Effects of Temperature and Duration of Leaf Wetness On Infection of Celery by Septoria Apiicola and Cultivar Screening For Partial Resistance

EI'I'ICTS 01' 'fDIPEO'1'UU BD DURATIOK or LDI' R'l'nSS
OK IUECTION 01' CEUItY BY SEP'1'ORIA UIICOLl.,
AND
CULTIv.aR SCREIKING l'Olt PARTIAL RESISTANCE
by
Danielle Mathieu CV
A thesis submitted to the Faculty of Graduate Studies and

Research in partial fulfilment of the requirements for the
degree of Master of Science.
Department of Plant Science 27 Septernber, 1991

Macdonald College
McGjll University
(
r
Montreal
ii
(
SHORT TITLB
INI'aCTION NODaL rOR.!.. UIICOLA
ABD RKSISTARCa SCR&aHIHG
Danielle Mathieu~
iii
1
~ORBWORD
There are four parts to this thesis. The first part

introduces the problem, states the goals of this research
and describes the general approach followed to achieve them.
Parts two and three are presented as complete manuscripts
that cover the entire research project. Part four is a
general discussion and conclusion.
The thesis format has been approved by the Faculty of

Graduate Studies and Research of McGill University and
follows the conditions outlined in the "Guideline Concerning
Thesis Preparation", section 7, "Manuscripts of authorship"
which are as follows:
"The candidate has the option, subject to the approval

of the Department, of including as part of the thesis
the text, or duplicated published text (see below), of
an original paper, or papers. In this case the thesis
must still conforrn to aIl other requirements explained
in Guidelines Concerning Thesis Preparation.
Additional rnaterial (procedural and design data as
weIl as descriptions of equipment) must be provided in
sufficient detail (e.g. in appendices) to allow a
clear and precise judgement to be made of the
importance and originality of the research reported.
The thesis should be more than a mere collection of
manuscripts published or to be published. It must
include a general abstract, a full introduction and
literature review and a final overall conclusion.
Connecting texts which provide logical bridges between
different manuscripts are usually desirable in the
interest of cohesion.
c It is acceptable for theses ta include as chapters
authentic copies of papers already published, provided
iv
these are duplicated cle~rly on reguLation thesis

stationery and bound as an integral part of the
thesis. Photographs or other materials which do not
duplicate well must be included in th~ir original
form. In such instances, connecting \~xts are
~atory and supplementary explanatory material is
almost always necessary.
The inclusion of manuscripts co-authorel'/' by the
candidate and others is acceptable but the candidate
is required to make an explicit statemer.t on who
contributed to su ch work and to what extent, and
supervisors must attest to the accuracy uf the claims,
e.g. before the Oral Committee. Since th~ task of the
Examiners is made more difficult in these cases, it is
in the candidate's interest to make the
responsibilities of authors perfectly clear.
Candidates following this option must inform the
Department before it submits the thesis for review."
(~
All the work reported here was the responsibility of the
candidate who conducted it under the supervision of Dr. A.C.
Kushalappa, Associate professor of the Plant Science
Department, Macdonald College of McGill University. The two
manuscripts will be submitted in the appropriate format to
Phytopathology or Plant Disease.
(
v
ABSTRACT
M. Sc. Danielle Mathieu Plant Science

zrrZCTS or TEMPZRATtJRI: AND TU DORATION or LDI' 1fI:'1'HI:SS
ON TU l:NJ'EC'l'ION 01' C&L&l\Y BY S&PTOl\IA APIICOLA, AND
CULTIVAR SCREZHING rOR PARTIAL RESISTANCE
Growth chamber studies were condllcted with the celery

cultivar Florida-683 to deterrnine the effects of ternperature
(10, 15, 20 and 30 OC) during wet periods (12, 24, 48, 72
and 96 h) on the epidemiology of Septoria blight. Results
show that the number of lesions increased with increasing
temperatures over the range of wet periods except at 30 oC,
..... where their nurnber decreased with increasing wetness
duration. The highest number of lesions was recorded at 25
oC, 72 and 96 h wetness. A polynomial regression equation
with nine terms, Adj-R 2 =74.8, was fitted to the arcsine
transformation of the proportion of the maximum number of
lesions. Cluster analysis of the predicted values yielded
four disease severity indices (DS1) ranging from very low
(1) to very high (4) infection probabilities. Cultivars
were evaluated for partial resistance under field and
greenhouse conditions. In the field ranking was based on
cluster analysis of the standard area under the disease
progress curve (SAUDPC) for intervals between sampling
dates. In the greenhouse, the cultivars were evaluated on
the basis of their response relative to five components of
vi
partial resistance: the SAUDPC, mean lesion area (MLA) ,
pycnidial density (PCD) , spore density (SPD), and the
latency period defined as the time from inoculation to 50 %
and 75 % disease (T so and T75 ) . In the greenhouse, overall
ranking was based on cluster and principal component
analysis of responses to SAUDPC, MLA, peD and SPD. Tso and
T75 were not significant. Three cultivars, Golden Plume,
Superdora and Summit, were rated as moderately resistant in
both field and greenhouse trials. The others ranged from
moderately susceptible to very susceptible. Florida 683
used to develop the infection model ranked among susceptible
cultivars.
(
"
vii
M. Sc. Danielle Mathieu Plant Science
I:I'I'E'1'S DI LA 'l'DCPERA'1'UU ET DI LA
DUUI DS LA MOUILLURE SOR L' INI'ECTION DU CtLl:RI PAR LI:
CHAMPIGNON SI:P'1'ORIA APIICOLA, 1:'1'
ÉVALUATION DE CUL'1'I~ POUR LA RtSIS'1'ANCE PAR'1'II:LLE
Des expériences à atmosphère contrôlée avec le cultivar de
céleri Florida-683 ont été effectuées afin de déterminer les
effets de la température (10, 15, 20, 25 et 30 OC) au cours
de diverses durées de mouillure (12, 24, 48, 72 et 96 h) sur
l'ép~démiologie de la brûlure septorienne. Les résultats
indiquent que le nombre de lésions augmente avec la durée de
la mouillure à toutes les températures sauf à 30 oC où la
relation inverse est observée. Le nombre maximum de lésions
est apparu à 25 oC, 72 et 96 hr de mouillure. La variable
indépendante (transformation arcsin de la proportion du
nombre maximum de lésions) est adéquatement décrite par un
modèle de régression polynomiale à neuf variables (R 2-
Ajusté=74.8). Une analyse par groupements hyérarchiques des
valeurs prédites a réduit celles-ci à quatre niveaux ou
indices de sévérité de la maladie (ISM) correspondant à des
probabilités d'infection variant de très faibles (1) à très
fortes (4). Des cultivars de céleri furent évalués afin de
déterminer leurs niveaux de résistance au champ et sous
serre. Au champ, l'aire sous la courbe de progression de la

viii
c maladie standardisée (SAUDPC) a servi de critère
d'évaluation. Les niveaux de résistance au champ ont été
charactérisés à l'aide d'une analyse de groupements
hyérarchiques des SAUDPC pour les ~ntervalles entre les
dates d'échantillonnaa p • Les critères d'évaluation pour les
deux essais sous serre étaient l'SAUDPC, la surface moyenne
des lésions (MLA), la densité des pycnides (PCD) , la densité
des spores (SPD) , et les intervalles 0-50% et 0-75%
d'apparition des lésions (T~ et T7s ) depuis l'inoculation.
Les analyses par groupements et ordination ont permis de
charactériser le niveau de résistance des cultivars pour
l'ensemblp. des paramètres suivants: SAUDPC, MLA, PCD, et
SPD. Les cultivars n'ont présenté aucune différences
( significatives en ce qui a trait aux Tso et T7s ' Trois
cultivars, Golden Plume, Superdora et Summit, se sont
systématiquement montrés moyennement résistants au cours des
trois essais. Les autres cultivars se sont montrés
modérément susceptibles à trèes susceptibles. Le cultivar
Florida 683 utilisé pour l'élaboration du modèle d'infection
a été jugé susceptible.
(
il<
ACICNOWLJ:DGMlI:N'1'S
l am very grateful to my advisor, Dr. Ajjamada C.

Kushalappa, for his advice and enccuLagement throughout this
project.
Others whose contribution was greatly appreciated are Dr. M.

A. Fanous and Dr. M. Waterway from the Plant Science
Department, and Dr. R.I. Cue, from the Department of Animal
Science, for their assistance with statistical analyses. My
gratitude goes also to M. Pikarnegar and G. Rimmer for their
cooperation and skills in maintaining our eJcctronic
equipment in good working condition; to H. Rimmer for help
in photography but also for her kindness and encouragementi
C. Rafuse for her help and patience in the greenhousei Dr.
D. Donnely for letting us use her image analyzer, Dr. R.D.
Reeleder and Dr. T. Paulitz for their suggestions, advice
and borrowed equipment.
Special thanks to the workers of the Agriculture Canada

experimental farm and of les serres Louise Lefort of Ste-
Clothilde, to my field assistant, France Leduc, and my lab
assistant, Catherine Jaynes, for their good work and
cheerfulness .
.....
x
Finally may deepest appreciation goes to my colleagues from

the epidemiology lab and graduate students from the Plant
Science Department for their friendship, support and
encouragement.
This project has been made possible through a scholarship

from the Fonds pour la fomration de chercheurs et l'aide à
la recherche (F.C.A.R.).
(
xi
:OBD l CATION
À toi, Lionel, pour m'avoir enseigné que la connaissance

s'acquiert à tout âge.
A mes enfants, Robert et Tobi, à mes soeurs et surtout à

toi, Claude, sans qui ce projet ne se serait concrétisé,
merci ~e votre compréhension et de votre soutien.
xii
'l'DLI or CONTENTS
rOU1fORD iii
US'J.'RACT ........... . ... . ... v
· ·· · ··· ···· vii

ACDOWLBDGMl:NTS · ··· ··· · · ·· · ··· ix
DEDICATION · ··· ··· · · ·· ·.. xi
'l'ABLI or CONTENTS
· ··· ··· · · ·· · ··· xii
LIST or 'l'ABUS · · · · · · · · · · · xiv
LIST or rIGURES .
· ·· · · ··· · · · · ··· xv
ABBRZVIATIONS USED . · ··· ···· · ·· xvii
1 GI:NJ:RAL INTRODUCTION AND LI'1'I:RATOU OVIn 1
1.1 GENERAL INTRODUCTION . . . . . . . . 1
1.2 REVIEW OF LITERATURE . . . . . . . . 5
( 1.2.1 Crop - Importance and Value . . .
1.2.2 Disease - Historical Perspective
5
5
1.2.3 The Pathogen . . . . . . . . . . 6
1.2.4 Symptoms and Etiology . . . . . . . . . 7
1.2.5 Epidemiology of Septoria Blight . 8
1.2.6 Control of Septoria Blight 9
1.2.7 Disease Resistance in Crops . . 11
1.2.8 Disease Forecasting . . . . . . . . 15
1.2.9 Multivariate Statistical Methods in
Epidemiological Studies . . . . . . . 17
LITERATURE CITED . . . . . . . . . . . . . . . . . . 22
2 ErrEeTS OF TEMPERA'1'UlUI: .AND LDI" 1IJ:'1'NJ:SS DURA.'1'ION
ON 'l'BI: INFECTION 01' CZLI!RY BY SI:P'1'ORIA UIICOLA. 29
2.1 ABSTRACT........ ....... 29
2.2 INTRODUCTION............... 31
2.3 MATERIALS AND METHODS . . . . . . . 34
2.3.1 Production of Plants • . . 34
2.3.2 Inoculum Production. . . • . . . . . . 35
2.3.3 Treatments and Inoculation . . . . . . . . 37
2.3.4 Data Analysis and Regression Model Selection 39
2.4 RESULTS . . . . . . . 43
2.6 DISCUSSION ........... . •. 49
2.7 CONCLUSION.. . . . . . . . . . . . 59
I.ITERATURE CITED . . . . . • . . . . . . . 61
(
" PREFACE '1'0 CBAPTER 3 64
xiii
.1, 3 rIZLD AND GREBNBOUSB ZVALOATIOH or CBLBRY CULTIVARS
rOR PARTIAL RESISTANCB Ta SEP'1'ORIA BLIGB'l'
. · ····· ·
ABSTRACT . · · · 65
3.1
· ··· 65
3.2
· ··· · · · ·· ··· · · ·· ·
INTRODUCTION 67
3.3 MATERIALS AND METHODS
····
· · ···· · · ··
3.3.1 Selection of cultivars
69
69
··· · ·
3.3.2 Field trial
· ·· ·
· 69
·
Transplant production
· · ·
Field plot establishment and layout· · · ·
· · ·
69
70
· · · ·· · ···
Field inoculation · 71
Disease assessment
·
3.3.3 Greenhouse trials ··· · · ·· 71
· · · · ·· · · · ·
· 82
·· · · ·· · ···
Plant Production · 82
·
Inoculation · · ·
· ·
Evaluation of components of resistance
· · 83
84
Disease estimation (SAUDl?C)
· ·· · 84
Mean les ion area (MLA)
··· · ·· ·
· 84
· · · ··· · · ·· ·
Spore density (SPD) · 85
· · · ·· · · · ··
Pycnidial density (PCD)
Spores per pycnidium (SPPC)
85
· ·· · · · ·
Latency period (T so and T7s )
86
· ··· 86
3.3.4 Data analysis
··· · ··· · 87
Field data · · ·
· · ·· · ···
· 87
...
Greenhouse data
·· · · · · ·· 87
.,.. 3.4 RESULTS
· ·
· · · · · · ·
· · · 88
...... 3.4.1 Field trial
·
3.4.2 Greenhouse trials· ·· ·· ·
· · 88
91
SAUDPC . · · ·
· · · · · · · · ·
· 99
· · · ··· ·
Mean lesion area (MLA)
Pycnidial density (PCD)
· 99
· ·· · · · · 100
Spore density (SPD)
· ·· · ·· · 108
· ·
Density of spores per pycnidium (SPPC) · 108
·· ······
Latency period (T so and T7S )
3.4.3 Multivariate analyses.
· 109
110
3.5 DISCUSSION AND CONCLUSION
·· · · ·
· · 121
LITERATURE CITED ·
· ·· · ·
· · · · · ·· ·
· 127
. GBNERAL DISCUSSION AND CONCLUSION · · ·
····· 130
LITERATURE CITED · ·· · · · · 137
APPBNDIX . . · ·· · · · · ·· · ··· · · · 139

xiv
LIST or TABLES
Tabl. 2.1. Parameter cstimates of the regression

equation to describe the effects of temperatures and
leaf wetness duration on infection levels, including
coefficients of determination for transformed values
(R; unadjusted, and R2a adjusted for degrees of
freedorn) and untransformed values (Rt2 adjusted for
degrees of freedom). . . . . . . . . . . . . . . . . 46
Tabl. 2.2. Effects of temperature and leaf wetness
duration during the infection period on the number of
days between inoculation and appearance of the first
lesions. ..................... 58
Tabl. 3.1. Partial correlation coefficients among
components of partial resistance for greenhouse trial
1, Surnrner, 1989. ................ 101
Tabl. 3.2. Partial correlation coefficients among
components of partial resistance for greenhouse trial
2, Winter, 1990. ................ 102
Tabl.3.3. Kendall's tau rank correlation coefficients
between field cultivar rankings for SAUDPC and
greenhouse rankings for individual cornponents of
resistance, and between the two greenhouse trials (GH-
1 and GH-2) . .................. 103
Tabl. 3.4. Principal component analysis of the data on
the SAUDPC, MLA, PCD, and SPD as pararneters used in
the evaluation of 17 celery cultivars for partial
resistance in the first greenhouse trial (GH-1).. 114
Tabl. 3.5. Principal component analysis of the data on
the SAUDPC, MLA, PCD, and SPD as pararneters used in
the evaluation of 17 celery cultivars for partial
resistance in the second greenhouse trial (GH-2). 118
(
xv
·.1 LIST or rIGOUS
rigure 2.1. Effect of temperature and leaf wetness
duration on infection of celery by Septoria apiicola 45
rigure 2.2. Proportion of maximum infection as a
function of temperature and leaf wetness duration. 47
rigura 2.3. Effect of leaf wetness duration on the
proportion maximum number of lesions (PML) at 10, 15,
20, 25 and 30 oc . . . . . . . . . . . . . . . . . . 50
rigura 2.4. Oisease severity indices (051 = 1 - 4)
categorized by cluster analysis of predicted disease
proportions . . . . . . . . . . . . . .. 52
J'iqura 3.1. Field plot layout 73
J'iqura 3.2. Standard diagram based on the modified
Horsfall-Barratt (HB) scale for a lesion size of 10
mm 2 to estimate of the proportion of celery leaf area
covered by â. apiicola lesions . . . . . . . . . . . 75
riqura 3.3. Standard diagram based on the modlfied
mm 2 to estimate the proportion of celery leaf area--
covered by les ions of ~. apiicola . . . . . . 77
J'iqura 3.4. Standard diagram based on the modified
mm 2 to estimate the proportion of celery leaf area--
covered by lesions of ~. apiicola . . . . . . 79
J'igura 3.5. Ranking of cultivar with respect to the
standard area under the disease progress curve
(SAUDPC) under field conditions . . . . . . 92
rigura 3.6. Grouping of celery cultivars into 5 levels
of partial resistance based on cluster analysis of the
data on the standard area under the disease progress
curve (SAUDPC) for the sampling intervals 0-25 days,
26-40 days, and 41-55 days following the introduction
of inoculum . . . . . . . . . . . . . . . . . . . . 94
rigura 3.7. Principal component analysis (PCA) of
cultivar response under field conditions relative to
the standard area under the disease progress curve
(SAUOPC) for the sampling intervals 0-25 days (S1),
26-40 days (52), and 41-55 days (53) after
inoculation . . . . . . . . . . . . . . . . .. 95
-
,
xvi
rigure 3.8. Disease progress curves of celery cultivars
tested for partial resistance under field condition 96
rigure 3.9. Comparison of celery cultivar rankings
relative ta the SAUDPC between the first and second
greenhouse trials (GH-l & GH-2), and with the field
ranking for SAUDPC . . . . . . . . . . . . . . . . 104
rigure 3.10. Comparis~.l of celery cultivar rankings
relative to the mean lesion area (MLA) between the
first and second greenhouse trials (GH-1 and GH-2),
and with the field ranking for SAUDPC . . . . . . 105
relative to the pycnidial density (PCD) in the first
and second greenhouse trials (GH-1 and GH-2), and with
the field ranking for SAUDPC . . . . . . . . . . . 106
relative to the spore density (SPD) in the first and
second greenhouse trials (GH-1 and GH-2), and with the
field ranking for SAUDPC . . . . . . . . . . . . . 107
rigure 3.13. Hierarchical classif~cation of 17 celery
(~
cultivars based on their response in greenhouse trial
1 relative to SAUDPC, MLA, PCD, and SPD . . . . . 113
rigure 3.14. Principal component analysis (PCA) of
cultivar response in greenhouse trial 1 relative ta
SAUDPC, MLA, PCD and SPD. . . . . . . . . . . . . 115
rigure 3.15. Hierarchical classification of 17 celery
cultivars based on their response in ghreenhouse trial
2 relative ta SAUDPC, MLA, PCD, and SPD . . . . . 117
rigure 3.16. Principal component analysis (PCA) of
cultivar performances in greenhouse trial 2 relative
ta SAUDPC, MLA, PCD and SPD. .......... 119
(
xvii
ABBRBVIATIONS USEn
Abbreviation Ter.m repr•••nt.d
AUDPC Area under the disease progress curve

CI Condition index
CN Condition number
Cp Mallow' s Cp
DSI Disease severity index
GH-1 First greenhouse experiment
GH-2 Second greenhouse experiment
IPM Integrated pest management
IQR Interquartile range
ML Maximum likelihood
MLA Mean les ion area
MLR Multiple linear regression
MR Moderately resistant
MS Moderately susceptible
OLSR Ordinary least squares regression
PC(1) Principal component l
PC(2) Principal component II
PCA Principal component analysis
PCD Pycnidial density
PLAD Proportion of the leaf area diseased
PMD Proportion of maximum disease
PML Proportion of the maximum number of les ions
S Susceptible
S1 Interval from 0-25 days post-inoculation
SAUDPC Standard area under the disease progress curve
SPD Spore density
SPPC Spores per pycnidium
T 50 Time from inoculation to 50 % disease
T 75 Time from inoculation to 75 % disease
UPGMh Unweighted pair-group method using arithrnetic
VrF Variance inflation factor
VS Very susceptible
WLSR Weighted least squares regression
.......
1
( 1 GENERAL INTRODUCTION AND LITaRATOR& REVIBW
1.1 GENZPAL INTRODUCTION
Public demand for innocuous food is rising with the

awareness of increasing environmental pollution and health
risks posed by modern agricultural practices. Soil
degradation, reduced yields and rising costs of production
are the legacy of decades of monoculture and indiscriminant
use of chemical fertilizers and pesticides resulting in the
obligation to use ever higher doses of chemically active
ingredients to control pests that have become resistant.
c Not surprisingly, a growing number of farmers have converted

or are in the process of converting to biological
agriculture. As an example, the Montreal daily 'La Presse'
of August 3D, 1991 reports an increase in their number from
100 to 400 between 1987 and 1991 in Quebec alone. But for
the majority of farms, the situation can only be improved
through long-term investments in more sustainable methods of
agriculture.
In the area of pest control, there has been progress made

toward the rationalization of pesticide use since the
concept of integrated pest management (IPM) emerged in the
late 1950'5. It promotes the use of aIl suitable
techniques, be they chemical, cultural or biological, within
(
2
limits set by environmental conditions to maintain pest

populations below levels at whlch economic los ses occur
(Zadoks & Schein,1979; Metcalf & Luckmann, 1982). In
botanical epidem~ology, this concept has led to increased
research into pathogen population dynamics and the
development of models. The information gajned through these
models on the ecology and biology of pathogens forms the
basis of man y disease forecasting or warning systems which
allow a more precise and ~fficient use of fungicides
(Zadoks, 1984).
Nowadays, the number of IPM programs through which farmers

hire the services of extension specialists who provide
technical support is on the rise. Many vegetable growers in
the muck soil area south of Montreal subscribe to an IPM
program through which they receive the services of scouts
who monitor pest populations in their fields and warn them
of the necessity to apply treatments. Their main crops are
carrot, lettuce, onion and celery. A sequential sampling
procedure to evaluate the severity of Botrytis leaf blight
of onions has been implemented through this program for the
timing of fungicide treatments (Boivin & Sauriol, 1984).
These growers are thus expected to be receptive to the
implementation of forecasting models such as that being
developed for Cercospora blight of car rot (Carisse, 1989;
......
3
{ Carisse and Kushalappa, 1990) and an eventual management
program to control Septoria blight on celery.
Late blight or Septoria blight of celery caused by Septoria

apiicola Speg. is a destructive disease against which
farmers apply fungicides on a regular basis from transplant
recovery until harvest (CPVQ, 1987, Paulus, et al., 1970;
Ryan, et al., 1972). There are no known cultivars of celery
with high levels of resistance to Septoria blight although
sorne are said to have low levels of resistance or tolerance
but none have ever been tested under Quebec growing
conditions. It is felt that Septoria blight could be
adequately controlled with fewer fungicide applications
( early in the season if initiated on the basis of sound
knowledge of blight favourable conditions.
Forecasting involves quantitative studies in the

interactions among host, pathogen and the environment that
result in disease increases (Campbell & Madden, 1990, Ch.
15; Zadoks & Schein, 1979, Ch. 3). Plant diseases can be
monocyclic, allowing a single pathogen generation, or
pOlycyclic and allowing many. Each disease cycle can be
divided into parts corresponding to components of the
pathogene life cycle that can be studied separately. These
are the infection, colonization, sporulation and
( dissemination processes. But the process that has been most

4
1 successful in the development of forecasting systems is the
infection process (Jones, 1986; Leonard & Fry, 1986, Ch.4;
Madden & Ellis, 1988, Ch. 13).
Models can he improved by taking into account other factors

su ch as host resistance. Partial resistance is another
means ~f improving the efficacy of fungicides. Its
inhibiting effect on the rate of disease increase can be
used to vary doses or frequency of fungicide applications
(Fry, 1978; Fry, Apple and Bruhn 1983) .
The work described in this thesis was conducted within the

framework of a more comprehensive program leading te
forecasting and management of Septoria blight of celery. It
consists in two parts. The first part invelves the
quantification the effects of temperature and leaf wetness
duration on the infection of celery by Septoria apiicola
Speg. The second part consists in a screening of celery
cultivars available in Quebec and Ontario for partial
resistance to S~ptoria blight under field and greenhouse
conditions. A number of resistance parameters were
evaluated as potential predictors of cultivar field
performance.
5
• 1.2
1 .2 . 1
REVIB. or LITBRATURE
Crop - Importance and Value
Almost half of the celery produced in Canada is grown in the

province of Quebec. In 1989 and 1990, 17,364 and 15,306
tonnes were sold at a farm value of $5.4 million and $4.4
million, respecti vely (Statistique Canada, 1991). It ranks
close to cucumber, peas, beans and cauliflower in terms of
farrn value even though the area cultivated is much smaller,
making it a highly valuable crop.
1.2.2 Di ••as9 - Historic.l Perspective
Septoria blight of celery has been known for almost as long

as celerv has been cultivated (Cochran 1932, Sutton &
Waterston, 1966). Briosi and Cavara were the first to
report its occurrence on cultivated celery in Italy in 1890.
By the turn of the century, it had been reported throughout
Europe and in Great Britain. In North America, it was first
noted in Delaware, U.S.A. in 1891 and by 1915, los ses as
high as $ 1 million were reported for the State of Michigan.
In Quebec, the disease was said to be on the decline in
1967, presumably on account of crop rotation and sanitation
m~asures (Simard, Crête & Tartier, 1968). ln recent years,
it has been on the rise again although the severity of
(
6
1 epidemics varies considerably from year to year and
meteorological conditions, being worse in wet years
(Brodeur, 1990, personal communication).
1.2.3 The Pathoqen
Taxonomy. The causal organisrn of late blight of celery is

Septoria apiicola Speg. first described by Spegazzini in
1887. There is a fairly long list of synonyms but the two
most commonly encountered in the !iterature until the work
of Gabrielson and Grogan (1964) and Sheridan (1968) were:
Septoria apii (Br. and Cav.) Chester

Septoria apii-graveolentis Dorogin
These were long thought to be two distinct species, the

former being associated with larg~ les ions affecting the
leaves only and the latter with srnaller lesions occurring on
petioles and leaves alike (Cochran, 1932). It was
demonstrated that differences were simply due to variability
in syrnptom expression.
Bost Ranqe. Septoria apiicola Speg. affects wild celery

Apiurn australe Thouars, the cultivated~. graveolens var.
dulce L. and celeriac ~. graviolens var. rapaceum.
Inoculation on other related members of the Umbelliferae
7
have failed to produce infection (Cochran, 1932; Gabrielson

& Grogan, 1964; Sheridan, 1966).
1 .2.4 Symptoma and Btioloqy
The fungus penetrates ho~t tissue directly (Cochran, 1932)

or through the stomata (Alippi, 1987) and grows
intercellularly. It produces greyish lesions which become
bronze colored as they mature and are sometimes surrounded
by a chlorotic halo. The shape and size of lesions can be
quite variable, being roundish on leaves and more elongate
on petioles measuring from 1 to 10 mm in diameter. From the
time they are first visible, lesions are covered with
globose amphigenous black pycnidia measuring between 55-190
J.1m in diarnteter (Sherf and Macnab, 1986, Ch. 7.). Sometirnes
pycnidia appear even before lesions and they tend to forrn in
greatest numbers on the rnost illuminated side of leaves
(Cochran, 1932). Spores are extruded through the ostioleE
in a gelatinous matr~ ~alled cirrhus. The pycnidiospores
are hyaline, though sometimes granulated, filiform and
slightly tapered at their apex. They can measure from 22 to
56 ~m in length and have one to seven septa (Sutton &
~aterson, 1988). Sporulation is profuse and the number of
spores per pycnidium has been estimated at 3,675 (Lin,
1939). The fungus has no known sexual stage, Cochran (1932)
(
8
1 tried unsuccessfully to induce its formation using ultra
violet-light.
The disease is superficially seed-borne (Sheridan, 1963;

Maude, 1964). It can also be transmitted by infected crop
debris in the soil where the fungus can survive up to 9
months (Maude & Shuring, 1970; Hewett, 1968; Sheridan, 1966)
but where it appears unable to grow saprophytically
(Cochran, 1932). Seedling infection is initiated by spores
washing from pycnidia on the seed coat to the cotyledons or
hypocotyl (Maude, 1964). Early disease symptoms are most
noticeable on outer petioles and leaves at the soil level.
.~, Secondary spread to new growth occurs through the action of
splashing rain drops, overhead irrigation, and by movement
of machlnery and workers through fields.
1.2.5 Epidemiology of Saptoria Blight
Germination of spores sprayed on celery leaves has been

shown to require a minimum of 94 % RH between 15 and 17 oC,
while optimal conditions for infection require the presence
of free water for a period of 72 hr at 22 oc (Be~ger, 1970;
Sheridan, 1968; Sheridan 1968a). Overhead irrigation and
high relative humidity can thus promotc infection in the
nursery. In the field, very low initial inoculum can
trigger an epidemic (Maude, 1970). Berger, in southern
9
( Florida, reports rates of disease increase as high as
r=0.44/day associated with heavy rainfall and wet periods of
over 24 hr at temperatures ranging from 13 to 27 oC. In
Quebec, these temperatures are typical of conditions
occurring in June, late July and August. In August when the
crop approaches maturity, even with low rainfall the
presence of dew on the foliage until almost noon promotes
infection which is readily spread during cultural
operations.
1.2.6 Control of Septor1a B11ght
Chem1cal and cultural .ethod•. The first control measure

( against any pest is avoidance. With Septoria apiicola,
avoidance is accomplished by use of clean seeds and crop
rotation. Seed disinfection is performed by immersion for
24 hr in a solution containing 0.2 % thiram and heated at 30
oc (Maude, 1964). It was once thought that storing seeds
for three years was enough to kill the pathogen (Cochran,
1932) but Sheridan (1966) observed renewed spore production
from old pycnidia exposed eight days to high relative
humidity and warned against using old seeds as a method of
control. Navaratnarn, Shuttleworth and Wallace (1980)
report 100 % control of Septoria blight on seeds using
aerated steam.
(
10
l Effective control of late blight in the field depends on
weather conditions, initial disease incidence and early
detection. In Canada, the only fungicides currently
registered for use on celery are chlorothalonil, anilazine,
maneb, mancozeb, zineb, metiram and fixed copper in
combinat ion with mancozeb, aIl of which are applied as
protectants at 7- to 12-day intervals (CPVQ, 1987). Sorne
chemicals used with suceess elsewhere include organotins and
benomyl (Kavanagh& Ryan, 1971), Ciba-Geisy 64251 10W, as
weIl as mixtures of benomyl and ehlorothalonil, and of
benomyl and anilazine (Paulus, Nelson & Otto, 1980).
However, sorne observers have reported fungal resistance to
benomyl (Dullahide, 1979; Gladders & McKeown, 1985). In
Italy and Australia, the triazole propiconazole is said to
have 2-day eradication power (Marco, 1987; Wicks, 1989).
Petroleum oils have also been used in the past for its
fungistatic properties which are sa id to reduce the
production of pycnidia and secondary infection (Wilson,
1961) .
Di ••• a. re.iatane. in eelery. Literature on the resistance

of celery to Septoria blight is seant and sorne, while
describing low levels of resistance in certain cultivars,
dates back more than 20 years (Munger and Newhall, 1953:
Nelson and Mooar, 1954). More recently, there have been
11
( reports of low leveis of resistance and tolerance (Sherf and
MacNab, 1986, Ch. 7) but cultivars with the most desirable

agronomie characters are said to be most susceptible
(Bedlan, 1985). In a report to growers, Zitter (1983) lists
the hybrid Green Giant as resistant to Septoria blight
without elaborating on its character or degree. There has
been attempts by Wright and Lacy (1988) to produce resistant
plants to a number of diseases including Septoria blight by
methods of somaclonal variation. Though cultivars for these
experiments had been selected on the basis of resistance to
Fusarium yellows, the authors nonetheless noted an increase
in the frequency of resistance to foliar diseases among
( plants regenerated from parents with moderate resistance.
1.2.7 Di •• ase R•• i.tance in Cropa
Plant resistance to diseases ls of two types originally

termed 'vertical' and 'horizontal' by Van der Plank (1963)
and performing distinct roles in disease management.
Vertical resistance is controlled by one or few genes

resulting in disease only if a given virulence gene in a
particular parasite matches exactly a specifie resistance
gene in a host. Because of the gene-for-gene character of
vertical resistance it is said to control 'unmatching
12
1 infection' (Robinson, 1987, Ch. 6). It is also termed
differential or race specifie because it involves
interactions at the sub-speeies level among pathogen
physiological races (vertical pathotypes) and host species
cultivars (vertical pathodemes) (Parlevliet, 1979; RObinson,
1987; Van der Plank, 1969). Vertical resistance can be
complete and allow no disease at al1 or be partial and allow
sorne disease to develop. It is relatively easy to breed
into crops because it obeys Mendellian laws of segregation
hence its great initial popularity (Robinson, 1987).
However, it has been plagued with sudden failures resulting
in important crop losses and its reputation for being of
short duration has caused it to be abandoned as a disease
management alternative (Zadoks & Schein, 1979).
Horizontal resistance operates against aIL races, it is what

remains once vertical resistance has been matched and for
that reason is said to control 'matching infection'
(Robinson, 1987, Ch. 6). It is quantitative in nature and
exhibits continuous variation in both the host and the
parasite. It is the result of an evolutionary process
making it more difficult to select for and assess.
Furthermore, it is often obtained at the expense of more
agronomicallv desirable characters. It has been designated
by a number of terms including race nonspecific,
- nondifferential, field, general, polygenic, uniform,

13
incomplete, quantitative and partial resistance (Browning,

Simons & Torres, 1978; Fry, 1982; Parlevliet, 1979). These
terms have different connotations and sorne could even apply
to vertical resistance because it can also be partial and
rate-reducing (Fry, 1982, Ch. 10). The major interests of
horizontal resistance are its apparent durability and its
ability to reduce the rate of epidemics by interfering with
sorne or aIl subprocesses in the parasite life cycle.
Relative assessment of horizontal resistance in cultivars is

accomplished by ranking them with respect to one or more
cornponents of rate-reducing resistance. The common
parameters used in its evaluation are infection frequency,
latency period or incubation period, lesion size, infectious
period and sporulation (Parl~vliet, 1979). Variations in
one or more of these paramel:ers become cumulative and are
most evident in polycyclic diseases (Fry, 1982, Parlevliet,
1979) .
Field tests are important in resistance trials as they also

reflect cultivar suitability to the climate of the area
where they are being performed (Robinson, 1987). Because
environmental factors have a profound effect on ho st and
pathogen alike, and also because cultivars can react
differentially to changing environments it is strongly
14
" recommended to test them in the geographical area of their

commercial production.
Interplot interference is a phenomenon that needs

consideration when planning field resistance trials (Van der
Plank, 1963). It involves the export of inoculum from plots
of susceptible cultivars to neighbouring more resistant
cultivars resulting in what Van der Plank termed 'cryptic
error'. It often causes resistant cultivars to appear more
susceptible and in susceptible ones to appear more resistant
than they would if grown in normal size fields (Van der
Plank, 1963). This is particularly true with small plots
and pathogens whose propagules are windborne. It is not
considered as critical if the experiment is not meant to
represent the field situation.
The cultivar screening described in chapter 3 was designed

to test their ability to reduce the rate of blight
development under the conditions prevailing in southern
Quebec. Not knowing ahead what mechanism of resistance may
be at work, the term 'partial resistance' is used throughout
to describe the gradation in celery cultivar resistance to
Septoria blight. The components of partial resistance
studied were the disease severity, lesion size, pycnidial
density and pycnidiospore density per square centimetre ~f
1esion, and the latency period.

•
15
1.2.8 Di ••• s. For.c.sting
Forecasting future disease outbreaks or issuing warnings

that infection has already occurred constitutes an efficient
and economical approach to disease management if accurate
and easy to implement (Krause & Massie, 1975). The primary
objective of disease forecasting is to reduce the number of
fungicide applications while maintaining adequate control.
The achievement of this goal necessarily calls for a
thorough understanding of the interactions among host,
pathogen and the environment that result in disease (Zadoks
& Schein, 1979).
The literature on the various types of models developed for

different purposes including disease forecasting in the last
fifty or sixt Y years i~ very extensive and description is
beyond the scope of this work (Campbell & Madden, 1990;
Kranz & Hau, 1980; Krause & Massie, 1975; Seem & Haith,
1986; Van der Plank, 1963; Zadoks, 1984). Models can be
simple, involving single components of the disease cycle
such as reproduction, dissemination, infection or
colonization. They can also be complex and involve a
combinat ion of processes into larger models that mimic
entire epidemics (Fry, Apple & Bruhn, 1983; Kushalappa et.
al., 1983; Zadoks, 1971).
16
Once the relationships between disease and weather factors

have been quantified and described in terms of mathematical
or r~gression equations these can be applied in different
ways towards disease forecasting. The equations can be used
directly to predict disease by substitution of weather
parameters (Carisse & Kushalappa, 1990; Chuang & Jeger,
1987; Madden et.al., 1991), or predicted values from the
equations can be transformed into disease severity indices
or infection equivalents for the environment (Danneberger,
Vargas & Jones, 1984; Gillespie & Sutton, 1979; Sutton,
James & Rowell, 1986; Kushalappa, et.al., 1984). Once the
validity of the model is established, it can be used in
conjunction with a predefined critical disease level or
threshold to time the first and subsequent spray
applications (Gillespie & Sutton, 1979; Madden, Pennypacker
& MacNab, 1978; Vincelli & Lorbeer, 1988).
Celery is not a major crop but a valuable one as

demonstrated earlier. Thus, an adequate warning system for
the control of Septoria blight will need ta be simple. It
would be expected to save a few fungicide applications at
the beginning of the season where the reduced plant size
provides sufficient aeration to render conditions less
favourable for disease development. Later on, the
microclimate in the maturing crop is almost always conducive
to low levels of infection because of the presence of dew
17
which often does not evaporate until midday due to poor
aeration.
One aspect that is easily incorporated into forecasting

models is the effect of ho st partial resistance (Fry, 1977;
Matyae & Bailey, 1988). Partial resistance, particularly if
horizontal, can be used to complement and increase the
efficiency of fungicides (Browning, et al., 1978;
Parlevliet, 1979). In sorne applications it has been used to
vary spray intervals or fungicide dosage adjustments (Fry,
1978) to account for the different level of resistance. In
others a disease index or infection equivalent generated by
the infection model cal. be corrected for host resistance
(Kushalappa, et al., t 984) .
1.2.9 Multivariate Stati.tieal ~thod. in Epidemiologieal

Studie.
Traditionally, multivariate statistical methods have been

the domain of ecological research and systematics (Sneath &
SokaI, 1973; Sokal & Rohlf, 1981). They are gradually
gaining acceptance in plant pathology because of their
capability to handle multiple dependent and independent
variables, to summarize and help in the overall
interpretation of results (Kranz, 1990).
f,
L
18
An example of applications in efidemiology which are related
te the work described herein is the use of cluster and
discriminant analysis in the classification of disease
favourable weather into severity values (Sutton, James &
Rowell, 1986). Other examples include the use of cluster
and principal component analyses to characterize cultivar
resistance based on their disease progress curve and other
resistance parameters (Anderson et.al., 1990; Jeger, 1980;
Lebeda & Jendrulek, 1988; Lancashire & Jones, 1985; Platt &
Tai, 1984; Thompson & Rees, 1979).
In addition to normal statistical procedures involving

multiple regression and analyses of variance, the
multivariate techniques used in chapters 2 and 3 to
summarize and interpret results were cluster analysis and
principal component analysis. Cluster Analysis is a
technique which groups objects into mutually exclusive
classes based on their similarity such that within cluster
variances are smaller than between cluster variances. The
role of principal component is to provide a concise summary
of the data in addition to helping in the interpretation of
the structure of the clusters (Dillon & Goldstein, 1984).
elu.ter analyaia. In cluster analysis, there are a number

of methods for calculating similarity or dissimilarity
coefficients and clustering algorithms whose description is
19
beyond the scope of this work and the subject of
comprehensive text books (Dillon & Goldstein, 1984; Legendre
& Legendre, 1984; Romesburg, 1984). The euclidian distance
was used as a measure of similarity among objects in the
analysis of the data in chapters 2 and 3, and the selected
clustering algorithm was the average linkage method.
According to this procedure the most similar objects are
grouped into progressively larger clusters. At each step in
the clustering process, the distance between a newly formed
cluster and outside elements is calculated as the average of
the distances of each object within the cluster and these
outside elements. Furthermore, at each step the distortion
introduced by calculating the average distances can be
( assessed by the cophenetic correlation coefficient. The
cophenetic correlation coefficient is obtained by comparison
of the elements of the cophenetic matrix i.e., that composed
distances recalculated after the last clustering step, with
the individual elements in the original distance matrix. It
becomes ~vident then that the larger the number of objects
within clusters and the fewer the number of clusters
remaining, the less the resemblance between the two matrices
and hence the lower the RZ • A variation of R2 is the
formula in the SAS r::LIJSTER procedure (SAS/sat, 1988)
according to which R2 is the ratio of the sum of within
cluster variances to total variance subtracted from one.
The graphie representation of results from cluster analysis
f
L
20
1 is a tree or dendrogram with the length its branches
corresponding to distances at which individual objects or
clusters area being joined. The optimal number of clusters
to retain by cutting the tree at a particular distance
coefficient is a function of the desired R2 which in turn is
a very suggestive matter that can only be resolved in
relation to the research objectives (James & McCulloch,
1990; Romesburg, 1984, Ch.16).
Principal compon.nt analyais. Principal component analysis

transforms a set of correlated variables into a reduced
number of orthogonal or mutually independent variables which
are linear combinations of the former and explain as much of
,.,#-
the total variation as possible. Principal components are
extracted in a manner such that the first component has the
largest variance and each successive component represents
the largest portion of the remaining variance (Dillon &
Goldstein, 1984). Weights or eigenvalues associated with
each principal component are proportional to their
respective share of the total variance. Each principal
component has an eigenvector with coefficients referred to
as component loadings. These reflect the contribution of
each original descriptive variable to the linear combination
that makes up a principal component. Component lcadings are
equivalent to the product-moment correlation of each
original predictive variable and the respective principal
21
components (Dillon & Goldstein, 1984). Together, the first

two principal components often account for almost 80 to 90 %
of the total variation. This has powerful graphie
implications because where n objects were originally
envisioned in a space of p dimensions corresponding to the
number of descriptive variables, they can now be depicted in
a two- or three- dimensional space with almost as much
resolution, which is sufficient to permit identification of
distinct groups. The objects' positions in the two-
dimensional space are determined by their respective
component scores which are a function of their performance
with respect to each of the original descriptive variables
in conjunction with the loading of those variables on the
( specifie components.
Principal component analysis is most valuable in

interpreting the structure of the groups obtained by cluster
analysis when the outline of latter are traced on a graph
showing the position of the n objects in the space described
by principal component l and principal component II as
coordinate axes.
(,
.
22
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26
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1
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! pycnidiospores of Septoria apiicola Speg. N.Z. Jl Bot.
1
1 6:315-322.
Sheridan, J.E. 1968. The causal organism of celery leaf
spot, Septoria apiicola. Trans. Br. mycol. Soc. 51:207-213.
Sheridan, J.E. 1968a. Conditions for infection of celery by
Septoria apiicola. Plant Disease Reporter 52:142-145.
Simard, T., Crête, R" et L. Tartier. 1968. Les maladies des
légumes de sol organique en 1967. Phytoprotection 49:49-54.
,
J$
c Sneath, P.H.A., and R.R. SakaI. 1973. Numerical Taxonomy -

The principles and Practice of Numerical Classification.
27 ~
4
1
i
~
W.H. Freeman, SanFrancisco. 573 pp.

Sokal, R.R., and F.J. Rohlf. 1981. Biometry - The Principles
and Practice of Statistics in Biological Research. 2nd Ed.
W.H. Freeman, San Francisco. 859 pp.
Statistique Canada. 1991. Fruit and vegetable production.
Catalogue 22-003 Seasonal.
Sutton, B.C., and J.M. Waterston. 1966. Septoria apiicola.
C.M.I. Descriptions of Pathogenic Fungi and Bacteria No. 88.
Commonwealth Agricultural Bureaux.
Sutton, J.C., James, T.D.W., and P.M. Rowell. 1986. Botcast:
a forecasting system to time the initial fungicide spray for
rnanaqinq Botrytis leaf blight of anions. Agriculture,
Ecosystems and Environment, 18:123-143.
Thompson, J.P., and R.G. Rees. 1979. Pattern analysis in
epidemiological evaluation of cultivar resistnace. Letter to
the editor. Phytopathology 69:545-549.
Van der Plank, J.E. 1963. Plant Diseases: Epidemies and
( Control. Academie Press, New York & London. 349 pp.
Van der Plank, J.E. 1969. Pathogenic races, host resistance,
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52.
Vincelli, P.C., and J.W. Lorbeer. 1988. Comparison of
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Wicks, T.J. 1989. Fungicidal control of leaf spot (Septoria
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Wright, J.C., and M.L. Lacy. 1988. Increase of disease
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259.
Zadoks, J.C. 1971. Systems analysis and the dynamics of
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(
28
1 Zadoks, J.C. 1984. A quarter century of disease warning,
1958-1983. Plant Disease 68:352-355.
Zadoks, J.C. and R.D. Schein. 1979. Epidemiology and Plant
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Plant Pathology. New York. 6 pp.
29
(
2 arraCTS or TDIP.u.'1'UU AllI) UAI' nTNJ:SS DURATIOR OR
TU IU&CTIOR 01' CaLaR! BY Sap'1'ORIA UIICOLA
2. 1 ABSTRACT
Infection of celery (Apium graveolens L.), cv. Florida-683,

by Septoria apiicola was studied by inoculating plants with
a suspension of 20,000 spores Iml, and incubating at five
temperatures (10, 15, 20, 25 and 30 OC) and five leaf
wetness periods (12, 24, 48, 72 and 96 hr). The entire
experiment was replicated once. Observations on the number
of lesions per leaf, two leaves per plant, were taken every
three days starting on the 9 th until the 24 th day after
inoculation. Lesions increased in number with temperature
up to 25 oC and then diminished. The number of lesions
increased increased monotonically at most tempe ratures
except at 25 and 30 oc where they diminished after 72 hr
and 48 hr wetness, respectively. The highest number of
lesions was recorded at 25 °C/72 hr and the lowest at 10 oC
and 30 OC. The incubation period was found to be to be
shortest at optimal infection temperatures. The angular
transformation of the proportion of maximum number of
lesions for each replicate was related to centered
temperature and wetness duration in a weighted least squares
polynomial regression, (R 2-adj.=74.8). Cluster analysis was
(
'f' 30
<If' used to divide the response surface into disease severity
indices corresponding to increasing infection probablities.
-
31
2.2 INTRODUCTION
Septoria blight or late blight caused by Septoria apiicola

Speg. is a serious foliar disease of celery. The fungus
produces greyish lesions that turn bronze as they get old
and dry. These are more or less circular on leaves and
elongate on petioles which they sometimes cover in their
entirety as they coalesce. Lesions are dotted with black
pycnidia exudinq masses of pycnidiospores in a gelatinous
matrix, the cirrhus, which protects them against desiccation
and inhibits germination until diluted by ra in (Louis, &
Cooke, 1985; Fitt, et al, 1989). Disease spreads from foci
characteristic of splash dispersal. Heavy precipitation,
conditions of cool weather, high relative humidity and dew
favour infection particularly as the crop matures and covers
the rows preventing evaporation of dew until almost mid-day.
At that time, the movement of machinery and workers in the
fields becomes an important source of secondary spread
(Linn, 1939). Although no figures are available for Quebec
and Ontario, Septoria blight is known to cause severe losses
in wet years, in addition to the requirement for labour
intensive trimming. Furthermore, the relatively long
incubation period of this disease (Berger, 1970) inevitably
means that les ions continue to appear in st orage provoking
further losses.
,
32
In addition to using treated (Maude, 1970) or 3 year-old
seeds (Cochran, 1932), farmers are advised to apply
fungicides on a weekly basis from transplant recovery until
harvest (MAPAQ AGDEX 250-605, 1987; OMAF Publication 363,
1988) ta protect their crop valued at $5,5 million for 364
ha harvested in Quebec (Statistics Canada, 1988).
Increasingly, health, environmental and economic concerns

are motivating research into forecasting systems to
determine the appropriate timing for initiation of fungicide
applications 50 as to hit pathogens at the most vulnerable
stage in their life cycle. The benefits are often a
reduction in the number of applications with equivalent
control ta that obtained with the regular schedules
currently used while addressing these pressing issues.
The investment in the development of forecasting systems is

justified for pathosystems characterized by sporadic disease
outbreaks affecting high value crops. It also presupposes
the availability of adequate management technology and
extension services (Fry, 1982, p.123). In Quebec, the
latter condition is met by a provincial government
phytosanitary advisory service and/or by grower subscription
to integrated pest management programs which provide the
services of scouts to monito~ their fields and advise them
on the necessity to protect their crop.
33
1 Environmental factors play a key role in the infection of a
host by a pathogen as illustrated by the disease triangle
(Zadoks & Schein, 1979). Consequently, the first steps into
the development of a forecasting model for a given
pathosystem often consist in fundamental studies of the
effects of leaf wetness duration and temperature during
these wet periods on infection (Campbell & Madden, 1990;
Fry, 1982, p.83, Jones, 1986). Earlier work with Septoria
apiicola Speg. showed that a minimum of 12 hr at 100 % RH
and temperatures ranging between 5 and 25 oc were required
for spores to germinate on water agar whereas optimal
conditions occurred between 20 and 22.5 oc (Sheridan,
1968b). Other studies indicated that a minimum of 36 hr at
no less than 94.5 % RH were required for leaf infection
while temperatures exceeding 30 OC were inhibitory
(Sheridan, 1968a). Berger (1970) who studied the
epidemiology of late blight under field conditions reported
high rates of infection (r=0.44) in association with periods
of heavy rainfall and average monthly temperatures below 25
oc. To date, no reports have been published on the
influence of temperature and leaf wetness duration for the
full range of conditions conducive to Septoria blight
epidemics.
This study was conducted to quantify the relationship
( between disease severity and leaf wetness duration at

\
34
various temperatures as a preliminary step in the
develop~ent of a management system for Septoria blight of
celery.
2.3 MATBRIALS AND METBODS
2.3.1 Production of Planta
Three year-old seeds of the cultivar Florida-683 obtained

from a commercial grower were sown in groups of 50 in 12.5
cm pots filled with a substrate made up of organic soil (27-
30 % organic matter), peat moss and perlite (4:1:1 v/v)
supplemented with horticultural lime. The pots were
enclosed in a plastic bag ta maintain high RH and placed in
a growth cabinet set at 20 oc. Four week-old seedlings were
transplanted individually into 13 cm pots using the same
substrate and were moistened with a solution of 400 ppm
phosphorus as 10-52-10. A di lute fertilizer solution (100
ppm N in the form of 15-15-17) was administered in the
irrigation water, and a foliar spray of calcium chloride
was applied at the 3-leaf stage to suppress black heart
which tended ta affect the 4th leaf in absence of the
treatment. The pH of the irrigation water was maintained
around 5.6 by addition of dilute H2 S0 4 as required. Plants
we~e placed in a growth cabinet with a 14-hour photoperiod
35
( supplied by a mixture of fluorescent and incandescent
fixtures providing a light intensity of 250 ~Em-2 sec- l •
Temperature was maintained at 22 ± 2 oC during the light

period and at 18 ± 2 oC in the dark while relative humidity
ranged from 45 to 60 % in day time increasing to 62 to 80 %
in the dark. The wider RH range was largely due to
variation in plant leaf area at different growth stages.
Relative humidity was monitored with the aid of a
hygrothermograph (Bendix, model 594) located among plants.
Aphid control in the nursery was achieved with by-monthly
applications of pirimor, orthene and Safer's Soap in
rotation.
2.3.2 Inoculum Production
An isolate of Septoria apiicola from diseased plants

originating in a commercial field south of Montreal was
cultured on 15 % celery juice agar (CeJA) supplemented with
0.4 % potato dextrose broth (PDB). The celery juice was
prepared by grinding 50 9 fresh celery leaves in 500 ml
water for two minutes in a blender set at high speed. The
mixture was strained, and the juice was boiled and strained
again to remove coagulated material. Agar (7.5 g) and PDB
(2 g) were added and the volume was adjusted to 500 ml by
addition of distilled water before autoclaving. In
v-a
( preliminary studies, this medium compared favourably to
36
1
' ,
juice agar (Tuite, 1969), PDA and Czapek-Dox for the
uniformity and appearance of pycnidiospores which are
generally filiform and tapered (CM! description ,No.SS)
whi1e those formed on PDA and Czapek-Dox tended to be blunt
and rod-shaped. The addition of POB to CeJA accelerated
germination without affecting spore quality.
Fresh cultures were maintained by regular isolations from

diseased plants, preferab1y from petioles where sporulation
was more abundant. This was done by touching extruded
cirrhi with a flamed sterile needle under a dissecting
microscope, and then dipping the needle in sterile
distilled water in an small petri dish to release adhering
cirrhi. These steps were repeated until the suspension
became turbid. Plates were incubated in the dark for 6 days
at 20 oC (optimum temperature deterrnined by preliminary
tests) and then exposed for 8 days to a regime of 12-hr
light provided by cool white fluorescent (Sylvania
F15T12/CW) and near UV (GE 15T8-BLB) light fixtures to
promote sporulation (Cooke & Jones, 1970). At the end of
that period, cultures were either used irnmediately or sub-
cultures were made.
Inoculum was prepared by repeatedly f100ding a culture plate

with a 5-ml solution of 0.01 % Tween 80 in distilled water
and dislodging cirrhi by gentle syringe outflow. The mean
37
( of 12 spore counts, measured with a hemacytometer, was used
as a basis for dilutions to obtain 20,000 spores/ml.
2.3.3 Tr.atments and Inoculation
The experiment, which was r~plicated once, consisted of five

separate inoculations conducted over tirne at ternperatures of
10, 15, 20, 25 and 30 oc. Each temperature treatment (main
plot unit) was adrninistered to 40 plants (sampling units),
eight for each leaf wetness treatment (subplot units)
consisting of 12, 24 48, 72 and 96 hr.
In the first replicate, the 4th and 5th true leaves (in
( order of appearance) of la week-old plants were inoculated
whereas in the second replicate, the 5th and 6th leaves of
12 week-old plants were used. In both cases, the youngest
leaf was not fully expanded at the time of inoculation to
allow four weeks for observation before the oldest
inoculated leaf senesced.
To ensure a unique time at which all sarnples from a given

wetness treatment would be removed from the rnist chamber,
plants were inoculated in groups of eight. Thus for each
wetness treatment, eight plants were selected at randorn,
their leaves were tagged and labels were randomized to pots.
(• The marked leaves were inoculated with an artist air brush

38
1 (Badger-350) operated at a pressure of 100 kpa sa as to
deposit a film of discrete droplets without runoff.
Inoculated plants were distributed randomly in the mist
chamber. The latter was assembled in a growth cabinet
(Conviron model 15E) with plexiglass sheets to protect
electrical circuitry. Intermittent mist was provided by a
timer-operated humidifier (Sovereign model 707ms-llO)
located at the center of the chamber and above the plant
canopy. Plants were dried with an electrical fan. Drying
time averaged 20 minutes as verified visually with a hand-
held lens.
As mentioned earlier, main plots were tested over time,

meaning that inoculum and temperature effects were
confounded. To minimize the effect of inoculum, percent
spore germination and spore deposition density were
determined by inoculating two CeJA plates during each
inoculation and incubating them at 20 oC for 48 hr. AlI
inoculations with less than 95 % spore germination were
discarded.
Lesions were counted every three days from the 9th to the
27th day after inoculation. However, the count of the 24th
day was retained for the analysis for two reasons: 1) the
incubation period varied with infection conditions and
initial lesions did not appear until the 24th day in sorne
39
c treatments; 2) by the 27 th day, lesions formed under
optimal conditions for infection were coalescing and their
continued increase in number aroused skepteticism regarding
the possibility of secondary lesions.
2.3.4 Data Analy.i. and ..gre •• ion Model Selection
Multiple linear regression (MLR) was used to describe the

influence of temperature and leaf wetness duration on the
number of lesions using SAS PRoe REG (SAS Institute Inc.,
1987) .
Infection was expressed as a proportion of the maximum

number of lesions (PML) observed in each replicate and
defined as follows:
PML k = no. lesions WjkT;Jlc

-
MAX no. lesions lc
(2.1)
where W1 stands for wetness, i = 1,2, ... ,5, Tj stands for

ternperature, j = 1,2, ... ,5, and k = 1,2 stands for the
replicate.
The angular transformation of PML was used to stabilize the

variance because it produced a better R-square and randorn
(
40
pattern of residuals than either the logistic or Gompertz
transformations. It is defined as:
yi = arcsin{1iR!. (2.2)
The first step in model selection was done by ordinary

least squares regression (OLSR) and Mallow's Cp (Kleinbaum
et al, 1988, chapt. 16) as selection criterion (see Appendix
A for details) . Analysis of residuals was done by means of
SAS UNIVARIATE procedure. Removal of outliers was based on
indications from the boxplots and stem-and-Ieaf diagrams of
jacknife residuals (In SAS, studentized residuals with the
current observation deleted). Following removal of
.',
outliers, the selection process was repeated requesting the
ten best solutions for each replicate separately and for
the pooled data (Appendix A-l) .
The next step was aimed at reducing collinearity among

pre~ictor variables inherent to polynomial regression (see
Appendix B for details). Predictor variables were centered
as a first measure to reduce collinearity. Diagnostic
statistics such as the variance inflation factor VIF,
eigenvalues of the predictor variables correlation matrix,
their associated condition number (CN), and variance
proportions were used to detect near collinearities and to
- delete troublesome predictors (Kleinbaum et al, 1988, Ch.

12) •
41
( Homogeneity of variances of the data from the two replicates
was verified by fitting the selected model to both sets of
data (F 186 ,189 = 1. 04 < P < 0.05).
A weighted least squares regression (WLSR) was conducted to

address a problem of heterocedasticity brought to light by
the analysis of residuals. Variances tended to increase
with increasing mean values of the response variable,
particularly as a function of temperature (see plots of
jacknife residuals vs PY for each temperature, Appendix C,
and interquartile range (IQR) of box-and-whiskers plots,
Appendix 0). Determining the right weights is not always
clear and may require a fair amount of trial and error
( (Dillon and Goldstein, 1984). Generally, the inverse of the
squares of Y, or PY from a previous OLSR serve as weights in
the next step (Freund & Littell, 1986). However, in this
case y was not used due to the presence of zero's and
negative values of arcsin square root PML in the
denominator. On the other hand, the inverse of PY squared
reduced R-square too much. So did the inverse of the
variance as determined by the UNIVARIATE procedure in step
one. The decision was made to use the inverse of the
squared standard error of residuals from the previous OLSR
output as weights in the next iteration.
(
42
.1 Analysis of residuals also pointed out sorne departures from
the normality assumption at the extrernes of the growth range
of ~. apiicola where conditions for infection allow only a
few les ions to develop (Appendix E). In this case,
residuals follow a binomial distribution. Computer programs
sueh as the Generalized Linear Interactive Modelling System
software (GLIM, The Nurnerical Algorithms Group, Ltd., 1987)
would be more appropriate to handle such problems by the
method of maximum likelihood (ML). However, the WLSR
solution was retained because residuals were normally
distributed at most temperature/wetness combinat ions 50 that
not much improvement was expected in going to such
refinement particularly when considering that the data was
highly variable.
Cluster analysis was used to categorize back-transformed

predicted values generated by the WLSR model into disease
severity indices (DS1). Clustering was done based on the
unweighted pair-group method using arithmetic averages UPGMA
(Dillon and Goldstein, 1984; Romesburg, 1984), SAS's
METHOD=AVERAGE.
43
( 2.4 RlSUL~S
In general, the number of lesions increased with increasing

duration of leaf wetness except at 25 oC and 30 OC where
fewer lesions developed after 72 hr and 48 hr, respectively
(Figure 2.1). For aIl leaf wetness periods, les ions were
more numerous at 25 oc. The largest number occurred at 25
°C/72 hr and the lowest numbers were recorded at 10 °C/12 hr
and 30 °C/96 hr leaf wetness. The data was quite variable
and differences occurred between the two replicates,
particularly at 10 and 15 OC where the response which had
been similar in the first replicate differed considerably in
the second. Otherwise, the trends were similar.
(
The equation for the final weighted least squares regression
was of the form:
~=~+~T+~W+~~+~~W+~~+~~W+~~+~~
(2.3)
where y' is the angular transformation of PML (eq. 2.1).
The shortest suggested equation using Mallow/s Cp was in

the second replicate. However, diagnostic tests for
cOllinearity were performed on the first equation of the
pooled data (Tables B-la and B-1b, Appendix B) which
contained a maximum number of terms common to equations from
both replicates. Deletion of correlated predictors reduced
44
the model to the second equation suggested for replicate 2

minus the term T2 (Appendix A-1) .
Estimates of all parameters included in the final pooled

model were significant at the 0.01 level and their values
were intermediate between those of the first and second
replicates (Table 2.1 and Appendices F-l & F-2).
The model accounted for 75 % of the total variation. The

intercept of the regression of observed values on back-
transformed predicted values was not significantly different
from zero and the slope of the regression line was 1 which
is expected when the fit is adequate (Appendix F-3) .
Solution of the regression equation produced the three-
dimensional response surface illustred in Figure 2.2.
Predicted values for increments of 6 hour wetness and one

degree temperature were back-transformed as follows:
PML = sin2f(~ (2.4)
The resulting proportions of maximum number of lesions (PML)

were plotted as a function of leaf wetness and temperature.
Equation 2.3 underestimated infection at 25 °C/24 hr and

45
1
Replicate 1
•. oo..-----...,....--------~---___,
~"HC
",
/
0.10 ................................ -/.......

/ :....... .....
0.10 . . . . . . / :/
.....
/ ""
/'
P
M . /'
/'
L
0.40 .
/'
,... ...... .
/'
/'
0.20 .. ,..... .
0.00 ~--~;::::'--.....;..-=--=-;;;;:::;~:::::::=-=:..:iil
11 14 41 11
Lea' Wei.... Durallon (br)
.
Replicate 2
'.00 ~----.------:-----~---=_=-_--,
/ IIC
010
. / .....
... / :. ............. .
"
./
/....--_----.(
/'
0.10 /'
P
M 20C
L
0.40
0.20
0.00 ~==:=::::::;::::::=-=~::..,=-=~;=;::::::::=Ml
11 24 41 7.
Leaf Wetne.. Duration (br)
..
ri9\1re 2.1. Effect of temperature and leaf wetness
duration on infection of celery by Septoria apiicola. PML
= Proportion of the maximum number of les ions observed in
each replicate (mean of eight plants, 2 leaves/plant, for
each temperature/wetness combinat ion)
(
46
"
'l'able 2.1. Parameter estimates of the regression equation
to describe the effects of temperatures and leaf wetness
duration on infection levels, including coefficients of
determination for transformed values (R2 unadjusted, and R2•
adjusted for degrees of freedom) and untransformed values
(R*2 adjusted for degrees of freedom).
Parameter estimates
Parameters
Replicate 1 Replicate 2 Pooled data
0.416322 0.436833 0.42673
(0.014905) (0.01535896) (0.11040)
0.093899 0.062716 0.078674

(0.004827) (0.005015) (0.0035896)
0.006685 0.004316 0.005517

(0.000546) (0.000558) (0.000403)
0.000489 0.000260 0.000374

(0.000146) (0.0001495) (0.0001077)
-0.0000606 -0.0000376 -0.0000492

(0.0000079) (0.000008) (0.0000058)
-0.000941 -0.000603 -0.000776

(0.0000504) (0.0000521) (0.0000374)
-0.00000596 -0.00000335 -0.00000465

(0.00000158) (0.00000161) (0.00000116)
-0.00003469 -0.0000365 -0.00003559

(0.0000023) (0.0000023) (0.00000166)
-2.32182e-9 -1.7J886e-9 -20.5387e-9

(0.000000) (0.000000) (0.000000)
0.8110 0.7216 0.7540
0.8030 0.7096 0.7489
Error df 189 186 384
R*2 0 7069<
.. bo 1S the value ot arCSln 'VY when T=U ana W'-'U.

47
c Prop. Mu.
• lesions
1.0 r - - - - - - - - - - - - - T - - - -__
0.8
o.e
o...
( 0.2
96~~~~
~:~ ~ ~ ~
tOe,.~ 15
0(1rh) Temperature (C)
l'iqure 2.2. Proportion of maximum infection as a function

of temperature and leaf wetness duration. Proportions are
back-transformed predicted values (PML=sin 2 f(TW)) where
f(TW) is the solution of the weighted least ~quares
( regression. R2 -adj.= 74.8.

48
25 °C/72 hr wetness duration whereas overestimations
occurred at 20 oc under 48 hr wetness and at 25 °C/96 hr
wetness (Figure 2.3). Note also the high variability of
observations at 25 oC.
Cluster analysis of the back-transformed predicted values

yielded five clusters by cutting the tree at R2 = 92 %
defined as:
(2.5)
where,
~Wj= summation of within cluster variance up to and
including the jth level in the hierarchy;
WT = total sample variance.
Graphical representation of the clusters was accomplished by

tracing their limits as contour lines on a two-dimensional
version of the response surface (Figure 2.4) previously
illustrated in three dimensions. The contour lines join
the points at which changes occur to the next cluster
corresponding te a lower disease severity index (DS1). A
given DSI should be interpreted as affecting the points on
the line and the area between it and the next lower DSI
line. The prediction limits are indicated by the double
contour lines.
49
f
\ Note that cluster 5, the area delineated by the top of the
graph and the fine contour line within OIS 4 (figure 2.3)
was merged with OSI 4 because values above the contour line
were overestimated by the model.
OSI 1 to 4 represent the following proportions of maximum or

probabilities of infection (Details in Appendix G) :
DS! 1 ~ 0.08 < OSI 2 ~ 0.19 < OSI 3 ~ 0.32 < OSI 4
2.6 DISCUSSION
1
Sheridan (1968a) had reported heaviest infection of celery
by Septoria apiico:.a at 72 hr wetness when temperatures were
maintained between 15-17 oC. In this experiment, optimal
conditions for infection occurred at 25 oC. The increase in
the number of les ions at various wetness periods was
monotonie exeept at 25 and 30 oc where fewer les ions
developed after 72 and 48 hr wetness, respeetively.
The fact that sorne les ions did appear at all tested
temperature-wetness combinations underlines the threat of
occurrence of low levels of infection throughout the growing
season given the presence of inoculum and a minimum of 12 hr
leaf wetness. Epidemies would then develop when conditions
50
l'igure 2.3. Effect of leaf wetness duration on the

proportion maximum number of lesions (I:-'t.1L) at 10, 15, 20, 25
and 30 oc. PML was obtained by back-transformation of
predicted values from eq. 2.3 (PML=sin 2 f(T W». Predicted
PML : ' - - 1; Observation means: ,- - -'; olJservations of
rep 1: '.1; observations of rep 2: '+'.
51
15 C
... 10 C
.
.................
... , ,..
M Il
Lu L ....
1.' .. ...
o........+.............. .
1
Il ..* .. .. ... ft .. . 01-
Il
.:-
M
.L
.. MI
•
ft .. ..
WetDell duratloD (hr) WeID'" darailoD (hr)
20 C 25 C
I.. I..
.'
+
"
+ +
*
• ... ,.. ... . : .. ~
+
M .;.
( L u L 0.4
Dt Dt *.. , "+ ' . , .. ,
0
12
•• M .. 611 ft
Wetne•• d .. ·atloD (hl)
. . 0
12 . M .. 611
Weln... daratloD (br)
ft . ..
01
30 C .. - ... ,. .....................
p
u
M
L 0.4
0.2
a
12 ..
t
M ..• to
Wetneu duratlon (hr)
'2 .. "
{
52
96 -96
0 90 90
U 84 84
r
a 78 -78
t 72 72
i
0
66
60
:4 66
60
n
54 54
0 48
t 48
42 2 42
w 36 36
e 30 30
t
n 24 24
e 3 1 18
18
"'t' 8
8 12 12
.L....
6 6
(hr)
9 10 11121314 15 16 17 18 19202122232425262728293031
Temperature du ring wet period (C)
Figure 2.4. Disease severity indices (DSI = 1 - 4)

categorized by cluster analysis of predicted disease
proportions. Predicted values are those obtained by
solution of eq. 2.3 for increments of 6 hr leaf wetness and
1 degree rise in temperature within the range of
combinations tested. The area delineated by the top of the
graph and the fine contour line was a fifth cluster absorbed
into DSI 4 (see text for explanation) .
•
53
became more favourable.
The ~roportion of disease at ZOC and 25C at 12 hr leaf

wetness indicate that wetness periods of 0, 3 and 6 hr
should have been tested to cover the entire infectious range
of Septoria apiicola. Nonetheless, it may be assumed that
infection would decline very rapidly below 12 hours wetness
judging from the shape of the response surface so that a
graphical approach could be taken to assign to the untested
combinat ions to likely disease severity indices to be
confirmed by field testing.
Overall, the model developed by weighted least squares

regression adequately described infection as influenced by
temperature and wetness (R2a = 74.8 %). However its utility
in predicting future infections is undermined by the
presence of a high degree of variability in the data.
Consequently, cluster analysis was used to classify
infection into rneaningful disease severity indices that
respect the shape of the response surface while absorbing
sorne of the variability. Cluster analysis has been used
previously to facilitate field application of complex models
(Sutton et al., 1984).
Many terms were required in the equation for adequate

representation of the relationships under study. For
(
54
instance, the term T4 was required to force the curve down

to almost zero at 30 °C/12 hr wetness. Likewise, TW 4
improved the fit by correctly preuicting the declining
nurnber of lesions with increasing wetness duration at 30 oc.
Without it, the response surface did not go down below PML =
0.2 at 30 °C/96 hr wetness.
The sine function has no biological cause-and-effect

implications but can and has been used to represent the
growth of pathogens in response te temperature (Butt &
Reyle, 1990). It is intrinsically linear, being linearized
by the angular transformation, and as such, is amenable te
regression analysis. The sine functien attains a maximum
of 1 when conditions are optimal for growth of the pathogen,
and a minimum of zero when temperature limits its growth.
There are other successful accounts where it was modified to
include the effects of wetness and temperature-wetness
interactions (Grove, et al, 1985). If the cardinal
temperatures for the growth and development of a pathogen
are tested, it is considered a satisfactory model.
The decline in the number of les ions with increasing leaf

wetness at 30 oc may be due to a loss of viability of
spores that had not successfully penetrated the host after
48 hr. At 25 oC, however, it is not clear whether the
obs~rved decline at 96 hr wetness can be attributed to
55
biological causes or whether it is simply an artifact due to
variability and removal of outliers. One extr~me high value
was removed at 25 oc / 96 hr wetness in each replicate,
artificially reducing variability at that combinat ion and
forcing the curve down (Figure 2.3). It was necessary to
remove these outliers because their values were more than
double that of the next highest ones in both replicates.
Their leverage was su ch that, when included in the
regression, they flattened the entire response surface
because they were the highest values by which all others
were divided.
Variabillty can be attributed to various causes, sorne

controllable with improved technology and technique and
others escaping our control. Sorne of the avoidable causes
include uneven spore distribution on the foliage brought
about by manual operation of the airbrush and confirmed by
the standard error of spore density measurements taken on
plates used in the germination tests (39 spores/cm 2 ± 16,
mean over all plates and inoculations). A second probable
cause may be uneven mist distribution in the moisture
chamber resulting in larger drops on plants nearest to the
vaporizer leading to a 10ss of spores through dripping.
Lastly, it was thought that the use of different growth
stages in the two replicates (leaves 4 and 5 in rep. 1 and
( leaves 5 and 6 in rep. 2) might be responsible for much

56
1 variation but a regression of PML including only data from
leaf 5 in both replicates did not improve the coefficient of
variability. This is understandable in view of the
transformation of the number of lesions to a proportion of
the maximum number of les ions in each replicate. Among
probable but less avoidable causes are differences in spore
vigour and fluctuations in individual plant response. In
the former, there may still be a degree of prevention such
as the use of fresh cultures only rather than first
subcultures. However since the type of culture was not
noted with each inoculation, there is no way of knowing
whether first subcultures were weaker than cultures made
from spores isolated from diseased plants. Using solutions
made by soaking diseased leaves would be ideal except that
the end of an experiment did not always coïncide with the
next inoculation and keeping a constant stock of diseased
plants requires additional space which is often at a
premium. Finally, variation may aiso arise through
fluctuations in individual plant response due to the nature
of the crop. Florida-683 is an open pollinated cultivar and
the expected out-crossing rate in commercial celery seed
production is said to range from 50 and 90% (Orton, and
Aru s , 1 984) .
The effects of over- and underestimation at given

temperature/wetness combinat ions by the regression model on
57
( the resulting DSI's rnay be evaluated by cornparison of
Figures 2.3 and 2.4 while recalling the disease probability
ranges associated with each OSI. The polynomial model
slightly underestimated infection at 10, 15 and 30 oC but
the corresponding OSI is still representative except at 15
°C/ 84 and 90 hr wetness where a DSI of 2 may have been more
appropriate. Sirnilarly, predicted infection at 25 °C/ 24
hr was assigned to DSI 3 while the rnean of observed values
(0.34) would place it in OSl 4. The underestimation at 25
°C/ 72 hr and overestimation at 25 °C/ 96 hr wetness may not
seriously affect prediction since aIl are placed within DSI
4 corresponding to the highest probability of infection
(>0.32). Again, field testing may point out where
( corrections are necessary.
An interesting relationship emerged from having taken lesion

counts at three day intervals for the duration of les ion
development. It became appearent that the date of
appearance of the first les ion on each leaf was also a
function of temperature and leaf wetness during the
infection process (Table 2.2). For example, under optimal
conditions it took exactly 12 days for first lesions to
appear on aIl sarnples in both replicates. In aIl other
cases first lesions appeared between the 12th and 25th day
post-inoculation. Ail plants were kept under the sarne
{ conditions of temperature and relative humidity during

58
Table 2.2 Effects of tempe rature and leaf wetness duration during the
infection period on the number of days between inoculation and
appearance of the first lesions.
Temperature Wetness duration (hr)
(C) 12 24 48 72 96
10 15-1S" 15-24 15-24 12-21 12-24

15 15-18 12-24 12-24 12-14 12-15
20 12-21 12-24 12-15 12 12
25 12-1S 12-15 12 12 12
30 15-21 15-24 15-24 15-24 21-24
The range represents the period over which first les ions appeared
on the 16 sample celery plants (S/replicate).
59
( symptom development so that observed differences may be
attributed solely to treatment effects. Failure by Sheridan
(1968a) and Berger (1970) to observe spore germination
and/or infection at 12 hr wetness may simply be the result
of an underestimation of the incubation periode
In the field, relating disease to weather conditions that
occurred 12 to 24 days prior to evaluation will be a fairly
complex task made worse by knowledge that incubation periods
are also subject to variations according to postinfection
temperatures as reported for Septoria blotch of wheat (Hess
and Shaner, 1987). The difficulty will come after a few
infection cycles where disease no longer increases at the
end of discrete periods but as a continuum of infections
( promoted by a wide range of conditions and resulting in
overlapping incubation perlods. Nevertheless, the problem
may be solved by taking the average of recorded temperatures
and leaf wetness between the 12th and 24th day prior to the
date of observation as was done in a similar situation by
Danneberger et al., (1983).
2.1 CONCLUSION
Based on this study, low levels of infection of celery by

Septoria apiicola may be expected where periods of 12 hr
leaf wetness occur at temperatures ranging from 10 and 30
(,
60
oc. Conditions most conducive to epidemic development occur
at leaf wetness durations of 72 hr concurrent with
temperatures between 23 and 26 OC. The model developed
adequately reflected the general tendencies observed. It
was simplified to reduce the effect of variability by
grouping predicted infection levels into disease severity
indices which will be tested under field conditions.
Incubation periods also varied as a function of temperature
and leaf wetness during infection, being shortest under
optimal conditions.
Obviously, a model based strictly on uninterrupted leaf

wetness is incomplete. In the field, leaf wetness periods
of 72 hr required for heavy infection are not too cornmon but
shorter wet periods interspersed with intervals of high RH
certainly are. Knowing that spores sprayed on celery leaves
can germinate and cause infection 94 % RH under controlled
conditions, it may be ?ssumed that such intervals in the
field could sustain ongoing infections by spores deposited
on leaf tissue by rain or dew. This is why the effects of
high relative humidity should be included in future studies.
61
t LIT8RATURI CITID

Florida State Hort. ~oc. Proc. 83:208-212
Butt, D.J., and D.J. Royle 1990. Multiple regression
analysis in the epidemiology of plant diseases. In: Kranz,
J. (ed.). Epidemies of Plant Diseases. Mathematical
Analysis and Modeling. 2nd Ed. 268 pp.
Cochran, L.C. 1932. A Study of two Septoria leaf spots of
celery. Phytopathology 22(10):791-812.
Cooke, B.M. and D.G. Jones. 1970. The effect of near-
ultraviolet irradiation ~nd agar medium on the sporulation
of Septoria nodorum and~. tritici. Trans. Br. mycol. Soc.
54 (2) :221-226.
CPVQ. 1987. Légumes, Protection. Gouvernement du Québec.
AGDEX 250-605.
Danneberger, T.K., Vargas, J.M. Jr., and A.L. Jones. 1~83. A
model for weather-based forecasting of anthracnose of annual
bluegrass. Phytopathology 74:448-451
Dillon, W.R. and M. Goldstein. 1984. Multivariate Analysis,
Methods and Applications. John Wiley & Sons, Inc. NY. Ch.
7•
Fitt, B.D.L., McCartney, H.A., and P.J. Walklate. 1989. The

role of rain in dispersal of pathogen inoculum. Annu. Rev.
Phytopathol. 27:241-270
Freund, R.J. and R.C. Littell. 1986. SAS System for
Regression. SAS Institute Inc., NC, USA.
Fry, W.E. 1982. Principles of Plant Disease Management.
Academie Press, Inc. 378 pp.
Grove, G.G., Madden, L.V., Ellis, M.A., and A.F.
Schimtthenner. 1985. Influence of temperatura and wetness
dration on infection of immature strawberry fruit by
Phytophthora cactorum. Phytopathology 75:165-169.
Jones, A.L. 1986. Role of wet periods in predicting foliar
diseases. In: Plant Disease Epidemiology, Population
Dynamics and Managerment. (Leonard, K.J. and W.E. Fry)
Macmillan Publishing Co. NY. p. 87.
,
62
Kleinbaum, D.G., Kupper, L.L., and R.E. Muller. 1988.
Applied Regression Analysis and Other Multivariable Methods.
PWS-KENT Publishing Co. Bonston. 718 pp.
Linn, M.B. 1939. Dissemination of celery blight pathogens on
the clothing of farm laborers. Phytopathology 29:553-554.
Louis, I. and R.C. Cooke. 1985. Conidial matrix and ~pore
germination in sorne plant pathogens. Trans. Br. mycol. Soc.
84(4), 661-667.
MAPAQ AGDEX 250-605. 1987. Légumes: Protection. Conseil
des Productions végétales du Québec. Gouvernement du
Québec, Ministère de l'Agriculture, des Pêcheries et de
l'Alimentation. 112 pp.
Ann. appl. Biol. 65:249-254.
Numerical Algorithms Group, Ltd. 1987. The Generalized
Linear Intpractive Modelling System Release 3.77. 2nd Ed.
OMAF Publication 363. 1988. vegetable Production
Recommentations. Ontario Ministry of Agriculture and Food.
78pp.
- Orton, T.J. and P. Arus. 1984. Outcrossing in celery (Apium

graveolens). Euphytica 33:471-480.
Romesburg, H.C. 1984. Clust!r Analysis for Researchers.
Lifetime Learning Publications. Belmont, CA. 334 pp.
SAS Institute Inc. SAS/Stat. 1988. Guide for Personal
Computers, Version 6. Cary, SAS Institute Inc., 1028 pp.
Sheridan, J.E. 1968b. Conditions for germination of
pycnidiospores of Septoria api icola Speg. N. Z. J. Bot.
6:315-322.
Statistics Canada. 1988. Fruit and vegetable production.
Catalogue 22-003 p.16.
Sutton, J.C., Rowell, P.M., and T.D.W. James. 1984. Effects
of leaf wax, wetness duration and temperature on infection
of onion leaves by Botrytis sguamosa. Phytoprotection 65:65-
68.
Tuite, J. 1969. Plant Pathological Methods: Fungi and

Bacteria. Burgess Publishing Co., Minneapolis, Minn. 239
pp .
.....
63
( Zadoks, J.C. and R.D. Schein. 1979. Epidemiology and Plant
Disease Management. Oxford University Press. NY. 427 pp.
(
64
plU:I'ACa 'rO CRAPDR. 3
The Septoria blight forecasting model is being developed

using the cultivar Florida 683 which is very popular among
growers for its high marketable yield and low trim loss
(Valk et. al., 1987). The long-term objectives, however,
are to include ho st partial resistance in the model to
permit even greater economies of fungicides where cultivars
with rate-reducing resistance are being used. The next
chapter is a report of a preliminary screening conducted to
test the levels of partial resistance in cultivars adapted
to growing conditions in southern Quebec.
65
c 3 rIELD AND GREE.BOOSI KYaLOATIOH or CEL&RY
PARTIAL RESISTANCE TC S.PTORXA BLIGBT
COLTI~ rOR
3.1 ABSTRACT
A preliminary test was conducted in the field and greenhouse

to determine the level of partial resistance to Septoria
blight in celery cultivars available in Quebec and Ontario
and to investigate the rale of compcnents of partial
resistance in predicting field response. The standard area
under the disease progress curve (SAUDPC) was used as a
resistance criterion in the field. In the greenhouse, the
components evaluated were the SAUDPC, the mean lesion area
( (MLA), pycnidial density (PCD), spore density (SPD), the
quantity of spores per pycnidium (SPPC) and the latency
period defined as the time from inoculation until disease
severity reaches 50 % and 75 %. Analyses of variance and
means comparisons tests were conducted for indivia~ll
parameters. Non-parametric correlations and multjvariate

analyses were performed ta summarize and compare results
between experiments. Three cultivars, Stokes Golden Plume,
Superdora and Summit performed best in the field and over
all resistance parameters in the greenhause. They are
considered to be moderately resistant. Most other cultivars
had intermediate reactions between the first group and the
( susceptible cultivars Calmario and Florida-683 and very

66
t susceptible cultivar Surepak. There were significant
differences among cultivars with respect to all components
of partial resistance studied in the greenhouse except for
the latency periode The SAUDPC was the most efficient at
identifying cultivars with above average rate-reducing
resistance but it overestimated susceptibility in cultivars
with large lesions. In general, resistance parameters were
poor predictors of the field response for the more
susceptible cultivars.
67
3.2 INTRODUCTION
Septoria blight of celery caused by Septoria apiico1a Speg.

is a polycyclic disease which attacks aIl above ground plant
parts and causes extensive damage in wet year (Berger, 1970;
Sheridan, 1968). It is most1y seed transmitted (Berger,
1970; Hewett, :968; Maude, 1964) but is also able to survive
up to 9 months in crop debris in the soi1 (Cochran, 1932;
Maude and Shuring, 1970).
Measures to reduce primary inoculum include the use of clean

seeds and seedbeds, and crop rotation. In southern Quebec,
control in the field is effected by regular applications of
{
protectant fungicides from transplant recovery until harvest
(CPVQ, 1 987) .
Most celery cultivars, are said to be susceptible to

Septoria blight or tolerant at best (Bedlan, 1985; Munger
and Newhall, 1953; Nelson and Mooar, 1954). Zitter (1983)
lists the hybrid Green Giant as having resistance to late
blight but does not elaborate on its degree or character.
Wright and Lacy (1988), using single cell culture techniques
and working with the cultivars Florida 683 and Tall Utah 52-
70 HK, have had success in generating plants with improved
resistance to four diseases including Septoria blight.
r
• Their choice of parent material was based on susceptibility
•
68
to fusarium yellows. It is not unlikely then that clones
with even more resistance to leaf blights rnay be produced
from parents demonstrating higher levels of resistance than
Florida 683 and Utah 52-70.
Partial resistance is often but not necessarily controlled

by multiple genes and thus cannot be termed horizontal
unless it has been demonstrated (Parlevliet, 1979; RObinson,
1987). It slows disease progress rnuch like chemical control
does by interfering with subprocesses in the pathogen life
cycle causing reductions in the infection frequency, in the
rate of les ion expansion, of sporulation, in the duration
the infectious period, or by increasing the latency period
(Browning, et al., 1978; Parlevliet, 1979; Zadoks & Schein,
1979) .
Because of its epidemiological impact, rate-reducing

resistance is now being incorporated into disease management
strategies. Studies have shown that it can be used to
reduce the frequency and rate of funglcide appllcations
while maintaining good disease control (Fry, 1978; Fry,
Apple and Bruhn, 1983). Such measures have the advantage of
lowering production costs and extending the useful life of
iungicides by retarding the development of pathogen
resistance. The',' also help to reduce fungicide residues in
food and environmental pollution.
69
( The objectives of these preliminary field and greenhouse
studies we~e to establish the levels of partial resistance
to Septoria blight among celery cultivars adapted to
conditions in southern 2uebec and Ontario, and to assess the
potential of resistance parameters tested under greenhouse
conditions as predictors of cultivar response in the field.
3.3 ~T&RIALS AND NlTBODS
3.3.1 Selection of cultivar.
Fourteen cultivars were selected for the field trial based

( on the~r availability to commercial growers in Quebec and
Ontario. The same cultivars plus three others were tested
in the two greenhouse trials.
3.3.2 ri.ld trial
Transplant production. AIl seeds were given a hot-water-

thiram treatment (Maude, 1970). Transplants were produced
at a commercial gre8nhouse. Seeds were sown manually on
April 25 in 400 cell trays filled with a commercial sterile
medium. Seedlings were thinned out after two weeks. They
received fertigation through the over-head irrigation
(
70
1 system. On June 8, seedlings were transplanted manually in
the field at 6 weeks.
ri.ld plot •• tabli.ha.nt and layout. The field experiment

was conducted during Lhe surnmer 1988, at the Ste-Clothilde
Agriculture Canada experimental farm in the muck soil area
south of Montreal. The fourteen cultivars were assigned
randomly co plots in each of four blocks laying
perpendicular to the wind gradient (Appendix H). The 4 mx
4 m plots consisted of six rows, 79 cm apart, with 20 plants
set 20 cm apart within rows. Spacing between plots was also
4 m with 2 m bands seeded to soybean at the centre to help
reduce inter plot interference.
The field was fertilized four weeks prior to transplant by

application of 55 kg N/ha, 100 kg P205/ha and 150 kg K20/ha.
A second application of 55 kg N/ha in the form of ammonium

nitrate was sidedressed five weeks after transplant.
Calcium nitrate (10 kg/ha) and solubor (1.5 kg/ha) were
applied as foliar spray when plants reached 15 and 20 cm
respectively as preventative measures against blackheart and
cracked stem.
One application of Dithane M45 (2.25 kg/ha) and Kocide 101

(2.8 kg/ha) with a hand sprayer was sufficient ta control a
light infection af bacterial blLght in early July.

,
i
71
( Tarnished plant bugs were suppressed by applications of
Malathion 25 WP (4.5 kg/ha) on July 15, August 2, and August
19. Weeding was done by hand.
ri.ld inoculation. Initial inoculum was provided by

transplanting three plants, one with sporulating lesions and
two others inoculated but symptomless, at the centre of each
plot on July 28 (Figure 3.1). Overhead irrigation was
supplied three days a week after 4 pm to favour disease
spread. Timing for the introduction of diseased plants was
based on statistics furnished by the Réseau de dépistage et
de recherche du sud de Montréal at the growers' annual
meeting and according to which infestations were discovered
after July 19 during the 1987 growing season.
Di •• a ••••••• sm.nt. Disease assessment was conducted four

times during August and September on each of three marked
and two randomly selected plants per plot (Figure 3.1).
Disease severity was assessed on leaves including petioles

using the modified Horsfall-Barratt scale published by
Berger (1980) which assigns codes for interclass
interpolation. Standard diagrams were developed depicting
blight severities of 75 %, 50 %, 25 %, 12.5 %, 6.25 %, 3.12
%, and 1.56 %. Three sets of diagrams were prepared based
on standard lesion sizes of 10, 20 and 40 mm 2 (Figures
(
72
1 3.2 - 3.4). Septoria lesions can be very small with no
obvious outline or medium to large when they occur on ~lder
leaves or leaves located closer to the heart. The three

sets were thus used in combinat ion to assess disease
severity in the field.
To design the standard diagrams, the projected outline of a

dried celery leaf and its petiole was drawn and photocopied.
An acetate was placed on the outline and lesions of the
selected size were drawn on it with a washable felt pen.
Individual lesions were drawn at low percentages of disease
whereas individual and groups of coalescing les ions were
used at higher percentages to mimic natural infection. The
acetate, without its outline, was photocopied periodically
and the lesion area was measured with an area meter (Delta-T
devices, Model PM-930, Cambridge, England) equipped with a
video camera (Model ITC-48), an rl compared to the total area
of the projected leaf measured with the same instruments.
Lesions were added or removed from the acetate, photocopied
and measured again repeating this process until the desired
percentages were obtained after which a final photocopy was
made with the leaf outline under the acetate.
The computer program DISTRAIN was used by the author and a

technician as a training tool in disease assessment and to
73
(
rigure 3.1. Field plot layout with 4 m rows each with 20

plants set 20 cm apart (circles). The small dark circles
represent marked plants evaluate1 at all sampling dates.
The dark square, star and triangle represent the three
( randomly selected plants to be sampled on the day
corresponding to each.
(
74
• marked • 25 days * 40 days ... 56 days

.. inoculated plants
0 0 0 0 0 0
0
0
0
...
0
0
0
0
0
0
0
0 ..•
0 0
0
0
0 0 0 0 0 0
0 0 0
0
• * 0
..
0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0
0 •0
0
*0
a
0
0
0
0
0
0 • 0 0 0
0
0
a
0
0
0
0
0
0
•
0
0
0
0
0
0
0
0
a a 0 0 0 0
75
rigur. 3.2. Standard di~gram based on the modified

Horsfall-Barratt (HB) scale for a lesion size of 10 mm 2 to
estimate of the proportion of celery leaf area covered by ~.
apiicola lesions. The three-digit numbers situated below

illustrations are the modified HB code and corresponding
c percentage area diseased. Codes between illustrations are

for inter-class interpolation.
76
111 112 222 263 333

122 226
1.I~a " 3.12 "
688 888 881 811 177

666
26 " 60 "
178 888 880 800 OOg
788
16 "
77
Figure 3.3. Standard diagram based on the modified

Horsfall-Barratt (HB) scale for a lesion size of 20 mm 2 to
estimate the proportion of celery leaf area covered by
les ions of ~. apiicola. The three-digit numbers situated
beloN illustrations are the modified HB code and
corresponding percentage area diseased. Codes between
illustrations are for inter-class interpolation.
L
7B
111 112 222 2BB BBB B .. 4

122 22G BB"
3.12 • B.26.
1.6G S
~66 666 688 888 687 677 771

~~6 668
12.6 S 26 S 60 ..
718 GGG GGS GSS SSS

T88
76 S
79
riqure 3.4. Standard diagram based on the modified

Horsfall-Barratt (HB) scale for a lesion size of 40 mrn 2 to
estirnate the proportion of celery leaf area covered by
lesions of ~. apiicola. The three-digit nurnbers situated
below illustrations are the rnodified HB code and
corresponding percent age area diseased. Codes between
illustrations are for inter-class interpolation.
80
111 112
228
3.12 •
,
41515 1566 DDD DDD DD7 D71 777
446 D6D
12.6 • 26. 60.
71e eee Dee 89U 9UU

788
(, 76 •
81
compare their ability to evaluate disease similarly,
accurately and consistently (Tomerlin, 1988).
In the field, workers alternated taking notes and assessing

disease on a plot by plot basis, initialling the data sheet
to ensure that the same plot would not be assessed by the
same persan on successive visits.
Horsfall-Barratt codes were converted ta proportions of leaf

area disf~ased (PLAD) using the software programm DISPAR
developed ta process epidemiological data and adapted for
that purpose (Carisse, 1989). The mean proportion of leaf
area diseased (PLAD) per plot over the observation period
was converted to the are a under the disease proqress (AUDPC)
using the formula of Shaner and Finney (1977):
r"",nC = ~
A vue ~
( Y1 +2 Y1+1) (t 1+1 - t 1) (3.1)
where the summation is over all sampling dates, Yl = disease

proportion at time i, and tl~ - t 1 represents the interval
between sampling dates. The units of AUDPC are proportion-
days.
In this experiment, each black was assessed on a different

day resulting in a three-day difference in the duration of
- the AUDPC between block 1 and block 4. Ta facilitate

82
t comparison of disease progress curves of different
duratiuns, the AUDPC is standardized through division by its
total duration. The resulting quantity has no units and has
been termed standard area under the disease progress curve
(SAUDPC) (Campbell & Madden, 1990), relative area under the
disease progress curve (RAUDPC) (Padgett, et al., 1990) or
normalized ADPe (Fry, 1978). The abbreviation SAUDPC will
be used throughout this document.
3.3.3 Greenhouse trials
Plant Production. Seventeen cultivars, those used in the

field plus three more, were grown in the greenhouse using
the sail mix and cultural methods outlined in chapter 2.
There were six experimental units per cultivar consisting in
individual plants.
In the first greenhouse experiment (GH-1), seeds were sown

on March 24 and plants were inoculated on July 21, 1989. In
the second greenhouse experiment (GH-2), seeding took place
on August 24, 1989 and inoculation on January 13, 1990.
GH-1 experiment was conducted under natural summer lighting

conditions. In GH-2 conducted during the winter, the
photoperiod was increased to 16 hr by use of 400 watt sodium
(, lamps (Lumiponic, Inc.) set 1.5 m apart and 20 cm above the
l
83
1 canopy to provide a luminosity of 250-300 J,1Em- 2 sec- 1 under a
16 hr light regime.
Inoculation. Inoculation for the greenhouse studies was

carried out in a spray chamber (Boston Gear, InC'om
International, Inc.) which could accommodate six plants at a
time with their leaves spread out. The inoculum was applied
as a fine mist using a full cane nozzle (TG 0,7, Spraying
System, Ca.) with a reservoir mounted on a trolley in the
chamber. The sprayer pressure was set at 1800 kPa and
trolley speed at 0.75 kmph to deliver 200 ml inoculum
containing 200,000 spores/ml with each pass. The density of
pycnidiospores deposited on agar plates placed at canopy top
was 850-1030 spores/cm 2 , and 115-140 spores/cm 2 at pot
level. Spore germination, measured according to the
procedure outlined in chapter 2, was 100 % in GH-1 and 98.25
% in GH-2.
Inoculated plants were placed in wet plastjc bags and

incubated for 72 hr on a growth bench set at 20 oc and 14 hr
photoperiod. At the end of the incubation period, the
plants were transferred to the greenhouse for symptom
development. They were placed in plastic trays partly
filled with water ta prevent wilting during hot days.
Relative humidity was measured in the second trial only and
ranged from 55 to 80.
84
In GH-l, temperature was manitoreo with maximum-minimum

thermometers placed amang plants whereas in GH-2,
temperature and relative humidSty were monitored using a
hygrot:hermograph located among the plants on the bench.
Temperature in the two greenhous\~ trials varied as follows:
GH-l) maximum temperature = 31.21,\ minimum temperature =
18.71, mean temperature = 24.99; GH-2) maximum temperature
= 24.18, minimum temperature = 17.11 6, arll.l mean temperature =
20.97.
Evaluation of component. of re.i.t.nc\l'
Di ••••••• timation (SAODPC). Assessment af disease in the

greenhouse and the conversion of raw data ta SAUDPC was
conducted in the manner outlined for the field trial.
Assessments were made every three days fram the 10 th to the
25 th day post-inoculation in GH-1 and fram the 13 th to the
25 th day in GH-2.
M.an l •• ion area (MLA). At the completion of the

observation period, two lesions were cut at random from each
of the three uppermost leaves per plant (6 plants per
cultivar). Their width and length were measured in
millimetres with the aid of a stereo microscope. Lesion
area, expressed in square centimetres, was calculated by the
(
85
formula for the area of a circle (xr 2 ) , where r = 1/4 (width
+ length).
Spore density (SPD). Lesions used for the determination

of MLA were then immersed for 24 h in a 10 ml solution
containing 0.1 % formaldehyde to stop spore production and
for preservation. The timing was based on preliminary tests
showing that a minimum of 18 hr was required for the spore
concentration to stabilize following the immersion of
pycnidia. The density of pycnidiospores expressed as '/cm 2
lesion, was determined by averaging the results of four
spore counts with a haemacytometer on the solution from each
tube, multiplying this amount by the total volume and
dividing by the total area of lesions in the tube. This was
repeated five times for a total of 6 replicates (plant) per
cultivar.
Pycnidial density (peD). Pycnidia on les ions used ta

assess MLA and SPD were counted on the top side of the leaf
tissue. Imbedded pycnidia whose shape was visible with the
aid of a stereo microscope were included. In the second
experiment only, lesions were cleared by immersing 24 hr in
a solution of equal parts 95 % ethanol and glacial acetic
acid (Benedict, 1970) to facilitate counting. The total
number of pycnidia was divided by the total area of the six
lesions per plant and expressed as t/cm 2 lesion.
86
( Spore. par pyenidium (Sppe). The number of spores per
pycnidium was obtained by dividing total pycnidiospores by
the total number of pycnidia for each plant and cultivar.
Lateney pariod ('1'50 and '1'75) • The time from inoculation

until lesions covered 50 % and 75 % leaf area was used to
estimate the latency periode Incubation and latency period
for Seotoria apiicola are synonymous since a qualitative
test involving all cultivars showed that when lesions less
than twenty-four hours old are immersed in water for one
hour they release pycnidiospores that germinate on celery
juice agar.
(~ Tso and T75 were estimated by first transforming the leaf

area diseased on each sampling date to a proportion of the
maximum disease (PMD) observed on the 25 th day post-
inoculation. PMD was then regressed on time using the
angular transformation selected after comparison with the
Gompertz and logistic transformations for the value of R2
and the randomness of residual distribution in aIl
cultivars.
T50 and T75 were calculated as outlined in the following

steps:
(
87
1 1) relJression of y' on time, where y' = sin- 1 VPMD, and
estimation of the intercept and slope of the regression line
for the six experimental units (plants) per cultivar;
2) for each plant and cultivar, solving the equation
(3 .2)
for PMD=O.5 and PMD=O.75.
3.3.4 Data annlysis
Field data. Analyses of variance were performed on the

final disease severity, total SAUDPC and current SAUDPC
(SAUDPC for each interval between sampling dates), for
marked, random and the total (marked + random) number of
plants per plot, followed by Duncan's New Multiple Range
Test for means comparisons. The effect of the sampling
procedure on the disease severity was tested by treating the
experimer.t as a split plot wi th marked and random as
subplots. Cluster analysis and principal component analysis
were used to group cu~tivars according to their SAUDPC for
each sampling interval.
Greenhous. data. SAUDPC, MLA, PCD, SPO, SPPC, T50 and T75
were submitted to analyses of variance followed by Duncan's
..... New Multiple Range Tests. Multiple linear correlations were
88
( performed on components of resistance for each greenhouse
experiment to evaluate their interrelationships. Rank
correlations were used to compare results on individual
resistance parameters between greenhcuse trials, as well as
to verify their relationship with field results. Cluster
ana1ysis and principal component analysis were used to
summarise the greenhouse data for SAUDPC, MLA, PCD, and SPD
and get an overall classification to be compared with that
obtained for the field.
Statistical analyses were performed using SAS GLM, CLUSTER

and PRINCOMP procedures. The average linkage method was
used for cluster analysis and principal component analysis
( was performed on the correlation matrix of the descriptive
variables (Legendre and Legendre, 1984).
3.4 RESULTS
3.4.1 Field trial
Significant differences were observed among cultivars with

respect to final disease severity and the SAUDPC (Appendix l
and Figure 3.5). The cultivar S~okes Golden Plume,
Superdora and Summit had the lowest values for the SAUDPC.
The first two a1so had the lowest final proportion of 1eaf
89
area diseased (PLAD). Cultivars Surepak, Florida 683 and
Calmario had the highest SAUDPC and final blight severity.
Results of the split-plot analysis showed a significant
effect of the sampling procedure emphasized by the large gap
in SAUDPC between marked and random plants (Figure 3.5),
where the SAUDPC for randomly chosen plants was consistently
about two thirds lower than for marked plants. Superdora
had the lowest amount of disease among the randomly selected
plants. Its more average performance in the marked plants
indicates that it is sensitive to alloinfection to which the
markcd plants were subjected because of repeated
manipulation. Stokes Golden Plume, however, performed best
in the marked plants, under high inoculum pressure. The
ranking order, however, WdS not affected except in the
middle group of cultivars which did not differ significantly
from one another.
Cluster analysis of the SAUDPC for the three intervals from
0-25 days (SI), 26-40 days (52), and 41-55 days (S3) (see
data Appendix J) produced five classes in order of
increasing resistance indicated by the lower case characters
from 'a' through 'e' on the dendrogram (Figure 3.6; see
statistics in Appendix K). These groups were obtained by
sectioning the tree at 0.85 < R2 < 0.91 (see definition, Eq.
2.5) .
90
The position of the cultivars and their cluster affiliation
is shown in Figure 3.7 relative to the coordinate axes
represeT'l"'~d by principal component l (PC(l») and principal
component II (PC(2}). Together, PCn) and PC(2) account for 99
% of the total variation (Appendix L-l). The position of
cultivars in the two-dimensional plane reflects their
standard component scores relative to each principal
component (see component scores, Appendix L-2). Clusters
are indicated by the da shed lines on the graph. PC(l)
represented 74 % of the tccal variation. It was influenced
by the SAUDPC for tne three sampling intervals but mostly
during the second interval. PC(21 represented by the
vertical axis accounts for 25 % of the total variation. It
is mostly influenced by the SAUDPC during the first sampling
interval and to a certain extent by that of S3 (see
coefficients of the eigenvectors, Appendix L-l). The
negative sign next to S3 indicates that the effect of SAUDPC
during that interval 1S in the opposite directio~ relative
to PC(2). It separates Summit from cultivars Stokes Golden
Plume and Superdora on account of the former's large disease
increase during the last interval.
Characterization of cultivar field reactions from moderately

resistant through to very susceptible was done on the basis
of their position relative with respect to the axes formed
by PC n > and PC (2) According to this, cultivars Stokes
( •
91
1 Golden Plume, Superdora and Summit were considered
moderately resistant; Tendercrisp, Bishop, Vicar, Stokes
Improved Utah, Green Giant, Deacon, Ventura and Utah 52-70
were considered mùderately susceptible; Florida 683 and
Calmario, were susceptible, and Surepak, very susceptible.
The disease progress curves of the 14 cultivars are
illustrated in Figure 3.8 in groups corresponding to their
cluster affiliation. It shows the similarities and
differences in blighc increase during successive sampling
interval. Flgure 3.8a ~hows a Jarge difference in the rate
of disease increase during the second interval separating
the most susceptible ultivar Sur~pak (cluster 'a') from
Calmario and Florida 683 (cluster 'b'). The curves in
figure 3.8b and figure 3.Bc all belong to the moderately
susceptible cultivars (cluster 'c'). They differ. only
slightly in disease proportion on the 40 th day following
inoculation. Moderately resistant cultivars in figure 3.8d
differ from the latter by having less disease on either or
both of the last two sampling dates.
3.4.2 Greenhouse trials
Significant differences were found among cultivars relative
to all components of partial resistance except for the
latency period (T 50 and T75 ) (Appendices M and N) .

92
(
rigure 3.5. Ranking of cultivar with respect to the

standard area under the disease progress curve (SAUDPC)
under field conditions. The line represents the total
SAUDPC for marked and randornly selected plants. The filled
portion of the bars represents the SAUDPC for random plants
(~
only, and the clear portion, that of th~ marked plants
showing the effects of the sampling procedure.
93
Stk.Gold.Plume
- ~otal
Summit
Superdora o Marked
(22J ~.ndom
Tendercrlap
BIshop
Vicar
Ventura
.. '",
Deacon
... Utah 52-70

Stk.lmp.Utah
Green Glant
Florida 683
Calmarlo
Surepak
0.00 0.02 0.04 0.06 0.08 0.10
SAUDPC
94
A••~age Dbteaa. aet_ _ ClaRu.
1.. 1.2 1 O., O., O.. 0.2 0

+--------+--------+--------+--------+--------+--------+--------+. Florida 683
1
1 b L Calmario
1 Surepak
• 1
Superdora
L
• Stk.Gold.Plume
Summit.
d
Utah 52-70
k Deacon
Ventura
Green Giant
Bise op
~
a
Vicar
( Stk.lmpr. Utah
Tendercrisp
rigure 3.6. Grouping of celery cultivars into 5 levels of

partial resistance based on cluster analysis of the data on
the standard area under the disease progress curve (SAUDPC)
for the sampling intervals 0-25 days, 26-40 days, and 41-55
days following the introduction of inoculum. Lower case
characters from 'a' to 'e' identify clusters in order of
increasing partial resistance.
(
95
1 I I (81, -83)
--+---------+---------+---------+---------+---------+--
MS
4 + ~ +
1 !œ. ........ ----, 1
2 +, (P S J'11
.... ______ ,___ a +,
,,-, ,....-----, ".. ........ - ,,.,,
o +------\-M-~-7c-----DNU-~----~F-----Cj~--------~R~----+ l
1 d -." I.T BV K 1GJ ~ - - - -b .. ... 1
- '.... ,1 -
-2 + ~~------r VS +
MS 1
--+---------+---------+---------+---------+---------+--
-2 -1 0 1 2 3
(81, S2, 83)
., Figure 3.7. Principal cornponent analysis (peA) of cultivar

response under field conditions relative to the standard
area under the disease progress curve (SAUDPC) for the
sarnpling intervals 0-25 days (S1), 26-40 days (52), and 41-
55 days (S3) after inoculation. PCA was based on the
correlation matrix of descriptor variables (SAUDPC S1 -
SAUDPC S3 ) ' The coordinate axes l and II correspond to the

first two principal components. Variables in parentheses
had the highest loadings on the respective principal
components. ,MR=moderately resistant, MS= moderately
susceptible, and VS= very susceptible. Underlined lower
case characters identify cluster affiliation. R=Surepak,
F=Florida 683, C=Calmario, D=Deacon, N=Ventura, U=Utah 52-
70, G=Green Giant, K=Stokes Improved Utah, V=Vicar,
B=Bishop, T=Tendercrisp, S=Superdora, P=Stokes Golden Plume,
and M=Surnrnit.
96
Figure 3.8. Disease progress curves of celery cultivars

tested for partial resistance under field condition and
grouped according to cluster analysis of the SAUDP~ for
intervals between sampling dates. Fig. 3.8 a) represents
the rnost susceptible cultivars, clusters 'a': -0- Surepak,
and cluster 'b': -*- Florida 683 and -0- Calmario. Fig. 3.8
b) represents some moderately susceptible cultivars in
c cluster 'c': -0- Utah 52-70, -*- Deacon, -0- Ventura and -0-
Green Giant. Fig. 3.8 c) represents the balance of

moderately susceptible cultivars in cluster 'c': -0- Bishop,
-*- Vicar, -0- Stokes Improved Utah and -0- Tendercrisp.
Fig. 3.8 d) represents the curve of cultivars considered
moderately resistant, cluster 'd': -0- Summit, and cluster
'e': -*- Superdora and -0- Stokes Golden plume.
{
97
a) b)
PLAD - Cluater. 'a' & 'b'
0.4 .;....::.:...;::...-~:;.:.:.~.,..;:.;....;:....;;.---~-----, 04 ;.P~L~AD=--_C~Q~.t=.~r~·c~·________.____________~
0.1 . • . 0.3
0.1 . . . . . . . . • 0.1
0.1 . • •. •• .,. 0.1
~---------=~~--------~--------~ 0.0 ' - - - - - - - - - , : . . . - - - - - - - - . - - - - - - - - "

10 .. 40 Il 10 Il 40 Il
/1 day. trom Inoculation /1 dlYs trom inoculation
c) d)
PLAD - Cluater 'c'
~4';"":~:"'-~:;':':'~~--------------------, 04T
p.~l~AD=--_CW==.~œ=r~.~'~d·~'~·.~·____~--------_,
o.a " •.•. • 0.3 •• • ••••
0.1 •.•••••••••.• 0.2
~1 •••.•.•• .• 01
O~'------------~---------~--------~Iii OO'------------~------_,r_----·----J
.11
10 Iii 40 10 Iii 40
/1 daya trom Inoculation /1 deys 'rom Inoculation
98
{ Correlations among resistance parameters within individual
greenhouse trials are reported in table 3.1 and table 3.2.
In GH-l, there was a positive relationship between MLA and
SAUDPC which was not observed in GH-2. This suggests a
stronger influence of lesion size on the SAUDPC in GH--l,
while in GH-2, this parameter was more dependent on
illf~ction frequency. There was a strong positive
relationship between spore density (SPD) and the number of
spores per pycnidium (SPPC). On the other hand, the
relationship was weak but significant between SPD and
pycnidial density (PCD) in GH-1 and even weaker in GH-2
indicating more variability in PCD than in SPD among
cultivars.
(
In the comparison of results between the two greenhouse
trials using the Kendall tau rank correlation coefficient,
only MLA showed a significant positive relationship (Table
3.3) . When compared to field ranks, GH-l ranks showed no
significant correlation except for SAUDPC. In GH-2 results
showed a positive correlation with field ranks in the case
of SAUDPC, MLA and PCD.
Rankings of cultivars with respect to SAUDPC, MLA, PCD, and

SPD in both greenhouse trials were compared graphically to
each other and to field rankings (Figures 3.9 to 3.12). On
{ these graphs, the cultivars are ordered according to

99
1 increasing field susceptibility to highlight any
relationship which may exist between component of resistance
studied in the greenhouse and cultivar reactions in the
field.
SAODPC. In GH-l and GH-2, cultivars Stokes Golden Plume,

Superdord and Summit had the lowest SAUDPC and Utah 52-70
the highest (Figure 3.9). There was poor correlation
between the two greenhouse trials for the SAUDPC (Table 3.3)
which is evident here from the frequent 5witches in ranks
between GH-l and GH-2 but both showed positive correlation
with field results. What is striking here, is that it is
mostly the cultivars with moderately resistant reactions in
the field that are adequately represented by the greenhouse
SAUDPC. Sorne moderately susceptible (Utah types) and very
susceptible cultivars (Surepak) in the field ranked almost
opposite in the greenhouse. It should be remembered though
that the greenhouse SAUDPC comprises only one dlsease cycle
while the field SAUDPC comprises many.
Mean lesion area (MLA). There was a weak but significant

correlation in cultivar rankings between greenhouse trials
with respect ta MLA but only the results in GH-2 were
positivel:" correlated to cultivar response in the field.
Comparison of results from GH-1 and GH-2 (Figure 3.10) shows
that in GH-l, lesions tended to be smaller, ranging in size
100
( from 0.24 to 0.44 cm 2 , whereas in GH-2, their sizes varied
between 0.31 and 0.76 cm 2 • T~is difference could possibly
have resulted from the differences in mean temperatures and
lighting conditions between experiments. The Utah cultivars
and Green Giant had the largest lesions in both experiments.
They, along with Calmario and Surepak showed a larger than
average increase in lesion size between experiments, while a
few others showed little or no variation, suggesting
differential cultivar responses to environmental changes.
Comparison of greenhouse results for MLA and SAUSPC (Figures
3.9 and 3.10; Tables 3.1 and 3.2) shows a positive
relationship between MLA and SAUDPC in GH-1 but not in GH-2,
indicating that in GH-2, SAUDPC might have depended more by
{ infection frequency.
Pycnidial denaity (PCD). The most striking aspect of

pycnidial densities (Fig~re 3.11) is the large discrepancy
in results between greenhouse trials. PCD ranged from 117
to 354 per cm 2 in GH-1, and from 544 to 902 per cm2 in GH-2.
A good portion of the observed difference may be explained
by the fa ct that les ions were cleared before counting
pycnidia in GH-2, which probably resulted in the counting of
imbedded or small pycnidia that would have otherwise been
missed. However, the effect of environmental factors, in
particular lighting which is known to affect pycnidial
(• formation in S. apiicola (Corchran, 1932) should not he
101
,J
'l'able 3.1. Partial correlation coefficients arnong components of

partial resistance for greenhouse trial l, Summer, 1989.
SAUDPC MLA PCD SPD SPPC T~o
SAUDPC 1 0.613 0.368 0.348 0.332 -0.168

0.0001 0.001 0.002 0.003 0.135
MLA 1 0.209 0.264 0.373 -0 .190
0.061 0.017 0.001 0.085
PCD 1 0.462 0.219 0.075
0.0001 0.049 0.505
SPD 1 0.705 -0.027
0.0001 0.814
SPPC 1 -0.131
0.244
T~o 1
The number on the second line represents the Prob > Irl. df = 79.
102
(
Table 3.2. Partial correlation coefficients among components of

partial resistance for greenhouse trial 2, Winter, 1990.
SAUDPC MLA PCD SPD SPPC T!o
SAUDPC 1 0.194 0.187 0.158 0.101 0.174

0.083 0.094 0.158 0.370 0.121
MLA 1 -0.124 -0.240 -0.177 0.037
0.269 0.031 0.114 0.741
PCD 1 0.293 -0.126 -0.046
0.008 0.261 0.684
SPD 1 0.856 0.450
0.0001 0.0001
SPPC 1 0.479
(~ 0.0001
T so 1
The nunber on the second line represents the Prob> III. df - 79.
(
103
1
'fable 3.3. Kendall' s tau rank correlation coefficients bet"'een field
cultivar rankinqs for SAUOPC and qreenhouse rankings for individual
components of resistance, and between the two qreenhouse trials (GH-1
and GH-2) .
Components Field SAUDPC Field SAUDPC GH-1

of resistance vs GH-1 vs GH-2 vs GH-2
SAUDPC 0.420 0.736 0.229

0.037 0.0002 0.201
MLA 0.187 0.648 0.353
0.352 0.001 0.048
PCV 0.121 0.582 0.147
0.547 0.004 0.41
SPD 0.187 0.253 0.118
-
N ....
SPPC
0.352
-0.01l
0.956
0.208
0.077
0.702
0.510
-0.176
0.323
Tso -0.297 0.275 0.059
0.139 0.171 0.741
TH -0.099 0.253 0.162

0.622 0.208 0.365
Numbers on the second line represent the Prob > IRI under Ho: Rho-O,
N=14.
104
Stk.Gold.Plume
Summit .0 Field
Superdora GH-1
Tendercrisp GH-2
Bishop
VI car
Deacon
Ventura
Stk.lmp.Utah
Utah 52-70
Green Glant
{ Florida 683
Calma rio
. 1
Surepak
0.02 0.04 0.06 0.08 0.10
SAUDPC
Figure 3.9. Comparison of celery cultivar rankings

relative to the SAUDPC between the first and second
greenhouse trials (GH-l & GH-2) , and with the field ranking
for SAUDPC. ~lltivar order follows increasing field
susceptibility (bars).
l
105
0.02 0.04 0.06 0.08 0.10

Gold. Plume _-:/r
Summlt
Superdora
Tendercrlsp
.oor;;;
--
~
"-
0
-
Field SAUDPC
GH-1
z.J - GH-2
Bishop ~
"-
Viear \.
Ceaeon
-
, )
-
'---
"-
Ventura '-
~
Stoke's Utah
Utah 52-70 ".,>
Green Glant ., .,
Florida 683 /' ~
-~
/
Calmario <
'\.
Surepak -'
~
Gld. Self. Blnch.
Florlda K-str.
Utah R-str. 1
0.10 0.20 0.30 0.40 0.50 0.60 0.70 0.60
Mean les ion area (cm2)
Figure 3.10. Comparison of celery cultivar rankings

relative to the mean lesion area (MLA) between the first and
second greenhouse trials (GH-l and GH-2) , and with the field
ranking for SAUDPC. Cultivar order follows increasing field
106
Stk.Gold.Plume
Summlt CJ FI.ld SAUDPC
Superdora GJi-1
Tendercrlsp
G~-2
BIshop
Vlca"
Ventura
Deacon
Utah 52-70
Stk.lmp.Utah
Green Glant
Florida 683
( Calmario
Surepak
Gld.Self Blnch.
Floride K-atr.
Utah R-str.
100 200 300.00 500 800 700 800 900100011001200
Pycnidial density (lcm2 lesion)
l'iqure 3.11. Comparison of celery cultivar rankings

relative to the pycnidial density (PCD) in the first and
second greenhouse trials (GH-l and GH-2), and with the field
ranKing for SAUDPC. Cultivar order follows increasing field
(,
107
Stk.Gold.Plume
Summlt Fld SAUDPC
Superdora
I----.-;-----;--~~~=- OH-,
Tendercrlsp
OH-~
BIshop
Vicar
Ventura
Deacon
Utah 52-70
Stk.lmp.Utah
Oreen Glant
Florida 683
Calmario
Surepak
Gld.Self Blnch.
Florlda K-atr.
Utah R-str.
0.00 0.50 1.00 1.50 2.00 2.50
Spore density (. 10"6 Icm2 lesion)
l'igura 3.12. Cornparison of celery cultivar rankings

relative to the spore density (SPD) in the first and second
greenhouse trials (GH-l and GH-2), and with the field
ranking for SAUDPC. Cultivar order follows increasing field
- susceptibility (bars).
108
entirely dismissed as a source of variation. Only GH-2
cultivar rankings for PCD showed a positive relationship
with field results. In this experiment, the three most
resistant cultivars in the field, namely Stokes Golden
Plume, Summit and Superdora, had the lowest PCD. In GH-1,
cultivars were basically divided into two groups, one group
having less than 200 pycnidia per cm2 lesion and the
majority having between 280 and 350 pycnidia per cm 2 lesion.
GH-l cultivar ranking for PCD showed no pattern
corresponding to results in either GH-2 or the field.
Spore denaity (SPD). The average spore density did not

vary between greenhouse experiments in response to
environmental changes as did other components of resistance
(Figure 3.13). However, sorne cultivars showed very large
differential responses to the change in environmental
conditions between trials while others varied little or not
at aIl. There was no correlation between the two greenhouse
trials with regards SPD and no relationship with field
results. This is likely due to the enormous amount of
inoculum being produced by aIl cultivars, in the order of
10 6 /per cm 2 lesion, such that statistical differences do
not translate into epidemiological differences.
Denaity of spore. par pycnidium (SPPC). There were

significant d!fferences among cult~.vars in both greenhouse
109
experiments. Values ranged from 4095 to 6992 pycnidiospores
per pycnidium ln GH-1 and from 1054 to 2604 in GH-2. SPPC
was strongly correlated to spore density within experiments
(Tables 3.1 and 3.2) . In GH-1 Summit, Vicar and Golden
Self Blanching had the lowest spore density per pycnidium
and Deacon follawed by Vicar, Utah R-strain and Green Giant
had the lowest in GH-2. Since there is no apparent
relationship between spore density and cultivar
susceptibility in the field, SPPC does not present any real
interest as will be discussed later.
Lat.ney period ('1'50 and T,S). There were no significant

differences among cultivars with respect to T~ and T75 in
either greenhouse experiment. In GH-1, Golden Self
Blanching had the shortest T50 (16.06 days), followed by
Surepak (16.28 days) which had the shortest among cultivars
tested in the field. The same two cultivars had the
shortest T75 in GH-1 (18.90 and 19.10 days). The longest T50
and T75 in GH-1 were recorded for Bishop (16.90 and 19.69
days) . In GH-2 the sit~ation was different. Utah 52-70
had the shortest T50 (14.59 days), but Summit had the
shortest T75 (17.54 days) with Utah 52-70 close second
(17.69 days). Tendercrisp had the longest T50 and T75 in GH-
2 (15.68 and 18.90 days) .
110
( Overall, in GH-1, the mean values for T50 and T75 were 16.52
± 0.50, and 19.34 ± 0.56, respectively whereas in GH-2, the
values were 14.90 ± 0.57 for T50 , and 18. 08 ± 0.63 for T75 •
It appears from these results that variation between
experiments was greater than within suggesting that
environmental conditions during colonization were more
influential than cultivars in determining the duration of
the latent period for Septoria blight.
3.4.3 Multiv.riate an.ly •••.
The resistance parameters selected as descriptive variables

for multivariate analyses were the SAUDPC, MLA, PCD and SPD.
The density of spores per pycnidium (SPPC) was not included
because of redundancy with pycnidial density and spore
density. Tso and T75 were also excluded for lack of
significance.
Cluster analysis of the GH-1 data categorized cultivar

responses into six resistance classes (Figure 3.13) by
sectioning the dendrogram at 0.66 < R2 < 0.78 (Appendix K).
Clusters are identified by the lewer case letters from 'a'
te 'f' in order of increasing partial resistance.
Interpretation of the dendrogram is best accomplished by

reference to results from the principal component analysis
(
111
(PCA). For GH-l, the proportion of the total variation
accounted for by each principal component and the component
loadings (contribution of the resistance parameter to each
component) are listed in Table 3.4. Together, principal
component l (PC(l) and principal component II (PC(2))
accounted for 85 % of the total variation. The parameter
with the greatest influence on pe(l) and pe(2) are indicated
on the graph showing the position of cultivars and cluster
affiliation (Figure 3.14) .
From the position of the cultivars along the coordinate axes

described by PCu) and PC(2) and the angles they forrn with
them, Superdora (cluster 'f') would be considered
moderately resistant. The cultivars in clusters 'c', 'd',
and 'e', would be termed moderately susceptible, those in
cluster 'b', susceptible, and Utah 52-70 (cluster 'a'), very
susceptible.
Six classes were aiso obtained from cluster analysis of data

for the same resistance parameters in GH-2, by sectioning
the dendrogram at 0.82 < R2 < 0.87 (Figure 3.15, Appendix
Q). The clusters are again identified by lower case letters
in order of decreasing susceptibility.
Cumulative proportions of each successive principal

components and contributions of the resistance parameters to
112
t each eigenvector in GH-2 PCA results are listed in table
3.5. The cumulative proportion of the total variation
accounted for by the first two principal components was 85
%. PC(1) was influenced equally by SAUDPC, MLA, and PCD and
to a lesser extent by SPD. PCD had by far the greatest
influence on PC(2) with a component loading of 91 %.
An overall ranking of cultivars for resistance in GH-2 was

determined on the basis of their positions relative to the
axes described by the first two principal components (Figure
3.16). Accordingly, cultivars in cluster 'f' appear
moderately resistant, those in clusters 'e' and 'd' would be
termed moderately susceptible while cultivars in clusters
'c', 'b', and 'a' are considered susceptible. Because spore
density contributed most to PC(2) but showed no correlation
to field results while SAUDPC, MLA, and PCD were positively
correlated to field results in GH-2, more importance is
given to the separation of clusters along the horizontal
axis of the graph in the characterization of cultivar
resistance. What differentiated cultivars in clusters 'a'
and 'b' from each other and those in group 'c' is their
extreme values for SPD particularly in the case of Stokes
Improved Utah, resulting in high leverage (Figure 3.12) .
113
1.. 1.2 1 0.1 0.1 O.. 0.2 0

+----+----+ 1 - '--~I------+I----~I~"'---+.
Florida 683
~ Call1lario
Florlda K-.traln
b
Stokes Improved Utah
Utah R-strain
Deacon
~
c
~ Vlcar
Ventura
Blahop
Tendercr1sp
Summit
d
Superdora
t
Surepak
~
Green Gi&nt
- •
Stokes Golden Plume
Golden Sel! BlanchlnQ
Utah 52-70
Figure 3.13. Hierarchical classification of 17 celery

cultivars based on their response in greenhouse trial 1
relative to SAUDPC, MI,A, PCD, and SPD. Lower case
characters from 'a' to 'f' identify clusters in order of
increasing partial resistance.
114
-(
'l'able 3.t. Principal ccmponent analysis of the data on the SAUDPC,

MLA, PCD, and SPD as parameters used in the eva1uation of 17 celery
cultivars for partial resistance in the first qreenhouse trial (GH-1).
Eiqenvalues of the Correlation Matrix

Eiqenvalue Difference Proportion Cumulative
PRIN1 2.13769 0.887258 0.534423 0.53442
PRIN2 1.25043 0.846576 0.312608 0.84703
PRIN3 0.40386 0.1951334 0.100964 0.94799
PRIN4 0.20802 0.052006 1. 00000
Eiqenvectors
( PRIN1 PRIN2 PRIN3 PRIN4
SAUDPC 0.549458 0.349680 -.646856 -.396733
MLA 0.144326 0.837930 0.318601 0.418970
PCD 0.570562 -.390380 -.158710 0.704893
SPD 0.593064 -.152317 0.674449 -.412544
(
115
..
l'iqure 3.14. Principal component analysis (PCA) of

cultivar response in greenhouse trial 1 relative to SAUDPC,
MLA, PCD and SPD. PCA was performed on the correlation
matrix of descriptor variables. MB = moderately resistant,
H[ = moderately susceptible, and ~ = very susceptible.
Dashed circles represent clusters identified by underlined
f
lower case letters. U=Utah 52-70, Y=Utah R-strain, K=Stokes
f
Improved Utah, F=Florida 683, X=Florida K-Strain,
1 C=Calmario, D=Deacon, T=Tendercrisp, B=Bishop, N;;:::Ventura,

1 V=Vicar, M=Summit, G=Green Giant, R=Surepak, P=Stokes Gclden
Plume, Z=Golden Self Blanching, S;;:::Superdora.
116
II (MLA)
--+---------+---------+---------+---------+---------+---
2+ MS 1 a +
1 1;0' 1
1 l '...~I 1
,.. ,1
~ ",--_ ...... '" VS 1
... - G ~ _---... , 1
,\ ...s'.1 ",;' Z 1 ... , " y ; 1
... , , 'K ... " 1
( R " : /,tI' !!. 1
1 \'P ...." : F { 1
~~---.~~ l'
o +-------------------------------+---T---~--------------+ l
1 .......1-" \ x \ 1
D ',\ c J
1
Ma , .," 1 , ...... 1
1 l ,
S IV 1 \.
(~
lM'.
' . . ' " ',N 1
1
T', \ c
...... .J
1 -.. B
......... _-_ .. ., ,1
-2 + MS +
--+---------+---------+---------+---------+---------+---
-3 -2 -1 0 1 2
(SAUDPC, PCD, SPD)
117
1 •• 1.2 1 o•• o•• o.• 0.2 o

+----+-- 1 1 1 1 1
•
Floria.. 1513
~
r-
Utah ~2-70
a Utah R-Itnln
Calmarlo
b
Oe.con
•
Bhhop
d
---c-t Tendeccc isp
Ventun
Floc ia. K-at ra ln
Golden Selt BlanchlnQ
Stokes Improved Utah
• Superdora
- f
1
•
SUl!llllt
Stokes Golden Plume
riqur. 3.15. Hierarchical classification of 17 celery

cultivars based on their response in ghreenhouse trial 2
relative to SAUDPC, MLA, PCD, and SPD. Lower case
characters from 'a' to 'f' identify clusters membership in
order of increasing partial resistance.
c 118
'l'able 3.5. Principal component analysis of the data on the SAUDPC,

MLA, PCD, and SPD as pararneters used in the evaluation of 17 celery
cultivars for partial resistance in the second greenhouse trial (GH-2).
Eigenvalues of the Correlation Matrix

Eigenvalue Difference Proportion Cumulative
PRIN1 2.52101 1.62565 0.630252 0.63025
PRIN2 0.89536 0.53959 0.223841 0.85409
PRIN3 O. 3557~' 0.12791 0.088943 0.94304
PRIN4 0.2278ti 0.056964 1.00000
Eigenvectors
PRIN1 PRIN2 PRIN3 PRIN4
( SAUDPC 0.557576 -.295025 0.093711 0.770252
MLA 0.544187 -.287544 0.544047 -.570257
PCD 0.547889 0.081261 -.787720 -.269649
SPD 0.304611 0.907566 0.273363 0.093858
119
Figure 3.16. Principal component analysis (PCA) of

cult var performances in greenhouse trial 2 relative to
SAUDPC, MLA, PCD and SPD. PCA was based on the correlation
matrix of descriptor variables. MB=moderately resistant,
Mli= moderately susceptible, S=susceptible and Yli= very
susceptible. Underlined lower case characters identify
cluster affiliation. K=Stokes Improved Utah, U=Utah 52-70,
R=Surepak, Y~Utah R-strain, C=Calmario, F=Florida 683,
G=Green Giant, Z=Golden Self Blanching, X=Florida K-Strain,
N=Ventura, T=Tendercrisp, B=Bishop, V=Vicar, D=Deacon,
S=Superdora, M=Summit, P=Stokes Golden Plume
120
c
I I (SPD)
-+------------+------------+------------+------------+--
U~ ,-\
2+ no .... ' tR,.!, +
1 ,'Z i', '-" 1
1 \ 1 \ 1
1 .4 \ 1 \ 1
1
1
1 1
1 1 :
X' 1
1
1 1 NI 1 1
," 1
1 ... "" ' .. l, .... _ - __ " VS 1
1 lS \ !. :8 l /F U" 1
o +---~---~----------~--T-L-+---J------------~----------+
\ ' ... _.... 1 \ l
1 \ PM , 1 1 R 1 1
MIl \."'..... 'J 1
.!. - ... --, " Y./ ~ 1
,'" V, ',C 0#'" 1
.'~D ,,1 ' ..... __ ., 1
,_~, 1
1
~ 1
-2 + MS l'a\
'.1 +
-+------------+------------+------------+------------+--
-2 -1 0 1 2
(SAUDPC, MLA, PCD)
(
121
3.5 DISCUSSION AND CONCLUSION
Three cultivars, Stokes Golden Plume, Summit and Superdora,

appeared more resistant to Septoria blight than others under
field conditions. The same cultivars tended to perform
weIl also under greenhouse conditions and were among the
least affected by changes in environmental conditions
observed between the two greenhouse experiments. The
difference between Summit and the other two cultivars was a
sudden increase in disease during the last sampling interval
which could be due to the increased rate of the epidemic or
most likely to a 10ss of resistance with maturity, two weIl
recognized phenomena (Parlevliet, 1979; Van der Plank, 1963,
Ch. 20). These three cultivars were most efficient at
slowing the rate of epidemic. They are thus potential
candidates for future investigations into the variation of
fungicide dosages and frequency to account for their level
of partial resistance relative to Florida 683 used in the
development of the infection model described in chapter 2.
Another potential use of resistance would be to correct the
predicted disease severity index of the infection model for
host resistance.
122
There were cultivars whose field and greenhouse reactions

appeared poorly related. The first, cultivar Surepak, had a
highly susceptible reaction under field conditions but was
ranked moderately susceptible in GH-l, and susceptible in
GH-2. Another is, Utah 52-70, moderately resistant under
field conditions but susceptible under greenhouse
conditions. These are cultivars with very large lesions
with a similar SAUDPC in the second greenhouse experiment.
That parameter was based on a single disease cycle in the
greenhouse whereas it represents many in the field. Thus,
it would appear that Surepak has a much higher infection
frequency than Utah 52-70 under field conditions, which is
compounded over successive disease cycles.
{
Of the cultivars tested only under greenhouse conditions,
Golden Self Blanching was moderately resistant in both
trials. Florida 683 K-strain was moderately resistant in
GH-l and susceptible in GH-2, while Utah R-strain was ranked
among susceptible cultivars in both trials. No prediction
can be made, however, about their potential field behaviour.
The component of partial resistance most highly correlated

to field results in both greenhouse trials was the SAUDPC .
There was also a positive relationship between rankings for
MLA and PCD in the second greenhouse trial and field
( ranking. The SAUDPC in conjunction with lesion size could

123
1 he used with caution to screen for above average rate

reducing resistance under greenhouse or controlled
environment studies. Its disadvantage, however, was to
overestimate the susceptibility of cultivars with large
les ions particularly used with uniform coverage and high
spore densities.
The enormous sporulation potential of ~. apiicola on aIl

cultivars was seen as the reason why, in spite of
statistically significant differences among cultivars, their
ranks with respect to SPD in either greenhouse trials
appeared unrelated to those in the field. Another reason
may be the protective nature of pycnidia and the conidial
matrix which prevent pycnidiospore germination unless
diluted (Louis & Cooke, 1985). Furthermore, only a portion
of spores enclosed in pypcnidia are likely to be released at
a given time in bouncing or dripping raindrops resulting in
the continuous presence of available spores regardless of
the level of production. This in turn, suggests that
differences in pycnidial density may have greater
epidemiological implications than differences in spore
density. The positive association of rankings for PCD in
GH-2 with field rankings supports this.
Groupings achieved by cluster analysis and ordination of the

components of partial resistance from the second greenhouse
.....
124
( study were in closest agreement with those in the field.
It is thought that the poorer relationship between the first
greenhouse results and field results may be due to the
warmer conditions experienced in the greenhouse during the
summer, where the mean maximum temperature was 31°C in
comparison to 24 oC during the winter experiment. ~.
apiicola being a cool climate pathogen, it is inhibited at

temperatures above 30 oC as determined in chapter 2. Thus,
cultivar response in the first greenhouse trial may be more
a reflection of pathogen inhibition than actual resistance.
Temperature also had a greater influence on the latency

period than did cultivars as demonstrated earlier. Although
it is one of the most important components of rate-reducing
resistance (Berger, 1977; Parlevliet, 1979), none of the
commercial cultivars tested differed significantly in their
ability to interfere with the growth of Septoria in the
hosto Differences of even one day in the latency period
are said to have an ep1demiological impact (Fry, 1982, Ch.
11). Could it be that in this case statistically
insignificant differences may have a significant
epidemiological impact? It may be in certain cases where
the average latency period is shorter, but it is doubtful
though not impossible under optimal conditions for the
pathogen, that enough disease cycles occurred during one
(
125
1 eropping season to make a differenee in the final disease

level.
Marking plants for repeated assessments over the duration of

the epidemic, though helpful in following disease
progression on a given plant, does not constitute a good
sampling method because of the excessive inoculum pressure
ereated by repeated manipulation. Fortunately, it did not
affect cultivar ranking or their classification into
specifie resistance categories. A better sampling method
would make use of spatial distribution by selecting plants
randomly in concentrie areas equidistant from foei of
infection. It would increase the quantity and quality of
the information eolleeted and would allow spacio-temporal
analysis of disease spread. (Campbell & Madden, 1990, Ch.
11) .
Multivariate analyses of the field data helped to get around

the usual overlapping that results from means comparison
tests and to classify disease progress curves according to
their shape. With the greenhouse data, principal component
analysis was most helpful as an aid to the interpretation
of groups formed by cluster analysis by providing an overall
picture of cultivar responses to the selected components of
partial resistance. The use of elusters also helped to
overcome a weakness of non-parametric correlations which,
126
1 because they are based on ranks rather than resemblance,

sometimes fail to detect a true association. This was
demonstrated with the parameter SAUDPC, common to field and
greenhouse studies, where the ranks between greenhouse
trials had nothing in common but in both cases agreed with
field results. This was attributed to cultivars with no
significant differences exchanging ranks between trials.
Sorne major criticisms of principal component analysis are

that it considers only linear relationships among
predictorsi that it gives equal weight to aIl predictors,
particularly when performed on correlation matrices as was
the case here to deal with different scales of measurements,
{ and that it is sensit~ve te outliers (James & McCulloch,
1990; Legendre & Legendre, 1984). AIso, the interpretive
value of PCA can sometimes be undermined by a high degree of
correlation among descriptors resulting in high component
loadings on more than one principal components. This could
be improved by assignment of weights to components of
resistance according te their epidernielogical importance but
the difficulty would then be te determine appropriate
weights. In spite of these weaknesses, man y workers find
multivariate analyses very attractive techniques in the
synopsis of resistance trial results and as an aid to their
interpretation (Jeger, 1980; Lebeda and Jendrulek, 1988;
Platt and Tai, 1984; Thompson and Rees, 1979).
-{
127
l LITZRATURZ CITZD
Bedlan, G. 1985. [Septoria leaf spot disease of celery.] Die

Septoria-Blattfleckenkrankheit des Selleries.
Pflanzenschutz 7:5-7.
Benedict, W.G. 1970. DifferentiaI effect of light intensity
on the infection of wheat by Septoria tritici Desm. under
controlled environmental conditions. Physiol. Pl. Path.
1:55-66.
Florida State Horticultural Society 83:208-212.
Berger, R.D. 1980. Measuring disease intensity. Proc. E.C.
Stakman Commemorative Symposium on Crop loss Assessment
(P.S. Teng and S.V. Krupa. eds.) Univ. of Minnesota Mise.
Publ. 7. St. Paul. pp 28-31.
Browning, J.A., Simmons, M.D., and E. Torres. 1978. Managing
host genes: Epidemiologie and genetic concepts. In: "Plant
Disease: An Advanced Treatise" (J.G. Horsfall and E.B.
Cowling, eds.), Vol. 1, pp 191-212. Academie Press, New
York.
Campbell, C.L. and L.V. Madden. 1990. Introduction to Plant
Disease Epidemiology. John Wiley & Sons. New York and
Toronto. 532 pp.
Carisse, O. 1989. Spore Production, Factors Influencing
Infection and Determination of a Disease Threshold for
Cercospora Blight of Carrot. M. Sc. Thesis, McGi11
1l, University. 128 pp.
1 Cochran, L.C. 1932. A study of two Septoria leaf spots of
1 celery. Phytopathology 22:791-812.
1
CPVQ. 1987. Légumes - Protection. AGDEX 250-605. Conseil
des Productions Végétales du Québec. Ministère de
l'Agriculture, des Pêcheries et de l'Alimentation du Québec.
112 pp.
Methods and Applications. John Wiley & Sons, Inc. NY.
Fry, W.E. 1978. Quantification of general resistance of
potato cultivars and fungicide effects for integrated
control of potato late blight. Phytopathology 68:1650-
1655.
' .
128
( Fry, W.E., A.E. Apple and J.A. Bruhn. 1983. Evaluation of

potato late blight forecasts modified to incorporate host
73:1054-1059.
Hewett, P.D. 1968. Viable Septoria spp. in celery seed
samples. Ann. appl. Biol. 61:89-98.
Holley, J.D., Hall, R., and G. Hofstra. 1983. Identification
of rate-reducing resistance to early blight in potato. Cano
J. Plant Pathol. 5:111-114.
Jeger, M.J. 1980. Multivariate models of the components of
partial resistance. Protection Ecology 2:265-269.
Kleinbaum, D.G., Kupper, L.L., and K.E. Muller. 1987.
2nd Ed. PWS-KENT Publishing Co. Boston. 718 pp.
Lebeda, A. and T. Jendrulek. 1988. Application of methods of
multivariate analysis in comparative epidemiology and
research into field resistance. Journal of Plant Diseases
and Protectjon (PfIKrankh.) 5:495-505.
Legendre, Land P. Legendre. 1984. Écologie numérique. 2. La
(~ structure des données écologiques. 2nd Ed. 335 pp.
Louis, l., and R.C. Cooke. 1985. Conidial matrix and spore
germination in some plant pathogens. Trans. Br. mycol. Soc.
84:661-667.
Maude, R.B. 1964. Studies on Septoria on celery seed. Ann.
appl. Biol. 54:313-326.
Ann. dppl. Biol. 65:249-254.
Maude, R.B. and C.G. Shuring. 1970. The persistence of
Septoria apiicola on diseased celery debris in soil. Pl.
Path. 19:177-179.
Munger, H.M. and A.G. Newhall. 1953. Breeding for disease
resistance in celery and cucurbits. Phytopathology 43: 254-
259.
Nelson, R., and M. Mooar. 1954. Comparative resistance to
leaf blight diseases in sorne commercial varieties of celery
in 1952 and 1953. Michigan Quarterly Bulleting. 36 (3) 275-
279.
( Padgett, G.B., Nutter, F.W. Jr., Kuhn, C.W. and J.N. AlI.
1990. Quantification of disease resistance that reduces the
129
t, rate of Tobacco Etch Virus epidemics in bell pepper.
phytopathology 80:451-455.
Parlevliet, J.E. 1979. Components of resistance that reduce
17:203-222.
Platt, H.W. and G.C.C. Tai. 1984. Assessment and analyses
for the interpretation of potato late blight response in
field studies. American Potato Journal 61:599-609.
Macmillan Publishing Company. New York. London. 263 pp.
Shaner, G., and R.E. Finney. 1977. The effect of nitrogen
fertilization on the expression of slow-mildewing resistance
in Knox wheat. Phytopathology 70:1183-1186.
Sheridan, J.E. 1968. Conditions for germination of
pycnidiospores of Septoria apiicola Speg. N.Z. JI Bot.
6:315-322.
Thompson, J.P. and R.G. Rees. 1979. Pattern analysis in
epidemiological evaluation of cultivar resistance.
Valk, M., Knibbe, E., Burbidge, H., and P. Flinn. 1987.
Muck Vegetable Cultivar Trials and Research Reports.
Research Report # 37. Horticultural Research Institute of
Ontario. Ministry of Agriculture and Food of Ontario. 72
pp.
Wright, J.C., and M.L. Lacy. 1988. Increase of disease
resistance iD celery cultivars by generation of whole plants
from cell suspension cultures. Plant Disease 72:256-259.
Zadoks, J.C. and R.D. Schein. 1979. Epidemiology and Plant
Disease Management. Oxford University Press. New York. 423
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Zitter, T.A. 1983. A Listing of Vegetable Varieties with
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Cooperative Extension, Cornell University Department of
Plant Pathology. Ithaca, New York. p.6
130
(
4 G.NZRAL DISCUSSION AND CONCLUSION
Successfu1 disease management calls for the use of all

available methods to reduce the amount of initial inoculum
and slow down the rate of reproduction of pathogens such
that their populations are maintained below economic in jury
levels for the duration of a crop (Fry, 1982). It implies
that a strategy is deve10ped to discourage the emergence of
pathogen populations that become resistant to treatment and
overcome plant resistance. Ultimately, it must be conducted
economically and in a manner which lirnits environmental and
health risks.
(
Septoria blight of celery is a disease with a high
destructive potential in humid temperate regions. Rapid
increases in disease developrnent have been linked to cool
rainy weather (Berger, 1970). Sorne environmental conditions
which limit or favour spore germination and infection are
known (Berger, 1970; Corchran, 1932; Sheridan, 1968) as weIl
as measures to control primary inoculum (Maude, 1970) and
disease in the field (CPVQ, 1987). However, to reduce the
dependency on fungicides in the field without taking the
risk of losses, it is necessary to have more precise
information on the factors that promote rapid disease
( increase. Models which serve to elaborate strategies for

131
disease management can be derived empirically from existing

historical data or developed from fundamental studies
involving experimentation under controlled conditions. In
the first part of this work it was decided to quantify the
infection as a function of temperature and leaf wetness
under controlled conditions as this basic step has proven
highly successful in model development {Jones, 1986).
Results have shown that the development of Septoria is
optimal at 25 oC in combinat ion with leaf wetness periods of
72 hr or more. Growth was severely limited at temperatures
below 10 oc regardless of wetness duration and again at 30
oC where increases in the duration of wetness beyond 48 hr
completely inhibited growth.
There are numerous methods, varying in complexity, now being

used in epidemiology to describe biological relationships
mathematically, or statistically (Hau & Kranz, 1990).
Where the objectives are to develop a simple forecast model,
linear regression can be used provided that the data be
suitable. The data collected for the temperature and leaf
wetness experiment was amenable to su ch treatment and the
relationship of infection of celery by ~. apiicola as
affected by temperature and leaf wetness duration was
adequately described by a polynomial regression involving
interactions. It was simplified to disease severity indices
using methods of cluster analysis. Four disease severity
132
( classes were obtained which respect the original

relationship and, pending verification of the accuracy of
the model under field conditions, we will be able to predict
disease levels to be expected for any temperature and
wetness combination within the range tested.
Increasingly, hest resistance is included in the strategies

for integrated disease management (Fry, Apple & Bruhn.
1983; Robinson, 1987). Low ta moderate levels of
polygenically inherited resistance are preferred for their
cumulative effects in polycyclic diseases, helping to reduce
their rate of increase. If their effect is sufficiently
large, they can have large economic impacts by replacing
{ fungicides in low value crops. But even low levels of
partial resistance requiring the complementary use of
fungicides can reduce their frequency of application
particularly when applied in conjunction with a forecasting
model (Fry, 1982).
Host resistance can be used in various ways ta complement

other control measures. It can act alone ta replace
chemical treatment; its levels can be used ta either reduce
the doses of active ingredients required for control or to
extend the intervals between spray applications, thus
relaxing selection pressure on the pathogen and extending
the useful life of fungicides, or model predictions can be
1
133
l corrected to account for host re~~stance (Kushalappa et.

al., 1982).
A forecasting model for the control of Septoria blight may

save fungicides early in the season but its effectiveness
would be enhanced by use of even moderate levels of partial
resistance. Partial resistance may be even more useful for
late celery whose growth period coincides with the most
favourable conditions for blight development. There,
fungicides may not be saved but the crop quality and
economic value may be improved considerably.
Cultivars available in our regions were tested to establish

levels of partial resistance to Septoria blight in celery.
They were tested for their ability to reduce disease in the
field and in the greenhouse, and various parameters were
investigated to determine WhlCh part of the pathogen life
cycle is affected hy resistance. A moderate degree of
partial resistance was found in three cultivars, two golden
varieties, Stokes Golden plume and Superdora, and a short
green cultivar Summit. Their ability to reduce the rate of
epidemics in the field which results from a combination of
factors such as lower infection frequency, small lesion size
and reduced pycnidial den8ity (Parlevliet, 1979), is
potentially large enough to justify incorporation into a
•
134
management strategy pending further testing with the various

methods proposed.
Another objective of the resistance trial was to investigate

the relative value of resistance parameters observed in the
greenhouse in predicting cultivar reactions in the field.
Here, it was clearly demonstrated that only the standard
area under the disease progress curve (SAUDPC) had
consistent predictive v~lue but it was limited to the most
resistant Ieaction types. This is because SAUDPC in the
greenhouse describes one disease cycle under a limited set
of conditions that are not necessarily representative of the
field situation where conditions vary continuously.
( Furthermore it evaluates cultivars at a specifie point in
crop development whereas levels of resistance are known to
vary according to crop developmental stage (Fry, 1982,
Robinson, 1987).
Resistance expressed as lowered production of pycnidia is

suspected of being more valuable in predicting disease under
field conditions than is spore density because spores
enclosed in pycnidia are protected against environmental
factors which ensures a long infectious period thus
eliminating the effect of differences in spore production.
This needs further testing however since the relationship
was established for only one of the greenhouse experirnents.
(
135
t
By definition, resistance which allows disease to develop i5
subject to environmental conditions which affect the
development of the pathogen inside the ho st (Fry, 1982).
Results here tend to show that components of partial
resistance are affected by changes in environmental
conditions during the incubation period and that sorne
cultivars react differentially in response to those changes.
SAUDPC could be used with extreme caution to screen
cultivars for better than average resistance but it is more
representative and just as easily measured in field plots.
The reasoning applies to pycnidial production which varies
according to conditions of lighting and temperature. One of
the major arguments in favour of field trials is that they
test resistance of cultivars adapted to the local
conditions.
Analysis of results in aIl experiments using multivariate

methods as a complement to analysis of variance and
regression confirmed reports of other researchers to the
effect that these methods provided improved interpretation
of results in epidemiology and resistance studies (Kranz,
1990; Lebeda & Jendrulek, 1988). They are often subjective
though and results must be considered carefully in the light
of research objectives.
136
( "Epidemiology provides plant pathologists with refined and

deepened knowledge of the behaviour of diseases in the field
for improved disease control or management" (Kranz, 1990).
Sorne new knowledge has been acquired through this work into
the epidemiology of Septoria blight of celery which also
brings forth suggestions for future research. Aside from
the necessary verification and validation of the infection
model, sorne points which need further investigation before
the model can be successfully implemented include: a study
of the effects of periods of high relative humidity on the
infection process, the establishment of an action threshold
and adequate sampling methods. The final model can further
be improved by determination of the effects of the initial
( number of foci of infection and of cultural practices on the
spread of disease in time and in space as is increasingly
being done in relation to diseases with aggregate patterns
of spread (Campbell & Madden, 1990). Finally, further
field studies are required to test resistance levels for
their value as 'fungicide equivalents', or to determine
coefficients of resistance which can serve to correct the
predictions of Lhe infection model developed using the
susceptible cultivar Florida 683.
(
137
LITEaATURZ CITBD

Florida State Hort. Soc. Proc. 83:208-212.
Campbell, C.L., and L.V. Madden. 1990. Introduction to
Plant Disease Epidemiology. John Wiley & Sons. N.Y.,
Toronto. 532 pp.
Corchran, L.C. 1932. A study of two Septoria leaf spots of
celery. Phytopathology 22:791-812.
CPVQ. 1987. Légumes: Protection. AGDEX 250-605. Conseil
des Productions végétales du Québec. Gouvernement du
Québec, Ministère de l'Agriculture, des Pêcheries et de
l'Alimentation. 112 pp.
Fry, W.E. 1982. Principles of Plant Disease Management.
Academic Press, inc. 378 pp.
Fry, W.E., Apple, A.E., and J.A. Bruhn. 1983. Evaluation of
potato late blight forecasts modified to ineorporate host
73:1054-1059.
Hau, B., and J. Kranz. 1990. Mathematics and statistics for
analysis in epidemiology. In: Kranz, J. (ed.). Epidemies of
Plant Diseases. Mathematieal Analysis and Modeling. 2nd
Ed. Springer-Verlag. London, NY, Berlin. 268 pp.
diseases. In: Leonard, J.J. and W.E. Fry (eds.). Plant
Disease Epidemiology, Population Dynamics and Management.
Macmillan Publishing Co. NY. 372 pp.
diseases. In: Leonard, K.J., and W.E. Fry (eds.). Plant
Disease Epidemiology: Population Dynamics and Management.
Vol 1. 372 pp.
Kranz, J. 1990. Epidemies of Plant Diseases. Mathematieal
Analysis and Modeling. 2nd Ed. Springer, Verlag. London,
N.Y., Berlin. 268 pp.
Kushalappa, A.C., Akutsu, M., and A. Ludwig. 1982.
Application of sUlvival ratio for monocyclic proeess of
Hemileia vastatrix in predicting eoffee rust infection rate .
........., Phytopthology 73:96-103.
•
138
Lebeda, A., and T. Jendrulek. 1988. Application of methods

of multivariate analysis in comparative epidemiology and
research into field resistance. Journal of Plant Diseases
and Protection (PflKrankh.) 5:495-505.
Ann. appl. Biol. 65:249-254.
Parlevliet, J.E. 1979. Components of resistance that reduce
17:203-222.
Macmillan Publishing Co. N.Y. 263 pp.
Sheridan, J.E. 1968. Conditions for for germination of
pycnidiospores of Septoria apiicola Speg. N.Z. J. Bot.
6:315-322.
f,
139
1
APPENDIX A. SELZCTIOH 01' POLYROMIAL UGUSSIOR UNIS
BAS&D OH NaIJ.QW' S C;
A. 1 DErINITION
Mallow's Cp is defined as:
c = SSE(p> - [n - 2 Cp + 1)] (A.l)

l' MSE(k)
where p represents the number of parameters in the

restricted model, k the number of parameters in the maximum
model and n, the number of observations. The particularity
of Cp is that it tends ta a value of p + 1 as MSE(p)
approaches MSE(k), in other words, when the correct model is
of size p or if a larger model containing the correct model
is being considered. Cp may assume values less than p + 1
when R2p is close to R\ (Kleinbaum, et al., 1987).
A.2 SELECTION OF THE BEST EQUATION
In the initial step of the analysis, the maximum model

consisting of 24 terms including the intercept was submitted
......
140
t in order of interactions 1 to unweighted least squares

regression using SAS PRoe REG and requesting the 10 best
equations cornprising 8 to 16 parameters. The order of
subrnission was the following for which Cp = 25:
T W TW T2 W2 TW2 T2W T2W2 T3 W3 TW3 T2W3 T3W T3W2

TJW3 T4 W4 T2W4 T3W4 T4W T4W2 T4W3 T4W4
Appendix A-1 lists the 10 best equations selected for each

of the replicates and the pooled data. The only equations
satisfying the requirernent of Cp S P + 1 were those listed
for the second replicate. The first equation of the pooled
data was used as a starting point for diagnostics (see
( Appendix 2) to avoid the possibility of excluding
significant terrns also because it contained the terrns of
rnany suggested equations in Rep 2 and in anticipation of the
necessity to delete predictors on account of near
collinearities.
The order in which terrns are submitted affects the

selection process (see Kleinbaurn et al., Ch. 16). It is
recommended to order terrns according to interactions if
.
o
they are suspected of occurring.

141
1 APPENDIX A-1 'l'ZN 81S'1' IQUATIONS FOR INDIVIDUAL UPlalCATI:S AIID

POOLm DA'rA 1fI'rH MAI.laOII' S c;.
AS SII.2C'1'IOH ClUDJUOH
FULL MODEL
C(p) R-square In Variables in Model
17 .1799 0.1960415 16 T N TW T2 W2 TN2 T2N T3 T3N T3W2 T4 W4 TN4 T3W4 T4W T4W4
17.3045 0.1959140 16 T W TW T2 TW2 T2W T2W2 Tl T3N T3W2 T4 TN4 T2W4 T1W4 T4W2 T4W4
17.8972 0.1956526 16 T M TW T2 W2 TM2 T2M T3 T3M T3M2 T4 M4 TW4 T3N4 T4N T4W2
17.9427 0.1956280 16 T W TW T2 W2 TM2 T2M T3 T3N T3W2 T4 M4 TW4 T2M4 T1N4 T4W
17.9736 0.1945269 15 T W TW T2 T2W T2W2 T3 TlW T4 TW4 T2N4 T3N4 T4W T4N2 T4W4
18.0240 0.7955839 16 T W TW T2 '112 TW2 T2W T3 T3W T3W2 T4 N4 TW4 T3W4 T4N T4W3
18.1471 0.7944328 1~ '1' N TW T2 '112 TN2 T2N T3 T3N T3W2 T4 W4 TW4 T3W4 T4W
18.1662 0.1944224 15 T N TW T2 TW2 T2N '1'3 T3W 'l'3W2 T4 '1' 114 T2W4 T3N4 T4W T4W4
18.4231 0.7953612 16 T W TW T2 '112 TW2 T2N T2W2 T3 'l'3N '1'3'112 'ft W4 TW4 T3N4 T4N4
18.4285 0.7953646 16 '1' W TW T2 '112 TN2 T2W T3 T2W3 T3W T3W2 T4 N4 TW4 T3W4 T4W
----------------------------------- REP-1 ------------------------------------

N - 198 Reqress 10n Models for Dependent Variable: Y
C(p) R-square In Variable. in Model
19.1389 0.8631707 13 T W TW T2 T2N T3 T3W T4 TW4 T2W4 T3N4 T4W T4W4
19.9208 0.8626050 13 T W TW TW2 T2W T3 T2W3 T3W T3W2 T4 TW4 T3W4 T4W3
20.8686 0.8604725 12 T W TW T2 T2N T3 T2W3 T3W T4 TW4 T3N4 T4W3
20.8966 0.8618991 13 T N TW T2W T3 TW3 T2W3 T3W T3W3 T4 TW4 T3W4 T4W3
21.0296 0.8618029 13 T N TW T2 T2N T3 T2W3 TlN T4 N4 TW4 T3W4 T4W3
21. 0"150 0.8617700 13 T W TW T2W T2W2 T3 T2W3 T3W 'f4 W4 TW4 'l'3W4 14W3
21.3561 0.861566'1 13 '1' W TW TW2 T3 T2W3 T3W T3W2 T4 TW4 T3W4 T4N T4~3
21.4721 0.8600359 12 T N TW T2 T3 T2W3 T3W T4 TW4 T3W4 T4W '1'4'113
21. 6565 0.8613494 13 T W TW '1'2 T2W T3 W3 T2W3 T3W 14 TN4 13W4 T4W3
21. 83 62 0.8612193 13 T N TW T2W T3 T2W3 T3N T4 '114 TW4 T3N4 'l'4W2 T4W3
----------------------------------- REP-2 ------------------------------------

N - 195 Reqression Modela for De.>endent Varhble: y
C(p) R-square In Variable. in Madel
... 8.5128
8.8642
8.9676
9.4578
0.7985736
O. '195~642
0.795e495
0.7975249
10 T 'II TW T2 T2W T3 T3N T4 TW4 T3W4
9 T W TW T2 12W T3 T3W T4 TW4
9 T W TW
10 1 'II TW
T2
T2
1'112 T2W T3 T3W T4
TW2 T2W 13 T3W T3W2 T4
9.7125 0.7972423 10 T W TW T2 1'112 T2W T3 T3W T4 T3W4
9.8305 0,7993308 Il T 'II TW T2 W2 T2W T3 T3W T4 TW4 T3W4
9.8925 0.7970425 10 T W TW T2 TZN T3 T3W '1'3'112 T4 TW4
9.9375 0.7992119 Il T W TW T2 T2W T3 T3W T4 TW4 T3W4 T4W
10.0093 0,8013511 12 T W T'II T2 '112 T2W T3 T3W T4 W4 'l'W4 T3W4
10.1105 0,7968006 10 T 'II TW T2 W2 T2W T3 T3W T4 '1' 114
-
142
( APPJ:NDIX B. COLLINZARlTY DIAGNOSTIC AND DJ:LlTIOR or
CO_LA'l'ID HIUIS rROM 'fD MODJ:L.
Problems of cOllinearity and scaling often arise in

polynomial regression and can lead to inaccurate estimates
of the regression coefficients (Kleinbaum et al., 1984). A
good measure of collinearity can be removed by centering the
predictor variables and it proved necessary in this case.
Secondly, use of higher than second degree polynomials and
interactions introduces near collinearities as illustrated
in Appendix B-la by the very high values (over 400) of the
variance inflation factors (VIF) observed for TW2, T3W2, TW4
AND T3W4. The VIF is of the form:
(
j -1,2, ... ,k (B.l)
where R2 j is the squared multiple correlation coefficient

of one predictor with the others. It is based on regressing
Xj on the remaining k. - 1 predictors. As R2 j approaches the
value of 1 it appears obvious from equation B.1 that the VIF
can become qujte large. The tolerance is simply the
reciprocal of VIF or 1 - R2j' It is recommended to pay
attention to any value with a VIF above 10.
The next diagnostic statistics originating from the

technique of principal component analysis are the
143
eigenvalues of the predictor variable correlation matrix,

and the variance proportion of estimated regression
coefficients associated with a given eigenvalue (Legendre &
Legendre, 1984, Ch. 9; Dillon & Goldstein, 1984).
Basically, principal component analysis reduces a set of
predictor variables to a smaller set of mutually independent
variables that are a linear combinat ion of the original
predictors while retaining aIl the original information. On
Appendix B-lb, eigenvalues represent the variance of the
principal components and a property of eigenvalues is that
their sum always equals the number of original predictors.
The variance proportion is the contribution of each original
predictor variable to this variance.
The condition number (CN) is the ratio of the highest and

lowest eigenvalues or ratio of the standard deviations of
the first and last principal components expressed as:
(B.2)
where lambda i5 a symbol for the eigenvalue and k is the

number of original variables. The ratio of the highest
eigenvalue and any other value of lambda whith j < k is a
condition index (Cl j ) although SAS PROC REG does not make
this distinction, grouping CN and CI's under the heading
-- 'CN'. Near zero eigenvalues are instrumental to the
144
( detection of near collinearities in that they represent

extremely small variances adding little information about
the predictors. Eigenvalues of a signal the presence of
exact collinearity. Therefore, the presence of two or more
predictors contributing more than 50% to the variance
(component loading) of a meaningless component indicates a
high degree of linear relationship among them and they are
usually associated with high VIF's. Kleinbaum et al (1984)
suggest that CI's between la and 20 represent mild to
moderate collinearity while values over 30 point out to
severe collinearity.
The following procedure was followed in the process of

( deletion of correlated predictor variables. It should be
noted that tne model was refitted after each deletion.
1) Removal of parameters whose estimates were not
significant at the 0.05 level (ex. Appendix B-la, T4W4
with Prob >ITI = 0.09) or became insignificant
following a deletion.
2) Of two predictors with the highest VIF's, and high
variance proportion (loading) on a component with eN <
10 (ex: Appendix B-1b, CN=lOl, TW2: VIF=563, Var prop.

= .97 and T3W2: VIF=564, Var prop. = .97), removalof
the predictor with the least significant parameter
estimate, or the highest power (T3W2 in this case) and
refitting. Although TW4 and T3W4 also demonstrated high
145
1 component loadings they were dealt with in a subsequent

step, having the next highest VIF.
3) Alternately removing and replacing mernbers of a problem
pair, refitting and deleting the one causing the
large st drop in adjusted R2 • This was the case with T2
and T4 where deletion of T4 first, caused a 5% drop in
Adj R2 -adj. whereas deletion of T2 keeping T4 in only
resulted in a 2% drop.
This process was repeated until an equation was obtained

that had a satisfactory R2 with one VIF > 10 and CN < 10
(Appendix B-2). Interestingly enough, this coincided with
the last equation selected for Rep 2 (Appendix A-1) minus
T2. Appendix B--3 lists the results of the same equation
obtained by weighted least squares regression where
weighting appears to have increased the VIF's slightly but
did not affect the CN which remained below 10.
146
APPIHDIX 8-1. COLLINURITY DIAGNOSTIC - lat t; SELECTION, POOLJ:D

DATA
IOUM ar
AIIalJ'd.
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1
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147
l
.. APfDDIX a-l.b COLLINBAR%TY
DATA
D~STIC - lat c; SELECTION, fOO.~.I:D
Col11nearlty DlaQnoltlcs
Condition Var Prop Var Prop Var Prop Var Prop Var Prop Var Prop
No EiQenvalue NUlIIber INTEReEP T 'II T'II Tl '112
1 5.18529 1.00000 0.0000 0.0006 0.0000 0.0002 0.0000 0.0000
2 4.46198 1.01801 0.0034 0.0000 0.0001 , .0000 0.0004 O. 0006
3 2.64624 1.39982 0.0014 0.0000 0.0155 0.0000 0.0002 0.0000
4 1. 979471. 61850 0.0000 0.0005 0.0000 0.0l?? 0.0000 0.0000
5 0.SS264 2.4101S 0,0014 0.0000 0.0003 0.0000 0.0023 0.0041
6 0.59904 2.94210 0.0000 0.0111 0.0000 0.0039 0.0000 0.0000
1 0.50110 3.21489 0.0678 0.0000 0.0261 0.0000 0.0005 0.0000
8 0.37299 3.72853 0.0265 0.0000 0.2598 0.('000 0.0002 0.0002
9 0.14551 5.96824 0.0001 0.0204 0.0005 0.0048 0.0000 0.0000
la 0.10129 7.15501 0.3505 0.0000 0.0745 0.0000 0.0041 O. 0019
11 0.05600 9.62260 O.COOl 0.0175 0.0001 0.4535 0.000 n 0.0000
12 0.0177917. 07281 0.0000 0.0202 0.0000 0.0087 0.0000 0,0000
13 O. 01709
17.41951 0.0005 0.3892 0.0000 0.1253 0.0001 0.0000
14 0.0091523.19954 0.2141 0.0000 0.1163 0.0000 0.0079 0.91611
15 0.0066521.91450 0.0940 0.0001 0.4080 0.0001 0.2364 0.0068
16 0.00~60 28.02932 0.2400 0.0001 0.0982 O.OOOJ 0.1479 0.0029
11 0.0005059 101. 24314 0.0001 0.5403 0.0001 O.J'IOF 0.0000 0.0004
Colline~rity Diagnostics (Cont'dl
Var Prop Var l'rop Var Prop Var Prop Var Prop Var Prop Var Prop Var P10p Var Prop Var Prop Var Prop
No TW2 T2W T3 T3W T3W2 T4 W4 TW4 T3W4 T4W T.W.
1 0.0001 0.0000 0.0006 0.0002 0.0001 0.0000 0.0000 0.0001 O.COOl 0.0000 0.0000
2 0.0000 0.0001 0.0000 0.0000 0.0000 0.0005 0.0008 0.0000 0.0000 0.0001 0.0086
3 0.0000 O. 0014 0.0000 0.0000 0.0000 0.0002 0.0000 O. 0000 0.0000 0.0018 0.0002
4 0.0000 0.0000 0.0005 0.0126 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
5 0.0000 0.0002 0.0000 0.0000 0.0000 0.0044 0.0063 0.0000 0.0000 0.!)OO4 0.00~1
6 0.0001 0.0000 0.0111 0.0038 0.0001 0.0000 0.0000 0.0004 0.0004 0.0000 0.0000
7 0.0000 0.0001 0.0000 0.0000 0.0000 0.0001 0.0007 0.0000 0.0000 0.0000 0.2151
8 0.0000 0.0020 0.0000 0.0000 0.0000 0.0011 0.0005 0.0000 0.0000 0.0088 0.0006
9 0.0022 0.0000 0.0203 0.0046 0.0022 0.0000 0.0000 0.0027 0.0027 0.0000 0.0000
.".. 10 0.0000 0.0009 0.0000 0.0000 0.0000 O. 0158 0.0238 0.0000 0.0000 O. 0031 0.699,
11 0.0001 0.0000 0.0171 0.4525 0.0001 tl.OOOO 0.0000 0.0002 0.0007 0.0000 0.0004
12 0.0281 0.0000 0.0115 0.0141 0.0268 0.0000 0.0000 0.0260 0.0282 0.0000 0.0000
~ 0.0000 0.3982 0.1215 0.0048 0.0000 0.0000 0.0169 0.0144 0.0000 0.0002
13 0.0035
14 0.0000 0.0016 0.0000 O. 0000 0.0000 O. 0060 0.9563 0.0000 0.0000 o.oon 0.0053
15 0.0000 0.7509 0.0001 0.0000 0.0000 0.2304 0.0013 0.0000 0.0000 o.nas 0.0003
16 O. 0000 0.2~26 0.0001 0.0002 0.0000 O.lH5 0.0039 0.0000 0.0000 0.2451 O. 0047
17 0.9659 • 0.0000 0.54 05 0.3905 0.9659 • o.uOOO 0.0004 0.9531 0.9539 0.0000 0.0000
t
148
( OJ.DlHUa 1&U'l' SQUUZS SOL'U'l'ION lII'l'H CO'RULAftD

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0.0001
0.0000
"O.lf!!11
eooo 0.0000
0.004)0
0.0001
O."U
0.0000
0.0000
0.0000
0.0000
0.000'
0.11'1
0.0001
O."IS
0.0011
0.0005
•• 0.05511
0.05211
' ... U5
'.00005
0.0000
0.0010
0.1010
O.""
0.(",0.
0.0001
0.12"
O.""
0.0002
0.0003
0.1214
0.7113
O.""
O.lltl
0.0000
0.000'
O.ooos
0.0114
(
149
,1 UPDDIX 1-3 1II!IGHT8n LDST SQUAUS SOLU'l'ION VITH CORRlLAT8n

PREDICTORS lœJfIOVm)
ANGULAR TRANSFORMATION - equat10n 2

USING INVERSE SQUARtO etd err ree1d
Model: MODELl
Dependent Variable: Y
Analyei. of Variance
Sum of Mean
Source OF squares Square F Value Prob>F
MOdal 8 1204.50196 150.56275 147.146 0.0001
Error 384 392.91113 1.02322
C Total 392 1597.41909
Root MSE 1.01154 R-square 0.7540
Oep Mean 0.26102 Adl R-sq 0.748'}
C.V. 387.54120
Paramater Eat1mates
Parameter Standard T for HO: Vadance
Variable OF Estimate Error Parameter-O Prob > ITI Tolerance InflatIon
INTERCEP 1 0.426730 0.01104040 38.652 0.0001 0.00000000
T 1 0.078674 0.00358958 21. 917 0.0001 0.08913791 11. 21857093
W 1 0.005517 0.00040300 13.691 0.0001 0.36910085 2.70928664
TW 1 0.000314 0.00010769 3.410 0.0006 0.09388856 10.65092444
T2W 1 -0.000049234 0.00000583 -8.431 0.0001 0.36848823 2.71379086
T3 1 -0.000176 0.00003738 -20.741 0.0001 0.0~868902 10.13213'147
T3W l -0.000004647 O. 00000116 -3.991 0.0001 0.09589115 10.42849058
T4 1 -0.000035591 0.00000166 -21.417 0.0001 0.99631670 l . 00 36':16'11
TW4 1 -2.0S3874E-9 0.00000000 -2.876 0.0043 0.38790690 2.57793817
Collinearity Dia9no.tlcs
Condit!on Var Prop Var Prop Var Prop Var Prop Var Prop Var Prop Var Prop Var Prop Var Prop
Humber El0envalue Number INTERCEP T W TW T2W T3 T3W T4 TW4
l 2.77006 1. 00000 0.0000 0.0081 0.0000 0.0037 0.0000 0.0088 0.0038 0.0000 0.0395
2 1.86139 1. 21194 0.0405 0.0000 0.0618 0.0000 0.0689 0.0000 0.0000 0.0421 0.0000
3 1.83214 1.22960 0.0001 0.0067 0.0001 0.0185 0.0001 0.0073 0.0188 0.0001 0.0006
4 1.62779 1.30450 0.1097 0.0000 0.0353 0.0000 0.0335 0.0000 0.0000 O.lOn 0.0000
5 0.30098 3.03374 0.5238 0.00113 0.0063 0.0022 0.0055 0.0193 0.0048 0.5222 0.3363
6 0.29914 3.04305 0.3136 0.0153 0.0015 0.0044 0.0008 0.0310 0.0070 0.3125 0.~ll0
7 0.20437 3.68163 O. 0115 0.0001 0.8883 0.0000 0.8908 0.0000 0.0000 0.0155 0.0010
8 0.04975 7.46219 0.0008 0.5612 0.0006 0.4218 0.0003 0.5148 0.3831 0.0004 0.0316
9 0.04838 7.5664Y 0.0000 0.4003 0.0000 0.5493 0.0000 0.4188 0.51123 0.0000 0.0039
150
(
APPalDIX C-l. PLO'1' or JACDIO NSIDOALS vs PRmICHD vu.UZS SBOIrIRG
IHCNUKD VAJtIAIIICK trI!B IIICNASIlIG 'l'IDIPDAmN.
(T-15 ·C)
A-l, B-2, etc. Plot ot RSTY*PY. A-l, 8-2, etc. Plot of RSTY*PY.
(NOTE: 21 ob. hldden.) (NOTE: 22 oone.)
S 1 S 1
t 5 + t 5 +
U 1 u 1
d 1 d 1
e 1 8 1 BD D
n 1 AB C n 1 F8E 0 A
t o + ZN 1 t 0 + HIG G 0
1 0 1 1 oEG
z 1
e e 1
d d
-5 + -5 +1
R
e -0.5
-+---------+---------+---------+
0.0 0.5 1.0
R
e
-+---------+---------+---------+
-0.5 0.0 0.5 1.0
Il Il
Predlcted Value of Y Predlcted Value of Y
(T-20 ·C) (T-25 ·C)

A"l, B-2, etc. Plot of RSTY*PY. A-l, 8"2, etc. Plot of RSTY*PY.
(NOTE: 22 oon8.) (NOTE: 211 gone.)
S 1 S (
t 5 + t 5 + AA
u 1 u 1 8
d 1 d 1 AA AA
e 1 C C e 1 AC A8
n 1 A B A n 1 o8 8 B
t 0 + DA E 0 E t 0 + EG ECG
( 1
e
1
1
1
L N l
A A
r
A
E
8
1
e
1
1
1
CA
8
F CA
A A
0
d 1 d 1
-5 + -5 +
R
e
-+---------+---------+---------+ R -+---------+---------+---------+
-0.5 0.0 0.5 1.0 e -0.5 0.0 0.5 1.0
s
Predlcted Value of Y • Predlcted Value of Y
(..
151
APPDDIX C-lb PLOT 01' JACDIft DSIDlJAl,S VS PRJ:J)IC'1'&J) V'ALOIIS SROIJIIJlG

INCIUI:AS&D VUIAItCK WIT. IIfCRIiUIltG 'rDIl'DA'rOU
(T-30 OC)
A-l, a-2, etc. plot of RSTY·PY.
(NOTE: 32 90ne.)
S 1
t 5 +
u 1
d 1
e 1 A
n 1 MC
t O. OZQ
1 1 L
1
e 1
d 1
-5 +
R
e
-+---------+---------+---------+
-0.5 0.0 0.5 1.0
a
Predlcted Value of Y
-
152
( APPENDIX D BOX-UD-1IJIISDRS PLOTS 01' JACICNIJ'E USIDOALS
SBOWIRG IRCRKASIRG SPR&AD or IHTSRQOARTILK
RAHGE .IH IRCRDSIRG TDlPERATURJ:
The interquartile range (IQR) of a box-and-whiskers plot is

a measure of spread of a distribution. In Figures D-la) to
D-lc), it is the area enclosed by the box where the upper
horizontal dashed line represents the value below which 75 %
of the observations lie, and the lower line, the value below
which 25 % of the observations lie.
(
153
1
UP&NDIXI: D-1. BOX-JUlD-nlsaas PLOTS or JACIOflrJ: USIDUALS SBOIfIIIG
llICaDSIIIG .Pa&AD or III'fDQUAR'fIU DIIGa lIl'fH
IHCNUIIIG DIIl'UA'fOM
UNIVARtATE PROCEDURE
Schemat1c Plots
Variable-RSTY Studentlzed Residud wlthout Current Ob.
3 +
1
1
1
+
1
1 +-----+ +-----+
1 1 1 1 1 1
l + +-----+ 1 1 1 1 1
1 1 1 1 1 t-----+ 1 1
1 0 1 + 1 1 1 1 1 +-----+ 1 1
1 1 +-----+ + __ +__ + +--+--,
e ____._ +-----+
6 __ + __ -
*-----*
+ _____ +
*-----* 1 + 1 1 1 1 1
o + +-----+ *--+--* *_ ... _--* *--+--* 1 + 1
1 *--+--* +-----+ *----_. +-----+ *._--*
1 +-----+ t-----+ +-----+
1 1 1 +-----+
-1 + i 1
1
1
1
-2 +
1
1
1
-3 +
W
------+---------+---------+--------+---------+---------t---------+---------+---------+---------+------
-38.4 -26.4 -2.4 2~.6 45.6 -38.4 -26.4 -2.4 21.6 H.6
(12) (24) (48) (12) (96) (12) (24) (48) (12) (96)
T -10 -10 -10 -10 -10 -5 -5 -5 -5 -5
(10) (10) no) (l0) (10) (15) ilS) (15) ilS) 115)
Contlnued ••••
,
j
154
( UPDDID D-lb BOX-~ISD1t8 I»LO'rS or JACDIft USIDUALS SROWING
IIfCUASDIG SPItUD or IIft'DQUAR'l'ILa UIIG& WI'l'H
IIfCUASIIIG . , . .. . .'l'UU.
UNIVARIATE PROCEDURE
Sch_nie Plota
Vuhbl .. ·liSTY Studentized Re.ldual wlthout Current Ob.
1
S+
1 0
1 1
1 1
4 + 1
1 1
1 1
1 1 1
l + 1 1
1 1 1 +---+ 1
1 1 1 1 1 1
1 1 1 1 1 1
2 t 1 1 1 1 1
1 1 +----+ 1 1 1
1 1 1 1 1 1 1
1 t----+ 1 1 1 t----+ *---* 1
1 1 +-----+ 1 1 1 1 t 1 1
1 1 +----+ 1 1 1 + 1 1 1 1 1
1 1 1 1 1 1 1 1 1 1 1 1
1 1 1 1 1 + 1 *----* 1 1 1 +---+
0 ...---t 1 t 1 *---* *----* +----t *--+-* 1 1 1
1 1 1 1 t 1 1 1 1 1 1 1 1
+-----+ 1 t 1 *----* 1 1 1 1 1 1 *---*
1 + 1 +-----+ *-----* 1 1 1 1 1 1 1 +
-1 + *----* *--+--* t---+ +----+ 1 1 +-----+ 1 t----+ +----+
1 +----+ 1 1 +---t 1 1 1 1
1 1 1 1 1 1 1 1
1 1 1 1 1
-2 t 1 1 1
1 1 1
(~
1 1
-3 t
1 +----t
1
,
-----t--------t---------t--------+------t------t--------t--------t--------t-------t---- ,l
W -l8.4 -26.4 -2.4 21.6 45.6 -38.4 -26.. -2.4 21.6 45.6
(12) (24) (41) (72) (96) (12) (24) (48) (72) (96)
T o 0 0 0 0 5 5 5 5 5
(20) (20) (20) (20) (20) (25) (25) (25) (25) (25)
Continued ••••
1
155
1 AlIPmfDIXI: D-10 BOX-UI)-WHISDItS PLO'fS or JACDID MSIDUALS SROWING

IHCUlUING S'BAD or IftDQUAI\'rILS IWIGK Wl'rR
INCJUIIASllfG 'r....SItA'l'OM.
UNIVAaIATE ,aOCEDuaE
Sch• •Uc Plot.
Varlable-RSTY Studentlzed Realdual wlthout current Obs
3 +
1
1
1
2 +
1 o
1
1
+ 1
1 1 o
1 1 1
1 +----+ +-----+
ft __ +__ *
*--+--*
o + *--+--* +-----+ +-----+
1 +-----+ *--+--* +
1 +-----+ +-----+ -----_.
1
-1 +
1
1
1
-2 +
1
1
1
-3 +
--------_ .._-+ --------_._+ -----------+-----------+ -----------+ -----------
W -38.4 -26.4 -2.4 21.6 45.6
(12) (24) (48) ('72) (96)
T 10 10 10 10 10
(30) (30) (30) (30) (30)
156
UP&KDIX & S'1'&M-JJn)-LUJ' DIAG1UU(S or JACDlrZ USIDOALS
A stem-and-leaf diagram is a one-dimensional display of

residuals showing their interrelationships. The stem is
represented by the numbers located to the left of the line
(Apendices F-la and F-2a), and the leaves by those on the
right. Together, the stem and leaf represent the value of
the residual rounded to two significant digits. For
example, at T=25/W=72 (Appendix F-1), there are four
residuals with values between -0.9 and 0.1, the stem being -
o and the leaf corresponding to the decimal. They are: -
0.4, 0.0, -0.9 and 0.3. The distribution of the residuals
takes on the typical bell shape and the statistics in part b
confirm this. At T=10/W=12 (Appendix F-2), there are 12
residuals with the value -2.3, two with value 2.1 etc.,
giving a skewed distribution.
(
157
1
UPENDIX .-1 ANALYSIS or USIDUAI.S WITB SAS UlfIVAlUAft 'ROCSDOU S'l'DI-
A!m-LDI' DIAGII.... or JACIaIln USlDUALS BY 'RIIPUA'l'UM
------------------------------- T-25 W-72 ----------------------------"---
a)
VUiable-RSTY
Studentlzed Re.lduel wlthaut Current Ob.
Stem Lee!
• Baxplat
o~ 1 ;1189
1 1
4 +-----t
15347 5 *-+--*
-0
-2
40'.13
4
4
l
+-----+
1
----+----+----+----+
Hultlply Stem. tee! by 10**-1
b)
Moment a
N 15 SUIn WQtll 15
Meen 1.108099 Sum 16.62149
Std Oev 2.152109 Varhnce 4.631574
Skewnllu 0.165199 Iturtoels -0.86978
USS 113.2603 CSS 64.84204
CV 194.2163 Std Meiln 0.555672
T:Mean-O 1. 99416 Prob> ITI 0.0660
sQn RilOk 30 Prob>ISI 0.0946
Num "'. 0 15
W:Normal 0.977341 Prab<W 0.9187
......
158
( UItINDID 1:-2 ANALYSIS or DSIDUALS WI'fR SAS UNIVARD.H ItJtOCa)URJ:
SRIC-AllD-LaI' DI.IIaAII or J&CJaIIn USIDUALS BY
'HM»1M'fUIt&
- - - - - - - - - - - - - - - - - - - - - T-10 W - 1 2 - - - - - - - - - - - - - - - - - - -
a)
Variable-RSTY
Studentized Rea1dual without Current Ob.
Stem Leer
49 •
2
Boxplot
o
JI
-2
11
333333333333
---t--+---+--+
2
12
1
1
t--+--+
*----*
Multlply Stem.Lear by 10**-1
b)
Moments
N 16 Sum WQts 16
Me.n -0.16138 Sum -2.58211
Std Oev 0.305172 Variance 0.09313
Skewne.s 1.457366 Kurto.is 0.431156
USS 1.813651 CSS 1. 396!146
CV -189.099 Std Mean 0.076293
T:Mean-O -2.11529 Prob>ITI 0.0515
5Qn R.lnk -34 Prob> 1S 1 0.0630
Nwn A_ 0 16
W:Normal 0.589021 Prob<W 0.0001
(
159
UPZIŒIIX 1'-1 nIGHUD L&AST SQuu.s NODEL l'ITTED '1'0 DATA l'ROH
UPLICAft 1
WEIGHT • INVERSE SQUAREO STO.ERR. REstO.

USUIG CENTERED TEMPERATURES AltO "ETNESS
----------------------------------- REP-1 ------------------------------------

Model: MODELl
Analysis of Variance
Sum ot Mean
Source OF Squares Square F Valub Prob>F
Model 8 765.45903 95.611238 101.347 0.0001
Error 189 178.43694 0.94411
C Total 197 943.89597
Roct MSE 0.97165 R-square 0.8110
Dep Mean 0.25665 Adj R-sq 0.8030
C.V. 378.58880
Parameter Est1mates
Parameter Standard T tor HO:
Variable OF tstlmate trror Parameter-O Prob > ,T'
INTtRctP 1 C 416322 0.01490547 27.931 0.0001
T 1 0.093899 0.00482651 19.455 0.0001
W 1 0.006685 0.00054637 12.235 0.0001
TW 1 0.0004119 0.00014574 3.355 0.0010
T2W 1 -0.000060551 0.00000792 -7.645 O. 0001
T3 1 -0.000941 0.00005040 -18.676 0.0001
T3W 1 -O.OO~\:C:!'!;l 0.00000158 -3.782 0.0002
T4 1 -0.000034690 0.00000225 -15.4~0 0.0001
TW4 1 -2.321823E-9 0.00000000 -2.397 0.0175
........
\,
c DPDOIX r-2 DlGaUD LZAS'f SQUAUS IIODZL rI'fHD 'fO DA'fA l'ROII
160
UPLlCAH 2
WEIGHT - INVERSE SQUARED STD.ERR. RESID.

USING CENTERED TEMPERATURES AND WETNESS
----------------------------------- REP-2 ------------------------------------

MOdel: MODELl
Sum of Mean
Source DF Square. Square F Value Prob>F
Model e 471.36136 51.92017 60.267 0.0001
Error 116 111.14261 0.97765
C Total 194 653.20404
Root MSE 0.91876 R-aquare 0.7216
Dep Mean 0.26544 Adj R-.q 0.7096
C.V. 372.49627
Parameter Eat1mate.
Parueter Standard T for HO:
Variable DF Eatim.ate Error Parueter-O Prob > ITI
INTERCEP 1 0.436833 0.01535896 28.442 0.0001
T 1 0.062116 0.00501475 12.506 0.0001
W 1 0.004316 0.00055122 7.732 0.0001
TW 1 0.000260 0.00014946 1.737 0.0841
T2W 1 -0.000037623 0.000001107 -4.U1 0.0001
T3 1 -0.000603 0.00005207 -11.5115 0.0001
T3W 1 -0.000003354 0.00000161 -2.078 0.0391
T4
( TW4
1
1
-0.000036461
-1.738861E-9
0.00000230
0.00000000
-15.873
-1. 759
0.0001
0.0802
(
161
1 APPENDIX 1'-3 DBT&RNI~TIOH or a-SQuaRa roa BA~~SrORNSD

PUDICHD VALmlS
Madel: MODEL1
Analysis ot Variance
SUIn of Mean
Source OF Square. Square F Value Pr > F
Madel l 12.59961 12.59961 946.550 0.0001

Error 391 5.20463 0.01331
C Total 392 17.110424
Root MSE 0.11537 R-square 0.7077
Dep Mean 0.12862 Adj R-sq 0.7069
C.V. 89.70139
Paramater Eatlmatea
Pilrilmeter St andard T for HO:
Variable OF Estimate Error Paramater-O Prob> ITI
INTERCEP 1 0.004142 0.00708802 0.584 0.5593
PY 1 1.013758 0.03295052 JO.766 0.0001
A-l, 8-2, etc. Plot of Y·PY.
1.0 + C A
1 A
1 A
1 A A
Y 1 A B
1 A 8
1 A AA C A E
1 AA A A 8
0.5 + BB A A A
1 A AA 0
of"o> 1 A E 8 A
1 AC 0 C A
.. ~
1
lA
E B
C C
BC
D
B
F A
0
1 AC A CA CA B
ILOR H H C B
0.0 +ZKZ R A A
-+-----_ •• _-+ ---------+ ---------+---------+
0.6 0.8
0.0 0.2 0.4
py
162
( U.DlDIX G 'RO.oaifIOH or llU'aC'flOlf COuaS'ONDING 'fO Dell DCII DlSUS.
S&WRlft INDU AS D.ftlUlIlII:D BY CLUS'l'ZIl AHALYSlS - SAS
CLUSHIl 'ROCmua.
OSI 1 OSI 2 OSI 3 OSI 4

MAX MIN MAX MIN MAX MIN MAX MIN
0.0782'11 0.000006 0.19251 0.014563 0.320609 0.221388 0.621998 0.393723
---------------------- --------- ------- ---------------------- --------------
T W py T W PY T W PY T W PY
----------------------
14 96 0.064535
-----------------------
17 96 0.172226
----------------------
19 96 0.311725
----------------------
20 96 0.393723
14 90 0.046916 17 90 0.145764 19 90 0.280232 21 96 0.474298
IS 90 0.067397 17 84 0.123969 19 84 0.251023 22 96 0.544691
1S 84 0.052954 18 84 0.180842 19 78 0.223761 23 96 0.596481
15 78 0.04221 18 78 0.15792 20 78 0.299413 24 96 0.621998
16 18 0.067542 18 12 0.137233 20 72 0.269553 25 96 0.613298
16 72 0.0~5893 18 66 0.118335 20 66 0.240702 26 96 0.560315
16 66 0.046002 19 66 0.174126 21 66 0.312803 27 96 0.451211
17 66 0.075713 19 60 0.151413 21 60 0.280909 23 96 0.596481
1'1 60 0.063127 19 54 0.130193 21 54 0.JSOO05 23 90 0.572162
17 54 0.051732 19 411 0.110309 22 54 0.31418 24 90 0.602604
17 48 0.041436 20 48 O.lfi1448 22 48 0.281102 24 84 0.577409
18 48 0.070099 20 42 0.137848 22 42 0.249152 24 78 0.547913
lB 42 0.056652 20 16 0.115835 23 42 0.302068 24 12 0.515463
18 36 (l.044616 21 36 0.165114 23 36 0.268963 25 72 0.523589
19 36 0.075033 21 30 0.13976 24 30 0.275225 25 78 0.553141
19 30 0.05985 21 24 0.ll!l9S5 25 30 0.296031 26 78 0.517899
19 24 0.046446 22 24 0.160317 26 30 0.290439 26 84 0.539024
20 24 0.076947 22 18 0.132919 26 36 0.3194111 26 90 0.553942
20 18 0.060243 23 111 0.113245 27 36 0.274155 26 96 0.560315
20 12 0.045465 23 12 0.141121 27 42 0.297649
21 12 0.073153 24 12 0.17a4 27 48 0.320609
29 12 0.04703 25 12 0.192286 28 48 0.221388
29 lB \).060211 26 12 0.19251 28 54 0.236402
29 24 0.070429 27 12 0.167256 28 60 0.251633
29 30 0.078271 28 12 0.1150118 28 66 0.266546
30 la 0.00413 28 18 0.138965 28 72 0.280102
30 36 0.004241 28 24 0.159051 28 18 0.290743
30 42 0.004101 28 30 0.116329 211 84 0.296398
lO 48 0.003871 29 36 0.084563 211 90 0.294547
( 30
30
30
54
60
66
0.003647
0.003403
0.003069
29
29
29
42
48
54
0.090074
0.095375
0.:'00752
28 96 0.282358
30 72 0.002'H9 H 60 0.106165 SUMHARY:

lO 78 0.001651 29 66 0.111198
la 84 0.000588 29 12 0.11503 OSI l S 0.08
lO 90 0.000006 29 18 0.116432 0.08 < DSI 2 S 0.19
30 96 0.001893 29 84 0.113829 0.19 < DS! 3 S 0.l2
29 90 0.105491 0.32 < OS! 4 < l
29 C)f> 0.019945
(
163
1 APPDlDIX H rIKLD LAYOU'l III CKLaRY CULTIVAR 'l'IUAL rOR PARTIAL

USIS'rAHC&
vinci
D C
• A
1 9 1 5 1 6 III 1
1 5 1 4 1 7 1 10 1 1) Florida
2) Superdora
3) Surepak
1 10 1 1 11 1 1 13 1 1 8 4) Surnmit
5) Utah Giant 52-70
6) Calmario
1 7 1 3 1 1 1 9 7) Deacon
B) Bi5hop
9) Venlura
1 3 1 6 1 11 1 114 1 10) Tendercrisp
11) Stk. Imp. Utah
12) Vicar
1 6 1 14 1 1 9 113 1 13) Green Giant
14) Stk.Golden Plume
1 12 1 1 2 1 3 1 7
'" 1 2 1 8 1 10 1 1 12 1
1 14 1 1 10 1 1 5 1 6
1 4 1 1 1 12 1 1 3
1 8 1 13 1 1 2 1 1
1 13 1 1 12 1 1 4 1 6
1 1 1 9 1 Il 1 2
III 1 1 7 1 14 1 1 5
164
UPalDIX 1. SZP'rOl\IA B1.IGB'r 'aoGaKSS OH l ' CZI.Bl\Y CUI.TIVARS - l'IRAI.

DISDS& S.v&JlI'l'J' &III) POUOaTION 01' L&U' AR&A DISDSIID
'rRAHSl'oauam 'l'O SAUDPC- - l tee l':n:w 'r1lIAL
Cultivar b Marked Random Mean

Plants Random & marked
Plants
Final disease severityC

Stokes Gold. Plume 0.1981 Cd 0.1543 cd 0.1806 e
Summit 0.2854 ab 0.1490 cd 0.2309 cd
Superdora 0.2455 bc 0.1193 d 0.1950 ed
Tendererisp 0.2848 ab 0.1646 cd 0.2367 cd
Bishop 0.3072 ab 0.1855 abc 0.2585 abc
Vicar 0.2999 ab 0.1938 abc 0.2575 abc
Ventura 0.2966 ab 0.1685 bcd 0.2454 bc
Deacon o 3035 ab 0.1638 cd 0.2476 be
Utah 52-70 o.J166 a 0.1807 be 0.2622 abc
Stokes Imp. Utah 0.3128 a 0.1947 abe 0.2656 abc
Green Giant 0.2852 ab 0.2048 abc 0.2530 bc
Florida 6b3 0.3084 ab 0.1809 be 0.2574 abc
Calmario 0.3352 a 0.2236 ab 0.2905 ab
Surepak 0.3443 a 0.2380 a 0.3018 a
SAUDf~
Stokes Gold. Plume 0.0521 d 0.0344 d 0.0432 e
Swnmit 0.0552 d 0.0363 cd 0.0457 de
Superdora 0.0615 cd 0.0361 cd 0.0488 cde
( Tendercrisp
BiShop
0.0592
0.0652
bcd
bcd
0.0446
0.0439
bcd
bcd
0.0519
0.0545
cde
cde
viear 0.0667 bcd 0.0456 bed 0.0562 bede
Ventura 0.0705 bcd 0.0421 cd 0.0563 bcde
Deacon 0.0689 bed 0.0437 bcd 0.0563 bede
Utah Giant 52-70 0.0735 bcd 0.0417 cd 0.0576 bcde
Stoke' s Imp. Utah 0.0713 bcd 0.0442 bcd 0.0577 bcde
Green Giant 0.0730 bcd 0.0495 be 0.0612 bed
Florida 683 0.0814 abc 0.0460 bcd 0.0637 bc
Calmario 0.0848 ab 0.0571 ab 0.0110 ab
SUJ:'epak 0.0972 a 0.0658 a 0.0815 a
Standard ar~a under the disease progress curve. Mean over four ?lots
for marked plants, random plants and ~verage for rnarked and random.
b
Cultivar ocder follows ranking for inereasing value of SAUDPC for
the average of marked and random plants.
c
Disease rating is the proportion of leaf area diseased. Values are
the mean of four plots. Each plot comprised three marked plants
revisited and two plants ehosen at random for each visit.
Mean! within eolumns followed by the same letter are not
significalltly different (p .. O. 05) as determined by DNMRT.
165
..
UPIJilDIX J. S&P'l'OJU:A 8LIGH'1' PaQGImSS III U C&LIUa CUL'l'IVARS '1'IWISrOJ'UŒ)
'1'0 'l'HI: S'1'ANDUD AUA Ulmu. '1'8 DIS"". PIlOGUSS (SAOD'C) FO.
S~LING IN'1'&RVALS 0-25 rAYI, 2'-40 »AYS, AND '1-55 DAY.
rOLI.OWING 'l'a IN'l'RODUC'l'lOH or llIOCULtJII.
Cultivar SAUDPC for intp~v~l~·

0-25 daysb 26-40 da' scc b 41-55 days
Stokes Golden Plume 0.0037 0.0454 cd 0.13~O f

Superdora 0.0040 0.0545 bcd 0.1480 tir
Summit 0.0018 0.0399 d 0.1535 def
Tendercrisp 0.0016 0.0498 bed 0.1';;66 cdef
Ventura 0.0024 0.0614 bed 0.1765 bede
BiShop 0.0018 0.0518 bcd 0.1792 bede
Deacon 0.0024 0.0581 bed 0.1794 bede
Viear 0.0017 0.0546 bed 0.1816 bede
Utah 52-70 0.0027 0.0584 bcd 0.1868 bede
Stokes Improved Utah 0.0018 0.0562 bcd 0.1872 bede
Green Giant 0.0021 0.0660 bcd 0.1905 bed
Florida 683 0.0032 0.0717 abc 0.1972 be
Calmario 0.0037 0.0769 ab 0.2185 ab
Surepak 0.0045 0.0955 a 0.2418 a
Values represent plot means for marked and randorn plants.

b
Means in column with no letters were only signlficant at the 0.09
level.
c
Means within eolumns followed by the sarne letter are not
siqnifieantly different (P=0.05) as determined by DNMRT .
.......
166
1 AP»IHDIX K CLOS TER ANALYSIS or !RI SAUD»C rOR IHTlRVALS BE!WZIN

SANPLING DADS IN '1'D FlaLD avALUATION or 14 CZLlRr
CUL'l'IVARS rOR »U'l'IAL USIS'l'AIfC& - StHID 1~88
AveraQe LinkaQe C1u.ter Ana1yeie

EiQenvalue. ot the Correlation Hatrix
E1Qenvalue DHterence Proportion Cumulative
2.22185 1.46889 0.740615 0.74062
0.75296 0.72776 0.250985 0.99160
0.02520 0.008400 1.00000
The data have been .tandardlzed to mean 0 and variance
Roct-Hea -Square Total-sample Standard Deviation -
Root-Hean-Square Diatance 8et~een Obaervationa
1
'.44949 - Norm T
RMS i
NCL Cl usters Jolned rREQ SPRSQ RSQ psr PST2 Dist e
13 Bshp Vicr 2 0.00074 0.999 113.2 0.09779
12 Deak Vnta 2 0.000114 0.998 115.1 0.10457
11 CL13 StkU 3 0.001114 0.997 17.4 2.5 0.14273
10 Utah CL12 3 0.00352 O. "3 63.6 4.2 0.19238
9 CLll Tend 4 0.00111 0.914 31.9 6.9 0.28733
8 CLI0 GrGt 4 0.00911 0.974 32.5 4.5 0.30127
'1 Spda GldP 2 0.01011 0.963 30.8 0.37486
6 Flda Calm 2 0.01267 0.951 30.9 0.40587
5 CU CL9 • 0.04201 0.909 22.4 9.8 0.42223
4 Smit CLS li 0.05762 0.851 19.1 6.0 0.69016
3 CL6 Supk 3 0.01401 0.767 18.1 6.6 0.92753
2 CL7 CL4 11 0.2U83 0.537 13.9 15.2 1.01911
1 CL3 CL2 14 0.53736 0.000 13.9 1.38114
{
'"
167
1 APPDmIX L-l PRINCIPAL COIIPOIIIII'l' ADLYSIS or 'l'D SAODPC l'OR SANPLIIIG

IN'fDVALS IH 'l'D l'IELI) &VALUA'l'ION 01' U C&uaY COL'l'IVUS
l'OR PU'l'XAL USII'l'ARCa - SUMMER 1988
Simple statistlcs
DAn DAY2 DAYJ
Mean 0.0026714286 0.0600142857 0.1813428511

Std 0.0009746231 0.0141126706 O.027743J766
Correlation Matrlx
DAn DAY2 DAYJ
DAYl 1.0000 0.5982 0.2715 l'ur _ PC day 25.

DAn 0.5982 1.0000 0.9058 Cur S~_:lPC doay 40.
DAY3 0.2715 0.9058 1. 0000 Cur SAUDPC ctay 53.
Eiqenval~es of the Correlation Matrix
Elqenvalue Di tterence Proportion cumuliltive
IN1 2.22185 1.46889 0.740615 0.14062

1N2 0.75296 0.72776 0.250985 0.99160
IN3 0.02520 0.008400 1.00000
Elgenvect.ors
PRINl PRIN2 PRIN)

n 0.456496 0.842891 0.284863 Cur SAUDPC day 2~ •
Y2 0.663202 -.108930 -.740471 Cur SAUOPC day 40.
YJ 0.593106 -.526944 0.608733 Cur SAUDPC day 53.
-
168
Al'PIDIDIX L-2 PIUIICIPAL COIIPOIID'l' AllALYIII or 'fB IAtmPC roa. Ilft'ZRVALS

U'1'IIDII UllPLIIIG œnl IH 'l'BI rIKLD IWALUA'fICIf or 1.
CUL'fIVUS roa .U'fIAL aSII'fAIICK - su.ma, 1988
STD SCORES
oas CV DAYl DAY2 DAY3 PRINl PRIN2 PRIN3
1 rlorld. 683 .0032 0.0111 0.1972 0.16193 0.07571 -0.69743
2 Superdor. .0040 0.0545 0.1480 -0.23459 2.10302 -0.33987
3 Surep.k .0045 0.0955 0.2418 2.56043 0.18350 -0.00510
4 SWMlit .0011 0.0399 0.1535 -1. 30729 -0.08016 1.19534
5 Ut.h Ghnt 52-70 .0021 0.0584 0.1868 0.03635 -0.07661 1. 34047
6 C.llUrl0 .0031 0.0769 0.2115 1.381141 0.06162 1. 44851
1 Deacon .0024 0.05111 0.1794 -0.17351 -0.21097 -0.13558
B Bishop .0018 0.0518 0.1792 -0.56353 -0.14855 0.81436
9 Vent.ura .0024 0.0614 0.1765 -0.11106 -0.11685 -1. 62711
10 Tendercrlsp .0016 0.0498 0.1666 -0.81014 -0.65430 -0.63443
11 Stoke Imp Utah .0018 0.0562 0.1812 -0.31007 -0.96280 0.46581
12 Vlc.r .0017 0.0546 0.1816 -0.47226 -0.92566 0.03649
13 Green Gl.nt .0021 0.0660 0.1905 0.14049 -0.82321 -1.16486
14 Golden Plume .0037 0.04 S4 0.1320 -0.84522 2.23519 -0.09599
r
169
APPIDIDIX K. PARTIAL DSIS'fAIIC3 or 17 CUL'fIVARS BASIm OH DSIS'1'ANCK

tAIWŒ'l'ZllS IN GUJ:NHOUS& 'flUAI. 1 (SUIIMSIl, 19U)
CULTIVAR COMPONENTS OF PARTIAL RESISTANCE-

SAUDPC .ILA PCD
Stokes Golden Plume 0.0454 db 0.3430 abcd 178.88 be
Superdora 0.0504 cd 0.3465 abcd 116.63 c
Ventura 0.0540 bcd 0.2457 d 283.07 a
Summit 0.0562 bcd 0.2534 d 302.83 a
Green Giant 0.0609 bcd 0.3907 abc 197.53 b
Vicar 0.0611 bcd 0.2518 d 271.25 a
Surepak 0.0657 abed 0.3296 bcd 186.02 be
Florida K-strain 0.0664 abed 0.324U bcd 318.76 a
Tendercrisp 0.0680 abcd 0.2456' d 281. 42 a
Bishop 0.0680 abcd 0.2442 d 353.47 a
Stoke's Improved Utah 0.0705 abcd 0.3869 abc 294.73 a
Calrnario 0.0708 abcd 0.3054 cd 288.29 a
Deacon 0.0713 abed 0.2711 d 275.75 a
Golden Self Blanching 0.0737 abc 0.3262 bcd 172.01 be
Florida 683 0.0760 abc 0.3400 abcd 314.'77 a
Utah R-strain 0.0775 ab 0.4092 ab 323.29 a
Utah 52-70 0.0920 a 0.4383 a 298.80 a
SPD SPPC T50 c T75
Stokes Golden Plume 1135810 bed 6466 abc 16.6920 19.4742
Superdora 495822 e 4330 bcd 16.3986 19.1864
Ventura 1558882 abc 5743 abcd 16.7088 19.5519
.f'> Summit 709166 de 3816 d 16.3827 19.1259
Green Giant 1278772 abcd 6747 ab 16.4630 19.3946
,,.J Vicar 1101066 cd 4161 cd 16.5051 19.2509
Surepak 1263360 abcd 4551 bcd 16.2874 19.1080
Florida K-strain 1595557 abc 5087 abcd 16.7778 19.6259
Tendercrisp 1789070 a 6992 a 16.6902 19.4993
Bishop 1746823 ab 6161 abcd 16.9084 19.6916
Stokes Improved Utah 1619130 abc 5691 abcd 16.3785 19.1448
Ca1rnario 1776623 ab 6180 abcd 16.3888 19.2175
Deacon 1336673 abcd 4696 abcd 16.4291 19.2864
Golden Self Blanching 729981 de 4095 cd 16.0563 18.9052
E'lorida 683 1295869 abcd 4344 bcd 16.6087 19.4816
Utah R-strain 1778129 ab 5693 abcd 16.7369 19.5731
Utah 52-70 1644810 abc 5798 abcd 16.3607 19.2104
The components are the standard area under the disea~le progress curve
(SAUOPC), rnean le5ion area (MLA), pycnidial density (PCD), spore
density (SPD), spores/pycnidiurn (SPPC, and the time from 0-50 % (T50)
and 0-75 % (T75) disease. Values are the mean of six ~eplicates.
b
Means within the ~ame column foL.vwed by the sarne letter are not
significantly different (P=O. 05) as determined by DNMftT.
c Means with no letter following were not significantly different.
170
c UPDlDIX N. PARTIAL USIS'fAllC& or 17 CZLIRY CULTIVARS BASED ON

USIS'fANCZ PARAM&'l'KJ\S IN GUDlllOUS. 'fIUAL 2 (WJ:N'fSa, 1990)
CULTIVAR COHPONENTS OF PARTIAL RESISTANCE-

SAUDPCti MLA PCD
Superdora 0.04042 e 0.3519 d 638.7 bcd
Summit 0.05052 de 0.3141 d 613.5 cd
Stokes Golden Plume 0.05097 de 0.3585 d 543.9 d
Deacon 0.05377 cde 0.3707 d 845.9 abc
Gld. Self Blnch. 0.05545 cde 0.3558 d 836.8 abc
Florida K-str. 0.05852 bcd 0.4551 cd 841.4 abc
Tendercrisp 0.06145 abcd 0.3933 d 791.6 abc
Bishop 0.06183 abcd 0.3315 d 762.5 abcd
Stokes Imp. Utah 0.06470 abcd 0.6272 ab 897.0 a
Ventura 0.06497 abcd 0.4066 d 758.1 abcd
Vicar 0.06507 abcd 0.3786 d 816.4 abc
Utah R-str. 0.06793 abc 0.6460 ab 901. 4 a
Utah 52-70 0.07170 ab 0.5770 bc 901.2 a
Florida 683 0.07248 ab 0.4481 cd 847.8 abc
Calmario 0.07387 ab 0.5820 bc 763.9 abcd
Surepak 0.07547 a 0.6126 abc 885.3 a
Green Giant 0.07755 a 0.7644 a 855.3 ab
SPD SPPC T50C
T75
Superdora 1245466 bcd 1899 abc 14.80891 18.12465
Summit 1187518 bcd 1970 abc 14.68043 18.54961
Stokes Golden Plume 1218374 bcd 2226 ab 14.43621 17.69051
Deacon 8j9480 d 1054 d 14.90020 17.64443
( Gld. Self Blnch.
Florida K-str.
1904897
1772182
ab
abc
2266 ab
2105 abc
14.94988
15.07798
18.10235
17.96925
Tendercrisp 1409497 bcd 1761 bcd 14.61266 17.85153
Bishop 1479558 bcd 1952 abc 15.10450 17.80015
Stokes Imp. Utah 2325416 a 2604 a 15.67580 18.02926
Ventura 1703526 abc 2234 ab 14.86365 18.90261
Vicar 1075339 cd 1318 cd 14.93661 17.95268
Utah R-str. 1395483 bcd 1303 cd 14.83820 18.17041
Utah 52-70 1685024 abc 1905 abc 14.59313 18.40070
Florida 683 1615711 abcd 1946 abc 15.28340 18.38613
Calmario 1397550 bcd 1840 abc 14.97911 17.97721
Surepak 1653200 abc 1881 abc 14.84545 18.03066
Green Giant 1092362 cd 1342 cd 14.69783 17.85895
Components are the standardized area under the disease progress curve
(SAUDPC), mean lesion area (MLA), pycnidial density (PCD), spore
density (SPD), spores/pycnidium (SPPC) and the time from 0-50 % (T50)
and from 0-75 % (T75) disease. Values are means of six replicates.
b
Means within a column followed by the same let ter are not
significantly different (p-O.OS) as determined by DNMRT.
Means not followed by lettera were not significantly different.
(
171
".- APPZHDIX 0 CLUS'fD AllALYSIS or C3LaU CUL'fIVAR DSPONsas 'tO saLaCftD

PUAlSftRS or USIS'rUCZ lB 'rB rIRS\' GUaBOusa t'RIAL
(GH-l - SUNNSR, 1"9)
GREENHOUSE EXP .1 - 11 CULTIVARS

SAUDPC, MLA, PCD, SPD
Averaoe Llnkaoe Cluster Analysis

tioenvalu •• ot the Correlation Matrlx
Eioenvalue D1tterence Proportion Cumulative
1 2.13169 0.887258 0.534423 0.53442
2 1.25043 0.846576 0.312608 0.84103
3 0.40386 0.195834 0.100964 0.94799
4 0.20802 0.052006 1.00000
The data hive ceen stand~rdized to mean 0 .nd variance
Root-Mean-Square Total-Sample Standard Deviallon • 1
Root-Mean-SfJuare Distance Between Observat ions • 2.828421
Horm T
RMS 1
NCL Clusters Jolned fREQ SPRSQ RSQ PSF PST2 Dlst e
16 Calm fiaI< 2 0.00519 0.995 12.8 0.28810
15 StkU UthR 2 0.00612 0.988 11.9 0.3280\
14 Supk GrGt 2 0.00925 0.979 10.7 0.38477
13 8shp Tend 2 0.00935 0.969 10.6 0.38678
12 Deak Vier 2 0.00993 0.960 10.8 0.39852
11 f"lda CL16 3 0.01620 0.943 10.0 3.1 0.46379
10 CL12 Vnta 3 0.02061 0.923 9.1 2.1 0.53514
9 CLll CLIS 5 0.03014 0.892 8.3 3.3 0.53142
8 CLIO CLB 5 0.034560.857 ~.7 2.6 0.59097
7 CL14 GldP 3 0.02901 0.828 8.0 3.1 0.62062
6 CL? GldS 4 0.04570 0.783 "}. '} 2.4 0.76781
5 CL9 CL8 la 0.127990.655 5.7 7.7 0.78921
4 Spda CL6 ~ 0.05292 0.602 6.5 1.9 0.83496
3 CL5 Smit 11 0.088290.514 7.4 3.0 0.9929~
2 CL3 CL4 16 0.35251 0.161 2.9 10.1 1.13141
1 CL2 Utah 17 0.16101 0.000 2.9 1.33718
172
DPDlDIX P-l PRINCIPAL COIIPORDI'l' ANALYSIS or CZLZRY CULTIVAR aSPORS&S
TC SIUC'1'&J) 'UAllKDU or USIST.AHC& IN TU rIRST
GRJ:&NllOUS& ftDl, (GII-1 - SUItICIR, 1989)
PRINCIPAL COMPONENT ANALYSIS

SAUOPC, MLA, PCD AND SPO
17 Ob.ervat ion.
4 Variable.
SAC PC MLA PCO SPD

Mean 0.0663470588 0.3207411765 262.2058824 1344443.706
Std 0.01111184493 0.0620806538 66.1802651 405165.598
Correlation Matrix
SAOPC MLA PCD SPO
SAOPC 1.0000 O.41U 0.4828 0.~819
HLA 0.41111 1.0000 -.1920 0.0142
PCD 0.4828 -.1920 1.0000 0.6940
SPD 0.4819 0.0142 0.6940 1.0000
El qenvalu.. of the Correlation Matrix

EiQenvalue Ditterence Proportion cumulative
PRINl 2.13769 0.117251 0.534423 0.53442
PRIN2 1. 25043 0.146576 0.312608 0.84703
PRIN3 0.40386 o.195U4 0.100964 0.94799
PRIN4 0.20802 0.052006 1.00000
( EiÇ/anvectors
PRINl PRIN2 PRIN3 PRIN4
SADPC a.5U.!»! 0.349680 -.646856 -.396733
MLA 0.144326 0.1137930 0.318601 0.418970
pcn 0.570562 -.H03BO -.158710 0.704893
SPD 0.593064 -.152317 0.674449 -.412544
'.
173
APPDDIX P-2 PRIHCIPAL CONPOIID'f AH&LYSIS or C&I&RY CUL'rIVAR "SPORS&S

oro sax..c'raD PARlMlftItS or USIS'rANC& IN 'rD rlU'r
GU&NHOUS& 'raDJ, (GB-l - SlHIIR, 1ge9)
SAUDPC, MLA, PCD AND SPD

STO SCORES
OBS CV SADPC HLA PCP SPD
1 F10rlda 683 0.0760 0.3400 314.77 1:195869

2 Superdora 0.0504 0.1465 116.63 495822
3 Surepak 0.0657 0.3296 186.02 1263160
4 summlt 0.0562 0.2534 302.83 709166
5 UUI! Glant 52-70 0.0920 0.4383 298.80 1644810
6 Calmarl0 0.0708 0.3054 288.29 1776623
7 Deaeon 0.0713 0.2711 275.7~ 1336673
8 Bishop 0.0680 0.2442 353.47 1746823
9 VenturA 0.0540 0.2457 283.07 1 ~58882
10 TendereJ:"lap 0.0680 0.2458 281.42 1789070
11 Stoke Imp Utah 0.0705 0.3869 294. ï3 1619130
12 Vleu 0.0611 0.2518 271.25 1101066
13 Green Glant 0.0609 0.3907 197.53 1218712
14 Golden Plume 0.0454 0.3430 178.88 1135810
15 Fla K-straln 0.0664 0.3248 318.76 1595557
16 Utah R-straln 0.0175 0.4092 323.29 17 18129
17 Gld Sl! Blnch 0.0731 0.3262 172.01 729981
OBS CV PRINI PRIN2 PRIN3 PRIN4
1 Florlda 683 O.b1617 0.24131 -1.04825 0.81046

2 Superdora -2.20268 0.91843 -0.01472 0.11,86
3 Surepak -0.53806 0.51799 0.2Q551 -1.41675
4 summlt -0.84436 -1.09116 -1. 43804 2.1593&
5 Utah Giant 52-70 1. 56507 1.84195 -0.73574 -0.07086
6 Calmarl0 0.71166 -0.34361 0.50462 -0.92888
7 Deacon 0.15951 -0.52959 -0.92295 -0.78596
8 Bishop 0.87480 -1. 49439 -0.05890 -0.02810
9 Ventura -0.1"'fi33 -1. 43302 1. 00024 -0.14194
10 Tendercrlsp 0.(:/479 -1.10921 0.336~7 -1. 78131
Il Stoke Imp Utah 0.71147 0.65072 0.75324 0.80218
12 Vlear -0.47619 -0.94469 -0.75103 0.14235
13 Green Giant -0.51883 1.05544 1.13256 0.09490
14 Golden Plume -1.36840 0.19291 1. 85337 0.47776
15 Fla K-straln 0.59311 -0.33228 0.47231 0.81604
]6 Utah R-straln 1.30964 0.91143 0.60521 0.90014
11 CId Slf Bloch -0.89136 0.95377 -1. 89402 -1.22~4~
.,..,.,
174
UPDlDIX 0 CLUSTU. AllALYSIS or C3L8RY CUL'fIVAR HSPOHSES 'fO SIlLEC'l'm

PAlWll:ftRS or USIS'fA11C3 IH 'flll: SECOND GRDHBOUSE 'fIUAL
(GH-2 - WIN'fIa, 1990)
SAUDPC, MLA, PCD, SPD
EiQenvalue. of the Correlation Matrix

E1Qenvalue D1fference Proportion Cumulative
1 2.52101 1.62565 0.630252 0.63025
2 0.119536 0.53959 0.223841 0.85409
3 0.35511 0.12191 0.088943 0.94304
4 0.221116 0.056964 1.00000
The data have been standardized to mean 0 and variance 1
Root-Mean-Square Total-Sample Standard Deviation & 1
Root-Mean-Square Distance 8etween Observations - 2.828427
Norm T
RMS i
NCL C1usters Jolned FREQ SPRSQ R!.Q PSF PST2 Dist e
16 Supk Utah 2 0.00184 0.998 36.2 0.17161
15 Bshp Tend 2 0.00251 0.996 32.7 0.200~5
14 Smit Gl~P 2 0.00430 0.991 26.5 0.262. 1
13 FlaJ( GldS 2 0.00596 0.985 22.5 0.30869
11 CL15 Vnta 3 0.00813 0.977 19.5 3.2 0.32805
11 CLl6 UthR 3 0.010911 0.966 17 .2 6.0 0.37292
10 Spda CL14 3 0.01434 0.C}S2 1S.4 3.3 0.43511
9 Deak Vicr 2 0.01218 0.940 15.6 0.44138
8 Flda CLll 4 0.01949 0.920 14 .8 3.0 0.49205
7 CL12 CLl3 5 0.02887 0.891 13.7 5.2 0.49466
( 6
5
4
CLB
CL6
CL9
Calm
GrGt
CL7
5
6
7
0.02031
0.05080
0.08200
0.871
0.820
0.738
14 .9
13.7
12.2
1.9
3.9
7.1
0.51743
0.75621
0.76200
3 CLS CLC 13 0.22606 0.512 7.4 10.2 0.92588
2 CL3 StkU 14 0.10671 0.406 10.2 2.7 1.09913
1 CL2 CLIO 17 0.40552 0.000 10.2 1. 30072
(
115
APPDlDIX R-l PRINCIPAL COIG'OHa'l' AlfAI,YSIS or CELERY CULTIVAR USPONSJ:S

'r0 SJ:LJ:C'RD PAltAIœ'RRS 01' Us:IS'rANC& IN 'rU S.COHI)
GRUNHOUS. 'rRIAL (GB-2 - WIII'1'D., 1990)
SAUDPC MLA PCD SPD
Principal Compon.nt Analyei.

17 Observations
4 Variables
Simple Steti.tics
SADPC MLA PCD SPD
Mean 0.0627452941 0.4690235294 794.1588235 1414151.941
Std 0.0101905085 0.1365645064 105.5752519 353500.205
Correlation Matrix
SADPC HLA PCD SPD
SADPC 1.0000 0.7589 0.6751 0.2140
MLA 0.7589 1.0000 0.6133 0.2250
PCD 0.6751 0.6133 1.0000 0.4044
SPD 0.2140 0.2250 0.4044 1.0000
EiQenvalues ot the Correlation MatriK

EtQenvalue Difference Proportion Cumulative
PRIN1 2.52101 1. 62565 0.630252 0.63025
PRIN2 0.89536 0.53959 0.223841 0.85409
PRIN3 0.35577 0.12791 0.088943 0.94304
PRTN4 0.22786 0.056964 1.00000
Elqenvectors
PRIN1 PRIN2 PRIN3 PRIN4
SADPC 0.551576 -.295025 0.0'13111 0.1702~2
HLA 0.544187 -.287544 0.544047 -.570257
PCD 0.547889 0.081261 -.787720 -.269649
SPD 0.304611 0.907566 0.273363 0.093858
176
l UPDmIX a-2 PaINCIPAI. CClGtOllKll'f AlG.LYSIS or C3UP.Y CUL'!rIVAR. _SPORSJ:S

'!ro SJ:L&C'nD 'AP.aImDRS or USIS'!r'BCZ IN '!rD SJ:COIm
GuamOUS& ftlAl. (GB-2 - 1IIH'fD., 1110)
PAIJDPC MLA PCD SPD ONLY
oas cv SAUDPC MLA PCD SPD

1 Florida 683 0.07248 0.4481 847.8 1615711
2 Superdora 0.04042 0.3519 638.7 1245466
3 Surepak 0.07547 0.fil26 1185.3 l t ~3200
4 Summit 0.05052 0.3141 613.5 11117518
5 Utah Giant 52-70 0.07170 0.5770 901.2 1685024
'5 Calmario 0.07387 0.5820 763.9 1397550
7 Deacon 0.05377 0.3707 845.9 899480
8 ai.hop 0.061113 0.3315 762.5 1479558
9 Ventura 0.06497 0.4066 758.1 1703526
10 Tendercri.p 0.06145 0.3U3 791.6 1409497
11 Stoke Imp Utah 0.06470 0.6272 897.0 2325416
12 Vlcar 0.06507 0.3786 816.4 1075339
13 Green Giant 0.07755 0.7644 855.3 1092362
14 Stoke Gold.Plume 0.05097 0.3585 543.9 1218374
15 Fla K-strain 0.05852 0.4551 841. 4 17721112
16 Utah R-.train 0.06793 0.6460 901. 4 13954113
17 Gld Slf Blnch 0.05545 0.35511 836.8 19041197
OilS CV PRIN1 PRIN2 PRIN3 PRIN4
1 Florida 683 0.53510 0.11643 -0.47714 1. 51620
2 Superdora -1. 69551 0.19675 0.52169 -1.80592
3 Surepak 1.19390 -0.148111 0.24711 0.37084
4 Summit -1. 55614 -0.205111 0.66504 0.22664
5 Uuh Giant 52-10 1.043117 0.14497 -0.20636 0.01193
6 Calmario 0.52643 -0.112422 1. 20528 0.89254
7 Deacon -0.691112 -1.02374 -2.18736 -1.15757
8 Bishop -0.47723 0.32294 -0.52960 1. 21050
.....'" 9 Ventura -0.07337 0.66385 0.36581 1. 211186
la Tendercrisp -0.27813 0.03062 -0.57754 0.43504
...... 11
12
Stoke lmp Utah
Vicar
1.26246
-o. '9057
1.98156
-0.93390
0.90380
-1. 36337
-1.15096
0.811128
13 Green Giant 1. 24412 -2.09639 0.94128 -0.77915
14 Stoke Gld. Plume -1.63994 -0.29134 1.87916 0.29905
15 Fla K-strain 0.13560 1. 007~2 -0.36269 -0.63425
16 Utah R-strain 0.93065 ·0.67865 -0.18151 -1.34475
11 G1d Slf Blnch -0.16242 1. 61855 -0.84365 -0.15328
- ;r
177
(
LIHRATUU CIHD

Methods and Applications. John Wiley & Sons, Inc. NY.
Kleinbaum, D.G., Kupper, L.L., and K.E. Muller. 1987.
2nd Ed. PWS-KENT Publishing Co. Boston. 718 pp.
Legendre, Land P. Legendre. 1984. tcologie numérique. 2. La
structure des données écologiques. 2nd Ed. 335 pp.
(~

Effects of Temperature and Duration of Leaf Wetness On Infection of Celery by Septoria Apiicola and Cultivar Screening For Partial Resistance

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EI'I'ICTS 01' 'fDIPEO'1'UU BD DURATIOK or LDI' R'l'nSS

OK IUECTION 01' CEUItY BY SEP'1'ORIA UIICOLl.,

CULTIv.aR SCREIKING l'Olt PARTIAL RESISTANCE

A thesis submitted to the Faculty of Graduate Studies and

Department of Plant Science 27 Septernber, 1991

INI'aCTION NODaL rOR.!.. UIICOLA

ABD RKSISTARCa SCR&aHIHG

There are four parts to this thesis. The first part

The thesis format has been approved by the Faculty of

"The candidate has the option, subject to the approval

these are duplicated cle~rly on reguLation thesis

M. Sc. Danielle Mathieu Plant Science

ON TU l:NJ'EC'l'ION 01' C&L&l\Y BY S&PTOl\IA APIICOLA, AND

CULTIVAR SCREZHING rOR PARTIAL RESISTANCE

Growth chamber studies were condllcted with the celery

M. Sc. Danielle Mathieu Plant Science

DUUI DS LA MOUILLURE SOR L' INI'ECTION DU CtLl:RI PAR LI:

CHAMPIGNON SI:P'1'ORIA APIICOLA, 1:'1'

ÉVALUATION DE CUL'1'I~ POUR LA RtSIS'1'ANCE PAR'1'II:LLE

Des expériences à atmosphère contrôlée avec le cultivar de

céleri Florida-683 ont été effectuées afin de déterminer les

effets de la température (10, 15, 20, 25 et 30 OC) au cours

de diverses durées de mouillure (12, 24, 48, 72 et 96 h) sur

l'ép~démiologie de la brûlure septorienne. Les résultats

indiquent que le nombre de lésions augmente avec la durée de

la mouillure à toutes les températures sauf à 30 oC où la

relation inverse est observée. Le nombre maximum de lésions

est apparu à 25 oC, 72 et 96 hr de mouillure. La variable

indépendante (transformation arcsin de la proportion du

nombre maximum de lésions) est adéquatement décrite par un

modèle de régression polynomiale à neuf variables (R 2-

Ajusté=74.8). Une analyse par groupements hyérarchiques des

valeurs prédites a réduit celles-ci à quatre niveaux ou

indices de sévérité de la maladie (ISM) correspondant à des

probabilités d'infection variant de très faibles (1) à très

fortes (4). Des cultivars de céleri furent évalués afin de

déterminer leurs niveaux de résistance au champ et sous

serre. Au champ, l'aire sous la courbe de progression de la

l am very grateful to my advisor, Dr. Ajjamada C.

Others whose contribution was greatly appreciated are Dr. M.

Special thanks to the workers of the Agriculture Canada

Finally may deepest appreciation goes to my colleagues from

This project has been made possible through a scholarship

À toi, Lionel, pour m'avoir enseigné que la connaissance

A mes enfants, Robert et Tobi, à mes soeurs et surtout à

· ·· · ··· ···· vii

APPBNDIX . . · ·· · · · · ·· · ··· · · · 139

Tabl. 2.1. Parameter cstimates of the regression

Abbreviation Ter.m repr•••nt.d

AUDPC Area under the disease progress curve

1.1 GENZPAL INTRODUCTION

Public demand for innocuous food is rising with the

c Not surprisingly, a growing number of farmers have converted

In the area of pest control, there has been progress made

limits set by environmental conditions to maintain pest

Nowadays, the number of IPM programs through which farmers

Late blight or Septoria blight of celery caused by Septoria

Forecasting involves quantitative studies in the