sprout suppressants. 7.
Headspace and residue analysis of chlorpropham in a commercial box potato
store. Potato Res., 29, 217-23.
Brandl, W. and Herrmann, K. (1984) Ueber das Vorkommen der Chlorogensauren
in der Kartoffel. Z. Lebensm. Unters. Forsch., 178, 192-4.
Breyhan, Th., Heilinger, F. and Fischnich, O. (1959) Ueber das Vorkommen und
die Bedeutung des Prolins in der Kartoffel. Landw. Forsch., 12, 293-5.
Brown, W. and Reavill, M.J. (1954) Effect of tetrachloronitrobenzene on the
sprouting and cropping of potato tubers. Ann. Appl. BioI., 41, 435-47.
Brownell, L.E., Nehmias, J.V. and Bulmer, J.J. (1954) Utilization of the Gross
Fission Products, Univ. Michigan Engng. Inst. Progress Report 7.
Bruinsma, J. (1966) Plant growth regulators: toys and tools. Mededelingen van de
Rijksuniversiteit Landbouwwetenschappen te Gent, 31, 343-69.
Bruinsma, J. and Swart, J. (1970) Estimation of the course of dormancy of potato
tubers during growth and storage, with the aid of gibberellic acid. Potato Res. ,
13,29-40.
Burton, W.G. (1950) Studies on the dormancy and sprouting of potatoes. I. The
oxygen content of the potato tuber. New Phytol., 49, 121-34.
Burton, W.G. (1952) Studies on the dormancy and sprouting of potatoes. III. The
effect upon sprouting of volatile metabolic products other than carbon dioxide.
New Phytol., 51, 154-61.
Burton, W.G. (1955) Biological and economic aspects of the refrigerated storage of
potatoes. Proc. Inst. Refrigeration, 51, 168-72.
Burton, W.G. (1957) The dormancy and sprouting of potatoes. Fd Sci. Abslr., 29,
1-12.
Burton, W.G. (1958) The effect of the concentrations of carbon dioxide and
oxygen in the storage atmosphere upon the sprouting of potatoes at 10c e. Eur.
Potato 1., 1(2), 47-57.
Burton, W.G. (1961) The physiology of the potato: problems and present status.
Proc. 1st Trienn. Can! EAPR, Braunschweig, 1960, pp. 79-117.
Burton, W.G. (1963). Concepts and mechanism of dormancy, in The Growth of the
Potato (eds J.D. Ivins and F.L. Milthorpe), Butterworths, London, pp. 17-41.
Burton, W.G. (1964) The respiration of developing potato tubers. Eur. Potato 1.,
7, 90-101.
Burton, W.G. (1965) The sugar balance in some British potato varieties during
storage. I. Preliminary observations. Eur. Potato 1., 8, 80-91.
Burton, W.G. (1966) The Potato, 2nd edn, H. Veenam and Zonen, Wageningen.
Burton, W.G. (1969) The sugar balance in some British potato varieties during
storage. II. The effects of tuber age, previous storage temperature, and
intermittent refrigeration upon low-temperature sweetening. Eur. Potato 1., 12,
81-95.
Burton, W.G. (1972) The response of the potato plant and tuber to temperature, in
Crop Processes in Controlled Environments (eds A.R. Rees, K.E. Cockshull,
D.W. Hand and R.G. Hurd), Academic Press, London, pp. 217-33.
Burton, W.G. (1973) Physiological and biochemical changes in the tuber as
affected by storage conditions. Proc. 5th Trienn. Cant EAPR, Norwich 1972,
pp.63-81.
Burton, W.G. (1974) The oxygen uptake, in air and in 5% O 2 , and the carbon
dioxide output, of stored potato tubers. Potato Res., 17, 113-37.
Burton, W.G. (1978a) Post-harvest behaviour and storage of potatoes, in Applied
Biology, Vol III (ed. T.H. Coaker), Academic Press, London, New York, San
Francisco, pp. 86--228.
Burton, W.G. (1978b) Biochemical and physiological effect of modified atmospheres
and their role in quality maintenance, in Post-harvest Biology and Bio-
technology (eds H.D. Hultin and M. Milner), Food and Nutrition Press Inc.,
Westport, Connecticut, USA, pp. 97-110.
Burton, W.G. (1982) Post-harvest Physiology of Food Crops, Longman, London
and New York.
Burton, W.G. and Hannan, R.S. (1957) Use of y-radiation for preventing the
sprouting of potatoes. 1. Sci. Fd Agric., 12, 707-15.
Burton, W.G. and Mann, G. (1954) Late storage of potatoes in England and
Wales. Agriculture, 60, 466--72.
Burton, W.G. and Meigh, D.F. (1971) The production of growth-suppressing
volatile substances by stored potato tubers. Potato Res., 14, 96--101.
Burton, W.G. and Wigginton, M.J. (1970) The effect of a film of water upon the
oxygen status of a potato tuber. Potato Res., 13, 180--6.
Burton, W.G. and Wilson, A.R. (1978) The sugar content and sprout growth of
tubers of cv. Record, grown in different localities when stored at 100, 20 and
20°C. Potato Res., 21, 146--62.
Burton, W.G., Mann, G. and Wager, H.G. (1955) The storage of ware potatoes in
permanent buildings. II. The temperature of unventilated stacks of potatoes. 1.
Agric. Sci., 46, 150--63.
Burton, W.G., Horne, T. and Powell, D.B. (1959) The effect ofy-irradiation upon
the sugar content of potatoes. Eur. Potato 1.,2, 105-16.
Bushway, R.J., Bureau, J.L. and McGann, D.F. (1984) Determinations of organic
acids in potatoes by high perlonnance liquid chromatography. 1. Food Sci., 49, 75-81.
Caldwell, J.S., Brunstetter, C.W. and Ezell, B.D. (1945) Causes and control of
discoloration in dehydration of white potatoes. Canner, 100(13), 35; 100(14),
15; 100(15), 14.
References 713
Carlsson, H. (1967) The relationship between specific gravity, dry matter and
starch in potatoes. Lantbr. Hoegsk. Ann., 33, 695-702.
Carter, J.N. and Bosma, S.M. (1974) Effect of fertilizer and irrigation on nitrate
nitrogen and total nitrogen in potato tubers. Agron. J., 66, 263-6.
Chaika, M.T., Savchenko, G.E. and Maister, A. (1980) Characteristics of the
formation of a protochlorophyll pigment in greening potato tubers. Dokl.
Akad. Nauk. BSSR, 24, 457-60.
Chaube, S., Swinyard, e.A. and Daines, R.H. (1973) Failure to induce malforma-
tions in fetal rats by feeding blighted potatoes to their mothers. Lancet, i, 329-30.
Cherif, A. (1973) Metabolisme des lipides dans Ie tubercule de pomme de terre
(Solanum tuberosum L). Evolution des lipides au cours de la conservation des
tubercules. Potato Res. 16, 126-47.
Chin, C.K. and Frenkel, Ch. (1977) Upsurge in respiration and peroxide formation
in potato tubers as influenced by ethylene, propylene, and cyanide. Plant
Physiol. 59, 515-18.
Cho, J.L., Iritani, W.M. and Martin, M.W. (1983) Comparison of methods for
measuring dormancy of potatoes. Am. Potato J., 60, 169-77.
Chung, C.H. (1985) Effects of temperature and chloropropham on phosphofructo-
kinase, mitochondrial respiration and reducing sugars in dormant Nooksack
potato tubers, Ph.D. Thesis, University of Missouri, Columbia, USA.
Coleman, W.K. (1987) Dormancy release in potato tubers: a review. Am. Potato
J., 64, 57-68.
Coleman, W.K. and King, R.R. (1984) Changes in endogenous abscisic acid,
soluble sugars and proline levels during tuber dormancy in Solanum tuberosum
L. Am. Potato J., 61, 437-49.
Conner, H. W. (1937) Effect of light on solanine synthesis in the potato tuber. Plant
Physiol., 12, 79-98.
Cornforth, J.W., Milborrow, B.V. and Ryback, G. (1966) Identification and
estimation of (+) abscisin II (dormin) in plant extracts by spectropolarimetry.
Nature, 210, 627-8.
Corsini, D., Stallknecht, G. and Sparks, W. (1979) Changes in chlorpropham
residues in stored potatoes. Am. Potato J., 56, 43-50.
Craft, C.e. (1963) Respiration of potatoes as influenced by previous storage
temperature. Am. Potato J., 40, 289-98.
Cronk, T.C., Kuhn, G.D. and McArdle, F.J. (1974) The influence of stage of
maturity, level of nitrogen fertilization and storage on the concentration of
solanine in tubers of three potato cultivars. Bull. Environm. Contam. ToxicoL,
11(2), 163-8.
Dalziel, J. and Duncan H.J. (1980) Studies on potato sprout suppressants. 4. The
distribution of tecnazene in potato tubers and the effect of processing on residue
levels. Potato Res., 23, 405-11.
Davidson, T.M.W. (1958) Dormancy in the potato tuber and the effects of storage
conditions on initial sprouting and on subsequent sprout growth. Am. Potato J.,
35,451-65.
Davies, A.M.e. (1977) The free amino acids of potato varieties grown in England
and Ireland. Potato Res. 20, 9-21.
Davis, R.M. (1961) Ingrown sprouts in potato tubers: factors accompanying their
origin in Ohio. Am. Potato J., 38, 411-13.
Dean B.B., Kolattukudy, P.E. and Davis, R.W. (1977) Chemical composition and
ultrastructure of suberin from hollow heart tissue of potato tubers (Solanum
tuberosum). Plant Physiol., 59,1008-10.
Denny, F.E. (1926) Hastening the sprouting of dormant potato tubers. Am. J.
Bot., 13, 118-25.
Denny, F.E. (1945) Synergistic effects of three chemicals in the treatment of
dormant potato tubers to hasten germination. Contr. Boyce Thompson Insl. Pl.
Res., 14, 1-14.
Dent, T.M. (1985) Review of current usage of pesticide chemicals for the control of
post-harvest losses in stored potatoes. Chemistry and Industry, Feb., 84-7.
Desborough, S. and Weiser, c.J. (1974) Improving potato protein I. Evaluation of
selection techniques. Am. Potato J., 51, 185-96.
Dizengremel P. (1985) Potato respiration: electron transport pathways, in Potato
Physiology (ed. P.H. Li), Academic Press, New York, London, pp. 60--109.
Drauschke, W., Steinbeiss, C.D. and Hohnstadter, J. (1981) Untersuchungen zum
Einsatz von Nonylalkohol als temporarer Keimungsinhibitor bei Pflanzkartoffeln.
2. Mitt. Makroautoradiografische Untersuchungen der mit 14C-markiertem
Nonylalkohol behandelten Pflanzkartoffeln. Nahrung, 25, 35-8.
Duncan, H.J. (1984) Recent developments on sprout suppressant chemicals at
Glasgow. Proc. 20th Ann. Conf. PCIRG, Heelsum (Neth.), pp. 47-9.
Duncan, H.J. and van Es, A. (1988) An assessment of DMN as a sprout
suppressant treatment for stored potatoes. Proc. 22nd Ann. Conf. ECSA-PRC,
Heelsum (Neth.), pp. 110--23.
Dwelle, R.B. and Stallknecht, G.F. (1978) Respiration and sugar content of potato
tubers as influenced by storage temperature. Am. Potato J., 55, 561-71.
Effmert, B., Meinl, G. and Vogel, J. (1961) Atmung, Zuckerspiegel und Ascor-
binsaure - Gehalt von Kartoffelsorten bei verschiedenen Lagertemperaturen.
Zuechter, 31, 23-32.
El-Antably, H.M.N., Wareing, P.F. and Hillmann, J. (1967) Some physiological
responses to d; I abscisin (dormin). Planta, 73, 74-90.
Elwood, J.H. and MacKenzie, G. (1973) Associations between the incidence of
neurological malformations and potato blight outbreaks over 50 years in
Ireland. Nature, 243, 476--7.
Emanuel, I. and Sever, C.E. (1973) Questions concerning the possible association
of potatoes and neural-tube defects, and an alternative hypothesis relating to
maternal growth and development. Teratology, 8, 325-31.
Emilsson, B. (1949) Studies on the rest period and dormant period in the potato
tuber. Acta Agric. Suec., 3, 189-284.
Espelie, K.E., Franceschi, V.R. and Kolattukudy, P.E. (1986) Immunocyto-
chemical localization and time course of appearance of an anionic peroxidase
associated with suberization in woundhealing potato tuber tissue. Plant Physiol.,
81,487-92.
Farrimond, A. (1973) Heat transmission in and from potatoes tmder different air
velocities. Proc. 5th Trienn. Conf. EAPR, Norwich, 1972, p. 135.
Filmer, A.A.E. and Rhodes, M.J.C. (1984) An assessment of 1,4, 6-trimethyl-
naphthalene as a sprout .suppressant for stored potato tubers. Potato Res., 27,
383-92.
Fitzpatrick, T.J. McDermott, J.A. and Osman, S.F. (1978). Evaluation of injured
commercial potato samples for total glycoalkaloid contents. J. Food Sci. 43,
1617-18.
References 715
Frydecka-Mazurczyk, A. (1978) The effect of thermal conditions during storage on
respiration in potato tubers. Biuletyn Institutu Ziemniaka, 22, 113-24.
Gager, G.S. (1912) Ingrowing sprouts of Solanum tuberosum. Bot. Gaz., 54,
515-24.
GaJliard, T. (1973) Lipids of potato tubers. I. Lipid and fatty acid composition of
tubers from different varieties of potato. J. Sci. Fd Agric., 24, 617-22.
Goodwin, P.B. (1966) The effect of water on dormancy in the potato. Eur. Potato
J., 9, 53--64.
Gray, D. and Hughes, J.C. (1978) Tuber quality, in The Potato Crop, (ed. P.M.
Harris), Chapman and Hall, London, pp 504-44.
Green, W.P. Hukill, W.V. and Rose, D.H. (1941) Calorimetric measurements of
the heat of respiration of fruits and vegetables. Technical Bulletin US Depart-
ment Agric., 771, p. 22.
Gull, D.D. (1960) Chlorophyll and solanine changes in tubers of Solanum
tuberosum induced by fluorescent light and a study of solanine toxicology by
bioassay technique, Ph.D. Thesis, Cornell Uaiversity.
Hagberg, A. and Nylsom, N. (1954) Reaction of potatoes to X-radiation and radio
phosphorus. Acta Agric. Scand., 4, 578.
Hair, B.L. and Gould, W.A. (1979) Reconditioning of cold-stored potato tubers as
influenced by harvest maturity. Research Circular Ohio Agric. Research and
Development Center, no. 250, pp. 35-7.
Hajslova, J. and Davfdek, J. (1986) Sprout inhibitors IPC and CIPC in treated
potatoes. Nahrung, 30, 75-9.
Hak, P.S. (1980) Het reconditioneren van chipsaardappelen. IBVL-Publikatie 331,
Wageningen.
Hak, P.S. and Vermeer, H. (1979) Bewaring van aardappelen bij hoge tempera-
turen (with English summary). IBVL-Rapport 236, Wageningen.
Hammes, P.S. and Nel, P.e. (1975) Control mechanisms in the tuberization.
process. Potato Res., 18,262-72.
Hanes, C.S. and Barker, J. (1931) The physiological action of cyanide. I. The
effects of cyanide on the respiration and sugar content of the potato at 15°e.
Proc. R. Soc. B. 108, 95-118.
Hannan, R.S. (1954) The preservation of food with ionizing radiations. Fd Sci.
Abstr., 26, 121--6.
Harborne, J.B. (1962) The plant polyphenols. VI. The flavonol glycosides of wild
and cultivated potatoes. Biochem. J., 84, 100--6.
Harkett, P.J. (1975) The influence of temperature and skin colour on chlorophyll
synthesis in potato tubers exposed to light. Proc. 6th Triennial Conf. EAPR,
Wageningen, 1975, pp. 179-80.
Harkett, P.J. (1981) External factors affecting length of dormant period in potato.
J. Sci. Fd Agric., 32, 102-3.
Hartmans, K.J. and van Es, A. (1986) Sprout inhibitors for potatoes. Proc. 21st
Ann. Conf. ESCA-PRC, Heelsum (Neth.), pp. 110-18.
Hartmans, K.J. and van Es, A. (1988) Alternative sprout inhibitors. Proc. 22nd
Ann. Conf. ECSA-PRC, Heelsum (Neth.), pp. 102-3.
Heikes, D.L. (1985) Mass spectral identification of a metabolite of chlorpropham
in potatoes. J. Agric. Fd Chern., 33, 246--9.
Heilinger, F. (1961) Stofftiche Veraenderungen in der Kartoffelknolle wahrend
der Lagerung. Proc. 1st Trienn. Conf. EAPR, Braunschweig, 1960, pp. 241-2.
Heilinger, F. and Breyhan, Th. (1959) Zur Kenntnis der Aminosaeuren in
Kartoffelknollen. Landbauforschung Volkenrode, 9, 17-18.
Heisler, E.G., Siciliano, S., Krulick, S., Porter, W.L. and White, J.R. Jr. (1973)
Nitrate and nitrite content of market potatoes. 1. Agric. Fd Chem., 21, 970--3.
Hemberg, T. (1970) The action of some cytokinins on the rest period and the
content of acid growth-inhibiting substances in potato. Physiol. Plant, 23,
850-8.
Hemberg, T. (1985) Potato rest. in Potato Physiology (ed. P.H. Li), Academic
Press, New York, London, pp. 353-88.
Holmes, J.C., Lang, R.W. and Sing, A.K. (1970) The effect of five growth
regulators on apical dominance in potato seed tubers and on subsequent tuber
production. Potato Res., 13, 342-52.
Holst, V.B. (1971) Some properties of inhibitor-p from Solanum tuberosum
compared to abscisic acid. Physiol. Plant., 24, 392-6.
Houghland, G.V.e. (1966) New conversion table for specific gravity, dry matter
and starch in potatoes. Am. Potato 1., 43, 138.
Huelin, F.E. and Barker, J. (1939) The effect of ethylene on the respiration and
carbohydrate metabolism of potatoes. New Phytol., 38,85-104.
Hughes, J.C. (1963) Blackening of plant tissues, in Report Low Temperature Res.
Stn, Cambridge, 1962, pp. 23-5.
Hughes, J.C. and Fuller, T.J. (1984) Fluctuations in sugars in cv. Record during
extended storage at 100e. Potato Res., 27, 229-36.
Hunter, J.H. (1985) Simulated external heat and moisture balance in potato
storages. Transactions of the ASAE, 28(4), 1279-83.
Hunter, J.H. (1986) Heat of respiration and weight loss from potatoes in storage, in
Engineering for Potatoes (ed. B.F. Cargill), Publ. Michigan State Vniv. and
ASAE, pp. 511-50.
Hylmo, B., Persson, T., Wikberg, C. and Sparks, W.C. (1975a) The heat balance
in a potato pile. I. The influence of the latent heat of the removed water. Acta
Agric. Scand., 25, 81-7.
Hylmo, B., Persson, T., Wikberg, C. and Sparks, W.C. (1975b) The heat balance
in ventilation. Acta Agric. Scand., 25, 88-91.
Iritani, W.M. and Weller, L.D. (1978) Factors influencing reconditioning of Russet
Burbank potatoes. Am. Potato 1., 55, 425-30.
Isherwood, F.A. (1973) Effects of changes in storage temperature on the meta-
bolism of the potato tuber. Proc. 5th Trienn. Conf. EAPR, Norwich, 1972, pp
156--7.
Isherwood, F.A. (1976) Mechanism of starch-sugar interconversion in Solanum
tuberosum. Phytochemistry, 15, 33-41.
Isherwood, F.A. and Burton, W.G. (1975) The effect of senescence, handling,
sprouting and chemical sprout suppression upon the respiratory quotient of
stored potato tubers. Potato Res., 18, 98-104.
Isola, M.e. and Franzoni, L. (1986) Mechanism of the increase in ribonuclease
activity in potato tuber slices. Plant Cell Physiol., 27, 331-5.
Ives, J.V. (1955) An abnormal form of skin spot on potatoes. Plant. Path., 14,
17-21.
Jacob, W.C. (1959) Studies on the internal blackspot of potatoes, Cornell Vniv.
Agric. Expt. Sta. Mem. 368, pp. 1-86.
Jadhav, S.J. and Salunkhe, K.D. (1975) Formation and control of chlorophyll and
References 717
glycoalkaloids in tubers of Solanum tuberosum L. and evaluation of glycoalkaloid
toxicity. Adv. Food Res., 21, 307-54.
ladhav, S.l., Sharma, R.P. and Salunkhe, K.D. (1981) Naturally occurring toxic
alkaloids in foods. CRC Crit. Rev. Toxicol., 9, 21-104.
ladhav, S.l., Sikchi, P.G. Kadam, S.S., Chavan, 1.K. and Salunkhe, D.K. (1984)
Effects of mechanical injury on ascorbic acid content of potato tubers. Indian
Food Packer, 38(5), 66-9.
Jeppsen, R.B., Wu, M.T., Salunkhe, D.J. and Jadhav, S.Y. (1974) Some observa-
tions on the occurrence of chlorophyll and solanine in potato tubers and their
control by Ni-benzyladenine, ethephon and filtered light. J. Food Sci., 39,
1059-61.
Johnson, G. and Schaal, L.A. (1957) Chi orogenic acid and other orthodihydric
phenols in scab-resistant Russet Burbank and scab-susceptible Triumph potato
tubers of different maturities. Phytopathology, 47, 253-5.
Kader, A.A. (1987) Respiration and gas exchange of vegetables, in Post-harvest
Physiology o/Vegetables (ed. J. Weichmann), Marcel Dekker, New York and
Basel, pp. 25-43.
Kahl, G. (1973) Genetic and metabolic regulation in differentiating plant storage
tissue cells. Botanical Review, 39, 274-99.
Kahl, G. (1974) Metabolism in plant storage tissue slices. Botanical Review, 40,
263-314.
Kapoor, A.c., Desborough, S.L. and Li, P.H. (1975) Potato tuber proteins and
their nutritional quality. Potato Res., 18,469-78.
Kawakami, K. (1952) Physiological aspects of potato seed tubers. Mem. Hyogo
Agric. College. (Sassayama, Japan), 2, 1-114.
Keller, E.R. and Berces, S. (1966) Check-testing for virus Y and leaf-roll in seed
potatoes with particular reference to methods of increasing precision with the
Ab-leaf test for virus Y. Eur. Potato J., 9, 1-14.
Kolattukudy, P.E. (1984) Biochemistry and function of cutin and suberin. Can. J.
Botany, 62, 2918-33.
Korableva, N.P., Morozova, E.V. and Metlitskii, LV. (1977) Role of phenolic
and terpenoid inhibitors in regulation dormancy of potato, in Plant Growth
Regulation. Proc. 2nd Int. Symp., Sofia, (eds Kudrev, Ivanova and Karanov)
pp.224-7.
Korableva, N.P., Karavaea, K.A. and Metlitskii, L.V. (1980) Changes of abscisic
acid levels in potato tuber tissues during deep dormancy and germination.
Fiziol. Rast., 27, 585-91.
Krijthe, N. (1946) De invloed van de bewaring van aardappelknollen op de bouw
van de knoppen en op de ontwikkeling tot volwassen plant. Meded. Land-
bouwhogeschool, 47, 3-36.
Krijthe, N. (1962) Observations on the sprouting of seed potatoes. Eur. Potato J.,
5,316-33.
Krijthe, N. (1977) Onderzoek over de groei oans Kiemen op poot aardappelen.
Publikatie 295, IBVL, Wageningen.
Kuroki, G. and Conn, E.E. (1988) Increased chorismate mutase levels as a response
to wounding in Solanum tuberosum L. tubers. Plant Physiol., 86, 895-8.
Kushman, L.J. and Haynes, Jr, F.L. (1971) Influence of intercellular space
differences due to variety and storage upon tuber-specific gravity-dry matter
relationship. Am. Potato J., 48, 173-81.
Lampitt, L.H. and Goldenberg, N. (1940) The composition of the potato. Chem.
Ind., 748-61.
Lange, H. (1970) Atmungswege bei vernarbenden und proliferierenden Gewebe-
fragmenten der Kartoffelknolle. Planta, 90, 119-32.
Lange, H. and Rosenstock, G. (1963) Physiologisch-Anatomische Studien zum
Problem des Wundreizes. II. Kausalanalytische Untersuchungen zur Theorie
des Wundreizes. Beitr. BioI. Pfl., 39, 383-488.
Lange, H. and Rosenstock, G. (1964) Ueber Alterserscheinungen bei traumatisch
induzierter meristematischer Aktivitiit. Phytopathol. z., 51, 136-52.
Lange, H., Rosenstock, G. and Kahl, G. (1970) Induktionsbedingungen der
Suberinsynthese und Zellproliferation bei Parenchymfragmenten der Kartof-
felknolle. Planta, 90, 109-18.
Laties, G.G. (1978) The development and control of respiratory pathways in slices
of plant storage organs, in Biochemistry of Wounded Plant Tissues (ed. G.
Kahl), Walter de Gruyter, Berlin, New York, pp. 421-66.
Laties, G.G. (1982) The cyanide-resistant, alternative path in higher plant respira-
tion. Ann. Rev. Plant Physiol., 33, 519-55.
Liljemark, A. and Widoff, E. (1960) Greening and solanine development of white
potatoes in fluorescent light. Am. Potato J., 37, 379-89.
Linnemann, A.R., van Es, A. and Hartmans, K.J. (1985) Changes in the content
of L-ascorbic acid, glucose, fructose, sucrose and total glycoalkaloids in
potatoes (cv. Bintje) stored at 7, 17 and 28°C. Potato Res., 28, 271-8.
Lipton, W.J. (1967) Some effects of low-oxygen atmospheres on potato tubers.
Am. Potato J., 44, 292-9.
Lougheed, E.e., Rowberry, R.G. and Franklin, E.W. (1975) Ethylene increases
respiration of sprout-inhibited potatoes. Naturwiss., 62, 185-6.
Ludwig, J.W. (1970) De Aardappel en aardappelprodukten. Syllabus van de
studiestof voor de MO-akte NI8, IBVL-Wageningen.
Ludwig, J.W. (1972) Bepaling van het droge-stof genalte van aardappelen via
onderwaterweging. IBVL-Publikatie 247, Wageningen.
Lyon G.D. (1972) Occurrence of rishitin phytuberin in potato tubers inoculated
with Erwinia carotovora vaL atroseptica. Physiol. PI. Path., 2, 411-16.
McDole, R.E. and McMaster, G.M. (1978) Effects of moisture stress and nitrogen
fertilization on tuber nitrate-nitrogen content. Am. Potato J., 55, 611-19.
McGee, E., Jarvis, M.C. and Duncan, H.J. (1985a) Wound healing in potato tuber
tissue. I. Effects of maturity at harvest. Potato Res., 28, 91-9.
McGee, E., Jarvis, M.C. and Duncan, H.J. (1985b) Wound healing in potato tuber
tissue. 2. Varietal and anatomical variation. Potato Res., 28, 101-8.
McGee, E., Jarvis, M.e. and Duncan, H.J. (1986) The relationship between
temperature and sprout growth in stored seed potatoes. Potato Res., 29, 521-4.
McGee, E., Jarvis, M.e. and Duncan, H.J. (1987) Wavelength dependence of sup-
pression of potato sprout growth by light. Plant, Cell and Environment, 10, 655-60.
Maga, J.A. (1980) Potato glycoalkaloids. CRC Crit. Rev. Fd Sci. Nutr., 12,371-405.
Maine, M.J. de, Bain, H. and Joyce, J.A.L. (1988) Changes in the total tuber
glycoalkaloid content of potato cultivars on exposure to light. J. Agric. Sci.
Camb., 111, 57-8.
Malmberg, A. and Theander, O. (1984) Free and conjugated phenolic acids and
aldehydes in potato tubers. Swedish J. Agric. Res., 14, 119-25.
Malmberg, A., von Rosen, G., Schatz, B.A. and Theander, O. (1980) Antifungal
References 719
activity of phenolic compounds against Phoma exigua var. foveata in potato
tubers. Swedish l. Agric. Res., 10, 89-95.
Mann, G. (1963) The cooling of potatoes in bulk. Ann. Report Ditton and Convent
Garden Laboratories, 1962-3, pp. 30-I.
Mapson. L.W. and Burton, W.G. (1962) The terminal oxidases of the potato
tuber. Biochem. l. 182, 19-25.
Marinus, J. and Bodlaender, K.B.A. (1978) Growth and yield of seed potatoes
after application of gibberellic acid to the tubers before planting. Neth. l. Agric.
Sci., 26, 354-65.
Mazza, G., Hung, J. and Dench, M.J. (1983) Processing/nutritional quality
changes in potato tubers during growth and long term storage. Can. Inst. Food
Sci. Technol. l., 16, 39-44.
Meeuwse, B.J.D. (1975) Thermogenic respiration in aroids. Ann. Rev. Plant
Physiol., 26, 117-26.
Meigh, D.F., Filmer, A.A.E. and Self, R. (1973) Growth inhibitory volatile
aromatic compounds produced by Solanum tuberosum tubers. Phytochemistry,
12,987-93.
Meijers, C.P. (1972) Effect of carbon disulphide on the dormancy and sprouting of
seed potatoes. Potato Res., 15, 160-5.
Meinl, G. (1966) Untersuchungen zur Sorten- und Umweltbedingten Variation der
Lentizellen- anzahl von Kartoffelknollen. Flora, 156B, 419-26.
Meinl, G. (1967) Zur Bezugsgrosse der Respirationsintensitat von Kartoffel-
knollen. Eur. Potato l., 10, 249-57.
Michaels, W.H. (1932) Relation of lenticels and surface area to respiration in the
potato tuber. Bot. Gaz., 94, 416-18.
Milborrow, B.V. (1967) The identification of (+) abscisin «+)-dormin) in plants
and measurement of its concentrations. Planta, 76, 93-113.
Miller, R.A., Harrington, J.D. and Kuhn, G.D. (1975) Effect of variety and
harvest date on tuber sugars and chip color. Am. Potato l., 52, 379-86.
Misener, G.c. (1986) Airflow resistance due to soil on bulk potatoes. Can. Agric.
Eng., 28,43-4.
Mondy, N.I. and Leja, M. (1986) Effect of mechanical injury on the ascorbic acid
content of potatoes. l. Food Sci., 51, 355-7.
Mottley, J. (1980) Influence of light-induced greening on the browning components
of different varieties of potato tubers. Luso, 1, 1-8.
Mulder, E.G. and Bakema, K. (1956) Effect of the nitrogen, phosphorus potas-
sium and magnesium nutrition of potato plants on the content of free amino-
acids and on the amino-acid composition of the protein of the tubers. Plant and
Soil, 7, 135-66.
Muller, K. (1846) Zur Biologie der Kartoffeln. Bot. Zg., 4, 769-72.
Muller, K. (1975) Veranderungen wertgebender Inhaltsstoffe in der Kartoffel-
planze und -knolle im Verlauf von Vegetation und Lagerung, ihre Bedeutung
fUr die Qualitat der Knolle. Hefte fur den Kartoffelbau, 17, Schriftenreihe
Forderungsgemeinschaft Kartoffelwirtschaft e.V., Hamburg.
Muller-Thurgau, H. (1882) Ueber Zuckeranhaufung in Pflanzentheilen in Folge
niederer Temperatur. Landw. lb., 11, 751-828.
Muneta, P. and Kaisaki, F. (1985) Ascorbic acid-ferrous iron (Fe 2+)-
complexes and after-cooking darkening of potatoes. Am. Potato 1., 62,
531-6.
Munzert, M. and Lepschy, J. (1983) Zur Frage des NitratgehaItes in Kartoffel-
knollen. Kartoffelbau, 34, 163-8.
Nagy, D., Weidlein, W. and Hixon, R.H. (1941) Factors affecting the solubility of
corn protein. Cereal Chern. 18, 514-22.
Nash, M.J. and Lennard, J.H. (1973) Forced-draught ventilation with cold outside
air compared with recirculation of artificially cooled air. Potato Res., 6, 47-52.
Nielsen, N.K. (1968) An investigation of the regenerative power of periderm in
potato tubers after wounding. Acta Agric. Scand. 18, 113-20.
Nitsch, A. and Klein, K. (1983) Ertrage und innere Qualitaten der Kartoffel in
Abhaengigkeit von der Stickstoffdungung. Kartoffelbau, 34, 30-4.
Nnodu, E.C., Harrison, M.D. and Parke, R.V. (1982) The effect of temperature
and relative humidity on wound healing and infection of potato tubers by
Alternaria solani. Am. Potato J., 59, 297-311.
Nolte, Ph., Secor, G.A. and Gudmestad, N.C. (1987) Wound healing, decay and
chemical treatment of cut potato tuber tissue. Am. Potato J., 64, 1-9.
Okazawa, Y. (1959) Studies on the occurrence of natural gibberellin and its effects on
the tuber formation of potato plants. Proc. Crop. Sci. Soc. Japan, 28, 129-33.
Olssen, K. (1986) The influence of genotype on the effects of impact damage on the
accumulation of glycoalkaloids in potato tubers. Potato Res., 29, 1-12.
Olufsen, L. (1903) Untersuchungen ueber Wundperidermbildung in Kartoffel-
knollen. Bot. Zentralblatt., 15, 269-308.
Ophuis, B.G. (1957) The effect of ventilation capacity on weight losses in
ventilated potato stores. Neth. J. Agric. Sci., 5, 180-94.
Oser, B.L. (1959) An integrated essential amino acid index for predicting the
biological value of proteins, in Protein and Amino Acid Nutrition (ed. A.A.
Albanese), Academic Press, New York, pp. 281-95.
Ozeretskovskaya, O.L. and Vasyukova, N.!. (1965) New formation of phenols in
damaged tissues of potato. Dokl. Akad. Nauk. SSSR, 161,968-70.
Patzold, C. and Kolb, W. (1957) Beeinflussung der Kartoffel (Solanum tuberosum
L.) und der Topinambur (Heilianthus tuberosus L.) durch Roentgenstrahlen.
Beitr. Bioi. Pfl., 33, 437-58.
Patzold, C. and Weiss, H.M. (1957) Beeinflussung der Kartoffelknolle durch
Gammastrahlen radioaktiven Kobalts (C0 60). Angew. Bot., 31, 93-105.
Painter, e.G. and Augustin, J. (1976) The effect of soil moisture and nitrogen on
yield and quality of the Russet Burbank potato. Am. Potato J., 53, 275-84.
Parnell, A., Bhuva, V.S. and Bintdiffe, E.J.B. (1984) The gJycoaJkoJoid content of
potato varieties. Nat. Inst. Agric. Bot., 16, 535-41.
Paupardin, C. and Tizio, R. (1970) Action de quelques composes phenoliques sur
la tuberisation de Ja pomme de terre. Potato Res. 13, 187-98.
Penner, H. and Fromm, H. (1972) Versuche zum Nachweis einer erfolgten
Bestrahlung von Kartoffeln. II. Bestimmung der Chlorogensaure. Z. Lebens.
Unters. Forsch., 150, 84-7.
Perez-Trejo, M.S., Janes, H.W. and Frenkel, Ch. (1981) Mobilization of respira-
tory metabolism in potato tubers by carbon dioxide. Plant Physiol, 67, 514-17.
Petermann, J.B. and Morris, S.e. (1985) The spectral responses of chlorophyll and
glycoalkaloid synthesis in potato tubers (Solanum tuberosum). Plant Science,
39, 105-10.
Peterson, e.L., Wyse, R. and Neuber, H. (1981) Evaluation of respiration as a tool in
predicting internal quality and storability of potatoes. Am. Potato J., 58, 245-56.
References 721
Pol, G. and Labib, A.1. (1963) De voedingswaarde van aardappelen van verschil-
lende rassen en de invloed daarop van bemesting en bewaring. Voeding. 83,
479-99.
Politis, J. (1948) Du role de l'acide chI oro geni que dans la formation des mem-
branes suberifIees. Revue Cytol. Cytophys. Veget., 10, 232-3.
Ponnampalam, R and Mondy, N.1. (1983) Effect of cooking on the total
glycoalkaloid content of potatoes. f. Agric. Fd Chem., 31. 493-5.
Ponnampalam, R and Mondy, N.1. (1985) Effect of sprout inhibitors and nitrogen
fertilization on nitrate-N content of potato tubers. f. Food Sci., 30, 1246--8,
1256.
Porter, W.L., Fitzpatrick, T.J. and Talley, E.A. (1964) Studies on the relationship
of specific gravity to total solids of potatoes. Am. Potato f., 41, 329-36.
Potts, M.J., Albert, W.V.D., Rutab, F.R et al. (1983) An agro-economic
assessment of seed-potato storage technologies in the Philippines. Am. Potato
f., 60, 199-211.
Priestley, J.H. and Woffenden, L.M. (1923) The healing of wounds in potato
tubers and their propagation by cut sets. Ann. Appl. Bioi., 10, 96--115.
Putz, B. (1983) Kritische Betrachtungen der Bestimmung des Trockensubstanz -
und Starke - gehaltes von Kartoffeln. Starke, 35, 415-19.
Racca, RW. and Tizio, R (1968) A preliminary study of changes in the content of
gibberellin-like substances in the potato plant in relation to the tuberization
mechanism. Eur. Potato f., 11, 213-20.
Radatz, W. (1967) Die Wundkorkbildung der Kartoffelknolle in Abhangigkeit
von Lagerungsbedingungen. Landb. forsch. Voelkenrode, 17, 153-8.
Rappaport, L. and Sachs, M. (1967) Wound-induced gibberellins. Nature, 214,
1149-50.
Rappaport, L. and Wolf, N. (1969) The problem of dormancy in potato tubers and
related structures, in Dormancy and Survival, Symp. Soc. Exp. Bioi., 23, 219-40.
Rappaport, L. Lippert, L.F. and Timm, H. (1957) Sprouting, plant growth and
tuber production as affected by chemical treatment of white potato seed pieces.
Am. Potato f., 34, 254-60.
Rastovski, A. (1987) Heat balance in a potato store, in Storage of Potatoes (eds A.
Rastovski, A. van Es et al.), Pudoc, Wageningen, pp. 212-16.
Reeve, R.M., Forrester, L.J. and Hendel, C.E. (1963) Histological analysis of
wound-healing in potatoes treated to inhibit sprouting I. CIPC (isopropyl-N-
chlorophenyl carbamate) treatments. f. Food Sci., 28, 649-54.
variation within potatoes. II. Phenolics, enzymes and other minor components.
Am. Potato f., 46, 347-86.
Reid, M.S. and Pratt, H.K. (1972) Effects of ethylene on potato tuber respiration.
Renwick, J.H. (1972) Hypothesis: anencephaly and spina bifida are usually
preventable by avoidance of a specific but unidentified substance present in
certain potato tubers. Br. f. Social Preventive Med., 26, 67-89.
Reust, W. (1984) Physiological age of the potato. Potato Res., 27, 455-7.
Rexen, B. (1976) Studies of protein of potatoes. Potato Res., 19, 189-202.
Rhodes, M.J.C. and Wooltorton, L.S.C. (1978) The biosynthesis of phenolic
compounds in wounded plant storage tissues, in Biochemistry of Wounded Plant
Tissues (ed. G. Kahl), Walter de Gruyter, Berlin, pp. 243-86.
Ringel, B. (1976) Messungen zur Sauerstoffversorgung der Kartoffelknollen.
Nahrung, 20, 847-50.
Rodis, P. and Hoff, J.E. (1984) Naturally occurring protein crystals in the potato
(inhibitor of papain, chymopapain and ficin). Plant Physiol., 74, 907-11.
Rosa, J.T. (1923) Abbreviation of the dormant period in potato tubers. Proc. Am.
Soc. Hart. Sci., 20,180--7.
Rosenstock, G. (1963) Physiologische und anatomische Studien zum Problem
der Wundheilung. 1. Untersuchungen tiber Zusammenhange zwischen dem
Wassergehalt verletzten Kartoffelparenchyms und dessen Vernarbungsreak-
tionen, nebst Bemerkungen zur Theorie der Wundreizes. Beitr. BioI. Pjl., 38,
275-319.
Rosenstock, G., Kahl, G. and Lange, H. (1971) Beziehungen zwischen Entwick-
lungszustand und Wundatmung beim Speicherparenchym von Solanum tubero-
sum. Z. Pjlanzenphysiol., 64, 130--8.
Rumpf, G. (1972) Gaschromatographische Bestimmung loslicher Inhaltsstoffe in
bestrahlten und mit chemischen Keimhemmungsmitteln behandelten Kartoffeln.
Potato Res., 15,236-45.
Rychter, A. Janes, H.W. and Frenkel, C. (1978) Cyanide-resistant respiration in
freshly cut potato slices. Plant Physiol., 61, 667-8.
Rychter, A., Janes, H.W. and Frenkel, C. (1979a) Effect of ethylene and oxygen
on the development of cyanide-resistant respiration in whole potato mito-
chondria. Plant Physiol., 63, 149-51.
Rychter, A., Janes, H.W., Chin, c.K. and Frenkel, C. (1979b) Effect of ethanol,
acetaldehyde, acetic acid, and ethylene on changes in respiration and respira-
tory metabolites in potato tubers. Plant Physiol., 64, 108-11.
Sadler, E.M. (1961) Factors influencing the development of sprouts of the potato,
Ph.D. Thesis, University of Nottingham.
Salaman, R.N. (1926) Potato Varieties, Cambridge Univ. Press, Cambridge, 378
pp.
Salunkhe, D.K., Wu, M.T. and Jadhav, S.J. (1972) Effects of light and tempera-
ture on the formation of solanine in potato slices. 1. Food Sci., 37, 969-70.
Samotus, B., Niedzwiedz, M., Kolodziej, Z. et al. (1974) Storage and recondition-
ing of tubers of Polish potato varieties and strains. 1. Influence of storage
temperature on sugar level in potato tubers of different varieties and strains.
Potato Res., 17, 64--81.
Saunders, P. (1978) Phytohormones and bud dormancy, in Phytohormones and
Related Compounds - a comprehensive treatise, Vo!' 2 (eds D.S. Letham, P.B.
Goodwin and T.J.V. Higgins), Elsevier, Amsterdam, pp. 423--45.
Sawyer, R.L. (1956) Effect of irradiation on potatoes for processing. Proc. 7th.
Am. Potato Utilization Conf., pp. 3-5.
Sawyer, R.L. (1959) Sprout inhibition, in Potato Processing (eds W.F. Talburt and
O. Smith), Avi Pub!. Co., Westport, Conn, pp. 143-54.
Sawyer, R.L. (1961) Relation of chloro-IPC for sprout inhibition to internal
sprouting of potatoes. Am. Potato 1.,38,203-7.
Sawyer, R.L. and Dallyn, S.L. (1964) Internal sprouting of potatoes. Am. Potato
1., 41, 59--69.
Schippers, P.A. (1956a) De duur van de rustperiode van een 4O-tal aardappelrassen,
bewaard bij verschillende con stante temperaturen. Stichting voor Aardappel-
bewaring, Wageningen, Pub!. 112, serie A.
References 723
Schippers, P.A. (1956b) De invloed van de temperatuur op de duur van de
rustperlode. Stichting voor Aardappelbewaring, Wageningen, Pub!. 108, serle A.
Schippers, P.A. (1976) The relationship between specific gravity and percentage
dry matter in potato tubers. Am. Potato J., 53, 111-22.
Schippers, P.A. (1977) The rate of respiration of potato tubers during storage. II.
Results of experiments in 1972 and 1973. Potato Res., 20, 189-206.
Schwimmer, S. (1978) Enzyme action and modifications of cellular integrity in
fruits and vegetables: consequences for food quality during ripening, senescence
and processing, in Post-harvest Biology and Biotechnology (eds H.O. Hultin
and M. Milner), Food and Nutrition Press, Westport, pp. 317-47.
Sherman, M. and Ewing, E.E. (1983) Effects of temperature and low oxygen
atmospheres on respiration, chip colour, sugars, and malate of stored potatoes.
J. Am. Soc. Hort. Sci., 108, 129-33.
Short, J.L. and Shotton, F.E. (1970) Storage conditions affecting the sprouting of
seed potatoes and their yield. III. Temperature. Exp. Husbandry, 19, 69-77.
Sikka, L. and Bryan, J.E. (1980) Sprout stimulation of dormant intact tubers.
J.I.P.A., 7(3),107-18.
Sinden, S.L. (1972) Effect of light and mechanical injury on the glycoalkaloid
content of greening resistant potato tubers. Am. Potato J., 49, 368.
Sinden, S.L. and Sanford, L.L. (1981) Origin and inheritance of solamarine
glycoalkaloids in commercial potato cultivars. Am. Potato J., 58, 305-25.
Sinden, S.L., Deahl, K.L. and Aulenbach, B.B. (1976) Effect of glycoalkaloids
and phenolics on potato flavour. J. Food Sci., 41, 520-3.
Singh, B.N. and Mathur, P.B. (1937) Studies in potato storage. I. Investigation of
physiological and chemical changes during the development and ripening of
potato tubers. Ann. Appl. Bioi., 24, 469·,,74.
Sizer, e.E., Maga, J.A. and Craven, C.J. (1980) Total glycoalkaloids in potatoes
and potato chips. J. Agric. Fd Chern., 28, 578--9.
Smith, B.G. and Rubery, P.H. (1981) Changes in phenylalanine-ammonialyase
levels in excised potato tuber tissue: effects of the gaseous environment and
desensitization to cinnamate repression by in situ pre-incubation. Plant, Cell and
Environment, 4, 377-81.
Smith, W.L. and Smart, H.F. (1959) Relation of soft rot development to protective
barriers in Irish potato slices. Phytopathology, 45, 649-54.
Smith, C., Watt, M., Boyd, A.E.W. and Holmes, J.e. (1973) Anencephaly, spina
bifida, and potato blight in the Edinburgh area. Lancet, i, 269.
Solomos, T. (1977) Cyanide-resistant respiration in higher plants. Ann. Rev. Plant
Physiol., 28, 279-97.
Solomos, T. and Laties, G.G. (1975) The mechanism of ethylene and cyanide
action in triggering the rise in respiration in potato tubers. Plant Physio1., 55,
73-8.
Sowokinos, J.R. (1973) Maturation of Solanum tuberosum. I. Comparative sucrose
synthetase levels between several good and poor processing varieties. Am.
Potato J., 50, 234-47.
Sowokinos, J.R. (1977) Sucrose rating as an index of processing potato maturity.
Valley Potato Grower, Ann. Rep. 43, no. 101, p. 31.
Sparenberg, H. and Ulmann, R. (1973) Inhibition of sprouting in potatoes by
irradiation in the Netherlands: economic and commercial aspects. Comm. of the
EEC; Eurisotop Office Information Booklet 58.
Sparrow, A.H. and Christensen, E. (1954) Improved storage quality of potato
tubers after exposure to 60Co gammas. Nucleonics, 12(8), 16-17.
Stegemann, H. (1975) Properties and physiological changes in storage proteins, in
Chemistry and Biochemistry of Plant Proteins (eds J.B. Harborne and C.F. van
Sumere), Phytochem. Soc. Symp. II, Academic Press, London, New York, San
Francisco, pp. 71-87.
Stephen, N.H. and Duncan, H.J. (1984) Potato sprout suppressant activity of some
substituted naphthalenes. Proc. 9th Trienn. Conf. EAPR, Interlaken, 1984,
pp. 321-2.
Struckmeyer, B.E., Weis, G.G. and Schoenemann, J.A. (1981) Effect of two
forms of maleic hydrazide on the cell structure of the mid section stem and bud
ends of the cortical and peri medullary regions of Russet Burbank tubers. Am.
Potato 1., 58, 611-18.
Sussman, A.S. (1953) The effect of ionizing radiations upon the respiration and
oxidases of the potato tuber. 1. Cell. Camp. Physiol., 42, 273-84.
Synge, R.L.M. (1977) Free amino acids of potato tubers: a survey of published
results set out according to potato variety. Potato Res., 20, 1-7.
Szalai, I. (1957a) Physiological studies on potato plants. VI. The distribution of
free amino acids in potato tubers and their change during vernalization I.
Photometric determination of total amino acid content of potato tuber sections
with the ninhydrin reaction. Acta Bioi. (Acta Univ. szegediensis), 3, 33-40.
Szalai, I. (1957b) Physiologische Studien an Kartoffelknollen VIII. Die Verteilung
der freien Aminosauren in Kartoffelknollen und ihre Beeinflussung durch
'Jarowisation'. II. Papierchromatographische Untersuchungen der freien Amino-
sauren des Kartoffelsaftes. Acta Bioi. (Acta Univ. szegediensis), 3, 41-9.
Szalai, I. (1957c) Physiological studies on potato plants X. Changes of bound and
free tryptophan content in potato tubers during germination. Acta BioI. (Acta
Univ. szegediensis), 3, 51-4.
Szalai, I. (1959a) Physiologische Studien an Kartoffelknollen. Die Verteilung der
freien Aminosauren in den mit Rindite behandelten Kartoffelknollen in
verschiedenen Phasen der Keimung. II. Papierchromatograpische Unter-
suchungen der freien Aminosaeuren des Kartoffelsaftes. Acta Agron. Acad.
Sci. Hung. 9, 253-M.
Szalai, 1. (1959b) Tryptophane contents of new potato tubers formed by rindite in
the different phases of the germination (Physiological studies of the potato
XIV). Physiol. Plant., 12, 155-61.
Tagawa, T. and Okazawa, Y. (1953) Physiological and morphological studies on
potato plants. IX. Nitrogen metabolism in potato tubers during the storage
period. 1. Fac. Agric. Hokkaido (Imp) Univ., 50, 65-73.
Talburt, W.F. and Smith, O. (1967) Potato Processing, Avi Publishing Company,
Westport, Connecticut.
Talley, E.A., Fitzpatrick, T.J. and Porter, W.L. (1964) Chemical composition of
potatoes. IV. Relationship of the free amino acid concentrations to specific
gravity and storage time. Am. Potato 1., 41, 357-66.
Talley, E.A., Fitzpatrick, T.J. and Porter, W.L. (1970) Chemical composition of
potatoes. VIII. Effect of variety, location and year of growth on the content of
nitrogen compounds. Am. Potato I., 47,231-44.
Talley, E.A., Thoma, R.B. and Orr, P.H. (1984) Amino acid composition of
freshly harvested and stored potatoes. Am. Potato 1., 61, 267-79.
References 725
Taylorson, R.B. and Hendricks, S.B. (1977) Dormancy in seeds. Ann. Rev. Plant
Physiol., 28, 331-54.
Theologis, A. and Laties, G.G. (1978). Relative contribution of cytochrom.e-
mediated and cyanide-resistant electron transport in fresh and aged potato
slices. Plant Physiol., 62, 232-7.
Theologis, A. and Laties, G.G. (1982) Potentiating effect of pure oxygen on the
enhancement of respiration by ethylene in plant storage organs: a comparative
study. Plant Physiol., 69,1031-5.
Thomas, P. (1982) Wound-induced suberization and periderm development in
potato tubers as affected by temperature and gamma irradiation. Potato Res.,
25, 155-M.
Thomson, A.J., Hughes, J.e. and Starr, C. (1987) Breeding for long dormancy and
low reducing sugar content following extended storage at low temperature.
Proc. 10th Trienn. Conf. EAPR, Aalborg, pp. 22-3.
Timm. H., Rappaport, L., Bishop, J.e. and Hoyle, J.B. (1962) Sprouting, plant
growth and tuber production as affected by chemical treatment of white potato
seed pieces. IV. Responses of dormant and sprouting seed potatoes to gibberellic
acid. Am. Potato J., 39,107-15.
Timm, H., Hughes, D.L. and Weaver, M.L. (1986) Effect of exposure time of
ethylene on potato sprout development. Am. Potato J., 63, 655-64.
Tizio, R. (1966) Interaction du facteur radiculaire et de I'acide gibberelliqe sur la
croissance des stolons et la tuberisation. C.R. Hebd. Seanc. Acad. Sci. Paris,
262, ser. D., 767-70.
Tripathi, R.K. and Kahl, G. (1982) Stimulation of synthesis and translational
acitivity of polyadenylated messenger RNA in wounded potato tubers by 2, 4-
dichlorophenoxyacetic acid. Biochem. Biophys. Res. Comm., 106(4), 1218-25.
Twiss, P.T.G. (1963) Trials of new methods of applying sprout suppressants. Ann.
Rep. Ditton and Covent Garden Laboratories, Jan. 1962-May 1963, p. 21.
Varns, J.L. (1986) Carbon dioxide and wind monitoring in a commercial potato
. storage, in Engineering for Potatoes, (ed. B.F. Cargill) Pub!. Michigan State
University and ASAE, pp. 598-615.
Varns, J., Currier, W. and Kuc, J. (1971) Specificity of rishitin and phytuberin
accumulation by potato. Phytopathology, 61, 968-71.
Vendelbo, P. (1980) Styring of vegetabilielagring efter koncentration af dannede
gasser. Beretning no. 96, Bioteknisk lnst., Kolding.
Verma, S.C., Malhotra, V.P., Joshi, K.e. and Sharma, T.R. (1971) Specific
gravity and dry matter content of potato. Potato Res., 14, 94-5.
Verma, S.C., Sharma, T.R. and Verma, S.M. (1974a) Sucrose accumulation
during high temperature storage of potato tubers. Potato Res., 17,224-6.
Verma, S.e., Sharma, T.R. and Verma, S.M. (1974b) Effects of extended high
temperature storage on weight losses and sugar content of potato tubers. Indian
J. Agric. Sci., 44, 702-6.
Villa, L.G. and Bakker-Arkema, F.W. (1974) Moisture losses from potatoes
during storage. Paper 74-6510, Wintermeeting ASAE.
Wager, H.G., Burton, W.G. and Mann, G. (1952) The storage of ware potatoes in
permanent buildings. I. Storage in bins in a uninsulated store, 1946-7. J. Agric.
Sci., 42, 266-75.
Waggoner, P.E. (1955) Radiation and resistance of tubers to rot. Am. Potato J.,
32,448-50.
Wall, M.E., Wani, M.e., Cook, e.E., Palmer, KH., McPhail, AT. and Sin, G.A
(1966) Plant antitumor agents. I. The isolation and structure of camptothecin,
a novel alkaloidal leukemia and tumor inhibitor from Camp to theca acuminata.
J. Amer. Chem. Soc., 88, 3888-90.
Wang, e.Y., Buta, J.G., Moline, H.E. and Hruschka, H.W. (1980) Potato sprout
inhibition by Camptothecin, a naturally occurring plant growth regulator. J.
Amer. Soc. Hort. Sci., 105(1), 120-4.
Wareing, P.F., El Antably, H.M.M., Good, J. and Manuel, J. (1966) The possible
role and mode of action of abscisin (dormin) in the regulation of plant growth and
development. Wiss. Z. Univ. Rostock, Reihe Mathematik Naturwiss., 16, 667-72.
Warren, D.S. and Woodman, J.S. (1974) The texture of cooked potatoes; a review.
J. Sci. Fd Agric., 25, 129-38.
Wassink, E.C., Krijthe, N. and Scheer, C. van der (1950) On the effect of light of
various spectral regions on the sprouting of potato tubers. Proc. K. Ned. Akad.
Wet., 53,1228-39.
Weber-Dahlmann, M. (1957) Beitrage zur Einwirkung organisch chemischer
Substanzen auf die Lagerfahigkeit von Kartoffeln. Z. Bot., 45, 395-420.
Weis, G.G., Schoenemann, J.A and Groskopp, M.D. (1980) Influence of time of
application of maleic hydrazide on the yield and quality of Russet Burbank
potatoes. Am. Potato J., 57, 197-204.
WHO (World Health Organization) (1974) Toxicological evaluation of some food
additives including anticaking agents, antimicrobials, antioxidants, emulsifiers
and thickening agents. WHO Food Additives Series, no. 5.
Wiese, W., Bommer, D. and Patzold, C. (1975) Einfluss differenzieter Wasser-
versorgung auf Ertragsbildung und Knollenqualitat der Kartoffelplanze (Solanum
tuberosum L.) Potato Res., 18,618-31.
Wigginton, M.J. (1973) Diffusion of oxygen through lenticels in potato tubers.
Potato Res., 16,85-7.
Wigginton, M.J. (1974) Effects of temperature, oxygen tension and relative humidity
on the wound-healing process in the potato tuber. Potato Res., 17,200-14.
Wills, R.B.H., Wimalasiri, P. and Scott, KJ. (1979) Short pre-storage exposures
to high carbon dioxide or low oxygen atmosphere for the storage of some
vegetables. Hort. Science, 14, 528-30.
Wilson, A.M., McGann, D.F. and Bushway, R.J. (1983) Effect of growth-location
and length of storage on glycoalkaloid content of roadside-stand potatoes as
stored by consumers. J. Fd Protect., 46, 119-21.
Worobey, B.L. and Sun, W.-F. (1987) Isolation and identification of chlorpropham
and two of its metabolites in potatoes by GC-MS. Chemosphere, 16, 1457-62.
Worobey, B.L., Pilon, J.C. and Sun, W.-F. (1987) Mass spectral characterization
of a halogenated azobenzene (3,3'-dichloroazobenzene) from potato peels. J.
Agric. Fd Chem., 35, 325-9.
Wu, M.T. and Salunkhe, D.K (1977) Inhibition of wound-induced glycoalkaloid
formation in potato tubers (Solanum tuberosum L.) by isopropyl-N-(3-
chlorophenyl)-carbamate. J. Food Sci., 42, 622-4.
Wurr, D.C.E. and Allen, E.J. (1976) Effect of cold treatment on the sprout growth
of three potato varieties. J. Agric. Sci. Camb., 86, 221-4.
Yamaguchi, M. (1959) Effect of certain areas of production and storage on
chipping quality, chemical composition and specific gravity of California
potatoes. Proc. Am. Soc. Hort. Sci., 74, 649-60.
References 727
Yamaguchi, M., Timm, H., Clegg, M.D. and Howard, F.D. (1966) Effect of stage
of maturity and post-harvest conditions on sugar conversion and chip quality of
potato tubers. Proc. Am. Soc. Hort. Sci., 89, 456.
Zilva, S.S. and Barker, J. (1939) The ascorbic acid content of potatoes. Rep. Food
Invest. Bd, London, 1938, pp. 199-20l.
Zitnak, A. (1953) Factors influencing the initiation and rate of solanine synthesis in
tubers of Solanum tuberosum L., Thesis, Univ. of Alberta, Edmonto, Alberta,
Canada.
van Es, A. and Hartmans, K.J. (1969) The influence of abscisin II and gibberellic
acid on the sprouting of excised potato buds. Eur. Potato J., 12, 59-63.
van Es, A. and Hartmans, K.J. (1986) Carbohydrate metabolism during storage of
potatoes. Proc. 21st. Ann. Conf. ECSA-PRC, Heelsum (Neth), pp. 40-89.
van Es, A. and Hartmans, K.J. (1987) Starch and sugars during tuberization,
storage and sprouting, in Storage of Potatoes (eds A. Rastovski, A. van Es et
al.), Pudoc, Wageningen, pp. 79-114.
van Hiele, F.J.H. (1961) Unsprouted potato tubers treated with gibberellic acid
(GA3). Eur. Potato 1., 4, 26-39.
van Loon, C.D. and Miiller, K. (1984) Einfluss des Bodens und der Diingung auf
Ernaehrungswert der Kartoffel. Proc. 9th Triennial Conf. EAPR Interlaken,
1984, pp. 71-97.
van Steveninck, R.F.M. (1975) The washing or 'aging' phenomenon in plant
tissues. Ann. Rev. Plant Physiol., 26, 237-58.
van Vliet, W.F. and Schriemer, W.H. (1960) The sugar accumulation in potatoes
kept at a low temperature, as studied in a small selection of samples of Dutch
varieties. Eur. Potato J., 3, 263-71.
van der Plas, L.H.W. (1985) Respiration and morphogenesis in storage tissue, in
The Physiology and Biochemistry of Plant Respiration, (ed. J.M. Palmer),
Cambridge Univ. Press, Cambridge, pp. 59-65.
van der Plas, L.H.W. (1987) Potato tuber storage: Biochemical and physiological
changes, in Potato (ed. Y.P.S. Bajaj), Biotechnology in agriculture and forestry ,
Vol. 3, Springer Verlag, Heidelberg, Berlin, pp. 109-35.
van der Plas, L.H.W. and Wagner, M.J. (1983) Regulation of the activity of the
alternative oxidase in callus forming discs from potato tubers. Physiologia
Plant., 58, 311-19.
Vander Zaag, R. Chase, R.W. and Hammerschmidt, R. (1985) The effect of
hypochlorite wash treatments on potato suberization. Am. Potato J., 62,
409-18.
von Scheele, C., Svensson, G. and Rasmussen, J. (1935) Om bestiimming an
potatisens. Stiirelse och torrsubstanshalt med tilhjap an dess specifika vkt.
Nord. Jordbr. Forsk, 17-37.
CHAPTER 15
Potato production in the tropics

D .J. Midmore
15.1 INTRODUCTION
The cultivated potato is understood to have originated in the tropics,

predominantly in the Andes of South America where it remains of great
importance as both a subsistence and cash crop. Today, the potato is culti-
vated commercially in almost all climates of the world (Midmore and
Rhoades, 1988) with the exception of the tropical rain forest (AF on the
classification of Koppen) and the tropical rainy, short, dry season climate
(Koppen's MM). In the latter climate, however, potato has been grown
successfully under experimental conditions. This highly adaptive nature of the
potato has permitted it to take second place only to maize as the crop grown
in the greatest number of countries in the world (Horton and Fano, 1985).
The introduction of the Andigena potato to Europe in the last quarter of
the sixteenth century subsequent to the Spanish conquest of South
America (Hawkes, this volume, Chapter 1), was followed by conscious or
unconscious recurrent selection for the ability of the inherently short-day
plant to tuberize under longer photoperiods. This gave to the resulting
Tuberosum potato the same adaptive advantage that was years later
similarly conferred to wheat by N.E. Borlaug during the initial research in
Mexico of what was later to become the green revolution. Paradoxically, it
has been the tolerance to long days that has bestowed a degree of
adaptation, particularly with respect to tuberization, to the Tuberosum
type of potato that has favoured its cultivation in the lowland tropics. The
physiological basis of this, and other aspects of adaptation to warm tropical
conditions, will be discussed later in detail.
Severe levels of stress may be experienced by the potato crop in any of
the tropical environments under which it is grown. Within this chapter I
intend to expand upon the influence of stresses, both biotic and abiotic, but
with more emphasis on the latter, on potato production in the tropics.
Another stress, and more acutely sensed by the extensionist than by the
physiologist or agronomist, is the high cost of inputs. In particular, the cost
Production statistics for the tropics and subtropics 729
of seed tubers comprises from 20 to 50% of total production costs (van der
Zaag and Horton, 1983) and cheaper alternatives suitable for the tropics
will be discussed.
Since much of the physiology, breeding, and cultural practices of tropical
highland potato are similar to those of temperate potato, the discussion of
potato in the highland tropics will relate only to areas of specific interest such
as breeding for frost tolerance, breeding with species other than S. tuberosum
ssp. tuberosum, and rustic seed storage practices. Greater emphasis will be
placed on the approach for finding solutions to overcome limitations to
potato production in what were at one time considered as non-traditional
climates, i.e. climates generally considered too hot for potato cultivation.
Interpretations of physiological responses to high temperature, drought,
salinity, and low soil nutrient status (toxic soils) will act as a basis for
recommending: (a) breeding and selection criteria, and (b) implementing
agronomic practices favouring potato production in new areas of the
lowland tropics. Finally, examples of research, extension, and farmer
experience with growing potato in the lowland tropics will be presented.
15.2 PRODUCTION STATISTICS FOR THE TROPICS AND

SUBTROPICS
The potato is generally regarded as a crop of northern-latitude importance,

often implied in the use of the term 'Irish Potato'; however, the data
presented in Table 15.1 on potato production worldwide indicate other-
wise. The area planted to potato over the past two decades has increased in
developing countries with market economies, and together with substantial
increases in yield per hectare has resulted in a sharp increase (103%) in
potato production. In contrast, the area under potato production in
developed country markets has declined over the same time span by
approximately 2% annually. This decline in area was not offset by
increases in yield per hectare; consequently annual production over the
same period decreased by more than 20%. The major increase in volume
of developing country production has taken place in Asia. Much of this
production is found in the Asian potato-cereal food system, which
stretches from the Punjab of India through to China (Midmore and
Rhoades, 1988), in climates classified as 'tropical rainy' or 'tropical humid-
summer with a humid winter' by Troll and Paffen (1965). Potato produc-
tion over the same period also doubled in Africa, and increased by 40% in
Latin America. An important feature of the data in Table 15.1 is that the
yield per hectare in developing countries, with the exception of the Near
East, rarely reached 50% of yields in developed countries. Calculations by
Vander Zaag (1984) on yield potential of the potato in temperate and
tropical climates indicate that the difference should not be as great as
suggested by data in Table 15.1. Some factors limiting potato~yield in
730 Potato production in the tropics
developing countries are similar to those of north-temperate countries, but
a higher level of input in north-temperate countries enables their farmer to
overcome many such constraints. As examples may be cited the purchase
of fungicides to control Phytophthora infestans, of quality seed tubers to
replace degenerated seed stocks, and of water and nutrients to supplement
those provided by the environment. Economic factors play an important
role in the limitation of yield per unit area in developing countries, but
appropriate selection of genotypes, and production and storage practices,
can be instrumental in raising tuber yield there.
Table 15.1 Total annual production of potato averaged over 198011982, and % changes in
potato production in developed and developing market economies over the period
1961/65-1980/82
Total Area Yield

production harvested per unit area
Million tonnes % change Million hectares % change % change
Developed market
economies 72.34 -22 2.95 -41 31
Developing market
economies (all) 33.02 103 2.92 39 46
Africa 3.05 121 0.42 112 4
Latin America 11.38 49 1.07 2 46
Near East 6.30 127 0.43 71 32
Far East 12.19 177 1.00 68 65
Centrally planned
economies
Asia 59.03 177 5.85 108 34
Source: International Potato Center (1984b).
Much of the increase in the volume of potato production in developing

countries with market economies has, with the exception of those in Latin
America where the potato is a native and often staple crop, resulted from
large increases in area planted to potatoes. The increase in total production
reflects both an increase in demand by the consumer and an increased
desired by the farmer to produce this profitable crop. Another reason for
an increase in production is the convenience with which the potato can fit
into existing cropping systems. The development of early maturing potato
varieties permitted their cultivation during the 80-100 day lapse between
rice and wheat in India (Swaminathan and Sawyer, 1983), while the
ingenuity of Chinese farmers led to the inclusion of the potato as a relay
crop, predominantly with maize in southern China (Liu Jieming and
Midmore, 1990).
Statistics collected by the International Potato Center (1984b) show that
70-75% of potato production in developing countries is used directly for
consumption. Potato production costs per kilogram of output are higher in
Tropical highland potato production (> 1500 m) 731
developing than developed countries (Vander Zaag and Horton, 1983) and
the resulting high market price, relative to the price of alternative
foodstuffs in tropical countries, is generally a key factor resulting in low per
capita potato consumption in developing countries. In 1983, Poats classi-
fied tropical potato consumption into three groups:
1. staple, generally in the centre of origin of the potato (Andean countries)
with per capita consumption ranging from 60 to 200 kg per year;
2. as a complementary vegetable with consumption in the range of 15-50
kg per year and typified by the highlands of the Philippines;
3. as a luxury vegetable with annual per capita consumption of less than 10
kg per year which are generally consumed during festive ocasions, e.g.
during Lebaran celebrations following the Moslem month of fasting.
Stimulation of consumption in areas where potato is not consumed as a
staple food will depend upon the ability to reduce the retail price of
potatoes. Low-cost technology is required to enable the farmer in tropical
countries to produce more potato at a lower unit cost. Only then will the
trend in increased consumption per capita be maintained at the same rate
as over the past two decades in developing countries.
The remainder of this chapter will discuss the scientific principles upon
which low-cost technology may be based in order substantially to increase
production and consumption of potato in the tropics.
15.3 TROPICAL HIGHLAND POTATO PRODUCTION (>1500 m)
The influence of latitude on mean annual and seasonal temperatures is

modified by altitude and proximity to water masses. Consequently the zone
'Tropical Highlands' as it relates to temperate weather conditions favour-
able for potato production defies definition in terms of altitude alone.
Distribution of the potato crop in the tropical highlands is delimited both in
time and place by the lower end of the temperature scale, primarily by the
incidence of frosts. Here the potato crop is grown as a single crop per year.
Cold tolerance among wild species appears to be related positively to the
altitude in which they are found (Smillie et al., 1983), but Li and Fennell
(1985) suggest that such a relationship can occur only within a species,
since potato species possess different inherent frost hardiness. Avoidance
of terminal frost through the evolution of short-season species does not
appear to have occurred. It is probable that the unpredictability of frost
incidence in the highland Andes has favoured evolution of the frost
tolerance rather than frost avoidance type of adaptation to those climatic
conditions. At the other end of the temperature scale, potato production in
the lower tropical highlands has traditionally been delimited primarily by
the lack of potato species adapted to high temperatures. For example, the
Andigena group cultivated in the high Andes will not produce an accept-
able yield below 3000 m (Dodds and Hawkes, 1952).
In the tropical highlands, tuber yields are limited mainly by the following
pests and diseases: virus, nematodes and late blight (Phytophthora infestans),
which are discussed in other chapters in this volume. Bacterial wilt
(Pseudomonas sotanacearum), which is a serious constraint to highland
potato production, will be discussed in the section on tropical lowland potato
production. In the following sections, recent developments are presented
on selection for frost tolerance, the use of diffused light storage for seed
tubers, and the application of the crop-growthiradiation-interception
model in the context of highland tropical potato production.
15.3.1 Frost tolerance

Leaf and stem injury due to frost are brought about by the freezing of
extracellular water resulting in desiccation, and following thawing, ion
leakage due to disruption of cell membranes and a loss of turgor. Li and
Fennell (1985) have defined frost resistance as the ability to survive
freezing temperatures of -4°C or below, without injury. They dispute the
conclusion that rupture or loss of semipermeability results in the efflux of
ions following the freeze-thaw cycle, and favour the theory of inactivation,
or altering, of membrane proteins involved in active ion transport. The
sharp increase of K+ permeability of injured cells could then be attributed
to a passive transport of K+. Irrespective of the mechanism of injury, the
measure of electrolyte leakage (by way of conductivity tests) from leaves
subjected to subzero temperatures for specified periods could provide
plant breeders with a methodology to screen germplasm for frost tolerance.
In practice, this test, in common with others involving vital stains (e.g.
TIC (Levitt, 1980» or plasmolysis (Li et at., 1979), has not provided the
plant breeder with the tools to screen for frost tolerance either more
objectively, or more quickly, than the screening of frosted plants based
upon visual symptoms. Added complications, under laboratory conditions,
such as the non-involvement of supercooling and the ability of some
species to acclimatize at low temperature and enhance their tolerance,
have resulted in discrepancies in the ranking for frost tolerance based upon
field and laboratory conditions (Richardson and Weiser, 1972; Li and
Fennell, 1985). Estrada (1982), however, did report a close correlation
between field and growth chamber (excised leaflet test) evaluations for
frost tolerance, but stressed the need to ensure that moisture conditions in
the growth chamber were adequate to induce ice nucleation.
Newer avenues of research, at the cellular level, based upon the role of
abscisic acid (ABA) in soluble pwtein synthesis during acclimatization
(Chen et at., 1983) may provide suitable techniques for the unequivocal
identification of frost-tolerant clones in breeding programmes. In common
with other species, frost tolerance in the potato has been reported to be
positively correlated with leaf proline content (van Swaaij et al., 1985). The
potential for increased frost tolerance through the selection of clones with
high proline levels has been demonstrated (van Swaaij et al., 1987).
Initially, hydroxyproline resistant (hyp) lines were selected in cell culture
and subsequently regenerated. Although leaves of the hyp lines were more
tolerant to frost (withstanding -3.6 to -4.2°C compared to -3.0°C for the
wild type) this was only weakly related to increased levels of either proline
or total levels of amino acids. It is clear that knowledge is limited on the
subject of responses elicited at the cellular level prior to, during, and
following frost in terms of applications to a plant breeding programme.
Other lines of research, reviewed by Li and Fennell (1985), have
attempted to relate inter-specific differences in frost tolerance to morpho-
logical (rosette habit) and anatomical adaptations. Controversy surrounds
the relationship between cell size and stomatal density and frost hardiness.
Firmer relationships have been observed between the development of
palisade parenchyma and frost hardiness. A highly developed palisade
parenchyma was found by Estrada (1982) to be closely related to frost
hardiness; and Palta and Li (1979) reported that frost hardy species had
two layers of parenchyma cells compared to the single layer in non-tolerant
species.
Deficiencies in the relationships between the previously mentioned
characters and cold tolerances (Smillie et al., 1983) and in particular their
inability to quantify non-freezing cold injury (Greaves and Wilson, 1986)
has led these researchers to experiment with the analysis of chlorophyll
fluorescence. Changes in chlorophyll a fluorescence following stress injury
in chloroplast membranes are monitored, which indicate changes in
photosystem II activity. This technique has been applied to illustrate
differential responses to various stresses (Smillie and Hetherington, 1983).
Cold tolerance was expressed as the time for a 50% decrease in the
maximum rate of the induced rise in chlorophyll fluorescence (FR ) of
leaves kept in the dark at O°e. Greaves and Wilson (1986) cautioned,
however, that other factors including antenna size, the rate of oxidation of
the electron acceptor, and the photochemistry of photosystem II may
affect FR , thereby making the objective use of the method for screening
questionable. Nevertheless, close relationships between altitudinal range
and tolerance to frost were found (Smillie et al., 1983). The technique also
permitted identification of some new sources of cold tolerance (e.g. S.
chacoense and some Chilean accessions of S. tuberosum ssp. tuberosum
- Greaves and Wilson, 1986), highlighting its potential use to screen
effectively for cold tolerance in breeding programmes.
A number of techniques exist for evaluating frost tolerance under
experimental conditions, but from a practical viewpoint only the visual
evaluation method of seedlings subjected to sub-freezing temperatures in
growth chambers lends itself to the scale required by breeding programmes
to test tens of thousands of genotypes per year. Mendoza and Estrada
(1979) have indicated that the visual assessment proposed by Chen et al.
(1976) for the excised leaflet test of Sukumaran and Weiser (1972) permits
both the rechecking of genotypes selected at the seedling level in growth
chambers particularly for escapes, and the definition of killing temperature
for potential parents. The test is based on a visual assessment of leaf colour
following thawing, instead of the measurement of conductivity of leachate,
following 1°C h- I our cooling to the desired temperature, and remaining
there for 30 min. The unpredictability in timing, intensity, and duration of
naturally occurring frosts, limit the extent to which they may be useful in a
screening programme. Final evaluation should be made under farmers'
field conditions, but efforts must continue to ensure that measures of
resistance in the laboratory or growth chamber correlate well with levels of
resistance required in the field.
15.3.2 Diffused-light storage of seed potatoes

Potato cropping in the tropical highlands is largely annual which requires a
storage period for seed tubers ranging from 7 to 8 months. Even with two
main cropping seasons per year, as found at lower elevations in the tropical
highlands (Potts et al., 1983), seed tubers are stored from the harvest of
one crop until planting of the same season crop in the following year.
Dark-storage of tubers between harvest and planting results in the develop-
ment of long, etiolated sprouts which need to be removed prior to planting.
Storage in diffused-light - although not affecting the length of dormancy -
restricts sprout growth which results in less weight-loss during storage,
eliminates the need to desprout prior to planting, and results in an earlier
and more uniform field emergence, often associated with greater tuber
yields (Table 15.2). Although the inhibitory influence of light on sprout
growth has been recognized for many years (Burton, 1966), planting of
green-sprouted tubers has not been popular in developed countries, since it
has been difficult to mechanize. Hand-planting, a common practice in
many developing countries, has facilitated the spread and adoption of
diffused-light storage (DLS).
Table 15.2 Effect of 180 day diffused-light storage (DLS) on storage behaviour of
seed tubers and their subsequent field performance in the tropical highlands of Peru
(means of eight clones)
Treatment Mean sprout Mean sprout Mean % Emergence % Tuber
length (cm) number per weight loss 20 dap yield
tuber (t ha- 1)
Dark stored 18.8 1.6 15.3* 1.2t 24.6
DLS stored 1.4 4.0 8.3 58.8 28.8
* Includes weight of sprout removed prior to planting.
t Desprouted prior to planting
Source: International Potato Center (1979).
20
E 15
3-
.c
g> 10
..Q!
"5
~ 5 ~~
~--=-====
•
CJ) .....
0.01 0.1 1.0 10

Solar radiation (W m·2 )
Figure 15.1 Sprout length following 6 months storage in DLS as influenced by

solar radiation and site of storage. 0 cool site; • warm site. (Source: International
Potato Center, 1983).
Studies on the interrelationship of temperature, light intensity, and the

rate and pattern of spout growth indicate that the light intensity necessary
to effect a 50% reduction in sprout length is almost unaffected by
temperature, and for European varieties it varies between 0.1 and 0.25 W
m- 2 (International Potato Center, 1984a). Under tropical conditions (Fig.
15.1) sprout growth was more extensive and more sensitive to light
quantity under cooler highland than warmer lowland conditions, suggest-
ing that the warmer temperatures were supra-optimal for sprout growth.
Further studies (International Potato Center, 1985) have shown that the
active wavelength to effect light inhibition of sprout growth is principally in
the blue range below 550 nm, with some response to red light (710 nm).
Additionally it was shown that inhibition of sprout growth may be effected
by conditions of high temperature (33/20°C day/night) alone. Increase in
the duration of daily exposure to natural diffused light effected propor-
tional reductions in the length of sprouts following a 6 month storage under
cool tropical conditions, with half an hour daily exposure effecting more
than 50% reduction in sprout length (International Potato Center, 1983).
Further data (McGee et al., 1987) from experiments in which the same
total light energy falling on tubers was applied either as high intensity for a
short duration (21 W m-2 for 1 h) or low intensity for a long period (2.1 W
m- 2 for 10 h), confirmed that the total light energy falling on the tubers was
the dominant factor controlling sprout growth. From this result and earlier
findings, the authors concluded that the suppression of sprout growth by
light (once dormancy was broken) belongs to the high irradiance class of
photomorphogenic reactions, i.e. is characterized by a non-photoreversible
nature, has complex action spectra with peaks as mentioned before in the
red and blue regions, and requires moderate light intensities over an
extended period.
Purpose-built diffused-light stores are not essential for the successful
adoption of the green-sprouting principle by farmers. Even farmers with
very limited economic means can benefit from the diffused-light tech-
nology, provided that a few simple requirements are met (Booth and
Shaw, 1981). Light penetration through transparent walls or windows, with
wire or open plastic mesh to prevent entry for rodents, yet permitting
ventilation, is essential. A well insulated and overhanging roof prevents
penetration by direct sunlight and rainfall, and shelves designed to contain
only two layers of tubers permit light to reach each tuber. Since tubers are
not isolated from the exterior, one major disadvantage of DLS is the
frequent need to control pests within the store. In particular potato tuber
moth (Phthorimaea opercuZelia (Zeller) (Raman and Booth, 1984), and
aphids (Parker et aZ., 1983) have been observed in rustic diffused-light
stores, but both may be controlled effectively through the correct use of
pest management components (e.g. repellant weeds, biological insecti-
cides, synthetic sex pheromones) in an integrated pest-management pro-
gramme (this volume, Chapter 11). The widespread adoption of DLS for
seed potato in the tropical highlands reflects the practical application of the
simple, yet sound, principle upon which it is based (Rhoades et af., 1983).
The simple relationship between light and percentage inhibition of sprout
growth is essentially independent of location, cultivar, and temperature
(provided the latter permits sprout growth) and can be used to predict the
minimum diffuse-daylight intensity required for adequate sprout growth
suppression (McGee et aZ., 1988).
15.3.3 Physiology of growth and development

Detailed studies on the physiology of growth and development of the
potato in tropical highland climates - the original niche of the potato - are
almost non-existent in the literature. With emphasis on fertilization
practices, Ezeta and McCollum (1972) quantified the accumulation of dry
matter in an Andigena clone, Renacimiento, throughout the growing
season at 3300 m in Peru (12°S). The average daily mean temperature
ranged from 9 to 12.4°C with a diurnal range from 10.6 to 18°C. Growth
was reportedly not limited by lack of water, and tuber yields ranged from
10.1 to 41.1 t ha- 1 for the zero and complete (160:160:160 N:PzOs:KzO)
fertilizer regimes, illustrating the phenomenal response to fertilization as
recorded over the earlier decade by McCollum and Valverde (1967; as
cited by Ezeta and McCollum, 1972). Maximum crop growth rates for the
well-fertilized plots were in the range of 19.8-20.5 g m- z day-I, close to the
upper limits reported by Allen and Scott (this volume) and final harvest
index was 0.7, not unreasonably lower than published values for potato in
northern latitudes. When irrigation was employed at the same site (Nelson
and Midmore, 1986; Midmore, unpublished), tuber yields of 70 t ha- 1 were
achieved in experimental plots.
Studies by Nelson (1987), along the lines of the neo-classical analysis of
growth (Monteith, 1977) in terms of cumulative solar radiation interception,
efficiency of conversion to dry matter, and partitioning to tubers
(Allen and Scott, 1980) confirmed the high yield potential of Andigena
clones in the tropical highlands of Peru, although water-logging and the use
of physiologically old seed resulted in abnormally early senescence (117
days, versus 165 days in the study by Ezeta and McCollum, 1972). The
radiation conversion coefficient (RCC - the slope of the relationship
between total plant dry weight and cumulative intercepted solar radiation,
g MJ- 1)at 1.99 g MJ- J was greater than those reported for potato in
southern England (1.0-1.5 g MJ- J , Burstall and Harris, 1984), Wales «1.5
g MJ- 1 , Bean and Allen, 1981) and Scotland (1.85 g MJ- 1 , MacKerron and
Jefferies, 1984). The greater efficiency at the highland tropical site was not
only characteristic for Andigena clones but also for Tuberosum clones.
That the efficiency was greater in the tropical highlands is perhaps
surprising since instantaneous values of intercepted radiation during a
large proportion of the day were greater than the 400 W m- 2 considered by
Sale (1974) to effect light saturation of photosynthesis in potato canopies.
Canopy size varied markedly between Andigena and Tuberosum clones
(the latter with a much smaller canopy) and could not be directly
responsible for the more effective use of intercepted solar radiation.
Leaves were thicker at the highland compared to the lowland site in the
study by Nelson (1987), and although comparable data between leaf
thicknesses of temperate and tropical cool sites are not available, it is
possible that the anatomical response of the leaf to high levels of irradiance
may confer on it a greater photosynthetic efficiency at higher irradiance
(Charles-Edwards, 1978).
Growth physiology of Andigena clones has been studied in Japan by
Isoda et al. (1984, 1985). No difference was observed between the RCC
of Andigena clones and that of a standard Tuberosum clone at high
plant density (11.1 plants m- 2 ) but at low density (4.9 plants m- 2 ) the
Tuberosum clone was always more efficient (by 27-40%). The authors
cited low photosynthetic activity, less source strength, and a canopy
structure at the lower populations leading to a more uniform distribution of
radiation over all leaf layers, as responsible for the lower RCC in the
Andigena clone. The RCC is but one component in the neo-classical
growth analysis model. Andigena clones appear well adapted to their
natural environment with respect to interception of solar radiation -
another of the components of the neo-classical growth analysis model
(Table 15.3). The subspecies Tuberosum is, compared to the Andigena
subspecies, poorly adapted to the highland tropics in this respect. Cool
temperatures and in particular the short daylength, effect a very rapid
onset of tuberization in Tuberosum clones and foliage growth ceases early
(Table 15.3), particularly in Tuberosum clones characterized by a long
critical photoperiod.
The introduction of potato to tropical highlands, other than those of
South America, has primarily been through the northern-latitude route,
Table 15.3 Influence of climate in Peru on maximum crop cover
and total tuber yield of four clones of contrasting genetic background
(Cool site - Huancayo; 3200 m above sea level, average J9.3/7.2°C
day/night; warm site - Lima: 240 m above sea level, 26. 0117. SOC)
Clone (+ pedigree*) Maximum crop cover Tuber yield

(% of best clone) (kg m o2 )
Cool Warm Cool Warm
DTO-33 (T x P) 32 68 0.79 2.34
Desiree (T) 54 89 1.06 2.65
Revolucion (A x(T x A)) 60 100 1.69 2.56
Mariva (T x A) 100 77 2.88 1.62
* T = Tuberosum, P = Phureja, A = Andigena.
Source: Nelson (1987).
and cultivated clones in non-Andean regions, almost without exception,

are derivatives of S. tuberosum ssp. tuberosum. Given the poor adapta-
tion of Tuberosum to highland tropics in terms of reduced potential to
intercept solar radiation (Table 15.3), this is an apparent anomaly. Until
recently, Tuberosum clones with the shortest critical photoperiods (i.e.
main crop or late varieties that do not tuberize under the longest days of
mid-summer in north-temperate climates) have been the source of material
planted in the tropical highlands of Africa and south-east Asia (Simmonds,
1971). With further distance from the equator, temperatures equivalent to
those reported above as suitable for the potato in the Peruvian Andes are
found increasingly at lower altitudes. Concomitantly, as the distance from
the equator increases, the delaying influence of longer summer photo-
periods on tuber initiation and maturity becomes more apparent, and the
precocious dwarf-plant habit of Tuberosum in the high Andes gives way to
the characteristically vigorous and later maturity crops of the same S.
tuberosum ssp. tuberosum cultivars in the mid-elevation tropics and
subtropics. When the potato is taken from high tropical altitudes
to the isohyperthermic environments of the lowland tropics, high tempera-
ture is responsible for delay of tuberization and, if not excessive,
permits the achievement of reasonable tuber yield there with the same
Tuberosum cultivars. This discussion is amplified in Sections 15.4.1 and
15.4.2.
Both the increase in mean growing-season temperature and the exten-
sion of the summer photoperiod at lower altitudes in higher latitudes are
conducive to a longer season Tuberosum crop but inhibitory for tuberiza-
tion in the Andigena crop. Temperature data during the season for the
tropical highland potato-producing areas in East Africa (Moreau, 1944)
vary between monthly means of 14 and 22°C, considerably higher than
those of the production areas in the high Andes, and sufficiently high to
retard the hastening effect of short photoperiods on tuberization of
Tuberosum clones, and, in combination with the long days, to inhibit
tuberization of Andigena clones.
As with the potato in the tropical highlands., published accounts of
detailed field studies on the physiology of growth and development in the
mid-elevation tropics were, until recently, non-existent. Moreau (1944)
was quick to defend the low latitude potato against the criticism that yields
could not match those of northern latitudes. His analysis, based upon
hours of sunlight, showed that low latitude crops receive between 1320 and
1625 h, compared to the estimated 2600 h in the Scottish lowlands. Yields
in the tropics under good crop husbandry ranged from 16.5 to 30.9 t ha- 1
and for the British official potato trial, illustrating the maximum yield
possibilities at that time, ranged from 26.7 to 44.5 t ha- 1 (Moreau, 1944);
the difference was proportional only ~o the number of sunlight hours during
the cropping season. A similar conclusion was obtained by Haverkort
(1986) when comparing potential crop yields for Rwanda against those
calculated for northern Europe (Vander Zaag, 1984). On an annual (as
opposed to a seasonal) basis, potential yields in central Africa are between
100 and 120 t ha- 1 , comparing favourably to the potential calculated by
Vander Zaag (1984).
A detailed study of tuber yield in the central African highlands was made
by Haverkort (1985). Using the framework of intercepted radiation, RCC,
and partitioning of dry matter to tubers, Haverkort and Harris (1987)
developed a regression model to predict total dry matter production. The
model differs from similar models for temperate conditions (MacKerron
and Waister, 1985) in that the RCC is weighted by a temperate correction
factor. Field data show that the uncorrected RCC ranged from 0.82 g MJ- 1
at 1350 m to 1.20 g Mrl at 2350 m, values somewhat less than those
reported earlier for temperate regions, but reasonable in view of the
seasonal average temperatures of 21.8°C and 15.0°C respectively. The
model adequately predicted total dry matter production to within a
precision of 13% and tuber dry yield to 10%, and permitted a careful
simulation of the influence of factors potentially limiting production e.g.
late blight in the rainy season. Data presented by Haverkort and Harris
(1986) illustrate the potential yield advantage of late crops in terms of both
greater cumulative radiation interception and higher RCC, but practical
considerations (e.g. delayed maturity coinciding with drought or heavy
rain; incompatibility with two potato seasons per year) were of overriding
importance in choosing suitable genotypes for the area.
Within the same framework of intercepted radiation, RCC, and tuber
partitioning, Haverkort and Rutayisire (1986a,b) interpreted the influence
of chemical fertilizers on tuber yield in tropical highlands. Chemical
fertilizers favoured leaf development early in the season, and nitrogen
additionally delayed senescence. The tuber RCC (i.e. the slope of the
relationship between tuber dry weight and cumulative intercepted radia-
tion) was especially responsive to phosphorus, and was greater in
treatments in which tuberization took place at greater leaf area index
(LAI). These data are interpreted through the influence of higher tuber
numbers per plant on sink strength, but could equally well be explained as
a result of reallocation of dry matter from the canopy to tubers.
The use of chemical fertilizers is but one of the avenues by which tuber
yield in the tropical highlands may be raised. It does represent a high-cost
input, in common with the use of pesticides, fungicides, irrigation, and cold
storage for seed tubers. Often, tuber yields may be raised substantially
(even doubled) by the implementation of low-cost inputs, such as healthy
seed, late blight and bacterial wilt resistant clones, soil moisture conserva-
tion practices and alternatives to tuber seed. A number of these low-cost
inputs are discussed in other chap,ters, and in later sections on lowland
tropical production. Those amenable to introduction through plant breed-
ing are briefly discussed below.
15.3.4 Plant breeding

Although the growth and tuberization of Tuberosum clones is acceptable
in intermediate altitudes of the tropics, and that of Andigena clones at
higher altitudes, it is obvious that specific breeding efforts to address
inherently tropical limitations to the achievement of potential yield are
called for (Haverkort, 1986).
Breeding objectives for the potato in the tropical highlands have largely
focused on the incorporation of pest and disease resistances and tolerance
to frost (International Potato Center, 1984a). Incorporation of beneficial
genes from various species is effected through elaborate techniques (e.g.
first division restitution 2n pollen production (Iwanaga, 1985), somatic
hybridization (Austin et al., 1985) and the use of bridging species
(Hermsen, 1987)). For the transfer of late blight resistance from S.
bulbocastanum and S. verrucosum, to Tuberosum cultivars both S. acaule
Table 15.4 Wild and cultivated (c) species

with known tolerance to frost and used in
breeding for frost tolerance
S. acaule S. demissum
S. ajanhuiri (c) S. juzepczukii (c)
S. boliviense S. megistacrolobum
S. brevicaule S. multidissectum
S. bukasovii S. raphanifolium
S. canasense S. sanctae-rosae
S. chomatophilum S. toralapanum
S. curtilobum (c) S. venturii
S. vernei
Sources: Estrada (1978, 1982): Li and Palta

(1978).
and S. phureja have been used (Hermsen, 1987), and efforts are con-
sistently made to select R-gene-free clones (International Potato Center,
1987). Similar efforts using wild and cultivated potato species are in
progress in the development of frost-tolerant clones (Table 15.4).
In order to maintain the valuable range of diversity, yet take advantage
of specific traits for incorporation into an agronomically suitable genetic
background, the population approach to breeding for improvement in
unadapted Solanum cultivated species has often been adopted (Plaisted et
al., 1987). Incorporation of resistances to root knot nematodes, bacterial
wilt, late blight and PVY to a lowland tropic population has been
successful, as has the incorporation of glandular trichomes from S.
berthaultii to cultivated potatoes. Although the degree of emphasis that is
placed upon various attributes for selection may vary between programmes
that breed potato cultivars for northern or tropical latitudes, and the route
through which selected parents are obtained may differ, the final step, that
of selected crosses between specific clones with desired characteristics for
introduction to the best adapted clones, is the same whether in pedigree or
population breeding. The population approach to breeding of new potato
varieties has only been in progress at the Internationl Potato Center for 15
years and its success in the generation of locally adapted varieties is
illustrated in Table 15.5, together with a list of specific characteristics for
which the clones were selcted.
Table 15.5 Germplasm distributed by elP, and released or named as

varieties by developing countries
No. of cultivars
Population Traditional Total no. Major attributes*
breeding breeding of countries
Latin America 16 17 10 BW, LB, PVY, PYX,
PLRV, E, HT, CQ,
Cyst, Fr
Africa 7 38 11 BW, LB, PVY, PLRV,
E,HT,Q
Asia 2 18 9 BW, LB, HT, PLRV,
PVY, W, E
* BW = Bacterial wilt resistance PLRV = Potato leafroll virus resistance
LB = Late blight resistance W = Blackwart resistance
Ff = Frost tolerance CQ = Cooking quality
HT = Heat tolerance Cyst = Globodera spp. resistance
PYX = Potato virus X E = Earliness
PVY = Potato virus Y resistance
Source: J.E. Bryan, personal communication, November 1988
15.4 TROPICAL LOWLAND POTATO PRODUCTION « 1500 m)-
CONSTRAINTS AND SOLUTIONS
Although the term 'Tropical Lowlands' has been generally considered

synonymous with the term 'hot tropics', due to the seasonal nature of
climate in all but the equatorial rain forests, there exist periods of the year
when climatic conditions are marginally more conducive to potato produc-
tion. For example, data in Fig. 15.2 from the tropical rainforest in the
Amazon basin, 60 S of the equator, and from the Philippines (14°N of the
equator) illustrate the monthly variation in night-time minimum tempera-
ture, its relation with rainfall, and the implied relationship with clear days
if rainfall can be considered proportional to cloudiness. The favourable
months for the potato based upon the premise that growth and yield of
potato are favoured by cool nights, would be June to August in Peru, and
December to February in the Philippines, which fortuitously are those with
the shortest days, additionally favouring potato tuberization.
24
E
::J
23
·§G
CO
22
'E-; 21
:E.a
_(1)
c~
20
o Ql 19
E~
C Ql 18
(ll-
o
~ 17
16+-~-,~~-,~--,----,
o 100 200 300 400
Mean monthly rainfall (mm)
Figure 15.2 Relationship between mean monthly minimum temperature and

mean monthly rainfall at Yurimaguas (Peru 60 S, 190 m above sea level, data from
1980-1982; 0) and Los Banos (Philippines 14°N, mean data 1959-1980; D).
From the outset it should be understood that, alhough one refers to

potato production in the lowland tropics, this does not imply a year-round
global production capability; this may only be achieved with truly lowland
tropical root or tuber crops (e.g. sweet potato, yams, cassava). Rather, the
term tropical lowland potato production refers to the judicious choice of
site, and particularly season, within the lowland tropics favouring the
production of potato.
15.4.1 Theoretical considerations and practical results, pre-1970s

In 1944, Moreau gave evidence of the potential for cultivating potato as a
low-elevation tropical crop, even though efforts were probably made to
cultivate the potato there prior to that date. Authors of early studies on the
Tropical lowland potato production «1500 m) 743
effects of high temperature (Bushnell, 1925; Werner, 1934), photoperiod
(Edmundson, 1941) and light and temperature (Beaumont and Weaver,
1931) on potato growth and yield under greenhouse conditions argued
against the potential of the potato as a lowland crop. In fact Bald (1941)
and Driver (1941) definitively stated that it would be impossible to
cultivate S. tuberosum in the tropics because of the short daylength.
Moreau (1944) and Rayner (1945) understood fully the important practical
implications of the interactions between temperature and photoperiod
in regulating tuberization, which were first reported by Roberts and
Struckmeyer (1938). Early studies involving high temperature were con-
ducted under long-day conditions (e.g. Bushnell, 1925). Rayner (1945)
assembled evidence to show that, although the optimum temperature for
yield was similar under short or long days, the decrease in yield due to
supra-optimal temperatures was much less accentuated under short than
long days. Rayner also recognized the value of main-crop potato varieties
as potential varieties for the warmer tropics, since they would not be so
dependent on long days for tuberization (i.e. varieties with a medium to
short critical photoperiod in present-day terminology).
During the 1950s, further discussion centred on the potential of the
potato as a tropical lowland crop. Dodds and Hawkes (1952) quoted
examples of successful potato production with northern latitude varieties at
sea level on the periphery of the tropics (Cuba - 20 to 23°N, Tamargo
(1948); Gangetic plains of India - 25° N, Pal and Pushkarnath (1951)),
during the winter growing season. Further evidence for the ameliorating
effect of short days on high temperature inhibition of tuberization was
obtained by Went (1957) and Gregory (1965). Gregory even defined the
upper temperature limit, under a lO-h daylength, at which no tuberiza-
tion occurred as being 30/24°C day/night air temperature and 30°C soil
temperature, consistent with present-day distribution of lowland potato
production.
Field studies on the cultivation of potato in the tropical lowlands in the
1960s were largely motivated by financial considerations; self-sufficiency in
production and the saving of foreign exchange were of paramount impor-
tance in many countries accustomed to importing potato. Successful
experimental crops in Trinidad in 1960/61 by Gooding (1961) with a
maximum yield of 26 t ha- 1 , and all but 2 of 16 cultivars yielding 15 t ha- 1 , in
a short-day dry environment with day and night temperatures of 3~31 °C
and 19.5-20.5°C respectively, provided the basis for further experimenta-
tion. Although lack of irrigation, nutrient supply and the presence of
Pectobacterium carotovorum (Erwinia carotovora ssp. carotovora) limited
yield in later experiments, Gooding (1964) concluded that potato could be
a potentially important crop at low latitudes and elevations in the Caribbean.
Additional experiments carried out by Chapman (1965) in Trinidad
identified problems related to the incidence of early blight (Alternaria
solani) and a species of Prodenia, to the erroneous use of tubers, harvested
in the wet season, for seed, and to the need to plant on soil with good
drainage and at close spacing (70 m x 30 cm).
The problems mentioned above, as well as others related to the
economics of production, served as guidelines for future field research on
the adaptation of potato to the lowland tropic. Aspects of this research will
be discussed in detail in later sections.
A greater degree of success than in Trinidad was achieved by Thavarajavel
(1969) in the Jaffna peninsula of Sri Lanka, also with a winter-planted crop
(November to January planting). Starting only with small observation plots
in 1957, the extent of production grew to 198 hectares in 1967/8 with orders
for seed tubers sufficient for 530 hectares for the 1968/9 crop (Thavarajavel,
1969). High seed costs and the dependence on the timely arrival of
imported seed were key factors that influenced potato production in
Jaffna. Research to find solutions to these types of limitations was also high
on the list of priorities favoured to stimulate self-sufficiency in production
of potato in the lowland tropics.
In the 1960s laboratory and greenhouse research was still addressing
environmental influence on potato growth and development. Data were
forthcoming that would have direct practical application on lowland
tropical potato production in the future. Optimal soil temperature favour-
ing rapid emergence (Yamaguchi et ai., 1964 - 21 to 24°C; Epstein, 1966-
22°C; Borah, 1959 - 22°C) was found to be greater than that favouring
tuberization and would have implications for the practical manipulation of
soil temperature. Jackson (1962) highlighted the difficulty in establishing a
crop from tubers planted in soil at > 18°C if subsequently flooded for more
than one day, which explained in part why the wet-season crop in Trinidad
(Chapman, 1965) was unsuccessful.
The tropical environment has a strong influence on plant morphology.
The lower leaf:stem ratio at higher temperature (Bodlaender, 1963) would
influence choice of optimum plant population for interception of solar
radiation. The characteristic decrease in relative rate of leaf expansion at
high temperatures later in the crop cycle (Borah and Milthorpe, 1962)
coupled with the shorter leaf longevity (Nelson, 1987) would have serious
implications for maintenance of crop cover during the latter part of tuber
bulking, offset in part by the role of fertilizer nitrogen applications (Burt,
1965). Although some variability between reports was apparent, most data
supported unanimously the hypothesis that the detrimental influence of
high soil temperature on tuberization and yield was greatest when air
temperature was also supra-optimal (Bodlaender, 1963). This, in combina-
tion with another conclusion by Bodlaender that the unfavourable in-
fluence of high temperature on potato tuber yield was more pronounced in
combination with drought, clearly defined the challenge that was to be
faced in fully adapting the potato physiologically to lowland tropical
conditions.
Already in the 1960s, scientists were breeding potatoes for the warmer
tropical climates (Khanna, 1966) with a certain degree of success. In India,
clonal selections able to tolerate 15 days post-planting at 45/30°C day/night
were field-tested at Patua and later in Coimbatore (India) where the
greatest yield (9.7 t ha- 1) exceeded that of control Tuberosum clones by a
factor of five.
15.4.2 Abiotic stress and growth of the crop, data 1970s onwards
In each of the following sections a short review of the principal effects of
abiotic stresses will be presented, together with an indication of the
appropriate agronomic practices and breeding strategies that have proved
useful in overcoming such stresses. It should not be forgotten that
important interactions between the various stress-inducing factors also
influence the potato crop in hot climates. For a fuller treatise on the
influence of environment in the lowland tropics on potato physiology, the
reader is referred to Midmore and Rhoades (1988).
(a) High temperature

Rates of physical and biological processes depend largely on ambient
temperature, and data presented in previous sections would tend to
confirm that optimum temperature for maximum potato yields varies over
the broad range of 15 to 22°C. High temperature is herein referred to as air
temperatures greater than 18°C minimum at night and greater than 28°C
maximum during the day, which are prevalent throughout the crop season.
This is in contrast to the potato-growing areas found in Idaho, USA, and
the Negev, Israel, where temperatures during bulking may reach similar
values, but planting and tuberization take place under more temperate
conditions.
Maximum and minimum soil temperatures at the depth where tuberiza-
tion occurs are on average 6 to 8°C higher than respective air temperatures
(Midmore et ai., 1986a) during the tropical winter. The principal effects of
high temperature are primarily manifest on emergence, tuber initiation,
net dry matter production, senescence, tuber quality, and replanting value.
Since successful emergence and establishment of the crop are of para-
mount importance to achieve maximum potential yield, much effort has
focused on this early phase in the potato crop development. The influence
of soil temperature on crop emergence in the lowland tropics is illustrated
with field data in Fig. 15.3, which were for presprouted tubers. A much
narrower temperature optimum and sharper slope at higher temperature
was evident for unsprouted tubers (Midmore, unpublished). Soil tempera-
ture during the day appears to be more important than in the night in the
regulation of the emergence rate (Midmore, 1984b). The involvement of
ethylene in reducing sprout elongation and by default, plant stand,
emergence, and vigour, has been implicated by data of Timm et al. (1980),
which show that at 30°C compared to 25°C or 20°C, sprouts thickened, root
growth was limited, and ethylene production increased. Further studies on
the role of plant growth regulators in the inhibition of emergence at high
temperature have not been reported. Genotypic variation for the rate of
emergence under high temperature has been reported (Midmore, 1984b),
and screening for the ability to emerge rapidly under high soil tempera-
tures is in progress (Gebre-Mariam, 1988).
..
Ql 30
()
cQl
•
• • •
... •
l?
~ 20
Ql
;J2. • •
0
l!) 10 •
.9
Ul
>-
co
0 0
30 50 70 90 110 130 150
Soil heating
(~oh>20°C, 08:00 - 18:00h)
Figure 15.3 Influence of daytime soil heating (7 cm depth) on crop emergence.

Soil heating was modified by shade treatments. Fitted regression y = 17.65 -
0.215x + 0.00196x2 , ? = 0.704. (Source: Midmore et at., 1988).
A more practical alternative approach to the selection of cultivars able to

emerge under high soil temperatures, is to cool the soil at the time of
planting. Data in Fig. 15.3 illustrate how this may be effected by an
associate crop shading the soil in a relay-cropping sequence. As an
extension of the shading principle, the use of organic mulches has been
widely employed to favour emergence and stand establishment under
harsh environmental conditions in the lowland tropics (Rashid et ai., 1981;
Taja et ai., 1984; Alkamper and Linde, 1985; Midmore et ai., 1986b),
Moreover, organic mulches also act as a barrier insulating the soil surface
from the atmosphere. This has important implications for the conservation
of soil moisture during the emergence period (Midmore et ai., 1986a). If
mulches are more reflective than the soil surface, they will also reduce the
potential sensible heat load at the soil surface. Soil reflect ants alone, for
which the shading and barrier effects were absent, reduced soil tempera-
ture and improved plant emergence and tuber yield (Midmore, 1984b).
The application of other simple principles relating soil temperature to
soil physical properties, e.g. manipulation of depth and form of planting,
planting position and orientation of rows, have been studied in relation to
the hastening of emergence (International Potato Center, 1981-1984).
Daytime soil temperature declines with increasing soil depth, but the effect
of this reduction in temperature must compensate for the greater distance
required for sprouts to emerge if deeper planting is to result in faster
emergence. Soil physical properties and soil water content may play an
overriding role in determining the planting depth. For example, in a sandy
soil in the Amazon (Alkamper and Linde, 1985; Midmore, unpublished)
plant emergence rate was optimal at 15 cm depth; whereas in the Philippines,
with a heavy clay loam, shallow planting (2 or 5 cm soil covering of the
tuber) was more favourable for quick emergence (Vander Zaag and
Demagante, 1987). The form of planting (ridged, in raised beds, on the
flat, in furrows) will also influence soil heating through direct effects on
the radiation balance at the soil surface. Tubers planted on the flat will
experience lower maximum soil temperature than at the equivalent depth
in ridges (Fig. 15.4). Planting in raised beds results in lower maximum soil
temperatures than if planted on the flat, but higher than if planted in
ridges. Planting in furrows results in lower maximum soil temperature than
if planted on the flat. Irrigation, when used, will also influence the choice
of planting form, mainly through considerations of field drainage and run-
off characteristics. Shallow plantings, if not mulched, should be followed
by one or more earthing-up operations before maximum crop cover is
achieved, in order to cover emerging stolons, and to reduce soil temperature
to favour tuberization.
~36
()
2...-
~ 32
"§
8. 28
E
2
'6 24
(f)
20+-~~~~~~~~~~--~
08:00 12:00 16:00 20:00 24:00 04:00 08:00

Time of day
Figure 15.4 Diurnal course of soil temperature (5 em depth) as influenced by

planting on the flat (e) or in ridges (0). Solid symbol could also represent mulched
soil if open symbol were non-mulched control. (Source: Midmore, unpublished).
Although the sun angle is high throughout much of the day in the
tropics, marked variation in soil temperature due to row orientation and
positioning within a ridge are experienced (International Potato Center,
1984a). Judicious choice of row orientation, and tuber placement, to
minimize soil heating in the vicinity of the tuber, lead to faster emergence,
but with little benefit on tuber yield. Under very hot soil conditions,
planting on the cool side of a ridge might represent the difference between
success or failure of the crop, and this technique is being studied in the
Sudan (El-Hassan, 1984). If, however, mulching or some other agronomic
practice to reduce soil temperature is available, then yields may be greater
if planting is made on the warm side of the ridge, since the foliage on that
side receives more irradiance available for conversion to dry matter, an
effect particularly important when complete crop cover is not achieved.
Once emergence has taken place, the foliage itself effects self shading
over the soil, with a corresponding reduction of soil temperatures (as crop
cover increases with time (Fig. 15.5a). Due to the short-season nature of
the potato crop in the lowland tropics (Midmore, 1983), the amount of
radiation effectively intercepted between planting and harvest, as a pro-
portion of that available over the whole crop cycle, is much less than that of
the corresponding temperate crop (Trebejo and Midmore, 1990). Hasten-
ing of emergence and achievement of full cover are therefore important
objectives not only from the viewpoint of reducing soil temperature, but
also to improve the efficiency of intercepting available solar energy
(Midmore, 1984b). Closer between-plant spacing, as a result of higher
populations, effected quicker crop cover over the soil and resulted in
greatly enhanced tuber yields (Midmore, 1988a), primarily as a con-
sequence of the extra irradiance intercepted. Mulch (Midmore et al.,
1986b) and irrigation (Trebejo and Midmore, 1990) treatments that
increased intercepted radiation over the crop season resulted in greater
tuber yield. Canopy duration (a cumulative measure of crop cover) was
also closely related to yield over a range of experiments in the lowland
tropics (Vander Zaag and Demagante, 1987) and in the semi-arid environ-
ment of Israel, (Levy et al., 1986).
(a) (b)
38
2: 34 1
....
Q)
.....~::J
....
Q) 30
c..
E
.....
Q)
·0 26
C/)
22+-~~--~-~--~~
08:00 12:00 16:00 20:00 10:00 14:00 18:00
10:00 14:00 18:00 08:00 12:00 16:00 20:00
Time of day
Figure 15.5 Soil temperature (5 cm depth) as influenced by crop cover (a) before
hilling (0, 15 dap; 6,20 dap; \l, 27 dap; 0, 32 dap) or (b) after hilling (0, 50 dap;
6,78 dap; 0, 84 dap) on clear days. (Source: Midmore, unpublished).
A clear understanding of the process of tuberization under high tem-
peratures has been the goal of many researchers, and it is perhaps this
process that has received more attention than any other related to the
adaptation of potato to hot climates. High temperature per se (28-30°C)
inherently retards the tuberization process (Nowak and Coleborne, 1989),
but if it is accompanied by short days, tuberization may take place in clones
characterized by a long critical photoperiod. These clones respond to the
shortening of daylength by an increase in the intensity of induction (Ewing,
1981) which overrides the negative influence of high temperature on
tuberization. Clones characterized by a long critical photoperiod are more
strongly induced to tuberize under short days than are clones with a shorter
critical photoperiod. In fact, clones with an extremely long critical photo-
period, when subjected to short-day conditions in the lowland tropics,
overcompensate for high temperatures and pass through their develop-
mental cycle with extreme rapidity, resulting in a short-season crop with a
very compact foliage and low yield potential (Midmore, 1983). This relates
to the earlier discussion on the type of genetic background adapted, in
terms of tuberization, to high temperatures. Andigena clones have a short
critical photoperiod, and hence no overriding influence of short days on
tuberization is apparent, whereas Tuberosum and Neo-tuberosum respond
to short days by a hastening of tuberization. Although the ability to
tuberize is therefore an important consideration in the adaptation of potato
to hot climates, selection for clones with extremely long critical photo-
periods might be counter-productive. Breeders should aim to achieve a
balance between the ability to tuberize under high temperature conditions,
and the foliage capacity to effect complete crop cover and maintain high
bulking rates.
Reynolds and Ewing (1986), working with whole plants and leaf bud
cuttings, were able to demonstrate clearly that the potential induction of
leaves to tuberize is principally affected by air rather than by soil temp-
erature, whereas the expression of the signal to tuberize may be blocked by
high soil temperature. These conclusions are supported by Sato (1981)
who, by varying air and soil temperature independently, observed that
tuberization at 30°C air temperature occurred when the soil temperature
was 24°C but not when it was 35°C. Data from field experiments in which
soil temperature was varied independently of air temperature (Midmore,
1984b) also indicated an earlier tuberization under cooler soil conditions
(30°C versus 37°C maximum): confirmatory evidence that the expression of
the tuberization stimulus is effected earlier under cooler soil conditions.
In a detailed study on the influence of high temperature, long photo-
period, and low irradiance on tuberization, Menzel (1984) concluded that
all three factors may suppress tuberization by increasing gibberellin
production. That plant morphology (small leaves, elongated internodes,
low tuber to shoot ratio) characteristic to plants grown under such condi-
tions may be mimicked through the exogenous application of gibberellic
acid (GA) led Menzel to this conclusion (Menzel, 1980). Although it is
unlikely that GA is the only plant hormone controlling tuberization
(Ewing, 1987), it does playa central role in the induction and expression of
tuberization. In his review of the role of hormones in tuberization, Ewing
(1987) has cited the non-exclusive role of gibberellins, their inhibitors,
cytokinins, and even auxins as components of the tuberization stimulus.
Certainly under the stress conditions mentioned herein, the plant response
most closely relates to the response to increased gibberellin content;
however this is only a result and not necessarily the cause of the suppressed
tuberization stimulus.
The interactions between temperature, photoperiod and irradiance do
have important practical applications. For example, in addition to the
long-day delay of tuberization, particularly when combined with high
temperature (i.e. the original basis for assuming that Tuberosum potato
clones would not adapt in the lowland tropics), the adverse effect of high
temperature on tuberization will be accentuated under conditions of low
irradiance (Menzel, 1985). Hence one should manipulate the interplay of
these three factors to improve potato production in the lowland tropics.
Correct choice of season (tropical winter with short days and high
irradiance (Fig. 15.2)), and of genotype (long critical photoperiod, absence
of symptoms of applied GA) are of paramount importance. Intercrop
shade should also be minimized prior to the time of tuberization. The
practice of shading the soil after planting potato (Fig. 15.3), which favours
emergence, should, if it results in shading of plant be terminated soon after
emergence and before the induction of tuber initiation, otherwise tuberiza-
tion is delayed (Midmore et al., 1988b).
Since the influence of high temperature on plant form and tuberization
may be mimicked by the exogenous application of GA, it is important to
enquire whether the application of growth retardants (e.g. CCC, ABA)
could reverse this process, as it can for long-day conditions (Hammes
and Nel, 1975). Studies in growth chambers (Menzel, 1980) confirmed that
the inhibitory effect of high temperature on tuber production under a
photoperiod of 14 h was almost completely reversed by the application of
CCC, and partially reversed by the application of ABA. Large scale field
applications of growth retardants under high temperature conditions
cannot be justified due to their inconsistent results (Menzel, 1984; Nelson
and Midmore, unpublished). A further factor that strongly influences the
timing of tuberization is the physiological age of the seed tubers, or more
specifically the sprouts, at the time of planting. Under temperate condi-
tions, plants from physiologically young tubers tuberize later than those
from old tubers of the same varieties. When temperatures are higher, data
of Menzel (1985) suggest that physiologically young plants have a greater
capacity for increased gibberellin synthesis in their buds than those of
physiologically older plants. Therefore the delay of tuberization by high
temperature will be proportionately greater in physiologically young
plants. This has important implications during the process of field selection
of clones for heat tolerance (as measured by the ability to tuberize), since
physiologically young plants with a long critical photoperiod will be
inseparable from physiologically old plants with a shorter critical photo-
period. Indeed, inferences on heat tolerance have already been influenced
by the physiological state of tubers at planting (Khedher and Ewing, 1985).
Screening clones for the ability to produce the tuberization stimulus
under high temperatures and long photoperiods, through the use of the
leaf-cutting technique (Ewing, 1981), aids in identifying clones that will
tolerate such conditions. Evaluation of the tuberization index in cuttings,
planted in heated soil, should improve the application of this technique
since screeening would not only be for the production of the stimulus, but
also for its expression. Screening progenies from reciprocal crosses of
parents with long or short critical photoperiods using leaf-cuttings (Lazin et
al., 1979), has illustrated the importance of maternal inheritance in the
transfer of heat tolerance. The methodology for screening many seedlings
in segregating populations for heat tolerance has also been studied by
Sattelmacher (1983). Periodic evaluation for tuberization in seedlings
momentarily removed from their small pots permiued classification of
seedlings into groups, based upon the earliness of tuberization. Ensuing
field experiments with the same material confirmed that the screening
technique was a good predictor of timing of tuberization and plant maturity
in the field. Similar techniques were used successfully by Levy (1984) in his
search for clones with the ability to tuberize in hot climates.
Various sources, other than S. tuberosum ssp. tuberosum and Neo-
tuberosum, in which tuberization is said to be favoured by short days and
high temperatures have been reported. S. commersonii (Davis, 1941), S.
antipoviczii, S. chacoense, and S. kesselbrenneri (Stelzner and Torka,
1940, as quoted by Dodds and Hawkes, 1952). S. phureja (Dodds and
Hawkes, 1952), and S. hygrothermicum (Ochoa, 1984) may all prove to be
useful in this respect. The crossing of S. phureja with Tuberosum has
already resulted in a number of heat-tolerant clones (Mendoza and
Estrada, 1979).
Both Ewing (1987) and Sale (1979) suggest that the net assimilate
balance available for plant growth may influence the tendency to tuberize;
high temperature reduces vigour and therefore delays tuberization. This
possibility extends the concept of heat tolerance to include not only the
ability to tuberize, but heat tolerance as it relates to net productivity under
high temperature conditions. In its simplest terms, the concept of selecting
for increased plant vigour, i.e. large leaves, haulms and thick stems (Ewing
et al., 1983), integrates the influence of high temperature on productivity.
Selecting for increased vigour considerably improves the success of selec-
tion for high tuber yield under hot conditions when combined with
selection for tuberization.
In 1925, Bushnell developed a schematic approach to the influence of
temperature on the daily carbohydrate balance, which has been little
improved upon over the last 60 years. Referring to night temperatures,
Bushnell concluded that of the total carbohydrate produced during the
day, the amount available for growth and storage was seven times greater
at 20°C than at 29°C. Since the accumulation of carbohydrate during the
day was essentially similar at 20°C and 29°C, the night temperature,
through its effect upon respiration, had an overriding influence on the net
dry matter accumulation. More recent data collected by Winkler (1971)
indicate a doubling of dark respiration per 10°C rise over the range of 10 to
30°C, and led Winkler to suggest that the net assimilation rate would be
negative at temperatures greater than 37°C. Coupled with the higher rate
of dark respiration at higher temperature is the tendency for stem growth
to be favoured at the expense of leaf growth (Burton, 1966). The leaf area
per unit total dry weight is less; therefore photosynthetic production per
gram of respiring haulm decreases with age, an effect accentuated by high
temperature (Winkler, 1971). Genotypic differences in maintenance res-
piration identified with the dark starvation method have been reported by
Lorenzen and Ewing (1987), and work in progress (Midmore, unpublished)
is directed in the search for intra- and inter-specific variation in this
character.
The rate of photosynthesis is less responsive than respiration to tem-
perature over the range within our discussion. Only when the leaf
temperature exceeds 32°C do gross photosynthetic rates decline (Dwelle et
at., 1981), although the net rates of photosynthesis are optimal over the
range 16-25°C (Ku et at., 1977).
More recent data based on the RCC provide an indication of tempera-
ture effects on growth efficiency. That high temperature reduces the RCC
has already been suggested through the reworking of Sales' (1973a,b, 1974)
data by Scott and Wilcockson (1978). The reduction of the RCC at higher
temperatures was attributed to enhanced rates of respiration. Halving the
rate of respiration during the final 30 days prior to harvest brought the
RCC in line with those of cooler climates. Lower RCC than those of cool
climates have been reported in other hot environments (Midmore, 1984a;
Nelson and Midmore, 1986; Trebejo and Midmore, 1990). Allowance for
the influence of temperature on the RCC through an empirical tempera-
ture correction factor (Haverkort and Harris, 1987) or with the 'photo-
thermal quotient' concept «MJ m- 2 day-l)/(mean to - base to) - Trebejo
and Midmore, 1990) permits a normalization of data collected in environ-
ments of contrasting temperatures.
The concept of the RCC has also been applied to the slope of the
relationship between tuber dry matter and cumulative radiation inter-
ception (Midmore, 1987; Vander Zaag and Demagante, 1988). In the
Philippines, in a group of superior clones for which the harvest index did
not vary greatly (Vander Zaag and Demagante, 1988), the tuber RCC was
30% less than under cool climates; whereas in the mid-elevation jungle of
Peru (Midmore, 1987), tuber conversion efficiency, although less than in
cool climates, was largely dependent upon maturity class. Early maturing
clones, due to greater partitioning to tubers, were apparently more
efficient than later maturing clones. In formulating guidelines for the
selection of heat-tolerant clones, techniques that highlight differences
between clones in their net productivity under high temperatures and that
relate to low rates of respiration might have wide application.
The quest for characters related to heat tolerance, as they apply to net
productivity at high temperatures, has involved numerous screening tech-
niques. Some of these have been documented by Ewing et al. (1987) and
range from measures of membrane integrity to the screening of biomass
production under non-inductive conditions for tuberization. However, in
common with other techniques relating to morphological characteristics
(International Potato Center, 1987), photosynthetic rates (Wivutvongvana,
1979), and heat-killing time (Chen et al., 1982), no positive results of direct
use to the plant breeder have been obtained. Holding more promise is th,e
selection for low rates of respiration (Wivutvongvana, 1979) and the
selection for fast rate of callus growth of undifferentiated cells on culture
medium (International Potato Center, 1987). Results with the use of the
chlorophyll fluorescence technique, although relating well to known tole-
rances to heat based upon species distribution (Smillie et al., 1983) and to
tuber yield under high temperature conditions (Hetherington et al., 1983)
might, together with the heat killing time method (Chen et al., 1982), be
more useful in the selection of clones with combined tolerance to heat and
drought. This is suggested since the temperatures at which assessments are
made are greater than those which leaves might experience under well-
watered field conditions. Short term high temperature (>35°C) shocks
effect a physiologically different response than does longer-duration high
temperature «35°C) stress (Wu and Wallner, 1984), which may have
important implications for the selection of appropriate screening tech-
niques to evaluate heat tolerance. The involvement and level of heat shock
protein synthesis at temperatures >35°C far exceeds their production level
at lower temperatures (Wu and Wallner, 1984). It is difficult to reconcile
the various approaches used when the chlorophyll fluorescence technique
has been applied as a screening method. Screening of plants grown at a
constant high temperature (35°C) might identify clones that acclimatize
well to high temperature (Sipos and Prange, 1986), due perhaps to a
moderate increase in heat shock protein synthesis, whereas the screening
of non-acclimatized plants after a short high-temperature stress (>40°C)
would identify clones which incurred cellular injury in the absence of heat
shock protein synthesis (Smillie et al., 1983). Both techniques have
correlated well with measures of growth or known heat tolerance, but
definitive relationships with dry matter accumulation under high temp-
eratures have yet to be proven.
More recent data (Prange, pers. comm.) suggest that chlorophyll
fluorescence studies are able to differentiate between the effects of 30/20°C
and 20/10°C day/night temperature regimes, long before marked dif-
ferences in dry weight accumulation occur. The T, or terminal, value was
particularly responsive to growing temperatures, as in a previous study
(Sipos and Prange, 1986). Another feature (Prange, pers. comm.) was the
insensitivity of the rate of respiration to growing temperature over the
above-mentioned range, somewhat in contrast to the widely held opinion
that respiration, and not photosynthesis, limits dry matter accumulation at
supra-optimal temperatures.
The possible interrelationship between heat and cold tolerance in potato
species or cultivars has caught the attention of a number of researchers.
Chen et al. (1982), using the electrolyte leakage method, found no
systematic relationship between freezing and heat tolerance among the
tested species, neither at normal growing temperature (20/15°C, day/night)
nor after heat (35/35°C, 1 day) or cold (2/2°C, 14 day) conditioning. Heat
acclimatization occurred in all species but did not enhance cold tolerance,
and, likewise, cold acclimatization did not increase heat tolerance with the
exception of S. commersonii. In another study by Smillie et al. (1983) in
which plants were neither heat nor cold acclimatized, heat and cold
tolerance (using the chlorophyll-fluorescence technique, at 41°C and O°C)
of wild potato species were not highly correlated, suggesting that cold and
heat tolerance can vary independently of each other in potatoes in the wild,
and that the degree of stress tolerance is largely dictated by the environ-
mental influence upon evolution. Data from similar studies, confined to the
cultivated species Tuberosum and its crosses with other cultivated species
(Hetherington et al., 1983), indicated positive correlations between heat
and cold tolerances, in contrast to the relationships in wild species.
Apparently, warm-adapted clones (i.e. selected for their ability to produce
tubers in warm climates) were not selections with specific heat tolerance,
but possessed an enhanced generalized stress tolerance. The range of
variation in heat tolerance was limited in the Tuberosum crosses, and the
authors concluded that only limited potential exists for improving this
character. Of the cultivated primitive species, S. phureja was identified as
the most heat-tolerant, confirming reports of Moreau (1944), and of
considerable practical importance since heat-tolerant selections DTO-2
and DTO-33 contain S. phureja in their parentage (Mendoza and Estrada,
1979).
High temperature not only influences the RCC in the potato crop, but
also the amount of canopy light interception, and the partitioning of dry
matter to tubers (Midmore and Rhoades, 1988). Crop cover of clones that
tuberize rapidly in hot conditions declines rapidly following the achieve-
ment of maximum cover (Fig. 15.6; Vander Zaag and Demagante, 1988),
with concomitant soil heating (Fig. 15.5b). Since differences between
clones in the conversion efficiency are not significant (Nelson, 1987;
Vander Zaag and Demagante, 1988) which tends to confirm the conclusions
100
80
~
Qi 60
1)
~ 40
e
() 20
O+----,~---.----T_---,--~~
o 20 40 60 80 100
Days after planting
Figure 15.6 Time course of crop for four clones grown in a warm site in Peru.
o early maturing clone DTO-33; /':,., early maturing clone Desiree; D, medium
maturity clone Revolucion; \j, late maturity clone Tomasa Condemayta. (Source:
Nelson, 1987.)
of Hetherington et al. (1983) on the genetic variability for heat tolerance

(based on chlorophyll-fluorescence readings) present in the tetraploid
potato, a realistic objective for yield increase may be to maximize the
interception of radiation during tuber bulking. Short leaf longevity result-
ing in a low maximum number of green leaves under high temperature field
conditions (Nelson, 1987) restricts the size and duration of crop cover, but
initial studies have not revealed genetic variation for this character (Fig.
15.7). High temperature alters the morphology of the potato plant (see
Midmore and Rhoades, 1988) and results in a less efficient interception of
solar radiation per unit of leaf area index (Midmore, 1988a), leading to
incomplete crop cover and inefficient use of available solar energy. High
average air temperature (28°C) under natural long days in The Netherlands
(Marinus and Bodlaender, 1975) led to earlier senescence of leaves than at
22°C or 16°C. However, due to the inhibition of tuberization at the highest
temperaure, leaves were formed over a longer period than at 22°C or 16°C.
The particularly high temperatures (40/30°C, day/night) imposed in the
study of Khedher and Ewing (1985) also reduced leaf longevity but delayed
tuberization substantially; hence canopy senescence was later than under
cooler temperatures due to the extended production of leaf nodes.
The rapid decline following maximum crop cover represents a waste of
solar energy available to the potato crop, and an important heat source
inhibiting the accumulation of dry matter in the tubers (Midmore, 1984b).
For this reason, practices that shade or cool the soil during that period such
as soil reflect ants (Midmore, 1984b), mulches (Midmore et al., 1986b), and
shade (Midmore, 1988b; Midmore et al., 1988) often result in an increase
in fresh yield and greater percent of dry matter content. Detailed studies of
tuber growth under high temperatures have shown a decline in total tuber
dry weight long before complete maturity (Vander Zaag et al., 1986;
o 20 40 60 80 100
Days post-emergence
Figure 15.7 Leaf production (---) and leaf senescence (----) of three clones
grown in a warm site in Peru. D, Desiree; 0, Revolucion: 6, Mariva. (Source:
Nelson, 1987.)
Midmore, 1988a) which highlights the need to either harvest before crop
maturity or to reduce the soil heating during senescence.
Numerous reports have indicated the quantitative decline in tuber dry
matter content due to the increase in growth temperature (Marinus and
Bodlaender, 1975; Khedher and Ewing, 1985). Rates of 1% or 0.45%
decline per degree mean rise in ambient temperature (over the range 15 to
25°C and 15 to 22°C respectively) have been reported (Espinola et ai.,
1984; Haverkort and Harris, 1987). Apparently tuber growth is under
environmental control similar to that governing tuber initiation (Ewing,
1981); however, the hormonal control of starch accumulation is as yet
unknown. Based upon studies using 14C, Krauss and Marschner (1984)
suggest that high soil temperature inhibits the conversion of sugars to
starch, probably through depression in the enzyme activity involved in
starch metabolism. No evidence for differences between varieties was
found. A similar conclusion on the role of starch synthesis was reached by
Villareal (1976), who observed that the total activity of sucrose synthetase
was considerably lower at night temperatures of 20 to 25°C, than at
temperatures of 10 to 15°C. This effect on sucrose synthetase could be
reversed by changing from high to low night temperatures. Lower starch
accumulation in tubers at high versus lower soil temperatures (Randeni
and Caesar, 1986) would tend to confirm that the influence of temperature
on tuber dry matter content is mediated through the control of conversion
of ethanol-soluble carbohydrates to starch.
Krauss and Marschner (1984) noted no differences between heat-
tolerant and non-heat-tolerant clones in their response to high temperature
as measured by incorporation of 14C into ethanol insoluble fractions;
similarly no differences were observed by Espinola et ai. (1984) between
clonal response of percent tuber dry matter to increase in growing
temperature. In a larger study, Gaur and Gupta (1981) also found no
evidence for significant genotype-by-environment interactions for tuber
dry matter. The generally positive relationship between maturity class and
dry matter content observed by Maris (1969) and Simmonds (1974) in cool
climates was also evident to some extent in hot climates (Espinola et al.,
1984; Nelson, 1987), but in a study by Haynes and Haynes (1983) in whkh
sources of high specific gravity under hot conditions were identified, no
mention of their relationship to maturity class was made. Selections from a
S. phureja x S. stenotomum cross maintained their high dry matter content
over the two high temperature sites, and dry matter contents exceeded
those of commercial tetraploid cultivars.
Stress, whether due to heat, drought, cold, or insects, is known to
increase the accumulation of glycoalkaloids in tubers (reviewed by Morris
and Petermann, 1985). Bitterness is closely correlated with glycoalkaloid
content (Sinden et al., 1976) and was reported by Espinola et al. (1984) to
be a frequent comment in tests on taste assessments of potato produced
under hot conditions. Analysis of total glycoalkaloid content of a number
of clones grown under the hot jungle conditions of Peru (Midmore and
Bushway, unpublished) showed that in some cultivars total glycoalkaloids
exceeded 25 mg per 100 g fresh weight. Glycoalkaloid content of tubers
harvested from plants grown in uniform potting mixtures at three sites
within Peru ranged from 7.1 through 18.4 to 20.7 mg per 100 g fresh tuber
for mean growing temperatures of 17.1, 21.8 and 25.6°C, respectively. In
addition to the direct effect of high temperature on tuber glycoalkaloid
content, the tendency to tuberize close to the soil surface (Midmore,
unpublished), the incomplete crop cover over the soil at high tempera-
tures, and the presence of wild species in the genetic background (Mendoza
and Estrada, 1979) would all tend to favour the accumulation of glyco-
alkaloids in hot climates (Sinden et al., 1984).
The foregoing account of high temperature and its influence on growth
and development of the potato is not exhaustive, yet it demonstrates the
complexities faced by plant breeders in their search for heat-tolerant
clones. Although many of the reports on the physiological response of the
potato crop to high temperature give the breeder an insight into the
possible mechanisms of heat tolerance, they do little to supply techniques
that will take the screening process out of the uncertain field environmental
conditions and into the laboratory. In the final analysis, the evaluation of
clones for heat tolerance must be carried out in the field, yet, at present the
basic screening is also undertaken in the field. This practice has much to
favour it, since heat tolerance is not only the physiological ability of the
plant to grow and form acceptable tuber yield under adverse conditions of
high temperature, but also the facility for the plant to contend with
additional biotic and abiotic stress. For these reasons, most selection is still
made at the field level but the choice of parents for initial crossing is
intuitively built upon a foundation of physiological principles whenever
possible (Mendoza and Estrada, 1979). Much basic screening is required
for both cultivated and wild species in order to identify useful sources of
genes that may later be transferred to the potato to improve its adaptation
to hot conditions.
In spite of the lack of direct application of physiological selection
criteria, genetic advances have been made in the quest for heat tolerance.
Following the strategy for population breeding, Mendoza and Estrada
(1979) have enhanced the frequency of clones for adaptation, high yield
and pest resistance. In studies by Mendoza (1984), results showed that
heritabilities for yield per plant, average tuber weight, and foliage maturity
ranged from 0.16 through 0.45 to 0.79, and were considered conducive to
genetic advance. Various sources of germplasm were used (e.g. Tuberosum,
Neo-tuberosum, S. phureja, S. stenotomum, S. chacoense, and S. sparsipilum)
and widest adaptation and maximization of yield have been attributed to
enhanced genetic divergence between the germplasm of the adapted
parental clones. Plaisted et al. (1987) have identified numerous clones
selected for heat tolerance that were derived from genetic material of this
breeding programme.
After one cycle of phenotypic recurrent selection in diploid populations
of S. phureja and S. stenotomum increases in survival, tuberization, and
yield ranged from 3% through 15% to 27%, respectively; however,
heritability based on data for three combined locations gave a low
heritability value of 0.07 for tuber yield (Gautney and Haynes, 1983).
Working under high temperature conditions with the same population,
Haynes et al. (1986) reported a marked increase in tuber dry matter
content of offspring over parental values. This source of material should
therefore prove useful in attempts to raise tuber dry matter content in hot
climates.
Other approaches in the search for heat-tolerant clones have con-
centrated on the cultivated Tuberosum potato. Selection for tuberization
under high temperature in open-pollinated progenies of S. tuberosum
(Levy, 1984) has led to some success, and clones equal to or better than
control clones have been identified in Israel. Another example of success
from a breeding programme with major emphasis on S. tuberosum parental
material has been the selection of the cultivar Arma by Sunoschi et al.
(1987). Based upon a cross between a female heat-tolerant parent and the
cuJtivar Maris Piper, the resistant cultivar Arma exceeded the local control
cultivar Desiree, which itself is well known for a degree of heat tolerance
(Midmore, 1984b, 1987).
Field-screening of a range of clones with varied genetic background
(Midmore, 1987; Vander Zaag and Demagante, 1988) has also indicated
further sources of heat tolerance, ready for wide-scale testing.
(b) Water
Water stress may take one of two forms: excess or deficit. During the hot,
tropical rainy season, the inhibition of emergence due to waterlogging at
high temperatures (Jackson, 1962), coupled with the sensitivity of potato
tuber growth to lower levels of oxygen in the soil (Cary, 1985) preclude the
possibility of cultivating potatoes then. Exceptions are rare and exploit local-
ized edaphic conditions, e.g. highly porous soil or raised beds, as for tropical
vegetable production (Tumuhairwe and Gumbs, 1983). A more common
limitation, particularly during the lowland tropical winter, is the lack of water.
In addition to the direct physiological effects, for example, on the expansion
of leaves (Gregory and Simmonds, this volume, Chapter 5), drought com-
pounds high temperature effects on photosynthesis and respiration.
The field response of the potato crop to drought may be interpreted
within the framework of the light-interception/crop-growth model. Inter-
ception of radiation by the crop is inevitably reduced by the lack of water.
Emergence is delayed, or even suppressed, depending upon the degree of
drought stress (Midmore and Rhoades, 1988), but no guidelines as yet
relate successful emergence and minimum soil water availability. The
result of reduced leaf expansion is manifest in a smaller crop cover,
resulting in less cumulative interception of radiation throughout the crop
season. Although the physiological processes involved in the reduction of
leaf expansion (e.g. osmotic control of cell expansion, reduced distribution
of dry matter for leaf growth) are probably similar to those under cooler
climates, the rate of increase in soil moisture deficit under high tempera-
tures, especially where the saturation vapour pressure deficit (SVPD) is
high, develops faster than under cooler conditions with associated faster
decline in the rate of leaf expansion (Midmore and Rhoades, 1988). Local
control of evaporation from the soil surface through the use of vegetative
mulches or shade favoured crop emergence (Midmore et al., 1986a,b,
1988) and crop cover, although this effect was not separable from the
simultaneous benefit due to reduction of soil temperature. Once the
depletion of soil moisture in mulched plots matched that of non-mulched
plots, the earlier advantage in crop cover attributed to the conservation of
soil moisture by mulch was not maintained, since the size of the crop
canopy was apparently adjusted according to the available soil mositure
(Devaux and Haverkort, 1987). Tuber yield, however, was proportional to
the cumulative radiation intercepted, which was almost doubled by the
application of mulch.
Failure to achieve maximum crop cover and the quick reduction in cover
due to drought, lead to an increase in soil exposed to incoming radiation. If
the soil surface is dry, no appreciable loss of soil moisture through
evaporation is effected, but the increase in reflected sensible heat results in
enhanced heat stress (Bidinger, 1980). Upon re-wetting of the soil surface,
however, evaporation from the soil surface increases in importance tem-
porarily as a component of evapotranspiration (ET). Closer between-row
spacing and higher fertilizer application rate favouring rapid canopy
development, and the use of vegetative mulches, would tend to increase
the water use efficiency under such conditions.
It is generally recognized that under temperate conditions reduction in
the rate of leaf expansion through reduced turgor pressure due to drought
precedes measurable effects upon transpiration and photosynthesis. In-
deed, under temperate field conditions reduction of tuber yield due to
drought is more closely related to the reduction in radiation interception
(Bean et al., 1984; Jefferies and MacKerron, 1987) than to modification of
the RCC. Once the rate of transpiration cannot supply the demand along
the vapour pressure gradient from the leaf to the air, stomates close and
leaf temperature rises above air temperature. During the midday period,
when the vapour pressure gradient is greatest, leaf temperatures of 8 to
lOoC greater than air temperature have been reported (Lomas et al., 1972;
Stark and Wright, 1985). In the presence of drought, raising the leaf
temperature resulted in a decrease in the rate of net photosynthesis
(Bodlaender, 1980). Drought at high temperature may result in longer
term effects attributable to permanent damage to the photosynthetic
apparatus, but in temperate climates this seems unlikely, since rates of
photosynthesis and transpiration in plots following release from drought
equalled those of non-stressed plots (Vos, 1986) as did bulking rates
(O'Brien and Allen, 1984). Under well-watered conditions photosynthetic
rates were maximum at 24 to 30°C (Dwelle et al., 1981) and, despite no
associated reduction in stomatal conductance up to 35°C, carbon assimila-
tion was reduced substantially at that temperature. The photosynthetic
quantum efficiency of barley declined at leaf temperature >25°C (Parkinson
and Day, 1983), and the same effect was probably responsible for
reduction in photosynthesis of potato when leaf temperature exceeded
30°C. Some evidence for a reduction of the RCC due to drought may be
had from data of Khurana and McLaren (1982) which show a lower
conversion value (1.32 g total dry weight Mrl) in the year with drought
than in the year without (1.81 g MJ- 1). Data reported by Trebejo and
Midmore (1990) confirm that drought stress can reduce the RCC. Using
the line-source design, a significant reduction of the RCC was effected by
the droughted treatments (Table 15.6). Harvest index (HI) was greater in
droughted than control plots under hot conditions (Table 15.6), whereas in
temperate climates HI is generally not significantly altered by drought
(Jefferies and MacKerron, 1987). Drought under high temperature has-
tened senescence (Table 15.6), but paradoxically Shimsi and Sunoschi
(1985) reported that the lowest yielding cultivars were those which
maintained foliage longer than other cultivars, even under water stress,
and attributed this to the diversion of assimilates to foliage instead of to
tubers.
Restricted canopy expansion and lower quantum efficiency severely limit
yield potential when potato is subjected to drought in hot climates.
Average reduction (50%) in tuber yield due to drought was greater than
the reduction (35%) in irrigation (Trebejo and Midmore, 1990). Water use
efficiency (WUE) in a desert environment, and calculated as final tuber
Table 15.6 Influence of irrigation supply on fresh tuber yield and
yield-water supply ratio in a desert climate of southern Peru
Applied irrigation water (mm)
970 840 760 690
(a) Tuber yield (t ha-1) 62.1 55.0 37.4 25.2
Yield/water ratio (kg m-3 ) 6.4 6.5 4.9 3.7
Applied irrigation water (mm)
584 453 378
(b) Tuber yield (t ha-1) 25.4 18.6 12.1
Harvest index (0 --49 dap;
56 dap%) 22.0 25.3 28.7
RCC* (g MJ- 1) 1.74 1.31 0.93
Maturity (25% crop cover,
- dap) 92.0 87.0 80.0
Yield/water ratio (kg m- 3) 4.7 4.4 3.4
• Radiation conversion coefficient.
Sources: (a) Versteeg (1985), means of two varieties; (b) Trebejo (1987), means
of three varieties.
yield at harvest per unit volume of irrigation, was greater in well-irrigated

than droughted plots (Trebejo and Midmore, 1990, and Table 15.6). The
efficiency of interception of radiation was greater per unit of applied water
in droughted plots (Trebejo and Midmore, 1990): therefore, based upon
the slope over the linear phase of the regression of dry matter accumula-
tion on cumulative applied irrigation water, droughted plots had a higher
WUE (i.e. transpirational WUE). The observation (Vos and Oyarzun,
1987) that stomatal conductance declined faster than photosynthetic capa-
city in the initial stages of drought may in part also explain the above noted
improvement in transpirational WUE of droughted plots. WUE values
based on final yield may put at a disadvantage treatments that induce early
maturity, and early maturing clones, in which tuber yield reaches a
maximum and then declines before final maturity (Trebejo and Midmore,
1990). Nevertheless, WUE is often quoted on the basis of final tuber
yields. Data from hot dry conditions show that WUE, depending on the
degree of drought, was on average two to seven times lower than WUE of
identical clones in cooler conditions with a much lower SVPD (Trebejo
and Midmore, 1990). Under well-watered conditions the WUE based upon
final yield data was approximately one-third that of cooler conditions (4.4
kg m- 3 versus 12.4 kg m- 3 ). The value under hot conditions compares well
with the recalculated 4.2 kg m- 3 of Vander Zaag and Demagante (1985) in
tropical Philippines, both somewhat lower than the 5-7 kg m- 3 calculated
by Doorenbos and Kassam (1979) as representative for the potato crop,
and much less than the reported values in areas of low night temperature
and high daily solar radiation, e.g. 11.7 kg m- 3 in Victoria, Australia (Rab
and Willat, 1987) and 9.0 kg m- 3 in Israel (Shalhevet et ai., 1983).
Incomplete canopy cover and frequent wetting cycles of the soil surface
are conducive to significant water loss through evaporation< from the soil
surface, with a concomitant reduction in WUE. Drip irrigation systems,
besides improving WUE through reduction in the soil evaporation com-
ponent of ET (Vander Zaag and Demagante, 1985), also modify root
distribution such that plants may withstand a greater degree of drought
before yield reduction is evident (Shalhevet et al., 1983). Evaporation
comprised 13% of total ET with a sprinkler irrigation system but only 4%
under drip irrigation (Shalhevet et al., 1983). Nevertheless, under certain
circumstances, for example where deep percolation of localized water
application exists, water requirements for trickle irrigation may be greater
than those for sprinkler irrigation (e.g. as reported in Cyprus by Stylianou
and Orphanos (1981)). Additional considerations such as the increase of
tuber moth and growth cracks under drip irrigation might be of overriding
importance in the choice of irrigation systems. Frequent irrigation, if water
is readily available, may be of considerable benefit to the potato under hot
conditions, due to the direct cooling effect on the soil, and if applied by
sprinklers to the foliage.
Burgers and Nel (1984) suggested that the high irrigation requirements
of potatoes may more often be ascribed to the cooling effect of the
frequent water applications, than to sensitivity to soil-moisture stress, and
concluded that the cooling effect could be obtained more cheaply by
mulching. Mulch reduced the estimated water use by 30% in well-watered
plots but only 17% in plots which were less frequently imgated. Ironically,
mulch has the greatest benefit in terms of conservation of soil moisture
when irrigation frequency is high, whereas the major demand for economies
in conservation of soil moisture is when irrigation frequency is low. Under
very well-watered conditions mulch was ineffective in increasing tuber
yield whereas under conditions of reduced irrigation, where soil cooling
was not effected by irrigation, tuber yield was significantly greater in
mulched than control plots (Vander Zaag and Demagante, 1985).
Another practice that has found favour amongst investigators is the use
of antitranspirants: chemicals which due to their physical nature, reduce
the rate of transpiration. Water consumption in pot experiments with
Folicote (a wax emulsion antitranspirant) was reduced by 20--40% (Lipe
and Thomas, 1980). Fresh tuber yield increased by 2.3-5.0 t ha<l in
experimental plots and by 5.3 t ha<t in farmer's fields, in response to
maintenance of soil moisture following applications of Folicote. Applica-
tions close to maturity gave maximum yield response, at a time when
stomatal conductance was still high, but when photosynthetic activity was
suboptimal due to the inherent age of the canopy.
Selection for drought tolerance is a high priority in the adaptation
process of the potato to the lowland tropics. Advance is slow because
the type of drought tolerance sought will influence the choice of tech-
niques suitable for the identification of drought tolerance. For example,
drought-tolerant clones in temperate climates might have only to contend
with short-term periods of drought after which water supply resumes. In
contrast, in many tropical climates, where the infrastructure for irrigation
is non-existent, drought-tolerant clones must often tolerate a drought
stress that increases in intensity with time. An example would be the winter
crop in the Gangetic plains of Bangladesh which exploits a receding water
table. Clones tolerant to transient drought may be characterized by
'conservative' avoidance mechanisms, such as an extensive root system
(Coleman, 1986) and osmotic adjustment (Jefferies and MacKerron,
1987), which permit maintenance of high leaf areas. Clones tolerant to a
progressive increase in drought are best characterized by 'opportunist'
mechanisms, such as earliness (Sunoschi and Shimsi, 1985; Versteeg, 1985)
and high stomatal conductance. Evaporative cooling of the leaf surface as a
result of high stomatal conductance should prevent heat damage of the
photosynthetic apparatus at a time when leaf temperature would otherwise
rise markedly above air temperature. Studies aimed at relating single
physiological indices (e.g. rate of photosynthesis, leaf conductance) to
clonal yield response under field water stress have largely been unsuccess-
ful due to the complexity ofthe problem (Levy, 1983; Shimsi and Sunoschi,
1985). Nevertheless, an indirect measure of the size of root systems (i.e.
the force required to pull plants out of the soil) does appear to be related to
yield ability under drought conditions (Ekanayake and Midmore, 1989),
and this methodology is being developed in the search for drought
tolerance. With our present state of knowledge, field experiments aimed at
screening in situ for yield per se under conditions of heat and drought stress
(Levy, 1985; 1986; Ekanayake and Midmore, 1988) are likely to be of more
use to the breeder in identifying tolerant parental clones than any single, or
combination of, physiological indices. A similar conclusion for the selec-
tion of drought-tolerant potato clones in temperate climates has been
reached (Beekman and Bouma, 1986). Their simple technique consists of
screening in the greenhouse for the potential for recovery following a
period of severe drought stress.
(c) Mineral nutrition and toxicities

Inasmuch as the potato crop in hot climates is of shorter (70-100 day)
duration than potato in temperate climates, timing and absolute require-
ments for mineral nutrients may be expected to differ. One study,
conducted in an isohyperthermic environment of Hawaii (Manrique et al.,
1984) did show that nitrogen (N) and potassium (K) uptake exceeded
application rates by 1.5 and 2.5 times respectively, and highlighted the
lack of information on the nutrient requirements for the potato crop in hot
climates. There is a dearth of information on the mineral requirements of
potato in hot climates; what is available almost exclusively relates to
fertilizer N response. Optimal levels of soil N at the time of planting, or
shortly thereafter, are important for rapid early canopy growth which
intercepts solar radiation for conversion to dry matter, reduces soil
heating, and reduces the soil evaporation component of evapotranspira-
tion. An adequate supply of soil N later in the season may be effective,
following achievement of maximum cover, in the delay of the characteristic
rapid decline in crop cover in hot climates. Soil, and later staggered foliar
applications of N after tuberization (40 kg ha- 1 at planting plus four foliar
applications of 9.2 kg ha- 1 each over the period 40 to 70 dap), has been
shown effectively to delay senescence under hot field conditions in
Bangladesh (Chaudhury et al., 1984) and resulted in a significant 10%
tuber yield increase. Splitting N application into two or three soil applica-
tions resulted in no yield benefit over a single N application, presumably
due to dry conditions prevalent after planting. Positive yield benefits due to
the splitting of N into two or more applications are rare, and generally only
occur under conditions conducive to rapid leaching. For example, in the
wet season in the high jungle of Peru (International Potato Center, 1987),
although no differences in yield were observed between single and split
applications for an early harvest, splitting the N in two applications
significantly increased final tuber yield compared to a single application,
for each of the four N rates (60,120,180 and 240 kg ha- 1). In the dry season
on the same soil type, although there was again a yield response to
increasing levels of fertilizer N, split application of N did not result in yield
greater than that of a single N application. Mid-season tuber yields of an
irrigated crop (Table 15.7) were greater in treatments with little initial N
application. Maximum tuber yield was achieved 80 days after planting in all
but the treatment in which N application was split into four parts (Table
15.7). Splitting the N application into four parts did not increase the total
intercepted radiation by the crop canopy, but did delay achievement of
maximum crop cover and the rate of foliar senescence. A similar response
to single applications of fertilizer N was observed in the dry season in
Trinidad (Chapman, 1965). In a comparison between 0, 75 and 150 lb N
Table 15.7 Effect of single or split N

applicatiO<lil (total 160 kg ha-1) on tuber yield
(t ha-1) 60, 70, and 80 dap in a warm climate
of Peru
Harvest date (dap)

Number of split
applications 60 70 80
1 18.1 33.9 30.4
2 16.5 31.0 27.7
3 15.5 28.3 28.2
4 19.9 26.3 29.9
Source: International Potato Center (1986).

Table 15.8 Influence of rate of nitrogen fertilizer (kg ha-I )
on (a) leaf area index (LAI) and (b) tuber yield (t ha-I fresh
weight) of the cultivar Patrones planted in Trinidad
(a) LAI (b) Tuber yield
N rate N rate
Sample date 0 84 168 0 84 168
13 Feb. 0.73 0.83 0.71 0.0 0.0 0.0
25 Feb. 2.80 3.75 3.57 2.4 2.4 1.9
10 Mar. 2.81 4.28 5.39 9.1 14.1 11.5
24 Mar. 1.81 2.40 2.62 20.2 20.6 20.6
07 Apr. 0.51 1.25 1.31 20.3 26.0 32.8
Source: Chapman (1965); by permission of the publishers,
Butterworth & Co. Ltd.
acre- 1 peak LAI was greater for the intermediate and higher rates than for
the control (Table 15.8) and the decline in LAI was delayed relative to that
of the control, resulting in greater tuber yield in the latter stages of
bulking.
On soils where there was no economic response to N fertilizer rates
above 30 kg ha- 1 (Mauritius Sugar Industry Research Institute, 1984) no
yield increase with split application of N was recorded. On red lateritic
soils of Cuba where potato is grown as an irrigated 'winter' crop, although
a positive response of tuber yield to 100 kg N ha- 1 was noted, split N
application was without significant effect (Argotte and Herrepa, 1987).
When application of a vegetative mulch at planting is warranted, obvious
practical benefits accrue with the single applications of fertilizer. Addition-
ally, the application of all fertilizer at planting, in combination with a
mulch cover, would obviate the need to earth-up, thereby preventing
infection by Pseudomonas solanacearum or other pathogens which gain
entry through damaged roots. Root growth is thermophobic and, in soils
that heat considerably, root growth would be directed away from the upper
fertilizer-rich zone of the soil. A mulch covering the soil surface, through
reduction of soil temperature close to the surface, will concentrate root
growth in the upper horizon of the soil (Midmore, 1984b) and, in
combination with conservation of soil moisture there, facilitate an efficient
uptake of mineral nutrients.
The efficiency of uptake of fertilizer is also influenced by the method of
application, as illustrated by data of Ahmad et ai. (1982). Broadcasting
NPK on a heavy soil following paddy rice gave greater tuber yields (18.5 t
ha- 1) than application below the seed tuber (12.9 t ha- 1), on one side of the
row (10.9 t ha- 1) or on both sides of the row (10.5 t ha- 1). Besides early
growth advantages, the broadcast treatment also resulted in the greatest
proportion of relatively large tubers. Source of nitrogen (ammonium
sulphate, ammonium nitrate, or urea) has been reported not to influence
the efficiency of uptake (Valdes et at., 1982) although a slight economic
benefit was evident with the sulphate source. The effect of high N rates on
harvest index and on the percentage tuber dry matter are consistent with,
and in addition to, those of high temperature (International Potato Center,
1988). Particular care must therefore be exercised to effect the appro-
priate balance through N application between foliar growth and tuberization.
The relationships between plant density, cuitivar, and water and fertil-
izer supply deserve further attention before generalized relationships
between fertilizer application and crop cover can be derived. As indicated
by Harris (this volume, Chapter 4) for temperate climates, fertilizer
response may interact with the other three factors.
In the lowland and mid-elevation tropics 30% of soils are known to be
acidic, with pH <4.5 and a high aluminium (AI) saturation of the cation
exchange capacity (Sanchez, 1976). The potato root system, although
considered relatively tolerant amongst crop species to soil acidity, is
damaged by soluble Al (Lee, 1971); hence mineral nutrition and water
uptake are restricted, and yield potential reduced. Lime application is
necessary to neutralize exchangeable aluminium, to permit the develop-
ment of a more extensive root system. Little benefit of lime is to be
expected if soil pH is >4.9, or if the concentration of extractable Al is less
than 0.9 meq per 100 g soil (Lierop et at., 1982). Deep liming, as with deep
cultivation (Rowse and Stone, 1980; Ross, 1986), may be particularly
beneficial under conditions of drought stress especially if they remove
constraints to geotropic root growth towards a receding water table. In a
large-scale experiment in the Amazon jungle of Peru on a Typic Paleudult
(pH 4.5,65% Al saturation of the cation exchange capacity (CEC); Roca
and Midmore, 1983), liming was done, prior to planting the potato crop, to
a depth of 5, 15 or 30 cm. The rate of lime application was equivalent to the
base saturation by Al on a soil volume basis. Root growth, the a,chieve-
ment of complete crop cover, and tuber yield were favoured in the limed
over the non-limed plots (Fig. 15.8). Double the lime rate was supra-
optimal at all depths of incorporation, causing perhaps deficiency of
micro nutrients , such as iron, manganese, copper, zinc and boron and a
restricted uptake of phosphorus. Soil cultivation to a depth of 30 cm
without lime application also effected yield increase, which suggests an
amelioration of soil acidity through the mixing of surface and subsurface
soils (the Mg + Ca CEC was 2.70,0.97 and 0.07 meq per 100 g soil for the
0-10, 10-20 and 20-30 cm profiles of undisturbed soil). Loosening of the
deeper soil also resulted in less visible waterlogging, and probably reduced
ethylene production associated with anaerobic soil conditions (Campbell
and Moreau, 1979). As a continuation of the experiments in the Amazon
jungle, potato was planted in the following year (Roca and Midmore,
unpublished) into plots which had been limed 12, 8 or 4 months prior to
potato, at the same depths and rates as previously mentioned. Averaged
over rates and depths, there was no difference in potato yield between
o
o 2
Applied lime
Figure 15.8 Tuber yield as influenced by rate and depth of incorporation of lime
on an acid soil in lowland Peru. Applied lime as multiples of the quantity on a
volumetric basis necessary to neutralize the Al effect in the CEC, /':, incorporated
to 30 cm; 0 incorporated to 15 cm; 0 incorporated to 5 cm. (Source: Roca and
Midmore, 1983.)
plots limed immediately prior to the potato crop, or limed 4 or 8 months

earlier. As with the fresh application of lime, averaged over rates of liming,
deep incorporation of the intermediate lime rate resulted in maximum
yield.
The need to lime to reduce toxicities due to Al (and probably also due to
Mn) represents an added financial investment for the farmer in the tropical
lowlands, one that he is often ill-suited to make. Therefore the selection of
potato clones with tolerance to soil acidity appears a deserving objective,
but one that has received little attention. The laboratory screening out of
susceptible potato seedlings is feasible, i.e. those whose roots stain with
haematoxylin following 1 day in 0.13 mM Al nutrient solutions (Midmore
and Espinola de Fong, 1983). Pot tests compared the tuber yield reduction
due to high Al saturation of the CEC for tolerant and susceptible clones
based upon the seedling screening performance. Fifty percent of the
variation in response to high Al was due to the known inability of the
clones to tolerate aluminium as seedlings, which indicates the potential
advances that could be made in the search for increased tolerance to acid
soils.
15.4.3 Biotic stresses

Crop damage due to pests and disease can limit the success of potato
cultivation in lowland tropical areas. In particular the potato tuber moth
(Phthorimaea opercu[ella), the leaf miner fly (Lyriomyza huidobrensis),
and mites and thrips are pests of economic importance, and the integrated
approach to their control is discussed by Raman and Radcliffe (this
volume, Chapter 11).
The incidence and intensity of bacterial diseases (e.g. Pseudomonas
solanacearum, Erwinia carotovora ssp. carotovora) is of greater impor-
tance than fungal diseases, although early blight (Alternaria solani) ,
particularly in warm humid areas of Brazil (Reifschneider et al., 1984), is a
severe problem.
(a) Early blight

Although early blight can be controlled chemically with fungicides (e.g.
Dyrene (anilazine) and Dithane M-4S), and economic damage may be
offset by the use of high rates of N fertilizer (MacKenzie, 1981), due to
economic and environmental criteria, selection of resistant genotypes is a
high priority (International Potato Center, 1988). In general, susceptibility
to early blight is highly correlated with earliness, but the later maturing
apparently resistant clones are often unadapted to hot tropical conditions,
largely as a result of their low HI (Midmore, 1984a). Efforts to break the
linkage between earliness and susceptibility to early blight have been
rewarded, and medium (90 day) maturing clones with tolerable early blight
disease ratings have resulted from crosses between the relatively early
resistant clone Maine-47 and early non-tolerant material (International
Potato Center, 1988).
Choice of season and agronomic practices which minimize abundant
humidity may also be employed to reduce the severity of infection. The
intercropping of potato with sugarcane, although effective in Brazil
(Strohmenger and Hidalgo, 1986), perhaps by exclusion of inoculum, was
ineffective during the dry season in Bangladesh for the reduction of early
blight incidence (Imam et al., 1990). The importance of early blight will
continue to increase with the further extension of potato cultivation to the
warm humid areas of the hot tropics. To date the use of fungicides and
tolerant genotypes appear to be the only acceptable methods to keep the
disease under control.
(b) Bacterial diseases

Chemical control of bacterial diseases in infected soil is not feasible in
developing countries; hence control measures are generally directed
towards preventing the initial introduction, and, if present, reducing the
persistence and spread of bacterial diseases in the soil.
In the highland tropics, bacterial wilt of the potato (Pseudomonas
solanacearum) is principally caused by race 3 (biovar 2) and in the lowland
tropics by race 1 (biovars 1, 3, and 4). Race 3 is primarily tuber-borne and
specific to the potato, and as such can be readily eradicated through
rotation with pastures (Lloyd, 1976) or reduced to manageable levels
through a single rotation with non-host crops (Devaux and Bicamumpaka,
1984). Simple positive seed selection, which includes a period of high
temperature storage to promote symptomatology in latently infected
tubers, has been effective in reducing spread of inoculum via planting
material (Gonzales, 1977).
Intercropping potato with haricot beans, or with maize, practices
common with farmers in the Nile-Zaire divide, significantly reduced the
incidence and rate of disease development in a potato crop planted with
infected tubers (Autrique and Potts, 1987). Greater distance between
individual potato plants, and the presence of root systems of other spedes
between adjacent potato plants, reduced plant-to-plant root transmission
of the bacterium. Intercropping, together with the planting of moderately
resistant cultivars, can supplement the requirement for clean tuber seed,
until adequate quantities are available.
Race 1 has a more cosmopolitan distribution and greater variation than
found for race 3. For these reasons, integrated control practices involving
disease resistance, appropriate field sanitation (including rotation and
choice of disease free fields), and in particular the use of clean seed, are
essential to the success of potato production in the lowland tropics.
Traditionally resistance to bacterial wilt in commercial cultivars has been
derived from the diploid S. phureja, but this resistance apparently breaks
down under intermediate and high temperature conditions (Schmiediche,
1988). Germplasm with a Tuberosum origin is surprisingly more tolerant
under such conditions, which is believed to be related to its greater
physiological adaptation and more vigorous growth under such tempera-
ture conditions. Breeding at the diploid level with new sources of resis-
tance derived from S. sparsipilum, S. chacoense and S. microdontum has
been effective in widening the genetic base for resistance to bacterial wilt,
and selections are at present undergoing field evaluation (Schmiediche,
1988). An alternative approach to breeding for resistance, that is the
insertion of genes which code for potent bactericides, is also underway.
This methodology opens the opportunity for achieving simultaneous
resistance to both Pseudomonas and Erwinia species, a difficult task using
conventional breeding methods (Schmiediche et al., 1988).
Rotation as a method to eradicate race 1 is essentially ineffective,
given the wide range of cultivated and weed species that are alternate hosts
for race 1 e.g. other Solanaceae, Compositae. Rotation as a practice to
reduce wilt incidence is more effective, and in particular rotation with
graminaceous crops (e.g. upland rice, maize, Brachiaria decumbens (a
tropical pasture) (International Potato Center, 1985)). At temperatures
>40°C survival of bacterial wilt is poor (Seneviratne, 1988). In the upper
soil horizons during periods of dry weather in the tropics, maximum
temperature of bare soil exceeds this critical temperature: an effective
control measure which can be enhanced by covering the soil surface with
clear plastic sheets. Anaerobic conditions do not favour survival of P.
solanacearum, and fields previously planted with flooded rice crops are fre-
quently free of bacterial wilt. Insufficient duration of flooding (Seneviratne,
770 Potato production in th tropics
1988) and infstd sourc of irrigation watr may rduc th ffctivnss
of this control.
Planting at shallow dpth, and rstraining from hilling-up ar also usful
componnts of intgratd disas managmnt, and togthr with mulching
and intrcropping can rduc incidnc and sprad of actrial wilt (Kloos
t ai., 1986). Th rsidual ffct of intrcropping with non-host plants is
also notal: in potato filds iercropped with maize or beans, subsequent
potato crops had less disease symptoms than those following sole potato
crops (Autrique and Potts, 1987), and the incidence of latent infection was
also less (Kloos et ai., 1986) - an important factor if a reliable supply of
clean tuber seed is not availabLe and farmers are obliged to re-use their
own seed.
Of the three soft rot erwinias, E. carotovora ssp. atroseptica, ssp.
carotovora and E. chrysanthemi, the latter two are of greater importance in
warm tropical climates (Hidalgo, 1988). Although the disease is seed-
borne, and infected tubers are the primary source of field contamination in
temperate climates, other sources (e.g. contaminated irrigation water -
Elphinstone et ai., 1988) are prevalent in warmer climates, where alternate
host plants, and potato crop debris, are continuously present. For this
reason Erwinia is a disease with severe potential impact on potato
production in the lowland tropics. A reduction in the incidence of the
disease through the use of clean tuber seed is a useful strategy to control
Erwinia, because effective resistance is not presently available in commer-
cially grown varieties.
Considerations of the roles of calcium and water, as contrasting factors
which respectively increase resistance (calcium through its role in cell wall
formation has been implicated in the prevention of infection) or increase
susceptibility (water enhances opening of lenticels, and reduces oxygen-
dependent host resistance mechanisms - Perombelon, 1988), deserve
further attention when formulating agronomic practices to reduce the
incidence of Erwinia.
15.4.4 Planting materials

The potato crop in temperate climates is traditionally established from
seed tubers, and sophisticated seed production programmes exist in which
physiological and pathological quality of seed tuber is guaranteed, mainly
through a certification system. Tuber seed from these programmes is
readily available for use in areas of tropical potato production, but the
combination of high cost, lack of timely supply, and non-availability of
locally selected genotypes, often limits the appeal that the use of good
quality tuber seed would otherwise have in such areas. The potential for
use of various alternative planting materials has been studied.
(a) Tuber seed
Independent of disease factors which can reduce vigour of plants produced
from warm-produced seed tubers, tuber yield from seed produced in cool
climates was inherently greater than from seed of the same varieties
produced in warm tropical climates (Wiersema and Booth, 1985; Midmore
and Roca, 1991). Similar but less marked trends have also been reported in
temperate climates, when extremes of temperature are less notable (Wurr,
1978).
Lowland tropical potato production is largely confined to a single crop
per year; climatic constraints exist during the remainder of the year. Cold-
storage (CS) is a costly but effective method to conserve seed tubers during
hot tropical summers, and Reust (1982) has claimed that CS reconditioned
warm-produced tubers. This has not been subsequently substantiated. As
an alternative to CS, storage for 9 months in DLS in the warm tropics
(Midmore and Roca, 1991) did not reduce stand establishment upon
replanting stored tubers, although plant vigour, crop cover and tuber yield
were less than those with CS stored tubers. Vigour, cover, and yield were
markedly influenced by source of seed tuber (cool versus warm production
sites), in contrast to HI and the RCC which were independent of seed
tuber source. Markedly lower dry matter content of seed tubers produced
in warm climates, and stored in DLS, may in part be responsible for the
noted effects on vigour and cover (Midmore and Roca, 1991). Genotypic
variation in yield decline following replanting of warm-produced seed
tubers exists (Do Xu an Khuong, 1987; Midmore and Roca, 1991), which,
provided tubers are harvested under dry conditions and screened for latent
bacterial infections, can obviate the need for the import of cool-produced
seed tubers.
Given that at present cool-produced seed tubers are the main commer-
cial source of tuber seed for planting in warm areas, economies of use have
been made by cutting the seed tubers, often into one-eye pieces (Rashid,
1979; Sikka, pers. comm.) Yields in lowland areas of Vietnam and the
Philippines from plants derived from cut seed we.re as great as those of
plants coming from whole seed (International Potato Center, 1988).
Greater potential yield with cool-produced seed tubers may not necessarily
result in greater net returns for farmers, especially if the lower yield with
warm-produced seed tubers is offset by their lower costs. Economic
analyses of these two alternatives have yet to be made. Alternatives to
tubers as sources of planting material also deserve research attention, and
are discussed below.
(b) Rooted cuttings

Rooted cuttings, i.e. rooted portions of a stem that contain an active
terminal or axillary bud and a functional root system, in general consti-
tute the intermediate, or bridging, stage between techniques of rapid
multiplication (e.g. in vitro) and nursery or field production of seed or
consumption potatoes. Rooted cuttings are actually rooted plantlets, the
'cutting' term referring to the cutting of stems, of whatever form, into
smaller units, from which new plants will develop. Rooted cuttings may be
derived from in vitro produced plants (Escobar and Vander Zaag, 1988),
from sprouts (van Ho et al., 1988), from tuber-origin plants (Benz et al.,
1987) or from seedlings derived from true potato seed (Midmore and Benz,
unpublished), all sources which have been tested under warm tropical field
conditions (Table 15.9).
Table 15.9 Influence of type and origin of planting material on tuber yield and
some of its components at warm sites in Peru
(a) Single stem planting material of the progeny Atzimba x DTO-28
Plants harvested (%) Tubers per hill Yield (t ha-1)
Seedling tubers' 84 9.0 19.5
Stem cuttingst 75 7.4 17.7
Apical cuttingst 61 16.8 20.8
Seedlings§ 67 18.1 21.8
(b) Mean tuber yield~ (g per plant) for contrasting sources and origins of planting
material.
[Origin] (Warm/cool) x 100%
Source Cool Warm
Seed tubers 511 287 56
Tuberlets from cuttings 424 300 71
Seedling tubers 501 354 71
Seedlings 419
* From nursery production in a cool climate.
t From mother plants derived from cool-produced seedling tubeTS.
:j: Apical stem cuttings (6-8 nodes) from young seedlings.
§ Seedling transplants.
~ Seed tubers and tuberlets means of three clones; seedling tubers and seedlings means of
two progenies.
Sources: Benz et al. (1987); Benz (unpublished).
Exponential increases in availability of planting material are feasible,

since plants derived from rooted cuttings can act as mother plants for
further production of rooted cuttings, or as the basis for nursery produc-
tion of seed tubers or field production of ware tubers. This is particularly
important if, due to climatic constraints, only a short favourable period for
the production of planting material is available before the planting season.
As an example, starting with in vitro plants, apical cuttings are taken and
rooted. Fifteen days later the apical shoot of the rooted cutting can be
removed, and itself used to establish another mother plant, or taken to a
nursery bed or to the field for tuber production. Lateral branches develop
and grow from decapitated in vitro and mother plants, and a continuous
flush of apical shoots can be removed every 5 days (Escobar and Vander
Zaag, 1988). In the absence of a source of in vitro plants, tubers can be
stored between harvest and the planting season, in DLS, and either the
sprouts detached, cut into single or two-nodal segments and potted, or the
tuber planted in a pot directly. Plants derived from the nodal segments of
sprouts can then be transplanted directly to the field (van Ho et al., 1988)
or used as mother plants for further production of root cuttings. The apical
and axillary branches can be removed from tuber-derived mother plants,
and rooted and transplanted as required. The poor vigour associated with
DLS warm-produced seed tubers is not apparent, neither with the use of
rooted sprout cuttings, nor with axillary branches derived from such
tubers. Another effective source of mother plants, as an alternative to in
vitro or tuber-derived mother plants, are seedlings derived from true
potato seed (TPS). One hundred seedlings growing on one square metre of
nursery bed provided sufficient rooted cuttings to plant one hectare over a
6 month period, without recourse to the use of new mother plants
(Midmore, unpublished).
To prolong the effective period of production of rooted cuttings by
mother plants, management practices that favour vegetative growth and
inhibit tuberization, e.g. low irradiance, high nitrogen and high tempera-
tures, have been successful. The same practices suppress tuberization in
cuttings excised from mother plants (Benz, unpublished) - an important
feature since premature tuberization in rooted cuttings, transplanted to the
field, limits canopy growth and yield. Short days and relatively cooler
temperatures at the winter solstice strongly induce tuberization, and result
in lower yield potential compared to rooted cuttings transplanted earlier or·
later in the year (International Potato Center, 1988). Indeed, field prac-
tices that compensate for limited canopy development of rooted cuttings,
e.g. increased plant population, will result in greater tuber yield (Escobar
and Vander Zaag, 1988), particularly for rooted cuttings of clones char-
acterized by a long critical photoperiod, which tuberize early under warm
conditions.
Yield from rooted cuttings transplanted directly to warm field conditions
compared favourably with that from good quality cool-produced seed
tubers (Table 15.9). Yield potential with tubers produced in warm-climate
nurseries with rooted cuttings was 3~50% less than similar material
produced in a cool climate. These results, in combination with the low
multiplication rate and the high proportion of commercial-sized tubers,
suggest that, subject to availability of irrigation, rooted cuttings should
have greater potential for use in warm climates as field-transplants than for
rapid multiplication to produce seed tubers.
(c) True potato seed
True potato seed (TPS), sexually produced seed of the potato is, in terms
of multiplication rate between generations, a more efficient propagule of
the potato plant than the seed tuber. Economies of size, storage, transport
costs, and with limited exceptions, freedom from disease, all favour TPS
over seed tubers. In common with rooted cuttings and in vitro plants, there
are many potential uses for TPS. To date, however, readily available and
reliable supplies of good quality TPS are as elusive as are those of good
quality seed tubers in lowland tropical conditions.
Potato flowering and berry set are in general favoured in cool long day
climates, with some evidence to suggest that the length of day should be
diminishing (Pallais, pers. comm.). In the mid- and lowland-tropics varied
flowering response has been recorded from poor (Malagamba, 1988) to
acceptable (Sikka, 1987), but with correct choice of heat-tolerant parents,
and manipulation to prevent tuberization (e.g. high N, artificially extended
photoperiod, application of gibberellic acid (GA» flowering and berry set
are more acceptable.
Antagonism may exist between growth of tubers and development of
embryos (Pallais, 1987), and factors which favour the balance away from
tubers tend to improve TPS quality. Quality TPS combines a high level of
germination with the ability to produce vigorous uniform seedlings which
perform well under a range of environmental conditions (Dickson, 1980).
Grown under optimal conditions for TPS production, high N rates to
female parent plants (6 x 40 kg ha- 1 sequential applications) in addition to
that for recommended tuber crops, inhibit tuberization and delay sene-
scence. This permits full achievement of seed maturity and improves the
rate of emergence and early seedling growth of resultant TPS (Pallais and
Fong, 1989). Evidence suggests that TPS quality is less under warm
tropical conditions, those which do not permit complete seed maturity due
to a short ripening period (Pallais, 1987). In general terms, short days
result in less heavy seed than that produced in long days (Taylrarson, 1982).
Short days also hasten tuberization, and may be additionally responsible
for poorer quality of TPS produced in warm tropical climates.
Post-harvest practices influence dormancy and vigour of TPS, and the
worth of TPS seed lots. Although dormancy control of germination can be
released by the use of GA it does not release seedlings from a secondary
dormancy, expressed as slower growth rates (Pallais, pers. comm.). The
length of dormancy is genetically determined (Pallais, 1987), and is
naturally released in a shorter period at room temperature than at 5°C
(D'Antonio and McHale, 1988). TPS quality declines rapidly under
ambient tropical conditions (high relative humidity and temperature).
To avoid this TPS must be dried after extraction to 5-7% moisture (Inter-
national Potato Center, 1982), and maintained at that level. Since storage
of dried seed at 5°C preserves dormancy (Pallais, 1987) room temperature
storage at <Z5°C is recommended, unless long term gene-bank storage is
the objective. Prior to sowing, a 5-day seed priming period is advan-
tageous, especially if sowing is to take place under high temperatures
(30°C). The genotype is, however, often of overriding importance in
determining seedling vigour (Pallais et al., 1988).
Germination of TPS is favoured by daytime temperatures of 15-Z0°C,
and adequate soil moisture, and is rarely successful when sown directly to
the field (Martin, 1988). For these reasons sowing TPS in warm climates is
usually done under semi-controlled conditions, often in nurseries. Once
germinated, seedlings can be utilized in a number of ways: transplanted to
the field, to nursery beds for the production of seedling tubers, or used as
mother plants for the production of rooted cuttings in the same way as with
plants originating from in vitro or sprout cuttings.
Transplanting potato seedlings to the field, as with seedlings of other
crop species, results in an interruption in their growth rate, which in part
is related to the reduction in effective root area (Sattelmacher and
Minzenmay, 1984). Screening for root regeneration following a simulated
transplant shock using polyethylene glycol (International Potato Center,
1986) is an effective method to identify progenies tolerant to transplanting
shock. Agronomic practices, e.g. mulching and shading, can also improve
transplant survival (Malagamba, 1984).
A reliable supply of water during field establishment is indispensable for
the successful production of potato using transplanted seedlings. Tuber
yields in warm climates can then match those from good quality seed
tubers, or rooted cuttings (Table 15.9). One major disadvantage of TPS
transplants, when used for the production of consumer potatoes, is the
large number of small-sized tubers which predominate at harvest, a
character which to date has defied genetic improvement (Golmirzaie, pers.
comm.). This disadvantage for field production of consumption tubers is
turned to advantage when TPS is used to produce tuber seed (known as
seedling tubers, i.e. the first generation tubers from TPS seedlings).
For seedling tuber production TPS is generally established at 100 plants
m-2 in nursery beds. At this population farmers in the lowland tropics have
produced from 400 to 850 tubers m-2 . Competition between genotypes in
densely sown beds favours vigorous genotypes, which impart a yield
advantage in the ensuing tuber generation (Atlin and Wiersema, 1988).
Provided care is taken during early field crop management, yields of plants
grown from small « 10 g) tubers equal those from larger sized tubers
(International Potato Center, 1983). The yield potential of seedling tubers
produced in situ in warm climates is 3G-50% less than that of the same
progeny produced in a cool climate (Table 15.9), reductions similar to
those reported with the use of warm-produced tubers from rooted cuttings
and from seed tubers. Impartial economic analyses are required to
compare the worth of these sources of planting material.
Successful adoption of transplanting practices using TPS will depend to a
large extent on the quality of TPS, and the genotype, whereas for
production of seedling tubers and rooted cuttings the genotype will be of
greater importance. Direct seeding of TPS is unlikely to be feasible in the
near future, because early seedling growth is very sensitive to the vagaries
of the physical and biotic environment in the field. In temperate climates
fluid drilling may hasten the commercial use of direct-seeded TPS (Martin,
1988).
Open pollinated (OP) and hybrid progenies are the two alternative
sources of TPS and their worth has been routinely compared. Rates of
selfing in potato range from 55 to 90% (quoted by Atlin, 1985). Therefore,
continued use of OP generations does not invariably result in yield
reduction, yet hybrid progenies are, with few exceptions, superior in their
field performance, unless in OPs only vigorous (i.e. heterozygous) seed-
lings are selected for transplanting, and consequently for production of OP
seed for the next generation (Golmirzaie and Mendoza, 1986). Various
breeding strategies for the production of TPS have been proposed (Atlin,
1985; Jackson, 1987) and the reader is referred to these for a full appraisal
of the current situation.
15.5 TROPICAL LOWLAND POTATO PRODUCTION - TWO

PRACTICAL EXAMPLES
The extension of potato cultivation from the traditional to the non-

traditional or warmer areas is taking place in the tropics and subtropics.
Two examples are given of tropical production areas, where gradual or
abrupt transition has occurred over the last decade. In both examples
potato either replaces a crop, or fills an unused niche within an existing
cropping system.
In Bangladesh, a subtropical country without highland areas, the transi-
tion has been gradual. Potato is grown traditionally in rotation with rice as
a non-irrigated winter crop utilizing residual soil moisture.
Plantings start from October in the north, to late November or early
December further south in the Dhaka area. The planting seasons extend
for 3 months in the north but only for 1 month in the south. High
temperature limits the period suitable for planting in the south; earlier
plantings often fail due to detrimental effects of supra-optimal temperature
and moisture during plant emergence, and tuberization in later-than-
optimal plantings is delayed. If tuberization does occur, growth deformities
in tubers reduce the marketable value of the harvested product. Efforts by
the Potato Research Centre, in Joydebpur, to extend the period suitable
for planting, which concomitantly extends the supply and prevents market
gluts, have been rewarded by selecting and naming as varieties heat-
tolerant clones that can withstand higher than traditional temperatures at
the time of tuberization and that can be planted later than normal.
Tropical lowland potato production - two practical examples 777
A seed multiplication scheme has been in existence in Bangladesh for 18
years which permits the multiplication of imported seed once, but more
recently three times before severe virus buildup. Virus-free nuclear stocks
of the newly named varieties are to be maintained for seed production
purposes. Seed storage without cold-storage facilities is very difficult in
Bangladesh, and most improved varieties (as opposed to 'indigenous'
varieties) are cold-stored. Seed costs are high for farmers and they are
accustomed to using cut seed tubers and often plant single eye pieces at a
close spacing (Rashid, 1979). This type of planting favours rapid ground
cover, more efficient use of limited soil moisture, and a quicker cooling of
the soil. Since seed requirements per unit area are not so great in
Bangladesh as in other traditional potato growing areas, sufficient quan-
tities of improved cultivars with heat tolerance should soon be available
from the national potato programme for later plantings of potato.
To achieve a successful crop from later planting, farmers are planting
potato simultaneously with sugarcane. Large areas of traditional potato
production in the north are now intercropped with sugarcane, but the cane
does not compete with the potato since temperatures are too low for rapid
cane growth (Imam et al., 1990), until after potato harvest. When cane and
potato are planted in late January or early February, the cane shades the
potato prior to the potato harvest and reduces the detrimental effects of
high temperature on tuber bulking late in the potato season. Research in
Bangladesh (Imam et al., 1990) and Maharashtra State of India (Nankar,
1990) has demonstrated the mutual benefits that can accrue from the
associate planting of potato and sugarcane. In addition to the benefit of
shading for late potato plantings, the practice of weed control, efficient use.
of fertilizer and shared cost of cultural practices common to both crops
(e.g. harvest of potato and earthing of cane is done as one operation) also
favour this intercropping system.
The success of earlier planting over traditional planting in Bangladesh is
somewhat more limited. In the absence of excessive soil moisture, mulch-
ing after planting sucessfully reduces soil temperature to the optimal for
plant emergence; and, largely because of the conservation of soil moisture
- a factor of great importance during the dry winter - mulching with rice
straw or water hyacinth is a common practice during the traditional potato
season. Cultivars which can withstand both high temperature and high soil
moisture content during the emergence and establishment phases are as yet
non-existent, and their absence precludes the possibility of commercial
early plantings.
Climatic conditions similar to those found in Bangladesh exist elsewhere
where potato is also grown as a traditional winter crop such as in Egypt,
Cuba, Senegal, central southern India, El Salvador, and East Java. Indeed
most of the efforts to extend the planting seasons in Bangladesh have also
been duplicated in these countries, but with local variations, for example,
using maize as the shade crop in Egypt. Undoubtedly this concentration of
effort to extend the planting season will result in a step-by-step expansion
of potato production into hotter seasons.
The lowlands of the Philippines and Sri Lanka are examples of tropical
areas where an abrupt transition to non-traditional potato-producing areas
has occurred. The Lesser Antilles and the smaller Pacific Islands are
examples of production in new areas far isolated from traditional potato-
production areas.
Although appropriate genotypes and agronomic practices are suitable
for potato production in these non-traditional areas, the lack of informa-
tion, practical experience, marketing networks, and planting material,
have until recently precluded the possibility of growing potatoes as a
commercial crop there. With these limitations, a unified approach is called
for that encompasses research and extension, together with credit facilities,
local government involvement and farmer participation.
To illustrate one approach to the introduction of the potato as a new
crop in the lowland tropics, the development of the lowland potato project
in the Philippines will be discussed. Over a six-year period SAPPRAD
(Southeast Asian Program for Potato Research and Development) imple-
mented the project whose objective was to introduce potato as one
component of a diversified farming system, on 6000 to 10 000 small
lowland farms (Page and Horton, 1987). Virtually none of the farmers and
only a few technicians participating in the programme had grown potatoes
before. Therefore, it was necessary to produce a package of instructions
based on previous research experience, covering the complete manage-
ment of the crop, and which was later upgraded to provide the basis for
publishing a farmer production guide (PCARRD, 1985). Initially, in each
of five farming communities near market centres, a few farmer cooperators
were chosen on the basis of a farming systems survey that quantified land
suitability, availability of irrigation, and interest in growing a new crop.
Seed produced in the Philippine highlands was provided by SAPPRAD,
the cost of which was discounted at harvest from the farmers' returns.
Through a single training course, farmers were taught how to plant potato
and technicians how to monitor the crops during the growing season, as
well as to arrange field days. Three-quarters of the farmers succeeded in
growing the crop on plots ranging from 50 m2 to 1000 m 2 ; the small plot
size was used to reduce the financial loss should the crop fail and to avoid
saturating the market.
In the following year, the technicians conducted the training and 120
farmers cooperated, but of these, 25% failed due to floods, typhoons, and
particularly bacterial wilt (Page and Horton, 1987), illustrating the impor-
tance of suitable site selection. Other problems, such as thrips and mites
(Batugal et al., 1985), provision of credit and planting materials, and a
guaranteed market price have also been reported (Yabis et al., 1986). All
of these problems were discussed by technicians and farmers, and potential
solutions were tested under farmer's conditions. For example, to overcome
General comments and future 779
the limitations with seed supply, besides an institutional incentive for
highland potato producers to supply suitable cultivars to the lowland
farmers, alternatives to seed tubers such as rooted cuttings and TPS were
field-tested. The latter two systems could resolve the problem of how to
distribute small lots of highland seed to a large number of farmers;
however, for systems based on TPS or rooted cuttings, sources of TPS or
mother plants for cuttings still have to be resolved. Research by SAPPRAD
along these lines is in progress, as well as the selection of cultivars for
which seed can be produced in the lowlands, and cold-stored for a short
period prior to DLS storage. The third year of the lowland project had 250
new cooperating farmers, with most well surpassing the minimum break-
even yield level of 7 t ha l .
Even though financial and marketing arrangements are not central to the
success of the lowland potato project in the Philippines, they can accelerate
the spread of potato production once the essential technological conditions
are met (Page and Horton, 1987). If the close institutional linkages of the
type mentioned above are established from the start of lowland potato
production programmes, and the technological conditions are in place,
lowland potato production will be a success for farmers. Much valuable
research is taking place with the potato in non-traditional areas (e.g.
Barbados, Maynard et at., 1985; Vanuatu, Lebot, 1988) and this should be
strengthened by more aggressive support from governments and local
communities.
15.6 GENERAL COMMENTS AND THE FUTURE
The potato is truly a tropical crop even though the tropical environment for
production is one of stress. Low temperatures at high altitudes and high
temperatures at low altitudes clearly define seasonality in potato produc-
tion, which, combined with the bulky perishable nature of the harvested
product, lead to logistic difficulties for a uniform distribution and market-
ing throughout time and space. Other sources of stress, particularly biotic
and edaphic, also limit the economic viability of the potato crop during the
climatically favourable periods of the year at these altitudinal extremes.
These stresses can be of overriding importance at the intermediate
altitudes in the tropics, where temperature is favourable for year-round
potato production. Faced with these constraints, and little opportunity to
repel them, the course of least resistance to improve potato productivity in
tropical regions is through the improvement of stress tolerance.
Stresses, in general, reduce the ability of the crop to assimilate CO2 ,
through reduction of the assimilating surface, through reduction of the
efficiency of the biochemical processes involved in photosynthesis, or
through a combination of the two. Research on the effect of frost, heat,
drought and pathological stress on potato growth and development as
presented in this chapter provides some understanding and perhaps the
basis for improvement of stress tolerance in potato.
The time is now ripe for the potato, evolved in the tropical highlands and
later in northern latitudes and distributed worldwide, to return to the
tropics as a lowland crop. The fortuitous overriding effect of short days,
pronounced in cultivars with a long critical photoperiod, on the delay or
inhibition of tuberization by high temperature, has made the return a
reality, and the interplay of temperature, daylength, light intensity and
mineral nutrition on the physiology of the crop will be of prime importance
in the development of alternative sources of planting material for the
lowland tropics. Farmers should be encouraged to use alternative methods
of propagation, especially if they have n.o a priori reason to plant seed
tubers. These alternatives would alleviate potential problems with seed
health, seed availability (with few exceptions, heat-tolerant clones are not
presently under commercial scale multiplication), and seed distribution.
Clearly the potato production systems for lowland tropical areas will not
resemble those of northern temperature latitudes.
Expansion of potato production in time, away from the lowland tropical
winter season, will be faced with physiological and, of no insignificant
importance, logistic problems. Cloudiness and longer daylengths will
suppress the tendency to tuberize, heavy wet soils will hinder land
preparation, and the duration of preceding crops (such as rice) will have
to be reduced to accommodate the earlier potato planting.
Solutions do exist for these problems, and the expansion of potato
production to the non-traditional areas attests to this.
REFERENCES
Ahmad, K.U., Rashid, A. and Habib, A.K.M.A. (1982) Effect of methods of

fertilizer application on the performance of potato. Bangl. J. Agric. Res., 7,
80-3.
Agric. Sci., Camb., 94, 583-606.
Argotte, J.V. and Herrepa, J.A. (1987) Estudio sobre fertilizacion nitrogen ada en
el cultivo de la papa. Rev. Centro Agric. Cuba, 14, 87-93.
Atlin, G. (1985) Inbreeding in TPS progenies: implications for breeding and seed
production strategy, in Present and Future Strategies for Plant Breeding and
Improvement, International Potato Center, Lima, pp. 71-85.
Atlin, G.N. and Wiersema, S.G. (1988) Selection against inbred seedlings in
mixtures of inbred and hybrid true potato seed. Potato Res., 31, 105-12.
Austin, S., Baer, M.A. and Helgeson, J.P. (1985) Transfer of resistance to potato
leaf roll virus from Solanum brevidens into Solanum tuberosum by somatic
fusion. Plant Sci. Letters, 39, 75-82.
Autrique, A. and Potts, M.J. (1987) The influence of mixed cropping on the
control of potato bacterial wilt (Pseudomonas solanacearum). Ann. Appl. BioI.,
111, 125-33.
References 781
Bald, J.G. (1941) A report on agricultural features of the Australian potato
industry. Aust. Counc. Sci. Ind. Res. Pamph., 106.
Batugal, P.A., Acasio, R.F., Macaso-Khwaja, A., Balaki, E.T., Sano, E. and
Balaoing, V. (1985) Development of lowland potato technology for small
farmers. Philip. J. Crop. Sci., 10, 107-12.
Bean, J.N. and Allen, E.J. (1981) Effects of physiological age on development of
the leaf canopy and light interception, in Proc. 8th EAPR Trien. Conf, Munich,
pp. 142-4.
Bean, J.N., Sherwood, H. and Allen, E.J. (1984) Irrigation, light interception and
yield in maiN crop potatoes, in Proc. 9th EAPR Trien. Conf, Interlaken, pp. 90-1.
Beaumont, J.H. and Weaver, J.G. (1931) Effects of light and temperature on the
growth and tuberization of potato seedlings. J. Amer. Hart. Sci., 28, 285-90.
Beekman, A.G.B. and Bouma, W.F. (1986) A possible screening technique for
drought tolerance in potato, in Potato Research of Tomorrow (eds A.G.B.
Beekman, K.M. Louwes, L.M.W. Dellaert and A.E.F. Neele), Pudoc,
Benz, J.S., Midmore, D.J. and Keller, E.R. (1987) Comparison of different types
of planting materials for warm climate potato production, in Proc. 10th EAPR
Trien. Conf, Aalborg, pp. 359-60.
Bidinger, F.R. (1980) Water stress effects on crop-environment interactions, in
Proc. Int. Workshop Agroclimat. Res. Need Semi-Arid Tropics, Hyderabad, pp.
147-53.
Bodlaender, K.B.A. (1963) Influence of temperature, radiation and photoperiod
on development and yield, in The Growth of the Potato (eds J.D. Ivins and F.L.
Milthorpe), Butterworths, London, pp. 199-210.
Bodlaender, K.B.A. (1980) Effect of water stress on photosynthesis and transpira-
tion (abstract). Potato Res., 23, 251-2.
Booth, R.H. and Shaw, R.L. (1981) Principles of potato storage, International
Potato Center, Lima, 105 pp.
Borah, M.N. (1959) The effect of light intensity, length of day, and temperature on
growth and tuber formation in the potato, Ph.D. Thesis, University of Nottingham.
Borah, M.N. and Milthorpe, F.L. (1962) Growth of the potato as influenced by
temperature. Ind. J. Pl. Phys., 5, 53-72.
Burgers, M.S. and Nel, P.e. (1984) Potato irrigation scheduling and straw
mulching. South Afr. J. Plant Soil, 1, 111-6.
Burstall, L. and Harris, P.M. (1984) The intercropping of contrasting varieties of
potato, in Proc. 9th EAPR Trien. Conf, Interlaken, pp. 301-2.
Burt, R.L. (1965) The influence of reduced temperatures after emergence on the
subsequent growth and development of the potato. Eur. Potato J., 8, 104-14.
Burton, W.G. (1966) The Potato, H. Veenman and Zonen, Wageningen, 382pp.
Bushnell, J. (1925) The relation of temperature to growth and respiration in the
potato plant. Tech. Bull. Minn. Agric. Expt. Stn, No. 34, 29pp.
Campbell, R.B. and Moreau, R.A. (1979) Ethylene in a compacted field soil and its
effect on growth, tuber quality, and yield of potatoes. Am. Potato J., 56, 199-210.
Cary, J.W. (1985) Potato tubers and soil aeration. Agron. J., 77, 379-83.
Chapman, T. (1965) Experiments with Irish potatoes (Solanum tuberosum) in
Trinidad. Trap. Agric. (Trinidad), 42, 189-98.
Charles-Edwards, D.A. (1978) An analysis of the photosynthesis and productivity
of vegetable crops in the U.K. Ann. Bot., 42, 717-31.
Chaudhury, E.H., Kabir, H. and Sikka, L. (1984) Nitrogen fertilization as
influenced by method and time of application. 1. Ind. Potato Assoc., 11, 67-72.
Chen, H.H., Shen, Z.Y. and Li, P.H. (1982) Interrelationships in freezing and heat
tolerance in tuber-bearing Solanum species. HortSci., 17, 248-9.
Chen, H.H., Li, P.H. and Brenner, M.L. (1983) Involvement of abscisic acid in
potato cold acclimation. Plant Physiol., 71, 362-5.
Chen, P.M., Burke, M.I. and Li, P.H. (1976) The frost hardiness of several
Solanum species in relation to the freezing of water, melting point depression
and tissue water content. Bot. Gaz., 137,313-7.
Coleman, W.K. (1986) Water relations of the potato (Solanum tuberosum L.)
cultivars Raritan and Shepody. Am. Potato 1., 63, 263-76.
D'Antonio, V.L. and McHale, N.A. (1988) Effect of storage temperature and
extraction methods on dormancy and germination of true potato seed. Am.
Potato 1., 65, 573-81.
Davis, G.E. (1941) The effects of certain environmental factors on tuberization in
the wild potato Solanum commersonii. Am. Potato 1., 18, 266-9.
Devaux, A. and Bicamumpaka, M. (1984) Les methodes de controle limitant Ie
developpement de la bacteriose (Pseudomonas solanacearum) au Rwanda, in
Proc. 9th EAPR Trien. Conf., Interlake:n, pp. 27-8.
Devaux, A. and Haverkort, A.I. (1987) The effects of shifting planting dates and
mulching on late blight (Phytophthora infestans) and drought stress of potato
crops grown under tropical conditions. Expl. Agric., 28, 325-33.
Dickson, M.H. (1980) Genetic aspects of seed quality. HortSci., 15, 771-3.
Do Xuan Khuong (1987) Evaluation of potato (Solanum spp.) for storability
under high temperatures, MS Thesis, Beuguet State University, Philippines, 72
pp.
Dodds, K.S. and Hawkes, I.G. (1952) Potatoes in the tropics, in Rep. 13th.
Internat. Hort. Conf., Vol. II, pp. 1132-7.
Doorenbos, 1. and Kassam, A.H. (1979) Yield response to water. FAG Irrigation
and Drainage Paper, 33, 172pp.
Driver, C.M. (1941) Recent advances in potato breeding. NZ 1. Sci. Tech., 23,
180-4.
Dwelle, R.B., Kleinkopf, G.E. and Pavek, 1.1. (1981) Stomatal conductance and
gross photosynthesis of potato (Solanum tuberosum L.) as influenced by
irradiance, temperature, and growth stage. Potato Res., 24, 49-59.
Edmundson, W.e. (1941) Response of several varieties of potatoes to different
photoperiods. Am. Potato 1., 18, 100-12.
Ekanayake, 1.1. and Midmore, D.I. (1988) Drought response of potatoes in warm
tropical areas, in Proc. Int. Conf. Dryland Farming (in press).
Ekanayake, 1.1. and Midmore, D.I. (1989) Root pulling resistance of potatoes in a
drought environment (abstract). Am. Potato 1.,66,519.
EI-Hassan, H.S. (1984) Effect of side of planting on yield and quality of potatoes,
in Ann. Rep. Hudeiba Res. Stn Sudan, pp. 75-7.
Elphinstone, 1., Lindo, L. de and French, E.R. (1988) Control of Erwina diseases
in San Ramon, in Rept. plan. Conf. Bacterial Diseases of Potato, International
Potato Center, Lima, pp. 193-201.
Epstein, E. (1966) Effect of soil temperature at different growth stages on growth
and development of potato plants. Agron. 1., 65, 400-4.
Escobar, V. and Vander Zaag, P. (1988) Field performance of potato (Solanum
References 783
spp.) cuttings in the warm tropics: influence of planting system, hilling, density
and pruning. Am. Potato 1.,65, I-tO.
Espinola, N., Midmore, D.J. and Poats, S. (1984) Dry matter content, total protein
and consumer acceptability of potato (Solanum spp.) produced under hot
conditions, in Proc. 6th Symp. Int. Soc. Trap. Root Crops, Intemational Potato
Center, Lima, pp. 499-507.
Estrada, N. (1978) Breeding frost-resistant potatoes for the tropical highlands., in
Plant Cold Hardiness and Freezing Stress (eds P.H. Li and A. Sakai), Academic
Press, New York, pp. 333-4l.
Estrada, N. (1982) Breeding wild and primitive potato species to obtain frost
resistant cultivated varieties, in Plant Cold Hardiness and Freezing Stress:
Mechanisms and crop implications, Vol. II (eds P.H. Li and A Sakai),
Academic Press, New York, pp. 615-33.
Ewing, E.E. (1981) Heat stress and the tuberization stimulus. Am. Potato 1., 58,
31-49.
Ewing, E.E. (1987) The role of hormones in potato (Solanum tuberosum L.)
tuberization, in Plant Hormones and their Role in Plant Growth and Develop-
ment (ed. P.J. Davies), Martinus-Nijhoff, Dordrecht, pp. 515-38.
Ewing, E.E., Kahn, B.A., Lazin, M.B., Benkheder, M., Mendoza, H.A. and
Plaisted, R.L. (1983) Selecting for heat tolerance within populations derived
from the Cornell Andigena collection, in Research for the Potato in Year 2000
(ed. W.J. Hooker), International Potato Center, Lima, pp. 81-2.
Ewing, E.E., Lorenzen, J.H., Reynolds, M.P., Gebre-Mariam, S., Snyder, R.G.
and Ben Khedher, M. (1987) Evaluating heat tolerance in potato, in Improved
Vegetable Production in Asia (eds W.N. Chang, P.W. MacGregor and J. Bay-
Peterson), Food Fertilizer Technology Center for the Asian and Pacific Region,
Taipei, pp. 10--120.
Ezeta, F.N. and McCollum, R.E. (1972) Dry-matter production, and nutrient
uptake and removal by Solanum andigena in the Peruvian Andes. Am. Potato
1.,49, 151-63.
Gautney, T.L. and Haynes, F.L. (1983) Recurrent selection for heat tolerance in
diploid potatoes (Solanum tuberosum subsp. phureja and stenotomum). Am.
Potato 1., 60, 537-43.
Gaur, P.C and Gupta, P.K. (1981) Evaluation of potato germplasm for tuber dry
matter and protein content. 1. Ind. Potato Assoc., 8, 92-5.
Gebre-Mariam, S. (1988) Effect of high temperature on sprout growth in potato
(Solanum tuberosum L.) and genetical differences in degree of tolerance, Ph.D.
Thesis, Cornell Univ., 196 pp.
Golmirzaie, A.M. and Mendoza, H.A. (1986) Effect of early selection for seedling
vigor on open-pollinated true potato seed (abstract). Am. Potato 1., 63, 426.
Gonzales, L.c. (1977) Determinacion de niveles de infeccion por Pseudomonas
solanacearum en tuberculos de semilla de papa. Fitopatologia, 12, tOl-4.
Gooding, H.J. (1961) Irish Potatoes (Solanum tuberosum) in Trinidad. 1. Agric.
Soc. Trin. Tobago, 61, 193-211.
Gooding, H.J. (1964) Yield and storage experiments with the potato, Solanum
tuberosum, in Trinidad, in Proc. XVIInternat. Hort. Congr., Brussels, pp. 551-8.
Greaves, J.A. and Wilson, J.M. (1986) Assessment of the non-freezing cold
sensitivity of wild and cultivated potato genotypes by chlorophyll fluorescence
analysis. Potato Res., 29, 509-20.
Gregory, L.E. (1965) Physiology of tuberization in plants (tubers and tuberous
roots). Handbuch d. Pflanzen Physiologie, 15, 1328-54.
Hammes, P.S. and Nel, P.C. (1975) Control mechanisms in the tuberization
process. Potato Res., 18, 267-72.
Haverkort, A.J. (1985) Relationships between intercepted radiation and yield of
potato crops under the highland conditions of central Africa, Ph.D. Thesis,
University of Reading, 259 pp.
Haverkort, A.J. (1986) Yield levels of potato crops in central Africa. Agric. Syst.,
21,227-35.
Haverkort, A.J. and Harris, P.M. (1986) Conversion coefficients between inter-
cepted solar radiation and tuber yields of potato crops under tropical highland
conditions. Potato Res., 29, 529-33.
Haverkort, A.J. and Harris, P.M. (1987) A model for potato growth and
development under tropical highland conditions. Agric. For. Met., 39, 271-82.
Haverkort, A.J. and Rutayisire, C. (1986a) Utilisation des engrais chimiques sous
conditions tropicales. 1. Effet de l'application d'azote, phosphore et potasse sur
les rendements de la pomme de terre en Afrique centrale. Potato Res., 29,
347-55.
Haverkort, A.J. and Rutayisire, C. (1986b) Utilisation des engrais chimiques sous
conditions tropicales. 2. Effet de I'application d'azote, phosphore et potasse sur
la relation entre radiation intercepte et Ie rendement de la pomme de terre en
Afrique centrale. Potato Res., 29, 357-65.
Haynes, F.L., Haynes, K.G. and Moll, R.H. (1986) Maintenance and improve-
ment of tuber dry matter in a diploid breeding population (abstract). Am.
Potato J., 63, 430.
Haynes, K.G. and Haynes, F.L. (1983) Stability of high specific gravity genotypes
of potatoes under high temperatures. Am. Potato J., 60, 17-26.
Hermsen, J.G. Th. (1987) Efficient utilization of wild and primitive species in
potato breeding, in The Production of New Varieties: Technological Advances
(eds G.J. Jellis and D.E. Richardson), Cambridge University Press, Cambridge,
pp. 172-85.
Hetherington, S.E., Smillie, R.M., Malagamba, P. and Huaman, Z. (1983) Heat
tolerance and cold tolerance of cultivated potatoes measured by the chlorophyll-
fluorescence method. Planta, 159, 119-24.
Hidalgo, o. (1988) Soft rot and blackleg (Erwinia spp.) in warm climates, in Rept.
Plan. Conf Bacterial Diseases Potato, International Potato Center, Lima, pp.
179-86.
Horton, D.E. and Fano, H. (1985) Potato Atlas, International Potato Center,
Lima, 135 pp.
Imam, S.A., Delwar Hossain, A.H.M., Sikka, L.C. and Midmore, D.J. (1990)
Agronomic management of potato/sugarcane intercropping and its economic
implications. Field Crops Res., 25, 111-22.
International Potato Center (1979) Annual Report CIP 1979, Lima, Peru, 134 pp.
International Potato Center (1984a) Annual Report CIP 1983, Lima, Peru, 164 pp.
International Potato Center (1984b) Potatoes for the Developing World. A collabo-
rative experience, International Potato Center, Lima, 150 pp.
References 785
International Potato Center (1986) Annual Report CIP 1985 Lima, Peru, 175 pp.
International Potato Center (1987) Annual Report CIP 1986-1987, Lima, Peru, 210
pp.
Isoda, A. Nakaseko, K. and Gotoh, K. (1984) Some characteristics of two
andigena (s. tuberosum spp. andigena) strains in terms of dry matter production
and canopy structure. Japan J. Crop Sci., 53, 416-22.
Isoda, A., Nakaseko, K. and Gotoh, K. (1985) Effects of planting density on
growth of the andigena strains (S. tuberosum spp. andigena). Japan J. Crop Sci.,
54,311-17.
Iwanaga, M. (1985) Ploidy level manipulation approach: development of diploid
populations with specific resistance and FDR 2n pollen production, in Present
and Future Strategies for Potato Breeding and Improvement, International
Jackson, L.P. (1962) Effect of soil water and temperature on the growth of potato
sets. Am. Potato J., 39, 452-5.
Jackson, M.T. (1987) Breeding strategies for true potato seed, in The Production of
New Potato Varieties: Technological Advances (eds G.J. Jellis and D.E.
Jefferies, R.A. and MacKerron, D.K.L. (1987) Aspects of the physiological basis
of cultivar differences in yield of potato under droughted and irrigated
Khanna, M.L. (1966) Breeding potato varieties tolerant to higher thermoperiods.
Curro Sci., 35, 143-4.
Khedher, M.B. and Ewing, E.E. (1985) Growth analyses of eleven potato cultivars
grown in the greenhouse under long photoperiods with and without heat stress.
Am. Potato J., 62, 537-54.
Khurana, S.C. and McLaren, J.S. (1982) The infillence of leaf area, light
interception and season on potato growth and yield. Potato Res., 25, 329-42.
Kloos, J.P., Fernandez, B.B., Tumapon, AS., Villanueva, L. and Tulog, B.
(1986) Bacterial wilt research in Mindanao. CIP Circular (Lima), 14(3), 1-7.
Krauss, A. and Marschner, H. (1984) Growth rate and carbohydrate metabolism of
potato tubers exposed to high temperatures. Potato Res., 27, 297-303.
Ku, S., Edwards, G.E. and Tanner, C.B. (1977) Effects of light, carbon dioxide,
and temperature on photosynthesis, oxygen inhibition of photosynthesis and
transpiration in Solanum tuberosum. Plant Physiol., 59, 868-72.
Lazin, M.B., Ewing, E.E. and Mendoza, H.A. (1979) Progress in screening for
heat tolerance through the use of potato stem cuttings (abstract). Am. Potato J. ,
56,471.
Lebot, V. (1988) Les performances de la pomme de terre a Vanuatu. L'Agron.
Trap., 43, 37-46.
Lee, C.R. (1971) Influence of aluminium on plant growth and tuber yield of
potatoes. Agron. J., 63, 363-4.
Levitt, J. (1980) Responses of Plants to Environmental Stresses. Vol. 1. Chilling,
freezing, and high temperature stress, Academic Press, New York, 497 pp.
Levy, D. (1983) Varietal differences in the response of potatoes to repeated short
periods of water stress in hot climates. 1. Turgor maintenance and stomatal
behaviour. Potato Res., 26, 303-13.
Levy, D. (1984) Cultivated Solanum tuberosum L. as a source for the selection of
cultivars adapted to hot climates. Trap. Agric. (Trinidad), 61, 167-70.
Levy, D. (1985) The response of potatoes to a single transient heat or drought
stress imposed at different stages of tuber growth. Potato Res., 28, 415-24.
Levy, D. (1986) Genotypic variation in the response of potatoes (Solanum
tuberosum L.) to high ambient temperatures and water deficit. Field Crops Res.,
15,85-96.
Levy, D. Livesku, L. and Vander Zaag, D.E. (1986) Double cropping of potatoes
in a semi-arid environment: the association of ground cover with tuber yields.
Potato Res., 29, 437-49.
Li, P.H. and Palta, J.P. (1978) Frost hardening and freezing stress in tuber-bearing
Solanum species, in Plant Cold Hardiness and Freezing Stress (eds P.H. Li and
A. Sakai). Academic Press, New York, pp. 49-7l.
Li, P.H. and Fennell, A. (1985) Potato frost hardiness, in Potato Physiology (ed.
P.H. Li), Academic Press, New York, pp. 457-79.
Li, P.H., Palta, J.P. and Chen, H.H. (1979) Freezing stress in potato, in Low
Temperatures Stress in Crop Plants (eds J.M. Lyon, D. Graham and J.K.
Raison), Academic Press, New York, pp. 291-303.
Lipe, W.N. and Thomas, D.G. (1980) Effect of antitranspirants on yield of
Norgold Russet potatoes under greenhouse and field conditions. Am. Potato J.,
57,267-73.
Liu Jieming, and Midmore, D.J. (1990) A review of potato intercropping practices
in Western Hubei, China. Field Crops Res., 25, 41-50.
Lloyd, A.B. (1976) Bacterial wilt in a cool temperate climate of Australia, in Proc.
Plan. Cant Workshop Ecol. Contr. Bact. Wilt P. Solanacearum (eds L.
Sequeira and A. Kelman), NC State Univ., pp. 134-5.
Lomas, J. Schlesinger, E., Zilka, M. and Israeli, A.(1972) The relationship of
potato leaf temperatures to air temperatures as affected by overhead irrigation,
soil moisture and weather. J. Appl. Eco/., 9, 107-19.
Lorenzen, J.H. and Ewing, E.E. (1987) Genotypic differences in maintenance
respiration in potato (abstract). Am. Potato J., 64, 448.
MacKenzie, D.R. (1981) Associations of potato early blight, nitrogen fertilizer
rate, and potato yield. Plant Dis., 65, 575-7.
MacKerron, D.K.L. and Jefferies, R.A. (1984) Potato: light interception and
growth, in Scottish Crop Research Institute Third Annual Report 1983, SCRI,
Dundee, 148 pp.
MacKerron, D.K.L. and Waister, P.D. (1985) A simple model of potato growth
and yield. Part I. Model development and sensitivity analysis. Agric. For.
Meteorol., 34, 241-52.
Malagamba, P. (1984) Agronomic management for transplanting TPS seedlings, in
Innovative Methods for Propagating Potatoes Rpt. 27th Planning Cant, Inter-
national Potato Center, Lima, pp. 63-93.
Malagamba, P. (1988) Potato production from true seed in tropical climates.
HortSci., 23, 495-500.
Manrique, L.A., Tsuji, G.Y., Uehara, G. and Fox, R.L. (1984) Winter and
summer performance of potato (Solanum tuberosum) in isohyperthermic
regimes. Am. Potato J., 61, 41-56.
Marinus, J. and Bodlaender, K.B.A. (1975) Response of some potato varieties to
temperature. Potato Res., 18, 189-204.
References 787
Maris, B. (1969) Studies on maturity, yield, under water weight and some other
characters of potato progenies. Euphytica, 18, 297-319.
Martin, M.W. (1988) Cultural practices for using true seed in potato production
under temperate climates. HortSci., 23, 505-10.
Mauritius Sugar Industry Research Institute (1984) Annual Report 1963, MSIRl,
Reduit, 77 pp.
Maynard, A., Ojeda, V. and Clarke, B. (1985) Potato (Solanum tuberosum)
variety and cost production trial at Mount Wilton, Barbados, in Proc. 20th.
Ann. Meet. Carib. Food Crops Soc., St. Croix, 1984, pp. 208-11.
McCollum, R.E. and Valverde, e. (1967) The fertilization of potatoes in Peru. A
summary and interpretation of data from field experiments completed from 1959
through 1964 in the sierra. Tech. Bull. No. 185, NC Agric. Expt. Stn, Raleigh.
McGee, E., Booth, R.H., Jarvis, M.e. and Duncan, H.J. (1987) The inhibition of
potato sprout growth by light. I. Effects of light on dormancy and subsequent
sprout growth. Ann. Appl. BioI., 110, 399--404.
McGee, E., Booth, R.H., Jarvis, M.C. and Duncan, H.J. (1988) The inhibition of
potato sprout growth by light. II. Effects of temperature and light intensity.
Ann. Appl. Bioi., 113, 137--47.
Mendoza. H.A. (1984) Selection of parental materials for yield and earliness, in
Proc. 9th EAPR Tien. Conf, Interlaken, pp. 194--5.
Mendoza, H.A. and Estrada, R.N. (1979) Breeding potatoes for tolerance to
stress: heat and frost, in Stress Physiology in Crop Plants (eds H. Mussel and R.
Staples), Wiley, New York, pp. 227-62.
Menzel, C.M. (1980) Tuberization in potato at high temperatures: responses to
gibberellin and growth inhibitors. Ann. Bot., 46, 259-65.
Menzel, e.M. (1984) Potato as a potential crop for the lowland tropics. Trap.
Agric. (Trinidad), 61, 162-6.
Menzel, C.M. (1985) Tuberization in potatolllt high temperatures: interaction
between temperature and irradiance. Ann. Bot., 55,35-9.
Midmore, D.J. (1983) Factors affecting development and yield of potato under hot
climates, in Research for the Potato in the Year 2000 (ed. W.J. Hooker),
International Potato Center, Lima, pp. 130-1.
Midmore, D.J. (1984a) Papa para climas caJidos, in Proc. I Reunion Latino
Americana de Olericu/tura, pp. 51-9.
Midmore, D.J. (1984b) The potato (Solanum spp.) in the hot tropics I. Soil
temperature effects on emergence, plant de¥elopmen.t and yield. Field Crops
Res., 8, 255-71.
Midmore, D.J. (1987) Screening potato clones for adaptation to warm tropical
conditions. Tests of Agrochemical and Cultivars, 8, (Ann. Appl. Bioi, 109,
supplement), pp. 172-3.
Midmore, D.J. (1988a) Potato (Solanum spp.) in the hot tropics VI. Plant
population effect on soil temperature, plant development and tuber yield. Field
Crops Res., 19, 183-200.
Midmore, D.J. (1988b) Intercropping potato (Solanum spp.) with maize in warm
climates, in Proc. 7th. Symp. Int. Soc. Trap. Root Crops (ed. L. Degras),
INRA, Guadaloupe, pp. 837-51.
Midmore, D.J. and Espinola de Fong, N. (1983) Screening potato for aluminium
tolerance in nutrient solution, in Research for the Potato in the Year 2000 (ed.
W.J. Hooker), International Potato Center, Lima, pp. 132-3.
Midmore, D.J. and Rhoades, R.E. (1988) Applications of agrometeorology to the
production of potato in the warm tropics, in Agrometeorology of the Potato (ed.
C.J. Stigter), Acta Horticulturae, 215, 103-36.
Midmore, D.J. and Roca, J. (1991) Influence of production and storage conditions
on subsequent growth and tuber yield of potato (Solanum spp.) in the hot
tropics. J. Agric. Sci. (Camh.) (submitted).
Midmore, D.J., Berrios, D. and Roca, J. (1986a) The potato (Solanum spp.) in the
hot tropics II. Soil temperature and moisture modification by mulch in
contrasting. environments. Field Crops Res., 15, 97-108.
Midmore, D.J., Roca J. and Berrios, D. (1986b) Potato (Solanum spp.) in the hot
tropics III. Influence of mulch on weed growth, crop development, and yield in
contrasting environments. Field Crop Res., 15, 109-24.
Midmore, D.J., Roca, J. and Berrios, D. (1988) Potato (Solanum spp.) in the hot
tropics IV. Intercropping with maize and the influence of shade on potato
microenvironment and crop growth. Field Crops Res., 18, 141-57.
Proc. R. Soc., Series B., 281, 277-94.
Moreau, R.E. (1944) The climatic background to the problem of potato varieties
for East Africa. Part I. E. Afr. Agric. 1., 9, 127-44; Part II. E. Afr. Agric. J.
9, 203-12.
Morris, S.c. and Petermann, J.B. (1985) Genetic and environmental effects on
levels of glycoalkaloids in cultivars of potato (Solanum tuherosum L.). Food
Chemistry, 18, 271-82.
Nankar, J.T. (1990) Scope and prospects for intercropping of potato with sugar-
cane in Maharashtra State, India. Field Crops Res., 25, 123-32.
Nelson, D.G. (1987) Light interception, dry matter production and partitioning of
the potato crop in tropical environments, M.Sc. Thesis, University of Wales,
135 pp.
Nelson, D.G. and Midmore, D.J. (1986) Light interception and crop productivity
in contrasting environments (abstract). Potato Res., 29, 258.
Nowak, J. and Coleborne, D. (1989) In vitro tuberization and tuber proteins as
indicators of heat stress tolerance in potato. Am. Potato J., 66, 35-45.
O'Brien, P.J. and Allen, E.J. (1984) Some effects of water regime on leaf
emergence, morphology and growth, in Proc. 9th EAPR Conf. Trien. Conf.,
Interlaken, pp. 31-3.
Ochoa, C. (1984) Solanum hygrothermicum, new potato species cultivated in the
lowlands of Peru. Econ. Bot., 38, 128-33.
Page, O.T. and Horton, D.E. (1987) SAPPRAD 5-year review, International
Potato Center, Lima, 161 pp.
Pal, B.P. and Pushkarnath (1951) Potato-breeding investigations in India. Emp. J.
Exp. Agric., 19, 87-102.
Pallais, N. (1987) True potato seed quality. Theor. Appl. Genet., 73, 784-92.
Pallais, N. and Fong, N. (1989) A compromise strategy for storing high quality true
potato seed (abstract). Am. Potato J., 66, 538.
Pallais, N., Garcia, R. and Fong, N. (1988) Factors affecting seedling vigor in
potatoes: II. Genotype, after-ripening and seed priming (abstract). Am. Potato
J., 65, 495.
Palta, J.P. and Li, P.H. (1979) Frost hardiness in relation to leaf anatomy and
natural distribution of several Solanum species. Crop Sci., 19, 666-71.
References 789
Parker, B.L., Booth, R.H. and Bryan, J.E. (1983) Myzus persicae (Sulzer) in
diffuse light and dark rustic stores and resultant PLRV transmission. Am.
Potato J., 60, 65-74.
Parkinson, K.L. and Day, W. (1983) The influence of water stress on photo-
synthesis in a barley crop, in Effects of Stress on Photosynthesis (eds R.
Marcelle, H. Clijsters and M. Van Poucke), Nijhoff/Junk, The Hague, pp.
65-74.
PCARRD (1985) Lowland potato Technology, PCARRD/SAPPRAD, Los Banos,
20 pp.
Perombelon, M.C.M. (1988) Ecology of Erwinias causing stem and tuber diseases,
in Rept. Plan. Conf. Bacterial Diseases Potato, International Potato Center,
Lima, pp. 143-77.
Plaisted, R.L., Mendoza, H.A., Thurston, H.D. et al. (1987) Broadening the range
of adaptation of Andigena (Neo-Tuberosum) germplasm. CIP Circular, 15(2),
1-5.
Poats, S. V. (1983) Beyond the farmer: potato consumption in the tropics, in
Research for the Potato in the Year 2000 (ed. W.J. Hooker), International
Potts, M.J., Albert, W.V.D., Rutab, F.R., Sano, E.O., Mariano, P.P. and Booth,
R.H. (1983) An agro-economic assessment of seed-potato storage technologies
in the Philippines. Am. Potato J., 60, 199-211.
Rab, M.A. and Willatt, S.T. (1987) Water use by irrigated potatoes on a duplex
soil. Aust. J. Exp. Agric., 27, 165-72.
Raman, K.V. and Booth, R.H. (1984) Integrated control of potato moth in rustic
stress, in Proc. 6th Symp. Int. Soc. Trop. Root Crops, International Potato
Center, Lima, pp. 509-15.
Randeni, G. and Caesar, K. (1986) Effect of soil tempemture on the carbohydrate
status of the potato plant (S. tuberosum L.). J. Agron. Crop Sci., 156,217-24.
Rashid, A. (1979) Role of plant popUlation on the production of potato, in Proc.
2nd Workshop Potato Res. Workers, Potato Research Centre, BARI, Joydebpur,
Bangladesh, pp. 63-8.
Rashid, A., Ahmad, K.U. and Habid, A.K.M.A. (1981) Effect of mulch on the
performance of potato. Bangl. Hortic., 9, 24-7.
Rayner, R.W. (1945) Notes on the effect of day),ength on potato yields. E. Afr.
Agric. J., 10, 25-8.
Reifschneider, F.J., Furumoto, O. and Filgueira, F.A.R. (1984) Illustrated key for
the evaluation of early blight of potatoes. FAO Plant Prot. Bull., 32, 91-4.
Reust, W. (1982) Contribution a l'appreciation de I'age pbysiologique des tuber-
cules de pommes de terre (Solanum tuberosum L.) et etude de son importance
sur Ie rendement, These, Ecole Polytechnique Federale, Zurich, 113 pp.
Reynolds, M.P. and Ewing, E.E. (1986) Effects of air and soil temperatures on
potato tuberization (abstract). Am. Potato J., 63, 451-2.
Rhoades, R.E., Booth, R.H. and Potts, M.J. (1983) Farmer acceptance of improved
potato storage practices in developing countries. Outlook Agric., 12, 12-15.
Richardson, D.G. and Weiser, C.J. (1972) Foliage frost resistance in tuber-bearing
Solanums. HortSci., 7, 9-21.
Roberts, R.H. and Struckmeyer, B.E. (1938) The effects of temperature and other
environmental factors upon the photoperiodic responses of some of the higher
plants. J. Agric. Res., 56, 633-77.
Roca, J. and Midmore, D.J. (1983) Effect of quantity of lime and its depth of
incorporation on growth and yield of potato in an acid soil, in Research for the
Potato in the Year 2000 (ed. W.J. Hooker), International Potato Center, Lima,
pp. 164--5.
Ross, C.W. (1986) The effect of subsoiling and irrigation on potato production.
Soil Tillage Res., 7, 315-25.
Rowse, H.R. and Stone, D.A. (1980) Deep cultivation of a sandy clay loam II.
Effects on soil hydraulic properties and on root growth, water extraction and
water stress in 1977, especially of broad beans. Soil Tillage Res., 1, 173-85.
Sale, P.J.M. (1973a) Productivity of vegetable crops in a region of high solar input
I. Growth and development of the potato (Solanum tuberosum L.) Amt. J.
Agric. Res., 24, 733-49.
Sale, P.J.M. (1973b) Productivity of vegetable crops in a region of high solar input
II. Yields and efficiencies of water use and energy. Aust. J. Agric. Res., 24,
751-62.
Sale, P.J.M. (1974) Productivity of vegetable crops in a region of high solar input.
III. Carbon balance of potato crops. Aust. 1. Plant Physiol., 1, 283-96.
Sale, P.J.M. (1979) Growth of potatoes (Solanum tuberosum L.) to the small tuber
stage as related to soil temperature. Aust. 1. Agric. Res., 30, 667-75.
Sanchez, P.A. (1976) Properties and Management of Soils in the Tropics, Wiley,
New York, 618 pp.
Sato, K. (1981) Growth and development of potato plants influenced by combina-
tions of air- and soil-temperatures. Japan. J. Crop Sci., 50, 267-75.
Sattelmacher, B. (1983) A rapid seedling test for adaptation to high temperatures.
Potato Res., 26, 133-8.
Sattelmacher, B. and Minzenmay, B.P. (1984) Effect of root temperatures and
fertilization on early growth of potato grown from TPS, in Proc. 9th EAPR
Trien. Cant, Interlaken, pp. 311-12.
Schmiediche, P. (1988) Breeding for resistance to Pseudomonas solanacearum, in
Rept Plan. Cant Bacterial Diseases Potato, International Potato Center, Lima,
pp.19-27.
Schmiediche, P., Jaynes, J. and Dodds, J.H. (1988) Genetic engineering for
bacterial disease resistance in potatoes, in Rept Plan. Cant Bacterial Diseases
Potato, International Potato Center, Lima. pp. 123-32.
Scott, R.K. and Wilcockson, S.J. (1978) Application of physiological and agro-
nomic principles to the development of the potato industry, in The Potato Crop
(ed. P.M. Harris), Chapman and Hall, London, pp. 678-704.
Seneviratne, S.N. de S. (1988) Soil survival of Pseudomonas solanacearum, in Rept
Plan. Cant Bacterial Diseases Potato, International Potato Center, Lima, pp.
85-91.
Shalhevet, J., Shimsi, D. and Meir, T. (1983) Potato irrigation requirements in a
hot climate using sprinkler and drip methods. Agron.l., 75, 13-16.
Shimsi, D. and Sunoschi, M. (1985) Growth and yield studies of potato develop-
ment in a semi-arid region. 3. Effect of water stress and amounts of nitrogen top
dressing on tuber yield and quality of several varieties. Potato Res., 28,
177-91.
Sikka, L. (1987) Review of TPS research in Bangladesh, in Proc. Work. True
Potato Seed Res. Bangladesh, Potato Research Center, BARI, Joydebpur,
Bangladesh, pp. 8-17.
References 791
Simmonds, N.W. (1971) The potential of potatoes in the topics. Trap. Agric.
(Trinidad), 48, 291-9.
Simmonds, N.W. (1974) Dry matter content of potatoes in relation to country of
origin. Potato Res., 17, 178-86.
Sinden, S.L., Deahl, K.L. and Aulenbach, B.B. (1976) Effect of glycoalkaloids
and phenols on potato flavour. J. Food Sci., 41, 520-23.
Sinden, S.L., Sanford, L.L. and Webb, R.E. (1984) Genetic and environmental
control of potato glycoalkaloids. Am. Potato J., 61, 141-56.
Sipos J. and Prange, R.K. (1986) Response of ten potato cultivars to temperature
as measured by chlorophyll fluorescence in vivo. Am. Potato J., 63, 683-94.
Smillie, R.M. and Hetherington, S.E. (1983) Stress tolerance and stress-induced
injury in crop plants measured by chlorophyll fluorescence in vivo. Plant
Physiol., 72, 1043-50.
Smillie, R.M., Hetherington, S.E., Ochoa, C. and Malagamba, P. (1983) Tole-
rances of wild potato species from different altitudes to cold and heat. Planta,
159,112-18.
Stark, J.C. and Wright, J.L. (1985) Relationship between foliage temperature and
water stress in potatoes. Am. Potato J., 62, 57-68.
Stelzner, G. and Torka, M. (1940) Tageslange, temperatur und andere umwelts-
factoren in ihren eihfluss auf die khollenbildung der Kartoffel. Zuchter, 12,
233-7.
Strohmenger, A. and Hidalgo, O. (1986) Batata consorciado com can de acucar na
epoca da chuva no Distrito Federal. Hart. Brasil., 4, 75.
Stylianou, Y. and Orphanos, P.I. (1981) Irrigation of potatoes by sprinklers and
trick Irs on the basis of pan evaporation in a semi-arid region. Potato Res., 24,
159-70.
Sukumaran, M.P. and Weiser, C.J. (1972) Freezing injury in potato leaves. Plant
Physiol., 50, 564-7.
Sunoschi, M. and Shimsi, D. (1985) Growth and yield studies of potato develop-
ment in a semi-arid region. 2. Effect of water stress and amounts of nitrogen top
dressing on growth of several cultivars. Potato Res., 28, 161-76.
Sunoschi, M., Costelloe, B., Lifshitz, Y., Lee, H.C. and Roseman, Y. (1987)
Arma: a potato cultivar resistant to heat stress. Am. Potato J., 64, 191-6.
Swaminathan, M.S. and Sawyer, R.L. (1983) The potential of the potato as a world
food, in Research for the Potato in the Year 2000 (ed. W.J. Hooker), Inter-
national Potato Center, Lima, pp. 3-4.
Taja, H., Cardona, A., Suetos, D., Acasio, R. and Vander Zaag, P. (1984) Potato
(Solanum tuberosum L.) tuber yield in Cagayan as influenced by planting date,
mulching and location. Phil. Agr., 67, 55-69.
Tamargo, M.A. (1948) Some observations on potato production in Cuba. Am.
Potato J., 25, 34-6.
Taylorson, R.B. (1982) Interaction of phytochrome and other factors in seed
germination, in The Physiology and Biochemistry of Seed Development, Dor-
mancy and Germination (ed. A.A. Khan), Elsevier, New York, pp. 323-46.
Thavarajavel, S. (1969) A study of potato cultivation in the Jaffna peninsula (Maha
1967-68). Trap. Agric. (Ceylon), 125, 101-9.
Timm, H., Timm, G.H. and Bishop, J.C. (1980) Ethylene production and growth
of potato sprouts under simulated high temperature soil conditions (abstract).
HortSci., 15, 383.
Trebejo, I. (1987) Study on the influence of water and light use in the potato
(Solanum tuberosum L.) crop (in Spanish), Thesis (Ing. Agr.) Universidad
Nacional Agraria, Lima, 108 pp.
Trebejo, I. and Midmore, D.J. (1990) The effect of water stress on potato growth,
yield, and water use in a hot and cool tropical environment. 1. Agrie. Sci.
(Camb.), 114,321-34.
Troll, C. and Paffen, K.H. (1965) Seasonal Climates of the Earth. World maps of
climatology, Springer Verlag, Berlin.
Tumuhairwe, J.K. and Gumbs, F.A. (1983) Effect of bed type and mulching on the
wet season production of cabbage (Brassiea oleraeea). Trap. Agrie. (Trinidad),
60, 11-16.
Valdes, c., Fraser, T. and Rosseaux, B. (1982) Estudio de distintas fuentes
nitrogenadas y su fraccionamiento sabre los rendimientos y la cali dad de la papa
(Solanum tuberosum). Ciene. y Teen. Agrie. (Cuba), 1-2,83-94.
Vander Zaag, D.E. (1984) Reliability and significance of a simple method of
estimating the potential yield of the potato. Potato Res., 27, 51-73.
Vander Zaag, D.E. and Horton, D. (1983) Potato production and utilization in
world perspective with special reference to the tropics and sub-tropics. Potato
Res., 26, 323--62.
Vander Zaag, P. and Demagante, A. (1985) Water requirements as influenced by
irrigation system and mulch for potato (Solanum spp.) grown in an isohyper-
thermic environment in the Philippines. Phil. Agr., 68, 571-83.
Vander Zaag, P. and Demagante, A.L. (1987) Potato (Solanum spp.) in an
isohyperthermic environment. I. Agronomic management. Field Crops Res.,
17,199-217.
Vander Zaag, P. and Demagante, A.L. (1988) The potato (Solanum spp.) in an iso-
hyperthermic environment III. Evaluation of clones. Field Crops Res., 19, 167--8l.
Vander Zaag, P., Demagante, A., Acasio, R., Domingo, A. and Hagerman, H.
(1986) Response of Solanum potatoes to mulching during different seasons in an
isohyperthermic environment in the Philippines. Trap. Agrie. (Trinidad), 63,
229-39.
van Ho, T., Hoa, N.T., Loan, T.T. et al. (1988) Techniques for using sprouts for
potato production in the tropics. Potato Res., 31, 379-83.
van Lierop, W., Tran, T.S., Banville, G. and Morissette, S. (1982) Effect ofliming
on potato yields as related to soil pH, AI, Mn and Ca. Agron. 1., 74, 1050-5.
van Swaaij, A.C., Jacobsen, E. and Feenstra, W.J. (1985) Effect of cold
hardening, wilting and exogenously applied proline on leaf proline content and
frost tolerance of several genotypes of Solanum. Phys. Plant., 64, 230--6.
van Swaaij, A.c., Nijdam, H., Jacobsen, E. and Feenstra, W.J. (1987) Increased
frost tolerance and amino acid content in leaves, tubers and leaf callus of
regenerated hydroxyproline resistant potato clones. Euphytiea, 36, 369-80.
Versteeg, M.N. (1985) Factors affecting the productivity of irrigated crops in
Southern Peru, in relation to simulation models, Doctoral Thesis, Wageningen,
182 pp.
Villareal, C.P. (1976) The influence of temperature on some carbohydrates in the
white potato tuber (Solanum tuberoSllm L.), M.Sc. Thesis, University of
Philippines, Los Banos, 84 pp.
Vos, J. (1986) The impact of transient water stress on potatoes (abstract). Potato
Res., 29, 264.
References 793
Vos, l. and Oyarzun, P.l. (1987) Photosynthesis and stomatal conductance of
potato leaves - effects of leaf age irradiance and leaf water potential. Photo-
synthesis Res., 11, 253--64.
Went, F.W. (1957) The experimental control of plant growth. Chronica Bot., 17,
109-12.
Werner, H.O. (1934) The effect of a controlled nitrogen supply with different
temperatures and photo periods upon the development of the potato plant.
Univ. Nebraska Agr. Expt. Stn Res. Bull., 75.
Wiersema, S.G. and Booth, R.H. (1985) Influence of growing and storage
conditions on the subsequent performance of seed potatoes under short-day
Winkler, E. (1971) Kartoffelbau in Tirol. II. Photosynthese vermogen und
respiration von verschiedenen kartoffelsorten. Potato Res., 14, 1-18.
Wivutvongvana, M. (1979) Physiological response of heat tolerant and heat
sensitive potatoes (Solanum species), Ph.D. Thesis, Cornell University, 121 pp.
Wu, M.T. and Wallner, S.l. (1984) Heat stress response in cultured cells. Heat
tolerance induced by heat shock versus elevated growing temperature. Plant
Physiol., 75, 778--80.
Wurr, D.C.E. (1978) 'Seed' tuber production and management, in The Potato
Crop (ed. P.M. Harris), Chapman and Hall, London, pp. 327-54.
Yabis, E., Taja, H., Tianza, G. and Vander Zaag, P. (1986) On-farm evaluation of
potato (Solanum spp.) in Cagayan during the 1983-1985 dry seasons. Phil. Agr.,
69,239-50.
Yamaguchi, M., Timm, H. and Spurr, A.R. (1964) Effects of soil temperature Olil
growth and nutrition of potato plants and tuberization, composition and
,periderm structure of tubers. Proc. Am. Soc. Hart. Sci., 84, 412-23.
CHAPTER 16
Potato production in the context of

the world and farm economy
D.E. Horton and J.L. Anderson
This chapter consists of three sections. Section 1 presents an overview of

global trends in potato production, use and nutritive value of the potato.
Section 2 briefly characterizes the systems of potato production, marketing
and use in developing countries - mainly tropical and subtropical areas.
Section 3 treats these same topics in developed economies, with special
reference to the situation in Great Britain.
16.1 GLOBAL PATTERNS AND TRENDS*
The potato is one of the world's most important, and widely grown crops.
Originating in the Andes of South America, potato cultivation has spread
throughout the world. The potato is now grown in more countries than any
other food crop save maize. As a cheap food source for the labouring
classes, the potato contributed substantially to Europe's industrial revolu-
tion. In many parts of the world, the potato has also provided famine relief
during periods of war and crop failure. In Ireland, on the other hand,
failure of the potato crop in the 1840s resulted in a devastating famine.
Due to its historical importance in Europe, the potato is often thought of
as a crop whose production and us.e are largely confined to industrial
nations. This is not true. Developing countries today produce about 25%
of the world's potatoes, and the share of these countries in global
production is increasing rapidly.
16.1.1 Origin and spread of the potato crop

Sailors returning from the Americas brought potatoes to Spain in the
sixteenth century. From Spain, the potato spread throughout Europe, first
• This Section is based on Horton (1987), Chapter 1. Readers are referred to this book for
detailed documentation and references.

Global patterns and trends 795
as a botanical curiosity, much later as an economic crop. In Europe

potatoes were fed to livestock long before they became a staple in human
diets. People rejected potatoes as being unclean, unhealthy, or even
poisonous. Only when potatoes became a low-cost source of food energy
(due primarily to the selection of adapted varieties) and when other foods
became scarce (primarily as a result of crop failures, wars, or famines), did
Europeans begin to replace their traditional staples such as buckwheat and
oats with potatoes. The popularity of the potato in Europe reached its
zenith around 1850 when it was the least expensive source of food energy,
the most important food crop after wheat, a major livestock feed, and the
principal source of starch and alcohol. Since then, per capita potato
production and use has declined in Europe, as other foods, feeds, and
sources of starch and alcohol have become cheaper, and as consumers have
diversified their diets, reducing dependence on low-cost staples.
Potatoes reached most other parts of the world through the European
colonial powers, rather than directly from South America. North America
first received potatoes from England via Bermuda in the 1620s. British
missionaries took potatoes to many parts of Asia in the seventeenth
century, and Belgian missionaries introduced them to the Congo in the
nineteenth century. Potatoes reached China around 1700 from the Dutch
East Indies (now Indonesia). Introductions were also made to northern
China from Russia. Portuguese traders took the crop to India in the 1600s.
Potatoes were introduced into Tibet from both China and India. They were
first carried to Japan around 1600 by Dutch traders from Java. Potatoes
were introduced into Persia in the late eighteenth century. About 50 years
later, they reached Syria, where they were known as 'colocasia of the
foreigners' .
Although the evidence is sketchy, it seems that, as in Europe, potato
production and consumption in the developing world was strongly in-
fluenced by the suitability of the environment for the crop, development of
production and post-harvest systems that were appropriate for specific
environments, and food habits and needs. Potato cultivation expanded first
in areas with cool, rainy climates. Often local food habits and myths
discouraged potato consumption long after its introduction, and until times
of food shortage, most potatoes were eaten primarily by European
colonists rather than by the indigenous population. In warm environments
for potato production - such as the Indo-Gangetic plain - potatoes were
restricted to garden plots until quite recently when a series of new supply
and demand factors - selection of adapted varieties, development of a seed
system, expansion of irrigation and cold storage, population growth,
urbanization, rising incomes, and changing food habits - induced the
spread of potato production and consumption.
796 Potatoes and the world and farm economy
16.1.2 Recent trends in potato production and use
Estimates of potato production and use are not very reliable, especially in
developing countries. However, the broad trends are known. This section,
based on recent, unpublished estimates of the Food and Agricultural
Organization (FAO), presents the most accurate figures currently available.
(a) Production
World potato production was about 135 million tonnes at the turn of the
century, 250 million tonnes around 1950, and in the late 1980s was about
200 million tonnes. During the first half of the century, Europe (including
the USSR) produced about 90% of the world's potatoes. Potato produc-
tion began to fall in Europe after World War II. Since 1960, production in
western Europe has dropped by more than a third. Potato production also
shrank by about 10% in eastern Europe and the USSR, but it has grown
elsewhere.
Since 1960, the total world potato area has fallen by nearly 20% (Table
16.1). This masks a 70% increase in developing countries taken as a group
and a 35% decrease in developed countries. Yields have also shown
marked changes over the period. The overall improvement of 50% hides a
doubling of yields in developing countries compared to an increase of less
than 40% in developed countries. The combination of trends in area and
yield results in a modest 8% increase in production at world level, which
breaks down into a 140% increase in developing countries and an 8%
decrease in developed countries.
Of total world production, eastern Europe and the USSR produce just
under half, and western Europe produces about 17% (Fig. 16.1) Nearly
15% of the world's potatoes are grown in the Far East, 10% in the
Americas and Oceania, and less than 5% in Africa and the Middle East.
Germay, once the world's leading potato producer, has registere a
Table 16.1 Trends in potato area, yield, and production (3-year averages)
World Developed nations Developing nations
Area Yield Prdn Area Yield Prdn Area Yield Prdn

(000 ha) (t ha- 1) (tx 106 ) (000 ha) (t ha- 1) (tx 106 ) (000 ha) (t ha- 1) (tx10 6 )
1961163 22143 12 265 18593 l3 236 3550 8 29

1965/67 21470 l3 283 17276 14 247 4193 9 36
1971173 20314 14 287 15669 15 240 4645 10 47
1975177 19439 14 275 14523 15 222 4916 11 53
1981183 18808 14 267 l3156 15 204 5652 11 63
1985/87 18330 16 286 12310 18 216 6020 12 70
Source: FAO Basic Data Unit. Revised, unpublished est,imates, May 1988.
(It = 1.016 tonnes)
350
300
250
00
c
.9
0
0 200
S.
c
0
n
:::l
ea.
"0
150
0
itl
'0
a..
100
50
o
1965 1970 1975 1980 1985
Year
Figure 16.1 Potato production in major regions 1961-87 (3-year moving average).
Source: FAO Basic Data Unit. Revised, unpublished estimates.
precipitous decline in production (Table 16.2). Since 1960, potato produc-

tion in the former West Germany has fallen by two-thirds. Potato
production has also dropped by half in France. In contrast, potato
production has remained nearly constant in the UK - a managed market
system - it has increased by 40% in the USA and has nearly doubled in The
Netherlands, building on comparative advantage. In many developing
areas, increases have been even greater. Potato production has quadrupled
since 1960 in India and Pakistan and more than tripled in Rwanda,
Bangladesh, Madagascar, Cuba, Mexico, and several north African and
Middle Eastern countries. As a result of these divergent trends, more
potatoes are now grown in Asia than in western Europe. China now
produces more potatoes than the USA, and India produces more than the
former West Germany.
Table 16.2 Potato production in selected countries
Production (000 t)
% change
1961/63 1985/87 production
Bangladesh 347 1110 220
China (exc. Taiwan) 12900 26750 107
Cuba 96 311 226
India 2844 11908 319
Madagascar 83 264 218
Mexico 366 950 160
Pakistan 119 585 391
Peru 1445 1596 10
Rwanda 79 253 221
France 14565 7003 -52
Germany, East 11533 11216 -3
Germany, West 24144 7996 -67
Netherlands 3842 7161 86
United Kingdom 7064 6709 -5
United States 12543 17454 39
Source: FAD Basic Data Unit. Revised, unpublished
estimates. May 1988.
In South America, where potaoes have long been a staple food, potato
production has grown less rapidly, in part because consumers are diversify-
ing their diets and moving away from traditional staples. Agricultural and
trade policies have also discouraged potato production in some countries,
like Peru.
Potato yields have risen throughout the world, most notably in develop-
ing regions. In Europe, increases in potato yields have not kept pace with
increases in cereal yields since World War II, reflecting greater improve-
ment in cereal varieties and production systems. As a result, potatoes have
become more costly than other energy sources, and use of potatoes for
livestock feeding and industrial purposes has fallen. Demand for fresh
potatoes has also dropped as diets have changed. For these reasons,
European farmers have reduced the area and production of potatoes.
In most developing regions potato yields have increased at least as
much as cereal yields and much more than yields of other root crops.
Improvements in varieties, seed systems and post-harvest technology have
helped bring down the cost of producing potatoes, improving their
competitive position on farms in many developing areas. Population
growth, rising incomes and changing food habits have also stimulated
potato production.
In conclusion, the balance of world potato production is gradually
shifting from the developed to the developing countries and from the
temperate to the tropical and subtropical zones. Only about 40% of the
world's potatoes are now grown in Europe; 35% are grown in other
developed countries and 25% in developing countries.
(b) Utilization
Potatoes are consumed by humans, fed to animals and serve as a raw
material for starch and alcohol. A rather large part of the havest is also
used for seed. In western Europe, as a result of declining feed and
industrial use of potatoes, per capita potato production has fallen by more
than 40% since 1960, but per capita potato consumption has dropped by
less than 15%. Declines in countries like France and Germany have been
partially offset by increases in others, like the UK and The Netherlands
(Figure 17.2). The share of the crop that is wasted and used as seed has also
fallen somewhat.
120 - , - - - - - - - - - - - - - - - - - - - - - - - - - - ,
United Kingdom ... ------
......... -- ---... - -----. _... _... -_.....

100
"0
ra
Q)
.r: W. Germany
CD N.etherlands
a.
Ci
:::.
c
aE
0
80
:J
France
'"c0
U
.... -111
~
(5
a.
60
o 4----.----.----.-----.---,r----.----.----,--~
1980 1985
Year
Figure 16.2 Trends in potato consumption in selected countries 1976-86. Source:

Table 16.2.
Use of potatoes in livestock feeding and industry has declined as
potatoes have become more expensive relative to alternative feedstuffs and
energy sources. Prior to World War II, nearly half the potatoes grown in
western Europe were fed to livestock, but this use is now less than 20%.
In most developing countries, few potatoes are processed or fed to
livestock, and consumption is rising. Since 1960, in most areas per capita
potato consumption has grown somewhat more rapidly than per capita
production, as better technology and rising yields have reduced the shares
of the crop that spoil and that must be reserved for seed.
In contrast to the immense international trade in the major cereal grains,
only a small proportion of the world's potatoes (about 3%) is traded on
international markets (Table 16.3). Potato exports are insignificant in all
but a few countries because their bulkiness and perishability make them
costly and risky to haul over long distances. Also, quarantine regulations
restrict international trade in potatoes in some instances.
Table 16.3 Potato trade by region, 1984 (000 t)

Potato
Country Imports Exports
World 7757 7695

Developing nations 1241 786
Developed nations 6516 6909
Africa 418 200
North and Central America 661 519
South America 55 5
Asia 754 604
Europe 5278 6161
Oceania 28 21
USSR 563 184
Source: Horton (1988) Table 30.
Over 80% of world potato trade occurs between developed countries,

with European trade responsible for over three-quarters of developed
country movement. The bulk of trade is in raw form although the
proportion of processed product French fries (chips in the UK) is on the
increase. Within the total movement of around 7.7 million tonnes, roughly
1 million tonnes is in the form of seed. The major seed trader is Holland,
which accounts for about half. Other important traders of seed are the UK
(Scotland and Northern Ireland), Canada, France and the former West
Germany. Countries bordering the Mediterranean arc the main recipients
of Dutch, French, and UK-produced seed. Canadian stocks move tradi-
tionally to the USA and South America.
Potato consumption levels vary greatly among countries and regions
U.S.S.R.
North & Central America
o 20 40 60 80 100 120 140

Potato consumption (kg)
per capita
Figure 16.3 Per capita potato consumption in major regions 1984 (kg).
Source: FAG Basic Data Unit. Unpublished.
(Fig. 16.3). Although potato consumption in Europe has generally been

falling (Holland and the UK excepted), it still exceeds 100 kilograms per
head per year in the UK, Poland, the USSR and the former East Germany.
And although consumption is expanding in developing countries, most
people still eat less than 20 kilograms of potatoes a year. Within the
developing world, average potato consumption levels are lowest in the hot
tropics and highest in countries that have significant temperate or highland
production zones. Potato consumption is rising particularly rapidly in those
areas where the demand for food is growing fast and technological change
is lowering the costs of potato production and marketing.
16.1.3 Nutritive value

Potatoes are a valuable source of nutrients (Table 16.4). Suffice to say that
potatoes come out quite well in terms of a balanced food if somewhat light on
Table 16.4 Composition of raw potato and other plant foods contrasted with composition when cooked (per 100 g edible portions)
Raw Cooked*
Food Total Total
Crude carbo- Dietary Crude carbo- Dietary
Energy Moisture protein Fat hydrate fibre Energy Moisture protein Fat hydrate fibre
(kcal) (%) (g) (g) (g) (g) (kcal) (%) (g) (g) (g) (g)
Potatoes 80 78.0 2.1 0.1 18.5 2.1 76 79.8 2.1 0.1 18.5 1.07t
Cassava 145 62.6 1.1 0.3 35.2 5.2 124 68.5 0.9 0.1 29.9
Sweet corn 96 72.7 3.5 1.0 22.1 3.7
Maize porridge 76 81.2 1.8 0.8 15.6
Wheat (hard) 332 12.3 13.3 2.0 70.9 12.1
Bread (unenriched
white) 278 32.7 8.7 1.6 55.7 2.7
Rice (milled
white) 364 12.0 6.8 0.5 80.2 2.4 135 67.9 2.3 0.3 28.0 0.8
Sorghum 342 10.9 10.1 3.4 73.2 9.0 85 79.6 2.7 0.5 17.0
Beans (Phaseolus
vulgaris) 338 11.8 21.9 1.6 61.2 25.4 118 69.0 7.8 0.5 21.4 7.4
* boiled, except bread.
t 1.3 including skin
Source: Woolfe (1987).
energy. Cooking (by boiling) reduces the disparity with other foods by
levelling up the moisture content, but in developed economies this is not a
bad thing since diets are generally high in energy. In developing countries
with more restricted diets, low energy content could be a disadvantage.
Roasting, French frying (chips-UK) or chipping (crisps in the UK)
corrects the energy factor through the elevation of the fat content (Table
16.5). Dietary fibre is generally reduced on cooking, largely because fibre
is lost with the peelings. Higher values for dietary fibre contained in
roasted, French fried or chipped (crisps-UK) potatoes result from the
lower moisture content of these products.
Table 16.5 Composition of potatoes cooked by different methods (per 100 g)

Total
Crude carbo- Dietary
Cooking Energy Moisture protein Fat hydrate fibre)
method (kJ) (kcaJ) (%) (g) (g) (g) (g)
Uncooked 335 80 78.0 2.1 0.1 18.5 1.7
Boiled in skin
(flesh only) 318 76 79.8 2.1 0.1 18.5
Boiled, peeled 301 72 81.4 1.7 0.1 16.8 1.6
Baked in skin
(flesh only) 414 99 73.3 2.5 0.1 22.9 1.9
Mashed (with milk
and margarine) 444 106 78.4 1.8 4.7 15.2
Roasted (in shallow
fat, flesh only) 657 157 64.3 2.8 4.8 27.3 2.7
French fried
(chips)* 1165 264 45.9 4.1 12.1 36.7 3.3
Chips (crisps)t 2305 551 2.3 5.8 37.9 49.7 11.9
Chips per 25 g
portion 576 138 2.3 1.4 9.5 12.4 3.0
* French fries = chips in the UK.
t Chips = crisps in the UK.
Source: Woolfe (1987).
Cooking often reduces mineral and vitamin constituents. In the case of

manufacture, it is possible to replace or boost missing or low ingredients in
order to enhance overall nutritional value. Fortification is not practicable
in domestic preparation, so shortfalls need to be met elsewhere in the diet.
This may be more difficult to do in a subsistence economy. In developed
areas, the dietary contribution of potatoes can be varied by eating different
forms of the commodity. In practice, most people do in fact consume
potatoes in various forms. The only difficulty of consequence may be
excessive consumption of manufactured products (French fries and chips/
crisps) by the young.
16.2 POTATOES IN THE FOOD SYSTEMS OF DEVELOPING
COUNTRIES*
16.2.1 Production areas

Some of the worlds largest and smallest, richest and poorest, most
progressive and most backward farmers grow potatoes. Subsistence potato
growers in isolated mountain areas in Rwanda and Bolivia are among the
world's poorest farmers. Commercial growers in northern Mexico and
Brazil are among the richest. Market garderners in parts of Guatemala and
the Philippines are among the world's most intensive and productive.
Uncounted households, both rural and urban, also grow potatoes, along
with other vegetables, in home or kitchen gardens.
Potatoes are grown under a wider range of altitude, latitude, and
climatic conditions than any other major food crop - from sea level to over
4000 m elevation and from the equator to more than 40° north and south.
The diversity of agro-ecological zones in which potatoes are grown
practically defies classification. However, three ideal types of production
zone can be identified.
1. Highland zones: The Andes, the Himalayas, and other mountainous
areas scattered throughout Africa, Asia, Latin America and Oceania.
2. Lowland tropical and subtropical zones: The Indo-Gangetic plain from
Pakistan through India into Bangladesh, Peru's coast, and northern
Mexico.
3. Temperate zones: Southern Argentina and Chile, the Korean peninsula,
northern Turkey and northern China.
In highland and temperature zones, most potatoes are planted in spring
and harvested in the autumn. Cold winter weather facilitates potato
storage for home consumption, sale later in the year, and for planting in
the next season. Climatic hazards like frost, hail, and drought are major
sources of production risk. Late blight (Phytophthora infestans) affects
most highland and temperate areas; insects and nematodes present prob-
lems in some localities.
In lowland areas, farmers generally plant potatoes at the beginning of
the cool season in areas with irrigation or abundant residual soil moisture.
Potatoes are subject to fewer climatic hazards but more pests and diseases
than in cooler areas. High summer temperatures after harvest make potato
storage difficult; costly refrigerated warehouses are used in some places.
Because of the high ambient temperatures and insect populations, it is not
easy for lowland farmers to produce and store high-quality seed tubers,
and many buy seed produced in highland or temperate zones. Traditionally,
few of the varieties available were suited to lowland growing conditions.
* This section is based on Horton (1987), Chapters 4-6.

Potatoes in the food systems of developing countries 805
Recent improvements in seed technology, varieties, pest management and
storage, have resulted in a significant expansion of potato production in
many warm lowland areas, like the plains of India and Bangladesh.
16.2.2 Marketing
Conventional demand projections assume that changes in income, prices,
and food habits have little impact on the demand for potatoes. Recent
studies show that in most developing countries rising incomes and changing
food habits, associated with economic development and urbanization, lead
to greater potato consumption. Improvements in production and post-
harvest technology have helped reduce relative potato prices, and this too
has stimulated demand. As a result, the demand for potatoes has grown
more rapidly than the demand for most foods in developing countries.
Since average potato consumption is still less than a quarter of that in
western Europe it is likely that the demand for potatoes will continue to
grow rapidly in the future.
Over 98% of the potatoes grown in developing countries are consumed
domestically, rather than exported. However, in some regions, like north
Africa and the Middle East, potatoes are an important export crop that
generates significant returns in foreign exchange.
Some governments have attempted to control potato prices or dampen
their fluctuations by buying potatoes at harvest time and storing them for
later sale. Most of these programmes have been short-lived because price
movements were difficult to predict, management was poor, and storage
costs were higher than expected. Potato farmers practically everywhere
store a part of their harvest, helping to stabilize market supplies and prices
throughout the year. Recent improvements in farm-level seed storage have
helped smooth the flow of potatoes to consumer markets by allowing
farmers to grow potatoes at different times of the year.
Manufacturing potato starch and alcohol is seldom practical in develop-
ing countries because of the relative high cost of potatoes. There are a few
exceptions, however, such as northern China and southern Chile, where
potatoes are cheap at harvest time and the cost of transporting them to
urban markets is high. Potato processing for human consumption is
expanding rapidly in many developing countries to satisfy the growing
demand for French fries and potato chips.
16.2.3 Consumption and nutrition

In rural areas, potato farmers generally eat more potatoes than non-
producers. In towns and cities, potato consumption depends upon relative
prices and income levels. Where potatoes are relatively cheap, such as in
the Andes, potato consumption is highest among poor households. In the
more common situation where potatoes are relatively expensive, as in most
of Africa and Asia, potato consumption is highest among the wealthy. The
implication is that as income levels rise in the future, so will potato
consumption in most places.
16.3 POTATOES IN DEVELOPED COUNTRIES
Section 16.1 reviewed the history of potatoes in the world context. Section
16.2 dealt with potatoes in the tropics and sub-tropics - broadly synony-
mous with production in the FAO-designated developing countries. This
section considers potatoes in the temperate zone where much of the crop is
produced in countries with developed economies, taking Great Britain
(GB) as being representative of the UK.
16.3.1 Changing patterns of consumption

Figure 16.4 refers to GB and provides detail behind the trend of rising
consumption illustrated in Fig. 16.2. Increased intake as measured in raw
equivalent terms, is clearly coming about through higher consumption of
processed product, particularly French fries. Consumption of raw potatoes
has held remarkably steady overall, assisted by a slowly increasing superior
fraction sold in prepack form (selected raw potatoes in packs of say 2 or 5
kg). Similar movements 'up market' to higher-value products are taking
place in other developed countries, although the UK is virtually alone in
increasing overall consumption. It is also the only country with a Market-
ing Board which is charged with the responsibility of managing production
in relation to perceived demand through the control of area and the
removal of surpluses which may arise.
Processed product manufacture started in the UK about 25 years ago
and has since become established on the Continent fiTst in Holland and
more recently in northern France. Both the latter countries use the
versatile Dutch-bred variety Bintje to a predominant degree. The extent of
the Dutch area expansion in recent years can be partly linked to the
development of the processing sector - seed production has also increased.
The fall in domestic ware production in neighbouring Germany (formerly
West Germany) is a direct consequence of Dutch competitive advantage.
Processed product consumption in the US is often cited as being more
than 50% of the total and it is suggested that this could happen in Europe
as well. It is worth remembering that total US consumption is only about
60 kilograms per head (raw equivalent basis) - 50% of this (30 kg) is
virtually the same as GB consumption of processed products in recent
years. Neither market appears to be advancing very rapidly from these
respective positions.
Potatoes in developed countries 807
7 ,--------------------------------------------,
Raw loose
2S
c
o
aE
iilc 4
o
()
o
1980 1985
Year
Figure 16.4 Estimates of potatoes used for human consumption in UK from all
sources 1977-87. Source: Potato Marketing Board.
16.3.2 Added value

In these days of declining unit values for farm products, it is not surprising
to find increasing interest being given to adding value. Taking GB as the
example, Table 16.6 illustrates the range in prices pertaining in the 1988/89
season (produce from the 1988 crop). Farmgate prices range from £57 to
£120 per tonne for ordinary ware retailed through greengrocers to pre-
packed bakers sold through supermarkets. Of the products which are sold
in the natural state apart from selection by size and/or freedom from
blemishes, it is interesting to note that prepacks of 'baby roasts' (small
potatoes for cooking or roasting) make about the same price as bakers
(potatoes for baking). With baby roasts, the problem of course, is to
accumulate enough to be commercially viable - one of the advantages of
larger 'packhouse' businesses. Processed product mark-up is understandably
Table 16.6 Rough guide to value added - GB (1988 season)
Producers' Retail Mark (%)
price price up mark
Product (£ t·!) (£ kg-i) (£ kg·!) (£ kg-i) up"
Ordinary ware 57E 0.057 0.18 0.123 + 216
Supermarket:
'Freeflow'll 80 0.080 0.26 0.180 + 225
Prepack 80 0.080 0.29 0.290 + 263
Bakers 120 0.120 0.87 0.750 + 625
Baby roasts 80E 0.080 0.87 0.790 + 988
French fries 60E 0.060 0.35t 0.290 + 483
'Oven ready'" 60E 0.060 0.38t 0.320 + 533
Chips (crisps-UK) 80E 0.080 0.93:j: 0.850 +1063
Dehydrate (powder) 40E 0.040 0.33§ 0.290 + 725
Canned 140 0.140 0.80~ 0.660 + 471
* To cover collection, manufacture where appropriate, waste, packaging,
storage, distribution and profit.
t Per tonne raw equivalent basis 1.9:1 conversion.
* Per tonne raw equivalent basis 4.3:1 conversion
§ Per tonne raw equivalent basis 6:1 conversion
~ Per tonne raw equivalent basis 1: 1 conversion.
II Potatoes of prepack quality exposed for personal selection, typically in
supermarkets.
** 'Oven ready' French fries which need only be warmed in an oven before
serving.
Sources: Producer prices - PMB or author's estimates (E), retail prices-
Edinburgh shops or supermarkets.
greater than ordinary ware or prepacks in view of the costs of

manufacture, packaging (of particular significance to chips (crisps-UK),
etc., all of which takes place away from the farm.
The total value of the GB crop has fluctuated between £252 and £566
million during the 1980s - Potato Marketing Board (PMB) estimates - the
retail value being of the order of £1.5 to £2.0 billion, depending on the
season. Typically producer returns amount to around 25% of the final
retail value but some of the seasonal fluctuations due to varying supply/
demand/quality relationships at the producer/trade interfa·ce are effectively
smoothed out to a significant degree by the time the retail level is reached -
supermarkets in particular tend not to change prices so often as more
traditional outlets.
16.3.3 Production economics

Turning to production economics, figures for the UK are again taken to
portray general relationships. These are not thought to differ greatly from
similar production situations among larger farms in northern Europe, but
will not be representative of prodl!l:ction on small family farms with little
mechanization and selling produce at the farm gate or in local markets. As
these smaller production units are much less significant in terms of overall
production, the lack of data about them is less critical to the overview.
Table 16.7 indicates that the contribution of potatoes to total agricultural
output is relatively unimportant within UK agriculture by comparison with
cereals and livestock. Nevertheless the effect of a good or a bad year is
evident. Table 16.8 focuses in to farm level, presenting data from the Farm
Business Survey for the 1984 and 1986 crop years which are broadly
comparable with the calendar year data given in Table 16.7. General
cropping farms in England grow much of the English ware crop. Scottish
figures for similar general cropping farms indicate the greater significance
of the potato crop as a result of over 60% of the enterprise being higher
value seed production. Even ware crops at 8% to 11 % of total output are
becoming significant contributors to farm incomes, but at 17% to 22% in
the case of Scottish farms, potato output clearly has a very significant effect
on total farm profitability from year to year.
Table 16.7 Value of UK output - calendar years

Output from: 1985 1987
fm % fm %
Cereals 2281 19.3 2041 16.7
Potatoes 318 2.7 534 4.4
Other crops 616 5.2 703 5.7
Horticulture 1290 10.9 1503 12.3
Total crops and horticulture 4505 38.2 4781 39.0
Livestock and live stock
products 7289 61.8 7472 61.0
Total 11794 100.0 12253 100.0
Due to rounding, percentages do not add exactly.
Source: MAFF (1989).
Table 16.8 Whole farm results for general cropping farms -

crop years (£ 000 per farm)
Output, input or NFl England Scotland
1984 1986 1984 1986
Total output 139.2 144.0 108.0 112.7
of which: cereals 60.2 -55.4 53.3 52.7
potatoes 10.9 15.7 18.5 24.5
(%)* (7.8) (10.7) (17.1) (21.7)
Total inputs 118.3 126.4 101.9 101.4
of which crop costs 29.7 31.8 30.9 29.7
Net farm income (NFl) 20.9 17.7 6.1 11.3
* Potatoes as percentage of total output.
Source: MAFF (1987, 1988).
Table 16.9 takes us down to enterprise level, outlining results from
enterprise studies undertaken in GB in the 1984 crop year - early and
maincrop ware in England, seed production in Scotland. Updates for 1986
are also provided for maincrop seed and ware production, but unfortu-
nately are not available for the first early ware sector. Suffice to say that
early ware production is traditionally a speculative activity owing much to
the general situation pertaining in the immediately preceding maincrop.
Maincrop supplies were short throughout northern Europe in 1983/84,
such that the 1984 new crop proved to be about twice as profitable as in
more typical years.
Comparison between 1984 and 1986 maincrop ware and seed costings
demonstrate the volatility characteristic of the enterprise in which produc-
tion in European Community countries (12) only varied by 3.7% between
Table 16.9 Enterprise costings for early ware, maincrop ware and
seed production in G B
England Scotland
Early ware* Maincrop ware Seed
1984 1984 1986E 1984 1986E
£ ha- I
Output
Seed 1897 2365
Ware 4051 2167 2953 735 1080
Stockfeed NA NA NA 39 40
Total output 4051 2167 2953 2671 3485
Variable costs 1325 1141 995 1828 1688
Gross margin 2726 1026 1958 843 1797
Fixed costs 916 1052 1126 1045 1108
Estimated surplus 1810 -26 832 -202 689
Total costs 2241 2193 2121 2873 2796
Average yield Tonnes ha-I

Seed 21.3 21.5
Ware 22.0 42.4 37.4 14.3 13.5
Stockfeed NA NA NA 2.7 2.7
Total yield 22.0 42.4 37.4 38.3 37.7
£ tonne-I
Average price
Seed 89.1 110.0
Ware 184.1 49.6 79.0 51.4 80.0
·South-west England - see text. E = estimated.
Sources: Early ware - Jenkins (1985), maincrop ware - Hinton (1987), seed
- Anderson (1988).
the years - 41. 7 million tonnes in 1984 down to 40.2 million tonnes in 1986.
Costs may then compound the problem. The 1984 crop was grown from
high value seed produced in shortfall 1983 which cost £472 per hectare for
ware crops and £678 for seed. In 1986 these figures had fallen to £316 and
£430 per hectare respectively.
It is also noteworthy that Table 16.9 indicates the significant ware
tonnage being produced by Scottish seed growers. If yields were to be
restricted to seed-size tubers (in the main 35-55 mm), then seed prices
would need to rise substantially because costs are generally not volume
related. The absence of a similar degree of ware tonnage from Dutch seed
crops largely explains the higher price for their product, although Dutch
pricing is also influenced by greater control in the pricing of varieties
through the long-standing breeders' rights policy which gives control to
individual seed houses for typically up to 25 years. This differs from much
of UK seed production which historically stems from freely released
stocks. Currently this UK situation is changing to controlled release similar
to the Dutch, but it is interesting to note that their traditional seed-only
production policy achieved through early burning down is not now being
applied so strictly, with the result that a greater ware fraction is now being
produced by the seed sector in Holland.
Table 16.10 highlights the main cost items of difference between seed
and ware production which necessitate seed returns to be higher in order to
achieve financial viability. Greater seed rates - 4.1 tonnes for seed
compared to 2.6 for ware - at higher unit costs to reflect the better health
Table 16.10 Comparison of average seed and ware variable costs in G B (£ ha-l )
1984
Variable cost Seed as
item Seed (Scotland) Ware (England) % of ware
Seed 678 472 + 44

Fertilizers 191 228 - 16
Casual labour 214 111 + 93*
Contract and materials 264 191 + 38
Royalties 32t +++
Levies, inspection
-fees:j:, bags, etc. 286 139 +106
Sub total 1665 1141 + 46
Rent of land 163§ +++
Total 1828 1141 + 60
* Not necessarily significant - see text.

t Specific to seed.
:j: Inspection fees and labels cost about £65 ha- 1
§ Specific to seed in Scotland (little ware grown on rented land in England).
Sources: Seed - Anderson (1988), ware - Hinton (1987).
status of seed for seed crops, plus extra expenditure on sprays (insecticides
and fungicides), royalties and seed inspection fees are obvious differences.
Average charges for royalties will be higher in Holland in view of the
greater proportion of controlled varieties.
Much of Scottish seed production is undertaken by merchant growers
who rent potato land on an annual basis for £350 to £500 per hectare
depending on land quality and the amount of work the leasee undertakes
as part of the deal. The average charge in the enterprise study (renters and
non-renters) is shown to be £163 per hectare, a fee perhaps exclusive to
Scotland although ordinary (longer term) rentals on the Dutch polders may
come close to comparable levels for rented land taken as a whole. Total
variable cost differences apart from the special rental figure are shown to
amount to 46% in 1984.
Financial viability of seed production is more variety sensitive than ware,
where skin condition, dry matter and sugars can be as important as variety.
The earlier discussion relating to added value and retail mark-ups in the
ware sector gave a broad indicator of the likely-to-be-more-profitable
sectors to aim for within the more general financial situation discussed in
necessarily brief form here. Note that bakers, baby roasts and, in a sense,
dehydrate are subfractions of crops grown for prepacking or more ordinary
ware respectively. Even going all out for bakers or canners will leave half
or more of the crop to be disposed of elsewhere. What is very clear is that
the general condition of potatoes for washed prepacking is not easily
achieved in practice due to disease, lack of skin finish and mechanical
damage, which is mainly why sales of this category continue to be a small
fraction of the total volume produced.
16.3.4 Marketing
Marketing of the GB crop is largely left to the trade (in the main relatively
small businesses) with a low but increasing proportion in the hands of
producer-controlled co-ops. Co-ops play a bigger part in north-west
Europe, being both larger and longer established. Marketing in Holland or
France is concentrated around a few large co-ops and merchanting houses
which account for the majority of trade. While Great Britain has a Potato
Marketing Scheme, this does not lead to a single price for potatoes as we
have already seen in Table 16.6; in fact the PMB only enters the market as
a buyer of last resort (and then only for limited quantities) when supplies
exceed demand and prices fall below pre-set trigger points. Similar buying
also takes place in France (ware) and in Holland (seed).
The GB market is loosely managed in the sense of setting target areas
which should produce requirements while constraining surpluses to
amounts which can be removed from the market at reasonable cost in the
event of over-supply. With the UK effectively becoming an open market
since 1979, the downside price limitation has been less effective, leading to
the system coming under review at the time of writing. Strongly held
producer views for retention of the main points of the present scheme are
being equally strongly countered by the free marketeers. The latter are led
by processors who feel that the present system can lead to inequitable
pricing of input for processing in what has become an international market
in which advantage falls to those able to obtain suitable supplies most
cheaply. Consumer interests could be said to benefit from lower prices but
this must be seen in the context of profitable production.
If low prices arise as a consequence of over-production then the benefit
to consumers can only be temporary as producers will find it uneconomic to
continue production. It is also relevant to the argument to remember that
producer prices, as already shown, amount to 30% or less of retail values.
New developments involving fresh investment will tend to come from an
industry confident in all its sectors, which means that each must be
generally profitable in the longer term. GB consumption increases when
consumption in other countries is at best only stable, begs the question as
to whether free marketing or managed markets are best. A managed
system on the GB model comes at a direct cost (roughly £2 per tonne),
while some would say it also brings inflexibility. The counter argument of a
free market place seems more likely to result in greater volatility from year
to year, which is not necessarily in the consumers' interests either.
Alternative means of income security include traditional forward con-
tracts and the use of futures markets, there being three of the latter in
Europe - the oldest Amsterdam, London and the most recent at Lille.
Traditional contracts have had limited appeal where the required product
is not unique and alternatives can be easily acquired on the free market.
Processors usually contract some supplies for processing into French fries
for example, in order to secure a base load for their factories, but retain the
freedom to obtain a significant proportion of needs from the open market,
in an attempt to even out the effects of inter-year price uncertainty.
Where requirements are more specific, forward contracting becomes
essential as with chip (crisps-UK) manufacture in GB. The variety Record
has little domestic ware appeal in GB so little is grown outside the
contracted needs of the processors. Similarly, canning requires forward
commitment by processors to give confidence to growers to restrict
potential yield in order to maximize production of immature, small-sized
tubers.
Futures markets are a useful means of spreading risk but unlike forward
contracts cannot specifically guarantee to provide a particular price.
Futures prices respond to current expectations concerning delivery at
specific times in the future. If marketeers are pessimistic about sup-
plies being adequate and feel that a shortfall could develop, then
users (buyers of potatoes) will hedge against the perceived price rise.
Producers become interested when the view is for a price fall. The actual
level of prices traded mayor may not be adequate in terms of being
financially viable to business. If it is, then viability can be secured with
confidence.
16.3.5 Future developments
(a) Ware
Production is becoming more market-led, with supermarkets well to the
fore in their demand for a consistent supply of quality products. Speciality
products such as bakers, baby roasts and organically grown potatoes are all
being given encouragement. Tighter sizing enhances appearance. The
result is increasing pressure to concentrate grading into larger units, be
these producer or trade controlled. Whole-crop movement off farms is
becoming more significant as the ability to combine lots from different
sources to meet orders to specific specifications over longer periods of
time, becomes increasingly important to long-term survival. Outgrades at
on-farm level become viable entities, baby roasts having already been
cited, while product enhancement, seen as tighter grading, may create an
out-grade of even greater value in the form of bakers. What is very clear is
that markets are becoming more sophisticated and all sectors need to work
together to make the most of the developing situation.
(b) Seed
Quality is one of the keys to this developing dimension and mini tuber
production (first generation seed of 10--15 mm produced in a greenhouse),
may be a significant step to improving the product for end users. Still in the
early stages of commercial development, it is too early to say where the
technique will find its level. Where quality - skin finish or whatever - is the
overriding factor and market forces combine to elevate price for supplies to
specific outlets, it is possible to foresee ware crops being grown directly
from relatively highly priced ex-greenhouse seed stock. Less profitable
sectors could still benefit from the technique but after perhaps two to three
multiplication years in the field. At the time of writing it looks more like
the latter scenario in the main. This would s,till be a major advance in the
sense that ware crops would be grown from material at most 3 years away
from the mother tuber rather than the 7-10 of today.
Seed-producing countries could be working with volume production of
very high grade material involving much the same area of land, but it is
very clear that production of seed material must meet the needs of the
market place in terms of quality and volume if the seed industry is to have
long-term viability. The growing of seed on contract for specific ware
objectives looks likely to become more important to the achievement of
this goal because development costs of new or cleaned-up old varieties are
such that over-release onto an unprepared market is likely to result in price
References 815
weakness and failure to meet financial objectives. With multinational
companies taking an increasing interest in plant breeding in general,
achievement of big business norms for return on capital, etc, seems likely
to become an increasingly important part of seed production in developed
economies.
REFERENCES
Anderson, J.L. (1988) Scottish Seed Potato Production, Scottish Agricultural

Colleges, Economic Report No.2.
Eurostat (various years) Crop Production Statistics.
Hinton, L. (1987) Potatoes in Surplus. Occasional Paper No. 37, Department of
Land Economy, Cambridge University.
Horton, D. (1987) Potatoes: Production, marketing, and programs for developing
countries, Westview Press, Boulder and IT Publications, London.
Horton, D. (1988) Underground Crops - Long term trends in production of roots
and tubers, Winrock International, USA.
Jenkins, J.N. (1985) Early Potato Production in Great Britain, 1984 Agricultural
Enterprise Studies in England & Wales, Economics Report No. 95, University
College of Wales.
MAFF (1987, 1988) Farm Incomes in the UK, HMSO, London.
MAFF (1989) Agriculture in the UK -1988, HMSO, London.
Potato Marketing Board (various years) Potato Statistics in Great Britain.
Potato Markting Board (various years) Annual Reports.
Wooife, Jennifer A. (1987) The Potato in the Human Diet, Cambridge University
Press, Cambridge.
CHAPTER 17
Principles of agronomy and their

application in the potato industry
E.]. Allen and R.K. Scott
This chapter reviews the extent of progress since the first edition of this
book in the strength of understanding of the principles governing yield
formation in the potato crop. The extent to which this knowledge has been
used in the industry is considered and the chapter ends with a critical
discussion of essential future requirements for the continuation of a
profitable industry. The objectives of this chapter are, therefore, not
materially different from those of the first edition and relate to:
1. the achievement of the greatest proportion of the potential yield in all
crops, irrespective of outlet;
2. evaluation and exploitation of available varieties and provision of
appropriate criteria for the breeding of improved varieties;
3. matching field husbandry to storage conditions to ensure the highest
yield and quality out of storage.
17.1 CROP GROWTH AND DEVELOPMENT
In the first edition Scott and Wilcockson (1978) questioned the utility of
conventional growth analysis in aiding understanding of crop growth and
showed that the alternative analysis of Monteith (1977) based on (a) the
amount of light intercepted during the growing season and (b) the
efficiency with which intercepted light is converted into carbohydrate had
advantages. They showed close relationships between crop growth rate and
amount of radiation intercepted for short intervals over the first two
months after emergence and cumulative relationships between dry matter
production and amount of radiation intercepted over the whole growing
season. Similar data were known for other crops, cereals (Biscoe and
Gallagher, 1977) and sugar beet (Scott and Jaggard, 1985) and the
potential value of this analysis was clear, both in research and for the
industry. It was argued that the basis for more effective analysis of crop
Crop growth and development 817
growth than hitherto had been established. One may therefore judge
subsequent progress against these two aspects of the analysis.
Since 1978 no comprehensive study of the carbon assimilation of the
potato crop has been published. Close cumulative relationships between
dry-matter yield and amount of radiation intercepted have been published
(Allen and Scott, 1980) but it is a fair criticism to suggest that reasonably
close cumulative relationships between the two accumulating parameters
are inevitable. The real progress in the testing of the relationship has
occurred in other crops. Glauert (1983) has clearly demonstrated that for
sugar beet, assimilation is indeed driven by interception of radiation as
shown in Fig. 17.1 for a cloud-free and an intermittently cloudy day. He
found that biomass increase throughout the season obtained from net gas
exchange agreed closely with measured crop dry weights from growth
analysis (Fig. 17.2). Similar data exist for spring barley (Biscoe et af., 1975)
and thus there is no doubt that the fundamental basis for the analysis of
crop growth is sound. It is necessary, however, to infer that a similar basis
exists in potatoes for total dry-matter production and to wonder why no
serious testing has been undertaken.
(a) cloud free
i~~'-~
~ ~-~':~------~~---
\-2~
(b) intermittently Cloudy
r ====-= I
~J"~ 4~02UPtake
~:
Cl - ----------------------------------- ---
~~--
-2 r
3 6 9 12 15 18 21
TimeGMP
Figure 17.1 CO2 assimilation in relati<OJn to irradiance in sugar beet (Glauert,

1983).
818 Applications of agronomy
2.0
1.5
1.0
••
• sequential harvests
0.5 o final harvest
••
O~~--~
Jun.
____
Jut.
~ ______
Aug.
-L----~
Sep.
______
Oct.
~ ______
Nov.
~~
Figure 17.2 Crop standing weight obtained from sequential harvests and from a
refined growth 'model' plotted against time in sugar beet (Glauert, 1983). The line
shows daily dry weight increments derived from the relationship of crop growth
rate to daily intercepted radiation obtained from gas-exchange results..
Much less is known about the factors influencing the efficiency of

conversion and the magnitude' of their effects. Where growth appears
unrestricted by resource limitations, mainly water and by pests and
diseases, total dry-matter production in potatoes is determined by the
amount of radiation intercepted and accrues at a rate of 1.4-1.6 g/MJ
(Scott and Wilcockson, 1978; Hogge, 1989). This rate can be reduced by
restricted water availability and in extreme cases, e.g. in 1976, the
restriction is sufficiently severe to arrest production. Detailed information
on any progressive influence of a decreasing water supply are not known
for potatoes but again for sugar beet and barley the manifestation of water
stress begins with a hysteresis-like response whereby the afternoon photo-
synthesis is clearly less than for similar irradiance in the morning (Glauert,
1983). As water availability becomes increasingly restricted photosynthesis
is arrested progressively earlier in the day and ultimately complete
cessation of photosynthetic activity will result. Diseases are also known
to have a similar effect in sugar beet. Figure 17.3 shows that a mixture
of Beet Yellow Virus and Beet Mild Yellowing Virus decreases the
efficiency of conversion by c. 20%. Similar effects may be anticipated in
potatoes although no virus disease is as widely distributed and as frequent
in occurrence as virus yellows in sugar beet. There is clearly, as yet, no
widespread appreciation amongst physiologists, pathologists, nematolo-
gists and virologists of the utility of this analysis in the study of potato
growth.
In potatoes total dry matter production is partitioned between canopy,
1600
Ii' 1200
E
E
of 800
~
g
"C
400
o 400 800 1200

Radiation intercepted (MJ mOo)
Figure 17.3 The effect of infection with virus yellows on the relationship between
crop dry weight and intercepted radiation (Scott and Jaggard, 1985)..
roots and tubers and as the timing of tuber initiation is not fixed in relation
to emergence in different varieties there is a considerable range in the
proportion that tuber dry-matter represents of the totaL For all published
data, close, linear relationships exist between tuber dry matter yield and
amount of intercepted radiation but the slopes vary (Fig. 17.4). Varieties
which initiate tubers early, e.g. Wilja, have a considerable period of
concurrent leaf, stem and tuber growth while varieties which initiate late,
e.g. Cara, may have long periods dominated by haulm growth (Fig. 17.4).
For the Wilja type the phase when growth is exclusively in the tubers
occurs earlier and represents a larger part of the bulking period than for
the iate initiating types.
17.1.1 Interception of radiation

The amount of radiation intercepted by a crop is determined by the
time course of leaf area index in relation to the seasonal drift in
incident radiation. This focuses attention immediately on the extent to
which the growth and persistence of the leaf surface parallel and coincide
with light receipts (Table 17.1). The months May-July have similar and
maximal receipts but there is progressive decline in receipts from July
onwards. April has receipts approaching those of August but the brightest
September rarely matches the dullest August. It is our impression that
growers generally may well not appreciate the potential for growth in
early summer and the extent of the decline in potential from late July
onwards.
2200
2000 •
•
1800
•••
•
1600 •
'"'E 1400 •
•
C)
~ 1200
E
.51000
CD I
.c I
:J
I- 800 I
I
I
600 I
I
I
I
400 I
200
200 400 600 800 1000 1200 1400 1600 1800

Intercepted radiation MJ m-2
Figure 17.4 Relationship between final total tuber dry matter yield and inter-
cepted radiation for two contrasting cultivars, Wilja (open symbols), Cara (closed
symbols) (Harris, 1991); fitted line, ---, Scott and Wi\Cockson, (1978).
Table 17.1 Monthly incident radiation (mJ m-2) for growing season
at Aberporth (Dyfed) and Broom's Barn (Suffolk) for 1970-1990
April May June July August Sept
Aberporth Mean 399 547 537 536 439 302
Brightest 509 690 691 685 606 360
Dullest 256 443 399 366 307 222
Broom's Mean 381 529 550 521 449 300
Barn Brightest 494 654 678 628 529 347
Dullest 322 417 421 437 377 235
Of all the major crops potatoes probably exhibit the widest range of
displacement in time of leaf area growth curves. There are numerous
reasons for this diversity.
1. The crop is grown over a very wide geographical range, from the
Channel Islands to the Orkneys and the Scilly Isles to Kent, which
creates a range in dates of planting from January to May and even June.
This results in emergence occurring from February to June in most
years. The crop is planted throughout the year over Europe as a whole
and the potential of crops planted over the whole range of dates has not
been analysed.
2. Varieties do not emerge at the same rate (Jones and Allen, 1983) nor do
they increase in leaf area at the same rate (Fowler, 1988).
3. For any area and date of planting, emergence can be altered by
sprouting of seed (Chapter 6), depth of planting (Allen et al., 1991), soil
moisture content of soil (Firman et al., 1991b) and use of floating plastic
mulch (Bean et al., 1991). As a result the onset of leaf growth usually
varies considerably from field to field within any area of production.
None of the potato crops has a leaf surface that persists into the autumn
like sugar beet, because (i) it is defoliated in order to facilitate harvesting,
(ii) it senesces naturally (and may be finally destroyed physically prior to
harvesting) or (iii) it is destroyed by frost. The leaf surface is frequently in
substantial decline before defoliation occurs and only in indeterminate
varieties such as Cara, on organic soils or in very late seasons such as 1983
is a near complete leaf surface maintained through September.
Figure 17.5 presents most advanced and most retarded leaf growth
curves for Pembrokeshire and Suffolk, sites at similar latitude. For average
incident radiation these convert into a huge range in values of intercepted
radiation by mid-September, when defoliation would be anticipated.
Although the coastal western environment has higher incident radiation
than inland in a sunny summer, the key to this variation is the proportion
of the May-July incident radiation which the crop is able to intercept. If
intercepted radiation is converted to tuber dry matter at 1.5 g Mrl the
range in tuber yield at 20% DM is 55-85 t ha- 1 for Suffolk and 98-100 t ha- 1
for Treftoyne. The values for the brightest and dullest year are even more
extreme: 65-119 and 45-79 t ha- 1 for Suffolk and 112-128 and 60-73 t ha- 1
for Trefloyne respectively. The challenge is clear: a complete and func-
tionalleaf surface should exist for as much of the available growing season
as possible. Any advance in the leaf growth pattern which improves inter-
ception at the beginning of the season will increase tuber yields more than
an increase in persistence of leaves at the end of the season. As incident
radiation decreases from July onwards it might be expected that bulking
rates would decrease even if a full leaf cover could be maintained. Figure
17.6, taken from Allen and Scott (1980), shows this general trend in many
sets of data (for which only tuber fresh weight is available).
Even for crops harvested as earlies the target is still to maximize
radiation interception before harvest. The very early planting of such crops
increases the risk of frost damage to foliage and this also applies over a
later range of planting dates in some maincrop areas, e.g. the Fens. To
delay planting in order to eliminate the risk by delaying emergence would
x 4
C])
"0
.!;;
<1l
~ 3
<1l
1ii
C])
...J 2
d
O~----------------------------------
Apr. May Jun. Jul. Aug. Sep. Oct.
Month
Figure 17.5 Leaf area index curves for early- and late-emerging crops grown
at Trefloyne (S. Wales) and Cambridge (E. Anglia). a-a, Trefloyne (early);
b-b, Cambridge (early); c-c, Trefloyne (late); d-d, Cambridge (late).
generally delay harvest unacceptably long in the case of earlies or shorten

the growing season too much for maincrops and reduce yields to un-
economic levels for it to be a realistic alternative.
For any intended date of harvest, potential yield can be calculated, with
sufficient accuracy to be effective, from average dates of emergence and
complete canopy formation using average incident radiation values. Simple
recordings of emergence and groundcover during the growing season can
detect the seasonal deviation from the average and by comparing actual
yields with the calculated potential the true efficiency of the husbandry can
be ascertained. Few growers, and few who work in the industry, yet think
in such a quantitative and flexible way, preferring simply to regard the
consistent achievement of certain yields, e.g. 20 t acre- 1 (50 t ha- 1) as the
measure of an efficient production system. As the calculations from Fig.
17.5 have shown, the achievement of 50 or even 60 t ha- 1 would in some
years be markedly inefficient in relation to potential while in others it
would be at the limit of efficiency, even unobtainable. In order to achieve
biological efficiency and maximize yield, growers must be able to identify
where loss of potential is occurring and then consider how husbandry can
be altered to rectify the shortfall. They must, therefore, be just as
concerned with interception of radiation as researchers. A second stage is
to consider an economic analysis when the biological efficiency is known.
17.1.2 Growth and development of leaves

At the end of their analysis of potato growth in 1980, Allen and Scott
quoted Watson (1968): 'It seems to me that of all topics in crop physiology
c
•
10
•
V
9
0
/I.
8
• • A
C
•
7
:¥
Q)
Q)
~
6 , • v •
';"111 /I.
..c: /I. b-
;t:.
5
v
./1.
Q)
~ X
Cl
c
:i2
"3
4
• b- •
+. y
•
Y
co
• ••
/I.
3 A
X
2 ~ + •
x + •
+
I
0
• x
May
Fi&Ure 17.6 The relationship between bulking rates and time. e, Darby (1974),
Pentland Crown; .. /:::,., Ifenkwe (1975); .. , Maris Piper 1973; /:::,., Maris Piper 1974; •
D, Allenetal. (1979);., Home Guard; D, Vanessa; x +, Dyson (1965) King Edward
(2 seasons); T 'V, Allen (1977); T, Desiree; 'V, Maris Piper (two dates of planting);
0, J. N. Bean (unpublished), Maris Bard; +,
Wurr (1971), Pentland Crown.
the internal and external factors controlling leaf production, growth and
longevity are most in need of further study ... It is obvious that much of
the future effort of crop physiologists must be directed to studies of
morphogenesis and towards understanding how plant form is determined
... ' These comments were made when the limitations to the utility of
conventional growth analysis (Watson, 1952) were becoming apparent but
before Monteith (1977) laid the foundation of the alternative approach.
Allen and Scott (1980) commented that 'Now that we recognize more
clearly how to analyse the growth of crops, the importance of Watson's
suggestions for research is all the more urgent'. A further measure of
progress since the first edition of this book is the extent of improvement in
understanding how to generate leaf surfaces of particular composition.
Figure 17.7 presents curves of leaf area index (L) against time that show
9
8
7
x
Q) 6
"'0
.!: 5
til
~
til 4
(ij
Q)
...J
3
2
0
100 150 200 250 300
Julian days
Figure 17.7 Leaf area indices for a range of varieties and date of planting:
o Home Guard, 11 March; D Desiree, 1 April; • Maris Piper, 18 April;. Desiree,
28 May; f::, Desiree, 10 June (Treftoyne); • Pentland DeU, 15 April and 0 Pentland
DeU, 27 May (Cambridge) (AUen, 1977; Jones, 1981; Hogge, 1989; and Bean,
unpublished data).
a wide range of emergence dates, rates of increase in leaf area, peak leaf
area, persistence of peak leaf area and rate of senescence. All these
features will influence interception of radiation which as Fig. 17.8 shows
increases with increases in L but at a decreasing rate. There is some
evidence that the arrangement of leaves on the stem influences the amount
of light intercepted per unit L (Fig. 17.9). Vanessa, which has few, large,
horizontally displayed leaves intercepts more radiation per unit L than the
other varieties that have more and erectly orientated leaves. In order to
understand these effects it is necessary to study the processes leading to the
formation of the complete leaf surface and its persistence.
Number of leaves
The mainstems of potatoes carry the leaves directly or indirectly (on
axillary branches or secondary stems) and also the tuber-bearing stolons.
Firman et al. (1991a) have established a clear developmental pattern within
the stem and quantified the rates and duration of initiation for primordia.
When growth resumes after dormancy the meristems in the main buds
differentiate primordia at a rate determined by prevailing temperature.
The differentiation of primordia may continue throughout a long sprouting
period in some varieties, e.g. Arran Comet, but is arrested in others, e.g.
Maris Piper, so that the number of primordia present in the sprout at
planting can be quite different in contrasting varieties (Table 17.2).
Unsprouted seed is planted with only 3 or 4 primordia per bud. From
Figure 17.8 Relationship between radiation interception and leaf area index
(L). 0, 12 000 sets ha- I ; 6,27000 sets ha- 1 ; 0, 109 000 sets ha- I (data of Williams,
unpublished, taken from Allen and Scott, 1980).
Table 17.3 it is clear that whilst the number of primordia at planting
directly influences the number of underground nodes it does not affect the
number of leaves below the first flower which remains remarkably constant
for a particular variety. Thus the environment experienced by primordia
forming the leaves of the final canopy will be different if from sprouted or
unsprouted seed. One would expect such diverse environmental conditions
Table 17.2 Effect of physiological age of seed tubers on number of

primordia initiated by sprouts at planting (14 April) in six varieties (from
Firman et aI., 1991)
Physiological Variety
age
Home Vanessa Estima Maris Desiree Cara SE
Guard Piper
Young 11.4 12.2 9.0 9.0 9.4 10.4
0.92
Old 30.2 25.8 18.2 20.5 24.4 25.8
90
85
\ I: p'
,~
6
SO
x
"
"
/' I:
/1
.I
? : ,: i
, J • :
75 : : i
, I:
I
Ii
, ,I I
: ~ Ii j
J,' / :
0'·· /
"
~ 70
,/
,. J
:
~ til /I.
~ :
/ ,:
6 65 / :
I
o
/ I
I
a: I
.I /
/
SO
/ /
/
/
./ I
I
i /
/
55 ;" I
I
I
/
I
/
/ X Vanessa
,
I
50 / o Pentland J-"n
/ Ii Arran Comet
/
/
/
d
45
0 2 3 4 5 6
Leaf area index
Figure 17.9 The relationship between percentage radiation interception and leaf
area index in three varieties (T.C. Gillison and E.J. Allen, unpublished).
Table 17.3 Effect of physiological age of seed tubers on number of (a)

below-ground nodes and (b) above-ground nodes to the first flower in six
varieties (from Firman et al., 1991a)
Physiological Variety
age
Home Vanessa Estima Maris Desiree Cara SE
Guard Piper
(a) Young 8.7 10.5 5.8 8.8 8.3 7.8
0.76
Old 15.7 14.5 9.5 12.7 14.2 12.2
(b) Young 17.0 17.5 15.7 17.3 17.7 18.7
0.56
Old 20.2 15.7 14.8 18.7 16.5 17.8
to influence leaf growth and development but no information on leaf sizes
and petiole lengths is available.
The initiation of flowers terminates leaf initiation on the main shoot and
further leaf initiation can only continue on branches. If a branch grows
from just below the flower it is taken to be an extension of the main shoot
rather than an axillary branch and will produce fewer leaves than the main
shoot before initiating flowers. This process may be repeated several times
producing layers of leaves as described by Shepers and Reestman (1975)
and leading to fertile varieties producing a sequence of flowers from June
onwards. The extent of leaf appearance above the first flower is taken as a
measure of the degree of indeterminacy in the main shoot. Varieties show
a wide range in this character ranging from complete indeterminacy in
Cara through to almost complete determinacy in some early varieties, e.g.
Home Guard. This character determines the number of leaves which
contribute to the canopy and as determinacy increases fewer leaves will
appear and the maintenance of a complete canopy can therefore become
dependent on the survival and functioning of only those leaves below the
first flower. The extent of determinacy can be altered within a variety by
husbandry. Increasing the rate of N fertilizer stimulates more leaves to be
produced above the first flower (Firman, 1987) while increasing physio-
logical age promotes determinacy and completely suppresses branching
above the first flower in early varieties such as Home Guard and Red
Craigs Royal (Firman et al., 1991a). Thus, the ageing of seed can markedly
restrict the total number of leaves per main shoot and contribute to the
reduction in leaf area index, as shown in Chapter 6. There has, therefore,
been some progress in understanding the basic morphology of the potato
main stem and this provides the basis for examining the growth and
development of the stem crop. There is some variation between varieties in
number of leaves to the first flower but for each variety this number is only
marginally affected by agronomic treatments, site and season (Table 17.4).
Varieties originating from continental Europe, especially Holland, of
second early classification, appear to be more determinate than traditional
maincrop varieties from the UK (Firman, 1987). The variation in deter-
minacy affects the period over which leaves appear and thus the period
when dry matter is used simultaneously for leaf, stem, root and tuber
growth (see later sections). In the 'model' of Ivins and Bremner (1965) it
was suggested that tuber initiation impairs leaf growth; more recent evidence
shows that in some varieties, e.g. Cara (Firman, 1987), leaf appearance
(and hence stem growth) continues throughout the life of the crop.
It now seems that the key developmental stage is flower initiation. This
does not simply mark the end of the first phase of leaf initiation for Firman
et al. (1991a) found that the event invariably preceded tuber initiation by a
fixed interval. Although flowering and tuberization are frequently taken as
analogous processes (e.g. Ewing, 1990) the real possibility exists that one
must precede the other and that flower initiation is the essential develop-
mental stimulus for the tuberization process in field-grown plants. The
Table 17.4 Leaf and stem characteristics of three varieties on two contrasting soil
types (Allen, unpublished)
Length of 10th
No. of leaves Area of Length of 10th petiole to Stem length to
to 1st flower 10th leaf petiole to basal terminal 1st flower
(em 2 ) leaflet (mm) leaflet (mm) (em)
Variety
Fen Silt Fen Silt Fen Silt Fen Silt Fen Silt
King Edward 16.6 16.0 163 124 4.95 4.1 18.3 15.4 76.4 57.2
Maris Piper 17.4 16.4 181 119 4.8 4.0 22.1 15.5 72.2 53.5
Diana 13.7 13.5 222 148 3.9 3.3 1S.8 15.0 51.2 40.2
SE 0.35 0.53 21.1 10.1 0.33 0.29 0.89 0.87 2.44 1.61
evidence from contrasting environments suggests that for a particular

variety the interval between emergence and tuber initiation is relatively
fixed although there is a considerable range from variety to variety. The
process of physiological ageing advances initiation by hastening emer-
gence, not by shortening the time from emergence to initiation.
There are many questions which remain unanswered in the elucidation
of the processes that control growth and development of the potato stem.
An obvious conundrum is how the sprout apex modulates the separation of
nodes such that a constant number of aerial leaves to the first flower is
always achieved despite a substantial range in the number of subterranean
nodes, the latter having a direct influence on the number of sites for stolons
and tubers. The need for rapid progress in these areas is urgent if improved
control over the growth of the canopy and its relation to tuber initiation is
to be achieved.
Rate of leaf appearance

Few studies of leaf appearance in potatoes have been reported. Firman
(1987) studied the process in the contrasting varieties Estima and Pentland
Crown. He found that the rate of appearance in both varieties was linear
with time and the fit was not improved by using thermal time (Fig. 17.10).
The rate of appearance of leaves above the first flower was usually slightly
slower than those below the first flower. This is not surprising as the
activation of axillary buds is necessary to sustain main stem leaf appear-
ance and the sequence cannot be regarded botanically as one continuous
system. Nonetheless, the entire sequence could be adequately described by
a single line where nutritional status was adequate; separate lines for each
sequence of initiation above a flower were needed when plants were N
deficient.
As the appearance of leaves is usually dependent on their attaining a
minimum length it is not surprising that treatments such as N fertilizers
slightly enhance the apparent rate of leaf appearance. It is not clear why
there should be a range of rates of appearance in different seasons and
35 (a) (b)
30
(J)25
Q)
>
<1l
..9120
15
'"
~ 15
E
:J First flower
Z 10
O+-r-~~~~T-r-~~~~
120 140 160 180 200 220 240 260 120 140 160 180 200 220 240 260
Julian days Julian days
Figure 17.10 Leaf appearance in (a) Estima and (b) Pentland Crown at three rates
of nitrogen: 00, D 90 and 6. 180 kg ha- 1 (Firman, 1987).
from different dates of planting which are not reconciled by the use of
thermal time (Table 17 .5).
Leaf appearance is not related to the rate of elongation of the mainstem
nor to the rate of growth or final morphology of the individual leaves.
Consequently the same number of leaves can be carried on long or short
stems (Table 17.4), with short or long petioles, and the whole appear as taU
or squat haulms. Planting date changes plant form, with early plantings
often producing short stems and late plantings producing tall ones. This has
implications for the interception of light per unit L. Determinate varieties
may fail to achieve a complete canopy if internodes are short, and mutual
shading of the limited number of leaves is increased.
Leaf size and morphology

It has been shown that the rate of leaf expansion is increased by delay in
planting (Firman, 1987; Fowler, 1988) and by increasing applications of N
fertilizer (Firman, 1987) (Table 17.6). Date of planting had only small
effects on final leaf size as the period of expansion was reduced but no such
compensatory effects accompanied the use of N so final leaf sizes were
markedly increased (Fig. 17.11). Delay in planting is usually associated
with an increase in temperatures but neither Firman nor Fowler found that
rates of leaf expansion could be wholly and satisfactorily reconciled
through use of thermal time. While it is clear that temperature has an
influence other factors as yet unravelled are involved.
Table 17.5 Rate of leaf appearance up to the first flower in chronological and
thermal time for several varieties planted on different dates in several years
Variety Date of planting Leaves/day R2 Leaves/ R 2*
day degree
Fowler (1988)
Pentland Dell 29/4/86 0.35 81 0.029 81
Pentland Dell 28/5/86 0.64 79 0.035 78
Estima 29/4/86 0.34 77 0.028 77
Estima 28/5/86 0.58 84 0.032 84
Stalham (1989)
Record 15/4/87 0.40 99 0.033 99
Record 6/5/87 0.41 98 0.032 98
Record 8/4/88 0.44 93 0.033 93
Record 6/5/88 0.48 95 0.033 93
Hogge (1989)
Pentland Dell 15/4~7 0.35 89 0.033 93
Pentland Dell 15/5/87 0.39 96 0.033 96
Pentland Dell 27/5/87 0.52 96 0.034 96
Firman (1987)
Estima 19/4/85 0.34 96 0.024 96
Estima 14/3/86 0.36 98 0.028 98
Estima 11/4/86 0.36 98 0.030 98
Estima 12/5/86 0.41 98 0.023 98
Pentland Crown 19/4/85 0.34 96 0.023 96
Pentland Crown 14/3/86 0.35 98 0.028 98
Pentland Crown 1114/86 0.33 98 0.027 98
Pentland Crown 12/5/86 0.41 98 0.023 98
• R2 = percentage variation explained by the regression parameters.
The position of leaves on the mainstem is known to affect the rate of

expansion and final size of leaves (Firman, 1987; Fowler, 1988). Above the
two or three juvenile leaves nearest the ground, leaf size increases to a
max mum and then decreases as the flower is approached (Table 17.7).
Table 17.4 shows that the length of petioles, both to the basal and terminal
leaflets, can vary considerably and tends to be consistently greater on
organic soils than on other soils.
Leaf longevity
Firman (1987) found that the life-span of individual leaves was principally
determined by the prevailing light environments for the longevity was
curtailed in shade. The continued 'piling on' of layers of leaves by
Table 17.6 Effect of rate of N application (kg ha- l ) and date of planting on the
initial rate of leaf extension (mm/day) of selected leaves in Estima and Pentland
Crown (Firman, 1987)
Date of planting
14 March 11 April 12 May
N (kg ha- 1): 0 90 180 0 90 180 0 90 180
Fifth leaf
Estima 7.5 8.4 10.8 8.9 9.6 10.9 12.1 11.7 11.8
Pentland
Crown 6.3 6.5 8.0 6.4 8.7 9.9 8.6 9.3 8.4
SE for all comparisons 1.09
Tenth leaf
Estima 6.9 11.7 15.3 10.3 10.7 11.7 10.7 9.9 14.6
Pentland
Crown 4.2 11.3 15,9 8.3 7.9 11.8 11.6 18.6 16.2
SE for comparisons at the same date of planting 1.57
SE for all other comparisons 1.80
Fifteenth leaf
Estima 2.5 3.7 7.8 1.2 4.5 10.1 5.2 11.7 8.6
Pentland
Crown 2.8 4.6 9.0 7.2 6.8 10.8 8.3 18.4 8.0
SE for comparisons at the same date of planting 1.14
SE for all other comparisons 1.10
Table 17.7 Leaf lengths (mm) at specific above ground nodes numbered
acropetally for several varieties (Firman, 1987)
Cara Diana Estima Maris Pentland Pentland SE
Piper Dell Crown
Fifth leaf
0 136 166 203 148 163 174 16.3
kgN ha- 1 90 186 209 204 190 181 169
180 191 198 204 223 213 172
Tenth leaf
0 171 149 216 164 189 211 14.6
kgNha- 1 90 240 203 205 227 187 210
180 264 213 226 259 229 234
Fifteenth leaf
0 151 74 94 115 140 152 17.6
kgN ha- 1 90 198 113 91 185 130 179
180 229 186 154 237 176 207
Twentieth leaf
0 110 26 3 70 80 111 15.5
kgNha- 1 90 151 21 20 107 90 111
180 188 136· 83 171 126 174
(a) Planted 14 March (b)

300
250
200
150
100
50
0
160 170 180 190 200 210
160 170 180 190 200 210
Planted 11 April
300
250
1.c
200
0, 150
c
~
(tj 100
.3
50
160 170 180 190 200 210 160 170 180 190 200 210
Planted 12 May
300
250
200
150
100
50
o
160 170 180 190 200 210 150 160 170 180 190 200 210
Julian days
Figure 17.11 Extension of the fifteenth leaf in (a) cv. Estima and (b) c.v. Pentland
Crown at three dates of planting with three rates of N application: 0 0, 0 90, l:,
180 kg ha- 1 (Firman, 1987).
indeterminate varieties results in a more rapid turn-over of leaves than in
determinate varieties (Table 17.8). On the other hand, the maintenance of
a substantial canopy and maintained growth rates in varieties such as
Estima is dependent on the upper leaves, just below the first flower and on
the first branch, remaining active for months rather than weeks. Fortunately,
Firman and Allen (1988) found only small decreases in photosynthetic
efficiency with increasing leaf age up to 6 weeks and illuminated leaves
retained photosynthetic capacity after 10 weeks. Nonetheless, it is clear
that the more determinate varieties are heavily and vulnerably dependent
upon a relatively small number of leaves for the attainment and main-
tenance of a complete groundcover. These leaves must function with
unimpaired efficiency over many weeks if the overall efficiency of con-
version of intercepted radiation is to be maintained.
Table 17.8 Leaf longevity (days) in six varieties at Cambridge University Farm
(Firman, 1987)
Cara Diana Estima Maris Pentland Pentland SE
Piper Dell Crown
Fifth leaf
0 70.0 85.7 71.7 85.7 77.0 92.7 6.44
kg N ha- 1 90 54.2 66.5 50.7 61.2 82.2 71.7
180 47.2 66.5 61.2 45.5 50.7 50.7
Tenth leaf
0 94.5 70.0 70.0 94.5 77.0 117.2 5.90
kgNha- 1 90 80.5 71.7 73.5 82.2 103.2 105.0
180 63.0 78.7 75.2 75.2 73.5 87.5
Fifteenth leaf
0 112.0 58.7 61.3 94.5 73.5 101.5 7.13
kgN ha- 1 90 101.5 66.5 66.5 85.7 92.7 96.2
180 71.8 73.5 68.3 96.2 82.2 98.0
The canopy
As the size of the leaf surface increases the proportion of incident radiation
which is intercepted also increases but at a decreasing rate, so that a leaf
area index of 3 intercepts approximately 85% of the incident radiation
(Fig. 17.8). Further increases in L only marginally increase interception
and complete light interception is never achieved. Increasing the leaf area
index above 3 can only therefore significantly affect interception of
radiation if it leads to increased persistence that prolongs the period of sub-
stantial interception. This is not usually the case for two reasons. First, the
continuing growth of leaves can only be achieved by increasing growth of
stems and branches which hastens the death of shaded leaves, and the net
effect is to have a stable number of layers of leaves. Second, markedly
indeterminate canopies are prone to lodge. Both factors limit the possibili-
ties for prolonging light interception and in practice very high peak leaf
area indices are frequently ephemeral (Fig. 17.7).
The rate of increase in L is of course influenced by the number of stems
as well as by the growth of leaves on individual stems. Crops grown at high
densities, e.g. varieties that have few tubers per stem, can achieve more
rapid rates of increase in L than crops grown at lower densities. Wurr et al.
(1991) have shown for five varieties that the number of tubers per stem is
not itself linked with any other character. Thus, the implications for
varietal improvement are that more rapid canopy closure should be
achievable by increasing the number of stems without unacceptable
increases in number of tubers which would adversely affect the grading of
the crop.
The initial rate of increase in L will not be influenced by the number of
leaves per stem but low temperatures, insufficient N and restricted water
availability will impair the rate of leaf expansion and shorten the inter-
nodes leading to restricted cover and reduced interception per unit L.
Varieties which produce leaves over a prolonged period, although re-
stricted by these factors initially, may eventually produce and maintain
complete cover. If more determinate types are restricted over their short
period of leaf appearance they will be particularly vulnerable and probably
fail to achieve complete cover. Such types will be much more sensitive to
early planting and hence emergence than indeterminate varieties; it is
likely that early planting, insufficient N and water shortage will exaggerate
determinacy, thus compounding the problem. In this respect water avail-
ability has effects which are additional to those relating to the evaporative
demand of the leaf surface. As Fig. 17.12 shows the emergence and growth
of the leaf canopy can be restricted where no water is added to soil supplies
after planting, even though the soil was at fi:eld capacity before cultivations
began. The root system had reached a depth of c. 20 cm by emergence and
had access to sufficient water to meet any atmospheric demand yet the rate
of increase in groundcover was reduced. If water was applied after
emergence the rate increased and within a few days leaf coveT was similar
to that resulting from very frequent watering and increasingly greater than
where no water was applied (Stalham, 1989).
The growth of the canopy has been considered so far primarily in
relation to the main stems and the upper branches. Varieties differ
markedly in the extent of branching and some develop branches at other
nodes than immediately below flowers. As growth proceeds the contribu-
tion of branches to crop leaf area increases and with some varieties
branches contribute the vast majority of the total leaf area for the latter
part of the season (Table 17.9). As the lower branches carry younger
leaves than the mainstem and its succeeding upper branches this may well
reduce the mean age of leaves within the canopy. The time of appearance
100
80
~
60
§
"'C
§ 40
e
(!l
20
O+-~--~~--~~--~-T--~-T--~~~
130 150 170 190 210 230 250
Julian day
Figure 17.12 Effect of irrigation regime on ground cover percentage of Record

grown under polythene rainshelters (Stalham, 1989). 0, No irrigation; ., irrigated to
maintain SMD less than 10 mm; +,
irrigated with 15 mm at an SMD of 20 mm
during tuber initiation, then 20 mm at 40 mm SMD.
Table 17.9 Component parts of leaf area index in two varieties grown at three rates
of N (Firman, 1987)
Pentland Crown Estima SE
Fraction kg N ha- 1 : 0 90 180 0 90 180
of leaf Date of
index harvest
Below 13 Jun 0.50 0.48 0.39 0.83 0.76 0.77 0.099
flower 27 Jun 0.91 1.61 1.79 1.71 2.60 2.78 0.269
11 Jul 1.08 1.47 1.72 1.74 1.63 2.01 0.206
25 Jul 1.32 1.63 1.71 1.61 1.60 1.88 0.206
15 Aug 1.57 1.96 1.84 1.39 1.69 1.59 0.265
12 Sep 0.70 0.72 0.76 0.09 0.19 0.21 0.117
Above 11 Jul 0.01 0.07 0.09 0.05 0.19 0.21 0.051
flower 25 Jul 0.13 0.26 0.33 0.10 0.41 0.47 0.069
15 Aug 0.43 0.95 0.85 0.23 0.44 0.65 0.168
12 Sep 0.77 1.03 1.09 0.08 0.49 0.25 0.304
3 Oct 0.30 0.22 0.33 0.00 0.00 0.03 0.080
Branch 11 Jul 0.09 0.27 0.36 0.06 0.53 0.45 0.208
25 Jul 0.18 0.18 0.69 0.09 0.73 0.89 0.176
15 Aug 0.44 0.82 0.77 0.23 0.70 1.18 0.165
12 Sep 0.82 0.91 1.43 0.06 0.21 0.50 0.160
3 Oct 0.15 0.17 0.29 0.00 0.00 0.02 0.083
and longevity of branches is of importance to the understanding of the
dynamics of the canopy. Branches are the only means by which certain
varieties, e.g. Desiree, and treatments, such as N can prolong the life of
the canopy. Planting density can reduce the degree of branching but the
appearance of branches does not seem to be predictable for its significance
to be properly assessed as yet.
Indeterminate varieties are better able to maintain a leaf area index
capable of intercepting most of the incident radiation for a considerable
part of the growing season than determinate varieties. Crop senescence
(decreasing interceptory capacity) and leaf senescence will occur syn-
chronously in determinate varieties but for indeterminate varieties a long
period of leaf senescence can occur without overall crop senescence.
Eventually for all varieties the canopy begins to decrease in size and
interceptory capacity and senescence is then usually inexorable, leading in
some varieties to a complete loss of leaf area before the end of the growing
season even in the absence of defoliation. Agronomic treatments that
markedly change the size of the leaf canopy often fail to change the timing
of the onset of senescence, and although rates of senescence may be
different, the complete death of canopies of very different maximum size
can be almost synchronous. Figure 17.12 shows an example of this:
severely restricted water supply considerably reduced peak L but canopy
senescence began at the same time whether or not the crop was affected by
water stress. The causes of this synchrony are not clear and the example
given is not readily explicable through N availability or leaf N content.
There is some evidence that the onset of senescence is linked to cessation
of root growth and it is perceived by growers as leading to increasing skin
set. However, Wilcockson et al. (1985) have shown that suitability for
harvesting is not dependent on visual signs of crop senescence nor on skin
set and it may be that termination of leaf appearance is the key event which
presages the final stage of the crop's life.
Efficiency
The recognition that dry matter production is determined by the amount of
radiation intercepted by crops represents a major improvement in the
analysis of crop growth. It has already been shown that for long periods the
conversion of intercepted radiation into dry matter proceeds at a constant
rate providing gross nutrient and water deficiencies are avoided. Con-
sequently, the practical application of the analysis has concentrated on
maximizing the radiation intercepted by crops. It must not be overlooked,
however, that efficiency of conversion can vary and in some environments
may be an important hictor in determining dry-matter production. In
temperate regions, limited water availability has been shown to reduce
efficiency (Scott and Wilcockson, 1978) and in hotter environments the
high temperatures lead to reductions in efficiency. Trebejo and Midmore
(1990) showed quite different efficiencies for two seasons differing by WaC
in maximum temperatures (18 and 28°C) from Australia. There is also
evidence that crop photosynthetic rate can be saturated at radiation
intensities experienced by many crops (Sale, 1973). The extent and
significance of these effects on efficiency of conversion of intercepted
radiation require study and will be of increasing importance as the crop
spreads into a wider range of environments.
17.1.3 Growth and functioning of roots

In order to maintain growth at the potential rate set by incident radiation
and by the size of the canopy, crop water uptake must meet the evapora-
tive demand from the leaf surface. Penman (1948, 1970) has summarized
the influence of meteorological conditions on the evaporation of water
from crop surfaces and his estimates of crop evapotranspiration for average
weather conditions are available for all areas of the UK (MAFF, 1967). As
leaf cover and incident radiation increase, the achievement of potential
growth rate requires the evaporation of more water. In order to meet this
demand the crop must be able to extract water from the soil at the required
rate then and throughout the remainder of the season. Thus, the extent
and capacity of the root system, the water-holding capacity of the soil and
the opportunity to supplement soil water with irrigation will determine
whether a crop maintains potential growth rate and hence achieves the
potential yield.
Studies of the form and function of potato root systems are few and
evidence is sparse to examine the activity of roots in relation to the creation
and maintenance of the leaf surface. There has been a general view for
many years that the potato crop is shallow rooting (Weaver, 1926;
Lesczynski and Tanner, 1976) and heavily reliant on supplementary
applications of water to maintain growth rates. As a consequence the crop
is seen as a prime risk in relation to nitrate pollution of ground water,
because with widespread irrigation any rain will fall on soils not far from
field capacity and hence vulnerable to leaching. Studies of root growth of
crops grown at the Cambridge University Farm during the 1980s and a
review of the published literature reveal a general pattern of growth that
contrasts somewhat with earlier perceptions.
Table 17.10 shows that for many varieties, grown over a number of sites
and seasons, roots reached a depth close to, or in excess of, 1 m.
Measurements in Holland by van Loon and Bouma (1978) show that root
penetration proceeds linearly with time unless slowed or prevented by
increasing resistance due to soil compaction. Stalham (1990, unpublished)
confirmed that, for a number of farm crops in England grown on soils free
from impedance, penetration was linear with time (Fig. 17.13) and generally
ceased around the time of cessation of leaf appearance. It is therefore
likely that root growth will cease earlier with determinate varieties and
Table 17.10 Maximum measured depths of root penetration for various
cultivars
Variety Depth (cm) Source
Bintje (Neth.) 97 BOOlle et al. (1978)
Cara 140 Stalham (1990, unpublished)
Desiree 100 Stone (1982)
Early Ohio (USA) 140 Weaver (1926)
Maris Piper 110 O'Brien (1984, unpublished)
Pentland Dell 140 Hogge (1989)
Record 120, 127 Allen and O'Brien (1985), Stalham (1989)
White Rose (USA) ,150 Wolfe et al. (1983)
they will have access to considerably less soil water than indeterminate
types. Limited evidence on maximum depth of rooting of different
varieties revealed that Cara, the most indeterminate, did root to the
greatest depth. However, more data are needed to properly establish
whether consistent varietal effects do occur.
Allen and O'Brien (1985) and Stalham (1989) found that maintaining
low soil moisture deficits early in the life of the crop reduced maximum
depth of rooting. The latter also found that density of rooting in the upper
horizons was increased by moist conditions and the effect was detectable
by emergence (Table 17.11). Similar effects have been reported by Squire
(1990) for groundnut and millet and it seems clear that the water status
of the soil from immediately after planting can affect density and depth
of rooting and as described on p. 834 the rate of increase in leaf area.
Thus, the study of root growth must be conducted with regard to the soil
Days after planting

20 40 60 80 100 120
0
20
40
E
~
.c 60 •
a
Q)
I!I
Q
80
• I!I
I!I
I!I
100
120
• •
Figure 17.13 Measured depths of rooting below ridge apex in Estima and Maris
Piper (Stalham, 1990, unpublished). 0, Estima, +,
Maris Piper.
conditions prevailing. For example, any measurements of root charac-
teristics made on crops irrigated from close to emergence would never
detect the capacity for deep rooting in drying profiles. The root system
overall produced 12-15 km m-2 of root in Stalham's experiments, a similar
value to those of Lesczynski and Tanner (1976), Asfary et al. (1983) and in
the middle of the 8--21 km m-2 range found by Parker et al. (1988). These
may be compared with 11.8 for wheat (Welbank et al., 1974),5.9-12.9 for
sugar beet (Brown et al., 1987) and 12.6 for barley (Welbank et al., 1974).
It does not seem therefore that potatoes can be properly regarded as
particularly shallow or sparse rooting.
Table 17.11 Effect of irrigation regime on rooting parameters of

Record grown under polythene rainshelters (Stalham, 1989)
Irrigation regime*
Dry Moist Wet SE
Number of root initials 39 43 61 6.2
per plant at emergence
Depth of rooting at emergence 21.6 21.7 18.5 1.00
(cm)
Maximum depth of rooting (cm) 118 99 88 8.4
* Dry, nO,t irrigated; Mo'ist, irrigated with 15 mm at 20 mm So'il mo'isture
deficit during tuber initiatio'n, then 20 mm at 40 mm soil} mo'isture deficit;
Wet, irrigated to, maintain So'il mo'isture deficit less than 10 mm.
Stalham (1989) found that the root system extended and ramified quite
rapidly both vertically and horizontally and initially the volume of soil from
which water is extracted increases rapidly as extension proceeds down-
wards and outwards. The density of the complete root system was uniform
to a depth of 40-60 cm after which rooting was less intensive (Fig. 17.14).
Of particular interest was the finding that throughout the profile root
density was little affected by position of sampling, i.e. whether in row or
furrow (Fig. 17.15). The root system thus ramifies the whole soil profi\.e
very uniformly.
Such knowledge of the growth of the root system allows the possible
contribution of soil water reserves to crop water demand to be estimated
for any soil profile if the water release characteristics are known. Stalham
(1989 and unpublished) used the neutron probe to measure soil water
changes and was able to show substantial uptake from the deepest layers of
the profile in which roots were found (Table 17.12). Interestingly, there
was greater uptake in many deeper horizons by Record than Cara despite
less intensive rooting. Stalham also showed that uptake from lower
horizons could occur simultaneously with uptake from re-wetted shallower
horizons. A greater depth of rooting will increase the contribution that soil
water reserves can make to crop need and thereby reduce the amount of
(8) Early-irrigated
Root length (em em-3 )
o 2 3 4 5
10
20
30
E 40
~
..c::: 50
a
~ 60
70
80 TRL = 13.4 km m-2
90
(b) Late/irrigated
o 2 3 4 5
10
20
30
E 40
~
..c::: 50
a
~ 60
70
80 TRL = 13.7 kmm- 2 ; SE = 1.16 km m-2
90
Figure 17.14 Effect of irrigation regime on root length density in Record

(Stalham, 1989). TRL = total root length.
o Core sampling position

X Plant position
4.13
1.58
2.51 3.47
2.17
1.63
0.36 0.24
0.17 0.18
0.14 0.20
WT WT
Depth S.E.
10 0.410
20 0.424
30 0.255
40 0.318
50 0.233
60 0.226
70 0.049
80 0.026
90 0.016
WT = water table 100
Figure 17.15 Effect of sampling position on root length density (cm cm- 3 ) in earJy-
irrigated Record (Stalham, 1989).
irrigation required. It also means that overall a higher soil moisture deficit
will be tolerable. To know that there is an increasing allowable deficit as
the season progresses is crucial, for the likelihood of unexpected rainfall
returning the soil to above field capacity and resulting in leaching is
automatically reduced if this schedule is followed. The increased though
still rudimentary knowledge of the time course and pattern of root growth
indicates that the capacity to extract water will increase over much or all of
Table 17.12 Effect of irrigation regime on maximum

horizon soil moisture deficits (mm) occurring under
polythene rainshelters
Record (Stalham, 1989)
Horizon Irrigation regime*
(em) Dry Moist Wet
0-10 4.5 4.3 1.4
10-20 10.9 8.9 3.3
20-30 12.1 8.0 5.7
30-40 12.5 7.9 6.1
40-50 13.1 7.1 6.3
50-60 12.3 6.0 5.0
60-70 11.7 5.9 4.4
70-80 10.0 6.1 4.2
80-90 7.0 6.0 4.4
Cara (Stalham, 1990, unpublished)

Horizon Irrigation regimet
(em) Dry-Dry Dry-Wet Wet-Dry
0-10 5.3 4.5 4.6
10-20 14.7 11.6 13.8
20-30 12.8 10.5 13.0
30-40 10.1 9.3 10.8
40-50 8.7 5.5 7.9
50-60 7.7 4.5 6.3
60-70 6.6 3.7 6.0
70-80 6.0 2.7 6.3
80-90 6.4 3.2 6.2
90-100 6.1 3.0 6.3
100-110 5.8 2.9 6.1
* Dry, not irrigated; Moist, irrigated with 15 mm at an SMD of

20 mm during tuber initiation then 20 mm at 40 mm SMD; Wet,
irrigated from planting to maintain SMD less than 10 mm.
t Dry-Dry, not irrigated; Dry-Wet, irrigated with 20 mm from
end of tuber initiation at an SMD based on easily-available
water within rooting zone; Wet-Dry, irrigated with 20 mm at 20
mm SMD only until end of tuber initiation.
5 /---... .......
/ "-
/ '\
/ \
4 / \
/ \
/
/
l-
f,
;;
f,
!J
Planting
I Tuber initiation
Emergence'Estima Cara
O+-~~-L--+-~L,----.----.--~
July Aug. Sept.
125
c::
o
~
~ 50
c::
Q)
a.
~ 75
.s:::
g.100
o
,
\
\
,-----
\
125
Figure 17.16 The time courses of leaf area index and depth of root penetration for
a determinate variety based on Estima ( ) and an indeterrninate one based
on Cara (----).
the period over which leaf cover is increasing (Fig. 17.16). As roots extend
and rainfall usually replaces some of the water used, it would seem unlikely
that irrigation would be necessary to maintain growth in the early stages.
The practical concern is how to estimate the deficits at which growth can
just be sustained, the essential objective of irrigation scheduling. When the
season is dry, irrigation will be needed earlier on sandier soils than on more
moisture-retentive ones. It is therefore necessary to establish the amounts
of water which can be removed from soils without prejudicing growth and
use these values as the trigger for starting irrigation. As all water is not
equally extractable the concept of easily available water, i.e. that water
held between field capacity and a limiting tension, is useful. The key
questions are the value of the latter and whether it is the same for all soil
horizons. Published values for the upper limit range from 0.25 bar (Epstein
and Grant, 1973) to 2 bar (Bailey and Minhinick, 1989) with permanent
wilting point at 7 bar (Campbell, 1972) or 10 bar (Cook and Dent, 1990).
As the scale of water release is logarithmic, the amount of water held
between 0.25 and 2 bar on even a light, sandy soil is considerable, of the
order of 5 mm per 10 cm depth. If these extremes were taken for such a soil
with a depth of rooting of 50 cm the easily available water would be 30 mm
at 0.25 bar and 57 mm at 2 bar. A considerable difference and the
difference would be larger on more moisture retentive soils. If the rooting
depth extended to 100 cm and water in all horizons was similarly available,
the easily available water would be double, and this has major implications
for irrigation scheduling (Section 17.2.6). It is not an appropriate strategy,
in very dry years, to exhaust first all water in the rooting depth and then
continue growth on water from irrigation. Growth would be checked by
the inevitable reduction in availability (increasing tension) from existing
rooting depth and the reduction would be likely to restrict the rate of root
penetration limiting access to additional supplies. The use of irrigation to
maintain growth will aid root penetration and the increase in available
water must be accounted for in the calculations of crop water use. If root
penetration becomes deeper than an assumed depth of rooting, the soil's
capacity to meet demand will be underestimated and irrigation applied at
any trigger deficit before the water is really needed, with attendant risks of
leaching. If the soil's capacity is related to depth of rooting it is inevitable
that allowable soil moisture deficits will increase as the season progresses.
17.1.4 Tubers
The initiation of tubers has long been regarded as a key developmental
occurrence in the life of the crop, although since tubers are truly an
extension of the main stem the process of swelling might be regarded as
developmentally neutral. As indicated in Section 17.1.2.1 more recent
evidence suggests that floral initiation is probably the developmental key.
It is important to make clear that all varieties initiate floral primordia but,
in some, development does not proceed beyond a certain stage and open
flowers are rare or absent. This view of developmental control through
floral initiation is supported by the observation that potato stocks tuberized
when tobacco stems that flowered were grafted onto them, but not when
the tobacco did not flower (Chailakhyan et at., 1981; Martin et at., 1982).
Whatever the cause of tuberization, the timing of the event has been taken
largely to predetermine the pattern of susequent growth. Ivins and
Bremner (1965) suggested that if tuberization occurred early, subsequent
leaf growth would inevitably be restricted. More recent knowledge makes
it clear why Ivins and Bremner (1965) arrived at their conclusion and why
Allen and Scott (1980) were justified in questioning whether this was the
general rule. Ivins and Bremner's 'model' related to the observed effects of
early tuberization resulting from planting aged seed of the early variety
Ulster Chieftain which we now know would be determinate. Thus, it would
The use of agronomic principles 845
initiate early but would not be able to produce leaves above the first flower.
If un sprouted seed had been used, emergence would have been delayed,
the interval to tuber initiation would have been much the same but a larger
canopy would have been formed from leaves on branches above the first
flower. The other varieties were not determinate, initiated later, probably
produced many leaves and hence had larger leaf areas. If aged seed of
Ulster Chieftain had not been included in the experiment the apparent
.effect of tuber initiation on subsequent leaf growth would not have been
suspected.
In many experiments tuber initiation is not measured or is measured so
infrequently that it is difficult to time its onset and hence duration with any
accuracy. Where tuber initiation has been recorded properly, it seems that
for any given variety it begins a fixed time after emergence and lasts no
more than 2-3 weeks during which time the number of tuber initials rises to
a peak. There is a considerable range in the interval from emergence to
onset of initiation from variety to variety; in some it begins at very low leaf
areas and ends before complete ground cover while in varieties such as
Cara it may not begin until complete ground cover is achieved. In the
former types initiation is over within 4-5 weeks of emergence while it may
be 10 weeks before it ends in Cara. It would be of interest to know if
indeterminate types exist which initiate flowers and tubers earlier, for this
would appear a most effective combination of characters and an appro-
priate selection pathway towards increasing yields over the whole growing
season.
As haulm, root and tuber growth continue for differing periods both
within and across varieties, the study of partitioning of dry matter in
potatoes, e.g. through the use of terms like harvest index, is fraught with
difficulties and not amenable to direct analysis. Indeterminate types
inevitably have a lower proportion of their dry matter in tubers than
determinate types. The general pattern of timing of leaf growth, root
growth and flower and tuber initiation is shown in Fig. 17.16 for a
determinate variety (based on Estima) and for an indeterminate type
(based on Cara). These patterns are now used to illustrate how this
understanding can explain effects of agronomic treatments.
17.2 THE USE OF AGRONOMIC PRINCIPLES IN UNDERSTANDING

THE EFFECTS OF AGRONOMIC TREATMENTS
17.2.1 General principles

The previous section has established that differences in yield between
crops or treatments only occur as the result of differences in amounts of
intercepted radiation, provided that the water requirement of these crops
is fully met and they are maintained free from diseases, i.e. that efficiency
is not impaired. The guiding principle for understanding tile effects of
agronomy is as stated by Scott and Wilcockson (1978): ' ... any factor
which leads to increased radiation interception is likely to increase
biological yield commensurately.' For each variety, tuber dry-matter yield
will also increase proportionately. The interpretation of the effects of
agronomic treatments is possible through measurement of amounts of
intercepted radiation and allows prediction of effects in other environ-
ments to be made. This cannot be done when only the yields of different
treatments are known; these are the end effects of unmeasured differences
in intercepted radiation. The essence of agronomy is to understand how
growth and development of the leaf and root system can be manipulated to
ensure that radiation interception is indeed maximized and efficiency
maintained. The effect of both environmental variables and many agro-
nomic treatments on the leaf surface has been established and this
information can be applied in different circumstances from those in which
the experiments were themselves conducted, It is possible to predict the
direction of the effect of many agronomic variables singly and in combina-
tion without further agronomic experimentation, and even to do so where
no experimental results exist. As we cannot predict incident radiation, for
progress to be made we must concentrate on the factors controlling leaf
growth. In practice manipulation of the leaf surface has to be the direct
influence on radiation interception and growers must seek to generate the
most effective leaf surface for their conditions. We are dealing with one
component of the cause of differences in potential yield and even without
knowledge of incident radiation we can identify the causes of treatment
effects and avoid wrong attributions. The latter is likely where only final
yields can be considered because as indicated earlier many factors may be
involved and effects may be in different directions in individual data sets.
The ability to predict reduces the need for experiments in numerous
disparate environments which in any case are often averaged to provide
advice. Some examples are necessary to illustrate the potential value of the
approach, i.e. to identify the limiting factor and react accordingly.
The previous sections have summarized current knowledge of the factors
influencing leaf growth and although understanding is imperfect it is
possible to explain the effects of many treatments on leaf growth, the
starting point for understanding their effect on yield. In Fig. 17.17 the
effect of sprouting (ageing) of seed and delaying planting on the time
course of leaf area indices are shown. Old seed and early planting
invariably produce leaf canopies that are smaller but earlier than those
from younger seed and later planting. However, the relative values of the
leaf area indices and hence their influence on radiation interception are
very variable and lead to a large range in effects on yield, presumably
through a similar range in amounts of intercepted radiation, In the case of
seed ageing this led to repeated debates as to the merits of the practice
basically because the size of the effect on yield ranged from insignificant to
very large but often averaging close to the minimum value required to
justify the expense of sprouting. Consequently the justification for the
practice seemed unclear to many growers who were apparently under the
impression that the effect of ageing should be a consistent number of
tonnes per hectare each year. The annual variation in leaf growth and
incident radiation make such variable effects inevitable and the utility of
sprouting cannot be assessed simply through the average effect. It has to be
assessed through the likely effects in a specific case. For example where
emergence is very rapid because planting is late and the soil rich in Nand
water as on the Fens, sprouting can only be really beneficial as an insurance
against very late planting because the benefits from small differences in
emergence only become significant when the growing season is greatly
shortened. Equally where harvesting is advanced the greater is the
likelihood of benefits from sprouting on all soils.
6
/..-- ......
/
I
/
/
/
I.
<--,,-->
(a) (b) (c)
March April May June July Aug. Sept. Oct.
Figure 17.17 Leaf area indices for (a) very early (February), (b) average (March)
and (c) late (May) planting of two ages of seed ( , old seed; ----, young
seed). Based on data for Desiree from several sites.
A similar interpretation can be drawn from the effects of date of planting

on leaf growth. Although gross leaf area may increase as planting is
delayed, all plantings may achieve near complete ground cover and effects
on yield will be dependent on overall radiation interception. Key factors
determining the outcome will be the extent of advance in early leaf cover
caused by early planting combined with effects on timing of senescence. In
a perfect growing season in which an early start to planting is possible one
might expect yields to decrease as planting is delayed and there was a time
when growers aimed to increase yield by lengthening the interval (notionally
the growing season) between planting and harvest. However, the perfect
season rarely occurs and some early plantings may not ever achieve
complete ground cover while in other seasons the smallest canopies may be
produced from intermediate planting dates but rarely from the last.
Therefore, the effects of planting date are like sprouting; they cannot be
summarized in any particularly useful way on the basis of an average loss in
yield per week's delay in planting. They have to be considered in relation
to how the causal factors are operating in individual circumstances. From
both observation and experiments delayed emergence and small leaf
canopies occur where planting takes place into cold, cloddy, dry, nitrogen
deficient soils and in practice the effect of these factors can be minimized
providing the grower accepts that planting on the earliest possible date is
not the universal objective. As cultivation when the soil is marginal in
terms of water content inevitably leads to subsoil compaction, reduced root
penetration and hence restricted nitrogen and water uptake, the separation
of causal factors reflected in final yield is extremely difficult. Although no
experiment has tested all these ·effects it is probable that they are additive
and certain that effects on determinate varieties such as Estima will be
more severe than with less determinate varieties such as Cara.
The magnitude of the effects in any environment will be determined by
the size and persistence of the leaf surface and by the prevailing seasonal
change in temperature and drift in radiation but all the grower needs to
appreciate is that the consequence of his actions will not change. Smaller
canopies will inevitably risk reducing yield to some degree and the risk will
be greatest in determinate varieties like Estima. A grower can therefore
adopt husbandry in respect of all these factors which minimizes the chance
of any of them limiting the achievement of potential yield. This also applies
where planting is interrupted for long periods for again planting into marginal
soil conditions may risk reducing potential yield especially with determinate
varieties. As a first consideration planting should not occur until after the
time which exposes the crop to unacceptable risks of frost damage and
friable but still moist, well fertilized soil conditions prevail and tempera-
tures are rising. This clearly runs contrary to the notion that length of
growing season is the determinant of final yield. In practice some growers
do tend to subordinate these considerations to ensure that each planter
covers the maximum area during the planting period which may stretch
from February to May. This may make economic sense but the inevitable
consequence is that the potential yield of many individual crops will be
reduced. As the capacity of planters on most farms is so much greater than
in the past, delays of some weeks in the onset of planting are unlikely to
lead to planting being incomplete by the time that yield is inevitably lost
because of restrictions on the available growing season, i.e. after mid-May.
Early planting on heavy soils such as the silts is clearly vulnerable to the
effects described and these are likely to be especially critical when
determinate varieties are used. The effect can be exaggerated further on
these soils because almost all crops are grown in wide rows (91 cm apart) in
an effort to minimize clod formation during cultivation. Consequently, the
limited number of leaves have to cover an increasing inter-row distance
before a complete leaf surface is to be achieved. Circumstantial evidence
and grower experience has led to such varieties not being grown on some
silt farms and where they are grown their planting is delayed until after the
planting of less determinate types. This occurs and is justified even when
the intention is to harvest the later planted variety first. This is a good
example of applying intelligence of how the system of yield formation
works. The solution to this specific example of inadequate leaf growth is
not to apply more nitrogen for this does not over-ride the underlying
restriction; however, more nitrogen is frequently adopted in such circum-
stances. This is bound to have a very limited benefit because it cannot
correct the intrinsic limitation, i.e. a fixed number of leaves.
An example from the processing industry illustrates this point. The
maincrop potato area is spread in latitude from the Isle of Wight to
Scotland and yields in the North are generally lower than those in the
Midlands and South. The northern crops are planted later, in moister soils,
where rotations are longer and include more grassland (and the use of
more animal residues or manure). Available N is likely to be plentiful, yet
during much of the 1970s and early 1980s they received increasing applica-
tions of N fertilizer in a vain attempt to increase their yields to the levels
achieved further south. The previous section provided evidence that such
an approach was fundamentally flawed. As the potential yield in most
years would be less, associated with a later spring and shorter growing
season, the target was probably unattainable in any case. The supply of
nitrogen from soil organic matter would be greater than in all arable
rotations prevalent further south so the application of fertilizer N was at
best unnecessary and at worst would delay tuber bulking and further
shorten the available growing season, while the prolongation of the canopy
substantially beyond the intended date of defoliation would create difficul-
ties in harvesting, risk reducing dry-matter content and impair processing
quality. For crops destined for crisping such a scenario could be interpreted
as evidence for the unsuitability of northern latitudes for the production of
high quality crisping potatoes. For one company (Smiths) a severe restric-
tion in the use of animal residues associated with reduced application of
fertilizer N has shown this to be untrue; crisps meeting the highest quality
standards are now regularly produced in North Yorkshire and Co. Durham
where less than ten years ago serious problems existed. No local experi-
mental data were available, indeed none was. necessary, to address the
problem. It was clear that an accurate knowledge of organic and inorganic
forms of applied nitrogen and an estimate of the contribution of soil organic
matter was all that was necessary as even conservative estimates of nitrogen
available from only organic sources amounted to two or three times the
quantities likely to be necessary to sustain a full leaf canopy for the
available growing season. The major initial impediment to improvements
was the grower's intuitive reluctance to accept that reducing something
could maintain yield and enhance quality. We have yet to meet a grower
who did not believe initially that substantial quantities of nitrogen must be
applied to guarantee substantial yield. Careful explanation of the prin-
ciples of crop growth and development combined with plots to demon-
strate the effects of different rates of N quickly overcame these reservations.
Although results were consistent with the understanding of how the system
works they were contrary to growers' expectations and a detailed explana-
tion of the causes was an essential step in their acceptance.
In some cases there was evidence that growers were also seeking to
emulate their southern counterparts in achieving early planting. On the
generally heavier soils and in the wetter springs the inevitable consequence
was cloddy, cold and inhospitable ridges overlying compacted subsoil. In
such circumstances of restricted rooting a grower's belief that ever increas-
ing rates of nitrogen were necessary to aid the apparently inadequate leaf
growth may well have had some validity, but yields were small in relation
to the amounts of nitrogen applied. The fundamental problem was
restricted root penetration, not an overall shortage of N. The key can now
be seen to concentrate on creating the right soil conditions at planting
which encourage root penetration and facilitate economy in nitrogen
application and in some cases improve both yield and quality. For a grower
without irrigation the increased root penetration also increases the likeli-
hood of meeting the water requirement for potential yield.
These examples represent the essence of agronomy: using our know-
ledge of causes to explain and predict effects and tailor husbandry to the
circumstances of the individual crop. Historically agronomy has been
concerned with measuring the effects of the husbandry variables not with
the causes of these effects. This can be appropriate during the initial stages
of agricultural development when knowledge is rudimentary and major
restrictions .abound; it can be relatively quick and reasonably effective.
However, as an industry develops the effectiveness of the approach
diminishes. This is often exacerbated by well-meaning attempts to resolve
contentious points by resorting to more and more experiments that still
deal exclusively with effects. Inevitably these simply restate the problem:
that the effects of most treatments will not repeat either numerically, in
percentage terms, or in some cases even in direction. In the latter case
growers generally feel that one direction must be correct and the other is in
some way wrong. Consequently, to many, agronomy seems an irreconcil-
able muddle, to pure scientists to have no theoretical base and to many
growers no practical use. The use of principles established in the first
section to disentangle the origin of effects should counter this impression.
The principles do not lead one to expect that effects of agronomic
treatments would repeat either numerically, in percentage terms or even in
some circumstances in the same direction. They will be determined by the
amounts of radiation intercepted by the respective crops and are wholly
reconcilable through it.
The examples given above were chosen to illustrate the utility of the
approach in explaining effects observed in practice and were not restricted
to a consideration of the effects of individual variables. This is crucial for
effects that the grower observes and knows to be important can be
explained through the principles. The explanation encompasses effects
resulting from the influence of a number of factors without inevitable
recourse to introducing the concept of interactions between treatments.
Interactions are a statistical reality but much less common than imagined
by non-statisticians and growers. An interaction between two factors
occurs when the effect of one is dependent upon the level of the second
(Table 17.13). It results from the factors not acting in an additive way and
therefore may arise from either an increase in the scale of effect or a
change in the direction of the effect. The latter occurrence is the one which
frequently mystifies farmers and in the absence of any real understanding is
frequently invoked to justify unexpected changes in effects. In most cases
there are no true interactions between the treatments involved and
attempts to use the concept of interactions in this loose way only leads to
confusion. This contributes to the conception that agronomy has no firm
principles and is not useful for the individual grower. This contrasts with
the present approach where changes in the direction of effects do not in
any way conflict with the basic integrity of the principles. Table 17.14
shows an interaction between variety, date of planting and level of nitrogen
in which the relative yields of the two varieties change as planting is
delayed and the level of nitrogen required also changes (Firman, 1987).
This is also consistent with the principles advanced.
As the demands of the industry become increasingly precise it becomes
more important that performance is first analysed in terms of biological
potential and then the economic implications can be assessed. In the past,
biologically inefficient systems have remained economically viable for
considerable periods but from now on biological efficiency in all forms
Table 17.13 The effect of physiological age and

timing of irrigation on number of tubers 4O-(j5
mm in Record (Stalham, 1989)
Timing of onset
of irrigation
Early Late SE
Physiological 0 I 340 279

age (number of 15.2
day degrees >40°C) 330 305 348
Table 17.14 Effect of rate of nitrogen application and date of
planting on tuber yield >40 mm (t ha- l ) in two varieties
(Firman, 1987)
Rate of N application (kg ha- 1)
0 90 180 0 90 180
Date of planting
Estima Pentland Crown
14 March 35.7 46.2 57.9 44.0 57.4 77.9
11 April 28.2 48.3 55.4 42.7 55.5 61.2
12 May 37.7 50.4 51.2 36.3 52.8 50.8
SE 4.13
(including use of N and water) will increasingly determine the profitability

and durability of the system. An example of the survival of a biologically
inefficient but economically viable system was the persistent use of only
physiologically old seed in traditional early areas (Kent and Pembrokeshire)
for the whole harvesting period which restricted yield to below the
potential for much of the harvest period. Nonetheless growers still made
money. In inland areas (Staffs, Shropshire, Suffolk) where the later arrival
of rising temperatures demanded the utmost skill to compete, a more
receptive response to new ideas led to the adoption of a sequence of
varieties and seed ageing treatments that led to increased yields at a
succession of harvests and improved returns. Whereas the areas of earlies
increased in the Midlands and East Anglia there was an eventual shrinkage
of the area grown in the more traditional areas. This would not have
occurred if growers in the traditional early areas had sought and applied
understanding of the principles of early growth, i.e. to arrange a sequence
of seed ages and/or varieties whose leaf growth patterns ensure yields were
close to potential for each date of harvest.
So far the discussion has centred on maximization of the interception of
radiation as the central objective in understanding and explaining hus-
bandry practices. This is justified but must not obscure the possible
contribution of changes in the efficiency of conversion of radiation. Much
less is known of these effects but sufficient reference has been made to
specific examples to illustrate the potential importance. Although com-
plete cessation of growth can be occasioned by severe water shortage, such
extremes are relatively rare and most effects on efficiency are smaller and
relatively short-lived, or both. Their cumulative effect may be large in
relation to total production and this is an area worthy of much greater
study. As we are dealing with the basic processes of plant growth the
necessary studies are specialized and quite distinct from conventional
agricultural experimentation. This should not delay their execution.
17.2.2 Variety
Growers quite rightly have a deep interest in variety as for many outlets
choice is crucial, in some cases obligatory. This interest relates partly to the
belief that varietal selection is a potent means of raising the total and
saleable yield of the crop and partly to the quest for improved disease and
quality characteristics. To these ends considerable resources are expended
annually in testing increasing numbers of varieties both in statutory trials
and those organized by processors, retailers, grower cooperatives and
growers themselves. For yield, the justification is hard to see for two main
reasons. First the belief that varieties with markedly superior growth
patterns leading to real increases in yield are likely to be forthcoming is
hard to sustain. Comparisons of old and new varieties do not demonstrate
any consistent advantage in gross yield for new varieties but show a trend
to decreased dry matter in newer varieties (Table 17.15). This is the most
significant change in varieties over the last twenty years. The introduction
of second earlies such as Estima has increased fresh yields from late June
onwards. This simply reflects the greatly increased water content of these
varieties for when compared on a dry matter basis they are actually lower
yielding than varieties such as Pentland Dell and Record (Fowler, 1988).
The popularity of the late maincrop Cara is essentially based on the same
effect. Its low dry matter content allows very rapid increases in fresh
weights despite late tuber initiation. These improvements are valuable to
the grower but unrelated to the principles discussed so far. However, while
further increase in fresh yield may be achievable through further reduc-
tions in dry matter content, as indicated by the advent of the variety
Nadine, the consumer is already reacting unfavourably against the culinary
consequences of this trend. Desirable culinary qualities seem to depend on
having a minimum dry-matter content so it does not seem likely that
further increases in fresh yield with acceptable quality can be anticipated
via ever lower dry-matter contents. It is hard to conceive how a markedly
Table 17.15 Effect of variety on (a) yields >40 mm (t ha,l) and (b) dry-matter
content on three sites (Allen and O'Brien, unpublished)
Maincrop Second early
Soil King Maris Cara Estima Romano Diana SE
Edward Piper
Introduced (1902) (1963) (1976) (1973) (1978) (1982)
Fen (a) 59.9 69.6 74.0 74.4 55.6 70A 7.73
(b) 21.1 22.6 19.6 19.0 20.0 18.3 0.52
Silt (a) 53.7 46.9 55.6 45.8 50.4 61.6 3.23
(b) 24.6 24.7 20.6 20.0 21.6 20.3 0.43
Clay
loam (a) 47.4 52.0 60.6 43.0 47.7 53.5 2.83
(b) 24.7 23.7 18.8 21.7 21.7 22.5 0.61
superior growth pattern can be expected without direct selection for those
morphological and developmental characteristics that we have shown to be
critical to leaf growth and dry matter production. More rapid rate of leaf
appearance of a greater number of leaves with reasonable persistence may
offer improvement. To our knowledge no such selection is in progress.
Secondly the examples in Section 17.3.1 indicate that varietal per-
formance would change according to the environmental conditions and in
the statutory system of variety testing varieties are compared over a
number of sites throughout the country. One would therefore expect
commercially important and consistent variation in a series of comparisons
of varieties caused by environmental and cultural factors. Indeed growers
firmly believe that this is the case and it is for this reason that the statutory
system uses numerous sites throughout the country. In practice such effects
are not frequently apparent in the short period (2-3 years) over which
comparisons are made and this apparent paradox requires explanation. For
a valid comparison the execution of each experiment must allow the
intrinsic characteristics of varieties to express themselves and be such as to
avoid a particular variety being atypically disadvantaged. This is not
readily achieved for while standardization of practice in respect of many
aspects of husbandry (source and size of seed, fertilizers, pesticides) is
attainable it is not possible to choose a single seed rate or irrigation regime
which is equally appropriate for each variety. Nor can decisions related to
cultivation and planting be guaranteed to be equally suitable for all
varieties from year to year and site to site. Examples of the consequences
of such uncontrollable variation can be illustrated by comparing the likely
performance of the two varieties already extensively referred to, Estima
and Cara at contrasting sites. In the mild south-west England and Wales
(e.g. Starcross or Trefloyne) the longer growing season would in general
favour indeterminate types more than in northern England (e.g. Cockle
Park). Estima's early tuberization and senescence before the intended date
of harvest would usually result in it being out yielded by less determinate
types. However, in late springs and a constricted growing season Estima
may perform better than any other type. In the north the balance would be
reversed; only if the intrinsic early advantage of Estima was nullified by an
unusually early spring or planting before soil conditions are really suitable
would less determinate types produce the highest yields. These 'atypical'
conditions do occur sufficiently frequently to obscure the general trend.
Even within a smaller geographic area where the aerial environment does
not normally change dramatically the water available to the crop from soil
and rainfall may differ sufficiently to create differences between varieties,
e.g. experiments comparing varieties at Gleadthorpe and Terrington
Experimental Husbandry Farms may produce similar yields when the
contribution of soil water and rainfall at Terrington can match the
contribution from soil, irrigation and rain at Gleadthorpe. In dry years,
Terrington may not provide sufficient water for all varieties because of
differences in root growth already described and yields and ranking order
of varieties would be different. It would be rare for emergence to be
synchronous at the two sites of such contrasting soil type even from the
same date of planting. This would lead to differences in the time course of
water demand and the ability of the soil to meet it at the two sites is likely
to differ between varieties leading to differences in yield. These effects
could not be expected to repeat in succeeding years.
The key here is to appreciate the low probability of the weather in two or
three consecutive years being representative of the long term climate at
each site and that husbandry in each year will be unlikely to be equally
suitable for all varieties. The chances of one effect consistently overriding
all others to determine performance in three successive years is extremely
unlikely. Consequently when data from several sites are combined over two
or three years it is rare for the yields of any two varieties to be completely
reversed. When considered in totality with seasons, the discrimination of
varietal performance by environmental conditions that we expect from
principles and growers expect from experience is not forthcoming. If the
objective were to exploit new varieties, either from an industry or grower
co-operation perspective, more effective discrimination could be achieved
from experiments at one site at which environmental conditions were
deliberately changed by date of planting and irrigation management, to
represent the range of conditions found in practice. Careful monitoring of
these experiments would provide the basis for effective interpretation of
the yield data and use agronomic understanding in a predictive sense.
Many growers will intuitively react unfavourably to the suggestion as they
believe testing varieties in many locations has paramount importance. This
has relevance primarily in a broader sense in ensuring that any defects,
unrelated to yield are detected. The approach can only be effective for
yield if the data are interpreted on the basis of agronomic understanding
and not at face value; i.e. variety A is superior to variety B because this
year it outyielded it.
Being realistic the production of a variety of consistently superior
performance will not require any extensive scheme for its merits to be
obvious. The arrival of Pentland Crown so obviously marked a step
forward in gross yield that no subtle testing was required to reveal its
merits. No subsequent concerns about its quality should diminish the
contribution of this variety.
There are of course many characters which have commercial value and
for a number of these breeders have made great improvements - nematode
resistance, resistance to fungal diseases, e.g. common scab, virus resistance
- but commercially important cultivars still exist that have weaknesses in
these areas. There is therefore every prospect of improved varieties
coming forward which are superior in one of these specific aspects but of
similar yield to existing cultivars. In these cases the objective of varietal
testing should be the validation of the commerical value of the improve-
ment once a comparability of yield has been established. Such testing
would be quite specific and may well require substantial areas (hectares) to
be grown rather than small plots. This would be especially true for
improvement in culinary characters likely to change over long-term storage
for which considerable tonnages may need to be stored.
Whatever variety a grower chooses and whatever information upon
which he bases that choice the way in which a variety is grown will have a
considerable influence on the yield obtained. The responses to soil and
environmental factors in terms of leaf growth have been much emphasized
but they are the basis on which yield is dependent and what is acceptable or
appropriate for one variety may be quite inappropriate for another.
Further, the effects of physiological age (Chapter 6), seed rate (Chapter 7),
and timing of harvests (Scott and Wilcockson, 1978) have major influences
on the saleable yield obtained and the quality of the produce. Thus, the
practical exploitation of the potential of any variety requires understanding
of growth and development and a willingness to change husbandry in
relation to a variety'S needs to a much greater extent than was previously
thought necessary.
17.2.3 Physiological age

In Chapter 6 the influence of physiological age on the growth and
development of the crop was explained on the basis of the principles being
used in this chapter. The ageing of seed advances emergence, tuber
initiation and frequently senescence. In principle the advancement of early
growth is highly desirable and ageing is an extremely important potential
influence on the production of all crops. Large responses are more likely
the shorter the growing season whether of earlies or maincrops; in very
long seasons seed of minimal age may out yield older seed. As the effects
can reverse over the harvesting period much suspicion of the utility of the
concept still remains. This is accentuated because the practical difficulties
of planting sprouted seed slow the planting operation and it seems that for
main crop production fewer growers now sprout than previously. This is
unfortunate for the potential benefits are being sacrificed. A rational
approach would be for growers to attempt to establish the contribution that
ageing could make to the growth pattern of their particular crops on the
basis of principles outlined in this chapter. The conclusion would be
controlled sprouting would have a place in many production systems and
the demand for planters capable of handling sprouted seed would stimulate
designers to overcome the mechanical difficulties.
17.2.4 Date of planting

Reference has already been made to some of the effects of this factor in
relation to growth and ultimate yield. The inter-relations with variety and
physiological age are clearly of great practical significance. There can be no
doubt that growers are increasingly questioning the advisability of planting
at the earliest possible opportunity because of the recognition that pursuit
of this objective frequently leads to poor soil conditions in the ridge,
deeper compaction into the subsoil and reduced leaf growth leading to
reduced yield. Subsoil compaction can occur even on light sandy soils if
planting is practised when soils are too wet. This has been seen on many
sandy soils in Suffolk where planting of earlies in late January and
February resulted in meagre canopies and poor yields and, unknowingly,
subsoil compaction. Where this occurs it also restricts root penetration and
water uptake. The compaction can be removed by subsoiling and applica-
tion of principles described here would ensure that it did not recur.
The risks associated with planting as early as possible into unfavourable
soil conditions are increased where growers use polythene in an attempt to
advance early leaf cover and increase yield. There is a belief that where
plastic covering is used it is desirable to plant even earlier than usual so that
benefits can be maximized. Observations and measurements of coverings
made from December onwards show that effects on soil and air tempera-
ture, and hence emergence and leaf growth are minimal until the end of
February when incident radiation becomes sufficiently intense to cause a
temperature increase. It would seem better to wait until soil conditions are
fit because the longer days and higher irradiance will increase temperatures
under the polythene more than earlier thereby hastening emergence and
ensuring a rapid production of a full leaf cover. If the crop can meet its
water requirement and is not checked after removal of the polythene it
would seem that the objective should be to leave the polythene in place
until as close to complete canopy as possible to gain maximum benefit from
the increased temperatures. However, Jenkins and Gillison (1991) reported
that removal of sheets covering only two rows should occur at no more
than 20% leaf cover if reductions in yield are to be avoided. It is not
immediately obvious why the timing of removal should be so critical unless
related to physical restrictions on leaf growth. Where sheets covering many
rows are used it is possible to delay removal until close to complete leaf
cover. It proved possible temporarily to remove the sheets to allow irriga-
tion to be applied and Table 17.16 shows the yields obtained in Kent when
planting was delayed until the soil was fit and wide plastic sheets were used..
The yields of Desiree harvested in late June were extremely high and of
pre-packing quality representing c. four weeks advance over crops planted
in the open. These comparisons highlighted the increased benefits from
ageing of seed the earlier the harvest.
17.2.5 Nitrogen
There can be no doubt that the use of increasing amounts of N fertilizers
has made a major contribution to the increasing yields of the national crop
especially in the early years of their use. Indeed many growers still see a
very direct relationship between the two as illustrated by the continuing
Table 17 .16 Effect of date ofplanting
and physiological age on tuber yield
(t ha- l ) in Desiree (Kent, 1990)
(Allen and O'Brien, unpublished)
Physiological age
Date of planting 0 300 600
Yield >30 mm on 30 May

20 March 7.9 13.6 12.6
6 April 0.3 1.9 2.9
SE = 0.67
Yield >40 mm on 21 June
20 March 32.6 32.9 33.7
6 April 12.2 18.6 22.1
24 April 0.9 5.0 7.8
SE = 1.16
Yield >40 mm on 10 July
20 March 46.9 45.6 43.7
6 April 26.0 34.0 38.9
24 April 12.0 25.4 19.8
SE = 1.23
increase in fertilizer use and national yields (Fig. 17.18). This view was
probably strengthened by the 'Blueprint' exercise which implied that large
increases in fertilizers and other inputs were necessary for increased yields
(Evans, 1975; Evans and Hield, 1981; Nield, 1977). The evidence was not
clear cut even at the time and with increasing concern over the fate of unused
N it is now essential that any applied N is justified. Unless increasing the
amount of N applied leads to the interception of more radiation there will
be no increase in yield (Fig. 17.19). On the basis of the effects of leaf area
on the interception of radiation, which is ofa distinctly diminishing
response form, there is no reason to expect continuing increases in yield
with ever increasing rates of N. Moreover, increasing use of N is known to
have undesirable effects delaying emergence (Firman, 1987), tuber initiation
(Firman, 1987), increasing stem length which may cause lodging (Ifenkwe,
1975) and by increasing the size of the canopy it may create an increased
risk of blight risk from the moister microclimate. None of these responses
can contribute to increased yields. In most experiments the simple aim has
been to plot yields against N applied in order to estimate an optimum rate.
The optima inevitably vary from site to site and year to year as the N
uptake of the crop (a combination of soil mineralization of N, use of N
fertilizer, soil water status and the functioning of the root system) and
incident radiation also varies. With such variation only broad categories of
optima can be drawn, usually by soil type and previous cropping (MAFF,
1984) and represent gross averages at best. From the characteristics of
indeterminate and determinate varieties one would expect the former to
need less N than the latter and Table 17.17 shows the distribution of
optima for a series of experiments in East Anglia. Variation still remains
although division by variety as well as soil type is possible.
The significance of this remaining variation must be put in context. In
most N response curves the final 50% of the optimum rate produces only
10-15% of the final yield (Fig. 17.20). Absolute precision in determining
the optima therefore, would not lead to great advances in yield but
increased precision is an essential component of a policy to reduce the risk
of pollution of groundwaters. The understanding of the dynamics of soil N,
50.-----------------------------r
40
300
"co;"
-as ~t30
.!!!.<:
>--
240
200
f~~"
iif[iD
~ 20 160 "';'zs.
120
10 80
O~~~~~~~~~~~~~~
1955 1965 1975 1985 1995
Year
Figure 17.18 National average potato yields and nitrogen fertilizer application,
1969-1989 (Potato Marketing Board and Fertilizer Manufacturer's Association)..
N rate TuberDM
1000
(kg ha") yield (t ha")
_300 12.2
~
E 750 -240 12.5
~ -180 12.3
i
a 500
_120 12.5
~ -60 11.5
.~ ~ 0 10.0
1: 250
C>
::J S.E. 0.45
0
May June July
Month
Figure 17.19 Effect of rate of nitrogen application on cumulative light intercep-
tion and final tuber dry matter yield of Estima at CUF (Fowler, 1990, unpublished).
Table 17.17 Distribution of optimum rate of nitrogen fertilizer
application for three varieties (CUPGRA, 1985-90)
Variety Nitrogen (kg ha- 1)
0 0-50 50-100 100-150 150-200 200-250

Estima 1* 0 2 8 11 4
Maris Piper 5 (4*) 1 1 1 3 1
Cara 2 (1 *) 0 5 1 1 0
• Peat soils
, its uptake by crops and use in the generation and maintenance of the leaf
surface are crucial to progress and here information is limited. Crop
nitrogen uptake usually proceeds linearly for much of the life of the crop
(Fig. 17.21) but the rates and hence the total uptake vary from year to year
(Table 17.18). Yields cannot be closely related to N uptake (Fig. 17.22)
and models using N uptake as a driving force for dry-matter accumulation
(Greenwood et al., 1985) are imprecise once moved from the data of their
creation (Fig. 17.23). Percentage N in foliage decreases as growth proceeds
and the rate of decline can be moderated by increasing nitrogen uptake.
The line derived from Greenwood's equation suggests that there are
critical minimum values which must be maintained if maximum dry matter
production is to be achieved. However, the data from the other experi-
ments show a wide range of dry matter production for any concentration of
nitrogen in the foliage. The proportion of applied N which is recovered is
also variable (Table 17.19) and the true source of N found in harvested dry-
matter is still not clear. Recent data from Cambridge University Farm
show large fluctuations in the quantities of N apparently available from soil
supply throughout the season (Table 17.20). This occurs both under crops
and in unplanted areas, presumably because N moves into and out of non-
available forms of the soil biomass. Such unpredictable changes render
attempts to establish the sources of N uptake by balance sheets of fertilized
and unfertilized crops potentially misleading.
More knowledge is urgently required on the way the potato crop takes
up N and uses it in growth and development. It is known that N excess to
current growth can be temporarily stored (Millard and Marshall, 1986) but
the relationships between leaf growth and development and N uptake have
not been generally sought. Some relationships are becoming clear. A
physiological definition of nitrogen requirement must begin with a descrip-
tion of the way applied N affects the crop's leaf area. Unpublished data of
J. N. Bean show a close relationship between leaf area and kg N ha- 1 in the
crop for a range of planting dates (Fig. 17.24). Similarly in winter wheat,
Sylvester-Bradley et al. (1990) have shown that leaf area indices were
directly related to N uptake and that yield response could be analysed in
these terms to provide a simple summary of the diminishing responses of
grain yield and grain protein concentration to applied N.
70 (a)
60
50
40
30
20
10
0
0 50 100 150 200 250 300 350
70 (b)
60
~
aI
50
~
~ 40
:2
Q)
':;' 30
Q;
.0
:::l
I-
20
10
0
0 50 100 150 200 250 300 350
70 (c)
60
50
40
30
20
10
0
0 50 100 150 200 250 300 350
Rate of N application (kg ha· 1 )
Figure 17.20 Response of tuber yield to rate of applied N in Estima grown in East
Anglia on (a) sandy loam, (b) loamy sand and (c) sand soil (Fowler, unpublished
data).
250
::- 200
'as
or:
l
~ 150
~
:::l
c:
~ 100
g
·c
~ 50
01972
A 1973
01974
0+-----------.----------,-----------.----------.
May June July August
Figure 17.21 Total nitrogen uptake in Desiree crops receiving similar fertilizer
dressings and planted at 37 440 tubers ha-1 and on similar dates in three years (from
Ifenkwe, 1975).
Table 17.18 Nitrogen uptake by a range of crops

Source Variety Rate of application Tuber Maximum N
of N (kg ha-1) yield uptake
(t ha- 1) (kg ha- 1)
Bean Maris Piper 0 47.6 131
(unpublished) 200 89.4 321
400 88.3 427
O'Brien Maris Piper 0 69.6 262
200 73.9 341
Fowler Pentland Squire 0 46.0 147
220 58.5 288
For potatoes it is known that where yield is increased by lengthening the

growing season this is frequently associated with greater uptake of N. This
is because the sustained leaf cover which allows the longer growing season
is the result of increased branching from increased N uptake.
The plethora of experiments aiming to establish N optima convince
many that 'we must know everything' about N. The foregoing shows that
this is far from the truth with previous work concentrating once again on
effects and with much less effort aimed at unravelling the causes of the
variation. Progress now requires fewer much more detailed experiments
which answer the questions raised above relating to soil supply and crop
20
•
• •
x
x x
';"co 15 x
• x
..r::.
•
• • [J
[J •
o [J •
o •
o
o 60 120 180 240 300 360 420

Total uptake of nitrogen (kg ha- 1 )
Figure 17.22 Relationship between final tuber dry matter yield and total uptake of
nitrogen in Maris Piper for different rates of N fertilizer at four sites over six years
(unpublished data). Key: 0, Treftoyne 1972; 0 Treftoyne 1973; !1, Treftoyne 1974;
x, Cambridge University Farm 1982; e, Treftoyne 1983;., Cambridge University
Farm 1984; .. , Prickwillow, Cambridgeshire 1986; T Manea, Cambridgeshire 1986.
Table 17.19 Efficiency of N recovery at the optimum rate of

application for several varieties grown on a range of soil types
Variety Soil type Optimum rate % recovery
Estima Loamy sand 180 30
Estima Loamy sand 200 68
Estima Sandy loam 130 55
Estima Sand 100 40
Maris Piper Sandy clay loam 200 97
Maris Piper Sandy loam 200 61
Pentland Squire Sandy loam 110 83
Pentland Crown Sandy loam 180 62
Cara Sandy silt loam 120 35
Cara Clay silt loam 60 12
(Uptake at optimum - uptake at No)
Efficiency = Rate at optimum
x 100
4 o
o
o
o
(]) 3
.~
S
.S:
c: o
(])
g
Ol
'c
#. 2
o
o o
tl
tl
tltl
x x
x
x x
x
o 5 10 15 20
Total plant dry weight (t ha· 1)
Figuie 17.23 Relationship between nitrogen content of plants and total dry matter
yield in Maris Piper (unpublished data). Key: Cambridge University Farm, 1982,
0; 1984, 6; Trefloyne, Tenby, 1983, 0; Marsham, Norfolk, 1987, x.
uptake and the fate of N. For the grower there are two essentially practical
points to consider in the interim. First, some recommended optima for N
(Table 17.21) are higher than the highest level used in most published
experiments. Secondly, if the strategy outlined in the following section is
used, growers do not need to change their rates of application in relation
to irrigation. As drainage of water through the profile should only occur
after excessive rainfall early in the season, there should not be any leaching
(and hence unavailability) of N through the use of irrigation. The similarity
of optima for a wide range of water regimes and yields is shown in Fig.
17.25. Using extra N as insurance cannot of itself reduce the risk and
Table 17.20 Available soil N from 0 to 90 cm (kg
ha-1) in three situatiollsat Cambridge University
Farm, 1990
Date Covered Uncovered Uncovered
bare soil crops receiving crop receiving
irrigation but irrigation
no nitrogen and 120 kg N ha- 1
21 Mar 115 150 374
2 Apr 129 139 455
8 May 162 182 574
4 Jun 176 133 349
3 Jul 365 153 228
28 Aug 270 63 166
5 x
4
Q
o
x
Q)
-0 Date of planting
.!: 3
co 03 March
Q) 05 April
n; 67 May
1ii 2 o x 10 June
Q)
...J
0
x
0
0
20 60 100 140 180 220
Total nitrogen uptake (kg ha- 1)
Figure 17.24 Relationship between leaf area index and total nitrogen uptake in
Desiree for four dates of planting in 1980 (Bean, unpublished).
Table 17.21 Published N recommendation (kg ha-1) for

maincrop potatoes grown on different soil types with
different N indices (MAFF, 1984)
Soil type
Mineral Organic Humus, moss and warp
N index 0 220 130 180
1 160 90 130
2 100 50 80
50
23
40
10
6
• • • • ..
o100
~------~------~--------~----~
150 200 250 300
Soil + fertilizer nitrogen (kg ha-')
Figure 17.25 Influence of total available (soil and fertilizer) nitrogen and seasonal
amount of water applied on total yield of Russet Burbank potatoes (Ojala et at., 1990).
incontrovertibly ensures the movement of more N if leaching occurs. There

seems no justification for the practice.
17.2,,6 Irrigation
The view that potatoes are shallow rooting and therefore highly susceptible
to water stress has greatly influenced attitudes to the application of water in
practice. There are two basic assumptions in current scheduling: a fixed
relatively shallow rooting depth leading to small limiting or trigger deficits
and the need to return the soil to close to field capacity over much of the
season (Bailey and Minhinick, 1989). Both are difficult to reconcile with
the evidence already considered. In any scheduling system demand is
calculated as evapo-transpiration (Penman, 1948) corrected for leaf cover.
This can be met from the soil supplies (replenished by rainfall) until a
certain 'trigger' soil moisture deficit is approached when irrigation is
applied unless rainfall is expected. The soil's supply is determined by its
water holding capacity and the rooting depth of the crop. The latter is
usually taken as 50-70 cm, considerably less than the rooting depth usually
observed in uncompacted soil during July. Sandy soils have low water hold-
ing capacity and therefore smaller limiting soil moisture deficits than heavier
soils. The application of water in practice is usually 20-25 mm and with the
smaller trigger deficits the soil will be returned to closer to field capacity on
sandy soils than heavier ones. It has become accepted by many that potatoes
produce their potential yield only if grown with small limiting deficits for
much of the season. This inevitably incurs the risk of unexpected rainfall
creating leaching of N. Moreover current knowledge of root growth
suggests this may well be unnecessary as the root system is capable of
extracting more water from deeper in the profile than previously accepted.
It seems that many potato crops may receive too much water and that
others would probably do so if only the mechanics of the system and the
energy of the operators allowed. Published values show a wide range of
responses of yield per mm of water applied as irrigation (Table 17.22). A
similar range has also been shown for sugar beet (Dunham, 1988). From
first principles water should be evaporated from the leaf surface at a rate
determined by incident radiation. Thus, for any environment there should
be a constant value for dry-matter production per mm water transpired
(efficiency) which for north-west Europe would be similar in all years;
published values suggest this is c. 5 g m-2 mm- 1 which equates in fresh
weight to c. 0.25 t ha- 1 mm- 1 . The exceptions are extreme years like 1976
when because of an extremely dry atmosphere the value decreased (Day et
al., 1978). However, as Table 17.22 shows, few crops approach even half
this efficiency value in relation to water applied through irrigation and
these were amongst the 13 crops grown in dry years when responses were
great. In the other years there are two clear groups: first, four crops
receiving only 50 mm or less with no yield response which probably reflects
a policy of starting irrigation around tuber initiation aimed to control
common scab not to increase yield. In addition, there will have been
occasions when there was no return because trigger deficits were being
approached, then the weather stayed dull and crops did not suffer much
water stress, and others when irrigation was applied but heavy rain soon
followed. These conditions inevitably lead to a reduction in response per
mm of water applied. However, it is difficult to see how these circumstances
can account for inefficiency in the second group of 13 crops whose yield
responses ranged from 0 to 8.6 t ha- 1 from applications of water from
50 to 150 mm and whose efficiency per mm of water applied was substantially
below those of the first grouping. From knowledge of the errors in such
experiments it is likely that only two of these responses would be
statistically significant and the conclusion must be that in half the years
(15 out of 30) there was no response to irrigation. Large responses to
irrigation occur less frequently than suggested if the values in Table 17.22
are taken at face value. It is surprising that few reports exist in which the
response to irrigation is assessed in relation to radiation incident and
the use of water after the application. For much of the season in many
Table 17.22 Yield response and efficiency of water
use from irrigation of maincrop potatoes at
Gleadthorpe E.H.F. (Bailey, 1990)
Year Irrigation Yield Efficiency
applied (mm) response (t ha- I ) (t mm- I )
1958 51 -1.5
1959 203 25.6 0.126
1960 150 7.3 0.049
1961 142 3.8 0.028
1962 132 3.3 0.025
1963 43 0.8 0.019
1964 86 2.8 0.033
1965 41 -0.1
1966 99 1.3 0.013
1967 152 16.1 0.106
1968 51 0.5 0.001
1969 102 14.8 0.145
1970 226 24.8 0.110
1971 84 3.5 0.042
1972 99 12.6 0.127
1973 64 0.8 0.013
1974 79 -1.3
1975 no data no data no data
1976 272 39.7 0.146
1977 111 23.0 0.207
1978 91 3.9 0.043
1979 120 18.9 0.158
1980 66 -0.8
1981 113 25.3 0.224
1982 115 8.6 0.075
1983 165 23.0 0.139
1984 137 23.9 0.174
1985 143 13.1 0.092
1986 154 17.9 0.116
1987 83 3.8 0.046
1988 142 2.9 0.020
scheduling systems the application of water is at a constant trigger deficit at

intervals determined solely by evapo-transpiration. This ignores extra
water which may be accessed in the interim as root penetration continues.
Consequently if dull dry weather follows an irrigation the consumption of
the applied water results in the trigger deficit being reached again but for a
crop whose soil supplies may sustain several more days of growth as roots
penetrate deeper into the profile. Thus, as long as root penetration
continues the trigger deficits should increase. This is particularly relevant
for early growers, who usually have irrigation facilities, for published
responses in early crops are small (Bailey, 1990). The value of thes.e crops
is high and small responses may be economic but the results suggest that
worthwhile responses to irrigation are uncommon. Early crops are grown
with a wide range of seed ages, varieties and on contrasting soils so it is
quite possible that their access to soil water varies and greater under-
standing is needed before consistent responses to irrigation will be achieved.
If irrigation does not begin until root penetration has ceased (this inevit-
ably means a large trigger deficit) continuing growth of the crop in the
absence of rainfall will be almost entirely dependent on water from
irrigation. If, for practical reasons, each application is a constant amount the
interval between applications will be determined entirely and justifi.ably by
evapo-transpiration. On the basis of the foregoing it would be expected
that irrigation would be related to depth of root penetration and would
be triggered by increasing deficits as the season progressed. In pra.ctice, an
application of a similar amount of water (25 mm) at each irrigation would
therefore leave an increasing soil moisture deficit as growth proceeded.
The possibility of scheduling irrigation in relation to measured rooting
depth and available water (held at less than 0.6 bar) per 20 cm horizon has
been tested in practice and the soil moisture deficits for such an approach
and those of the most widely used conventional system are shown in Fig.
17.26. The data are taken from a field-scale comparison of the two systems
in a crop of cv. Record grown for the Pepsico-Foods International Demon-
stration in the hot, dry year of 1990. The conventional approach maintained
a smaller deficit and 38 mm more water was applied than in the Cambridge
University Farm system for identical yield and quality. Figure 17.26 also
shows the soil moisture deficit created by removal of all water held at
tensions less than 0.6 bar from the measured rooting depth allowable. The
difference between this and the deficit allowed by the Cambridge University
Farm approach is large and the real targets for irrigation research should
be to establish how close commercial scheduling can be brought to this
deficit and whether for all varieties and horizons 0.6 bar is the limiting
tension. Progress in this direction might well ensure full yield for less water
and hence less risk of pollution.
The timing of onset of irrigation is determined by the need to control
common scab as well as to maintain growth of the canopy. As freedom
from common scab is an absolute prerequisite for markets that require the
highest quality tubers, and a number of popular varieties are susceptible, it
is important to distinguish water needed for this purpose from that required
to maintain growth. Lapwood (1968) established that the presence of free
water in the region of the stolon tips for up to 6 weeks from the onset of
tuber initiation would prevent the disease in susceptible varieties. Figure
17.16 indicates that it is unlikely that a soil moisture deficit sufficient to
limit growth would be created some time after tuber initiation of most
varieties. Thus, the amounts of water needed for control of common scab
must be established for individual varieties and individual soil types. The
Julian day
120 140 160 180 200 220 240
I -20
·0
~
"0
~ -40
~
.~
~ -60
-80
Figure 17.26 Effect of irrigation scheduling on weekly soil moisture deficit in
Record (Stalham, 1990, unpublished). +,ADAS; 0, CUF; ., allowable.
amounts involved may be sufficiently trivial to achieve the disease control

but inconsequential in relation to maintaining growth while minimizing the
risk of leaching.
There may well be real differences between varieties in their extraction
of water and in their response to decreasing water availability; there is an
urgent need for comprehensive study which links root and leaf growth,
water use and yield formation so that the limiting tensions can be
established throughout the life of individual varieties.
All this discussion associated with efficient use of irrigation should not
minimize the importance of irrigation to the maintenance of a profitable
industry. The quest for continued increase in yields, which will be essential
to maintain profitability, inevitably requires the transpiration of more
water per hectare. For all growers, particularly those without irrigation, a
desirable aim is to maximize recovery of soil water. Consideration of
current knowledge indicates that a potato industry with increasing yields
and still heavily dependent on irrigation need be no more, perhaps less of a
threat to the environment than conventional wisdom has it. Because the
effect of irrigation on yield and quality and hence value is so large, potato
growers are always going to ensure, and rightly so, that no real opportunity
for benefit is missed. In some crops, e.g. sugar beet, it has been suggested
that less than full irrigation may be the most economic (Dunham, 1988) but
for the potatoes with their high price elasticity this is unlikely ever to be so.
The evidence presented here suggests that less water could be applied
without missing the real opportunities.
17.2.7 Harvesting and storage practices
The principles of harvesting and storage are clearly different from those
associated with crop growth, development and yield formation and are
dealt with in separate chapters in this book. They tend to be considered
separately and few experiments have been reported in which the relation-
ship between growth in the field, suitability for harvesting and storage
performance have been inter-related. This is surprising for most growers
believe the interdependence to be great. The published evidence does not
support this view but it seems that the implications have been largely
ignored. Wilcockson et al. (1985) reported the results of a national research
programme funded by the Potato Marketing Board which studied this
relationship. They found that it was not necessary for crops to be visibly
senescing for there to be minimal weight loss in store, nor was defoliation
necessary for efficient storage, even for crops harvested in full leaf growth.
As early harvesting has been shown to be advantageous by others (Potato
Marketing Board, 1979) and is known to minimize the transfer of disease
inoculum from seed to daughter tubers (Firman, 1990) one would have
thought that these findings would have influenced practice. They offer
flexibility in timing of haulm death as well as lower costs. They have been
largely ignored except where applied by E.J. Allen working directly with a
group of growers supplying crisping potatoes to Smiths. In the belief that
defoliation was essential these growers in the North of England usually
defoliated their crops 14-21 days in advance of early dates of harvesting
(mid-September) and lost yield thereby. Thus, the opportunity to leave the
crop to grow for 2-3 extra weeks is often welcome. Long-term storage and
processing quality are not affected and from small beginnings of a few
hectares in 1987, over 5000 tonnes entered storage in 1990 without prior
defoliation. This is an example of the successful application of research
findings in the industry, but it is still only applied to a small fraction of the
crop stored for processing.
The research programme of Wilcockson et al. (1985) used treatments
chosen in consultation with the industry to create large differences in
storage potential. These did not materialize nor did they in the work of
Hogge (1989), Stalham (1989) and Colenso (1989) who examined the
relationships between field growth, harvesting practices and storage and
processing performance. Treatments which were expected to ruin fry
colour ·failed to do so. This led Hogge et al. (1991) to conclude that it is
difficult to demonstrate a large influence of the state of the crop at harvest
on performance in storage either in weight loss or processing performance
even where experimental treatments ranged much wider than ever ex-
perienced in practice. As modern stores provide stable environmental
conditions throughout the period it is perhaps not surprising that once
tubers have settled into the stable conditions few treatment differences
persist. The industry's perception may well be influenced by the problems
of an era already past when serious losses occurred due to deficiencies in
the stores. The modern store is reliable and provides a uniform environ-
ment so that over a much longer period than before tubers change only
slowly in their culinary and processing potential. Some differences do
persist and there is a need to study them: e.g. tubers seem as likely to
improve as deteriorate in processing potential during storage, a view
contrary to the industry's perception.
The real problems in harvesting remain the effects of the mechanical
operation on the tuber that create damage in all its forms and remove much
value from the crop as a whole. This has never been investigated in relation
to the effect of the forces involved in the harvesting operation on the
physiology of the tuber. The simple 'rule of thumb' that visual senescence
can be taken as a sign that skin thickening is occurring, maturity is
approaching and hence suitability for harvesting improving, was shown not
to apply by Wilcockson et al. (1985). They were driven to state 'complete
avoidance of the term maturity would probably help all future discussion of
factors influencing the suitability of tubers for storage'. There is still a need
to try to relate the changes in the tuber to the growth and development of
the canopy and to investigate how these relate to damage and wound
healing during the harvesting, loading and storage phase. The harvesting of
tubers from crops with substantial leaf canopies and weak skins inevitably
leads, at the very least, to some skin removal and although the tubers will
probably store with little yield loss the tubers may not achieve the visual
appearance required for the highest value unless a new skin of similar
appearance to the original skin can be grown quickly. Observations over
many years have shown that crops with substantial canopies and firm skins
do exist and the physiological conditions which control skin growth and
regeneration have not been defined. If this could be provided, harvesting
and storage environments could be chosen to ensure that each crop reaches
storage with minimum damage and may then be expected to store in a
predictable way.
There is massive potential return from improving our knowledge of the
principles governing the tuber's suitability for harvesting, for damage costs
many millions annually. Their elucidation requires an integrated study
involving physiologists, agronomists and engineers and continuing separa-
tion of these component disciplines will severely reduce the value of any
further research. Such research which must be on a large scale, will be
difficult to organize and hence costly but without it the industry increases
the risks that it will fail to improve the quality of its produce, as it must if it
is to continue successfully.
Relevance of agronomic understanding in the potato industry 873
17.3 RELEVANCE OF AGRONOMIC UNDERSTANDING TO
IMPROVEMENTS IN THE POTATO INDUSTRY
17.3.1 Changes in the industry since 1976

Over the period since 1976 potato production has become concentrated in
far fewer hands who are able to meet demand from fewer hectares (Table
17.23). There has been some regional reorganization as the area of
potatoes produced in Scotland has fallen. These structural changes in the
industry have occurred as the market has changed. The proportion of the
crop destined for processing and pre-packing has continued to increase and
in both sectors the target specification has become tighter over a broader
range of characters than before. Consequently, potato growers are now
faced by a diverse, but increasingly sophisticated market which can only
be supplied if crops are grown specifically for a particular outlet. This
represents a major shift in emphasis in the planning of production.
Table 17.23 Number of registered producers over time (PMB, 1990)

Year Number Year Number Year Number Year Number
1960 76825 1970 43346 1980 30225 1990 17745
1961 74933 1971 42756 1981 28760
1962 70056 1972 40756 1982 27498
1963 66380 1973 38753 1983 26499
1964 60940 1974 36872 1984 25480
1965 57725 1975 35478 1985 24948
1966 54700 1976 34938 1986 22565
1967 50310 1977 35286 1987 21439
1968 48239 1978 33387 1988 20155
1969 45130 1979 32149 1989 19266
In order to meet these changing requirements, husbandry methods have

changed considerably. The adoption of stone and clod separation, mechanical
harvesting, more sophisticated irrigation systems and controlled-
environment storage has been widespread. All these result from the
activities of the engineers and it is hard to escape the conclusion that the
major changes in mechanization have been driven by the need to meet
the physical demands of the system and the logistics of the practical
operations with less consideration for the demands of the crop itself. In
some aspects the machinery has been developed while the bases for its
efficient operation or even its justification are still unclear: e.g. the drawing
of ridges in autumn, timing of stone separation in relation to subsequent
soil conditions, windrowing systems for harvesting. When the agronomic
questions are resolved it may be that the ideal specification for the
machines will change.
In several aspects of husbandry it has been shown in this and other
chapters that growers still adopt practices which are not easily reconciled
with the current state of knowledge and often seem reluctant to develop
techniques whose principles seem firmly established but whose practical
utility requires further work. The slow uptake of the clear benefits derived
from the use of smaller seed, and slow recognition of the flexibility offered
by the discovery that defoliation a fixed interval before harvesting is not an
essential prerequisite for successful storage, are good examples. The
widespread reluctance even to consider reductions in nitrogen and water
use even when restrictions are already in place in certain locations is
indicative of a similar mistrust. At the other end of the spectrum growers
have been prepared to introduce a wide range of foliar sprays containing
many substances. For many of those there is little evidence to justify their
use. For a yield response to be obtained they would have to improve eit~er
the interception or the efficiency of conversion of radiation. Since leaf
cover is usually adequate when they are used, the route to any effect must
be either to increase the efficiency with which intercepted radiation is
converted to biomass, delay senescence or affect the partitioning of
assimilates in favour of the tubers. The general evidence from experi-
mental work is that these are not readily achieved. Thus, while there is no
doubt that changes and general improvements in husbandry have occurred
over the last 15 years, opportunities do seem to have been missed. This
may be a consequence of a major change during the period, the accelerat-
ing decrease in the research and development effort which has reduced
both the extent of research findings and their dissemination to growers but
the failure to adopt new techniques and innovate generally may also reflect
deficiencies in methods of transmission of information. This is a matter of
particular concern for the future as additional knowledge is only of
practical value if it can be utilized in improved systems of production.
Many of the developments in machinery, marketing (e.g. bakers), new
crops (out-of-season earlies) and store loading can be attributed primarily
to enterprising farmers not to the efforts of applied researchers.
17.3.2 Problems and future challenges

The very high consumption of potatoes in the UK necessarily means that at
best only small increases can be anticipated in the future and at worst
maintenance at current levels may be difficult. It is inevitable that existing
markets will tighten their specifications over a broader range of characters
and success in maintaining consumption will depend on continuing
improvement in overall quality. There are opportunities to develop new
markets and products which may be relatively small in relation to the
overall market. These products may be quite distinct from current pro-
ducts. For example, the production of small (20-40 mm) tubers as early
(new) potatoes throughout the year, for which there is a clear demand, will
require new methods of planting, harvesting and storage in addition to the
adoption of agronomy appropriate to growing seasons and harvest periods
quite distinct from existing ones. There are opportunities to develop new
markets overseas particularly for seed and if these are to be taken, seed
must be produced which meets the buyers' requirements. It is not simply a
matter of producing more seed and selling it to new customers overseas.
These new markets will have precise requirements for varieties, sizes,
diseases status and timing of end of dormancy which are different from
those currently accepted in the UK. The tightening of specifications
domestically and the need to supply a wider range of customers overseas to
precise and different specifications to those prevailing in the UK will
demand agronomic innovation for success. For the future, scientific
agronomy holds the keys to continuing progress for the whole of the potato
industry. It is therefore essential that mechanisms for acquiring and
transmitting agronomic knowledge are effective.
17.3.3 Meeting the challenges
Existing knowledge
Throughout this book there have been many examples that indicate that
growers do not use existing knowledge and indeed may not be aware of its
value. It has been suggested that the benefits from getting the agronomy
right will continue to increase. Even without any further research the
application of current knowledge could effect considerable improvement.
There are lessons to be learned. It is just as important to consider how best
to transmit information to the grower as to continue to aim for new
knowledge. We believe that the principles outlined earlier in this chapter
offer the way to apply existing knowledge in different contexts. This is
especially important as there is a general tendency to respond to new
products, markets or problems through the sanctioning of more experi-
ments that may do no more than duplicate existing knowledge. As the
industry faces markedly reduced funding for research and development,
more effort should be expended in obtaining the full value of current
knowledge and new programmes directed specifically where knowledge is
really lacking. In these, priority should be given to understanding causes
and not simply monitoring effects.
Acquisition of new knowledge

In this chapter many areas of limited knowledge have been identified
where greater understanding would contribute to continuing improve-
ments in the industry. The key is to appreciate that the ability to do this
derives from a long standing (25 years) study of how the potato crop grows
and develops, not from a disjointed series of short-term investigations
attempting to resolve the contemporary problems considered to be the
most pressing of the day. An understanding of growth and development
provides the basis for progress in matters as diverse as the formation of
yield, skinning of tubers during harvesting, ingress of disease in tubers, the
risk of N leaching and their inter-relationships. As future research is to be
funded directly by farmers it is crucial to progress that those charged with
using farmers' money are made aware that progress can be made in this
way. It is natural for research priorities to be dominated by the perception
of current problems experienced by the industry and for time limits to be
assigned. While we do not advocate that researchers should have a
complete free rein to range as they see fit, we do stress how vital it is that
research should have a strong underlying theme sustained in the long term.
For agronomy this should be to understand better the control of growth
and development. If knowledge is pursued in the right context, i.e. with
close association with the industry and its problems, benefits will inevitably
accrue, research needs will readily present themselves and a core of real
expertise will endure. If a new perspective is taken at every review, with
major alterations in objectives, research centre and group ,fragmentation
will result in every sense - most importantly there will be no coherent
appreciation of the current state of knowledge nor future needs.
Major reviews must be made from time to time but the notion that
reviews and funding should be on an annual basis is totally alien to
progress. All research must be accountable and its progress reviewed but
worthwhile agronomic research cannot be done on such a short term basis.
Similarly it has to be appreciated that expertise and experience cannot be
gained annually or re-created after a break of several years. It does seem
that the majority of growers seem to be firmly attached to the value of
organizing research on a short term basis. They simply want the solutions
and quickly. While this is understandable it is fraught with dangers. We
recognize that growers are likely to react intuitively and unfavourably to
the idea that progress depends on progressively improved understanding of
the system as a whole. This places a great responsibility on those charged
with administering research because they will have to argue a case that the
bulk of growers feel is neither needed or justified. We believe this to be the
greatest challenge for the future well-being of the potato industry and
indeed for applied research in all crops which is dependent on grower
funding. If research is allowed to be dominated by short term problems,
funded on the basis of annual or short term research contracts, it will lead
to very limited progress. Such a policy will force the remaining experienced
research workers out of agronomic research and thereby further erode
what has become a small research base.
Future dissemination
Throughout this chapter we have drawn extensively on the knowledge and
experience with the industry achieved through the formation of the
Cambridge University Potato Growers Research Association (CUPGRA)
in 1982 and through close associations with parts of the processing and seed
industries. The success of CUPGRA has derived from the close association
between growers and scientists which has meant that members have come
to know well the experiments in the field, the results and their implications
and can see how the overall programme has evolved. It has become natural
for growers to take the same approach and think in the same way as
researchers. The basis of the research has always been and continues to be
improved understanding of the crop's growth and development and to
apply this for commercial improvement. No compromise has been made in
this respect by the scientists and while this has caused a few growers to
leave claiming 'it's all too complicated', the vast majority (230 out of 250)
have remained members and through improved understanding have de-
rived benefit in the management of their crops. They have also become
much more effective in communicating their problems back to the researchers
and hence influencing future research.
There are several lessons here for the future. First, effective dissemina-
tion of research requires the closest possible contact between researcher
and grower. Where this is achieved the necessary development of new
ideas can be carried out by the growers with the research worker providing
the necessary 'back up' and scientific rigour for such work. Secondly,
growers who have a working knowledge of how the crop grows and
develops appreciate that agronomic management is more effective when
judgement is exercised in relation to particular circumstances rather than
when attempts are made to base it on a straightforward 'blueprint'. As the
range of specifications required by different markets becomes increasingly
exacting, it is inevitable that the achievement of potential will demand that
more and more decisions have to be taken. The value of basing those
decisions on a sound understanding will continue to rise. It is therefore
essential that the question of how to promote the most effective relation-
ship between research and the industry is addressed as part of the future
research and development strategy. Clearly in order to meet the needs of c.
15 000 growers it cannot be based on 'one to one' meetings between
growers and researchers. However, it will be vital to have sufficient well-
trained agronomists with research experience who can act as the essential
conduit between a small number of research scientists and a much larger
number of growers.
ACKNOWLEDGEMENTS
The authors wish to thank Sharon Parker, Susan O'Brien and Judy
Dickinson for their help (and forbearance) in the preparation of this
chapter. They also thank all members of the research group at Cambridge
University Farm and Dr J.N. Bean for access to unpublished data and help
in preparing the text.
REFERENCES
Allen, E.J. and O'Brien, P.J. (1985) Water use, irrigation need and the yield of
potato varieties. United Kingdom Irrigation Assoc. Conference. Irrigating
Potatoes. Silsoe College, Bedford, 19-24.
Agric. Sci., Camb., 94, 583-606.
Allen, E.J., O'Brien, P.J. and Firman, D.M. (1991) An evaluation of small seed
for ware potato production. J. Agric. Sci., Camb. (in press).
Asfary, A.F., Wild, A. and Harris, P.M. (1983) Growth, mineral nutrition and
water use by potato crops. J. Agric. Sci., Camb., 100,87-101.
Bailey, R.J. (1990) Irrigated Crops and their Management, Farming Press, Ipswich.
Bailey, R.J. and Minhinick, R. (1989) Irrigation News, United Kingdom Irrigation
Association, No. 15, pp. 19-30.
Bean, J.N., Allen, E.J. and O'Brien, P.J. (1991) Effects of floating plastic mulches
on soil and air temperatures and growth and yield of several potato varieties J.
Agric. Sci., Camb. (in press).
Biscoe, P.V. and Gallagher, J.N.G. (1977) Weather, dry matter production and
yield, in Environmental Effects on Crop Physiology (eds J.J. Landsberg and
C.V. Cutting), Academic Press, London, pp. 75-100.
Biscoe, P.V., Scott, R.K. and Monteith, J.L. (1975) Barley and its environment
III. Carbon budget of the stand. J. App. Ecology, 12,269-93.
Boone, F.R., Bouma, J. and De Smet, L.A.H. (1978) A case study on the effect of
soil compaction on potato growth in a loamy sand soil. 1. Physical measure-
ments and rooting patterns. Neth. J. Agric. Sci., 26, 405-20.
Brown, K.F., Messem, A.B., Dunham, R.J. and Biscoe, P.V. (1987) Effect of
drought on growth and water use of sugar beet. J. Agric. Sci., Camb., 109,421-35.
Campbell, M.D. (1972) The lower limit of soil water potential for potato growth.
Ph.D. Thesis, Washington State University, USA.
Chailakhyan, M.Kh., Yanina, L.J., Devedzhyan, A.G. and Lotova, G.N. (1981)
Photoperiodism and tuber formation in grafting tobacco onto potato. Doklady
Akademii Nauk. SSSR, 257, 127~0.
Colenso, J.M. (1989) Effects of potato harvesting practices on skin abrasion and
processing quality. Ph.D. Thesis, University of Cambridge.
Cook, H.F. and Dent, D.L. (1990) Modelling soil water supply to crops. Catena,
17,25-39.
Day, W., Legg, B.J., French, B.K., Johnston, A.E., Lawlor, D.W. and Jeffers, N. de
C. (1978) A drought experiment using mobile shelters: the effect of drought on
barley yield, water use and nutrient uptake. J. Agric. Sci., Camb., 91, 599-623.
References 879
Dunham, R.J. (1988) Irrigation of sugar beet: the main effect. British Sugar Beet
Review, 56(3), 34-7.
Epstein, E. and Grant, W.J. (1973) Water stress relations of the potato plant under
field conditions. Agron. J., 65, 400-4.
Evans, S.A. (1975) Maximum potato yield in the United Kingdom. Outlook on
Agriculture, 8, 184-7.
Evans, S.A. and Hield, J.R.A. (1981) The achievement of very high yields of
potatoes in the UK. J. Agric. Sci. Camb., 97,391-6.
Ewing, E.E. (1990) Induction of tuberization in potato, in The Molecular and
Cellular Biology of the Potato (eds M.E. Vadya and W.D. Park), C.A.B.
International, Wallingford, Chap. 3.
Firman, D.M. (1987) Leaf growth and senescence of the potato. Ph.D. Thesis,
University of Cambridge.
Firman, D.M. (1990) Cambridge University Potato Growers Research Association.
Annual Report.
Firman, D.M. and Allen, E.J. (1988) Field measurements of the photosynthetic
rate of potatoes grown with different amounts of nitrogen fertilizer. J. Agric.
Sci., Camb. 111, 85-90.
Firman, D.M., O'Brien, P.J. and Allen, E.J. (1991a) Flower initiation in potato
(Solanum tuberosum L.) sprouts and stems in relation to number of nodes and
tuber initiation. J. Agric. Sci., Camb. (in press).
Firman, D.M., O'Brien, P.J. and Allen, E.J. (1991b) Predicting the emergence of
potato sprouts. J. Agric. Sci., Camb. (in press).
Fowler, J.H. (1988) Effect of plant arrangement and density on growth, develop-
ment and yield of two potato varieties. Ph.D. Thesis, University of Cambridge.
Glauert, A.W. (1983) Carbon exchange of the sugar beet crop through a season.
Ph.D. Thesis, University of Nottingham.
Greenwood, D.J., Neeteson, J.J. and Draycot, Ann (1985) Response of potatoes
to N fertilizer: Quantitative relations for components of growth. Plant and Soil,
85, 163-83.
Harris, P.M. (1991) Mineral nutrition, in The Potato Crop', The scientific basis for
improvement (ed. P.M. Harris), Chapman & Hall, London, pp. 00·-00.
Hogge, M.e. (1989) Effects of site, season and husbandry on yield and processing
quality of the potato variety Pentland Dell. Ph.D. Thesis, University of
Cambridge.
Hogge, M.C., Stalham, M.A. and Allen, E.J. (1991) Effects offield treatments on
processing quality of Record and Pentland Dell potatoes during storage. J.
Agric. Sci., Camb. (in press).
Ifenkwe, O.P. (1975) Effects of row width and plant density on growth and
development of two maincrop potato varieties. Ph.D. Thesis, University
College of Wales, Aberstwyth.
potato. Outlook on Agriculture, 4, 211-7.
Jenkins, P. and Gillison, T.e. (1991). Get the best from plastic. Potato Review,
March/April 1991.
Jones, J.L. and Allen, E.J. (1983) Effects of date of planting on plant emergence
leaf growth and yield on contrasting potato varieties. J. Agric. Sci., Camb., 101,
81-95.
Lapwood, D.H. and Hering, T.F. (1968) Infection of potato tubers by common
scab (Streptomyces scabies) during brief periods when the soil is drying.
European Potato J., 11, 177-85.
Lesczynski, D.B. and Tanner, C.B. (1976) Seasonal variation of root distribution
of irrigated field-grown Russet Burbank potatoes. Am. Potato J., 53, 69-78.
Martin, C., Vernay, Rand Paynot, N. (1982) Physiologie vegetale photo-
periodisme, tuberisation, floraison et phenolamides. Comptes Rendus de
I'Academie des Sciences, Paris, 295, 565-8.
within the potato (Solanum tuberosum L.) crop, in relatiol!l to nitrogen
application. J. Agric. Sci., Camb., 107, 421-9.
Ministry of Agriculture, Fisheries and Food (1967) Potential Transpiration.. Tech-
nical Bulletin 16, Ministry of Agriculture, Fisheries and Food, London, HMSO.
Ministry of Agriculture, Fisheries and Food (1984) Fertilizer recommendations
1985-1986 Reference book 209.
Philosophical Transactions of the Royal Society of London, 281, 277-94.
Nield, J.R.A. (1977) The potato blueprint reviewed. Arable Farmer, May, 17-23..
Ojala, J.C., Stark, J.C. and Kleinkopf, G.E. (1990) Influence of irrigation and
nitrogen management on potato yield and quality. Am. Potato J., 67, 29--43.
Parker, c.J., Carr, M.K.V., Jarvis, N.J., Evans, M.T.B. and Lee, V.H. (1988)
Effects of subsoil loosening and irrigation on soil physical properties, root
distribution and water uptake of potatoes (Solanum tuberosum). Soil & Tillage
Research, 13, 267-85.
Penman, H.L. (1948) Natural evaporation from open water, bare soil and grass.
Proceedings of the Royal Society of London: Series A, 193, 120--45.
Penman, H.L. (1970) Woburn irrigation, 1960-8. iv. Design and interpretation vi.
Results for rotation crops. J. Agric. Sci., Camb., 75, 69-73; 89-102.
Potato Marketing Board (1979) Sutton Bridge Annual Review for 1978.
Potato Marketing Board (1990) Historical data summary from 1960.
Sale, P.J.M. (1973) Productivity of vegetable crops in a region of high solar input I.
Growth and development of the potato (Solanum tuberosum L). Aust. J. Agric.
Res., 24, 733--49.
Schepers, A. and Reestman, A.J. (1975) Mature plant resistance in relation to
some morphological characteristics of the potato plant. Proc. 6th Trien. Cant
Eur. Ass. Potato Res. Wageningen.
Scott, RK. and Jaggard, K.W. (1985) The effects of pests and diseases on growth
and yield of Sugar Beet. Proceedings of the 48th winter congress. Institute Inter-
national de recherches, Betteravieries, pp. 153-69.
Scott, RK. and Wilcockson, S.J. (1978) Application of physiological and agrono-
mic principles to the development of the potato industry, in The Potato Crop;
The scientific basis for improvement (ed. P.M. Harris), Chapman & Hall,
London, pp. 754.
Squire, G.R. (1990) The Physiology of Tropical Crop Production, C.A.B.,
Wallingford.
Stalham, M.A. (1989) Growth and water use of the potato variety Record on
contrasting sites. Ph.D. Thesis, University of Cambridge.
Stone, D.A. (1982) The effects of subsoil loosening and deep incorporation of
nutrients on yield of broad beans, cabbage, leek, potatoes and red beet. J.
Agric. Sci., Camb., 98, 287-306.
References 881
Sylvester-Bradley, R., Stokes, D.T., Scott, R.K. and Willington, V.B.A.
(1990) A physiological analysis of the diminishing responses of winter wheat to
applied nitrogen. 2. Evidence. Aspects of Applied Biology, 25, Cereal Quality
II.
Trebejo, 1. and Midmore, D.J. (1990) Effect of water stress on potato growth, yield
and water use in a hot and a cool tropical climate. J. Agric. Sci. Camb., 114,
321-34.
van Loon, C.D. and Bouma, J. (1978) A case s.tudy of the effect of soil compaction
on potato growth in a loamy sand soil 2. Potato plant responses. Netherlands 1.
Agric. Sci., 26, 421-9.
Watson, D.J. (1952) The physiological basis of variation in yield. Advances in
Agronomy, 4, 101-30.
Watson, D.J. (1968) A prospect of crop physiology. Annals of Applied Biology,
62, 1-9.
Weaver, J.E. (1926) Root Development of Field Crops, McGraw-Hill, New York
and London.
Welbank, P.J., Gibb, M.J., Taylor, P.J. and Williams, E.D. (1974) Root growth of
cereal crops. Rothamstead Experimental Station, Report for 1973, part 2,
26-66.
Wiicockson, S.J., Allen, E.J., Scott, R.K. and Wurr, D.C.E. (1985) Effects of crop
husbandry and growing conditions on storage losses of Pentland Crown
potatoes. 1. Agric. Sci., Camb., 105,413-35.
Wolfe, D.W., Fereres, E. and Voss, R.E. (1983) Growth and yield response of two
potato cultivars to various levels of applied water. Irrigation Science, 3, 211-22.
Wurr, D.C.E., Fellows, Jane, R. and Allen, E.J. (1991) Determination of
optimum tuber planting density in the potato varieties Pentland Squire, Cara,
Estima, Maris Piper, and King Edward. 1. Agric. Sci., Camb. (in press).
Index
Names of cultivars are indicated by (cv.). n denotes ireference to a footnote.
Abscisic acid (ABA) Agrotis ipsilon 477, 489

in potato plant 126, 127 Agrotis segetum 477, 489
and suberization 112 Ailsa (cv.), tuber quality 520
and tuber formation 112, 118, 120--2 Air flow in storage 620--1
Abscission of flower 136 Albumin 51<0
Acarina 490 Alcidodes westermanni 477
Acaulia Aldicarb 426
classification 17, 38 Allagash Russet (cv.), organic acid content
evolution 55, 56, 57 707
geographical distribution in South Alloxydim 381
America 45 Alloxydim-sodium 391
Acetaldehyde, influence on respiration 648 Allozyme analysis 54
Acetic acid, influence on respiration 648 Alopecurus myosuroides (Black-grass) 378,
Ackersegen (cv.) 392
tuber structure 112 Alpha (cv.) 204
water use 217 amino acid content 699
Aclonifen 393 tuber structure 112
Actipron 392 Alternaria solani
Added value 807-8 control of 423, 428
Africa geographical distribution 407
climate and yields 738-9 survival 420
consumption 801, 806 symptoms 407
imports and exports 800, 805 tuber infection at lifting 417
production 796-7 Altitudes for potato growing 4, 41-5
Ageing Amino acids 509
causes of premature senescence 407-8 2-aminobutane 427
stages in 261-3 Amitrole 384
see also Physiological age Amphibious bistort (Polygonum
Agricultural Development and Advisory amphibium) 377
Service (ADAS) 458 Amrasca biguttala biguttala 483
Agriotes 477, 486 Anatomy of potato plant 65
Agrobacterium rhizogenes as agent for Andean potato weevils (Premnotrypes) 476,
introducing genes 144 477, 478, 487
Agrobacterium tumefaciens 126, 114 Andes
Agromyzidae 490 consumption 805
Agronomy 816 origin of potato cultivation 794
and breeding 353-5 Angiosorus solani 404
general principles 845-53 geographical distribution 407
knowledge of 875-7 symptoms 407
Agrostis gigantia (Black bent) 376 Annual meadowgrass (Poa annua) 378, 387,
Agrostis stolonifera (Creeping bent) 376 389, 391, 392
Agrotis 477, 489 Anomala 487
Index 883
Anomala dimidiata 477 Bacterial diseases, in the tropical lowlands
Anther culture 141-2 768-70
Anthocyanins 515, 520 Bacterial ring rot, import restrictions 404
Apex, see Shoots, apex; Stolons, apex Bacterial wilt
Aphanes arvensis (Parsley piert) 391 and climate 403
Aphididae 480--3 in the tropical lowlands 768
Aphids 272, 480-3 Bangladesh
and diffused-light storage 736 example of production 776-7
leaf hairs as resistance mechanism 85--6 production 797--8, 805
trapping of 482-3 Bed planting 577--8
Aphis craccivora 481 Belonolaimus 442
Aphis fabae 477, 481 Belonolai¥tHlS longicaudatus 440
Aphis frangulae 481 Belt planters 576
Aphis gossypii 477,481 Benomyl426
Aphis nasturtii (Buckthorn aphid) 477, 481, Bentazone 391-2
482 Bintje (cv.)
Archaeological evidence of domestication amino acid content 699
2-3 carbohydrates 685-9, 690-1, 695, 696
Area under production 796-7 mineral nutrition 179-80, 193-4
Aristo (cv.), and fertilizers 206 organic acid content 706
Arma (cv.), heat tolerance 758 regeneration 69
Arran Banner (cv.) respiration 640
leafy shoot 71 root depth 838
tuber development 104 seed tubers 257
tuber quality 520 tuber quality 545, 667
Arran Comet (cv.) tuber structure 112
leafy shoot 76 use in France and Netherlands 806
leaves 824--6 wound healing 653
plant density 315 Black bent (Agrostis gigantia) 376
seed tubers 252, 258, 267, 281, 282 Black bindweed (Fallopia convolvulus) 378,
tuber shape and size 548 388, 391
Arran Consul (cv.) Black dot 405, 414, 416, 423, 431
growth 660 geographical distribution 407
respiration 638 symptoms 407
seed tubers 250 Blackening
Arran Pilot (cv.) after cooking 533--6
leafy shoot 70, 71 internal, effect of fertilizer use 207
seed tubers 250, 256 Black-grass (A,topecurus myosuroides) 378,
tuber development 100-4, 124 392
tuber structure 98 Blackheart 517
Ascorbic acid 511-12 Blackleg 414, 420, 421
content 512, 705 conaol of 426
Asia geographical distribution 406
consumption 801, 806 and plant spacing 410
imports and exports 800 seed tuber decay 404, 405
production 797 symptoms 406
Asparagine 510 Black nightshade (Solanum nigrum) 375,
Assia (cv.), and carotenoids 515 378,388
Atlantic (cv.), organic acid content 707 Black scurf, 416, 419, 420, 424, 426
Atriplex patula (Orache) 378 and cut seed 409
Aulacorthum circumfiexum 481, 482 geographical distribution 408
Aulacorthum solani 477, 481, 482 incidence after various treatments 274
Austroasca viridigrisea 477 seed tuber decay 405
Available Water Capacity (A WC) 221 and stems 310
Avenafatua (Wild oats) 378, 381, 392 symptoms 408
Blackspot 525, 527, 528-30,581
Bacteria Blemishes on tubers 415-16
geographical distribution 406 Blight 46,394,412,533, see also Early blight;
symptoms 407-9 Late blight
Bacterial brown rot 404 Blue spot 646-7
884 Index
Bolivia, potato production 804 geographical distribution in North and
Bonum (cv.), organic acid content 706 Central America 44
Boron, content 165-7
Bothinus maimon 477 Calcium
Botrytis cinerea content 165-7
geographical distribution 407 and hollow heart 518
and leaves and stems 414 and internal rust spot 519
symptoms 407 Cambridge University Potato Growers
Box storage and handling 593-4 Research Association (CUPGRA) 877
Brachycaudus helichrysi 481 Camptothecin 672
Brazil, potato production 804 Canada, seed trade 800
Breeding Cann.ing 547-8
behaviour 48-51 Canopy, see Leaves
history 335-7 Captafol 428
methods 337-52 Cara (cv.)
objectives 353-62 and growth 819
other techniques 362-4 and intercepted radiation 820
population breeding 350 and leaves 821, 827, 831, 833, 843
production of commercial varieties 361-2 and nitrogen 168-74, 181-2, 202, 860, 863
programmes 342-9 and physiological age 825-6
statutory trials 349-50 plant density 316, 328-9
in the tropics 740-1 and planting date 848
True Potato Seed (TPS) 351 and potato cyst nematodes 450-1
Bridgenorth (cv.), nitrogen and yield 179 and roots 838, 843
British Agrochemical Association (BAA) seed tubers 248, 263, 267
380 and tuber initiation 845
British Atlas of Potato Varieties 70 and water supply 218, 220
Broad mites (Polyphagotarsonemus latus) and yields 853-4
477,490 Carbendazim 427
Brodrick (cv.), tuber quality 543, 545, 546 Carbohydrate metabolism 124
Brown centre 517,518-19 Carbohydrates
Browning changes during storage 684
enzymic (discolouration) 530-2 content 802-3
non-enzymic, during processing 540-7 Carbon dioxide 641-2
Brown Oil of Vitriol (BOV) 396 and intercepted radiation 817
Brown rot 404,414,415 and wound healing 657
and climate 403 Carbon disulphide 675
and cut seed 409 Carotenoids 515
geographical distribution 406 Cells
symptoms 406 division and expansion in the tuber 105
Brown spot, see Rust spot division stimulated by wounding 111
Bruising 517, 524-6 Central America
causes 580 consumption 801
factors 581 geographical distribution of wild potatoes
internal 528-30 44-6
statutory and other standards 578-9 imports and exports 800
types of bruise 581 production 797
Buckthorn aphid (Aphis nasturtii) 477, 481, Ceramics depicting potatoes 2-3
482 Cereal cyst nematodes (Heterodera avenae)
Buds 456
adventitious 69, 74 Ccrtilfied seed 271-8, 405
development as leafy shoot 93-4 Charcoal rot 424, 425
development as stolon 93-4 geographical distribution 407
lateral 70 symptoms 407
meristems, changes at sprouting 130-3 tuber infection at lifting 417
Bulbocastana Charlock (Sinapsis arvensis) 378
chemotaxonomy 54 Cheiroplatys latipes 477
classification 15, 19 .Chemical composition, changes during
crossability 51 storage 677-79
evolution 56 Chemicals, effect on respiration 647-50
Index 885
Chemical treatments, effect on wound Coccinellids, leaf-feeding 476, 486, 493
healing 656 Coiled sprout 424
Chemotaxonomy 53-4 Coleoptera 485-7
Chenopodium album (Fat hen) 375, 378, Colletotrichum, and irrigation 416
391, 392 Colletotrichum cocco des 414, 419
Chickweed, common (Stellaria media) 377, geographical distribution 407
378 symptoms 407
Chile, marketing 805 Colorado potato beetle, see Leptinotarsa
China decemlineata
marketing 805 Columbia root-knot nematode, see
origin and spread of potato 795 Meloidogyne chitwoodi
production 797-8 Commersoniana
production figures 507 classification 15, 22-3
Chlorine, content 165 crossability 51
(2-chloroethyl)-trimethylammonium evolution 56
chloride (CCC), and tuber formation geographical distribution in South
113, 119-120 America 45
Chlorophyll 515-16 Common chickweed (Stellaria media) 377,
changes during storage 707-8 378
Chloropicrin 426 Common couch grass (Elymyus repens) 376.
Chloropropham 666--70 384, 391, 392
Chlorosis, causes 406, 408 Common fumitory (Fumaria officianalis)
Chocolate spot, see Rust spot 378, 387, 388, 389, 390
Chromosome numbers 51 Common reed (Phragmites communis) 376
of individual species 15-17, 18-41 Common scab
Chrysanthemum segetum (Corn marigold) and consumer quality 514
378,387,389,391 control of 427
Chrysomelidae 485-6 and cut seed 409
Cicadellidae 483-4 geographical distribution 408
CIP (International Potato Center) 41, 341, and irrigation timing 869
496,507 risk determination 431
Circaeifolia survival of pathogens 404
classification 15,23 symptoms 408
crossability 51 Composition, changes during storage
evolution 56 carbohydrates 684
geographical distribution in South chemical composition 677-79
America 45 chlorophyll 707-8
Cirsium arl'ense (Creeping thistle) 375, 376 dry matter 679-84
Citric acid 706, 707 hormones 708-9
Classification of potato species 13-41 lipids 708-9
Clal'ibacter, tuber infection during growth N-fraction 696--703
415 phenolif compounds 706
Clavibacter michiganensis subsp. sepedonicus pigments 706--7
414, 420, 421, 428 terpenoids 708
and cut seed 409 vitamins 704-6
geographical distribution 406 Compression bruising 525
symptoms 406 Conicibaccata
Cleavers (Galium aparine) 374, 378 chemotaxonomy 53
Climate classification 16, 26-8
and diseases 403 evolution 57
and yield 738 geographical distribution in North and
Clivia (cv.), nitrate content 703 Central America 44
Clonal reproduction 337-9 geographical distribution in South America
Clopyralid 393 45
Clostridium 418 Conidiobolus obscurus 493-4
Clumping 310--11 Conodorus 477. 486
Cobbler (cv.) Consumption
organic acid content 707 changing patterns 806--7
tuber structure 105 in the tropics 731
Coccinellidae 486 Control of Pesticides Regulations 382
886 Index
Control of Substances Hazardous to Health Cyanazine 390-1
382 Cyanidin 515
Convolvulus arvensis (Field bindweed) 377 Cycloxydim 392
Cooked potato, quality 533-9 Cystine 509
Cooking methods and their effects 802-3 Cytokinins
Copitarsia turbata 477 and lateral shoot development 95--6
Copper, content 165-7 in leaves and stems 126
Copper salts 426 in stolon tips 126-7
Corky ringspots 440 and suberization 112
Corn marigold (Chrysanthemum segetum) and tuber growth 122-3
378, 387, 389, 391 Cytology 52-3
Costs
of nematode control 460-2 Dalapon 384-5, 386
of preventing disease 404-5 Daleo (cv.), and water use 217
of production, in the tropics 728-9, 730-1 Damage
of production, United Kingdom 808-12 during harvesting 578-82
of seed 292 mechanical 432, 524-30, 580
Couch grass, common (Elymyus repens) 376, resistance to, see Damage resistance
384, 391, 392 statutory and other standards 578-9
Cracking 516-17, 524 Damage index 579
Craig's Defiance (cv.), growth 660 Damage resistance
Craig's Royal (cv.), mineral nutrition 166, breeding for 354, 360-1
187 measurement of 581-2
Creeping bent (Agrostis stolonifera) 376 Date of planting, effect on tuber yield 269
Creeping thistle (Cirsium arvense) 375, 376 Day length
Criconemoides 442 and leaf form 81
Crinkling of foliage 406 and shoot apical meristem 71
Critical photoperiod (CPP) 116-18 and tuber formation 113-18
Crop rotation, see Rotation of crops Dazomet 426
Crossability 48-51 and plant growth 452
Crossing, problems in producing seed 340-1 DD 426
Crude protein content 802-3 Defoliation
Crushing 524, 580 dry matter content 523
Crushing bruise 581 fungal causes 407
Ctenicera 477, 486 statutory dates 273
Cuba, potato production 797-8 time of 281
Cultivar characteristics 166 vi,ral causes 411
and breeding 341,354 Dehydration (manufacturing process) 547
and colouration 706-7 Delphinidin 515
and dormancy 659--60 Demissa
and drought 240-1 chemotaxonomy 53, 54
drought tolerance ranking 240 classification 17, 40-1
and respiration 651 evolution 55, 57
and tuber size 513 geographical distribution in North and
and wound healing 653 Central America 44
and yield 853--6 Desiccation 582, 583
see also individually-named varieties
Culture, experiments in 118-20 amino acid content 699
Cuneoalata heat tolerance 758
classification 16, 26 leafy shoot 76
evolution 55, 56, 57 leaves 823-4, 825--6, 836, 847
geographical distribution in South and nitrogen 858, 862, 865
America 45 plant density 309, 315, 319
Cup planters 575--6, 578 planting date 857
Curculionidae 487 regeneration 144
Cuts 524 root depth 838
Cut seed 247, 293-4 root growth 225
and disease 409 seed twbers 266,267,269,283
in the tropics 771 sprouting 661
Cutworms 489 tissue culture 139, 142, 143
Index 887
tuber quality 520, 531 geographical distribution 439
and water supply 241 resistance to 457
wound healing 653 and tubers 440
Desprouting 262, 266 Ditylenchus dipsaci (Stem nematode) 440
Development of potato plant 65 geographical distribution 439
Diabrotica 477 resistance to 457-8
Diagnosis and Recommendation Integrated Docks (Rumex) 377
System (DRIS) 188-9 Dr McIn10sh (cv.), evaporation 630
Diana (cv.) Domestication 1-4
and leaves 828,831,833 Dormancy 128-33, 657-64
and stems 828 seed tubers 250-2
and yields 853 Drop tests 581
Diclofop-methyl 392 Drought
Dietary fibre content 508, 508-9, 802-3 and cultivars 240-1
Dihaploids 351-2 sensitivity 217,228-36
Dimethipin 395 tolerance ranking of cultivars 240
Dimethylnaphthalenes (DMN), and sprout in the tropical lowlands 762-3
inhibition 665-6, 667 and yield loss 238-9
Dinoseb 380, 395 Dry matter
Diploids changes during storage 679-84
crossing 48 content 508, 521-4, 85.3
cultivated species 36, 42 efficiency of production 836-7
Diptera 490 and fertilizer use 206-7
Diquat 386, 388, 393, 395-6 and intercepted radiation 169-76, 180
Directive on the Marketing of Seed Potatoes and nitrogen content 864
275 and transpiration 216-17
Discolouration Dry rot
after cooking 533-6 control of 427
vascular 517, 520 geographical distribution 407
Disease control 421-2 seed tuber decay 404, 405
chemical 426-8 in store 418, 419
cost of 404-5 symptoms 407
cultural 422-6 Dye, geographical distribution and
future prospects 431-2 symptoms 406
growing conditions 423-5
legislation 428-9 Earlaine (cv.), flower 136
resistant cultivars 429-31 Early blight
seed certification 428-9 and climate 403
storage management 422-3 control of 423,427,428
Diseases and foliage 412
brief history 403 geographical distribution 407
and climate 403 sympt<vns 407
control, see Disease control in the tropical lowlands 768
detection by machine vision 603-4 Early death 408
effects of 405-19 Early Ohio (cv.), root depth 838
resistance to, considerations in breeding Early potatoes, areas planted in UK 268
354,357-60 Early wilt disease 442
and yield loss 404 Earth included in storage, effect on air flow
Distortion of foliage 406 620
Distribution, geographical East Germany, see Germany
diseases 406-8 Ecology 41-6
Globodera 445 Economic considerations, stem density
insect pests 477 323-30
nematodes 439 Economics of production, United Kingdom
potato cyst nematodes 445 808-12
potatoes 41-6 Ectoparasites, migratory 442
potato species 18-41 EEe regulations
Ditylenchus (Potato rot nematodes) 440 req uirements for certification 278
Ditylenchus destructor (Tuber-rot nematode) seed-producing land 453
control of 454, 456, 459 Eelworms 396
888 Index
Elateridae 486 and nitrogen 852, 859-61, 863
Elite (basic seed grade) 275-7 physiological age 825-6
Elymyus repens (Common couch grass) 376, plant density 307, 316, 321, 328-9
384, 391, 392 planting date 852
Embryo balance numbers (EBN) 51 roots 838--9, 843, 845
of individual species 14--41 seed tubers 258, 260, 270
Embryogenesis 137-8 tuber growth 819
Embryoids 141-2 tuber quality 514
Empoasca abrupta 484 yields 853-4
Empoasca arida 484 Estolonifcra, classification 14-18
Empoasca devastans 477, 483 Ethanol, influence on respiration 648
Empoasca fabae 477 Ethepon 519
Empoasca filamenta 484 Ethylene, influence on respiration 647-8
Empoasca solani 484 Etuberosa, classification 14-18
Endoparasites Europe
migratory 442-3 consumption 801
sedentary 443-7 imports and exports 800
Energy introduction into 5-9
content 802-3 production 507, 796-7
value 508 spread throughout 9-10
England, introduction into 5-6, 9 European Association fO.T Potato Research
Environmental concerns 551-2 (EAPR) 658
Environmental effects on tuber formation European corn borer (Ostrinia nubialis) 489
113-18 European Economic Community,
Epicauta 477 regulations
Epicauta hirticornis 477 req uirements for certification 278
Epicure (cv.), leafy shoot 72-3, 76 seed-producing land 453
Epilachna 477, 486 European Plant Protection Organization 457
Epitrix 477, 486 Evaporation from stored potatoes 629-32
EPTC 385 Evolution 56-7
Eremotermes 477 divergence in wild species 55-6
Erwinia relationships of cultivated species 57
control of 425 Explant
and cut seed 409 meristem culture 76-7
and general decay 415 somatic tissues 138
and irrigation 416 Exports, regional trade 800, 805
and Ostrinia nubialis (European corn Eyes 97-8
borer) 489 number and size, as measure of plant
and rain 420 density 293-5
resistance to 431 position, effect on yield 293-4
and volunteer potatoes 398
Erwinia carotovora subsp. atroseptica Fallopia convolvulus (Black bindweed) 378,
geographical distribution 406 388, 391
seed tuber decay 409 False root-knot nematode, see Nacobbus
and stems 414 aberrans
symptoms 406 Farmyard manure 194-7
in the tropics 770 Fat content 508, 509, 802-3
Erwinia carotovora subsp. carotovora Fat hen (Chenopodium album) 375, 378,
and blackleg 414 391, 392
geographical distribution 406 Feltia experta 477
seed tuber decay 405 Fentin acetate 428
symptoms 406 Fentin hydroxide 396, 428
in the tropics 770 Fertilization 136
Erwinia chrysanthemi Fertilizers
and blackleg 414 application 573-4
geographical distribution 407 application rate and irrigation 202
symptoms 407 economic considerations 192-4
in the tropics 770 factors influencing use 164
Estima (cv.) placement 197-8
leaves 828-33, 835, 843, 845 and plant density 203-5
Index 889
and potato quality 205-7 symptoms 407-9
and rainfall 200-3 Fusarium
requirements 177-94 control of 423, 424
timing of application 199-200 geographical distribution 407
trends in England and Wales (1958-86) infection at lifting 416, 417
167-8 resistance to thiabendazole 427
and tuber size 206 spread 420
and yield 178-82 in store 418
see also Nitrogen fertilizers; Potassium survival 419
fertilizers symptoms 407
Field bindweed (Convolvulus arvensis) 377 and wilting 414
Field Capacity (FC) 221 Fusarium caerulum 533
Field pansy (Viola arvensis) 388 Fusarium solani
Field pennycress (Thlaspi arvense) 378, 391 control of 425
Flame desiccation 583 geographical distribution 407
Flavines 515 at lifting 417
Flavonal glycosides 53 symptoms 407
Flavones 515 Fusarium solani subsp. coeruleum
Flavour 539 geographical distribution 407
Flea beetles (Epitrix) 477, 486 infection at lifting 417
Flesh colour 514-5 seed tuber decay 405
Floral initiation 257-8 symptoms 407
Flower, development and structure 133-7 Fusarium sulpkureum 673
Flowering time, effect on breeding 49 Fusarium wilt, geographical distribution and
Fluazifop-butyl 393 symptoms 407
Fluorochloridine 393
Fluroxypyr 398 Galium aparine (Cleavers) 374, 378
Foliage Galls
analysis of nutrient content 188 causes 408
destruction 412-13 on roots 444
disease 412-13 Gamma irradiation 647, 673-4
disease control, chemical 427-8 Gangrene
premature death 413-14 and climate 403
symptoms and causes of diseases 406--8 control of 422, 425, 426, 427
see also Leaf area growth; Leaf area index; disease in store 418-9
Leaves geographical distribution 407
Folic acid 705, 706 and growth 409
Fomesafen 393 and lifting 416
Food and Environmental Protection Act 382 and plant spacing 410
Fracture cracking 517, 524 and potato eyes 295
France seed tuber decay 404, 405
marketing 812 and stqms 310
production 798, 799 symptoms 407
seed trade 800 and volunteer potatoes 398
Frost Gas exchange 638-42
breeding for tolerance 732-4, 740-1 Gelechiidae 487-9
evaluation of tolerance 733-4 Genes
frost-hardiness and leaf structure 83, 84 HI, aond nematode resistance 456-7, 458,
resistance 46 462,465
tolerance, in tropics 729 H2, and nematode resistance 457,458,465
Fumaria officianalis (Common fumitory) Genetic engineering 144-5
378,387,388,389,390 Genetic variability 46--8
Fumaric acid 706, 707 Genome differentiation in wild potatoes 55
Fumitory, common (Fumaria officianalis) Geographical distribution, see Distribution,
378,387,388,389,390 geographical
Fungal diseases 272 Geographical spread of potato 1-11, 794
Fungi Germany
geographical distribution 407-8 consumption 801
interaction with nematodes 414, 452 production 796-8, 799
as nematode control 456 seed trade 800
890 Index
Germination 66 Green peach aphid, see Myzus persicae
Germination inhibitors, prohibited in seed Green top harvesting 583
for sale 275 Grey mould
Gibberellic acid (GA) 675--6 geographical distribution 407
in culture experiments 118-9 leaves and stems 414
effect on tuber formation 120, 125 symptoms 407
Gibberellins Groundsel (Senecio vulgaris) 378, 387,
and dormancy 663 389
extracted from leaves 126 Growing season, length, effect on stem
in leaves 126 density 318
in meristem culture 76 Growth
and stolon formation 95--6, 127 agronomic principles 816-44
and tuber formation 120, 125, 127 damage by disease 405-10
in tubers 133 and nitrogen 168-76
and wounding 133 and planting date 856-7
Globodera (Potato cyst nematodes) 441-2, and root-feeding nematodes 447-51
448-51, 453-4 in tropical highlands 736-40
Globodera pallida (White potato cyst arid water use 214-19
nematode) 444-7 Gro\\·,th cracking 516-7
geographical distribution 438-9 Growth rate, effect of stolon size 92
interaction with fungi 414, 452 Gryllotalpa africana 477
resistance to 46, 359, 456-7, 458, 459, Guardian (cv.)
461 tuber quality 516, 517
and volunteer potatoes 398 and water supply 240
Globodera rostochiensis (Golden nematode Guatemala, production 804
of potato) 444-7 Gypsum 426
geographical distribution 438-9
interaction with fungi 414, 452 HI Genes, and nematode resistance 456-7,
resistance to 46,359,456-7,458,461 458, 462, 465
and volunteer potatoes 398 H2 Genes, and nematode resistance 457,
Globulin 510 458,465
Glucosides 53 Habitat, individual species 14-41
Glufosinate 393 Hairs on leaves 84--6
GluJamine 510 Halticoptera patellana 494
Glutelin 510 Handling, effect on respira.tion 645-7
Glycoalkaloids 532-3 Hard rot 418
effect on irradiation 548-50 Harrows, reciprocating 571
Glycosides 53 Harvesters 586-8
Glyphosate 377, 385, 393, 397, 398 cleaning systems 588
Golden nematode of potato, see Globodera design and mechanical damage 526
rostochiensis types 589-90, 591
Golden Wonder (cv.) Harvesting
leafy shoot 74 grading 594
seed tubers 250 handling 593-4
tuber quality 531 inspection 598--604
Gouges 524, 580 mechanical damage 578-82
Grading 594 potato quality 578
electronic, and mechanical damage 526 practices 871-2
seed crops 285 preparation 582-5
Grafting to support tuber development 115 seed crops 281, 284--6
Graminicides, use against grass weeds 377 sizer types 596-8
Graphognathus leucoloma 477 sizing 595--6
Grass weeds 376 stages 585-92
Great Britain, see United Kingdom transport 592-3
Greening 515--6 and weeds 375
and depth of planting 575 Haulm
effect of irradiation 548 destruction 394--6
Green Mountain (cv.) pulling 583-5
mineral nutrition 166 pulverizers 582-3
organic acid content 707 Healing, effect of irradiation 548
Index 891
Heat Humidification 636
output and sprout growth in storage 626, Humidity
676 and formation of wound periderm 111
production in store 621-5 and sprout growth 662
and respiration 642-3 and wound healing 655
Heat treatment, and virus elimination 77-8 Husbandry methods
Helicobasidium purpureum changes in 873-4
geographical distribution 407 and nematode reduction 456
spread 419 seed production 281-4
symptoms 407 Hybrids., wild 49
tuber infection during growth 415 Hydrogen cyanide, influence on respiration
Helicotylenchus 442 647-8
Heliothis armigera 477 Hypolithus 477, 486
H elminthosporium
and irrigation 416 Imazalil 427
resistance to thiabendazole 427 Import restrictions 428
Helminthosporium solani United Kingdom 404
and cut seed 409 Imports, regional trade 800
geographical distribution 407 Improved Haswell (cv.), nitrogen and yield
infection of eyes and early harvesting 274 179
and spread 420 Incubation period 262
symptoms 407 India
Henosepilachna ocel/ata 486 experiments in 744
Henosepilachna sparsa vigintisexpunctata 477 origin and spread of potato 795
Henosepilachna vigintioctopunctata 486 production 797-8, 805
Herbicides Ingifolia 57
methods of treatment 382-3 classification 16, 29
recommendations for use 382-94 geographical distribution in South
specific 384-92 America 45
use against broad-leaved weeds 377 Insecticides 478-80
weed control 380-2 botanical 493
Heterodera avenae (Cereal cyst nematodes) carbamate 427
456 organophosphate 427
Heterodera glycines 464 and potato leaf roll virus 482
Heterodera schachtii 465 Insects
Heteronychus arator 477 attacking leaves 477
Highland zones attacking stems 477
production 804 attacking tubers 477
production in the tropics 731-41 biology of 480-90
History control of 490-7
of breeding 335-7 geographical distribution 477
of diseases 403 Inspection
of the potato 1-11 of crops 598-{i04
Holland, see Netherlands of seed crops 277-8
Hollow heart 517-8,522,652 Institute for Storage and Processing of
Holmemoor (cv.), nitrogen and yield 179 Agricultural Produce (IBVL) 545
Home Guard (cv.) Integrated Pest Management (IPM) 495-7
growth 660, 823-4, 825-7 Intercepted radiation, see Radiation
mineral nutrition 201 interception
plant density 298-9,303,319 Intercropping, effect on disease in the tropics
respiration 644, 649 769
seed tubers 250, 252-3, 258, 267, 280, Internal rust spot (IRS) 517, 519, 520
282 International Potato Center (CIP) 41, 341,
Homoptera 480-4 496,507
Hormones Iprodione 427
changes during storage 708-9 Irish Cobbler (cv.)
endogenous 125-{i flower 136
treatment of tubers 120-4 tissue culture 139
see also Gibberellins tuber structure 105
Host plant resistance (HPR) 490-2 Iron, concentration in plants 165-7
892 Index
Irradiation 548-51 and fungicides 405, 431
radioactive 673-4 geographical distribution 407
and respiration 647 ,\Ud heating 425
and wound healing 656 resistance to 430, 431
Irrigation 236--7 and seed tubers 404
and common scab 869 symptoms 407
in the tropical lowlands 761-3 and volunteer plants 404
and tuber yield 201 Leaf area growth and development
and yields 866--70 and mineral nutrients 176--7
Isopropyl N-(3-chlorophenyl) carbamate and nocturnal rehydration 236
(CIPe) 666--70 and root size 182
Isopropyl N-phenylcarbamate (IPe) 666--70 variations 820-2
Isoproturon 393 Leaf area index
Isozyme analysis 54 and intercepted radiation 220, 819-22,
Ivy-leaved speedwell (Veronica hederifolia) 378 824-6, 833-4, 847
and nitrogen fertilizers 765, 860, 865
J assidae 483-4 and planting date 821-2, 824-6, 847
J assids, see Leafhoppers and root depth 842-3
Jelly end 517 seed 260-1
JUldika (cv.), nitrate content 703 and temperature 264
Juglandifolia 51 and water 842-3
classification 18 Leaf drop streak 411
Katahdin (cv.) Leaf growth
amino acid content 698 and development 822-36
flower 136 and plant spacing 309
glycoalkaloid content 701 and yield 260
tuber development 101, 104 Leafhoppers (Jassids) 476, 478, 483-4, 492
vitamin content 705 Leafminer flies, see Liriomyza
Kennebec (cv.) Leaf roll virus
respiration 644, 649 control of 425,427
tuber development 102, 104 geographical distribution 406
tuber structure 105, 112, 113 resistance to 431
vitamin content 705 and spread 421
Kerr's Pink (cv.), leafy shoot 71 and stunting 411, 412
King Edward (cv.) symptoms 406
carbohydrates 693 and tuber discolouration 521
diseases 405, 425 and yield losses 404
flower 134 Leafy shoot 70
leafy shoot 70, 71-2 Leaves
mineral nutrition 177, 201 description 14
plant density 294, 301, 315, 316, 319, 326, development and structure 81-86
328-9 growth, see Leaf growth
respiration 638, 640, 645 leaf canopy 833--{)
root growth 225 longevity 830-33
seed tubers 250, 254 longevity, effect of nitrogen uptake 322
sprout growth 660-1 morphology 829-30
stolon 87 nematodes 440
tissue culture 142 number of 824-8
tuber quality 511, 529 rate of appearance 828-9
and water supply 225, 240 size 829-30
Kirsty (cv.), tuber quality 520 water potential 228-31, 233-4, 236
Knotgrass (Polygonum aviculare) 377, 378, and water stress 236
387, 388, 389 see also Foliage; Leaf area growth; Leaf
area index
Land preparation 571-2 Legislation
Langworthy (cv.), leafy shoot 74 against disease 428-9
Late blight control of nematode damage 453-4
and breeding 430 seed certification 275
and excessive nitrogen 423 Lenape (cv.), glycoalkaloid content 532
and foliage 412-3, 427 Lepidoptera 487-9
Index 893
Leptinotarsa decemlineata (Colorado Potato geographical distribution in South
Beetle) 476, 477, 478, 479-80 America 45
control of 493, 494 Magnesium
resistance to 490-1, 492 content 165-7
resistance to insecticides 482, 485 toxicity and hollow heart 518
Libertas (cv.), and water use 217 Magnesium sulphate 426
Light Magnum (cv.), organic acid content 706
and glycoalkaloids 533 Mainstems, number of, related to number of
and greening 515-6 sprouts 255
and sprout growth in storage 662 Majestic (cv.)
Lignicaulia carbohydrates 685
classification 15, 23 gas exchange 638, 650
crossability 51 leafy shoot 71
evolution 56 plant density 295, 311, 315
geographical distribution in South root growth 225
America 45 seed ttlbers 250, 254
Limonius 477, 486 storage temperature 616
Linguistic evidence of origin 4-5 water loss 632
Linuron 387-8 and water supply 214, 240
Lipaphis erysimi 481 wound heal,ill1lg 653
Lipids, changes during storage 708-9 Maleic hydrazide 399, 670-1
Liriomyza (Leafminer flies) 477, 478, 482, 490 and respiration 644, 649
resistance to insecticides 482 Malic acid 706, 707
in the tropics 767 Malvidin 515
Liriomyza huidobrensis 490, 492, 494 Mancozeb 428
in the tropics 767 Maneb 428
Liriomyza patagonica 477 Manganese, content 165-7
Liriomyza trifolii 477,490,494 Manures, organic 194-7
Listroderes obliquus 477 Marfona (cv.), seed tubers 270
Little potato disorder 262, 409, 424 Maris Bard (cv.)
Longidorus 442, 460, 462-3, 466 and growth 823
Longidorus elongatus 442, 448 seed tubers 263, 268
Longidorus leptocephalus 442, 455 tuber quality 520
Longipedicellata Maris Huntsman (cv.), root growth 225
and browning 531 Maris Page (cv.), and nitrogen 204
chemotaxonomy 54 Maris Peer (cv.)
classification 17, 38-40 plant density 315, 319
evolution 55, 56, 57 and pot3lt<o cyst nematodes 450-1
geographical distribution in North and tuber shape and size 548
Central America 44 Maris Piper (cv.)
Lowland zones development 141
production 804-5 leaves 823--6, 828, 831, 833
production in the tropics 742-79 nitrogen 860, 862-4
Lutein 515 plant density 294,304,308,315,316,319,
Lysine 509 327-9
and potato cyst nematodes 449
Machine vision root depth 838
sizing 597 root growth 225
tuber inspection 603-4 seed tubers 258, 259, 260, 280
Macrophomina phaseolina tuber quality 531, 548
geographical distribution 407 and water supply 238, 239, 240-1
symptoms 407 yields 853
tuber infection at lifting 417 Marketing
tuber infection during growth 415 in developing countries 805
Macrosiphum euphorbiae (Potato aphid) in Great Britain 812-14
477,481,482 Markets
Macrosteles fascifrons 484 changes in 873, 874-5
Madagascar, potato production 797-8 requirements 507
Maglia 57 sizing for 595
classification 16, 29 Markkrone (Peri medullary zone) 91, 99,105
894 Index
Matricaria matricarioides (Mayweeds) 377 shoot apical 71, 74, 75
Matricaria recutita (Mayweeds) 377 Methionine 509
Maturity Methyl bromide 426
respiration as index of 650 Metoxuron 395, 396
and wound healing 653 Metribuzin 381, 388, 390-1, 392
Mayweeds 377, 378 Metsulfuron methyl 393
MCPA 380 Mexico, production 797-8, 804
Mechanical damage, see Damage Micropropagation, in vitro 362
Mechanization of crop production 570-607 Microtubers 363,578
M egaulothrips usitatus 490 Middle East
Megistacroloba exports 805
classification 15, 25--6 production 796
crossability 51 Mineral content, effect of cooking 803
evolution 55, 56, 57 Mineral elements, see Nutrients, mineral
geographical distribution in South Mineral oil 427
America 45 Minitubers 578
Melanin 525, 581 Mites 490
Melanotus 477,486 in the tropics 767
Meloidogyne (Root knot nematodes) 443-4 Moisture content 802-3
control of 455 Molybdenum, content 165-7
geographical distribution 440 Monohaploids 351-2
and growth 450-1 Monolinuron 389
and soil temperature 441 Monona (cv.), organic acid content 707
Meloidogyne arenaria Mop top virus 406, 416
and galls 444 M()('elliformia
geographical distribution 441 chemotaxonomy 54
resistance to 458 classification 15, 19
and soil temperature 441 crossability 51
Meloidogyne chitwoodi (Columbia root-knot evol,l!Iltion 56
nematode) geographical distribution in North and
control of 455 Central America 44
and galls 444 Morphology
geographical distribution 439, 441 and breeding 354
and soil temperature 441 seed tuber 248--50
spread 466 tuber 97-8
Meloidogyne hapla (Northern root-knot Mosaic 411
nematode) geographical distribution 406
control of 455 symptoms 406
geographical distribution 439, 441 Mottling, causes 406
interaction with fungi 414 Mulches, plastic 394
resistance to 458 Multa (cv.), tolerance to potato cyst
and roots 444 nematodes 459
and soil temperature 441 Myllocerus subfasciatus 477
and wilting 452 Myzus ascalonicus 481, 482
and yields 448 Myzus persicae (Green peach aphid) 476,
Meloidogyne incognita 477,480,481
control of 456 control of 491-2
geographical distribution 441 resistance to insecticides 482
resistance to 46, 458
and roots 451 Nacobbus 441,455
and soil temperature 441 Nacobbus aberrans (False root-knot
Meloidogyne javanica nematode) 443
geographical distribution 441 geographical distribution 439, 440
resistance to 458 interaction with other organisms 452
and soil temperature 441 spread 466
Melolontha 477, 487 Nacobbus batatiformis 443
Mentor (cv.), mineral nutrition 166 Nacobbus serendipiticus 443
Mercury 426, 427 Nacobbus serendipiticus bolivianlls 443
Meristem Nadine and growth 853
culture 75-8 Names for the potato 5, 9
Index 895
Necrosis Norchip (cv.)
causes 406-7 organic acid content 707
low temperature 517 vitamin content 705
Nematicides 454-5 North America
Nematodes acreage and production 507
attacking roots 441-7 consl!lmption 801
attacking stems and leaves 440 geographical distribution in 44--6
attacking tubers 440-1 imports and exports 800
biology 438-47 introduction into 10
damage control and avoidance 453--62 production 797
effects on growth and yield 447-51 Northern root-knot nematode, see
interactions with other organisms 451-3 Meloidogyne hapla
research, future developments 462--6 Nutrients, mineral
Neomyzus circumflexum 481 accumulation and use 163-8
Neotylenchus vigissi (Pseudo stem essential elements 163-7
nematode) 440 leaching 190
Netherlands levels of 165
fertilizers and plant density 203-5 and rainfall 200-3
fertilizer rates recommended 185 relationship with growth and yield 168-77
haulm pulling 583-4 taken up and removed in the tubers
marketing 812 165-7
nematode control 460 in the tropical lowlands 763-7
production 797-8, 799 Nutritional value 508-9, 801-3
seed trade 800, 806, 811
statutory defoliation dates 273 Oceania
and virus infection 271, 273 consumption 801
winter rainfall and soil nitrates 203 imports and exports 800
Netted Gem (cv.) production 796-7
glycoalkaloid content 701, 702 Odontotermes obesus 477
tuber quality 533 Olmosiana
Newchurch (cv.), nitrogen and yield 179 classification 15, 23-4
New Potatoes Prescription 578 evolution 56
Niacin 512, 705, 706 geographical distribution in South
Nipaecoccus vastator 477 America 45
Nitrate pollution 837 Olympia (cv.), wound healing 654
Nitrate test strip 190-1 Oospora pustulans 452
Nitrogen Orache (Atriplex patula) 378
and blackening after cooking 535 Organic acids
and blackspot 529 changes during storage 706
content 165-7, 508, 509 content 707
and dry matter 179-81, 522 Organo-tin compounds 428
fertilizers see Nitrogen fertilizers Or.igin of potato 1-5, 794
and growth and yield 168-76 Ostrinia nubialis (European corn borer)
and leaf area development 176-7 489
and tuberization 118 Overheating in a stack of potatoes 624-5
yield prediction model 186 Oxalic acid 706, 707
Nitrogen fertilizers Oxygen 639-41
excess of 849-50 effect on wound healing 656-7
and leaf area 860, 865
and number of tubers 175 Pacheco (urns) 3
optimum economic rate of application 192 Paecilomyces lilacinus 456
pressure to restrict amount 193 Pakistan, production 797-8
recommendations for different soil types Palisade parenchyma, and frost hardiness
865 733
timing of application 190 Papaver rhoeas (Poppy) 378
in the tropical lowlands 763--6 Para crinkle 77, 78
and yields 857--66 Paracrinkle virus 406, 412, 425
see also Fertilizers; Nitrogen Paralongidorus 442
Noctuidae 489 Paralongidorus maximus 442
Nonanol672 Paraquat 380, 386, 388, 390, 393
896 Index
Paratrichodorus (Stubby root nematodes) production 798
geographical distribution 439, 440 Pest control
and growth 442 behavioural control 493-5
and stunting 441 biological control 493-5
virus transmission 440-1 botanical insecticides 493
Paratrioza cockerelli (Potato psyllid) 477, 484 cultural control 492
Parsley piert (Aphanes arvensis) 391 host plant resistance (HPR) 490-1
Pasteuria penetrans 456, 464 repellent plants 493
Patatin 510 Pests
Pathogens, survival and spread 419-21 control of, see Pest control
see also Bacteria; Insects; Nematodes; insects, see Insects
Pests; Viruses nematodes, see Nematodes
Pelargonidin 515 resistance to, breeding for 354, 356-60
Pendimethalin 390-1 Petonidin 515
Pendulum tests 582 Petota, classification 14-41
Pentaploids, cultivated species 37 Peunidin 515
Pentland Crown (cv.) pH, soil, effect on uptake of plant nutrients
growth 823 205
leaves 828-9, 830-3, 835 Phenacoccus grenadensis 477
and nitrogen 852, 863 Phenolic compounds, changes during storage
plant density 294-5, 301, 313, 319, 325-7 706
and potato cyst nematodes 450-1 Phenylamides 428
root growth 225 Philippines
seed tubers 278 climate and yield 742
tissue culture 141 example of potato production 776-7
and water supply 240, 241 production 804
wound healing 653 Phloem
yields 852, 855 leaf 82
Pentland Dell (cv.) stem 79-80
dry matter 853 stolon 91-2
growth 824 tuber 105
leaves 830-1, 833 Phoma, and irrigation 416
and mechanical damage 576 Phoma exigua
plant density 301, 307, 326 effect on potato eyes 295
and potato cyst nematodes 449-51 effect on stems 310
roots 838 Phoma exigua subsp. exigua
seed tubers 266 geographical distribution 407
tuber quality 514, 522, 531, 540, 548 seed tuber decay 405
Pentland Hawk (cv.), wound healing 653 symptoms 407
Pentland Ivory (cv.) Phoma foveata
plant density 301, 326 control of 423, 424
seed tubers 255 disease in store 418-19
Pentland Javelin (cv.) geographical distribution 407
seed tubers 263, 264, 267, 268 seed tuber decay 405
and storage 826 spread 420
tuber quality 531 symptoms 407
Pentland Squire (cv.) tuber infection at lifting 416, 417
and nitrogen uptake 862-3 Phosphorus
plant density 295, 316, 327-8 content 165-7
tuber quality 514, 520 and leaf area development 176-7
Penycuron 427 Photosynthesis 214-15
Perennial sowthistle (Sonchus arvensis) 377 Phragmites communis (Common reed) 376
Periclinal chimeras 73-5 Phthorimaea operculella (Tuber moth;
Periderm of the tuber 110-11 Tuberwl.J'rm) 476, 477, 478, 487-9
wound periderm 111-13, 652 and diffused-light storage 736
Perimedullary zone (Markkrone) 91,99, 105 and irrigation 762
Permanent Wilting Point (PWP) 221 trapping of 488
Peru in the tropics 736, 762, 767
climate and yield 738, 742 tuber infection during growth 415
early cultivation 4 Phyllognathus dionysius 477
Index 897
Phyllophaga 477, 487 Planting date, effect on growth and yield
Physiological age 260--70, 676-7 856-7, 858
effect on growth 856 Plastic mulches 394
effect on yield 858 Ploidy 50, 52
Phytophthora erythroseptica Ploughing 571-2
control of 423 PLRV 46, 358-9, 478, 480, 482
d,isease in store 419 Plusia orichalcea 477
geographical distribution 407 PMB (Potato Marketing Board) 812
post-lifting disease 417 Poa annua (Annual meadowgrass) 378, 387,
symptoms 407 389. 391, 392
tuber infection during growth 415 Poland
and wilting 414 consumption 800
Phytophthora infestans production figures 507
geographical distribution 407 Pollen sterility 136
and haulm destruction 394 Pollination 51
insect-transmitted 428, 430 controlled 339-40
and rainfall 200 Polyadenia
resistance to 46 classification 15, 21-2
seed tuber decay 405 crossability 51
survival 419 evolution 56
symptoms 407 geographical distribution in North and
and terpenoids 708 Central America 44
tuber infection during growth 415 Polygonum amphibium (Amphibious
tuber infection at lifting 417 bistort) 377
Pigmentation of tuber 74 Polygonum aviculare (Knot grass) 377, 378,
Pigments, changes during storage 706-7 387, 388, 389
Pink rot Polygonum persicaria (Redshank) 378, 391
and drainage 423 Polyphagotarsonemus latus (Broad mite)
geographical distribution 407 477,490
and orgalilic manures 424 Polyploidy 50, 52
seed tuber 404 Polyscytalum
symptoms 407 and irrigation 416
tuber infection during growth 415 resistance to thiabendazole 427
Pinnatisecta Polyscytalum pustulans
chemotaxonomy 54 and cut seed 409
classification 15, 19-21 and early lifting 424
crossability 51 geographical distribution 407
evolution 56 infection of eyes and early harvesting 274
Pit rot 418 and leaves and buds 416
Piurana and potato eyes 295
chemotaxonomy 53 spreading disease in store 419
classification 16, 28 and stems 310
evolution 57 and storage temperature 425
geographical distribution in North and survival of 420
Central America 44 symptoms 407
geographical distribution in South Pontiac (cv.), vitamin content 705
America 45 Poppy (Papaver rhoeas) 378
Plant analysis of nutrient content 187-92 Population breeding 350, 741
Plant density 292 Potash 184
effects 312-30 Potassium
and fertilizer use 203-5 and blackening after cooking 535
spatial arrangement 306-12 and blackspot 528-9
units 293-306 content 165-7
Planters effect on enzymic browning 531
cup feed, irregularity of spacing 309 effect on leaf area development 176-7
design 575-7 effect on mechanical damage 206
type, effect on tuber yield 270 Potassium fertilizers and dry matter COlJ;lltent
Planting 573, 574-8 523
seed crop, timing 281 Potato aphid (Macrosiphum euphorbiae)
in the tropics 770--6 477,481,482
898 Index
Potato blight, see Blight; Late blight; Early Production
blight areas and volumes 507
Potato cyst nematodes (Globodera) 441-2, changes in annual tonnage 730
448-51,453--4 recent trends 796-801
Potato, classification 15-17, 18--41 zones 804--5
Potato Marketing Board (PMB) 812 Progeny tests for diseases 360
Potato psyllid (Paratrioza cockerelli) 477, Prolamin 510
484 Prometryn 389, 399
Potato rot nematodes (Ditylenchus) 440 Propagation 334--5, 362
Potato sickness 452 Propham 666-70
Potato tuber moth, see Phthorimaea Propylene, influence on respiration 648
operculella Proteins 107-8
Potato tuberworm, see Phthorimaea content 508, 509-10
operculella electrophoretic studies 53--4
Potato viruses 482 Protoplasts 142-3
A 78 Prunus, and aphids 481
S 77, 78 Pseudomonas solanacearum 452
X 77, 78, 358 and cut seed 409
Y 46,78,358,412,478 geographical distribution 406
potato leaf roll virus (PLRV) 46, 358--9, resistance to 46
478, 480, 482 survival 419
see also Viruses symptoms 406
Pottery depicting potatoes 2-3 in the tropical highlands 732, 740--1
Powdery scab in the tropical lowlands 768, 769, 778
control of 423, 424, 426 tuber infection during growth 415
disease in store 419 and wilting 414
geographical distribution 408 Pseudo stem nematode (Neotylenchus
risk determination 431 vigissi) 440
seed tuber decay 405 Psyllidae 484
in soil 404 Psylloides plana 477
symptoms 408 Pulsar prototype harvester 587, 589
tuber infection during growth 415 Pungo (cv.), mineral nutrition 166
Pratylenchus (Root lesion nematodes) Purple Congo (cv.), tuber quality 515
control of 455 Pyralidae 489
diagnosis 462 Pyrallis farinalis 477
geographical distribution 440 Pyrethroids 427
and roots 441, 442 Pyridoxin 512, 705
Pratylenchus brachyurus 442-3, 455 Pythium ultimum
Pratylenchus coffeae 433 geographical distribution 408
Pratylenchus penetrans post-lifting disease 417
control of 456 symptoms 408
geographical distribution 439 tuber infection at lifting 417
interaction with fungi 414 Quality
interaction with other organisms 452 breeding considerations 354, 355-{i
resistance to 457 cooked potato 533-9
and roots 442 and fertilizer use 205-{i
and yield losses 448 during harvest 578
Pratylenchus pratensis 443 and irradiation 548
Pratylenchus solanacearum 452 losses in storage 608
Pratylenchus thornei 452 potatoes for processing 54:0-8
Premnotrypes (Andean potato weevils) 476, statutory standards 578
477, 478, 487 and water supply 236-41
Prescription for New Potatoes and Ware and weeds 375
Potatoes 578 Quarantine 404, 428
Pressure bruising 581 Quintozene (PCNB) 426
Prevention of Spread of Pests (Seed Quizalofop 393
Potatoes) (Great Britain)
Order 1974275 Radiation interception
Prochloray 427 and carbon dioxide 817
Producers, number of 873 and crop dry weight 819
Index 899
and dry matter yield 169-76, 180, 522, to frost 46
82(}-1 to pests 45-6
efficiency of conversion 836--7 and wound periderm 111
and growth 816--7 Respiration 636--51
and leaf area index 220, 822, 824--6, 833-4, chemicals, influence of 647-50
847 and cultivar type 651
in the tropics 737 gas exchange 638-42
and tuber yield 168 gelilc:ral 636--8
see also Gamma irradiation and handling 645-7
Radioactive irradiation 673-4 heat production 642-3
Radiocarbon dating of potatoes 3 and irradiation 548, 647
Rainfall reconditioning 644--5
effect on plant nutrients 20(}-3 and sprout growth 626, 651, 676
effect on wild hybrids 49 and storage period 643-4
Raphanus raphanistrum (Wild radish) 378 and storage temperature 612, 622, 642,
Recirculation of air in storage 635 643
Reconditioning and respiration 644--5 and sugar accumulation 544
Record (cv.) and tuber maturity 650
amino acid content 699 and tuber size 651
carbohydrates 692 and wounding 647
discolouration 520 Rest period 128-9, 658
in Great Britain 813 Rhigopsidius tucumanus 487
growth cracking 516--17 Rhizoctonia
internal bruising 529 and haulm pulling 585
and leaf growth 830 and nematicides 426
plant density 301, 319, 325-7 seed tubers 427
processing quality 515, 543, 545-6, 548 Rhizoctonia solani
root growth 225 control of 423, 424
seed tubers 240, 241, 255, 274, 280 and crop frequency 423
tissue culture 141 and cut seed 409
and water supply 240, 241, 835, 838-42, and early lifting 424
851, 869, 870 geographical distribution 408
wound healing 653 and humidity 419
yield 853 infection of eyes and early harvesting
Red Craig's Royal (cv.) 274
development 119 and potato sickness 452
and growth 827 and skin blackening 416
and mechanical damage 576 spread 420
seed tubers 268, 281 and stems 310
wound healing 653 survival 419
Red Pontiac (cv.), organic acid content 707 symptoms 408
Red Salad (cv.), tuber quality 515 Rhopalosiphoninius larysiphon 481
Redshank (Polygonum persicaria) 378, 391 Rhopalosiphum padi 481
Redskin (cv.), wound healing 653 Riboflavin 512, 705, 706
Reducing sugars 540, 541, 546, 547 Rindite 675
and irradiation 548 Ring rot
Reed, common (Phragmites communis) and cut seed 409
376 geographical distribution 406
Refrigeration in storage 619,635 import restrictions 404
Regeneration from roots 69-70 symptoms 406
Renacimiento (cv.), mineral nutrition 166 tuber infection during growth 415
Reniform nematode (Rotylenchulus Roller table research 599-602
reniform is) 444 Rolling of foliage 406, 408
Reproduction Romano (cv.), and growth 853
clonal 337-9 Root depth
methods 48 comparisons 225-7
Resistance and soil water stress 235-6
breeding for 354 Rooted cuttings, as planting material in the
to damage, measurement of 581-2 tropics 771-3
to disease 45-6 Root-knot nematodes, see Meloidogyne
900 Index
Root length water use 218

comparisons 225 Scab, common, see Common scab
density 182, 227 Scarabaeidae 486--7
effect on water flow 233 Sclerotinia sclerotiorum 414, 419
and size of crop canopy 182 geographical distribution 408
Root lesion nematodes, see Pratylenchus symptoms 408
Roots Scottish Centre of Agricultural Engineering
adventitious 68-9 (SCAE) 577, 579, 599
depth, see Root depth Scottish Classification Scheme, entry of
development 67 cultivars 276
distribution and soil water stress 234--6 Scottish Crop Research Institute (SCRI),
growth and functioning 837-44 and breeding 342, 345, 346
length, see Root length Scrobipalpopsis solanivora 477, 487-8, 489
nematodes 441-7 Scrobipalpula absoluta 477, 487-8, 489
regeneration from 69-70 Scuffing 524
shoots, response to water deficits 229 Seed
'size, see Root size certification 271, 272, 275, 277, 278, 405,
structure 67-9 428-9
Root size costs, as proportion of total costs 292
comparisons 225 multiplication 271-8
and leaf area 182 production, future developments 814-15
and water supply 223 production and utilization 278-86
Roseval (cv.), tuber quality 520 rate, see Seed rate
Rotary cultivators 571-2 see also Seed tubers; True potato seed
Rotation of crops 373-4, 396 Seedbed preparation 570--3
effect on disease development in the Seedlings
tropics 769 development 66
as nematode control method 455-{), 460, 461 as weeds 399
Rotylenchulus reniformis (Reniform Seed potatoes, see Seed tubers
nematode) 444 Seed rate
Rotylenchus 442 as measure of plant density 296
Row width 306--8 relationship with plant density and tuber
effect on damage during harvesting 321 yield 300, 302-3, 305--6
RUCIP rules (Rules of Usages of Intra Seed size 247, 273-4
European Trade in Potatoes) 578-9 and silver scurf 321
Rules of Usages of Intra European Trade in and yield 296--7, 305
Potatoes (RUCIP rules) 578-9 Seed tubers
Rumex (Docks) 377 ca,tegories 275
Russet Burbank (cv.) cutting of, see Cut seed
amino acid content 698, 699 damage by disease 405-10
mineral nutrition 166 depth, surveyed variations 310--11
organic acid content 707 disease control, chemical 426--7
respiration 649 dormancy 250--2
root growth 225, 226 floral initiation 257-8
solanine 703 growth and development 247-71
sprouting 670 health 272
tuber quality 511, 535, 545, 549 leaf growth and yield 260
tuber structure 105, 108, 110, 143 morphology 248-50
vitamin content 705 number of, as measure of plant density 296
water soluble vitamins 680 physiological age 260--71
water supply 239 planting time 281
Rust spot, internal 517, 519, 520 size, see Seed size
Rwanda, production 797-8, 804 sprout growth and development 252-8
Rye-grasses 392 stenn development 258--60
storage in diffused light 734--6
Sabina (cv.), organic acid content 706 surface area of, as measure of plant
Saskia (cv.), effect on oxygen concentration 656 density 296
Saturn a (cv.) in the tropical lowlands 771
carbohydrates 685--6, 687, 688, 695, 696 utility in seed production 271
seed tubers 280 see also Seed
Index 901
Senecio vulgaris (Groundsel) 378, 387, 389 symptoms 407
Senescence, premature, causes of 407-8 Sodium, concentration in plants 167
Sethoxydim 381, 392 Soft rots
Shape of tuber 513-4 and cut seed 409
and canning 548 geographical distribution 406
and mechanical damage 526--7 and grading 419
Shatter bruise 581 seed tuber decay 405
Shepody (cv.), and mechanical damage 576 and storage temperature 541, 549
Shistocerca gregaria 477 symptoms 406
Shoots and wilting 414
aerial 70 Soil
apex structure 71-5 analysis of nutrient content 182-5
apical meristem 71, 74, 75-8 application of chemicals to control disease
lateral, effect on yield 70 426
leafy shoot 70 and fertilizer requirements 182
root shoots, response to water deficit 229 moisture, see Soil moisture
Sieglinde (cv.), leaf 84 nitrogen index 185
Sieving 587-8 pH, effect on uptake·of plant nutrients 205
Silver scurf porosity 227
and caraway oil 673 potash, and bruising 581
and cut seed 409 sieving 587-8
and early lifting 424 temperature and hollow heart 518
geographical distribution 407 type, and bruising 581
and heating 425 water deficit 222-3
and humidity 423 water potential 230--6
and seed size 321 water uptake from 230--6
and seed tubers 420, 426, 427 and yield 184
and skin blemishes 405, 514 Soil moisture
in store 419,420 and brown centre 518
symptoms 407 and dry matter content 523
Sinapis arvenis (Charlock) 378 and growth cracking 516
Sirtema (cv.), glycoalkaloid content 532 Solanum (genus) 13
Size of tuber 513 Solanum abacayense 29
and bruising 581 Solanum acaule
and canning 548 and blight resistance 740-1
and dry matter content 524 classification 17, 38, 60
and mechanical damage 526--7 disease resistance 46, 431
statutory standards 578 evolution 55
Sizers 596--9 frost resistance 46, 740
Sizing 595--6 self-pollination 51
electronic, and mechanical damage 526 subsp. acaule 38
Skin subsp. aemulans 38
colour 514-5 subsp. punae 38
strength measurement 582 Solanum acroglossum 28
texture 514 Solanum acroscopicum 29
Skin spot Solanum agrimonifolium 16, 27
and climate 403 Solanum x ajanhuiri
and cut seed 409 classification 17, 36, 59
and early lifting 424 and frost tolerance 740
geographical distribution 407 geographical distribution in South
and heating 425 America 42
and humidity 423 Solanum alandiae 16, 30
and leaves and buds 416 Solanum albicans 17, 38
and potato eyes 295 Solanum albornozii 28
and seed tubers, 405, 420, 426, 427 Solanum ambosinum 30
and stems 310 Solanum andigenum, sprouting 675
and storage temperature 418, 425 Solanum andreanum 30
symptoms 407 Solanum antipoviczii, tuberization in high
Smut 404 temperatures 751
geographical distribution 407 Solanum apurimacense 37
902 Index
Solanum arnezii 15, 24 Solanum x chaucha
Solanum astleyi 15, 25 and breeding 50
Solanum avilesii 30 classification 17, 36, 59
Solanum berthaultii geograpJilical distribution in South
and breeding 741 America 43
classification 16, 30 Solanum chavinense 25
dormancy 660 Solanull,i chiquidenum 31
insect resistance 46, 491-2 Solanum chomatophilum
Solanum boliviense classification 16, 27
classification 15, 25 frost tolerance 740
frost tolerance 740 Solanum circaeifolium 15, 23
Solanum brachistotrichum 15, 19-20 subsp. circaeifolium 23
Solanum brachycarpum 17,40 subsp. quimense 23
Solanum brevicaule Solanum clarum
classification 16, 30 chemotaxonomy 53
disease resistance 464 classification 15, 19
frost tolerance 740 Solanum coelestipetalum 31
Solanum brevidens Solanum colombianum 16, 27
and breeding 51,431 Solanum commersonii
classification 18 and breeding 50
virus resistance 46, 431 chemotaxonomy 54
Solanum bukasovii classification 15, 22
classification 16, 30 heat and cold tolerance 754
frost tolerance 740 tuberization in high temperatures 751
Solanum bulbocastanum subsp. commersonii 22
blight resistance 740 subsp. malmeanum 22-3
chemotaxonomy 54 Solanum x curtilobum
classification 15, 19 and breeding 50
subsp. bulbocastanum 19 classification 17, 37, 60
subsp. dolichophyllum 19 cytology 52
subsp. partitum 19 frost tolerance 740
Solanum cajamarquense 30 geographical distribution in South
Solanum calvescens America 43
and breeding 50 Solanum demissum
classification 22 blight resistance 46, 357-8, 430
Solanum canasense and breeding 51
classification 16, 30 classification 17, 40
frost tolerance 740 cytology 53, 56
Solanum candolleanum 30 frost tolerance 740
Solanum capsicibaccatum 15, 23 So/an,um diemii 37
Solanum cardiophyllum So/anum x doddsii 31
and breeding 50 Solanum x edinense 41n, 50, 52
chemotaxonomy 53, 54 Solanum etuberosum
subsp. cardiophyllum 20 classification 18
subsp. ehrenbergii 20 resistance to leaf roll virus 46
subsp. lanceolatum 20 Solanum fendleri
Solanum chacoense chemotaxonomy 53
bacterial resistance 46 classification 17, 39
and breeding 55 evolution 56
classification 15, 24 subsp. arizonicum 39
cold tolerance 733 subsp. fendleri 39
heat tolerance 758 Solanum fernandizianum 18
insect resistance 46, 491 Solanum flahaultii 27
tuberization in high temperatures Solanum gandarillasii 31
751 Solanum goniocalyx 36
wilt resistance 769 Solanum gourlayi
subsp. chacoense 24 classification 16, 31, 52
subsp. muelleri 24 potato cyst resistance 457
Solanum chancayense 30--1 subsp. gourlayi 31
Index 903
subsp. pachytrichum 31 classification 16, 32
subsp. saltense 31 wilt resistance 769
subsp. vidaurrei 31 subsp. gigantophyllum 32-3
Solanum guerreroense 17, 40-1 subsp. microdontum 32
Solanum hastiforme 25 Solanum mochiquense 33
Solanum hintonii 20 Solanum molinae 37
Solanum hjertingii Solanum morelliforme
browning 531 and breeding 51
classification 17, 39 classification 15, 19
Solanum hondelmanni 16, 31 Solanum moscopanum
Solanum hoopessi 31 classification 16, 27
Solanum hougasii 17, 41 cytology 52
Solanum huancabambense 24 Solanum multidissectum
Solanum humectophilum 32 classification 16, 33
Solanum hygrothermicum, tuberization in frost tolerance 740
high temperatures 751 nematode resistance 457, 458
Solanum hypacrarthrum 28 Solanum multiinterruptum 16, 33
Solanum imite 32 Solanum navaritense 20-1
Solanum incamayoense 32 Solanum neocardenasii
Solanum X indunii 50 classification 16, 33
Solanum infundibuliforme 16, 26 resistance 46
Solanum ingifolium 16, 29 Solanum neorossii 33
Solanum iopetalum 17, 41 Solanum neovalenzuelae 27
Solanum jalcae 28 Solanum nigrum (Black nightshade) 375,
Solanum jamesii 378,388
and breeding 50 Solanum ochranthum 18
chemotaxonomy 53 Solanum okadae 16, 33
classification 15, 20 Solanum olmosense 15, 24
Solanum juglandifolium 18 Solanum oplocense
Solanum X juzepczukii classification 16, 33
and breeding 50 cytology 52
classification 17, 37, 59 Solanum oxycarpum 16,27
frost tolerance 740 Solanum pampasense 33-4
geographical distribution in South Solanum papita 17, 39
America 43 Solanum paucijugum 27
Solanum kesselbrenneri, tuberization in high Solanum paucissectufin 28
temperatures 751 Solanum phureja
Solanum kurtzianum 16, 32 bacterial resistance 46
Solanum laxissimum 27 blight resistance 741
Solanum leptophyes breeding I'Ise 351
ancestor of cultivated potato? 3 classification 17, 36, 57-9
classification 16, 32, 60 disease resistance 431
disease resistance 464 geographical distribution in South
Solanum leptostigma 37 America 42
Solanum lesteri 15, 21-2 heat tolerance 751
Solanum lignicaule 15, 23 Solanum pinnatisectum
Solanum longiconicum 27 chemotaxonomy 53, 54
Solanum lycopersicoides 18 classification 15, 21
Solanum maglia Solanum piurae 16, 28
and breeding 50 Solanum polyadenium 15, 22
classification 16, 29 Solanum polytrichon
Solanum marinasense 32 chemotaxonomy 53
Solanum matehualae 39 classification 17, 39
Solanum medians 16, 32 Solanum raphanifolium
Solanum megistacrolobum classification 25-<i
classification 15, 25, 60 frost tolerance 740
frost resistance 46, 740 Solanum raquialatum 29
Solanum X michoacanum 20 Solanum X sambucinum
Solanum microdontum chemotaxonomy 53, 54
and breeding 49 classification 21
904 Index
Solanum sanctae-rosae nematode resistance 456, 457, 458
classification 15, 26 source 6-7
frost tolerance 740 Solanum tuberosum subsp. tuberosum
potato cyst resistance 457 classification 17, 37, 60
Solanum sandemanii 34 geographical distribution in South
Solanum santolallae America 43
chemotaxonomy 53 nematode resistance 457
classification 16, 27 source 6-7
Solanum schenckii 17, 41 Solanum tuquerrense
Solanum X semidemissum and breeding 51
and breeding 50 classification 16, 28
chemotaxonomy 52 Solanum ugentii 34-5
classification 17, 41 Solan'um urubambae 28
Solanum x setulosistylum 34 Solanum x vallis-mexici
Solanum sitiens 18 and breeding 50
Solanum sogarandinum 26 chemotaxonomy 53
Solanum solisii 28 classification 17, 40
Solanum sparsipilum Solanum venturii
bacterial resistance 46 and breeding 50
classification 16, 34, 60 classification 35
heat tolerance 758 frost tolerance 740
nematode resistance 458 Solanum vernei
wilt resistance 769 and breeding 49
subsp. calcense 34 classification 16, 35
subsp. sparsipilum 34 frost tolerance 740
Solanum spegazzinii nematode resistance 457
classification 16, 34 subsp. ballsii 35
potato cyst resistance 457 subsp. vernei 35
Solanum stenophyllidium 21 Solanum verrucosum
Solanum stenotomum blight resistance 740
evolution 57, 60 chemotaxonomy 53
geographical distribution in South classification 16, 35
America 42 evolution 57
Solanum stoloniferum Solanum x viirsooi 50
chemotaxonomy 53 Solanum violaceimarmoratum 16, 28
classification 17, 39-40 Solanum virgultorum 35
evolution 55, 56 Solanum weberbaueri 35
insect resistance 46 Solanum ytmgasense 15, 25
virus resistance 46, 431 Solanum X zudaniense 25
subsp. moreliae 40 Somaclonal variation 143, 363
subsp. stoloniferum 40 Somatic hybrids 363-4
Solanum subpanduratum 27-8 Somatic tissues 138
Solanum X sucrense 16, 34 Sonchus arvensis (Perennial sowthistle) 377
Solanum tarijense 15, 24-5 South America
Solanum tarnii 15, 21 consumption 801
Solanum toralapanum distribution in 41-3, 45
classification 15, 26 imports and exports 800
frost tolerance 740 source of the potato 1-2
Solanum trifidum Southeast Asian Program for Potato Research
chemotaxonomy 53 and Development (SAPRAD) 778
classification 15, 21 Sowthistle, perennial (Sonchus arvensis) 377
Solanum x trigalense 25 Spacing 577
Solanum tuberosum Spacing of seed
classification 37, 59 effect on yield 297-8
sweetening 545 irregularities 309-11
Solanum tuberosum subsp. andigena row width 306-8
classification 17, 37, 59-60 Species concepts 46-8
geographical distribution in South Specific gravity 521-2, 537
America 43 and bruising 581
Index 905
Spindle tuber 412, 425 and growth 322
Spindle tuber viroid 406 as measure of plant density 312
Splits 580 and number of tubers 312-17
Spodoptera exigua 477 and nutrient uptake 322
Spongospora subterranea and quality and saleability 320-1
geographical distribution 408 relationship with seed rate and tuber yield
interaction with other organisms 419, 452 300,302-4
survival and spread 419, 420 and yields 317-20
symptoms 408 Stem-end blackening 533-6
tuber infection during growth 415, 416 Stem nematode, see Ditylenchus dipsaci
virus-transmitting 441 Stem rot, causes 407-8
Spraing disease 419, 440 Stems
Sprout growth aerial, anatomy 78-81
and development 252-8 clumping 310-11
in diffused-light storage 734---6 decay, causes 407
and irradiation 548 density, see Stem density
and respiration 651 development 258-60
in storage 626, 664-5, 676-7 and length of growing season 318
seealso Sprouting; Sprouting period and length of sprouting period 298
Sprouting as measure of plant density 301-6
changes in bud meristem 130-3 nematodes 440
changes in tuber 129-30 optimum densities 319
control in storage 664-5 types 298
and evaporation 629 water potential 232-3, 236
included in storage, effect on air flow 620-1 Stenotycha 477
inhibition, see Sprout inhibition Sterility 48-51
internal 670 Sthenaridea pusiUa 477
period, see Sprouting period Stina (cv.), organic acid content 706
see also Sprout growth Stolons
Sprouting period anatomy 88-92
effect on stems 298 apex 87-8
effect on tuber yield 270 development 93-6
Sprout inhibition formation 86--7
and respiration 649-50 Stoma, responses to water deficits 228-30
in storage 665-74 Storage
Sprouts allU blackening after cooking 535-6
growth, see Sprout growth canning potatoes 611
inhibition, see Sprout inhibition conditions, see Storage conditions
number of, as measure of plant density and evaporation 629-32
295 industrial potatoes 611
stimulation in storage 674---6 losses 608-9
see also Sprouting period, see Storage period
Sri Lanka, experiments in 744 practices 871-2
Stalk break, 414 seed crops 285
geographical distribution 408 seed potatoes 609-10
symptoms 408 seed potatoes in diffused light 734
Standards, potato quality 578-9 temperature, see Temperature
Staple foods 795 ware potatoes for consumption 610
Starch 611 ware potatoes for processing 610-11
content 108-10, 508, 522 Storage conditions
Statistics, production in the tropics 729-31 effect on sprout growth 660-3
Stellaria media (Common chickweed) 377, effect on wound healing 655-7
378 Storage period
Stem canker, 424, 426 and bruising 581
geographical distribution 408 and respiration 643-4
and plant spacing 410 Strepomyces scabies
symptoms 408 affected by fertilizers? 206
Stem density and cut seed 409
and damage during harvesting 321 geographical distribution 408
economic considerations 323-30 symptoms 408
906 Index
Stresses in the tropical lowlands and dry matter content 522
bacterial diseases 768-70 and evaporation 612
early blight 768 and flowering 134
high temperature 745-58 and formation of wound periderm 111
mineral nutrition and toxicities 763-7 and glycoalkaloids 533
water 758--63 and greening 515-16
Structure industrial potatoes in store 611
potato plant 65 and leaf form 81
root 67-9 and mechanical damage 525
Stubby root nematodes, see Paratrichodorus; and nematode activity 458
Trichodorus and number of sprouts 253
Stunting, causes 406, 411-14 and physiological age 265
Suberization 111-12, 652, 654, 655 in a potato stack 614-16, 617-21
Sucrose in tuber formation 124 and respiration 612, 622, 642
Sugar content 508 seed potatoes in stores 267, 609-10
and irradiation 548 and shoot apical meristem 71
and respiration 544 of a single tuber 613-14
and water deficits 239 and sprout growth 250, 255, 257, 263-5,
Sulphur 426 660-1,664,735
content 165-7 and sprout length 256
Sulphuric acid 396 and starch content 108
Super Elite (basic seed grade) 275-7 in the tropical lowlands 745-58
Superior (cv.) and tuber formation 113-15
development 139 ware potatoes for consumption 610
organic acid content 707 ware potatoes for processing 610-11
vitamin content 707 and wound healing 655
Surprise (cv.), and water use 217 Terbuthylazine 390
Survival of pathogens 419-20 Terbutryn 387-8, 389-90
Sutton Bridge Experimental Station 599 Terpenoids, changes during storage 708
Sweetening 540--4 Tetranychidae 490
senescent 545-7 Tetraploids, cultivated species 37, 43
Sweetening in storage Texture 536-9
canning potatoes 611 Thiabendazole 427
industrial potatoes 611 Thiamin 512, 705, 706
seed potatoes 609 Thistles 376
ware potatoes for consumption 610 Thlaspi arvense (Field pennycress) 378, 391
ware potatoes for processing 611 Thripidae 490
Symmetrischema plaesieosema 477, 487, 489 Thrips, 478,490
Symptoms of diseases 406-8 in the tropics 767
Synchytrium endobioticum Thrips palmi 477, 490
geographical distribution 408 Thysanoptera 490
and moisture 409, 415 Tilth requirement 570-1
spread 420 Tissue culture 138-43
survival 419 Tobacco flea beetle (Epitrix hirtipennis) 486
symptoms 408 Tobacco rattle virus (TRV) 406, 420, 440-1,
459,466
Tasso (cv.), tuber quality 536 Tolclofos-methyl 427
Taxonomy 46--8, 53-4 Toxicities in the tropical lowlands 763
TeA 380,385 Transpiration 214-16, 228
Tecnazene 671 efficiency 215-18
Telone 426 Transport, field to store 592-3
Temperate zones, potato production 804 Trapping of insects 482-3, 488
Temperature Trichodnridae 441
ambient 616-17 Trichodorus (Stubby root nematodes)
and Andean potatoes grown in Europe 7 control of 455, 460, 466
and brown centre 518 geographical distribution 439, 440
and bruising 528, 581 and growth 442
canning potatoes 611 interaction with other organisms 416, 419,
and cell layers in the periderm 110 440-1
and dimensions of stored mass 617-21 and roots 441
Index 907
and sugar beet 466 Two-stage harvesting 590-2
Trietazine 388, 390 Tylenchorhynchus 441, 442
Trinidad, experiments in 74~ Ukama (cv.), organic acid content 706
Tripleurospermum inodorum (Mayweeds) Ulster Chieftain (cv.)
377 and growth 844-5
Triploids, cultivated species 36-7, 43 leafy shoot 70, 71
Tropics Ulster Prince (cv.)
future production 780 diseases 421
highland production 731--41 plant density 320
lowland production 742-79 Ulster Sceptre (cv.), tuber quality 520
production statistics 729-31 United Kingdom
True potato seed (TPS) 9, 286-7, 351, 352 added value 807-8
as planting material in the tropics 774---6 consumption 507, 806-7
Tuber-bearing plants, distinguishing features import restrictions 404
14 marketing 812-14
Tuber flea beetle (Epitrix tuberis) 486 potato production 797-8, 799
Tuberin 510 production economics 808-12
Tuberinin 510 seed production 810-12
Tuberization in culture 118-20 seed production, choice of area 271
Tuber moth, see Phthorimaea operculella seed trade 800
Tuberosa United States Department of Agriculture
classification 16-17, 29-37 462
crossability 51 United States of America, potato production
cultivated 17, 36-7 797-8
evolution 55, 57 Up-to-Date (cv.)
geographical distribution in North and plant density 315
Central America 44 tuber structure 112
geographical distribution in South Urea formaldehyde 426
America 45 USA, see United States of America
wild 16, 29-35 USSR
Tuber-rot nematode, see Ditylenchus consumption 801
destructor imports and exports 800
Tubers production 796-7
changes at sprouting 129-30 Utilization 799-801
characteristics, see Cultivar characteristics
diseases at retail outlets 417-18 Vanessa (cv.)
diseases in store 418-19 and growth 823, 825-6
diseases during transport 417 root growth 225
experiments in culture 118-19 Varieties, see Cultivar characteristics
formation, factors controlling 113-28 Ventilation
formation and development 96-7, 98-113 and evaporation 629-31
infection during growth 414-16 in storage 619, 625-6
infection at lifting 417 Veronica hederifolia (Ivy-leafed speedwell)
initiation 844-5 378
morphology 97-8, 354 Verticillium
nematodes 440-1 control of 426
number of, effect of plant density 312-17 interaction with fungi 441
pigmentation 74 and irrigation 424
quality, see Quality risk determination 431
seed, see Seed tubers and wilting 413, 414
size, see Tuber size Verticillium albo-atrum
structure 105-13 control of 423
symptoms of disease 406-8 geographical distribution 408
yield, relationship with seed rate and plant interaction with other organisms 452
density 300, 302-6 symptoms 408
Tuber size and wilting 414
effect of fertilizer use 206 Verticillium dahliae
and respiration 651 control of 423
see also Seed size geographical distribution 408
Tuberworm, see Phthorimaea operculella interaction with other organisms 442, 452
908 Index
Trichodorus cont. and growth 214--19
and senescence 464 loss in storage 627-36
survival 419 potential 228-36
symptoms 408 and root growth 223-28
and wilting 414 and tuber quality 236--41
Verticillium lecanii 494 and tuber quantity 238
Verticillium nubilum 409 uptake from soil 230-6
Verticillium wilt 404,409,413,414,452,455 and yield 219-23
geographical distribution 408 Water deficits
symptoms 408 cause of tuber defects 239-40
Viola arvensis (Field pansy) 388 effects on sugar contents 239
Violaxanthin 515 in storage 627-36
Violet root rot stresses in the tropical lowlands 758-63
geographical distribution 407 see also Drought
symptoms 407 Water potential 228-36
tuber infection during growth 415 Water use efficiency (WUE), in the tropical
Virus elimination lowlands 760-2
by culture 78 Water vapour pressure deficit (WVPD)
effect of heat treatment 77-8 627-36
effect of size of explant in meristem Watery wound rot
culture 76, 77 geographical distribution 408
Viruses symptoms 408
diseases 272 tuber infection at lifting 417
elimination, see Virus elimination Wavelength of light, and sprout growth 735
geographical distribution 406 Weeds
in ~he shoot apex 77 annual 377-8
symptoms 407-9 and changes in cultural pattern 373-4
transmitted by aphids 421, 480 control of 379-94
see also Potato viruses effects of 374-6
Virus Tested Stem Cuttings (VTSC) 271, perennial 376--7
275-7 potato as cleaning crop 374
Visual assessment in breeding 362 potatoes as weeds 396--9
Vitamin B6 705, 706 Weight losses in storage 608
Vitamin C 510-11 Western flea beetle (Epitrix subcrinita) 486
Vitamins West Germany, see Germany
changes during storage 704-6 White grubs 486--7
content 510-13 White potato cyst nematode, see Globodera
losses in cooking and processing 511-12,803 pal/ida
losses in storage 511, 704-6 White Rose (cv.)
Voluntce.T cereals 378 dormancy 131
Volunteer oilseed rape 378, 388, 393 and growth 838
Volunteer plants 404, 423, 432 mineral nutrition 166
Volunteer potatoes 377,397-9 tuber development 120
effect on virus levels 396 tuber structure 108
VTSC (Virus Tested Stem Cuttings) 271, Whitewash 427
275-'7 Wild oats (Avena fatua) 378, 381, 392
Wild potatoes, geographical distribution and
Ware Prescription 578, 595 habitats 44-6
Ware production, future developments 814 Wild radish (Raphanus raphanistrum) 378
Wart Wilja (cv.)
certified seed 405 and intercepted radiation 820
control of 423 mineral nutrition 168-76, 181-2,202
and cut seed 409 seed tubers 270, 280
geographical distribution 408 tuber quality 514
and humidity 419, 429, 430 water use 218
resistance to 429 Wilting
survival of pathogens 404 ca uses 406--7, 413-14
symptoms 408 see also Bacterial wilt; VerticiHium wilt
Water Windrowing 572-3
deficits, see Water deficits and damage 591
Index 909
and mechanical damage 526 and depth of planting 575
Wireworms 476, 486, 492 and diseased plants 410
Witches' broom 406, 411-12 and farmyard manure 196
Wound healing geographical differences 854
changes in tissue 651-3 and herbicides 380
and chemical treatments 656 influence of lateral shoots 70
and cultivars 653 and irrigation 866--70
and humidity 655 and lime use 766
and irradiation 548 losses caused by disease 404
and maturity 653 measurement of 478
and oxygen and carbon dioxide 656--7 and mechanical weed control 379
periderm formation 111-13, 652 and nitrogen 168-76, 764-5
and physiological state of tuber 653-5 and nitrogen reduction 194, 849-50
requirements for 609 and planting date 856--7, 858
and storage conditions 655-7 and potato cyst nematodes 458
and temperature 655 and root-feeding nematode& 447
and tuber characteristics 653-5 and seed rate 304--6
Wounds and stem density 317-18
effect on respiration 647 and water supply 219-23
healing, see Wound healing and weeds 374-5
types 653 Yungasensa
classification 15, 24-5
Xiphinema 442 evolution 55, 56
geographical distribution in South
Yellow dwarf virus 406, 411, 416, 421 America 45
Yields
breeding for 354 Zinc, content 165-7
and choice of cultivar 853--6 Zineb 428

The Potato Crop

Uploaded by

Copyright:

Available Formats

You might also like

The Potato Crop

Uploaded by

Document Information

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

The Potato Crop

Uploaded by

Copyright:

Available Formats

THE POTATO CROP

World Crop Series

fiji Springer-Science+Business Media, B.V.

Chapter 13 © 1992 B.D. Witney and D.C. McRae

3 Structure and development of the potato plant 65

4 Mineral nutrition 162

5 Water relations and growth of potatoes 214

6 Seed tuber production and management 247

6.5 True seed 286

Part 2 Insect pests 476

12 Tuber quality 507

13 Mechanization of crop production and handling operations 570

14 The physics and physiology of storage 608

15 Potato production in the tre,pics 728

15.5 Tropical lowland potato production - two practical

17 Principles of agronomy and their application in the potato industry 816

History of the potato

1.2 THE STATUS OF THE POTATO IN SOUTH AMERICA

1.3 EVIDENCE FOR DOMESTICATION - ARCHAEOLOGICAL DATA

The most striking archaeological evidence for the antiquity of potato

1.4 THE POTATO AT THE TIME OF THE SPANISH CONQUEST

1.5 HISTORICAL AND LINGUISTIC EVIDENCE

1.6 THE INTRODUCTION OF THE POTATO INTO EUROPE

1.7 THE NATURE AND EXACT SOURCE OF THE EARLY EUROPEAN

I'SOLANvMcuberofumcfculentum:caulem ha- admodlll1l figura nonn lk Matthiolo & huicli-

1.8 THE SPREAD OF THE POTATO INTO OTHER PARTS OF THE

Bauhin, C. (1620) Prodromus Theatri Botanici, Frankfurt-on-Main.

Biosystematics of the potato

The potato of commerce belongs to a single species, Solanum tuberosum

2.2 BRIEF OUTLINE OF POTATO CLASSIFICATION

The genus Solanum, to which the cultivated potato belongs, is an extremely

2.2.1 Classification of potato species

Series II JUGLANDIFOLIA (Rydb.) Hawkes

Subsection POTATOE G. Don.

Series II BULBOCASTANA (Rydb.) Hawkes

Series III PINNATISECTA (Rydb.) Hawkes

Series IV POLYADENIA Buk.

Series V COMMERSONIANA Buk.

Series VI CIRCAEIFOLIA Hawkes

Series VII LIGNICAULIA Hawkes

Series VIII OLMOSIANA Ochoa

Series IX YUNGASENSA Corr.

Series X MEGISTACROLOBA Card. et Hawkes

Series XI CUNEOALAT A Hawkes

Series XII CONICIBACCAT A Bitt.

Series XIII PIURANA Hawkes

(Note: the other species in this series, S. raquialatum Ochoa, is not

Series XV MAGLIA Bitt.

Series XVI TUBEROSA (Rydb.) Hawkes

Note: The former species, S. goniocalyx, has now been placed as a

Series XVIII LONGIPEDICELLATA Buk.

Series XIX DEMISSA Buk.

2.3 DISTRIBUTION AND ECOLOGY OF POTATOES

The common cultivated potato, S. tuberosum, through the efforts of

Figure 2.1 Distribution of diploid cultivated potatoes in South America.

The frost-resistant species S. x ajanhuiri, S. x juzepczukii and S. x

I ~~~[; I "0 400

Figure 2.2 Distribution of triploid, pentaploid and tetraploid cultivated potatoes

II Bulbocastana and IV Po~adenia