Springer New York Botanical Garden Press

Haploid Angiosperms
Author(s): Gordon Kimber and Ralph Riley

Source: Botanical Review, Vol. 29, No. 4 (Oct. - Dec., 1963), pp. 480-531
Published by: Springer on behalf of New York Botanical Garden Press
Stable URL: http://www.jstor.org/stable/4353678 .
Accessed: 22/09/2014 21:55
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp
.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.
New York Botanical Garden Press and Springer are collaborating with JSTOR to digitize, preserve and extend
access to Botanical Review.
http://www.jstor.org
This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

All use subject to JSTOR Terms and Conditions
HAPLOID ANGIOSPERMS
GORDON KIMBER AND RALPH RILEY

Plant Breeding Institute, Cambridge, England
Introduction ....... .......................... 480

Terminology . .
................................ 493
Meiotic Behaviour ..... ............................ 496
Monoploids ................... . . .......... 497
Autopolyhaploids ..... ............................ 499
Allopolyhaploids ..... ............................ 501
Aneuhaploids .... ............................. 505
Secondary Pairing of Univalents ...... .......................... 507
Morphology and Anatomy ............................... . . 508
Origin................................ 509
Spontaneous Origin .... ............................ 511
Origin after Experimental Treatment ........... .. .................. 511
Origin as a Result of Polyembryony ................................ 513
Uses................................ 516
Conclusions . .
................................ 517
Addenda ............. ..... 518
LiteratureCited.... 519
Supplementary Citations .. . ................. 529
Citations Added in Proof . . ..................... 531
480

HAPLOID ANGIOSPERMS 481
INTRODUCTION
Haploid angiosperms are sporophytes with gametic chromosome
numbers. It is not only such numbers, however, which distinguish
haploids, for they also have the precise chromosomecomplements of
the gametes of normalindividualsof the species.As might be expected
with this constitution,they mostly arise through parthenogeneticfunc-
tioning of unfertilisedgametes.
The first haploid to be reportedwas found in Datura stramoniumin
1921 by A. D. Bergner (Blakeslee, Belling, Farnham and Bergner,
1922). The second to be discovered was also solanaceous,being in
Nicotiana tabacum (Clausen and Mann, 1924). Subsequently,haploids
were recorded in Triticum compactum (Gaines and Aase, 1924),
Oryzasativa (Morinaga and Fukushima,1931) and Hordeum vulgare
(Johanson, 1934). In 1928 haploids were reportedin the Compositae
and again in the Solanaceae,in Crepis capillaris (Hollingshead, 1928)
and Solanum nigrum (Jorgensen, 1928); also in the Cruciferae,in
Matthiola incana (Lesley and Frost, 1928). Soon afterwardshaploids
were detectedin Oenotherarubricalyx(Gates, 1929), Zea mays (Ran-
dolph, 1932a) and Gossypiumbarbadense(Harland, 1936).
Thus it was apparent from the early years of investigation that
haploids are not restrictedto any particulartaxonomic group but can
be collected from representativesof severalfamilies. It is not, therefore,
surprisingthat, in the literatureup to the present, haploids have been
recorded in at least 71 species, representativeof 39 genera in 16
families (Table I).

TABLE I
ANGIOSPERM SPECIES, IN WHICH HAPLOIDS HAVE BEEN FO
Somatic
Species Chromosome Origin Type Cy
No.
PAPAVERACEAE
Eschscholtzia californica 6 S M
CRUCIFERAE
Sisymbrium
y orio 14 S P 13.2 I, 0.411.
Matthiola incana 7+f H M association of
Brassica campestris 10 S M 101, occasiona
Brassica napella 19 S P up to 7II per
Brassica oleracea 9 S M 91 in 38 P.M.C
2 II in 4.
Brassica napus 19 S P
Brassica napus 19 S P 5-8II, some he
Brassica carinata 17 H P 17I, occasional
Generic and specific names are given in the first column; somatic chromosome numbers
letters in the third column give some indication of the origin of the haploids: S indica
ously; E, after experimental treatment; H, after hybridisation; T, as a member of a se
fourth column identify monoploids and polyhaploids, respectively. A brief description o
in the fifth column, when such information is available, and the following abbreviations
III, trivalent; IV, quadrivalent; P.M.C., pollen mother cell; MI, first meiotic metapha
refer to the references in the bibliography.

Som4tic
No.
PORTULACACEAE
Portulaca grandiflora 9 H M irregular meio
CHENOPODIACEAE
Beta vulgaris 9 E M bivalents and
in 40% of cel
Beta vulgaris 9 T M
LINACEAE
Linum usitatis-simuin 30 T P
ONAGRACEAE
Godetia 'whitneyi 14 S M II's in less tha
Godetia amonea 14 S M
Oenothera blandina 7 S M 71 in 80% of
II's and III's
Oenothera franciscana 7 H M no II's, associa
Oenothera franciscana 7 H M
Oenothera franciscana 7 S M 71 irregularly
Oenothera franciscana 7 H M 1 or 2 II in s
Oenothera Hookeri 7 H M
Oenothera Hookeri 7 H M 1 or 2 II in so
Oenothera lamarkiana - -
Oenothera rubricalyx 7 H M 7I.
Epilobium collinumn 18 S M no true chrom
Epilobium hirsutum 18 H M many II, III a
Epilobium luteum 18 H M no true chrom
Epilobium parviflorum 18 H M no true chrom

Somatic
Species Chromosome Origin Type C
No.
CUCURBITACEAE
Citrullus vulgaris 11 E P Ill in 7 cells
and 2II in 2
Cucumis sativus 14 E M
Cucurbita maxima 20 T P occasionally
TILIACEAE
Corchorus olitorus 14 E M
MALVACEAE
Gossypium barbadense 26 T P
Gossypium barbadense 26 T P 25.6I, 0.2II.
Gossypium barbadense 26 T P 26I, analysis
Gossypium hirsutum 26 T P up to 5 assoc
Gossypium hirsutum 26 T P 0.16II per P.M
Gossypium hirsutum 26 T P
Gossypium hirsutum 26 T P properties of

Gossypium davidsonii 13 T M
Gossypium sturtii 13 T M
EUPHORBIACEAE
Ricinus communis 10 S M 10I's, occasio
PAPILIONACEAE
Ficia faba 6 S M 6I's.
Medicago sativa 16 S P 8II in 43% o

Somatic
No.
Medicago sativa 16 S P good pairing,

Medicago sativa 16 S P many abnorma
Meliotus spp. 8 S M describes haplo
Sesbania aculeata 12 T P
SALICACEAE
Populus tremula 19 S M
ROSACEAE
Fragaria vesca 7 H M 7I's.
Prunus persica 8 T M
RUBIACEAE
Coffea 22 - P to 6 II's.
Coffea arabica 22 - P
VALERIANACEAE
Valeriana sambucifolia 28 T P 14II.
COMPOSITAE
Crepis capillaris 3 S M
Crepis capillaris 3 H M always 3I.
Crepis tectorum 4 E M (Androgensis).
Parthenium argentatum 36 HS P up to 17II's.
Parthenium argentatum 36 HS P 181I.
PRIMULACEAE
Primula vulgaris 11 H M
Primula veris 11 H M

Somatic
Species Chromosome Origin Type Cyt
No.
SOLANACEAE
Nicotiana glutinosa 12 E M
Nicotiana glutinosa 12 SE M 12I.
Nicotiana glutinosa 12 H M 121.
Nicotiana langsdorfii 9 H M 1II in 3% P.M
than 1% (And
Nicotiana repanda 24 H P 241 (Androgens
Nicotiana rustica 24 H P occasional biva
Nicotiana rustica 24 H P
Nicotiana rustica 24 HE P
Nicotiana rustica 24 H P occasional biva
Nicotiana sylvestris 12 H M lII in 6.8% P
(Androgensis).
Nicotiana tabacum 24 - (Review paper
up to 1942).
Nicotiana tabacum 24 H P 24I, occasional
Nicotiana tabacum 24 T P
Nicotiana tabacum 24 H P (Androgensis).
Nicotiana tabacum 24 S P terminal associ
Nicotiana tabacum 24 H P
Nicotiana tabacum var. angustifolia 24 H P
Nicotiana tabacum 24 H P 1 to 3II at Mi.
Nicotiana tabacum 24 E P 241 scattered o
Nicotiana tabacum 24 H P Duplication de
with higher pa
normal haploid
homologous pa

Somatic
No.
Nicotiana tabacum 24 H P up to 14 chrom

associated.
Nicotiana tabacum 24 H P 0-121I, highly
Nicotiana tabacum 24 HT P 0-2II's at MI.
Nicotiana tabacum 22 H P
Capsicum annuum 24 T M up to 12 chrom
associated.
Capsicum annuum 24 H M (Androgensis).
Capsicum annuum 24 E M 24I.
Capsicum annuum 24 T M
Capsicum frutescens 24 T M
Datura metel 12 H M
Datura pruinosa 12 H M
Datura stramonium 12 H M 12I, assorted a
Datura stramonium 12 H M
Datura stramonium 13 H M Primary and se
ln+1 types.
Lycopersicum esculentum 12 S M
Lycopersicurm esculentum 12 S M 121.
Solanum demissum 36 H P 4.7II.
39 H P 5.61I per P.M.C
Solanum demissum 36 H P 4.7II per P.M.C
Solanum demissumn 36 H P 6.5911 and 9.78
Solanum demissum 36 H P 5.43II per P.M
Solanum nigrum 36 H P ranges from 36
Solanurn tuberosum 24 T P
Solanum tuberosum 24 H P 11-12II.
Solanum tuberosum 24 H p 23 of cells with

Somatic
No.
Solanum tuberosum 24 H P
Solanum tuberosum 24 S P meiotic analysi

Solanum polytrichon 24 H P 7.9II.
Progeny of (4x Solanum chaucha x
S. tuberosum) 24 S 12II frequently
CONVOLVULACEAE
Ipomea 15 H M 15I's at M1.
Ipomea 15 H M
SCROPHULARIACEAE
Jntirrhinum majus 8 S M II's recorded.
Antirrhinum majus 8 H M 1 to 4 II per P
Antirrhinum majus 8 E M
LILIACEAE
Asparagus officinalis 10 T M
PALMAE
Cocos nucif era 16 T M
ORCHIDACEAE
Bletilla striata 16 T M occasional II.
GRAMINEAE
Zea mays 10 H M
Zea mays 10 E M

Somatic
No.
Zea mays 10 S M non-homologous

Zea mays 10 S M double nature
at prophase.
Zea mays 10 S M mutations in pr
Sorghum vulgare 10 S M II's in 10% of
Sorghum vulgare 10 T M 36 cells 10I, 8
4 with 2II and
Sorghum vulgare 10 H M 22 cells 10I an
F6 (Sorghum halepense X S. 'vulgare) 20 H P lOII with occa
Phleum pratense 21 T P
Phleum pratense 21 T P 7II+7I.
Dactylis glomerata 14 T P 71I regular.
Bromus inermis 28 S P 14II regularly.
Poa annua 14 S P temperature ri
pairing.
Poa alpina 14 T P
Poa pratensis T P
Poa pratensis 16
18 T P
Awvena strigosa 7 H M 102 cells with
7 cells with 1I
Aegilops longissima 7 H M 6.74I, 0.13II.
Jegilops ovata 14 H P 384 cells with
49 with 2II, 2
1III and 1 wit
Afegilops caudata 7 S M mainly 7I.
Secale cereale 7 TE M
Secale cereale 7 E M 3% of P.M.C.

Somatic
Species Chromosome Origin Type Cyto
No.
Secale cereale 7 T M II's observed,

statistical analy
Triticale 21
28 S P
Hordeum distichum 7 T M 1II in 5% of P
Hordeum distichum 7 H M
Hordeum vulgare 7 S M
Hordeum vulgare 7 T M
Triticum monococcum 7 S M 490 cells with 7
Triticum monococcum 7 E M 7I distributed
Triticum monococcum 7 E M
Triticum monococcum 7 SE M III in 3% of P

Triticium dicoccum 14 E P
Triticum durum 14 H P
Triticum durum 14 T P
Triticum durum 14 TH P
Triticum persicum 14 E P
Triticum persicum 14 H P Almost totally
Triticum timopheevi 14 H P 13.45I, 0.25II,
Triticum timopheevi 14 S P OII in 70 cells,
and 2II in 2 ce
Triticum turgidum 14 H P
Triticum spelta 21 S P
Triticum compactum 21 H P 2II, sometimes
Triticum aestivum 21 T P
Triticum aestivum 21 S P

Somatic
Species Chromosome Origin Type Cyto
No.
Triticum aestivum 21 E P II's and associa
Triticum aestivum 21 H P 25 cells with O
1II, 3 cells wit
Triticum aesticvum 21 S P up to 61I, usua
Triticum aesti'vum 21 H P 211 in 47.7% o
111 in 36.6%, 2
0.4% 4II in 0.4
Triticum aestivum 21 H P
Triticum aestivum 21 S P 57.6% of P.M.C
Triticum aestivum 21 E P up to 5II, statis
of chromosomal
Triticum aestivum 21 P review of whea
Triticum aestivum 21 - P analysis of hap
Triticum aestivum 21 S P 18.05I, 1.3811,
Triticum aestivum 21 S P genetic control
20 behaviour.
Triticum aestivum 20 E P 18.53I, 0.671I, 0

Triticum aestivum 22 E P 19.28I, 1.25II,
Triticum aestivum 21 E P use of cytoplasm
Agropyron desertorum 14 T P approx. 711.
Agropyron intermedium 21 T P 14.5I, 3.1II, 0.1
Oryza sativa 12 S M 12I.
Oryza sativa 12 T M
Oryza sativa 12 S M occasionally 1 o
Oryza sativa 12 S M
Oryza sativa 12 S M description of h
Oryza glaberrina 12 S M c.f. 0. sativa.

492 THE BOTANICAL REVIEW
Haploidy was first reviewed by Gates and Goodwin (1930) when

they described a new haploid in Oenothera. Subsequently,Ivanov
( 1938) reviewed the literature and described attempts at artificial
production of haploids. The most extensive review is that by Kostoff
(1942) who listed all existing reports of work on them. At the same
time he discussedtheir origin, morphology,anatomy,cell size, meiotic
behaviourand fertility.
A more recent review was producedby Kehr (1951) who gave a
table recording 56 species in which haploids have been found. In
addition,the origin of haploid-diploidtwin embryosin the angiosperms
has been reviewed by Maheshwari (1945). More recently, Avery,
Satina and Rietsema (1959) have given detailed considerationto the
many haploids of Datura discoveredby Blakeslee and his co-workers,
whilst Sears (1948) has reviewed the literatureon the haploids of the
Triticinae.
The study of haploids needs little justification,since they constitute
an importantgroup of individuals showing genetically anomalousbe-
haviour in many species. In addition, the study of haploids has borne
fruit in some theoreticaland practicalfields. First, since in the meiotic
divisions of a haploid each chromosomeusuallylacks its normalhomol-
ogous partner, new and hitherto undetected pairing affinities may be
revealed at meiosis. These may critically influence the attitude taken
on the cytogenetic structure and evolution of a species. Moreover,
regarding crop plants, opinions on the efficient utilization of genetic
variation may be modified as a result of the new information made
availableon the genetic structureof species by haploid studies.
A practical outcome of this work has been to make it possible to
obtain completely homozygous lines-by doubling the chromosome
numbersof haploids, using colchicine. In this way plants homozygous
for self-incompatibilityalleles have been obtained without resorting
to difficultprogrammesof inbreeding.The same means can also readily
provide genetically stabilized lines for breeding or experimentalpur-
poses. In addition, the differencesof the external phenotypes of hap-
loids, from those of normal individuals,have affordeda measureof the
influenceof genome dosage on morphogenesis.
The purpose of the present paper is to review the conclusionswhich
can be drawn from the latest work on the origin, cytogenetics and
agriculturalutility of haploids.

TERMINOLOGY
Before entering into a discussion of chromosomebehaviourin hap-
loids, it seems desirableto establish the sense in which certain terms
will be used. This requires, initially, a brief considerationof some
widely used terminology.
Many of the species in which haploids occur are members of poly-
ploid series, that is, of series of species between some of which there
are differencesin chromosomenumbers such that those with higher
numbers have multiples of the haploid numbersof the species lowest
in the series. The haploid number of species at the base of the series,
that is of the diploids, is known as the "basicnumber".The concept
of the polyploid series has, of course, been widely discussed (see Dar-
lington, 1937), and a satisfactoryclassificationof the types of poly-
ploids has been proposed by Stebbins (1947).
However, in order to provide a backgroundto the classificationof
haploids it is necessarybriefly to consider some of the kinds of poly-
ploid species. The foundation of the series is provided by diploid
species, with somatic chromosomenumbers which are twice a basic
number. If a plant with the gametic chromosomeconstitution is pro-
duced by a diploid, it may be known either as a "monoploid"(Mc-
Clintock, 1933) or simply as a "haploid".The haploid set of chromo-
somes of a diploid species is known as a "genome".Diploids which
have divergedgeneticallyand in chromosomestructure,to such a degree
that synapsis is considerablyimpaired at meiosis in hybrids between
them, may be said to have differentgenomes.
The polyploid species of a series,with chromosomenumbersexceed-
ing the diploid numbers, arise by multiplication of the chromosome
number, either in somatic or generative cells, of species lower in the
series. If the genome of only one diploid species is present in the
polyploid, so that, for example, in a tetraploid it would have the
genome structureAAAA, then it may be termed an "autopolyploid".
If, however, structurallydistinct genomes are present, whether the
genomes contain the same or different numbersof chromosomes,it is
termed an "allopolyploid".An allotetraploidof this type would have
the genome structure AABB. Stebbins has also applied the useful
description"segmentalallopolyploid"to those species derived by poly-
ploidy from parental species which have not diverged to the same
extent in chromosome structureas the parents of full allopolyploids.

More complex terminologyis requiredto categorizespecies with some

genomes representedtwice only and others more than twice.
Auto-, segmental allo- and allopolyploids are stages along a con-
tinuous series of types of polyploid species.The patternof chromosome
behaviour they display at meiosis affords the principal distinction be-
tween them. Theoretically, in autopolyploids each chromosome is
equally capable of synapsis with however many structurallyidentical
partnersmay be present. Consequently,multivalentsare frequent and
there is polysomic inheritance.At the other extreme, in allopolyploids,
every chromosomehas only one partner with which to pair, so that
there are only bivalentsand inheritanceis disomic. Segmentalallopoly-
ploids fall between these two extremes, forming fewer multivalents
than autopolyploids,but with some divergencesfrom the regularpair-
ing of allopolyploidsand with some polysomic inheritance.
The term "haploid"should be strictly interpreted as meaning an
individual "havinga single set of chromosomes",and hence ought per-
haps to be confinedto the haploids of diploid species. It should not be
applied to individualshaving the gametic chromosomeconstitution of
polyploid species, since they have more than a single set of chromo-
somes. For this purpose the word "polyhaploid"(Katayama,1934) is
NMonoploid
Euhaploid Allopolyhaploid
Polyhaploid
Haploid Autopolyhaploid
D)isomic haploid
Addition haploid
Aneuhaploid Nullisomic haploid
Substitution haploid
Misdivision haploid
Figure 1. Types of Haploids

useful in that it clearly indicates the multiplicity ofL-singlesets. How;

ever, for convenience in the present work the general term "haploid"
will be used when reference is not being made to a specific category
of plant. When a special category is referred to, the specific term
derived in the mannerdiscussedbelow and shown in Figure 1 will be
used.
The cytogenetic constitutionsof polyhaploidscan be very different,
dependingupon the type of polyploid from which they are derived. It
is necessary,therefore, to distinguish the derivativesof autopolyploid
species as "autopolyhaploids", since they have more than one dose of
most of the genetic material.These can be contrastedwith the deriva-
tives of allopolyploid species, "allopolyhaploids", in which supposedly
there is little if any duplication.
In discussingthe classesof haploids,referencemust be made to those
with chromosomenumbers which are not exact multiples of one of
the basic numbers of the group. These aneuploid haploids may be of
two kinds-with extra chromosomes,or deficient for chromosomes-
relative to the euhaploidcondition. The extra chromosomesin the first
group may be part of the gametic set of the species. Thus the plant
is disomic for some membersbut monosomic for the rest of the com-
plement. Such disomic haploids have been most frequently observed
in Datura (Avery et al., 1959) but were first reported in Matthiola
incana (Lesley and Frost, 1928).
The extra chromosomes, alternatively, may be different from all
those of the normal gametic complement of the species, to give an
"additionhaploid".An example of this was the plant with the poly-
haploid complementof Triticum aestivumplus a single chromosomeof
Secale cerealedescribedby Riley and Chapman (1957).
Occasionally,haploids have occurred deficient for a chromosome
from the normal gametic complementof the species. These have been
called "subhaploids"in Poa by Kiellander (1942) and "nulli-haploids"
in Triticum aestivum by Riley and Chapman (1958). Although the
latter term is useful when the symbol for the deficientchromosomecan
be inserted,as in "nulli-V haploid"for example, the full term, "nulli-
somic haploid",will be used in the present context.
Another sort of haploid has been mentioned by Riley (1960). This
so called "substitutionhaploid"had one of the chromosomesof Triti-
cum aestivumreplacedby a chromosomeof Secale cereale.
Finally, haploids can be formed with telocentricor iso-chromosome

misdivisionproductsof some chromosomes.These may be termed "mis-

division haploids".
MEIOTIC BEHAVIOUR
In haploid forms with little chiasma-formation,it is often difficult
to distinguish precisely first metaphaseof meiosis from first anaphase.
This is due to the absenceof a metaphaseplate and to the distribution
of univalentsto the poles. Consequently,some authors (Person, 1955)
have preferredto use the term "meta-anaphase" to denote the whole of
this period of division. In general,however,it seems that this difficulty
can be resolvedif those cells are said to be in anaphasein which partner
chromatidsfall apart and are held together only at the centromere.
Metaphaseis then the precedingstage, duringwhich partnerchromatids
lie close together,and it lasts from the time at which the nuclearmem-
brane disappearsand the apparent body repulsion between chromo-
somes, characteristicof diakinensis, ceases. Use of the criterion of
partner chromatidseparationto distinguish metaphasefrom anaphase
may not always strictly coincide with distinctionsbased on the move-
ments of chromosomespaired in bivalents.Nevertheless,in the haploid
organismsin which it is possible, recognitionof the phases of division
in this way will permit more accuratedescriptionsof meiosis.
It is widely assumed that the formation of a chiasma between two
chromosomesimplies that they possess a common segment, similar in
structureand gene content. Thus the observationof chiasmatais ac-
cepted as evidence of at least segmental homology. Levan (1942), in
consideringthe meiosis of monoploid Secale cereale,has examined this
premise in considerabledetail, only to conclude that the evidence was
inadequateto refute it. In the light of this and other evidence, in the
following discussionin which first metaphasebehaviourwill be mainly
considered,the hypothesisof the direct relationshipbetween homology
and chiasma-formationwill be accepted.
Since the chromosomestructureof a species embodiesits past history,
and the precise structureso far as internal homologies are concerned
is best exposed in haploids,materialof this type has been particularly
useful in evolutionarystudies.Indeed, haploid meiosis has on occasions
been used to determine the chromosomeconstitutions of species. Be-
cause of its reflection of evolutionaryhistory, in the following discus-
sion the meiotic behaviourof haploidswill be consideredin relation to
the supposed diploid or polyploid status of the species in which they
occurred.

HAPLOIDANGIOSPERMS 497
MONOPLOIDS
The occurrenceof bivalents or multivalents at first metaphase of
meiosis is uniformlylow in monoploids (Table I), but regularforma-
tion of one or more bivalents has been recorded in some species.
Indeed, of the 34 species on which data on monoploid meiosis are
available,17 formed occasionalbivalents.
For example,a monoploid of Hordeumdistichumwas found to have
a bivalentin 5.6 per cent of its pollen mothercells (Tomentorp, 1939).
Levan (1942) recorded bivalent formation in four monoploids of
Secale cereale,ranging from very low levels in one plant to a mean of
almost one bivalent per cell in another. In a sample of 500 cells of
monoploid Triticum monococcum, analysed by Katayama (1935a),
only ten contained a single rod bivalent, whereas Smith (1946) ob-
served a bivalent in three per cent of cells of anothermonoploid of the
same species.
From the results quoted and from others listed in Table I it is clear
that considerabledifferences,eithergeneticallyor environmentallydeter-
mined, may exist between different monoploids of the same species.
However, it is rare for many analyses to be available of monoploids
of a particularspecies, so that conclusionsbased on monoploid meiosis
must of necessity be drawn from the considerationof extremely re-
stricted samples.
The occurrenceof bivalentsis hard to explain in monoploidsderived
from species supposedly with no history of polyploidy which could
have left traces of residualhomology within the gametic complement.
A number of possible explanationshave been advancedto account for
the pairing. Thus it may arise from homologies due to an archaicpoly-
ploid origin, of which pairing in the monoploid is the only trace, the
remote diploid ancestorsnow being extinct. Alternatively,the homol-
ogies may indicate an aneuploid origin of the contemporarychromo-
some number,certain chromosomeshaving been duplicatedin the past
either completely or in part. Finally, the pairing may be due to dupli-
cated segments, which originally arose through translocation,being
carriedin the diploid species.
Reiger (1957) is amongst those who have proposed a polyploid
origin for an apparentlydiploid species. He drew this conclusionfrom
the occurrenceof bivalents in a monoploid of Antirrhinummajus; but
Ernst (1940), from other monoploids of the same species, considered
that there were only one or two chromosomes,more or less modified

in structure,present in double dose. Thus he postulated an aneuploid

origin of the present basic number. Similarly,from the observationof
one or two bivalents at meiosis in a nine-chromosomemonoploid of
Brassicaoleracea,Thompson (1956) was led to agree with the con-
clusion of Catcheside (1937), from observations of the secondary
pairing of bivalents in the diploid, that six is the true basic number
of the genus. Thompson suggested that in this monoploid three of the
six chromosomeswere representedtwice and three only once.
Nandi (1936) has comparedthe secondaryassociationof bivalents
in diploid rice (Oryza sativa) with the formation of bivalents in the
monoploid (Morinaga, 1943). He concluded from the two forms of
evidence together that this species, with 2n = 24, has resulted from
the combination in an amphidiploid of two species with five pairs
of chromosomesand that one member of each set is duplicated.
From the instances mentioned it is clear that some authors have
been significantlyinfluencedby meiotic pairing in monoploids to draw
conclusions about the origins of diploid species. It seems reasonable
that there should be instancesin which tracesof the origin of a species,
throughaneuploidyor polyploidy,remainin the chromosomestructures
of apparentlyvalid diploids. These may be compared with the allo-
polyhaploids, for example, of Nicotiana tabacum (Nettancourt and
Stokes, 1960), Triticum timopheevi (Riley and Chapman,1957), Gos-
sypium barbadense(Kimber, 1961a) and Sorghum vulgare (Brown,
1943) which contain no more bivalents at first metaphaseof meiosis
than many monoploids. These species would be regarded as good
diploids if no lower members of those polyploid series had persisted.
Extinction of the lower members in the series may therefore be the
only differencebetween some species with bivalent-formingmonoploids
and the polyploids quoted above.
Occasionally,the formationof bivalents in monoploids is due to the
presence of duplicated segments. For example, in monoploids of
Oenothera blandina, Catcheside (1932) showed, on the basis of
chiasma-formationbetween differently placed segments, that the oc-
currenceof bivalents and trivalentscould be ascribedto six duplicated
regions, and appropriatemultivalentswere to be found in the diploid
parent. Similarly,in Antirrhinummajus Rieger (1957) found that 40
per cent of the observedchiasmataoccurredin one duplicatedsegment
of the monoploid.
Little attention has been given to the study of prophasepairing in

monoploids, presumablybecause of the technical difficultiesinvolved.

However, in considering the prophase pattern of the Antirrhinum
majus monoploid, Reiger (1957) pointed out that there was a high
level of prophase pairing, the majority of which did not persist to
metaphasebecause of failure of chiasma-formation.These results led
him to postulate a general pairing tendency which is normally sup-
pressed in the diploid by homologous pairing. McClintock (1933)
observed a similarly high level of prophasepairing in the monoploid
condition of Zea mays, the majorityof which was also unaccompanied
by chiasma-formation.She pointed out that despitethe rarityof chiasma-
formation,the association,at pachytene,appearedto be as intimate as
homologouspairing. Results essentiallysimilar to these have been ob-
tained by Brown (1958) in certain Gossypium hybrids,where again
high prophase pairing was associated with an almost complete lack
of chiasma-formation.The lack of chiasma-formationfollowing inti-
mate prophasepairing, in materialof this type, is a problem greatlyin
need of detailed study.
In conclusionit shouldbe made clear that, whilst there is an apparent
anomaly in the occurrenceof meiotic pairing and chiasma-formation
in some monoploids,this behavior is not universal.Some monoploids
support the reputationof the species in which they occur in that they
give no evidence of possessinganythingother than the basic cytological
complement of the group. Meiotic pairing, however, when it does
happen, can sometimes be used to illuminate in new and interesting
ways the past historyand contemporarystatus of the species.
AUTOPOLYHAPLOIDS
Frequent chromosomesynapses,in some cases associatedwith high
fertility, are the most conspicuousfeatures of this group of haploids.
Complete synapsis has been reported in autopolyhaploidsof Valeriana
sambucifolia (Skalinska, 1954), Partheniumargentatum (Gerstel and
Mischanec, 1950), Medicago sativa (Stanford and Clement, 1955;
Lesins, 1957), Dactylis glomerata (Miintzing, 1943), Bromus inermis
(Elliott and Wilsie, 1948), Solanum tuberosum (Ivanovskaja,1939)
and Sorghumhalepense (Duara and Stebbins, 1939). The Parthenium,
Bromus and Sorghumautopolyhaploidswere self-fertile.
Whilst high meiotic pairing is a feature of autopolyhaploids,com-
plete synapsis does not necessarilyoccur, even though duplicated and
triplicated chromosome sets may be present. Nordenskiold (1945)

describeda situation in autohexaploidPhleum pratense (2n=6x=42)

where only bivalents and univalents were formed at meiosis, even
though some of the segregationratios obtained in the parent species
could be explainedonly by hexasomicsegregation.The autopolyhaploid
of this species (Levan, 1941) makes seven bivalents and seven uni-
valents at first meiotic metaphase,even though each chromosome is
triplicated.A "triploid"plant of Phleum pratense (3n=9x=63) ob-
tained from a polyembryonic seed by Muntzing (1938) made 28
bivalents and seven univalents at first metaphase. This and other
evidence (Nordenskiold, 1955; Miintzing and Prakken, 1940) indi-
cated that bivalents are formed only in multiples of the basic number,
seven. Consequently,there is apparentlysome mechanism which re-
stricts chiasmata-associations to the formation of bivalents.
Just as there is a range of chromosomebehaviourin polyploids,from
bivalent formation in allopolyploids,through the formation of some
multivalentsin segmentalallopolyploids,to frequentmutivalentforma-
tion in autopolyploids,so there is a correspondingrange in the chromo-
some behaviourof polyhaploids.Autopolyhaploidshave complete, or
almost complete, chromosome pairing, whilst allopolyhaploids have
only infrequent chromosome synapses. However, between these two
extremes there is a range of patterns of chromosomepairing, and it
is possible to relate this pairing to the polyploid statusof the parent.
Probablythe best range of types of chromosomebehaviourhas been
found in the Solanaceae.In addition to examples of allopolyhaploids
such as Nicotiana rustica, N. tabacum and N. repanda,and autopoly-
haploids such as Solanum tuberosum, there were polyhaploids with
intermediatebehaviours,for example, Solanum demissum, S. nigrum
and S. polytrichon.
Polyhaploidsof the hexaploid species Solanumdemissumhave been
recordedby several authors (Dodds, 1950; Bains and Howard, 1950;
Howard and Swaminathan,1953; Marks, 1955a), and all the meiotic
analysesagree well with each other (Table I). The numberof bivalents
in Solanum demissum polyhaploids is less than the number in poly-
haploidsof the tetraploidspecies Solanumtuberosum,and yet is higher
than would be expected in an allopolyhaploid.Clearly the status of
Solanum demissum lies somewherebetween those of the two extreme
types that give rise to allopolyhaploidand autopolyhaploid.It is prob-
able that the polyploid status of Solanum demissum can be most
accuratelydescribedby the term "segmentalallopolyploid" (Stebbins,

1947) which implies that there is some structuralsimilarity between

the componentgenomes but that they are not identical.The polyhaploid
of the hexaploid species Solanum polytrichon (Marks, 1955b) has a
mean frequencyof 7.9 bivalentsper cell at meiosis (Table I) and again
representsa type intermediatebetween autopolyhaploidsand allopoly-
haploids.
The behaviourof Solanumnigrumpolyhaploidsis somewhatdifferent
from that of the polyhaploidsof Solanumdemissumand Solanumpoly-
trichon. The polyhaploid forms approximately 12 bivalents and 12
univalents per cell (Jorgensen, 1928). Nakamura (1937) noted a
strong morphological resemblance between the hexaploid Solanum
nigrum and a diploid species, Solanum nodiflorum.Stebbins (1950),
therefore, regards Solanum nigrum as a probable auto-allopolyploid
with four genomes of Solanum nodiflorum or some closely related
species and a further two genomes from another diploid, as yet
unidentified.
The contrast between autopolyploidsand allopolyploids represents
more than a separation into two distinct classes; intermediatetypes,
such as segmentalallopolyploidsand auto-allopolyploids,are also found.
Recognition of these intermediatetypes is facilitatedby examinationof
the meiosis of their polyhaploids,where each chromosomemay lack
its complete homologue, as well as by the diminished multivalent for-
mation and modifiedpolysomic inheritancein the parent species.
ALLOPOLYHAPLOIDS
Meiotic data are available on haploids of 16 distinct allopolyploid
species (Table I), and in all except one there was a low level of bi-
valent formation. The data concerning the one exception, Triticum
compactum,in which there was no bivalent formation (Gaines and
Aase, 1926), are somewhat dubious, since polyhaploids of a closely
related species, Triticum aestivum, regularly form bivalents with a
low frequency.The results for allopolyhaploidscontrastratherstrikingly
with the data for monoploidsin which only 50 per cent of the species
for which data were availableshowed bivalent formation.
It has generallybeen assumedthat meiotic pairing in allopolyhaploids
is due to the association of corresponding,homoeologous (Huskins,
1931) chromosomes from the component genomes. This certainly
seems the most likely explanation but it is a hypothesis which it is
hard to test.

The only definite evidence on the relationshipsbetween the paired

chromosomeshas been provided by Okamoto and Sears (1956, 1962)
and Sears (1958). Their test depended upon the pollination of 21-
chromosomeallopolyhaploidindividuals of hexaploid Triticum aesti-
vum with pollen from normal 42-chromosome plants. Occasionally,
under these circumstancesmore or less unreducedegg cells functioned.
Where the egg cell was the appropriate segregant of a megaspore
mother cell meiosis in which bivalents had been formed, the non-
homologous chromosomes involved in the bivalents had undergone
reciprocaltranslocationrelative to the original structuralcondition as
a result of chiasma-formation.Translocationheterozygoteswere thus
derived, and by using appropriatetester lines the chromosomescould
be identifiedwhich were involved in the translocations,that is, between
which bivalent formationhad occurredin the allopolyhaploid.
In the 13 translocationsisolatedby this procedure,eight distinct pairs
of non-homologous chromosomes were involved. Nine independent
translocationsinvolved four pairs of homoeologouschromosomes,and
all of these had resulted from pairing between A and D genome
chromosomes.Of the four translocationsresulting from non-homoeol-
ogous pairing, two were between chromosomeswithin the A genome
and two between non-homoeologouschromosomes,respectively,in the
A and the B genomes. From this it seems clear that the majority of
pairing in allopolyhaploidsof Triticum aestivum is between homoeol-
ogues, and it may well be that this will generallybe found to be true
when more informationof this nature becomes available for allopoly-
haploids.
It has been pointed out by Riley (1960) and Kimber (1961a) that
pairing in allopolyhaploidscan be comparedwith that in hybrids be-
tween the putative diploid parentsof the species concerned.In this way
it should be possible to determine whether the meiotic affinityof the
chromosomesof the component genomes has been modified since the
origin of the polyploid state (Kimber and Riley, 1963). When this
is done, in some instancesthere is no essentialdifferencein pairing and
apparentlyno modificationhas occurred.This is true of the comparison
between the allopolyhaploidof the tetraploid species Brassica napus
(Olsson and Hagberg, 1955) and the hybrid between the diploid par-
ents of the tetraploid,Brassica campestrisand Brassica oleracea (U,
1935).
In most instances,however,pairing in the naturalallopolyhaploidis

considerablyreduced comparedwith that in the "raw"polyhaploid of

hybridorigin. For example,the allopolyhaploidof the tetraploidspecies
Nicotiarnatabacum (Lammerts, 1934) has a mode of zero and a
maximum of three bivalents,whereas the "raw"polyhaploid resulting
from hybridizationbetween Nicotiana sylvestrisand Nicotiana tomen-
tosiformis has a mode of three and a maximum of seven bivalents
(Goodspeed, 1934). Similarly, there is less pairing in the allopoly-
haploid of the tetraploid species Brassica juncea than in the hybrid
between its probable parents, Brassica campestrisand Brassica maja
(Ramanujamand Srinivasachar,1943). An analogoussituation occurs
in the allopolyhaploidof the tetraploidspecies Aegilops ovata (Matsu-
mura, 1940; Kihara, 1937). In addition,whilst allopolyhaploidsof the
tetraploidspecies Gossypiumbarbadenserarelyform bivalents, hybrids
between species closely related to the original diploid parents, Gossy-
pium arboreumand Gossypium thurberi,average eight bivalents per
cell (Skovsted, 1937).
Thus evidence can be derived from meiotic pairing in a number
of allopolyhaploidswhich indicates reduced affinitybetween homoeol-
ogous chromosomes.It should be pointed out that this evidence of
differentiationbetween the constituent genomes is more critical than
that depending upon the absence of multivalent formation in normal
plants of polyploid species. This is because,in the allopolyhaploidcon-
dition, there are no differentialaffinitiescausing complete homologues
to pair at the expense of pairing between homoeologues.
Two suggestions have been advancedto account for the differentia-
tion between chromosomesof the constituent genomes of polyploids.
Eitherthe differentiationis due to the accumulationof structuralmodi-
fications which cause divergence of the chromosomesets, or, alterna-
tively, it depends upon alterationsin the genetic regulationof meiosis.
Recent evidence indicates that the latter is the true cause of reduced
affinity,at least in some instances.
Riley and Chapman(1958) found that in 20-chromosomenullisomic
allopolyhaploidsof Triticum aestivum, deficient for chrososome5B of
the normal gametric complement, there was greatly increasedpairing
relative to that occurringat meiosis in normal21-chromosomeallopoly-
haploids. This was not solely due to the chromosome number, since
nullisomic allopolyhaploids deficient for chromosomes lB and 1D
(Riley, 1960) and 7B (Riley, unpublished) or for anotherunidentified
chromosome (Pai and Swaminathan,1959) did not differ in pairing

at meiosis from normal allopolyhaploids.The modificationof the pair-

ing pattern is evidently attributable,therefore, to the removal of the
activity of specific genes carried on chromosome 5B alone (Riley,
1960; see also ADDENDA).
In the 20-chromosomenullisomic-5Ballopolyhaploidsmean pairing
as high as 3.55 bivalents plus 2.00 trivalents per cell could occur,
whereasthe highest mean pairing observedin 21-chromosomeallopoly-
haploids was 1.69 bivalents and 0.02 trivalentper cell (Riley, 1960).
The high frequencyof trivalentswas taken as a strong indication that
the activity of chromosome 5B in the normal allopolyhaploidcondi-
tion was responsible for the limitation of pairing between homoeol-
ogous chromosomesof the differentgenomes. As an outcome of these
results it has been possible to demonstrate,by the observation of
40-chromosomenullisomic plants, that the activity of chromosome5B
is also responsible for the strictly bivalent-formingmeiotic regime in
42-chromosomehexaploid individualsof Triticum aestivum.
Kimber (1961a) has concluded that the diploid-like meiotic be-
haviour of tetraploidGossypiumspecies may also have a genetic basis.
He has comparedmeiotic pairing in allopolyhaploidsof the tetraploid
species Gossypiumbarbadensewith that in the "raw"polyhaploidsand
in the two kinds of triploid hybridinvolving the tetraploidspecies and
each of its diploid progenitors.The small amount of pairing in the
allopolyhaploidis inconsistentwith the affinitiesbetween the genomes
which can be adduced from pairing in hybrids. This can apparently
be explained only if homoeologouspairing is genetically restrictedin
the tetraploidspecies and its allopolyhaploids.
Endrizzi (1962) has pointed out that the multivalent formation in
synthetic allotetraploidcotton is much less than would be predicted
from the pairing of the F1 hybridand concludedthat a genetic mecha-
nism is unlikely to be responsiblefor this behaviour.Endrizzisuggests,
alternatively,that the reducedmultivalentformationmay be related to
the differential contraction of the A and D genome chromosomes.
However, chiasma formation takes place before the differential con-
traction,and it is therefore difficult to see how this contractioncould
be the basis of the reduction in multivalent formation. Furthermore,
the evidence of high pairing in the F1 hybrid versus low pairing in
the haploid must be considereda more sensitive test, for there is not
the complication that each chromosomehas a homologous partner as
in the synthetic allotetraploid.

A genetic isolation of homoeologues is suggested by the behaviour

of an allopolyhaploidof Nicotiana tabacum derived from a mutant,
coral, by Lammerts (1943). The parent of this allopolyhaploidwas a
translocationheterozygote,and the haploid resulted from the function-
ing of a gamete carrying a duplication-deficiencycondition for the
chromosomes involved in the translocation.Meiotic pairing in this
allopolyhaploidwas much higher than in normal allopolyhaploidsof
Nicotiana tabacum.Moreover, the extra pairing was considerablyin
excess of that which could be ascribed solely to the presence of the
duplicated segment. A possible interpretationof these observationsis
that the chromosomesegment deficient from the allopolyhaploidcar-
ried factors,like those on chromosome5B of Triticumaestivum,which
normallyrestrictpairing between homoeologouschromosomes.
From the preceding discussion it is clear that meiotic pairing in
allopolyhaploidsmay often be an uncertainindication of the true rela-
tionships between chromosomes of the sub-genomes of the species
concerned.The occurrenceof some pairing even in normal allopoly-
haploids of Triticum aestivum indicates that the genetic restrictionof
intergenomic pairing is incomplete. Nevertheless, the more detailed
knowledge of the system in Triticum must emphasisethe caution with
which conclusionsshould be drawn from pairing in allopolyhaploids.
ANEUHAPLOIDS
Haploid plants with aneuploid chromosomeconstitutionshave been
recorded in a small number of species. The best examples are in
Triticum aestivum, in which the complete range of aneupolyhaploids
listed in Figure 1 has been described.
Nullisomic allopolyhaploidsof Triticum aestivum lacking chromo-
somes lB and ID were not differentfrom the normal allopolyhaploids
in bivalent formation (Riley, 1960); neither was an unidentifiednul-
lisomic allopolyhaploid analysed by Pai and Swaminathan (1959).
However, as has been discussedearlier, all five nullisomic-5Ballopoly-
haploids had greatly increased pairing at meiosis, apparentlydue to
the absence of one or more genes, carriedon this chromosome,which
are responsiblefor the preventionof homoeologouspairing (Riley and
Chapman,1958; Riley, 1960).
A substitution allopolyhaploidof Triticum aestivum, with chromo-
some IV of Secale cereale replacing chromosome1B of Triticum, did
not differ at meiosis from normal allopolyhaploids of the species.

Neither did the addition allopolyhaploids which had chromosome

II or chromosomeIII of Secale cereale added to the normal gametic
complement of Triticum aestivum. The Secale chromosome merely
made an extra univalent (Riley, 1960). The meiotic behaviour of a
22-chromosomeaddition allopolyhaploidof Triticum aestivum, which
carriedan unidentifiedchromosomeof Aegilops longissima,was similar
to those with additionalSecale cereale chromosomes (Kimber, unpub-
lished).
A 21-chromosomemisdivision allopolyhaploidof Triticum aestivum,
with an iso-chromosomeformed from the long arm of chromosomeSB,
but completely deficient for the short arm of the same chromosome,
was describedby Riley (1960). The iso-chromosomeusually formed
interbrachialchiasmata,and so was present as a ring-shapedunivalent.
Except for this iso-chromosomering, meiosis was like that of a normal
allopolyhaploid,so that from this it appears that the activity of the
long arm alone is effective in preventing homoeologous pairing in
Triticum.
Finally,amongst the aneupolyhaploidsof Triticum aestivum,Kimber
(1961b) has analysed a 22-chromosomedisomic haploid which had
an unidentifiedchromosomeof the gametic complementin the disomic
condition. This pair of chromosomesregularlyformed a ring bivalent
but otherwise the meiosis was like that of a normal 21-chromosome
allopolyhaploid.
The remarkablestabilityof the controlof haploidmeiosis in Triticum
aestivum is emphasisedby the absence of any modificationof the pat-
tern of pairing in plants with these aneuploid conditions. Indeed, in
view of the similarity of all the other aneuploid types to the normal
allopolyhaploid,the influenceof the nullisomic-5Ballopolyhaploidstate
on meiosis is even more striking.
In addition to the forms described in Triticum aestivum, aneuhap-
loids have also been reportedin Matthiolaincana,Datura stramonium
and Poa pratensis.Lesleyand Frost ( 1928) discovereda disomic mono-
ploid in Matthiolaincana,in which the extra componentto the gametic
chromosome set was a fragment which always remained unpaired at
meiosis.
Disomic monoploids in Datura stramonium (Avery et al., 1959)
were derived from primaryand secondarytrisomics. Using the termi-
nology employed in Datura work, the primary disomic monoploid
investigatedwas ln + 13.14, whilst the secondarydisomic monoploids

were ln + 5.5, ln + 7.7 and In + 15.15. In a furtherdisomic mono-

ploid the disomic chromosomewas not identified.
Instead of 12 univalents, like the normal monoploid, the primary
disomic monoploid had one bivalent and 11 univalentsat meiosis. The
secondarydisomic forms usually had 13 univalents, although in some
instances the extra chromosomeassociatedwith the unalteredchromo-
some from which it had been derived by misdivision.
An aneuhaploid of a 72-chromosome Poa pratensis (Kiellander,
1942) had 18 chromosomesand was fertile, its progeny having 16 as
a typical chromosomenumber.
SECONDARY PAIRING OF UNIVALENTS

At first metaphaseof meiosis in haploids,in addition to associations
dependentupon chiasmata-bivalents or multivalents-secondary asso-
ciations of univalentsalso occur.They have been reportedin Triticum
aestivum,T. timopheevi and Aegilops longissima (Riley and Chapman,
1957), in Gossypium barbadenseand G. hirsutum (Webber, 1938;
Beasley, 1942; Kimber, 1961a), in Solanum demissum (Dodds, 1950;
Bains and Howard, 1950; Marks, 1955a) and in Citrullus vulgaris
(Swaminathanand Singe, 1958).
There are two kinds of secondaryassociationin Triticum, those in
which the associatedunivalentslie side-by-sideand parallel,and those
in which they lie end-to-end and often appear to be connected by a
chromatin thread (Katayama, 1935a; Kostoff, 1938; Krishnawamy,
1939; McGinnis and Unrau, 1952; Person, 1955; Riley and Chapman,
1957). Person (1955), Riley and Chapman (1957) and Kimber
(1961b), using different mathematicaltreatments,have all concluded
that the frequency of side-by-sideassociationwas inversely correlated
with the frequency of associations dependent upon chiasmata.Con-
sequently the chromosomesassociated in this way are probably seg-
mentally homologous,or homoeologous.
By contrast there was no relationship between end-to-end associa-
tions and chiasmataassociations,so that the relationshipsbetween the
univalents involved probably do not depend upon homology. It has
been suggested (Riley and Chapman, 1957) that the occurrenceof
both types of associationis due to the fusion of heterochromaticregions
and that the side-by-sidealignment results from such fusion following
prophasepairing in which no chiasmatawere formed.
Kimber (1961a) demonstratedthat, in allopolyhaploidsof Gossy-

pium barbadensein which chromosomesof the two genomes are of

different sizes, secondary association took place between large and
small univalentsmuch more often than should occur if the processwere
random.This suggested that the secondarilyassociatedunivalentswere
probably corresponding,homoeologous chromosomes from the two
genomes. Thus the secondarypairing of univalents in Gossypiummay
well be analogous to the formation of side-by-side associations in
Triticum.
MORPHOLOGYAND ANATOMY
Haploids are characterisedmorphologicallyby a reductionin size of
all vegetative and floral organs. The most notable exception was a
Datura pruinosa haploid (Satina, Blakeslee and Avery, 1937) which
approachedthe vigour of a normal diploid, but the foliage was dis-
tinctly finer; in addition, Duara and Stebbins (1952) found a poly-
haploid Sorghum that was taller, had fewer tillers, broaderleaves and
more numerouspanicle branchesthan the parentalstrain.
Kostoff (1943) gave tables of the results of his investigationsinto
the morphology of Nicotiana haploid material. He investigated the
general growth, leaf size and index (breadth divided by length), the
size of the flowers,the time of onset of flowering,and the length and
breadth of seeds produced on polyhaploid and tetraploid plants. The
leaves, flowers, leaf index and overall plant size were smaller in poly-
haploids than in normal individuals.The seeds from the polyhaploid
were of a size similar to that of the tetraploid,although the seeds from
the polyhaploid were slightly narrower.The polyhaploids, with one
exception, flowered somewhat earlier than the tetraploids, but the
differences were probably not significant. Kostoff also compared the
alkaloid content of polyhaploid and normal Nicotiana sylvestris, and
TABLE II
DIAMETERS OF POLLEN MOTHER CELLS IN PLANTS OF Datura stramonium AT
DIFFERENT LEVELS OF PLOIDY, FROM BELLING AND BLAKESLEE (1923).
No. of Observed Calculated

Plant P.M.C. diameter diameter*
x 100 1.00 1.00

2x 100 1.32 1.26
3x 40 1.43 1.44
4x 100 1.69 1.59
*The fourth column gives the cube roots of 1, 2, 3 and 4.

found that in autumn the alkaloid content of the polyhaploid leaves

was considerablygreater.
Belling and Blakeslee (1923) produced the series x, 2x, 3x and 4x
in Datura stramonium,measuredthe diameters of the pollen mother
cells and obtainedthe results shown in Table II, which indicate a close
dependenceof cell size on chromosomenumber.
Other workers have demonstratedan approximate2:1 relationship
between diploid and haploid cell volumes. Ivanov (1938) found that
the somatic cells of the root meristem were almost, and the nucleus
exactly,twice as large in diploid as in haploidNicotiana rustica.Cooper
(1943) recordeda similar relationshipof the volumes of the nuclei of
haploid and diploid embryosof Lilium; and Lamm (1938) noted that
the stomatalguard cells of a Solanumhaploid were smaller than in its
diploid twin.
Sinnott, Houghtaling and Blakeslee (1934), in a comprehensive
investigation of a polyploid series in Datura stramonium,found that,
in the series x, 2x, 3x and 4x, the volume of epidermalcells was closely
related to chromosomenumber. However, the cross-sectionalarea of
pericycle fibres and the size of starch grains were almost constant in
the four members of the series, and the area of the vasculartissue of
the pedicel increased at a rate greater than that expected from the
simple numericalrelationshipof the polyploid types.
Kimber (1961b) has shown that the length of the stomatal guard
cells in Gossypium barbadenseand G. hirsutum polyhaploids and
tetraploidsis related by the factor 1.26 (cube root of two) and that
the ploidy of the two plants derived from a polyembryonicseed can
be recognised by using this factor.
Darlington (1937) summarisesthe effect of polyploidy on cell size
as follows: "It seems as though doubling will increase the size of the
normal type in most cases, but that some organismsare more strictly
adapted to a specific size so that particular genetic conditions in-
fluencing size, such as chromosome number, may have a deleterious
effect."Observationsof haploidmorphologysupportthe generalvalidity
of this conclusion.
ORIGIN
It is generally assumed that haploid sporophytesarise from some
haploid component of the embryo sac. However, androgenichaploids
have been recorded on seven occasions. For example, Clausen and
Lammerts(1929) made the cross Nicotiana digluta (2n=72) X Ni-

cotiana tabacum (2n=48) and obtained a haploid Nicotiana tabacum

derivative with 24 chromosomes.Kostoff (1929) obtained a small
Nicotiana langsdorfiiplant with nine somatic chromosomesfrom the
cross Nicotiana tabacum var. macrophylla (3n=72) X Nicotiana
langsdorfii(2n= 18). More recently,from the crossNicotiana glutinosa
(2n=24) X Nicotiana repanda (2n=48), Kehr (1951) obtained a
haploid Nicotiana repanda, a haploid Nicotiana glutinosa, a diploid
Nicotiana repandaand F1 interspecifichybrids.It is suggested by the
authors that these paternal haploid forms resulted from the develop-
ment of a male gamete in female cytoplasm.See ADDENDA.
Kostoff (1942), however, obtained a haploid Nicotiana sylvestris
after pollinating F1 plants of Nicotiana tabacum X Nicotiana sylvestris
with pollen of Nicotiana sylvestris.He suggested a possible alternative
for the apparentlyandrogenicorigin of the Nicotiana sylvestris hap-
loid. If, during megasporogenesisin the F1 plant, all 12 chromosomes
of Nicotiana sylvestrissegregatedto one pole and gave rise to the egg
apparatus,an egg having the haploid constitution of Nicotiana syl-
vestris would be formed. The statisticalprobabilityof this occurrence
is very small; moreover,bivalents are formed between N. tabacumand
N. sylvestrischromosomesat meiosis in the F1 plant and interchange
of chromosome segments must take place. Charactersof Nicotiana
tabacum would therefore be expected in a haploid possessing mainly
Nicotiana sylvestrischromosomes,but they were not observed.Indeed,
Kostoff consideredit unlikely that the haploid originatedin this way.
Gerassimova(1936) emasculatedand treatedwith X-raysa plant of
Crepis tectorum homozygousfor certain dominant genes; it was then
pollinated with pollen from another plant of C. tectorum homozygous
for the correspondingrecessive genes and one haploid with the reces-
sive constitution was found. Whilst this haploid probably had an
androgenic origin like the four previously mentioned, it is also pos-
sible, although unlikely, that a multiple mutation could have taken
place in an egg cell which was stimulatedby the pollination to develop
parthenogenetically.Ehrensberger(1948) was able to induce paternal
haploidy in Antirrhinum majus by irradiation of the egg cells and
subsequentpollination with normal pollen. Ehrensbergerwas also able
to induce maternal haploidy by pollinating normal plants with irra-
diated pollen.
A recent use of irradiationin the productionof haploids is interest-
ing. Swaminathanand Singe (1958) induced a haploid branch on a

watermelonby irradiationof the parentalseed. The origin of the hap-

loid branchis ascribedto a reductionalmitosis; this is the only record
of a haploid being producedin this way.
Determination of the causal stimulation, and identification of the
particularnucleus of the embryo sac involved in the formationof the
embryo, is not easy in haploids formed from maternaltissue. The in-
formation available in the literaturemay, for convenience,be divided
into three groups,namely,spontaneousorigin, origin after experimental
treatment,and origin as a result of polyembryony.
SPONTANEOUS ORIGIN
Haploids having a spontaneous origin represent a heterogeneous
group comprising,in effect, all cases for which the causalstimulationis
unknown. For example, haploids have been found in field cultures of
tomatoes (Morrison, 1932) and in commercialstrainsof cotton under
cultivation (Harland, 1936). The examplesof the spontaneousproduc-
tion of haploids can give little, if any, information about the origin
of the haploid embryo.
ORIGIN AFTER EXPERIMENTAL TREATMENT

Haploids have been observedafter subjectingone or both parentsto
treatmentby one of the following agencies: X-rays,delayedpollination,
cross-pollinationeither intervarietalor interspecific,temperatureshocks,
colchicine treatments (see Table I).
In some cases the haploids have been observed after treatments
designed for other purposes. For example, a haploid of Beta vulgaris
(Levan, 1945) arose after colchicine treatment,and a haploid of Secale
cereale (Miintzing, 1937a) appearedafter low temperaturetreatment
designed to induce polyploidy.
X-rays have been employed on several occasions to induce haploids,
for example, in Nicotiana tabacum (Badenhuizen,1941), N. glutinosa
(Goodspeed and Avery, 1929; Webber, 1933), N. rustica (Ivanov,
1938), Triticum persicum and T. dicoccum (Yefeiken and Vasilev,
1936), T. monococcum (Katayama, 1934), Antirrhinum majus
(Ehrensberger,1948) and Crepistectorum (Gerassimova,1936).
The developmentof haploids after experimentaltreatmenthas most
commonly followed hybridisation,and the first critical account of the
phenomenon was by J0rgensen (1928). Some 90 flowers of Solanum
nigrum were pollinatedby Solanumluteum and 43 fruits were formed,

from which 70 seeds were extracted.Thirty-fiveof the resulting seed-

lings were Solanum nigrum and seven of these were haploid. A study
of the fertilisation processes showed that the Solanum luteum pollen
would germinateon the stigma and the pollen tube would grow down
the style of Solanumnigrum.When the pollen tube reachedthe embryo
sac, both nuclei entered,one fused with the endospermnucleus whilst
the other degenerated. Evidently the presence of these nuclei was
sufficientto stimulate developmentof the embryo.It is presumedthat
the 28 diploid plants from this particularcross had an origin similar
to that of the haploids but that chromosomalduplication had taken
place.
Whilst hybridisationmay increase the frequency of haploidy, there
is evidence demonstratingthat this may not always be the case in
Datura. Blakeslee, Belling and their co-workers (see Kostoff, 1942)
recorded211 haploid Daturaplants, 109 after cross-pollinationand 102
after self- or open-pollination. Satina, Blakeslee and Avery (1937)
found 13 haploids amongst 76,080 plants grown from seed produced
by self-pollinatedplants (0.017 per cent) and 14 haploids amongst
77,358 plants grown from seed produced by cross-pollinatedplants
(0.018 per cent). This differenceis not significant.
Kostoff ( 1942) thought that the frequencyof haploidsin J0rgensen's
work on Solanum should have been calculated from the number of
ovules available and not from the number of normal plants produced
at the same time, as the elimination of hybrid embryos caused an
overestimateof the frequency.He then reworkedJ0rgensen's (1928)
data on the assumption of 30 ovules per fruit and found that the
frequency of haploids was 0.26 per cent. He next calculatedsimilar
estimates for his own Nicotiana haploid material and obtained fre-
quencies of the same orderas those of Satinaet al. (1937) for Datura.
This method of calculationis applicableonly to haploids producedas
the result of hybridisation.
More recently, Chase (1947, 1949a, b, c; 1952a, b) has described
methods of detecting haploids in Zea mays. Stocks homozygous for
certain recessive marker genes affecting seedling characterswere pol-
linated with pollen from plants homozygous for the corresponding
dominant alleles. Seedlings showing the recessive phenotype were
selected,and tested for haploidy.By the use of characterswhose effects
were visible at the seedling stage, large numbers of plants could be
examined with a minimum of labour, although androgenic haploids

would not be detectedby this technique.It was possible to select strains

that were good producersof haploids irrespectiveof the male parent.
Male parents were found that could induce a high frequency of hap-
loids irrespectiveof the female parent. The frequencyof haploids was
increased twentyfold by the use of appropriatestocks, and the ratio
haploid: diploid changed from 1:900 to 1:55 (Chase, 1949a, 1952a).
Chase noted that the frequencyof haploids was greater in strains that
had undergone many years of agronomic selection, and he suggested
that the removal of lethal genes by inbreeding and selection was a
significantfactor in the developmentof the capacity for productionof
high frequenciesof haploids.See ADDENDA.
Coe ( 1959) has isolateda stock of maize which producesabout three
per cent haploidsand has shown that the pollen of this stock is effective
in inducing haploids in other lines. Furthermore,the effect is heritable
and can thereforebe transferredto other backgrounds.
Kihara and Tsunewaki (1962) describe the productionof haploids
in two lines where a nucleushad been substitutedinto alien cytoplasm.
One of these lines (developed from a Triticale producedby J. Taylor)
had a frequency of 1.7 per cent haploids, whilst the other line (de-
veloped from the cross Aegilops caudata X Triticum aestivum var.
erythrospermum)had a frequencyof 53 per cent haploids.No haploids
were found in either of these lines when the nucleus was not in alien
cytoplasm.Kihara and Tsunewaki suggest the use of alien cytoplasm
as a new means for the productionof haploids.Mazoti and Mualenberg
(1958) also reported a similar increase in haploidy when nuclei of
maize were substitutedin teosinte cytoplasm.
A stock of Triticum aestivumvar. Holdfast, monosomicfor chromo-
some I, has recently been describedthat in two successivegenerations
produced 35 per cent of haploids (Riley, 1963). About half the seeds
obtained by selfing this stock failed to germinateand had no embryos.
The genetics of the situation are not yet understood.
ORIGIN AS A RESULT OF POLYEMBRYONY

Most informationconcerning the origin of haploids is contained in
the literature dealing with haploids that arose from polyembryonic
seeds. The first recordof such a haploid concernsan embryoin a poly-
embryonicgrain of Oryzasativa (Ramiaha,Parthasarthiand Ramanu-
jam, 1933). Kappert (1933) describedhaploids from polyembryonic
seeds of Linum usitatissimum,whilst Lehman (1925) illustrated a

polyembryonicGossypiumseed which must have been a haploid-diploid

twin, although haploid-diploidpolyembryonicseeds in Gossypiumwere
not specificallyidentified until some time later (Harland, 1936).
Kappert (1933, 1950) showed that the frequencyof polyembryony
in Linum usitatissimumis controlledby recessive genes. The effect of
these genes was seen in two ways-a maternaleffect and a zygotic one.
The maternal effect was the more obvious, anidstrains were selected
that producedhigh or low frequenciesof polyembryony.However, the
maternalcontrol of polyembryonywas not complete, since, if a female
which had a high frequencyof polyembryonyon selfing was pollinated
with pollen from a plant which had a low polyembryonicfrequencyon
selfing, there was a reduction in the number of polyembryonicseeds,
demonstratingthe zygotic effect. The influence of the maternaleffect
was such that the frequencyof polyembryonicseed could not be raised
above the level found on selfing,even when pollination was with pollen
from male parentsknown to have a higher frequencyof polyembryony
when selfed (Kappert, 1950; Table III). The frequency of haploid-
diploid twins was higher in self-pollinated than in cross-pollinated
material (Kappert, 1950).
Chase (loc. cit.) found similar effects in the productionof haploids,
but not by polyembryony,in Zea mays. Morganand Rappleye (1950)
and Christensen and Bamford (1943) observed differences in the
frequency of polyembryonic seeds in various strains of Capsicum
frutescens.
The cell from which a haploid embryo develops, in haploid-diploid
polyembryonicseeds, is almost certainly a synergid. Embryos formed
from synergids were first recordedin Mimosa by Guignard (1881).
Cooper (1943) observed haploid-diploid twin embryos in ovarian
sections of Nicotiana and Lilium, and thought that the formation of
embryos from synergids could account for many of the haploids
recordedin the literature.
The origins of the embryos of some polyembryonicseeds, for ex-
ample, in Capsicumfrutescens,can be determinedby a combinationof
genetical and cytological techniques. By the use of suitable marker
genes, Morgan and Rappleye (1950, 1954) were able to distinguish
many types of polyembryonicseeds, including haploid-haploidtwins,
diploid-diploid-diploidtriplets, diploid-diploid twins and haploid-dip-
loid twins. Since the antipodalsdegenerateprior to fertilisationin this
species, Morgan and Rappleye thought that the maternal haploids

HAPLOID ANGIOSPBRMS 515
arose from synergids. More recently Campos and Morgan (1960)

produced evidence that the frequency of twinning in this species is
controlledby the genotype of the female parent.
Differences in the frequency of polyembryonyin Gossypium have
been recordedby variousworkers (Harland, 1936; Beasley, 1942), but
Kimber (1958) has shown that most of these differencescan be ac-
counted for in Gossypiumbarbadenseby the inadequacyof the earlier
methods of detection. Twins consisting of one large and one small
embryo were found in both Gossypium barbadenseand Gossypium
hirsutum.The frequencyof these so called cryptic twins in Gossypium
barbadensewas equal to that of "normal"sized twins, but, although
the frequencyof normal twins in Gossypiumhirsutumwas low, it was
higher than the frequency of cryptic twins. The frequency of cryptic
plus normal twins in Gossypium barbadensewas 0.74 per cent.
As the antipodalcells in cotton degeneratesome days prior to fertil-
isation (Gore, 1932), haploids most probably originate from un-
fertilisedsynergids.Silow and Stephens ( 1944) have producedevidence
which indicates a maternal origin of the haploid in haploid-diploid
twins in cotton. A cross was made between a female individual
homozygousfor a recessivegene giving green plant colour and a pollen
parent which was homozygousfor a dominant gene giving red plant
colouration.A seed found in the progeny was shown to have haploid
and diploid embryos.The diploid had the expected heterozygouscon-
stitution, whilst the haploid carried the gene for green plant coloura-
tion, like the seed parent.
Silow and Stephens (loc. cit.) also made the cross between a female
parent in the variety Acala Okra,which was homozygousboth for the
recessive gene giving white pollen and for the incompletelydominant
gene giving lacinateleaf, and a pollen parent,in the variety U4, which
was homozygousfor the alternativeallelesgiving yellowpollen andbroad
leaves. Amongst the progeny of this cross was a diploid-diploidtwin,
one member of which had yellow pollen and intermediateleaf shape,
while the other had white pollen and laciniate leaf. The first was the
expected hybrid type and the second was evidently of maternalorigin.
Whilst it is possible that the latter arose from sporophytictissue, Silow
and Stephens consider it more probable that this plant arose from an
unfertilised synergid and that doubling of the chromosome number
took place during development.

Wilson and Ross (1961) have recorded 77 twins of Triticum

aestivum,of which four had a haploid component.
Thus, whilst it is occasionallypossible to be quite sure of the origin
of haploids, on many occasions the evidence is open to several inter-
pretations.Moreover,little is understoodof the causal factors respon-
sible for the developmentof unfertilised eggs. In several instancesthe
productionof haploids has been attributedto externalstimuli-X-rays,
temperature, colchicine, hybridisation-but that these agencies are
effective in some but not in other instancessuggests that the effect of
the genotype is also significant.
USES OF HAPLOIDS
Perhapsthe most obvious practicaluse of haploidsis the exploitation
of the totally homozygous forms that can be produced when the
chromosomecomplementsof haploids are doubled by colchicine treat-
ment. However, despite the fact that this potential use of haploids has
been recognised many years, only one doubled haploid, the Marglobe
tomato, has been grown commercially.There are several difficultiesin
the use of doubledhaploidsas commercialcrops.First of all, there is the
low frequencyof occurrenceof haploids in many species, although,by
the use of polyembryonyor geneticallymarkedseedlings,large popula-
tions can be screened and haploids recoveredmore efficiently.In out-
breeding species it is probable that completely homozygous material
would show the equivalent of inbreeding depression.Furthermore,it
is possible that the genetic balance and phenotypic uniformity of an
outbreeding and consequently heterozygouscrop would be disrupted
by total homozygosity,and this could lead to an unexpected increase
in variability.
Employment of haploids and doubled haploids as tools in plant
breeding, rather than as commercially grown crops, may be more
successful.Chase (1952b), for example, has collected and used double
haploids of Zea mays as the equivalent of inbred parents in the pro-
duction of hybrid corn. Also, Hougas, Peloquin and Ross (1958) have
manipulated Solanum tuberosum haploids in crosses with diploid
Solanum species in attempts to transferdesirablegenes into the culti-
vated potato. Nei (1963) calculatedthe frequenciesof haploids, and
of their diploidisation,that are necessaryfor the haploid method to be
more efficient than a diploid method of breeding. He concluded that
the haploid method of breeding is advantageouswhen the number of

genes concerned is large and the frequencies of favourablealleles in

the population are small, even though the frequencyof haploid occur-
rence is low.
Sears (1954) described the use of haploids in the production of
monosomicsin Triticumaestivum.In the progeny of haploid X diploid
crosses he obtained 94 monosomic plants and was eventually able to
identify the chromosomes deficient from 66 of them. These plants
served in the productionof the monosomic series in wheat which has
been valuable in intervarietalchromosomesubstitution work as well
as in genetic investigations.
However, it is perhaps in studies not directed at the commercial
applicationof haploids that this type of materialhas proved most use-
ful. Haploids have been relied upon in the recognitionof the polyploid
status of the species from which they were derived.Recently, allopoly-
haploids in Triticum aestivum deficient for a particularchromosome
were the starting point of a series of investigations that have led to
the elucidation of the genetic mechanism controlling the diploid-like
chromosomebehaviourof that species (see Riley, 1960).
Also, comparison of the meiosis of an allopolyhaploid with the
meiosis of the F1 between the parentsof the allopolyploidcan indicate
whether there is any genetical control of the chromosomeisolation in
the allopolyploid.Kimber (1961) has made this comparisonfor Gossy-
pium barbadenseand has inferredthat there is genetical control of the
pairing in the polyploid.
CONCLUSIONS
Haploids are interestingcomponentsof the range of geneticallyaber-
rant types that are found when cytogeneticinvestigationsare made into
many angiospermspecies. As a consequenceof their unique constitu-
tion, they offer new possibilities for the investigationsof many cytol-
ogical and genetical problems. They also present a new range of
problems intrinsic to their own chromosomalstatus.
The study of haploidsover the last ten years has helped considerably
in the elucidation of problems concernedwith the evolution of crop
species and with the cytogenetic organisation of allopolyploids and
autopolyploidsinvolved in plant breeding. Continued use of haploids
as cytogeneticaltools will undoubtedlyassist in furtherstudies in these
fields.
It is perhaps the greatest obstacle to the wide use of haploids that

no techniqueexists by which forms with gametic chromosomeconstitu-

tions can be freely producedwhen required.
ADDENDA
Some papers that have considerablesignificancein the study of hap-
loidy have recentlyappearedand must be briefly mentioned.
The first two refer to the origin of plants of Sorghumvulgare with
diploid chromosomenumbers from synthetic tetraploidstreated with
colchicineunder particularconditionsof light, temperatureand humid-
ity. Sandersand Franzke (1962) and Ross, Chen and Simantel (1963)
have shown that in particulartetraploidgenotypes,some cells respond
to the treatmentby somatic changes of chromosomenumbersfrom te-
traploid to diploid, and that the latter may subsequentlytake over the
growing point completely and give rise to entirely diploid individuals.
Meiotic analysisand the use of chromosome-structural markersshowed
that the lower numbersrepresentedthe true diploid complement and
that some undeterminedform of reductiondivision must have occurred.
These results may clearly be of the greatest future significancein as-
sessing means by which polyhaploid conditions can be intentionally
produced.
Chedda and Harlan (1962) have used the meiotic behaviour of
20- and 21-chromosome polyhaploid plants of Bothriochloainter-
medium (2n = 40) to provide part of the evidence that leads them
to conclude that, in the Bothriochloiniae,there is a locus which reg-
ulates meiotic chromosomepairing. The dominant allele at this locus
causes chromosomesto associate in pairs-inhibiting multivalent for-
mation and causing chromosomesthat are not strictly homologous to
form bivalents. When the recessive allele is homozygous,only truly
homologous chromosomespair. This situation is obviously relevant to
our previous consideration of the genetic determination of pairing
specifiicityin wheat. However, there is a markedoperationaldifference
between the two systems.
Finally, a further indication has been provided by Goodsell (1961)
and Chase (1963) of the practicalvalue of monoploids in maize. Use
has been made of androgenetic haploids, recognised by appropriate
genetic markers,to transfer the genotypes of inbred lines into cyto-
plasm that causes male sterility. This system has the advantage,com-
pared with nuclear substitutionby backcrossing,that the chromosomal
system remains intact. Chase has further indicated that, when the

cytoplasmicdonor has a tetraploidnuclearconstitution,the androgenic

substitutionsmay well be diploid instead of monoploid, so obviating
the need for haploidy.
LITERATURE CITED
SEE ALSOSUPPLEMENTARYCITATIONS
AND
CITATIONS
ADDEDIN PROOF
1. AALDERS, L. E. 1958. Monoploidy in cucumbers. Jour. Hered. 49: 41-44.
2. AASE, H. C. 1930. Cytology of Triticum, Secale and Aegilops hybrids
with reference to phylogeny. State Coll. Wash., Res. Stud. 2: 3-60.
3. . 1946. Cytology of cereals. II. Bot. Rev. 12: 255-334.
4. AVERY, A. G., SATINA, S. and RIETSEMA, J. 1959. Blakeslee: The Genus
Datura. Ronald Press, New York.
5. BABCOCK, E. B., and NAVASHIN, M. 1930. The cytology of Crepis. Bib.
Genet. 6: 14-46.
6. BACCHI, 0. 1941. [Cytological observations on Coffea. VII. Macrosporo-
genesis in "monosperma" variety.] Bragantia 1: 483-490.
7. BADENHUIZEN, N. P. 1941. Experimentally produced haploids in Nico-
tiana tabacum by means of X-rays. Natuurewetenschap. Tijds. Neder-
land-Indie. 101: 240-242.
8. BAINS, G. S. and HOWARD,H. W. 1950. Haploid Solanum demissum
plants. Nature [London] 166: 795.
9. BEACHELL, H. M., and JONES,J. W. 1945. Tetraploids induced in rice
by temperature and colchicine treatments. Jour. Amer. Soc. Agron.
37: 165.
10. BEASLEY,J. 0. 1940. The production of polyploids in Gossypium. Jour.
Hered. 31: 39-48.
11. . 1941. Hybridization, cytology and polyploidy of Gossypium.
Chron. Bot. 6: 394-95.
12.- . 1942. Meiotic chromosome behaviour in species, species hy-
brids, haploids and induced polyploids in Gossypium. Genetics 27:
25-54.
13. BELLING,J. and BLAKESLEE, A. F. 1923. The reduction division in hap-
loid, diploid, triploid and tetraploid Daturas. Proc. Nat. Acad. Sci.
60: 106-111.
14. , and . 1927. The assortment of chromosomes in
haploid Daturas. La Cellule 37: 355-61.
15. BLAKESLEE, A., and AVERY,A. 1939. The induction of diploids from
haploids by colchicine treatment. Genetics, 24: 95. [Abs.]
16. , BELLING, J., FARNHAM, M. E., and BERGER, A. D. 1922.
A haploid mutant in Datura stramonium. Science 55: 646-647.
17. , MORRISON, G., and AVERY, A. G. 1927. Mutations in a
haploid Datura. Jour. Heredity, 18: 193-9.
18. , SATINA, S., BERGNER, A. D., and AVERY, A. G. 1933. Pri-
mary and secondary ln + 1 types; a new unbalance of the Jimson
weed. Science 77: 518.
19. BLEIER, H. 1933. Die Meiosis von Haplodiplonton. Genetica 15: 129-176.
20. BOLSUNOV, I. 1939. [An experimentally obtained haploid in Nicotiana
rustica L.] Jour. Inst. Bot. Acad. Sci. Ukraine 21-22: 197-199.
21. BROWN, M. S. 1943. Haploid plants in Sorghum. Jour. Hered. 34: 163-66.

22. . 1958. The implications of pachytene and metaphase pairing

for species differentiation. Proc. X Int. Cong. Genet. Vol. 2: 36.
23. BURD, L. W. 1924. A preliminary note on a sterile dwarf rogue in Sea
Island cotton. Emp. Cot. Grow. Rev. 1: 46-48.
24. CAMERON, D. R. 1949. Inheritance in Nicotiana tabacum. XXII. Investi-
gations on multiple seedlings. Amer. Jour. Bot. 36: 526-529.
25. CAMPOS, F. F. 1958. Haploid parthenogenesis in Capsicum frutescens L
following crosses with untreated and X-rayed pollen. Ph.D. Thesis,
Univ. Maryland.
26. , and MORGAN, D. T. JR. 1958. Haploid pepper from a sperm.
Jour. Hered. 49: 134-137.
27. CA1CHESIDE,D. G. 1932. The chromosomes of a new haploid Oenothera.
Cytologica 4: 68-113.
28. . 1937. Secondary pairing in Brassica oleracea. Cytologia,
Fujii Jub. Vol. 336.
29. CHAPMAN, V. 1960. Ae. caudata haploid. [Personal communication].
30. CHASE, S. S. 1947. Techniques for isolating monoploid maize plants.
Amer. Jour. Bot. 34: 582.
31. . 1949a. Monoploid frequencies in a commercial double cross
hybrid maize, and its component single cross hybrids and inbred lines.
Genetics, 34: 328-332.
32. . 1949b. The reproductive success of monoploid maize. Amer.
Jour. Bot. 36: 795-796.
33. . 1949c. Spontaneous doubling of the chromosome complement
in monoploid sporophytes of maize. Proc. Iowa Acad. Sci. 56: 113-115.
34. . 1952a. Monoploids in maize. In: Gowen, J. W. Heterosis
[ed.] Iowa State Coll. Press.
35. . 1952b. Production of homozygous diploids of maize from
monoploids. Agron. Jour. 44: 263-267.
36. CHIPMAN,R. H., and GOODSPEED,T. H. 1927. Inheritance in Nicotiana
tabacum. VIII. Cytological features of the purpurea haploid. Univ.
Cal., Publ. Bot. 40: 141-158.
37. CHIZAKI, Y. 1933. On the occurrence of haploid plant in monococcum
wheat. Proc. Crop. Sci. Soc. Jap. 5: 267-270.
38. . 1934. Another new haploid plant in Triticum monococcumn.
L. Bot. Mag. Tokyo 48: 621-628.
39. CHRISTENSEN, H. M., and BAMFORD,R. 1943. Haploids in twin seedlings
of pepers, Capsicum annuum. L. Jour. Hered. 34: 99-104.
40. CHRISTOV,M. 1929/30. A haploid tobacco plant. Year book. Univ. Sofia
Fac. Agr. 8: 285-296.
41. CLAUSEN,R. E., and LAMMERTS, W. E. 1929. Interspecific hybridisation
in Nicotiana. X. Haploid and diploid merogony. Amer. Nat. 43: 279-82.
42. , and MANN, M. C. 1924. Inheritance in Nicotiana tabacum.
V. The occurrence of haploid plants in interspecific progenies. Proc.
Nat. Acad. Sci. 10: 121-4.
43. COOK, R. C. 1936. A haploid Marglobe tomato. Practical application of
a "short cut" for making pure lines. Jour. Hered. 27: 432-435.
44. COOPER,D. C. 1943. Haploid-diploid twin embryos in Lilium and Nico-
tiana. Amer. Jour. Bot. 30: 408-413.
45. DARLINGTON,C. D. 1937. Recent advances in cytology. Churchill, London.
46. DAVIS, B. M. 1931. Some attempts to obtain haploids from Oenothera
Lamarckiana. Amer. Nat. 65: 233-242.

47. , and KULKARNI,C. G. 1930. The cytology and genetics of a

haploid sport from Oenothera franciscana. Genetics 15: 55-80.
48. DEANON, J. R. (JUN). 1957. Treatment of sweet corn silks with maleic
hydrazide and colchicine as a means of increasing the frequency of
monoploids. Philipp. Agr. 41: 364-77.
49. DODDS,K. S. 1950. Polyhaploid of Solanum demissum. Nature [London]
166: 795.
50. DUARA, B. N., and STEBBINS, JR., G. L. 1952. A polyhaploid obtained
from a hybrid derivative of Sorghum halepense x S. vulgare var.
sudanense. Genetics 37: 369-374.
51. EHRENSBERGER, R. 1948. Versuche zur Auslosung von Haploidie bei Blu-
tenpflanzen. Biol. Zentralblatt 67: 537-46.
52. EINSET, J. 1943. Chromosome length in relation to transmission fre-
quency of maize trisomes. Genetics 28: 349-364.
53. ELLIOTT, F. C., and C. P. WILSIE. 1948. A fertile polyhaploid of Bromuts
inermis. Jour. Hered. 39: 377-380.
54. EMERSON, S. H. 1929. The reduction division in a haploid Oenothera.
La Cellule 39: 159-65.
55. ENDRIZZI, J. E. 1959. Cytogenetics of four sets of twins in cotton. Jour.
Hered. 50: 222-226.
56. , and MORGAN, JR., D. T. 1955. Chromosomal interchanges
and evidence for duplication in haploid Sorghum vulgare. Jour. Hered.
46: 201-208.
57. ERNST, H. 1940. Zytogenetische Untersuchungen an haploide Pflanzen
von Antirrhinum majus L. Zeits. Bot. 35: 161-190.
58. FORD, L. 1954. A cytogenetic comparison of maize monoploid derivatives
and inbreds. Agron. Jour. 46: 34-36.
59. GAINS,E. F., and AASE, H. C. 1926. A haploid wheat plant. Amer. Jour.
Bot. 13: 373-385.
60. GATES, R. R. 1929. A haploid Oenothera. Nature [London] 124: 948.
61. -, and GOODWIN, K. M. 1930. A new haploid Oenothera, with
some considerations on haploidy in plants and animals. Jour. Genet.
23: 123-156.
62. GERASSIMOVA, H. 1936a. Experimentally produced haploid plant in
Crepis tectorum. Biologitcheski Jour. 5: 895-900.
63. . 1936b. Experimentell erhaltene haploide Pflanze von Crepis
tectorum. Planta 25: 696-702.
64. GERSTEL, D. U., and MISCHANEC, W. M. 1950. On the inheritance of
apomixis in Parthenium argentatum. Bot. Gaz. 112: 96-106.
65. GOODSPEED, T. H., and AVERY,P. 1929. The occurrence of a Nicotiana
glutinosa haplont. Proc. Nat. Acad. Sci. 15: 502-4.
66. GORE, U. R. 1932. Development of the female gametophyte and embryo
in cotton. Amer. Jour. Bot. 19: 795-807.
67. GUIGNARD, L. 1881. Recherches sur l'embryogenie des Legumineuses. Ann.
Sci. Nat. Bot. 6: 5-166.
68. HACHATUROV, S. P. 1937. [Nicotiana tabacum haploid with bivalents at
meiosis.] Bull. Appl. & Genet., Plant Breed. Ser. 2(7): 191-196.
69. HAKANSSON, A. 1940. Die Meiosis bei haploiden Pflanzen von Godetia
Whitneyi. Hereditas 26: 411-429.
70. HARLAND,S. C. 1936. Haploids in polyembryonic seeds of Sea Island
Cotton. Jour. Hered. 27: 229-231.
71. . 1939. The Genetics of Cotton. Jonathan Cape.

72. . 1955. The use of haploids in cotton breeding. Indian Jour.

Genet. & Plant Breed 15: 15-17.
73. HAQUE, A. 1946. Haploid-haploid polyembryony in Sesbania aculeata.
Cur. Sci. 15: 287.
74. HAYASE, H. 1954. [Studies on Cucurbita crosses. V. The occurrence of
twin plants with a haploid chromosome number in the F1 of C. maxima
x C. moschata]. Ikushugaku Zasshi/Jap. Jour. Breed. 4: 115-21.
75. HESLOP-HARRISON,J. W. 1931. The Northumberland and Durham primu-
las of the Section Vernales. Trans. North. Nat. Un. 1: 48.
76. HOLLINGSHEAD,L. 1928. A preliminary note on the occurrence of hap-
loids in Crepis. Amer. Nat. 62: 282-4.
77. . 1930. A cytological study of haploid Crepis capillaris plants.
UJniv. Calif., Pub. Agr. Sci. 6: 107-134.
78. HOvIN, A. W. 1957. Seasonal variation of meiotic chromosome pairing
in haploid Poa annua L. Genetics 42: 377-78. [Abstracts of papers
presented at the 1957 meetings of the Genetics Soc. of America held
at Stamford, Calif. Aug. 26-28/1957.]
79. . 1958. Meiotic chromosome pairing in amphihaploid Poa
annua L. Amer. Jour. Bot. 45: 131-138.
80. HOUGAS, R. W., and PELOQUIN, S. J. 1957. A haploid plant of the potato
variety Katahdin. Nature [London] 180: 1209.
81. , and . 1960. Crossability of S. tuberosum haploids
with diploid Solanum species. European Potato Jour. 3: 325.
82. , , and Ross, R. W. 1958a. Haploids of the com-
mon potato. Jour. Hered. 49: 103-106.
83. , , and . 1958b. The potential of hap-
loid potatoes for research and breeding. Amer. Potato Jour. 35: 442.
84. , , and . 1958c. Induced haploids in
Solanum tuberosum. Amer. Potato Jour. 35: 441.
85. HOWARD, H. W., and SWAMINATHAN, M. S. 1953. The cytology of hap-
loid plants of Solanum demissum. Genetics 26: 381-391.
86. HUMPHREY, L. M. 1934. The meiotic divisions of haploid, diploid and
tetraploid tomatoes with special reference to prophase. Cytologia 5:
278-300.
87. HUSKINS, C. L. 1931. A cytological study of Vilmorins unfixable dwarf
wheat. Jour. Genet. 25: 113-124.
88. IVANOV, M. A. 1938. Experimental production of haploids in Nicotiana
rustica L. Genetica 20: 295-397.
89. IVANOVSKAJA, E. V. 1939. A haploid plant of Solanum tuberosum L.
Comp. Rend. (Doklady) Acad. Sci. U.R.S.S. 24: 517.
90. JOHANSON, D. A. 1934. Haploids in Hordeum vulgare. Proc. Nat. Acad.
Sci. [U.S.A.] 20: 98-100.
91. J0RGENSEN, C. A. 1928. The experimental formation of heteroploid plants
in the genus Solanum. Jour. Genet. 19: 133-211.
92. KAPPERT, VON H. 1933. Erbliche Polyembryonie bei Linum usitatissimum.
Biol. Zentralblatt 53: 276.
93. . 1950. Botanishe untersuchiengen zur Erblichkeit der Poly-
embryonie. Moderne Biologie. F. W. Peters. Berlin.
94. KASPARAYAN, A. S. 1938. Haploids and haplo-diploids among hybrid
twin seedlings in wheat. Comp. Rend. (Doklady) Acad. Sci. U.R.S.S.
20: 53-56.

95. KATAYAMA Y. 1934a. Experimental formation of haploid plants in Tri-

ticum monococcum. Bull. Alumni Assoc. Utsumomiya Agr. Col. 1: 41-50.
96. . 1934b. Haploid formation by X-rays in Triticum monococ-
cum. Cytologia 5: 234-237.
97. . 1935a. Karyological comparisons of haploid plants from
AIegilotricum and diploid wheat. Jap. Jour. Bot. 7: 349-380.
98. . 1935b. Haploid plants in Japanese morning glory. Jap. Jour.
Genet. 11: 279-281.
99. KEHR, A. E. 1951. Monoploidy in Nicotiana. Jour. Hered. 42: 107-112.
100. KHOSHoo, T. N. 1957. A polyhaploid plant of the tetraploid race of
Sisymbrium irio. Jour. Hered. 48: 238-242.
101. KHRISTOFF, D. 1930. A haploid tobacco plant. Ann. Univ. Sofia v. Facul.
Agron. 8: 285-296.
102. KiDD, H. J. 1952. Haploid and triploid Sorghum. Jour. Hered. 43:
204-235.
103. KIELLANDER, C. L. 1942. A subhaploid Poa pratensis L. with 18 chromo-
somes and its progeny. Svensk. Bot. Tidskr. 36: 200-220.
104. . 1943. Die Entwicklung des Embryosacks und die Befruch-
tung bei Poa alpina. Bot. Not. 85-95.
105. KIHARA, H. 1936. Ein diplo-haploides Zwillingspaar bei Triticum durum.
Agr. & Hort. [Tokyo] 11: 1425-1433.
106. . 1940. [Formation of haploids by means of delayed pollina-
tion in Triticum monococcum]. Bot. Mag. Tokyo 54: 178-185.
107. , and KATAYAMA, Y. 1932. Uber das Vorkomman von hap-
loider Pflanzen bei Triticum monococcum. Kwagaku 2: 408-410.
108. , and . 1933. [On the maturation division of hap-
loid Triticum monococcum]. Agr. and Hort. [Tokyo] 8: 2725-2738.
109. , and YAMASHITA, K. 1938. [Artificial production of haploid
and triploid einkorn wheats by pollination with X-ray irradiated pol-
len.] In: Commemoration Papers on Agronomy Prepared in Honour of
Prof. Masas Akemine on the occasion of the Thirtieth Anniversary of
his Academic Service by his Friends and pupils. 9-20.
110. KIMBER, G. 1958. Cryptic twin plants in New World cotton. Emp. Cot.
Grow. Rev. 35: 24-25.
111. -. 1961a. The basis of the diploid-like meiotic behaviour of
polyploid cotton. Nature [London] 191: 98-100.
112. KING, E. E. 1957. [Personal communication].
113. KISCH, R. 1940. Morphologie und Zytologie haploide Pflanzen von Epilo-
bium hirsutum. Zeits. Bot. 36: 513-537.
114. KOMATSU, Y. 1936. [Investigations on the progeny of haploid plants and
embryo sac formation by diploid and haploid plants of Brassica na-
pus.] Proc. Crop. Sci. Soc. Japan 8: 364-72.
115. KOSTOFF, D. 1929. An androgenic Nicotiana haploid. Zeits. Zellforsch
9: 640-642.
116. . 1934. A haploid plant of Nicotiana sylvestris. Nature [Lon-
don] 133: 949.
117. I . 1936. Studies on polyploid plants. 13. Haploid Nicotiana
rustica. Comp. Rend. Acad. Sci. U.R.S.S. 1: 239-242.
118. . 1938. Heterochromatin at the distal ends of the chromo-
somes in Triticum monococcum. Nature [London] 141: 690-691.
119. . 1939. Frequency of polyembryory and chlorophyll defi-
ciency in rye. Comp. R. (Doklady) Acad. Sci. U.R.S.S. 24: 479-482.

120. . 1942. The problem of haploidy. (Cytogenetic studies in

Nicotiana haploids and their bearing on some other cytogenetic prob-
lems.) Bib. Genet. 13: 1-148.
121. . 1943. Haploide Triticium 'vulgare und die Variabilitat ihrer
diploiden Nachkommenschaften. Zuchter 15: 121-125.
122. , and AKSAMITAIA, A. 1936. Chromosome conjugation in hap-
loids and its genetical significance. Comp. Rend. Acad. Sci. (Doklady)
U.R.S.S., 1: 330-332.
123. KRISHNAWAMY, N. 1939. Cytological studies in a haploid plant of
Triticum vulgare. Hereditas 25: 77-86.
124. KULKARNI, C. G. 1932. Chromosome configurations during the hetero-
typic prophase and metaphase in certain Oenotherae and their genetic
significance. Pap. Mich. Acad. Sci., Arts & Litt. 15: 83-95.
125. KURIYAMA, H., and WATANABE, Y. 1955. Studies on the haploid plant
of Brassica carinata. Ikushugaku Zasshi/Jap. Jour. Breed. 5: 1-5.
126. LAMM, R. 1938. Notes on a haploid potato hybrid. Hereditas 24: 391.
127. LAMMERTS,W, E. 1934. On the nature of chromosome association in
Nicotiana tabacum haploids. Cytologia 6: 38-50.
128. LEHMAN, S. G. 1925. Two rare types of abnormality in cotton seeds.
Jour. Elisha Mitchell Sci. Soc. 41: 138-140.
129. LELIVELD, J. A. 1932. Cytological observations on the diploid offspring
of a haploid Oenothera franciscana. Cellule 41: 281-289.
130. LESINS, K. 1957. Cytogenetic study on a tetraploid plant at the dip-
loid chromosome level. Canad. Jour. Bot. 35: 181-196.
131. LESLEY, M. M., and FROST, H. B. 1928. Two extreme "small" Mathiola
plants-a haploid with one and a diploid with two additional chromo-
some fragments. Amer. Nat. 62: 22-33.
132. LEVAN, A. 1941. Syncyte formation in the pollen mother cells of haploid
Phleum pratense. Hereditas 27: 243.
133. . 1942. Studies on the meiotic mechanism of haploid rye.
Hereditas 28: 171-211.
134. . 1945. A haploid sugar beet after colchicine treatment.
135. LINDSTROM,E. W. 1929. A haploid mutant in the tomato. Jour. Hered.
20: 23-30.
136. . 1941. Genetic stability of haploid, diploid and tetraploid
genotypes in the tomato. Genetics 26: 387-97.
137. , and Koos, K. 1931. Cytogenetic investigations of a haploid
tomato and its diploid and tetraploid progeny. Amer. Jour. Bot. 18:
398-410.
138. MARKS, G. E. 1955a. Cytogenetic studies in tuberous Solanum species.
1. Genomic differentiation in the group Demissa. Jour. Genet. 53:
262-269.
139. . 1955b. A polyhaploid plant of Solanum polytrichon. Nature
[London] 175: 469.
140. MATSUMURA, S. 1940. Induzierte Haploidie und Autotetraploidie bei
Aegilops ovata. L. Bot. Mag. Tokyo 54: 404-13.
141. MCCLINTOCK,B. 1933. The association of non-homologous parts of
chromosomes in the mid-prophase of meiosis of Zea mays. Zeits.
Zellforsch. & Miknosk. Anat. 19: 191-237.
142. MCCRAY, F. 1932. Another Nicotiana tabacum haploid plant. Bot. Gaz.
93: 227-230.

143. McGINNIs, R. C. and UNRAU, J. 1952. A study of meiosis in a haploid

of T. vulgare and its progeny. Canad. Jour. Bot. 30: 40-49.
144. MEYER, J. R., and JUSTUS, N. 1960. Potential value of doubled haploids
in cotton. [Abs.] Genetics 45: 999.
145. MIDUNO, T. 1940. Chromosomenstudien an Orchidazeen. III. U0ber das
Vorkomman von haploiden Pflanzen bei Bletilla striata. Reichb. f. var
gebina Reichb f. Cytologia 11: 156-176.
146. MODILEWSKI,J. 1939. [Cytogenetic investigations of the genus Nicotiana.
IX. Cytology and embryology of the haploid N. rustica]. Jour. Inst.
Bot. Acad. Sci. Ukraine. Nos. 21-22: 201-15.
147. MORGAN, D., and RAPPLEYE, R. 1950. Twin and triplet pepper seedlings.
A study of polyembryony in Capsicurm frutescens. Jour. Hered. 41:
91-95.
148. , and . 1954. A cytogenetic study of the origin of
multiple seedlings of Capsicum frutescens. Amer. Jour. Bot. 41: 576-586.
149. MORINAGA, T. and FUKUSHIMA, E. 1931. Preliminary report on the hap-
loid plant of rice, Oryza sativa L. Proc. Imp. Acad. [Tokyo] 7: 383-4.
150. , and . 1932. Some observations on microsporo-
genesis of the haploid plant of rice. Proc. Imp. Acad. [Tokyo] 8:
403-405.
151. , and . 1933. Karyological studies on a spontan-
eous haploid mutant of Brassica napella. Cytologia 4: 457-460.
152. , and . 1934. Cytogenetical studies on the haploid
plant of rice. Jap. Jour. Bot. 7: 73-106.
153. MORRISON,G. 1932. The occurrence and use of haploid plants in tomato
with especial reference to the variety Marglobe. Proc. VI Int. Cong.
Genet. Vol. 2: 137.
154. MUNTZING, A. 1937a. Note on a haploid rye plant. Hereditas 23: 401.
155. . 1937b. Polyploidy from twin seedlings. Cytologia, Fujii Jub.
Vol. 211-227.
156. . 1938. Notes on heteroploid twin plants from eleven genera.
157. . 1943. Characteristics of two haploid twins in Dactylis
glomerata. Hereditas 29: 134-140.
158. , and PRAKKEN, R. 1940. The mode of chromosome pairing in
Phleum twins with 63 chromosomes and its cytogenetic consequence.
159. NAKAJIMA, G. 1935. Occurrence of a haploid in Triticum turgidum. Jap.
Jour. Genet. 11: 246-247.
160. NAKAMURA, M. 1937. Cytogenetical studies in the genus Solanum. I.
Autopolyploidy of S. nigrum L. Cytologia, Fujii Jub. Vol: 57-68.
161. NAKAMURA, S. 1933. [Some haploid plants in rice.] Jap. Jour. Genet.
8: 223-227.
162. NAMIKAWA, S., and KAWAKAMI, J. 1934. On occurence of haploid,
triploid and tetraploid plants in twin seedlings of common wheat.
Proc. Imp. Acad. [Tokyo] 10: 668-671.
163. NANDI, H. K. 1936. The chromosome morphology, secondary association
and origin of cultivated rice. Jour. Genet. 33: 315-336.
164. NATARAJAN, A. T., and SWAMINATHAN, M. S. 1958. Haploidy induced
by radiations in wheat. Experientia 14: 336.
165. NETTANCOURT,DE D. and STOKES,G. W. 1960. Haploidy in tobacco. Jour.
Hered. 51: 102-104.

166. NEWCOMER, H. 1941. A colchicine-induced homozygous tomato obtained

through doubling clonal haploids. Proc. Amer. Soc. Hort. Sci. 38:
610-612.
167. NIEL-SON, E. L. 1946. Breeding behaviour and chromosome numbers in
progenies from twin and triplet plants of Poa pratensis. Bot. Gaz.
108: 26-40.
168. NISHIYAMA, I. 1940. [Studies on artificial polyploid plants. IV. Com-
parative studies on 1x to 4x plants in Capsicum annuum]. Bot. &
Zool. 8: 81-89.
169. NORDENSKIOLD, H. 1939. Studies of a haploid rye plant. Hereditas 25:
204-210.
170. . 1945. Cytogenetic studies in the genus Phleum. Acta Agr.
Suecana 1: 1-138.
171. OKAMOTO, M., and SEARS,E. R. 1956. Structural relationshipsbetween
non-homologous chromosomes. Wheat Inf. Serv. 3: 6.
172. OKURA, E. 1933. [A haploid plant in Portulaca grandiflora]. Jap. Jour.
Genet. 8: 251-260.
173. OLSSON, G., and HAGBERG, A. 1955. Investigations on a haploid rape.
174. PAT, R. A., and SWAMINATHAN, M. S. 1959. [Cytological behaviour of
a nullihaploid of bread wheat.] Naturwiss. 46: 584-585.
175. PELOQUIN, S. J., and HOUGAS, R. W. 1958. The fertility of two hap-
loids of Solanum tuberosum. Science 128: 1340-41.
176. , and . 1959. Decapitation and genetic markers as
related to haploidy in Solanum tuberosum. European Potato. Jour. 2:
176-183.
177. , and . 1960. Genetic variation among haploids of
the common potato. Amer. Potato. Jour. 37: 289-297.
178. PERSON, C. 1955. An analytical study of chromosome behaviour in a
wheat haploid. Canad. Jour. Bot. 33: 11-30.
179. PISSAREV, V. E., VINOGRADOVA, N. M., and PODDUBNAYA-ARNOLDI, B. A.
1945. [A haploid barley plant produced by remote hybridization].
Comp. Rend. (Doklady) Acad. Sci. U.R.S.S. 49: 372.
180. POGLIAGA, H. H. 1951. Un nuevo caso de polihaploidia en Nicotiana
tabacum. Rev. Argent. Agron. 18: 221-30.
181. POOLE, D. D., and HADLEY, H. H. 1954. Haploidy in castor beans. Jour.
Hered. 45: 285-88.
182. PRATASSENJA, G. D. 1939. Production of polyploid plants, haploid and
triploids in Prunus persica. Comp. Rend. Acad. Sci. (Doklady) U.R.S.S.
22: 348-351.
183. RAMANUJAM, S., and SRINIVASACHAR, D. 1943. Cytogenetic investigations
in the genus Brassica and the artificial synthesis of B. juncea. Indian
Jour. Genet. & Plant Breed. 3: 73-88.
184. RAMANUJAM, S. 1941. A haploid plant in Toria (Brassica campestris).
Proc. Indian Acad. Sci. B. 14: 25-34.
185. RAMIAH, H. 1934. Haploid plant in rice. Giorn. Risicolt 24: 143-146.
186. . 1935. Polyploidy in rice. Indian Jour. Agr. Sci. 5: 119-124.
187. , PARTASARATHI, N., and RAMANUJAM., S. 1933. Haploid plant
in rice. Oryza sativa. Current Sci. 1: 277-278.
188. RANDOLPH, L. F. 1932a. Some effects of high temperatures on poly-
ploidy and other variations in maize. Proc. Nat. Acad. Sci. [U.S.A.]
18: 222-229.

189. . 1932b. The chromosomes of haploid maize with special

reference to the double nature of the univalent chromosomes of the
early meiotic prophase. Science 75: 566-567.
190. RANDALL, T. E., and RICK, C. M. 1945. A cytogenetic study of polyem-
bryony in Asparagus officinalis L. Amer. Jour. Bot. 32: 560-569.
191. RAW, A. R. 1937. Genetical studies of wheat-haploids of Triticum vul-
gare. Jour. Dept. Agr. Victoria 35: 300-306.
192. RICHARDSON, M. M. 1935. Meiosis in Crepis. II. Failure of pairing in
Crepis capillaris L. Jour. Genet. 31: 119-143.
193. RIEGER, R. 1957. Inhomologen Paarung und Meioseablauf bei haploiden
Formen von Antirrhinum majus L. Chromosoma 9: 1-38.
194. RILEY, R. 1960. The diploidisation of polyploid wheat. Heredity 15:
407-429.
195. , and CHAPMAN, V. 1957. Haploids and polyhaploids in
Jegilops and Triticum. Heredity 11: 195-207.
196. , and 1958. Genetic control of the cytologically
diploid behaviour of hexaploid wheat. Nature [London] 182: 1244-1246.
197. , KIMBER, G., and CHAPMAN, V. 1961. The origin of the ge-
netic control of the diploid-like behaviour of polyploid wheat. Jour.
Hered. 52: 22-25.
198. Roux, J. B. 1958. L'haploidie chez le cotonnier. Cott. et. Fibr. Trop.
13: 289-92.
199. RUTTLE, M. L. 1928. Chromosome number and morphology in Nicotiana.
II. Diploidy and partial diploidy in the roots of tabacum haploids.
Univ. Cal., Publ. Bot. 11: 213-32.
200. S. M. 1937. [Cytology of twin-embryoed plants.] Bot. & Zool.
5:146.
201. SATINA, S., BLAKESLEE, A. F., and AVERY, A. G. 1937. Balanced and
unbalanced haploids in Datura. Jour. Hered. 28: 193-198.
202. SEARS, E. R. 1939. Cytogenetic studies with polyploid species of wheat.
I. Chromosomal abberations in the progeny of a haploid Triticum
vulgare. Genetics 24: 509-523.
203. - . 1948. The cytology and genetics of the wheats and their
relatives. Adv. Genet. 2: 239-270.
204. , and OKAMOTO, M. 1958. Intergenomic chromosome relation-
ships in hexaploid wheat. Proc. X Int. Congr. Genet. Vol. 2.
205. SKALINSKA, M. 1954. Meiosis in polyhaploid twin plant and a hexaploid
hybrid of Valeriana sambucifolia. Acta Soc. Bot. Poloniae 23: 359-374.
206. SKOVSTED, A. 1935. Some new interspecific hybrids in the genus Gossy-
pium L. Jour. Genet. 30: 447-463.
207. . 1939. Cytological studies in twin plants. Comp. Rend. Lab.
Carlsberg. Ser. Physiol. 22: 427-445.
208. SILoW, R. A. and STEPHENS, S. G. 1944. Twinning in cotton. Jour.
Hered. 35: 76-78.
209. SIMONET, M., and CHESNEAUX, M. T. 1954. Etude cytogenetique de deux
cereales haploides. Comp. Rend. Acad. Sci. [Paris] 238: 147-149.
210. , and FLECKINGER, J. 1937. Sur la presence de deux plantes
haploides chez Triticum spelta L. Ann. Epiphytes et Phylogenetique
[Paris] 3: 23-24.
211. SMITH, L. 1946. Haploidy in einkorn. Jour. Agr. Res. 73: 291-301.

212. SPRAGUE, G. F., RUSSELL, W. A., and PENNY, L. H. 1960. Mutations

affecting quantitative traits in the selfed progeny of doubled mono-
ploid maize stocks. Genetics 45: 855-866.
213. STANFORD, E. H., and CLEMENT, W. M. 1955. A haploid plant of Medi-
cago sativa. Abs. Ann. Meet. Amer. Soc. Agron.
214. , and . 1958. Cytology and crossing behaviour of
a haploid alfalfa plant. Agron. Jour. 50: 589-592.
215. STEBBINS, G. L. 1950. Variation and Evolution in Plants. Columbia
University Press, New York.
216. , and KODANI, M. 1944. Chromosomal variation in guayule
and mariola. Jour. Hered. 35: 163-172.
217. STOMPS, T. J. 1930. Uber Parthenogenesis infolge Fremdbefruchtung bei
Oenothera. Zeits. Ind. Ahstamm. u Vererbungslebre 54: 243-245.
218. STRAUB, J. 1937. Untersuchungen zur Physiologie der Meiosis. VII.
Die Abhaingigkeit der Chiasmabildung bei Vicia Faba und Campanula
persicifolia von ausseren Bedingungen. Zeits. Bot. 32: 225-68.
219. . 1941. Die Cytologie der haploiden Epilobein und die Phy-
logenie der Gattung. Biol. Zentralbl. 61: 573.
220. SWAMINATHAN, M. S., and HOWARD, H. W. 1953. The cytology and
genetics of the potato (Solanum tuberosum) and related species. Bib.
Genet. 16: 1-192.
221. , and SINGE, M. P. 1958. X-ray induced somatic haploidy in
watermelon. Cur. Sci. 27: 63-64.
222. TAKAHASHI, H. 1936. [Rice plants grown from the seeds of haploid
individuals]. Proc. Crop. Sci. Soc. Japan 7: 355-63.
223. TAKENAKA, Y., and TANAKA, M. 1955. Cytogenetic studies in the genus
Nicotiana. VIII. A haploid plant of Nicotiana tabacum. Ann. Rep.
Nat. Inst. Genet. Japan. 6: 61-62.
224. TERNOVSKY, M. F. 1936. Polyploids and haploids in Nicotiana in inter-
specific hybridization. Proc. Tobacco and Makharka Genet. Breed. Vol
2: 59-108.
225. THOMPSON, K. F. 1956. Production of haploid plants of marrowstem
kale. Nature [London] 178: 748.
226. TOMENTORP, G. 1939. Cytological studies on haploid Hordeum disti-
chum. Hereditas 25: 241-254.
227. TOOLE, M., and BAMFORD, R. 1945. The formation of diploid plants from
haploid peppers. Jour. Hered. 36: 67-70.
228. TRALAV, H. 1957. Uber ein haploide Form von Populus tremula. Bot.
Not. 110: 481-83.
229. U., N. 1932. On the reappearance of haploid in the Japanese morning
glory. Jap. Jour. Bot. 6: 225-243.
230. . 1935. Genome analysis in Brassica with special reference
to the experimental formation of B. napus and peculiar mode of fer-
tilisation. Jap. Jour. Bot., 7: 389-452.
231. VALENTINE, D. H. 1952. Studies in British primulas. III. Hybridization
between Primala elatior (L) Hill. and P. veris L. New Phytol. 50:
383-399.
232. VASILYEV, B. 1936. A haploid plant of durum wheat, Triticum durum.
Comp. Rend. Acad. Sci. U.R.S.S. 10: 243-244.
233. WEBBER, J. M. 1933. Cytological features of Nicotiana glutinosa hap-
loids. Jour. Agr. Res. 47: 845-867.

234. . 1938. Cytology of twin cotton plants. Jour. Agr. Res. 57:
155-160.
235. . 1940. Polyembryony. Bot. Rev. 6: 575-598.
236. WILSON, J. A., and Ross, W. M. 1961. Haploidy in twin seedlings of
winter wheat, Triticum aestivum. Crop Sci. 1: 82.
237. YAMAMOTO, Y. 1936. Ein haplo-diploides Zwillingspaar bei Triticum
vulgare. Bot. Mag. Tokyo 50: 573.
238. YAMASAKI, K. 1934. The haploid plant of common wheat, Triticum vul-
gare. Cytologia 5: 305-307.
239. . 1935. [Cytological studies on haploid wheat plants.] Jap.
Jour. Genet. 11: 314-315.
240. . 1936. Some observations on the microsporogenesis of the
haploid plant of Triticum 'vulgare. Jap. Jour. Bot. 8: 151-153.
241. YASUI, K. 1941. Diploid bud formation in a haploid Oryza with some
remarks on the behaviour of the nucleolus in mitosis. Cytologia 11:
515-525.
242. YEFEIKIN, A. K., and VASILYEV, B. I. 1936. [Artificial induction of hap-
loid durum wheats with X-rayed pollen.] Bull. AppI. Bot., Genet. &
Plant Breed. II. 9: 39-45.
243. ZUKOV,N. I. 1939. Nicotiana rustica L. haploids. Symposium on breed-
ing, genetics and seed production of tobacco and Nicotiana rustica.
All-union Mikojan Research Institute of the Tobacco and Makharka
Industry. Krasnodar 139: 224-35.
SUPPLEMENTARY CITATIONS
The following papers have come to the attention of the authors since
this review was originallly completed in August, 1961. Appropriate addi-
tions to the text and to Table I have been made.
244. ALEXANDER, D. E. 1954. Evidence for crossing-over between non-homol-

ogues during megasporogenesis of monoploids. Maize Genet. Coop.
News Letter 28: 52.
245. . 1956. Further evidence for crossing-over between non-ho-
mologous chromosomes during megasporogenesis of haploids. Maize
Genet. Coop. News Letter 30: 37-38.
246. BURK, L. G., and GERSTEL, D. U. 1961. Haploid Nicotiana deficient for
two chromosomes. Jour. Hered. 52: 203-206.
247. CAMPOS,F. F., and MORGAN, D. T. JR. 1960. Genetic control of haploidy
in Capsicum fjrutescens L. following crosses with untreated and X-
rayed pollen. Cytologia 25: 362-372.
248. COE, E. H., JR. 1959. A line of maize with high haploid frequency.
Amer. Nat. 93: 381-382.
249. DEWEY, D. R. 1961. Polyhaploids of crested wheatgrass. Crop. Sci. 1:
249-254.
250. . 1962. The genomic structure of intermediate wheatgrass.
Jour. Hered. 53: 282-290.
251. ENDRIZZI, J. E. 1962. The diploid-like cytological behaviour of tetra-
ploid cotton. Evolution 46: 325-329.
252. FISHER, H. E. 1956. Untersuchungen an Zwillingen von Beta 'vulgaris
L. Zuchter 26: 136-152.

253. HOUGAS,R. N., and PELOQUIN, S. J. 1959. Hybrids of Solanum tuberosum

haploids and the tuber-bearing Solanum species. Amer. Potato Jour.
36: 296. [Abs.]
254. Hu, C. 1959. A comparison between diploid lines obtained from a hap-
loid plant and diploid lines from the original variety. Karyological
studies of haploid rice plants. III. Jap. Jour. Breed. 9: 135-139.
255. -. 1960. Karyological studies in haploid rice plants. IV. Chro-
mosome morphology and intergenome pairing in haploid plants of
Oryza glaberrina Steud, as compared with 0. sativa. Cytologia 25:
437-449.
256. JACOB, K. T., and SEN, S. 1961. Haploidy in jute (Corchorus olitorius
L.). Nature [London] 192: 288-289.
257. JARAOWSKI,J. 1961. Haploid-diploid twin embryos in Meliotus. Genet.
Polon. 2: 129-137.
258. KIHARA, H., and TSUNEWAKI, K. 1962. Use of alien cytoplasm as a
new method of producing haploids. Jap. Jour. Genet. 37: 310-313.
259. KIMBER, G. 1961b. Cytogenetics of haploidy in Gossypium and Triticum
Ph.D. Thesis. Univ. Manchester.
260. . 1962. The effect of chromosome 5B at prophase. Wheat.
Inf. Serv. 14: 3-4.
261. , and RILEY, R. 1963. The relationship of the diploid progeni-
tors of hexaploid wheat. Canad. Jour. Genet. & Cytol. 5: 83-88.
262. MATSUBAYASHI, M. 1960. Studies on haploid plants of Solanum tubero-
sum. II. Meiotic chromosome pairing in the polyhaploid plants. Jap.
Jour. Breed. 10: 195-202.
263. MAZOTI, L. B., and MUALENBERG, G. E. 1958. Natural haploids in
maize. Rev. Argent. Agron. 25: 171-178.
264. MEYER, J. R., and Jus-rus, N. 1961. Properties of doubled haploids in
cotton. Crop. Sci. 1: 462-464.
265. MUNTZING, A., HRISHI, N. J., and TARKOWSKI, C. 1963. Reversion to
haploidy in strains of hexaploid and octoploid triticale. Hereditas 49:
78-90.
266. NEI, M. 1963. The efficiency of haploid methods of plant breeding. He-
redity 18: 95-100.
267. NISHIYAMA, I. 1961. Studies in Avena. VIII. A haploid plant of sand
oats (Avena strigosa). Jap. Jour. Genet. 36: 72-75.
268. OKAMOTO, M., and SEARS, E. R. 1962. Chromosomes involved in trans-
locations obtained from haploids of common wheat. Canad. Jour. Genet.
& Cytol. 4: 24-30.
269. PELOQUIN, S. J., and HOUGAS, R. W. 1959. Haploidy in Solanum tuberosum
and the sub-species Andigena. Amer. Potato. Jour. 36: 302. [Abs.]
270. , and . 1961. Hybrids between S. tuberosum hap-
loids and diploid Solanum species. Abs. Ann. Meet. Amer. Soc. Agron.,
St. Louis, Missouri. P. 54.
271. RILEY, R. 1963. Cytogenetics and plant breeding. Proc. 11th Int. Cong.
Genet. [In press]
272. ROTHACKER,D. and SCHAFER, G. 1961. Some investigations on haploid
plants of S. tuberosum. Ziuchter 31: 289-297.
273. SAITo, K. 1958. [On a haploid plant obtained from the induced auto-
tetraploid plant of the common California poppy]. Jap. Jour. Breed.
7: 152-156.

274. SASTRY,G. R. K., and SWAMINATHAM, M. S. 1960. Chromosome asso-

ciations in haploid Gossypiumbarbadense.Canad. Sci. 29: 398.
275. SINNOT,E. W., HOUGHTALING, H., and BLAKESLEE, A. F. 1934. The
comparative anatomy of extra chromosomaltypes in Datura stramon-
ium. CarnegieInst. Wash., Publ. 451.
276. VISHVESHWARA,S. 1960. Occurrenceof a haploid in Coffee arabica L.
cultivar "Kents".Indian Coffee24: 123-124.
277. WHITEHEAD, R. A., and CHAPMAN, G. P. 1962. Twinning and haploidy
in Cocosnucifera Linn. Nature [London] 195: 1228-1229.
CITATIONS ADDED IN PROOF

278. CHASE, S. S. 1963. Cytoplasmic replacement through androgenesis in
maize. Proc. XI Int. Cong. Genet, Vol. 1: 229.
279. CHEDDA, H. R., and HARLAN, J. R. 1962. Mode of chromosomeassocia-
tion in Bothriochloahybrids. Caryologia15: 461-476.
280. GOODSELL,S. F. 1961. Male sterility in corn by androgenesis.Crop Sci. 1:
227-228.
281. Ross, J. G., CHEN, C. H. and SIMANTEL, G. M. 1963. Colchicine-induced
somatic chromosomereductionin sorghum.Proc. XI Int. Cong. Genet.
Vol.1: 120.
282. SANDERS,N. E., and FRANZKE, C. F. 1962. Somaticreductionof tetraploid
sorghum to diploid mutants following colchicine treatment. Nature
[London] 196: 696-698.


Springer New York Botanical Garden Press

Uploaded by

Document Information

Original Description:

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Springer New York Botanical Garden Press

Uploaded by

Copyright:

Available Formats

Haploid Angiosperms

Author(s): Gordon Kimber and Ralph Riley

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

GORDON KIMBER AND RALPH RILEY

Introduction ....... .......................... 480

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

Gossypium hirsutum 26 T P properties of

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

Medicago sativa 16 S P good pairing,

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

Nicotiana tabacum 24 H P up to 14 chrom

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

Solanum tuberosum 24 S P meiotic analysi

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

Zea mays 10 S M non-homologous

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

Secale cereale 7 T M II's observed,

Triticum monococcum 7 SE M III in 3% of P

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

Triticum aestivum 20 E P 18.53I, 0.671I, 0

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

Haploidy was first reviewed by Gates and Goodwin (1930) when

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

More complex terminologyis requiredto categorizespecies with some

Aneuhaploid Nullisomic haploid

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

useful in that it clearly indicates the multiplicity ofL-singlesets. How;

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

misdivisionproductsof some chromosomes.These may be termed "mis-

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

in structure,present in double dose. Thus he postulated an aneuploid

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

monoploids, presumablybecause of the technical difficultiesinvolved.

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

describeda situation in autohexaploidPhleum pratense (2n=6x=42)

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

1947) which implies that there is some structuralsimilarity between

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

The only definite evidence on the relationshipsbetween the paired

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

considerablyreduced comparedwith that in the "raw"polyhaploid of

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

at meiosis from normal allopolyhaploids.The modificationof the pair-

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

A genetic isolation of homoeologues is suggested by the behaviour

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

Neither did the addition allopolyhaploids which had chromosome

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

were ln + 5.5, ln + 7.7 and In + 15.15. In a furtherdisomic mono-

SECONDARY PAIRING OF UNIVALENTS

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

pium barbadensein which chromosomesof the two genomes are of

No. of Observed Calculated

x 100 1.00 1.00

This content downloaded from 157.181.127.228 on Mon, 22 Sep 2014 21:55:12 PM

found that in autumn the alkaloid content of the polyhaploid leaves