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Science in Horticulture Series
General Editor: Professor L. Broadbent, University of Bath

Published in collaboration with the Royal Horticultural Society and the

Horticultural Education Association.

This series of texts has been designed for students on courses in

horticulture at the Higher National Certificate or Diploma level, but care
has been taken to ensure that they are neither too specialised for
lower-level courses, nor too superficial for university work.

All the contributors to the series have had experience in both the
horticultural industry and education. Consequently, the books have a
strong practical flavour which should reinforce their value as textbooks
and also make them of interest to a wide audience, including growers and
farmers, extension officers, research workers and workers in the agro-
chemical, marketing and allied industries, and the many gardeners who
are interested in the science behind their hobby.

The authors are all British but they have illustrated their books with
examples drawn from many countries. As a result the texts should be of
value to English-speaking students of horticulture throughout the world.
C. North

Formerly Head of the Plant Breeding Section

Scottish Horticultural Research Institute
lnvergowrie, Dundee, Scotland

© C. North 1979

All rights reserved. No part of this publication

may be reproduced or transmitted, in any form
or by any means, without permission

First published 7979 by

London and Basingstoke
Associated companies in Delhi Dublin
Hong Kong johannesburg Lagos Melbourne
New York Singapore and Tokyo

Set I.B.M. by

British Library Cataloguing in Publication Data

North, C
Plant breeding and genetics in horticulture.
- (Science in horticulture series).
1. Plant·breeding 2. Plant genetics
I. Title II. Series
635'.04'3 SB123

ISBN 978-0-333-23581-2 ISBN 978-1-349-03707-0 (eBook)

DOl I 0.1007/978-1-349-03707-0

This book is sold subject to the standard conditions

of the Net Book Agreement

Preface ix
1.1 Early progress in plant improvement
1.2 Sexual reproduction
1.3 Mendelism
1.4 Genes
1.5 Chromosomes
1.6 Mitosis
1.7 Meiosis
1.8 Crossing over and linkage
1.9 Homozygosity and heterozygosity
1.10 Segregation
1.11 The cytoplasm and inheritance
1.12 Variegation
1.13 Chimaeras
1.14 The chemical structure of genes
2.1 Polyploids
2.2 Techniques for inducing polyploids
2.3 Chromosome counting
2.4 Haploids
2.5 Aneuploids
2.6 Accessory chromosomes
3.1 Natural pollination
3.2 Flower form
3.3 Monoecious and dioecious plants
3.4 Hand pollination and emasculation
3.5 Use of insects
3.6 Male sterility
3.7 Selective gametocides
3.8 Double flowers
3.9 The plant breeder's glasshouse
3.10 Control of flowering
vi Contents


4.1 Seed-bearing plants
4.2 Pollen and sperm
4.3 Embryo sac and egg cell
4.4 Fertilisation
4.5 Embryo and endosperm development
4.6 Fruit set
4.7 Seeds
4.8 Incompatibility
4.9 Overcoming incompatibility barriers
4.10 Embryo culture
4.11 Protoplast fusion
4.12 Apomixis
5.1 Forecasting segregations
5.2 Dihybrid segregation and gene interaction
5.3 Lethal genes
5.4 Certation
5.5 Goodness of fit
5.6 Progeny size
5.7 Complex segregations
5.8 Inbreeding depression and hybrid vigour
5.9 Combining ability
5.10 Sex ratios·
6.1 Changes in chromosome and gene structure
6.2 Artificial induction of mutations
6.3 Mutations of ageing seed
7.1 Clones and seed propagated cultivars
7.2 Problems of breeding vegetatively propagated cultivars
7.3 Soft fruits
7.4 Tree fruits
7.5 Perennial vegetables
7.6 Perennial ornamentals
7.7 Verifying hybridity
8.1 Fertility and stability
8.2 Self and cross fertilisers
8.3 Self pollinating vegetable and salad crops
8.4 Cross pollinating vegetable crops
8.5 Seed propagated ornamentals
8.6 Seed production
Contents vii


9.1 Selection
9.2 Objective assessment and instrumentation
9.3 Independent trials
9.4 Registration and Plant Breeders' rights
9.5 Naming and release of cultivars
9.6 'Virus-free' stocks
9. 7 Rapid propagation techniques
9.8 Gene pools

Index 143

Without a working knowledge of genetics, modern plant breeding techniques

are largely unintelligible. One of the major problems in preparing a book
of this length, therefore, is to condense an explanation of genetics theory
into a relatively small space and yet to leave enough room for plant
breeding matters. This has been tackled here by clothing, as it were, the
bare bones of genetics with the flesh and skin of practical plant breeding.
Whenever possible horticultural examples have been chosen to illustrate
the theory-especially examples of those species of greatest interest for
cultivation in the northern hemisphere.
Like the other texts in this series, this book is intended for students
of horticulture studying for Higher National Diploma or Certificate, for
National Diploma in Horticulture and as an introduction for university
degree students. I have had in mind also the idea of producing a pocket
reference book of the kind I would like to have had myself when I first
became interested in plant breeding. I hope, therefore, that it will prove
of interest to a wider public, indeed anybody who is seriously interested
in the improvement of horticultural crops and garden plants.
To help in making the text read fluently, literature references have
been kept to a minimum. Those references quoted have been chosen
whenever possible from papers in easily available journals in English and
of direct interest to a reader wishing to follow up a topic.
Discussion with my colleagues Drs J. R. T. Hodgkin, H. j. Gooding,
D. L. Jennings and A. B. Wills have been of considerable help in formulat-
ing the contents of this book and Mr R. L. Knight of the National Seed
Development Organisation Ltd., has kindly advised on some matters
relating to the testing and introduction of new cultivars. I am grateful to
many workers who permitted the quotation of their results, especially to
Dr 0. Banga, Dr S. Blixt, Professor J. Sneep and the late Dr W. Heydecker.
Figure 1.1 was reproduced with assistance from the St Andrews University
Library of Rare Books. I am grateful to the Literary Executor of the late
Sir Ronald A. Fisher, FRS, to Dr Frank Yates, FRS and to the Longman
Group Ltd., London, for permission to include Table 5.1 which is based
on a table in their book Statistical Tables for Biological, Agricultural and
Medical Research.

I should like to record my thanks also to Professor L. Broadbent for

his helpful comment and encouragement, to Professor j. Sneep for kindly
reading through the manuscript, and to my wife who suffered in silence
when I wrote the text and read it through to ensure that it was intelligible
to a member of the public who is not a plant breeder or geneticist.

Newmi/1 of Knapp, lnchture, 7979 C.N.



Plant breeding, using the term in its broadest sense, is a very old human
activity. Ever since crops were first cultivated, human selection pressures,
as distinct from natural selection, have been exerted on plants to modify
them and their progenies to suit human needs and whims. The achieve-
ments are stupendous. Many of the cultivated forms of the important
crop plants, such as the cereals, potatoes and some of the pulse crops, are
the basis of present-day western civilisation. Without these cultivars agri-
culture would not be sufficiently productive to feed the present world
population and modern technology probably could not have been develop-
ed. It has been said, for example, that the swede turnip introduced to
Britain in the early 1700s provided the basis for the industrial revolution.
It caused an improvement in agriculture by providing winter feeding
stuffs and thus permitted the over-wintering of larger numbers of cattle
to produce meat to feed the increasing population. The swede is a hybrid
between two wild species and probably could not have survived and would
not have been developed into a useful agricultural crop without the
discerning eyes of plant selectors.
In spite of these achievements and the tangible evidence of the varieties
themselves, which do not crumble away like buildings, tools and books,
plant breeding is rarely given due prominence in the social history of the
human race. Our world would be a very different place without our
inheritance of farm, vegetable and ornamental plant cultivars.
Purists may argue that early achievements merely resulted from
selection of individual plants or groups of plants and that this does not
strictly constitute plant breeding. Nevertheless, selection is a major part of
horticultural plant breeding, even when the most sophisticated modern
techniques and knowledge are used. To be able to carry out effective
selection it is essential for the breeder to know and to have a 'feeling' for
the crop being worked. The person who knows plants may 'pick a winner'
without having a scientific background, but the scientist who has no feel-
ing for plants has little hope of becoming a successful plant breeder.
Genetics are not synonymous with plant breeding but a knowledge of
genetics is essential if a breeder is to achieve his or her full potential.
Great strides have been made merely by selecting 'good' plants. The
cabbage illustrated by Leonard Fuchs over four hundred years ago (Figure
1.1) was undoubtedly derived by selectors without a knowledge of the
2 Plant Breeding

Figure 1.1 Reproduction of a woodcut of a cabbage published by Palmaising, Basel in

1549 in a herbal by Leonharti Fuchsii. Reproduced by courtesy of St
Andrews University Library of Rare Books.

fundamentals of heredity, yet it seems to compare favourably with some

of the best modern cultivars of the Dutch Langendijker type. Mackintosh
Red apple was found as a chance seedling by John Mcintosh in 1796 and
dominated the Canadian apple industry. Similarly, Lloyd George raspberry
was found in a wood in 1922 by J. J. Kettle of Corfe Castle, Dorset and
for some decades was the most important British cultivar of that fruit. In
all these cases the selector knew how to 'spot a winner' though no con-
:;cious effort was made to induce one.
Selection depends on having some variation of the plant material
from which to select, for clearly without variation the breeder has no
opportunity to exercise his talents. For centuries, variation has occurred in
cultivated crops because farm and garden techniques, by bringing different
species together and altering the physical conditions under which they
were grown, have encouraged hybridisation and mutation. True plant
breeding started when a conscious effort was made to induce variation and
to try to direct it towards the plant types being sought.
The Mechanism of Inheritance 3


The most important source of variation for the plant breeder comes
through hybridisation but the knowledge that plants, like animals, can be
mated together is a relatively recent discovery. Rudolph jacob Camerarius
from Ti.ibingen in 1694 seems to have been the first to assert that plants,
like animals, are sexually differentiated. The first recorded deliberate
species hybrid is credited to the Englishman, Thomas Fairchild, who in
1718 crossed Dianthus barbatus with the carnation D. caryophyllus. The
hybrid he called the Sweet William was sterile and should not be confused
with the present-day Sweet William-a name reserved for the species
Dianthus barbatus itself which is, of course, fertile. joseph Gottleib
Kolreuter in 1766 published several papers describing hybridisation with
54 different species. He also noted that only relatively closely related
species will hybridise and that the progenies from such crosses are often
much larger and stronger than their parents, although they may be sterile
'vegetable mules'.
The knowledge that plants could be hybridised, and the enthusiasm
for collecting 'new' species from all parts of the world, engendered a great
deal of activity amongst horticultural plant breeders in the 1800s. This is
the period when many of our garden ornamentals such as the dahlia,
rose and rhododendron were developed. The breeders knew little of the
scientific basis of inheritance but they were astute plantsmen. There were
often small fortunes to be made and the few available accounts of this work
with its rivalries and sometimes its chicanery make fascinating reading.


Since ancient times there have been speculations on the 'nature of heredi-
tary substance' in animals, and even Charles Darwin (1809-1882) accepted
the view held by Hippocrates in 400 Be that the male and female sub-
stances consisted of body extracts which mixed together at mating and
were in some way able to influence the development of the new individual.
These views were revolutionised by the monk Gregor johann Mendel
(1822-1884) who lived at Brno, now in Czechoslovakia, and by carefully
controlled experiments, mainly with peas, showed that the 'hereditary
substance' was not a uniform body-extract but was instead composed of
many independent and constant hereditary units. For example, when tall
peas were crossed with dwarf peas and all the seed collected and sown,
all the resultant seedlings grew into tall plants. The result of the crossing
was not a family of medium-height plants as one would have expected
from a uniform mixture of body extracts. Furthermore, when the tall
plants were allowed to self pollinate and all the resultant seeds were sown,
798 of the seedlings gave rise to tall and 266 to dwarf plants (a ratio of
3:1 ). Thus the dwarf character had not been lost; it was carried over to
4 Plant Breeding

the next generation as a hereditary unit. Moreover, repeated experiments

showed that it was possible in crosses of this kind to predict the approxi-
mate numbers of tall and dwarf plants which would occur amongst the
Mendel's later work included experiments with Hieracium sp. (hawk-
weed) which, for reasons now well understood, did not seem to support
his hypothesis. He died a disillusioned man doubting the validity of some
of his scientific work, knowing that it had not been accepted by scientists
in his lifetime and thinking that his term as abbot at the monastery had
been an administrative failure. Nevertheless, his findings are one of the
greatest scientific achievements of all time. No attempt will be made here
to elaborate on the full meaning of his work but its significance is embraced
in what follows.

The full significance of Mendel's work was not appreciated by scientists
until about 1900 but, once it was accepted, it engendered a period of
intense research on heredity, leading to the discipline of genetics-a
surprisingly young branch of science which originated during the child-
hood of present-day old-age pensioners.
In 1909 Johannsen first used the term gene to describe a unit of
inheritance. As we have seen from one of Mendel's experiments, a particu-
lar gene controls the height to which pea plants will grow. It exists in two
forms known as alleles, one for tall and one for dwarf growth. In other
instances there may be more than two alleles, as for example the 50 or so
known alleles of the incompatability gene in Brassica oleracea (see Section
4.8.2). Frequently one allele is dominant over another which is then said
to be recessive, as the tall pea gene allele is dominart over its dwarf reces-
sive counterpart. In some cases there is incomplete dominance of one
allele over its counterpart. For example, when 'fern-leaf' kale, which is a
type having a deeply indented lamina, is crossed to a normal kale with
entire lamina, the hybrid has the fern-leaf character but this is less pro-
nounced than with the true fern-leaf type. The fern-leaf allele is said to
be incompletely dominant. In yet other cases alleles of a gene are not
dominant over one another and each exerts an approximately equal
influence in the hybrid, as when a red-flowered sweet pea is crossed with
white; the hybrid has pink flowers.
A gene may control more than one character of a plant, in which case
it is said to be pleiotropic. For example cultivars of some plants which
have reddish or purple leaves or stems often also have red flowers. A more
complex example of pleiotropism is found in Aquilegia vulgaris which has
a gene, an allele of which gives red flowers, a dark-coloured endosperm,
anthocyanin in leaves, prolongs the stalks and increases seed weight. The
effect of a gene is influenced by the environment. For example, some
forms of Primula obconica carrying genes for red flowers may produce
white flowers at high temperature and low light intensity and a form of
The Mechanism of Inheritance 5

Brussels sprout carrying genes for premature 'bolting' will not run to seed
prematurely at high temperatures. In many cases, especially when a
physiological reaction is concerned, it is more correct to think of a gene
as inducing proneness to a certain reaction rather than causing its fulfil-
Genes frequently interact with one another in their effect on the plant.
An allele of one gene may induce pink flower colour but it may require
the presence of an allele of a completely different gene to intensify the
colour to red. Quite often a character such as yield may be controlled by
many genes, all of which have a small additive or interactive effect and
inheritance then is said to be polygenic. One gene may mask the effect of
another, in which case it is said to be epistatic to the other hypostatic gene.
This situation should not be confused with allele dominance, although the
term dominance is sometimes loosely used by breeders to refer to this
situation. In certain cases, two genes in their dominant forms may be
required for the expression of a certain character, and these are called
complementary genes.
The nature of an individual growing plant is determined by its entire
genetic constitution, the genotype, whose expression is modified by the
environment to produce the phenotype. Two plants rooted from cuttings
of the same individual, then one grown in the mountains and the other in
a lowland valley, may differ considerably in height of growth and size of
leaves and flowers. They are both of the same genotype but the phenotypes
are different. One frequently hears a comment that a certain plant charac-
teristic is 'genetic' or that it is 'environmental'. Such a statement is never
the complete truth as both genetic constitution and the environment
influence the phenotype.
The development of every plant is controlled by thousands or tens of
thousands of genes which ensure the hereditary characteristics so that a
primrose, for example, produces another primrose from seed and not an
oak tree. We cannot see the genes, even with the aid of an electron micro-
scope, and only become aware of their presence when alternative alleles
occur. It is these different alleles that ensure variation.
Little is known about the way in which genes exert their influence on
the growing plant but it is generally accepted that they produce enzymes
which control cell behaviour. The whole process is, of course, a complex
of interacting factors. The meristematic cells which give rise to the petals
have the same chromosome complement as those of the root tip yet by
virtue of their relationship to the surrounding tissues they develop very


Individual genes have not been recognised visually but it is known that
they occur on chromosomes, which can be observed fairly easily in most
species with the aid of a light microscope. Chromosomes are visible,
usually as rod-shaped bodies, within the nucleus at certain stages of cell
6 Plant Breeding

division. The word chromosome is derived from Greek, meaning a colour-

ed body, and indicates that it stains darker than most other cell contents
when the material is treated with certain chemical dyes. Chromosomes are
somewhat analogous to computer punched cards. They carry instructions
for the development and function of the cell, and the genes and their
alleles might be likened to the punched holes in the cards. With few
exceptions, each cell of a plant carries an identical set of chromosomes in
its nucleus and the number of chromosomes per cell is usually constant for
a given species. Examples of chromosome numbers of some of the most
important horticultural plants are given in Table 2. 7 but the significance of
this information will be more fully appreciated after reading Chapter 2.
As each nucleus of a plant divides, so do its chromosomes, by a process
known as mitosis, and each new cell thus carries an identical chromosome
complement. However, a special type of cell division known as meiosis or
reduction division occurs when the sex cells are produced, and the chromo-
some number is then halved. If this did not occur the chromosome number
would be doubled every time sexual fusion occurred. The somatic cells
(body cells as distinct from the sex cells) of the plant in fact each contain
one set of chromosomes from the mother and one set from the father and
retain these throughout the life of the plant.
The process of cell division is rather complex but the student should
make every effort to grasp the meaning of it in principle since this is
essential to understanding the mechanism of inheritance. It is similar in
nearly all living organisms whether they are animals or plants, except for
the very smallest forms of bacteria and viruses.

Chromosomes are not recognisable by microscopic examination, except
during the process of cell division when four phases can be distinguished
(Figure 7.2).

1.6.1. Prophase
The chromosomes first become recognisable as a coiled mass within the
nucleus. At this stage they are long and thin and each is made up of two
strands, chromatids, pressed close together, the chromosomes having
already divided longitudinally.

1.6.2 Metaphase
The chromosomes contract to become shorter and fatter, and take up a
position arrayed at the 'equator' of the cell. Each chromosome is attached
by a fibre, all of which are joined at the poles to form the spindle. The
point on the chromosome where it joins the spindle is a non-staining
constriction called the centromere. This site remains constant for the
chromosome and this feature can serve as an aid in identification of a

Figure 1.2 Mitosis -a diagrammatic representation in which chromosomes inherited

from the two parents are shown in black and white. 1, Cell with resting
nucleus. 2, Prophase when chromosomes become visible as l9ng paired
threads. 3, Metaphase, ch,·omosomes shorter and broader, divided longi-
tudinally but the halves not yet separated. 4, Late metaphase to early
anaphase with divided chromosome portions separating on spindle (s).
5, Anaphase, two new identical sets of chromosomes moving apart.
6 , Telophase, cell wall laid down between two new cells and chromosomes
becoming invisible.
8 Plant Breeding

chromosome or as an indication of its possible evolution from earlier types.

Sometimes secondary constrictions occur and these are usually associated
with the chromosome's regular point of attachment to a nucleolus.

1.6.3 Anaphase
The two chromatids split apart and one half of each chromosome moves
along the spindle to form two equal groups of chromosomes clustered at the
poles of the cell. The spindle then disappears.

1.6.4 Telophase
A new cell wall is formed between the two daughter nuclei and the
chromosomes uncoil and again become indistinguishable within the

This type of division only occurs with special cells designated to produce
the male or female sex cells known as gametes. The process of meiosis
always consists of two successive divisions so th:t one nucleus starting on
the pathway results in the formation of a tetrad of four nuclei, each having
only half the number of chromosomes which were present in the mother
nucleus (Figure 1.3). To appreciate the significance of meiosis for gene
inheritance, it has to be remembered that at the onset of the process the
nucleus contains two similar sets of chromosomes, one inherited from its
father and one from its mother. Each pair usually carries the same genes
arranged in the same order but these genes are frequently of different
allelic forms on the two chromosome partners of the pair.

1. 7.1 Prophase of first division

As in prophase of mitosis the chromosomes first become recognisable, but
in meiosis they come together in like pairs, divide lengthwise, and twist
around each other, break in places and then join together (Figure 1.4).
This produces four chromatids which have exchanged portions with each
other by a process known as crossing over, and all differ in their gene allele
constitution from the original chromosomes.

1. 7.2 Metaphase of first division

A spindle is formed as in mitosis. The rearranged chromosomes, each
consisting of a pair of chromatids, come together at the equator of the cell.

1. 7.3 Anaphase of first division

One chromosome from each pair moves along the spindle to the poles of
the nucleus.
The Mechanism of Inheritance 9

2 3

' I

~0 \

Figure 1.3 Meiosis--a diagrammatic representation in which chromosomes inherited

from the two parents are shown in black and white. 1, Metaphase of first
division, chromosomes divided but not separated. 2, Metaphase pairing of
chromosomes. 3, Metaphase crossing over of chromatids. 4, Early ana·
phase of first division with pairs of chromatids separating after crossing
over and exchanging parts. 5, Separation of paired chromatids into two
cells. 6, Metaphase to early anaphase of second division with chromatids
separating. 7, Tetrad of cells with half the chromosome members of the
original cells.
10 Plant Breeding

A a A a A a

8 b b

c c c c c

A a

8 b

c c

Figure 1.4 Diagram illustrating crossing-over of chromatids with one chiasma (above)
and two chiasmata (below).
The Mechanism of Inheritance 11

1. 7.4 Telophase of first division

As in mitosis two new cells are reconstituted, but with half the chromo-
some complement of the original, and the chromosomes again become
visibly indistinguishable. After a short resting period the second division of
meiosis occurs.

1.7.5 Second division of meiosis

This proceeds in a similar manner to mitosis and each pair of chromatids
separates to form two chromosomes but, as a result of the first division of
meiosis, these chromosomes differ from one another in gene alleles and the
two new cells formed are not generally identical as in mitosis.


As we have seen, the feature of meiosis of overriding importance for

inheritance is the exchange of chromosome parts to give a redistribution of
gene alleles. This process is called crossing over and the sites at which
chromosomes break and join are chiasmata (singular--chiasma).
Characters for which the genes are situated on different chromosomes
are inherited quite independently of one another, but this is not so for
genes located on the same chromosome. If crossing over did not occur
then all genes on a chromosome would be inherited together. Since cross-
ing over takes place, this usually does not happen, but it follows that the
closer together two genes are on a chromosome the less chance there is of
chiasmata occurring between them and the greater chance that the charac-
ters they control are inherited together. This closeness of genes is referred
to as linkage and genes on a chromosome are said to constitute a linkage
group, there being as many linkage groups for a genotype as there are
chromosome pairs. However, when only limited studies have been made on
the inheritance of a species it may not be possible to distinguish between
genes which are widely spaced on a chromosome and those on different
chromosomes. For practical purposes there may be more linkage groups
than chromosome pairs. By measuring the degrees of linkage through
growing large numbers of progenies from planned crosses and noting the
frequency of association of characters, it is possible to plot the relative
positions of genes on the chromosomes and to produce a chromosome
map for the material. This is arduous and time-consuming work and has
not been done in any detail for most horticultural crops. However, much
is now known about gene position in such crops as wheat, maize, tomato
and pea. A chromosome map for peas (Figure 1.5) shows genes represent-
ed as le-tters or abbreviations, which is the usual convention utilising
initial capital letters for the dominant and small letters for the recessive
Relatively little is known about gene position for most horticultural
crops and the term linkage group may then mean group of genes which
2 3 4 5 6 7 ...
a chi 5 uni 1a1
m 4 cp wla wsp
10 14 6
vi I 7 sbm lm
ram 25 m leu 4
Chi 3 12 24 p I 24
13 14
chi22 ch~~ 6 Ia z
wa ~ rl 9P 25 mine
li 9 ah 10 12
was II
If II 19 eri II xa I
10 ar 10 miu 5
miv mier dem 4 na 9 15
vacl 5 9 sc cr cana
dli 17 10 sip 25 cat 8 pa
y lr mp 6 to chi21 pi
pra 15 10
6 Cly COY 7
rup 8 16 It II r
8 er Bra PY 6 5
sru •• 7 Tra coh [bl] II
5 14 lob 6 com 5 II
d 5 Ia
8 all 12 rub
Pur Opo 6
5 suD cal 16
Him ch3 F 5 28 6
4 chi 23 cotr
25 6 8 each
.. 10 inc
2 Ina
9 aba
foe .112 10 sui 7
1om _ II bl
wb ld Ha
26 14 pt
18 chi 16 ce
Fw 10
pia 9 Is
Kpa 7 Br
pall 10 13 I 23
lov pat
beg och 6 8
ins 14 8 can
at In ca 4 II
pal I NP I 30
par 6 lru 5
vim 4
sal 5 yp 17 chi 1 pro alb
19 lum
.g 4 Lap I b
7 chi
den op chi9 ·~~ilv 5
II 7
red 14 rb 22 le
6 Ro 6
sir chi6 prae
gl 6
r30 rog
10 •
gri+ 20 Q:J
er un
•a early flowering ~
Figure 1.5 Chromosome map for pea {Pisum sotivum and P curvense) showing distribution ICj

of gene loci on the seven somatic chromosomes (after Blixt, 1972).

The Mechanism of Inheritance 13

seem to be inherited independently on the knowledge available. As we

have seen, in these cases there may be more recognised linkage groups
than chromosomes.


Each cell nucleus has one set of chromosomes inherited from the mother
and one from the father plant. In most fertile plants these chromosome
pairs usually carry the same genes but as a rule the alleles of many of
these genes differ between the two chromosomes of a pair. If the alleles
on each pair were all the same, the plant would be said to be homozygous
and it would always produce sex cells with the same alleles, in spite of
crossing over, and self-fertilisation of the plants would lead to all plants
with identical genotypes-the plant would be 'true breeding'. In practice
this very rarely occurs except in a few cases where plants have been
induced to develop from pollen or pollen-mother cells (Section 2.4).
However, plants like cultivars of the French bean, the garden pea and the
lettuce, which are normally self-fertilising and have been selected for
uniformity of the progenies, are for practical purposes homozygous and
true breeding.
Most plants have different alleles in their chromosome pairs, do not
breed true and are said to be heterozygous. Plant breeders talk rather
unprecisely about the 'degree of homozygosity', meaning capacity to
breed apparently uniform progenies.
The terms homozygous and heterozygous can also be used when one
is discussing one or a few specified genes instead of the entire gene content.
Varieties, families or individual plants are then said to be homozygous or
heterozygous for these particular genes or for certain characters such as
flower colour although they may be heterozygous for others.


It is possible with a knowledge of the genes governing particular characters

to work out the segregation from deliberate planned crosses of the number
of individuals carrying those characters. The easiest way of doing this is to
construct chequerboard diagrams, and a number of simple examples will
be considered here. More complex segregations are discussed in Chapter 5.

1.1 0.1 Monohybrid segregation

Monohybrid segregation is segregation for a single gene only, such as Wh
which governs flower colour in Brassica oleracea. Plants which carry the
dominant Wh allele have white flowers and only those homozygous for
the recessive wh allele have yellow flowers. (B. oleracea is the species
which embraces Brussels sprout, cabbage, cauliflower, kale and kohl rabi.
Most forms have yellow flowers but some cauliflower cultivars segregate
white flowers and Portuguese cabbage breeds true for this character.)
14 Plant Breeding

WhWh wh wh

Wh wh

S3 j \
Wh wh

Figure 1.6 Monohybrid segregatio n from white-f lowered ( Wh Wh) X yellow·flo wered
(whwh) Brassi ca o/eracea.
The Mechanism of Inheritance 15

Homozygous WhWh and whwh individuals produce only one kind of

sex cell, Wh or wh respectively, but heterozygous individuals Whwh
produce both kinds.
In Figure 1.6 we see that in the so-called F 1 generation, that is when
the two types are crossed, all the offspring are white-flowered hetero·
zygotes. In the F 2 generation, derived from self pollinating the F 1 , there
are 3:1 white:yellow-flowered plants. When the heterozygote is back
crossed to the dominant parent all the progeny are white-flowered and
when it is back crossed to the recessive parent there are 1:1 white: yellow·
flowered plants.

1.1 0.2 Dihybrid segregation

More complex cases involving two genes can also be worked out in this
way. As we have seen (Section 1.4), the fern-leaf character Fn is dominant
to normal leaf fn in Brassica oleracea. It is inherited independently to
white/yellow flower colour. Thus plants carrying both the dominant alleles
Wh and Fn have white flowers and fern leaf whereas only the homozygotes
whfnwhfn have yellow flowers and normal leaves. From Figure 1.7 it is
evident that when homozygous WhFnWhFn plants are crossed with homo-
zygous whfnwhfn plants all the F 1 progeny WhFnwhfn are white-flowered
and fern-leaved. From the chequerboard diagram of the F2 we see that
there are:
9 white fern leaf
3 white normal leaf
3 yellow fern leaf
1 yellow normal leaf
A slight complication is that fern-leaf is incompletely dominant and it
is possible to recognise those genotypes which are heterozygous for fern
leaf by their phenotype. Thus of the nine white fern-leaf types three are
fully fern-leaved and six are incompletely fern-leafed and of the three
yellow fern-leaf type one is fully fern-leafed and two are incompletely
Similar and more complex cases with three or more genes can be
worked out in this way and the reader will find it interesting to experiment
with other examples. Some typical segregation ratios, as they are called,
are discussed in more detail in Chapter 5.
The ratios represent the proportionate numbers of individuals in a
progeny. However, the proportions will never be exact unless infinite
numbers of individual plants are produced. It is unlikely, for example, that
exactly three of the plants would be yellow-flowered fern-leaf if only 16
plants were grown, but the segregations will approach these proportions
unless other factors intervene. It is possible, for example, that plants
carrying one type of allele combination may be weaker growing than the
16 Plant Breeding

p X

Wh wh Fn fn


// \~ Wh fn wh Fn wh fn


Figure 1.7 Dihybrid segregation from white-flowered, fern-leaved (WhWhFnFn) X

yellow-flowered, normal leaf (whwhfnfn) Brassica oleracea.
The Mechanism of Inheritance 17

others so that fewer of its seedlings survive, but such cases will be discussed
in more detail later. Chequer-board calculations of segregations involving
more than one gene only apply when these genes are inherited indepen-
dently; linkage will affect the proportion, though it is possible to obtain an
estimate of the numbers if the degree of linkage is known.


We have noted that inheritance and variation are controlled primarily by

the genes located on chromosomes which are contained within the nucleus.
However, the cytoplasm and its contents which surround the nucleus may
influence the expression of the genes. Notable examples of horticultural
interest are found in some cases of male sterility such as in forms of the
onion and maize in which certain gene allele combinations prevent the
development of fertile pollen but only in conjunction with certain cyto-
plasms. Knowledge of this phenomenon can be used as an aid to produce
hybrid cultivars, as we shall discuss in more detail in Chapter 8.
The male sperm cell carries very little cytoplasm with it when it
fertilises the female egg cell, which is itself endowed with a fairly generous
supply of cytoplasm from the mother plant. When two different homo-
zygous plants are crossed the progeny therefore will have the same geno-
type but the cytoplasm will differ according to which plant was used as
the mother. Such differences in offspring from reciprocal crosses are often
taken as evidence of cytoplasmic inheritance and in most cases this is the
correct explanation. However, similar effects due to other causes are
known. Apomixis (Section 4.12), in which seed is not the product of
hybridisation, results in progenies resembling the mother parent and is
fairly frequent with some genera. An unusual and interesting situation
which occurs in some species of the Canina section of Rosa is described
by Blackburn and Harrison (1921). The somatic chromosome number is
35 but by a complex process the male and female gametes carry different
numbers of chromosomes, namely 7 and 28 respectively. Thus the mother
parent contributes four times as many chromosomes as the father and the
offspring arc therefore largely maternal-like.
The effects of the cytoplasm are especially pronounced in certain
wide species crosses. Some chromosome combinations may be unable to
co-exist with the cytoplasm of one of the parents so that the hybrids will
thrive only if the cross is made one way. French bean x runner bean
(Phaseolus vulgaris x P. coccineus) hybrids are much easier to achieve and
survive better when french bean is used as the seed parent; similarly
radish x cabbage (Raphanobrassica spp.) crosses are most likely to succeed
when radish is used as the mother parent. However, gene/cytoplasm inter-
action is not the only cause of failure to achieve hybrids in one direction.
There may be several other causes, such as a blockage of the pathway to
fertilisation or abnormalities in the development of the seed after fertilisa-
tion (Chapter 4).
18 Plant Breeding

The striping and segmentation of different colours in leaves, and to a lesser
extent petals and fruits, is an important feature of some ornamental plants.
In some instances this is caused by virus infection, as in the variegated-
leaved Abutilon spp. and the 'breaking' of tulip, wallflower and viola
flowers. In these cases the character is maintained by vegetative propaga-
tion and can be transmitted to new cultivars by infection with the appro-
priate virus. However, expression of the symptom is gene-controlled and
it may be impossible to get 'broken' forms of some tulips even with virus
infection. Also it is questionable if the products of new forms of these
types of 'varieties' should be encouraged as they may act as 'disease'
reservoirs for other plants. For example, tulip breaking virus causes a
serious disease of lilies, as well as tulips; and wallflower breaking is caused
by cabbage black ringspot virus which damages brassicas such as cabbage
and cauliflower.
Most cultivars with variegated leaves owe their aesthetic attraction to
abnormalities inplastids which occur in the cytoplasm. The most important
plastids are the green chloroplasts, which are essential for the fixation of
energy from sunlight. Other white or pale-coloured plastids occur in the
leaves of some plants but they are only able to exist in plants which also
have some green chloroplasts. Plastids increase in number by division and
this is usually independent of nuclear division. When a cell divides the
plastids are distributed at random between the two new cells. Thus some
new cells may have no chloroplasts and when they divide they give rise to
pale-coloured segments in the leaves. Sometimes entire growing points
develop from cells which have received only green plastids in the random
distribution at cell division. These growing points are then all green and
shoots arising from them have to be cut out to prevent 'reversion' as all-
green shoots are usually more vigorous than those which are variegated.
june yellows is a 'disease' of strawberry in which the degeneration of
the green leaves through a variegated to a chlorotic form occurs over a period
of months or years {Wills, 1962). It is presumably partly gene-controlled
since it can be transmitted by the pollen as well as by the mother parent.
Cultivars like 'Howard 17', 'Blakemore', 'Auchincruive Climax', and
others known to develop this trouble, should be avoided by breeders.

Many popular variegated leaf forms have a more complex constitution,
especially those with darker leaf centres or margins and other fairly
regular patterns. Most plants have two, and sometimes three, well-defined
different layers of cells in the growing point which give rise to different
layers of leaf cells. Sometimes one or more of these layers may be devoid
of chloroplasts and/or other plastids to give green, white or transparent
tissues. These peric/inal chimaeras occur in Hosta, Hydrangea, Ligustrum,
Pelargonium, Spiraea, Vinca and many other genera.
The Mechanism of Inheritance 19

A more bizarre periclinal chimaera is the graft-hybrid Laburnum

adami obtained by M. Adam in France in 1825. It is a chimaera in which
layers of two different species occur and has separate layers of the yellow-
flowered Laburnum and the purple-flowered shrub Cytisus purpureus. It
grows as a tree carrying three types of branches; those of Laburnum and of
Cytisus and some which are intermediate in character between the two
species. As the egg cells arise from the inner core of Laburnum, all seeds
give rise to normal Laburnum plants. Crataegomespilus spp. are similar
graft hybrids with an inner core of hawthorn surrounded by medlar tissue.
Although genes may influence leaf patterning, nearly all forms of
variegation relate to the cytoplasm. Interesting new types are more likely to
be obtained using a variegated form as the mother rather than the father

False variegation
In some cases leaf patterning is not associated with plastid differences.
The 'variegated' leaves in Coleus spp. result from distribution of antho-
cyanin pigments and the patterns on leaves of Lamium species, Zebrina
pendula and Pulmonaria saccharatum are associated with small air blisters
just below the epidermis. These features are solely gene controlled, and do
not depend on differences in plastids.


The clarification by Watson and Crick (1953) of the chemistry of the basic
genetic material is a milestone which may prove to be as important to
plant breeders as the discovery of sex in plants and of Mendel's work on
Deoxyribonucleic acid (DNA) has been identified as the geneti;; base.
It is a threadlike molecule composed of two spirals of nucleotides linked
together in chains. The number of combinations of nucleotide arrange-
ments is virtually infinite and the combinations represent the genetic code
units. The genetic structure of some bacteria has been altered by introduc-
ing DNA from other organisms-a process known as transformation.
Similar effects have been claimed for higher plants (johnson and Grierson,
1976), but the average practical breeder of horticultural plants is unlikely
to be able to profit by such techniques at present.
Knowing that genes essentially consist of forms of DNA, doubt now
arises as to whether many classical genes are indeed indivisible units. In
marw cases they may be closely situated groups of chemically different
units. It is therefore usual nowadays to refer to th~ sites on chromosomes
as loci (singular-locus) or, when it is known they comprise several such
chemical units, as cistrons. However, for practical purposes they usually
can be called and can be envisaged as genes used in the classical sense as
20 Plant Breeding

BLACKBURN, K. B. and HARRISON, T. W. H. (1921). The status of the British
rose forms as determined by their cytological behaviour, Ann. Bot., 3S, 159-
BLIXT, S. (1972). Mutation genetics in Pisum, Agri. Hortique Genetica, 30, 1-293.
JOHNSON, C. B. and GRIERSON, D. (1976). The uptake and expression of DNA by
plants, in Commentaries in Plant Science (ed. Smith, H.), Pergamon, Oxford,
286 pp
WATSON, ). D. and CRICK, F. H. C. (1953). A structure for deoxyribose nucleic
acid, Nature, Lond., 171,737-738
WILLS, A. B. (1962). Genetical aspects of strawberry june yellows, Heredity, 11,

KIRK, T. C. and TILNEY·BASSETT, A. E. (1966). The Plastids, W. H. Freeman,
London and San Francisco, 608 pp
SWANSON, C. P. (1957). Cytology and Cytogenetics, Prentice·Hall, Englewood
Cliffs, 596 pp


We have seen that the nucleus of each cell of the plant, with the exception
of the sex cells, carries two sets of chromosomes; one inherited from the
father and one from the mother. The basic chromosome number of the sex
cells in which each chromosome is different and carries a different set of
genes is the haploid number. The cells of the body of the plant (somatic
cells) usually carry double this number and are diploid. However, plants
with more than two sets of the haploid number are not uncommon and are
said to be triploid (3x ), tetraploid (4x ), pentop/aid (5x ), hexaploid (6x ),
heptaploid (7x·), octaploid (8x), decaploid (1 Ox), dodecap/oid (12x ), or
referred to collectively as polyp/aids. For example (Table 2. 7) several
cultivars of apple and pear are triploid, leek tetraploid, plum hexaploid,
strawberry octaploid and black mulberry highly polyploid. Sometimes the
somatic cells may not contain an exact multiple of the haploid number so
then there are a few extra or fewer chromosomes; these are aneuploids.
Triploids and tetraploids and to a lesser extent higher polyploids
frequently have larger, broader and thicker-textured petals and leaves and
more intensely coloured flowers than diploids. It is not surprising, there-
fore, that these forms have been selected by plantsmen even when the
ploidy was not known. Many of our vegetatively propagated ornamentals,
especially those with large flowers which tend to win prizes at shows, such
as chrysanthemum, daffodil, dahlia, gladiolus, hyacinth, rose and tulip are
polyploids. However, polyploids of some genera do not have these attri-
butes and are not then of the same interest to horticulturalist$. Tetraploids
of some species are often less vigorous and later flowering than correspond-
diploid forms.
There are two basic types of polyploids. Autopolyploids are those in
which each set of chromosomes carries the same or nearly the same genes,
which is the usual situation in hyacinth, leek, etc. Allopolyploids are those
in which the sets of chromosomes are not all the same and may even have
different numbers. For example some, but not all, cultivars of the so-called
Poetaz narcissus such as 'Cheerfulness' and 'Golden Dawn' are triploids
with a somatic chromosome number of 24, made up of one haploid set of
10 from Narcissus tazetta and two haploid sets of 7 from N. poeticus.
Primula x kewensis is a tetraploid with a somatic number of 28 and has
two sets of chromosomes from P. floribunda and two from P. verticillata.
22 Plant Breeding

Table 2.1

Chromosome numbers of some cultivated fruits and vegetables (after Darlington and
Wylie, 1961)
(The figures in parenthesis are somatic numbers of the most frequently grown culti-
vars. x is the basic haploid number for the genus.)

Almond (15, diploid) Prunus X= 8
Apple (34, 51 diploid, triploid) Malus X= 17
Apricot (16, diploid) Prunus X= 8
Blackberry (mainly 28, tetraploid) Rubus X= 7
Black currant (16, diploid) Ribes X= 8
Blueberry, highbush (48, tetraploid) Vaccinium X= 12
Cherry, Morello (32, tetraploid) Prunus X= 8
Cherry, sweet (16, diploid) Prunus X= 8
Currant, red (16, diploid) Ribes X= 8
Fig, (26, diploid) Ficus X= J3
Gooseberry (16, diploid) Ribes X= 8
Grape vine (38, 57, 76, diploid, triploid, tetraploid) Vitis x=19,20
Mulberry, black (over 300, polyploid) Morus X= 14
Mulberry, white (28, diploid) Morus X= 14
Nectarine (16, diploid) Prunus X= 8
Peach (16,diploid) Prunus X= 8
Pear (34, 51, diploid, triploid) Pyrus X= 17
Plum (48, hexaploid) Prunus X= 8
Raspberry (14, diploid) Rubus X= 7
Strawberry (56, octaploid) Fragaria X= 7

Asparagus (20, diploid) Asparagus X= 10
Beetroot (18, diploid) Beta X= 9
Broad bean (12, diploid) Vicia x= 5,6,7
Brussels sprout (18, diploid) Brassica x= 8,9,10,11
Cabbage (18, diploid) Brassica x= 8,9,10,11
Carrot (18, diploid) Daucus x= 8,9,10,11
Cauliflower (18, diploid) Brassica x= 8,9,10,11
Celery (22, diploid) Apium X= 11
Chervil (18, diploid) Anthriscus x= 7,8,9
Chicory (18, diploid) Chicorium X= 9
Cucumber (14, diploid) Cucumis X= 7, 12
Endive (18, diploid) Chicorium X= 9
Fennel (22, diploid) Foeniculum X= 11
French bean (22, diploid) Phaseolus x =11, rarely 12
Garlic (16, diploid)A//ium x= 7,8,9
Globe artichoke (34, diploid) Cynara X= 17
jerusalem artichoke ( 1 02, hexaploid) Helianthus X= 17
Leek (32, tetraploid) Allium x= 7,8,9
Lettuce (18, diploid) Lactuca x= 8,9,17
Marrow (40, tetraploid) Cucurbita X= 10, 12
Melon (24, diploid) Cucumis X= 7,12
Onion (16, 32, diploid, tetraploid) Allium x= 7,8,9
Parsley (22, diploid) Petroselina X= 9,11
Parnsip (22, diploid) Pastinaca X= 11
Chromosome Number 23

Pea (14, diploid) Pisum X= 7

Pepper (24, diploid) Capsicum X= 12
Potato (48, tetraploid) Solanum x =mainly 12
Radish (18, diploid) Raphanus X= 9
Rhubarb (22, 44, diploid, tetraploid) Rheum X= 11
Runner bean (22, diploid) Phaseo/us x = 4, rarely 12
Seakale (30, diploid) Crambe X= 15
Spinach (12, diploid) Spinacia X= 6
Squash (24, 40, 48, diploid, tetraploid) Cucurbita X= 10,12
Swede (38, amphidiploid) Brassica x= 8,9,10,11
Tomato (usually 24, diploid) Lycopersicum X= 12
Turnip (20, diploid) Brassica x= 8,9,10,11
Watercress (32, 48, tetraploid, hexaploid) Nasturtium X= 8
Water melon (22, diploid) Citrullus X= 11

The sets of chromosomes from each species in P. x kewensis have the same
number but the chromosomes do not carry the same gene complement so
it is an allotetraploid.
Autopolyploids are frequently less fertile than diploids so that most
crops raised from seed are diploids (Table 2.1). The poor fertility is due to
difficulties which arise in chromosome pairing at meiosis. When the auto-
polyploids are of odd chromosome numbers such as triploids, pentaploids
and heptaploids, it is clear that pairing of all chromosome sets cannot
occur. However, although fertility is considerably reduced, some fertile
gametes may be formed and may produce for example a few haploid or
diploid egg and pollen cells. The autotriploid Narcissus tazetta cultivar
'Soleil d'Or', for example, sometimes produces seeds. With even numbered
autopolyploids pairing is also difficult because at meiosis each chromo-
some will be attracted to more than one like partner, confusion arises, and
the situation may not be resolved before the chromosomes part to the
poles. However, chromosome pairing is itself to some extent gene control-
led, and it is possible in many cases to select for improved fertility in
autopolyploids. For example, the cultivated raspberry is diploid but
autotetraploid forms can be induced. These are usually rather infertile, as
can be seen by the irregularly developed fruits, although they usually do
produce some seeds. Two or three generations of inbreeding and selecting
for uniformity of fruit development produces fertile tetraploids. One other
feature of autopolyploidy is that it tends to break down the natural
incompatibility system when it is gametophytic but not if it is sporophy-
tic (see Section 4.8).
Allopolyploids behave differently. When a wide cross is made between
two diploid parents the hybrid is frequently sterile, because pairing can-
not take place between the very different sets of chromosomes. If, however,
the somatic chromosome number of the hybrid is doubled then pairing
24 Plant Breeding

often readily takes place because there is no confusion over chromosome

partners at meiosis and in breeding they behave more or less as diploids.
Fertile chromosome-doubled forms of wide crosses are often referred to as
amphidip/oids. They are more or less constant breeding and progenies
show little if any variation because the chromosomes from the original
wide parents rarely pair and little crossing-over occurs. Sometimes it is
advantageous to the breeder to double the chromosome number of the
two original parents before making the wide cross. The hybrid then is
likely to be fertile and the like chromosome pairs may differ in gene allele
structure so that through crossing-over some variation may occur with
progenies from such a polyploid.


Colchicine, an alkaloid obtained from the autumn crocus (Colchicum

autumnafe) is by far the most important chemical used by breeders to
induce the doubling of chromosome numbers. Its application was first
demonstrated by A. F. Blakeslee and some other American investigators in
1937. It prevents spindle formation at mitosis and 'blocks' the movement
of the chromosomes. The chromosomes divide but no new cell wall is
formed so the cell gains an extra set of chromosomes (Figure 2. 1).
Colchicine, which is expensive to buy and a dangerous poison, must
be treated with respect. It is usually used as a 0.05-0.2 per cent aqueous
solution which is best used fresh or may be kept for a few weeks in a
refrigerator but not frozen. It should be made up by dissolving the powder
in a few drops of industrial alcohol and the water added slowly; with rapid
addition of water the colchicine may come out of solution. The optimal
strength of solution and time required for treatment vary with the species
and cultivar. A treatment which causes some damage to the plant without
serious debilitation usually gives the optimal chance of chromosome
Imbibed seed or seedlings may be immersed in a colchicine solution
but the usual way of treating seedlings of dicotyledons is to apply a drop
of colchicine to the growing point between the expanded cotyledons once
or twice a day for several days. It may be necessary to add a wetting agent
if the plant surfaces are waxy or hairy. Established plants or parts of them
also can be treated, but it is essential to apply the colchicine to a meristem
or a site where callus tissue and adventitious shoots may arise. Axillary
buds on stems, cut surfaces of stems and root portions of plants propagat-
ed by root cuttings are sometimes suitable. Lily bulb scales which normally
produce adventitious buds on the surface where they have been broken
from the bulb plate can readily be treated by soaking for about 8 hours in
0.1 per cent solution and yield a high proportion of bulbils with doubled
chromosome numbers. Some other bulbous species can be treated in a
similar way. Until recently Narcissus species were thought to be difficult

( A'\"1\
X~ I VJ . VI
\ W!}!)
'-___-) \______

I ./.
,x~, ("~ (~

Figure 2. 1 Diagra m illustrating thee ffect of colchicine on mitOSIS.

26 Plant Breeding

subjects but by cutting the bulbs into small portions, each with two pieces
of bulb scale and a portion of the base plate and applying the solution to
the latter between the scale portions, chromosome doubled forms may be
fairly readily obtained (North, 1976).
Not all the treated seedlings or plant portions will produce chromo-
some doubled shoots; the material will have to be screened. It may be
possible to obtain an indication of doubling by increased size and width of
leaves or particularly by the size of stomata and leaf epidermal cells.
Portions of the epidermis can be carefully torn off using a pair of forceps,
and that of the shoots from the treated material compared with untreated
shoots using a microscope. Chromosome doubled stomata will generally
show an increase of 10-30 per cent in linear size (Figure 2.2). For some
material, comparisons of stomata size may be made from imprints obtain-
ed by painting nail varnish thinly on to the leaf, then stripping it off when

A B c
Figure 2.2 Relative sizes of stomata and epidermal cells in Nicotiana sy!vestris X
tomentosiformis. A, diploid; B, tetraploid; C, octaploid. (After Greenleaf,

dry and examining under a microscope. This is done quickly with some
material and can be used to sample plants directly in the field and obtain a
more or less permanent record, but it is not always practicable with hairy
leaves. Pollen grains of polyploids are usually larger than those of corres-
ponding diploids but one then has to wait until the plants flower to be
able to make comparisons with known diploid pollen. The only sure way
of ascertaining whether chromosome doubling has occurred is to make
chromosome counts, as described later.
Chromosome doubling may sometimes be induced by frequent cut-
ting of stems and by heat shocks, but these treatments are far less reliable
than colchicine. One other method which has met with limited success
involves the use of nitrous oxide under pressure. It has been used to
produce seed with tetraploid rather than diploid embryos from crosses
between two diploid tulip parents, and was first used to produce tetra-
plaids in Crepis phalaris. It functions by preventing the development of a
Chromosome Number 27

cell wall after the first nuclear division of the zygote. Not only is nitrous
oxide useful for tulip, which is difficult to double with colchicine, but it
may have applications for direct production of fertile amphidiploids from
wide crosses of other species. Flowering plants at a specified time after
pollination are put into a pressure chamber and treated with nitrous oxide
at 50 atmospheres for 7 hours letting the gas in and out slowly (Zeilinga
and Schouten, 1968}.
Triploids can sometimes be obtained by crossing tetraploids with
diploids. This is not always successful, probably because of endosperm
imbalance, and to ensure the best chance of obtaining triploids the cross
should be made both ways. As a rule it is more likely to succeed if the
tetraploid plant is used as the female parent.
Polyploids may arise naturally through the formation of egg and
pollen cells with an unreduced chromosome number. Several triploid
varieties probably have arisen in this way rather than from crosses between
diploids and tetraploids, and they in turn may have produced unreduced
gametes which mated with other diploid gametes and gave rise to tetra-
ploid offspring.


Root tip squashes are the most suitable means for observing mitosis and
for counting chromosomes of most plants. The best material is from plants
growing vigorously in pots and usually taken at a specific time of the day,
say 8-9 a.m., depending on the species. The plants are turned upside down
and the pot removed to expose the ball of soil. Root tips 2-4 mm long are
broken off with a pair of forceps, taking care to include as little soil and
grit as possible, and transferred immediately to a saturated aqueous
solution of para-dichlorobenzene. After one and a half hours, but not
longer than three hours, at room temperature the root tips are transferred
to fixative made of 1 part glacial acetic acid and 3 parts absolute alcohol
(ethanol}. They are left in the fixative for 3-24 hours at room temperature,
but may be kept longer in it if stored in a refrigerator. If no refrigerator is
available, they should be transferred to 70 per cent ethanol and stored in a
tightly closed small glass tube.
To soften the material for squashing, the tips are transferred to 10 per
cent hydrochloric acid in shallow glass dishes and kept in an oven at 60°C
for 10 minutes or longer depending on the material. If no oven is available
they may be treated for 1-2 hours at room temperature. After a quick
rinse in distilled water they are ready for staining.
Aceta-orcein is a useful stain for most root tips. It is prepared by dis-
solving 1 gramme of orcein in 45 ml hot glacial acetic acid; this is cooled,
made up to 100 ml with distilled water, shaken well and filtered. Washed
tips are transferred directly to stain in shallow glass dishes for a half to
three hours. Overstaining should be avoided. To produce a squash, a root
28 Plant Breeding

tip is carefully lifted by the cut end with a pair of forceps and transferred
to a microscope slide. With a sharpened needle the end 1-2 mm tip is cut
off and a drop of 45 per cent acetic acid or stain placed over the tip. The
slide is heated over a spirit lamp for no more than two seconds and a clean
cover slip applied. The root tip will spread slightly with the weight of the
cover slip and is then gently tapped; the end of a pencil with an india-
rubber attached is useful for this operation. A filter paper is then placed
over the slide which is pressed gently with the finger taking care not to
move the cover slip sideways. Individual workers develop their own
techniques for squashing to separate the chromosomes without applying so
much pressure that they are fractured. It is a relatively simple technique
which forms the basis of successful practical cytology and needs practising
if one is to become proficient. The squash is examined first with a 10 x
objective and it may be necessary later to use a 100 x oil-immersion
objective to make accurate counts. The onion is a useful plant for practis-
ing on as it has fairly large chromosomes and roots can easily be made
available at any time of the year, whereas some species such as cabbage
and raspberry Rave very small chromosomes.
Chromosome counts may also be made at meiosis by anther squashes
using a technique similar to that described above, although the para-
dichlorobenzene treatment is unnecessary. It is much more difficult to
obtain anther material than roots at the right stage, and very young anthers
have to be taken. This is especially difficult with some bulbous plants
where meiosis may occur in flower buds within the bulb and before shoot
elongation has started. However, anther squashes are especially useful for
observing pairing of chromosomes at meiosis, in addition to making counts.
Regularity of pairing indicates fertility of the gametes and may give useful
information on similarities and differences of chromosomes from different
species in wide species crosses.


The sex cells of diploid plants carry the basic haploid number of chromo-
somes. In many cases it would be of considerable interest to the breeder to
obtain haploid plants and to double the chromosome number to produce
diploids, which then would be completely homozygous and true breeding.
There are several ways by which this has been achieved for a few species,
but it cannot be claimed that there is a ready, useful method which can be
applied as a routine for the majority of plants.
The most promising way is to induce pollen or pollen mother cells to
grow directly into haploid plants. This has been achieved so far with 17
genera and 23 species, mostly Solanaceae, and primarily for academic
interest rather than as a tool for plant breeders. As a rule intact anthers are
used and the precise stage of development at which they should be taken is
critical and varies according to the species. Whereas plants may develop
Chromosome Number 29

directly from the microspores or pollen, in some cases callus develops and
has to be transferred to another medium with a lower auxin content to
induce shoot growth. In some material the somatic tissue of the anther
wall or the end of the filament proliferates and tends to dominate the
material and to overcome this attempts have been made to culture free
microspores as with tobacco (Nitsch, 1974). Medium composition may be
critical, especially in relation to the balance of plant growth substances.
Anther culture has exciting possibilities but so far precise techniques
of use to a plant breeder have not been formulated, with the possible
exception of that for the potato (Dunwell and Sunderland, 1973) and
asparagus (Pelletier eta!., 1972). It would be especially helpful if haploid
plants could be raised from Brassica oleracea, which is often difficult to
obtain in a homozygous condition, but although anther culture has been
attempted with this species the results have not been encouraging.
Another way to obtain haploids is simply to look for them amongst
seedlings from diploid plants. Sometimes seeds are produced with two or
more embryos instead of the usual single one, and occasionally one of the
supernumeraries is a haploid. A notable example of this is asparagus, in
which a mean frequency of 0.23 haploids was found in 1,000 seeds (Marks,
1973). Haploid seedlings have also been found in Brassica oleracea-type
kales. One feature of haploids in seeds is that the percentage yield of
haploid embryos is gene controlled and depends very largely on the female
parent and on the source of seed. A large number of seed! ings have to be
screened to find the haploids, and an interesting technique is proposed for
cucumber (Aalders, 1958) where haploid seeds are lighter and float on
water whereas those of diploids sink.
Some plants which very rarely, if ever, produce haploid embryos can
be induced to do so by pollinating with another species with which fertil-
isation cannot occur, or by using dead or X-ray irradiated pollen. A
notable example of the use of foreign pollen is with barley, which gives a
relatively high number of haploid embryos when pollinated with the wild
diploid Hordeum bulbosum. These embryos need to be cultured in vitro
for them to survive and grow into plants but the technique has neverthe-
less been sufficiently productive for it to be recommended as a routine
in barley breeding. The haploids, whether produced in this way or as
naturally occurring twins, are nearly always sterile and have to be con-
verted into autodiploids (also called di-haploids or double haploids) by
colchicine treatment before they can be useful in breeding.
Of course, wide pollination often gives rise to seedlings which are
apomicts of a similar ploidy to the seed parent. If they are double haploids
they can be of great value to breeders but this is rarely the case. Brassica
oleracea pollinated with B. napus rarely hybridises but sometimes gives
germinant seeds which were thought at one time to be of doubled haploid
constitution. It is now known that they are not homozygous and that they
are of little use to breeders.
30 Plant Breeding


Aneuploids are plants that have chromosome numbers which are not an
exact multiple of the haploid number-they have one or more chromo-
somes more, or one or more fewer, than the normal. Aneuploids, especially
those lacking a chromosome, are less common in diploids than polyploids.
This is probably because the loss or duplication of a chromosome, which
in the balanced diploid is represented only twice, has more serious conse-
quences than when it is represented four times, as in an autotetraploid.
Trisomes are aneuploids with 2n + 1 chromosomes, monosomes 2n --- 1
are very rare as complete plants but monosomic tissue sometimes survives
in conjunction with diploid tissue in a chimcera, and nul/isomes 2n - 2
(that is, with one complete set of chromosomes missing) occur in the
godetia (Clarkia amoena). In some plants of high polyploidy such as the
laurel {Prunus laurocerasus) (176 chromosomes) and Kentucky bluegrass
{Poa pratensis) (36-123 chromosomes) aneuploidy is common. Although
aneuploids are not uncommon, their occurrence is often of little signifi-
cance to breeders of horticultural species. However, for the breeder of
cereal crops they are of great interest. By using aneuploids in hybridisation,
it is possible to transfer one or more extra chromosomes from one lot of
material to the next. Further, by making use of a gene which allows
indiscriminate pairing of chromosomes at meiosis, it is possible virtually to
transfer single genes from one genotype to another. It may be possible to
use such techniques with horticultural crops when we understand more
about their genetics.


The additional chromosomes of aneuploids are duplicates of the normal

somatic members but so-called 8 chromosomes are extras with a different
gene make-up. They may vary in number in the gametes and in different
parts of the plant and, as a rule, their presence is negligible or deleterious.
They occur in a wide range of species and are fairly common in some
monocots such as the leek, Lilium and Trillium spp.

AALDERS, L. E. (1958). Monoploidy in cucumbers, f. Hered., 49, 41-44
DARLINGTON, C. D. and WYLIE, C. D. (1961). Chromosome Atlas of Flowering
Plants (2nd Impression), University Press, Aberdeen, 519 pp
DUNWELL, ]. M. and SUNDERLAND, N. (1973). Anther culture of Solanum
tuberosum L., Euphytica, 22, 317-323
GREENLEAF, W. H. (1938}. Induction of polyploidy in Nicotiana by hetero-auxin
treatment,/. Hered., 29,451-464
MARKS, G. E. (1973}. Selecting asparagus plants as sources of haploids, Euphytica,
NITSCH, C. (1974}. La culture de pollen isole sur milieu synthetique C.r. hebd.,
Seanc. Acad. Sci., Paris, 278,1031-1034
Chromosome Number 31

NORTH, C. (1976). Artificial chromosome doubling in Narcissus and its implication

for breeding N. tazetta hybrids, Acta hortic., 63, 161-163
PELLETIER, G., RAQUIN, C. and SIMON, G. (1972). La culture in vitro d'antheres
d'asperge (Asparagus officina/is). C.r. hebd. Seanc. A cad. Sci., Paris, 274,848-851
ZEILINGA, A. E. and SCHOUTEN, H. P. (1968). Polyploidy in garden tulips. II: The
production of tetraploids, Euphytica, 17, 303-310

DARLINGTON, C. D. and LaCOUR, L. F. (1969). The Handling of Chromosomes
(5th Edition), Allen and Unwin, London, 272 pp
DERMEN, H. (1940). Colchicine polyploidy and technique, Bot. Rev., 6, 599-636
HASKELL, G. and WILLS, A. B. (1968). Primer of Chromosome Practice, Oliver and
Boyd, Edinburgh, 180 pp
KIMBER, G. and RILEY, R. (1963). Haploid angiosperms, Bot. Rev., 29,480-531
WEBBER, J. M. (1940). Polyembryony, Bot. Rev., 6, 575-598

By virtue of their colour and form, flowers are a constant source of

aesthetic pleasure and inspiration and they exert a very important, though
not always obvious, influence on the design of man-made objects. Biologi-
cally they are organs which make mating possible for static individuals
which, unlike most higher animals, are unable to move about in search of a


The male part, the pollen, is transferred from one plant to another by the
aid' of the wind, in a few cases by water, and most frequently by animals,
especially insects. Visitors are attracted to flowers by colour, scent and
sometimes by what might be called trickery--for instance by the shiny
nectary-like protuberances of Parnassia spp or the insect-like flowers of
Ophrys spp. To the insect visitor flowers offer nectar and, what is often
equally attractive to bees, pollen as a protein source. Although insects are
the most common pollinators, birds pollinate Aloe, Antholyza, Fuchsia,
Hibiscus, Kniphophia and Strelitzia spp in their natural habitats and slugs
may pollinate some Compositae such as Chrysanthemum !eucanthemum.
Honey bees and bumble bees are the most important pollinators of
cultivated plants in Europe, although other insects, especially flies, play a
significant role in pollination of many crops and are the major pollinators
of Umbelliferae such as carrot and parsnip. Butterflies play a minor part
and they tend to visit flowers with coloration resembling their own. Some
plants, such as Nicotiana species and the butterfly orchid P!atanthera
chlorantha, by virtue of their pervading scent, white flowers and long
nectar tubes, are adapted to pollination by night-flying moths. An especially
interesting adaptation occurs in Yucca spp which are pollinated by the
larvae of the moth Pronuba yuccasella. The adult collects pollen in a sticky
ball, transfers it to the stigma of another flower and pierces a hole in the
ovary of that flower into which it lays a batch of eggs. The larvae feed
within the developing ovary of the moth-fertilised flower but destroy only
about 20 per cent of the seeds; the others develop normally.
Wasps are the main pollinators of the snowberry (Symphoricarpus spp)
and the figworts (Scrophu!aria spp). Another group of Hymenoptera, the
ichneuman-flies, pollinate the twayblade orchid Listera ovata presumably
because the lip bears a marking which resembles a larva of a type frequently
Flower Form and Pollination 33

parasitised by these insects. Perhaps the most specialised association is

between the 8/astophaga wasp and the fig. Wild figs bear three kinds of
closed sac-like inflorescences or 'fruits': male, female and a specialised
sterile type known as the caprifig solely to nurture the larvae of the wasp.
Adult female wasps fly to all three kinds but the fertile types are unsuit-
able to support larvae. Most cultivated figs lack the caprifig type and when
fertilisation is necessary for fruit development as in the Smyrna fig used
for drying it is customary to tie some caprifigs in the plantation to provide
wasps to effect pollination. This is unnecessary for fruit development of
fresh fig type cultivars which produce edible 'fruits' without fertilisation.
Wind is the major agent of pollination in some Chenopodiaceae includ-
ing beetroot and spinach and in many catkin bearing trees such as hazel,
filbert and poplar but insects may also effect pollination of these species.
Some species, including such troublesome weeds as Oxalis corniculata
and Lamium amplexicaule, at times produce flowers which are self
pollinated and fertilised in the bud stage and never open fully. Several
species produce cleistogamous flowers with reduced petals which are
specially modified for self fertilisation without opening of the buds, in
addition to normal petalous flowers suited to insect pollination. This
group includes such well known species as the violet (Viola odorata), wood
sorrel (Oxalis acetosella) and balsam (Impatiens biflora).
Many species produce flowers which open and are exposed to insects
but which are largely self pollinated and self fertilised without outside
aids. They include members of the Compositae and Leguminosae such as
lettuce, chicory, French bean and peas. However, the flowers are not
protected from insect pollination and cross fertilisation also can occur.


Basically the flower consists of the perianth, androecium and gynaecium.

The perianth protects the other parts of the flower and often contains
brightly coloured components to attract pollinators. It may be composed
of sepals, petals or both and in some species it is impossible to distinguish
between these, and the perianth segments are then called tepals as in tulip
and lily. The androecium and gynaecium comprise the male parts {the
stamens) and the female parts or carpels {which collectively comprise the
pistil) respectively.
It is not possible here to elaborate in detail on flower form, which is
very diverse. The breeder will soon learn his way around the flowers of the
species being worked. However, as a word of warning to the uninitiated,
flowers of some genera such as Penstemon have staminodes {modified
sterile stamens) which look superficially like pistils. Also several species
produce stigmas which remain shut and unreceptive during part of the life
of the flower. As a rule both stamens and pistil of a flower are functional
at the same time but in some plants the stamens are ripe before the pistil
34 Plant Breeding

and vice versa and the flowers are said to pe protondrous or protogynous
respectively. Protandry occurs with Campanulaceae, Caryophyllaceae,
Compositae, Geraniaceae, Labiatae, Rosaceae and Umbelliferae, notably
in the genera Aqui/egio, Componulo, Doucus (carrot), Delphinium, Loctuco
(lettuce) and Scobiosa. Protogyny is much less common but occurs in
Colchium, Hel/eborus and Magnolia.


Plants of over 90 per cent of all species are always hermaphrodite and
function sexually as both male and female. Usually all flowers on such
plants have functional stamens and pistils but some species, such as sweet
corn and many members of Cucurbitaceae including cucumber, melon and
marrow, bear flowers which are separately male or female on the same
plant and the plants are then said to be monoecious. Occasionally a plant
may carry in addition to these types of flowers those which are hermaph-
rodite and also modified sterile flowers which are usually large, colourful
and conspicuous to attract insects. Aesculus spp have hermaphrodite and
male flowers, A triplex hermaphrodite and female flowers; some Viburnum,
Hydrangea and Centaurea spp have hermaphrodite and sterile flowers.
A few species have male and female flowers on separate plants. They
are then said to be dioecious from the Greek implying that the sexes
occupy separate houses. This group includes asparagus, spinach and species
of poplar and willow. Even more complex situations arise with some
species having not only plants which are separately male and female but
others which are also hermaphrodite. This occurs with certain forms of
spinach and is also found in Dryas, Caltha, Thymus and in the strawberry
where some old cultivars like 'Tardive de Leopold' produce only female
flowers whereas most are hermaphrodite.


To obtain deliberately planned matings it is often necessary to hand-

pollinate flowers. Before this is done the flowers of the seed parent usually
have to be emasculated by carefully removing the anthers or entire stamens
with a pair of forceps or scissors in order to prevent self pollination. This
must be done before the anthers have dehisced, either before the flowe~
buds have opened or shortly afterwards. Also the forceps must be dipped
in industrial alcohol and allowed to dry when different pollen genotypes
are used so that the risk of effecting unplanned matings is reduced to a
minimum. In some species notably of the Leguminosae and Compositae
the filaments are joined to form a tube around the style and it requires
considerable skill and practice to emasculate the flower without damaging
the female parts and rendering it unsuitable for crossing. Emasculation is
especially difficult with Phaseolus spp (French and runner beans) where
Flower Form and Pollination 35

Figure 3.1 Pair of scissors adapted for emasculation of Brassica oleracea flowers .
(After Weiring, 1958.)

the long style surrounded by the stamen sheath is coiled like a spring.
Various tools have been devised to aid emasculation, notably a pair of
scissors with a notch in the blade (Figure 3. 7) used by some workers to
emasculate Brassica flower buds. It removes part of the perianth and
stamens but leaves the pistil untouched. However, most workers prefer to
use a pair of forceps.
It may also be difficult to emasculate very small flowers like those of
lettuce or at least to do this without leaving some pollen grains. However,
in this genus the flowers are protrandrous and a useful technique is to
wash the small capitulae, each of which has 10-20 flowers, with a stream
of water shortly after they have opened. Sometimes attempts are made to
emasculate by hand first. The water removes or destroys most of the pollen
and the flowers can then be pollinated about one hour later when dry, by
which time the stigmas are usually receptive (Figure 3.2). A similar techni-
que may be useful with some other Compositae. A small portable electric
vibrator is used to aid self pollination in some plants such as tomato and
After pollination, the flowers usually must be protected from un-
wanted pollen by bagging, except in rare circumstances when the plants
are grown in special glasshouses. In the open, hand pollinated flowers
usually must be protected by bags which do not collapse on them. Those
made of muslin or fine nylon net on a wire framework and tied to a cane
support are most useful for insect-pollinated plants, although small insects
like thrips can sometimes find their way in and may cause unwanted
pollination. Stiff waterproof-type cellophane bags or those made of water-
proof paper with a cellophane window are also useful in the open. Poly-
thene bags are not often used because they are impermeable to moisture,
so that condensation occurs and they tend to collapse on the flowers.
Indoors, cellophane bags are useful, provided the plants are not watered
overhead. With some plants that have large flowers, such as Lilium, a small


Figure 3.2 Development of anthesis in lettuce florets. Water treatment to kill pollen should be applied at stage 3 when style
is extended but stigma is not yet receptive. a, Anthers; s, stigma; p, pollen; r, receptive surface of stigma. (After
Thompson, 1938.)
Flower Form and Pollination 37

piece of metal foil wrapped around the stigma after pollination prevents
contamination of flowers indoors or outside and bagging is then unneces-
sary. Especial care is needed with wind pollinated crops for the pollen is
very fine, and, of course, easily blown about. Beet pollen has been collect-
ed by aircraft at heights of up to 300m. In this case it is helpful to use a
pollen parent that carries a genetic marker which means that it is homo-
zygous for an easily recognisable character not found in the seed parent.
All hybrids can then be recognised in the offspring.
If possible, anthers should be collected before they have shed their
pollen and be ripened in open containers in a warm, dry, draught-free room.
Sometimes pollen may be collected by sucking it from open flowers into
a small glass container. This is not effective with some species adapted to
insect pollination many of which have pollen which tends to stick together
in clumps. Another method which has been used with fruit trees and
grapes is to collect flowers with unopened anthers and allow them to dry
for a short time on a sieve. The anthers are then rubbed through the sieve
on to a sheet of glass and in 12-24 hours when the pollen is shed it is
scraped with the anthers off the glass, using a razor blade, on to a finer
sieve and separated. A small electrically operated vibrator is useful for
some species, the pollen being shaken into a container. Stored pollen
(Section 4.2) may also be used.
Pollen is usually applied to the stigma with a fine 'camel-hair' paint
brush which is dipped in industrial alcohol and dried before using batches
of pollen of different genotype. However, it is sometimes more convenient
to transfer pollen by daubing an anther on to the stigma. Airborne pollen
such as that of beet may be injected by means of a coarse syringe into the
bag protecting the flowers of the seed parent.
Crosses should be labelled so that the writing does not fade or wash
off and the labels are not blown away or removed by birds. Valuable work
can be lost if adequate precautions are not taken with this simple operation.


It may be difficult or impossible to make crosses by hand if the flowers are

very small, as with the carrot and celery. Fortunately these species, like
many Umbelliferae, have flowers which are strongly protandrous. Flowers
on an umbel open more or less in synchronisation and at first only function
as males. After a few days the stamens of the upright-facing flowers drop
down and wither and the stigmas become receptive so that the flowers
then function solely as females. By bagging an umbel on a plant in the
female stage together with one in the male stage and introducing insect
pollinators to the bag, the desired mating may be effected. The pollen
donor may be a single cut umbel with the stalk standing in a tube of water;
it is not necessary to have awhole plant.
House flies have been used for pollinating Umbelliferae in this way
38 Plant Breeding

but blow flies are easier to procure in quantity. Larvae (jones and
Emsweller, 1934} can be obtained by exposing pieces of meat outside
under a roof to keep off the rain, although it is generally more convenient
to buy larvae sold as bait for fishermen. The larvae soon pupate and the
pupae are sieved out and stored just above freezing in a refrigerator.
Shortly before they are required they are taken into a warm room in a
container and generally hatch within a few days. The containers are then
put back into the refrigerator where the newly hatched adults are rendered
torpid with the cold and can be tipped out and quickly made into batches
for pollinating before they become active again. Flies can conveniently be
used to pollinate many species, even those like Brassica spp which are
normally bee pollinated, because they are reasonably active in dull weather
when hive bees tend to remain in the hive. If flies are used to pollinate
small batches of plants in a glasshouse it is desirable to confine them in a
cage with a ceiling just above the top of the highest flowers otherwise they
tend to fly upwards and away from the plants.
Hive bees are frequently used as pollinators, although it is not generally
convenient to use them in bags as described above for flies. Colonies of
hive bees can be used in small isolation rooms with a batch of plants to be
pollinated (Kraai, 1954). They should be provided with sugar water to
prevent them from starving when there are insufficient flowers. Many bees
may die trying to escape during the first few days but as a new 'brood'
hatches they tend to settle down. The hive may safely be transferred to
another compartmeflt when the bees have returned during the evening; the
bees clean themselves of pollen overnight and the risk of pollen contamina-
tion with the next batch of plants is negligible.
Certain kinds of bumble bees have been used (Kraai, 1954) and males
and nematode-diseased queens are preferred to healthy workers because
they make no attempt to return to a hive. Bumble bees have to be caught
individually and are freed from pollen by shaking in a bottle of water
which causes the pollen to burst by osmosis. They work in dull weather
and at lower temperatures than hive bees and flies and they visit flowers
deliberately rather than effect pollination as flies do simply by indiscrimin-
ate movement. However, some bumble bees 'rob' flowers like Antirrhinum,
puncturing a hole near the base of the corolla to obtain nectar rather than
entering the normal way and pollinating the flowers.


Occasionally species which are normally hermaphrodite produce some
plants which have no functional pollen and are therefore male sterile (MS).
This condition is gene controlled but in many cases it is only expressed in
certain cytoplasms. Genera of horticultural interest in which such cyto-
plasmic male sterility has been found include Allium (onion and leek},
Aqui/egia, Beta (beetroot}, Capsicum (sweet pepper}, Daucus (carrot},
Helianthus (sunflower}, Lycopersicum (tomato), Petunia and Raphanus
Flower Form and Pollination 39

(radish). In several other plants, genetic male sterility which is not modi-
fied by the cytoplasm is known; examples are Brassica oleracea (Brussels
sprout, cabbage), Foeniculum vulgare (fennel), Lactuca (lettuce) and
Vicia faba (field bean). Sometimes more than one gene governs the con-
dition as in carrot.
The type and degree of male sterility also vary. In MS onions and
one form of MS carrot the stamens are reduced to rudimentary swellings.
In MS Brassica and the so-called 'brown anther' form of MS carrot, anthers
develop but they lack fertile pollen which generally aborts at the tetrad
stage (see Section 1. 7). An interesting type of sterility occurs in one form
of MS tomato when normal fertile pollen is produced but the anthers
never open to release it-a condition known as functional male sterility.
As we shall discuss in Chapter 8, male sterility can be useful to the
breeder to control crossing of parent lines in the production of hybrid
cultivars. Cytoplasmic sterility is the most useful condition because it is
possible to produce from seed families which give all male-sterile plants.
Simple gene controlled male sterility is more difficult to make use of
because it is usually controlled by a recessive gene and to obtain entirely
male-sterile progenies it would be necessary to self-pollinate homozygous
MS types, but these types, of course, carry no pollen to make the 'selfing'.
Functional male sterility can be used even when it is not cytoplasmically
controlled by opening the anthers artificially and using the pollen to effect
self pollination but unfortunately this condition is rare. One can obtain
segregating MS progenies by crossing MS homozygous plants with pollen
from fertile MS heterozygous plants. If male sterility were linked to an
easily recognisable morphological marker then it would be possible to
'rogue' (identify and remove) male fertile plants from the seed parent line
before they flowered. Unfortunately no such useful linked characters have
been found in the most important vegetable crops with genetic male
sterility. Another possible way of utilising non-cytoplasmic male sterility
would be to spray the homozygous MS plants with a substance such as
gibberellic acid and to make them temporarily fertile, and to self them to
give all MS progenies. This technique has interesting possibilities but it has
not yet been used on a large scale.
Male sterility occurs from time to time in progenies of wide inter-
specific crosses but so far little use has been made of this source with
horticultural crops. Attempts to use the cytoplasmic male-sterile condition
which occurs with Raphanus x Brassica by transferring it to cabbage and
other Brassica oleracea forms have not yet been successfully used on a
commercial scale partly because of the development of accompanying
female sterility.


Several attempts have been made to induce male sterility by spraying with
chemicals generally referred to as selective gametocides. The substance
40 Plant Breeding

most commonly experimented with is FW 450 (sodium 2, 3-dichloroiso-

butyrate) (Wit, 1960). Used as a 0.2 per cent aqueous spray it has induced
male sterility in such crops as cabbage, chicory, eggplant, lettuce and
tomato. A 0.5 per cent solution of a similar compound, sodium dichloro-
acetate, also has induced ma.le sterility in Antirrhinum majus (Kho and
de Bruyn, 1962). These substances have a transitory effect and need to be
applied about every five days throughout the flowering period to keep the
plants male sterile. Unfortunately the effects are modified by unknown
factors and the treatments are not fully rei iable as they sometimes do not
kill the pollen and they may seriously damage the plants by killing other
tissues. As a rule they do not reduce female fertility. Selective gametocides
are not yet used as a routine commercial treatment in the production of
seed of hybrid cultivars but they have been used by breeders to emasculate
flowers of some species prior to hybridisation (see Section 8.3.3).
The relatiwe numbers of male and female flowers vary in most mon-
oecious plants according to genotype and to the environment. Daylength
treatments have been given to obtain all female plants as an aid to produc-
ing hybrids but with limited success. Short days tend to increase 'female-
ness' of most cucurbits. Spraying squashes (Cucurbita pepo) and cucumber
(Cucumis sativus) with Cepa or Ethrel (2-chloroethylphosphoric acid) as
0.05 per cent solutions has given very promising results. The treat-
ments must be applied once or twice whilst the plants are still quite small
and give rise to fully female (gynoecious) plants.


Double flowers are either (1) flowers which contain extra petals or petaloid
organs, or (2) in the case of Compositae such as Chrysanthemum, Dahlia
and Pyrethrum, flowers in which some or all of the disc florets are replac-
ed by ray florets. In either case the condition is gene controlled, but the
degree of doubleness may be reduced when the plants are growing weakly
as with double cultivars of Clematis which may produce only single
flowers until the plants are well established. In the first type both stamens
and carpels may be petaloid but frequently it is the stamens only which
are changed and the plants then exhibit male sterility. As a rule this is not
complete and some functional stamens with pollen are produced but the
so-called 'petaloid' male-sterile form of carrot is a true double in which all
the stamens are converted to petals.
Another true double flower is that of the double stock (Matthiola
incana) which has neither stamens nor pistil and is completely sterile.
Doubling is a recessive condition so that homozygotes are double and
heterozygotes single and one would expect on selfing the heterozygotes
to obtain one double to three singles. However, in certain 'ever sporting
single' forms those pollen grains which do not carry the recessive allele die
so that when heterozygotes are crossed between themselves they give
approximately 50 per cent singles to 50 per cent doubles. Furthermore,
Flower Form and Pollination 41

seed of the double types tends to germinate before the singles and the
seedlings are more vigorous so that it is possible to recognise and to rogue
out most single forms at an early stage. This is especially important when
the plants are forced for early flowering as it economises on glasshouse
Occasionally male-sterile flowers may be desirable for reasons other
than their immediate decorative appearance, for instance as double flowers
and for controlling pollination. Lilium pollen readily stains petals, skin and
clothing and it is customary to remove the anthers of some species when
they are sold as cut flowers, so cultivars like 'Corsage' which produce no
pollen have been developed.


Except when dealing with trees or large shrubs most plants used by a
breeder for pollinating to produce new hybrids are grown in a glasshouse,
although the pollen may have been collected from plants growing in the
open. A glasshouse permits some control over the environment and protec-
tion from unwanted insects and predators. It makes it possible to maintain
temperatures above the ambient and to a limited extent to control humid-
ity, both factors which are important for good seed set and seed ripening
of many species grown in the cooler parts of Europe.
A glasshouse built specifically for breeding should preferably have
insect-proof screening on the ventilators, although it should also be possible
to give thorough ventilation to prevent temperature rising too high in
sunny weather. For wind-pollinated species such as beet and spinach it is
useful to have forced draught ventilation, the incoming air being filtered
to remove air-borne pollen. For many species it is desirable to have a glass-
house divided into small insect-proof compartments, or one capable of
division in this way when necessary. Artificial heating may be important,
but it is not essential for all crops and facilities for shading in bright sun-
light and for increasing humidity by misting are useful in some cases.
Certain wide crosses succeed best at fairly high temperatures and with a
dull moist atmosphere. Artificial lighting to extend daylength and to
supplement daylight is a useful facility.
In addition to the main glasshouse for the more exacting controlled
matings it is helpful to have some unheated insect-proof small glasshouses
or cages with a degree of protection from rain. These may be movable
structures and are especially useful in the production of small-scale nuclear
seed batches of vegetables and some annual ornamentals.


In many breeding programmes it is desirable to have a measure of control

over flowering time: (1) to stagger the work of hand pollination and thus
to make the optimal use of the facilities available, (2) to obtain more than
42 Plant Breeding

one generation in a year and decrease the time taken to reach the objective,
and (3) to induce the species which does not flower under ambient con-
ditions to produce flowers so that a desired mating can be achieved. It is
only possible to make a few generalised statements on flower induction
here; the breeder will have to study the literature in more detail to deal
with a specific crop (see Bleasdale, 1973).
Soaked seed of beetroot, chicory and carrot can be vernolised by cold
treatment of a few weeks at 2-3°C to induce early flowering. Seed of
lettuce, pea, turnip and spinach also may be vernalised to a lesser degree,
but this treatment has little effect on many species. However, a chilling
treatment of young plants for 8-12 weeks at or below S°C hastens flower-
ing in many plants, especially when accompanied by long daylength
treatment of about 16 hours. Exposure at or below freezing is not neces-
sary and often may damage the plants and even delay flowerin~. Further-
more it is considerably more expensive to refrigerate plants to 2 C than to
about S°C. Plants of some species such as Brossico olerocea cannot be
induced to flower until they have reached a certain physiological stage
which for practical purposes is reached when they have grown to a 'good'
size for transplanting in the field. In some cases, as with lettuce and endive,
flowering has been induced or hastened by spraying young plants with
3-10 p.p.m. gibberellic acid.
A room or rooms designed for temperature and daylength treatments
are :t valuable facility for a breeder. Temperature control need not be so
accurate as that required for physiological experiments but the refrigera-
tion should be adequate to maintain the desired low temperature with full
lighting even during the height of summer. An additional facility to
maintain temperature below freezing may be useful as an environment to
screen for low-temperature tolerance.
The time taken to induce flowering of seedlings of some trees such as
apple may be reduced by keeping them growing continuously at high
temperatures and long photoperiods and generally giving them optimal
fertile growing conditions. Manual defoliation has induced early flower
bud break and bark ringing of seedlings growing in the open hastens flower-
ing. Grafting on dwarfing rootstocks also has been used but care must be
taken to ensure that the root stocks are not virus infected.

BLEASDALE, J. K. A. (1973). Plant Physiology in Relation to Horticulture,
Macmillan, London, 144 pp
JONES, H. A. and EMSWELLER, S. L. (1934). The use of flies as onion pollinators,
Proc. Am. Soc. hort. Sci., 31,160-164
KHO, Y. 0. and DE BRUYN, ]. W. (1962). Gametocidal action of dichloroacetic
acid, Euphytica, 11, 287-292
KRAAl, A. (1954). The use of honey-bees and bumble-bees in breeding work,
Euphytica, 3, 97-107
Flower Form and Pollination 43

THOMPSON, R. C. (1938). Genetic relations of some color factors in lettuce, USDA

Tech. But., 620
WIERING, D. (1958). Artificial pollination of cabbage plants, Euphytica, 7, 223-227
WIT, F. (1960). Chemically induced male sterility, a new tool in plant breeding?,
Euphytica, 9, 1-9

FREE, ). B. (1970). Insect Pollination of Crops, Academic Press, London, 544 pp
HUDSON, ). P. (1957). Control of the Plant Environment, Butterworths, London,
240 pp


The Spermatophyta or seed-bearing plants are the most highly developed

members of the plant kingdom and the processes which occur within them
on the pathway to the formation of new offspring are complex and
fascinating. It is important that the breeder should understand these
processes, at least in broad principle, so that he _may in some cases direct
them to achieve the end product he desires. The Spermatophyta include
both the gymnosperms, including the conifers and cycads, and the angio-
sperms, comprising those genera normally thought of as flowering plants
although in its widest meaning this term embraces all the Spermatophyta.
'Gymnosperm' means 'naked seed' and the process of fertilisation is less
complex in this group than in angiosperms in which ovules and seed are
enclosed in an ovary. The discussions here are confined to the angiosperms.


Pollen develops from specialised cells within the anther known as pollen
mother cells. At a predetermined stage of stamen development these cells
all undergo meiosis in synchronisation, to give pollen cells joined in groups
of four and known as tetrads (see Section 1. 7.5).
The nuclei in each pollen cell carry half the number of chromosomes
of the parent plant, and because crossing over has taken place those of
each pollen grain differ genetically, though all carry the cytoplasm of the
mother plant. The tetrads generally break apart before the anthers split
open so that the pollen grains are separated from one another before they
are released. However, in Rhododendron and other Ericaceae the pollen
grains remain joined together in fours after they are shed from the anther,
and in Mimosa even larger aggregates of pollen are released. Many terrest-
rial orchids and species of Asclepias release their pollen in large packages
known as pol!inia, which have sticky pads and become attached to insects
visiting the flowers.
The pollen grain is not in itself the male sperm. It is a specialised cell
with its own controlling nucleus, the vegetative nucleus, and containing
within its cytoplasm either one or two generative nuclei, each having a thin
layer of its own cytoplasm and in fact constituting individual cells which
are the male gametes or sperms. The pollen grain of some species contains
only one generative nucleus when it is released from the anther but this

Fertilisation and Seed Development 45

divides to produce two at pollination or within the pollen tube as it travels

down the style.
Most pollen grains have a relatively rigid external wall which may
contain degradation products from the parent plant and which is patterned
in a way that is characteristic of the species and sometimes can serve as an
accurate feature for identification, especially with the aid of a scanning
electron microscope. There are one or more pores in the external wall
through which the pollen tube can grow, and their number and position
also may be important diagnostic features. Pollen grains are generally fairly
constant in size, usually with a diameter of 25-100 /-(m but those of some
species such as Cucurbita pepo may be as large as 230 /-(m. Pollen grains of
tetraploids and other polyploids are usually larger than those of the
corresponding diploids.
Pollen of most species loses its capacity to germinate after days or
weeks and often in an even shorter time if it is wetted. That of some
grasses is especially short-lived and may remain viable for only a few hours
under normal conditions. However, the plant breeder can store pollen of
some species, especially those of Rosaceae, Primulaceae, Ranunculaceae,
Liliaceae and lridaceae, for much longer periods. To avoid contamination
with unwanted pollen grains, anthers should be collected just before
dehiscence and laid out to dry in open containers in a dry room without
draughts, or in a desiccator. The dried pollen is transferred to corked air-
tight specimen tubes and kept in a deep-freeze cabinet. This technique
enables pollen of some plants such as apple, plum, rose, raspberry, lily and
iris to be used at least a year after it has been collected, but many pollens
will not keep for more than six months, even under these conditions.
It may be important for the plant breeder to know if freshly collected
or stored pollen is viable. If there is some doubt with fresh pollen, it
should first be examined under a microscope in a drop of liquid paraffin
or glycerine jelly (1 part gelatine warmed for 2 hours in 6 parts water
then 7 parts glycerine added with 1 per cent phenol as a preservative if
the preparation is to be kept). Plump grains of uniform size indicate that
the pollen probably is viable, but if the grains vary in size and many are
shrunken it may have low fertility or be completely infertile.
A more sophisticated test for pollen viability is that described by
Heslop Harrison (1970) which indicates the integrity of the cell membrane.
Pollen is mounted in fluorescein diacetate dissolved in acetone (2 mg per
ml) and diluted with a sucrose solution of a concentration selected to
minimise bursting {usually 0.5 M). The slides are viewed with a microscope
fitted with an ultraviolet Iight source, of course using precautions to
protect the viewer. Live grains fluoresce noticeably more brightly than
those which are dead or have low vitality. The original tests were used on
several species, including Cucurbita pepo and Tagetes patula. Staining with
tetrazolium salts has also been used as a test for pollen viability as with
plum {Norton, 1966).
46 Plant Breeding


The embryo sac, which can be looked upon as the female equivalent of
the pollen grain is found within the ovule in a tissue of thin-walled cells,
the nucellus. Like the pollen grain, the embryo sac develops from a tetrad
of cells with a reduced chromosome number produced by meiosis. How-
ever in the embryo sac the nuclei of these four cells are not always separat-
ed by cell walls. After a fascinating and complex series of nuclear divisions
and specialisation which varies according to the species, the embryo sac is
ready for fertilisation (Figure 4.1).
It is usually more or less egg-shaped and contains three groups of
nuclei. The egg cell, destined after fertilisation to produce the new plant,
lies at the more pointed end of the embryo sac. In most species it is
accompanied by two specialised cells of characteristic pointed shape, the
synergids, whose function is incompletely understood but seems to be to
aid fertilisation. They usually die shortly after this takes place. There are
no true synergids in a few genera such as Ceratostigma. Near the centre
of the embryo sac are one to nine polar nuclei which, after fusion with one
of the generative nuclei from the pollen tube, give rise to the endosperm.
Embryo sacs of most plants have only two polar nuclei but there are four
in Li!ium, Fritillaria and some species of Ery thronium and a few other
genera. At the blunt end of the embryo sac there are a number of so called
antipodal cells which normally play little apparent part in the develop-
ment of the embryo, but under certain exceptional circumstances one or

Figure 4.1 Embryo sac of a typical angiosperm. s, Synergids; e, egg cell; p, polar
nuclei; a, antipodal cells.
Fertilisation and Seed Development 47

more of them may function as a sex cell. Very occasionally one of the
synergids also may function as an egg cell. Thus the antipodal cells and
synergids may act as stand-by reserves in the process of fertilisation if
the egg cell fails.


In angiosperms the egg cell is confined within the embryo sac, itself em-
bedded in the nucellus, which is part of an ovule enclosed in an ovary.
The conveyance of the male gametes from the receptive surface on the
outside of the ovary, the stigma, to the female gametes is achieved by the
pollen tube, and a pollen grain no more than 50 pm in diameter may
produce a tube 10 em or longer to achieve this objective.
The surface of the stigma is generally composed of finger-like struc-
tures, the papillae, and may be bathed in a stigmatic exudate as in the tulip
and other Liliaceae. Both these devices help to retain pollen on the stigma.
The exudate mainly contains sugars and its role is not fully understood,
but it seems that its primary function is to provide water at a satisfactory
osmotic pressure which will allow the pollen to germinate yet minimise the
risk of the pollen tube bursting. However, it may in some circumstances
aid germination in other ways, and some plant breeders claim that the
transference of exudate from stigmas which are known to be compatible
with a given pollen to those which are not can aid in the production of
hybrids which are otherwise unobtainable.
Pollen usually begins to germinate on a compatible stigma after a few
minutes. A pollen tube breaks through one of the pores in the outer

Figure 4.2 Diagram illustrating germination of pollen on stigmatic surface (st).

p, Pollen grain; t, pollen tube; s, sperm nuclei; tn, tube nucleus.
48 Plant Breeding

covering of the cell carrying with it the vegetative nucleus followed by

the sperm cells (Figure 4.2). Occasionally, as in some Cucurbitaceae, more
than one tube is produced but only one from each pollen grain can carry
the vegetative nucleus and the others soon discontinue elongation. The
functioning pollen tube may penetrate the papillae or grow down between
them and it usually then follows the line of least resistance. In many
monocots the style is hollow and the pollen tube travels down the cavity.
If the pollen contains only one sperm cell at pollination, as occurs with
some species, this divides to produce two during the passage down the
style, or in exceptional cases, after the tube has entered the ovary. The
pathway of the tube to the embryo sac varies according to the species and
may pass through the stalk of the ovule or through the ovary wall directly
to the embryo sac end of the nucellus.
The pollen tube usually approaches the embryo sac at the end closest
to the egg cell where it encounters the synergids and bursts to release the
sperm. The function of the synergids has been described as acting as
shock-absorbers against the impact of the pollen tube, or to rip it open,
though the precise function is still not fully understood. Frequently the
synergids disintegrate after fertilisation but sometimes one persists for a
time, although it performs no important function in the developing seed.
One of the sperms (generative nuclei) unites with the egg cell to form
a zygote which will develop into the new embryo and the other fuses with
the polar nuclei to give rise to the endosperm. Several polar nuclei are
involved in fusion with one sperm so that the endosperm contains a larger
contribution from the mother parent and is therefore not genetically the
same as the zygote. The synergids, vegetative nucleus and antipodal cells
usually die after fertilisation has taken place. An important feature to be
remembered is that a 'double fertilisation' occurs; that of the egg cell and
that of the polar nuclei. If this is not successful then a viable seed is rarely
produced. Another important feature of the process of fertilisation is that
the cytoplasm of the generative cell is only rarely known to be carried
through to the new hybrid plant which inherits its cytoplasm solely from
the egg cell. However the so-called 'tare-leaved rogue' character of cyto-
plasmic origin of some pea cultivars is transmitted through pollen and in
this case some cytoplasm must be transmitted with the sperm.
The time interval between pollination and fertilisation for most
species is generally 12-48 hours but it may be shorter in some Compositae
and longer in some tropical orchids. It is much longer for some trees and
shrubs; 3-4 months in the hazel Corylus ave/lana and 12-14 months in
some oaks (Quercus spp) and in Hamamelis. Pollen tubes over-winter in
the last two genera. The rate of growth of pollen tubes is also influenced
by the genotype combination of the pollen and the pistil and this will
be discussed in more detail under the section on incompatibility. Environ-
mental factors, especially temperature, affect the growth rate, and some
difficult crosses may be obtained more readily when the pollinated plants
Fertilisation and Seed Development 49

are kept at a relatively high temperature. For this reason, in the uncertain
climate of northern Europe it is usually advantageous for the breeder to
make his pollinations on plants protected in a glasshouse wherever possible.
For studies on pollen tube growth and for practical plant breeding
requirements, it is often desirable to be able to trace the progress of
pollen tubes in the style. This used to be done by the laborious method
of transverse sectioning of prepared material. A quicker technique was to
fix and stain styles and squash them on a slide with a cover slip for micro-
scopic observation, but difficulties were often experienced in distinguish-
ing the pollen tubes from surrounding cells, especially from the conducting
tissue. A modern squash technique described by Martin (1959) has greatly
facilitated the test. It depends on the presence of callose in the pollen
tubes which, stained with aniline blue dye, fluoresces bright yellow green
against the blueish colour of the stylar tissue when viewed under the
microscope with ultraviolet light. The test seems to be applicable to a wide
range of species.
In the updated technique used as a routine test in Brassica, the pistils
are fixed for at least 5 hours in a mixture of 60 per cent ethanol, 10 per
cent glacial acetic acid, 30 per cent chloroform. They are then transferred
to absolute ethanol and taken down through two or three intermediate
concentrations of ethanol in water to distilled water; softened for 1 hour
at 60°C in 0.8 M sodium hydroxide; stained for 3 hours in 0.1 per cent
aniline blue in 0.1 M potassium phosphate (K 3 P0 4 ); mounted in 80 per
cent glycerol and gently squashed. It is important to use the specified


After a short resting period, lasting usually for several hours or at the most
a few days, the fertilised egg cell begins to divide and the proembryo is
formed. This enlarges and elongates fairly rapidly, and within one or two
weeks the cotyledons begin to develop and the embryo proper is formed
(Figure 4.3).
Fusion of a generative nucleus within the polar nuclei starts off the
development of the endosperm, usually before the fertilised egg cell has
completed its resting period. In most angiosperms growth of the endo-
sperm is essential for seed development and without it the embryo fails or
aborts prematurely. However, although fertilisation of the polar nuclei is
essential for seed formation in orchids, the endosperm does not develop
in this group of plants and the embryo remains in the proembryo stage,
very small and undifferentiated. In the majority of angiosperms the
endosperm acts as an intermediary for the transference of food supplies
from the mother plant to the developing embryo and it is probably signifi-
cant that its genetic components (usually 2 parts female parent to 1 part
male parent (Figure 4.1), or in the case of some Liliaceae such as Lilium
50 Plant Breeding

Figure 4.3 Embryo development in Capsefla bursa-pastoris, typical of Cruciferae.

(Adapted from Johansen, 1950.)
Fertilisation and Seed Development 51

and Fritillorio 4 parts female to 1 part male, are intermediates between

those of the embryo and the mother plant. In most plants some endo-
sperm tissue is carried through to the ripe seed where it functions as a
food supply for the germinating seedling. However, in some genera the
endosperm is completely or almost completely absorbed and the ripe
seed contains no endosperm but a large embryo with a food supply for the
young plant stored almost entirely in the cotyledons. Such non-endospermic
seeds are found in Leguminosae including peas, beans, lupins; Cruciferae
including cabbage, turnip, wallflower and some Rosaceae, but not rasp-
berry or strawberry. Most monocots of horticultural importance have seeds
with copious endosperm but those of Alismataceae are non-endospermic.
The development of the endosperm does not usually proceed in the
organised manner of most other plant tissues. As a rule, in the early stages
it is non-cellular with free nuclei, sometimes of different chromosome
numbers. It can be seen as a milky fluid exuding from immature seeds if
these are squashed between the fingers, and the liquid nature of the endo-
sperm is clearly observed in young horse chestnut seeds or in the coconut.
Cell walls generally develop in the endosperm before the embryo reaches
full size and the endosperm of mature seeds is usually solid unless there
has been some malformation in its development.


The term fruit is difficult to define accurately but may very broadly be
described as the enlarged or altered wall of the ovary containing one or
more seeds. Other parts of the flower, especially the receptacle, may be
involved in false fruits such as the strawberry, apple, fig and rose hip.
The plant breeder requires a fairly detailed knowledge of the superficial
fruit morphology of the plant being worked on, but this is easily acquired
by experience and an extensive examination of fruit types is not possible
here. However, it is important to realise that some 'seeds' such as those
of the carrot, celery, Ronuncu/us spp, spinach, lettuce and marigold are
in fact dry one-seeded fruits. Beetroot 'seeds' as generally sold are also dry
fruits, usually containing more than one seed, but sometimes these clusters
are broken up by machine and single true seeds sold as an aid to singling
and precision seeding.
Fruit and seed development are often interdependent so that failure
of one or the other affects both, and this can be very important to the
plant breeder wishing to achieve a certain cross. In black currants, blue-
berries, apples and pears for example, the developing seeds produce
hormones which sustain fruit growth. If hormone production falls to a
low level because very few seeds have been set it may be insufficient to
maintain fruit growth, an abscission layer is formed, the fruit is shed and
the seed lost. This situation has sometimes been overcome by applying
naphthaleneacetic acid (~AA) or its sodium salt dissolved in either water
52 Plant Breeding

or lanoline. Other auxins, gibberellins and cytokinins have also been used
for the same purpose. Pear-apple hybrids were achieved by brushing a
40 p.p.m. solution of ,6-naphthoxyacetic acid on the pear ovary and apply-
ing apple pollen to the stigma (Crane and Marks, 1952). This prevented
fruit abscission before the apple pollen tubes, which are slow growing in
pear tissue, reached the embryo sac and effected fertilisation. Without
these aids this hybrid combination probably cannot be achieved.
Pollination alone induces sufficient hormone production in some
plants to allow fruit development without fertilisation and in some culti-
vars not even this stimulus is necessary. For example, the fig cultivars
grown for their fresh fruit do not require fertilising to develop ripe fruit
whereas those of the Smyrna fig grown for dried fruit will not develop
without fertilisation. Both are forms of .the same species, Ficus carica.
A similar situation occurs with the glasshouse cucumber and the outdoor
or ridge cucumber; both are forms of Cucumis sativus. The former regularly
produces parthenocarpic fruits without pollination whereas the latter
does not usually develop fruits unless it is fertilised, but some individuals
may occasionally do so under certain environmental conditions.


Seeds from a breeding programme represent much valuable effort stored

in a small bulk and should be treated with the utmost care.
Extraction of seed from dry non-fleshy fruits is not usually difficult
when small bulks are being handled as these can be rubbed or shaken out
by hand. With larger quantities, such as advanced selections of vegetable
stocks for field trials, some small threshing and cleaning machines may
be useful. A few genera such as Viola and Impatiens eject seed from ripe
capsules which must be stored in suitable containers with lids to prevent
seed loss.
The extraction of seeds from fleshy fruits such as those of Fragaria,
Lonicera, Rosa, Rubus, Ribes and Vaccinium can be done by macerating
a suspension of fruits in water in a domestic liquidiser for a short period.
The seed will then sink and the water with skins and pulp is decanted;
the seeds are washed and the operation repeated until they are clean and
can be laid out to dry. Strawberry fruits may be roughly skinned and the
skins which bear the seeds dried between folded sheets of paper; when dry
the seeds can be rubbed off by hand. Seeds of some species are surround-
ed by a mucilaginous pulp and this may be washed away after a 1-2 day
fermentation period of the crushed fruit. A more satisfactory method is
to treat the pulped fruits with acid. A common procedure with tomatoes,
cucumbers and marrow (Barson and Ballinger, 1944) is to add the equiva-
lent of 4 ml of concentrated hydrochloric acid to every pound of pulp
and to decant the skin and flesh with water after a 15 minute treatment.
Sulphuric acid (30 per cent), which may be used in the same proportion,
Fertilisation and Seed Development 53

Table 4. 7

Relative storage capacity of some seeds (after Heydecker, 7974)

Low: leek, onion, parsley, salsify.
Medium: asparagus, beans {broad, French, runner), calabrese, carrot, celery and
celeriac, kohlrabi, pea, spinach {true and New Zealand}, sweet corn,
sweet pepper, tomato.
High: beet {red and spinach), most brassicas {including Brussels sprouts, cab-
bage, cauliflower, kale, swede, turnip}, chicory, cucumber, egg plant,
endive, melon, radish, vegetable marrow.


Low: Cal/istephus, Celosia, Delphinium {perennial}, Festuca, Helichrysum,
lberis, Kochia, Lilium, Nemesia, Papaver {perennial}, Salvia.
Medium: Ageratum, Antirrhinum, Aster, Bellis, Campanula, Chrysanthemum
{perennial), Dahlia, Delphinium {annual, larkspur}, Digitalis, Godetia,
Lupinus, Myosotis, Poa, Penstemon, Petunia, Phlox, Pyrethrum, Tagetes,
High: Althaea, Alyssum, Arctotis, Calendula, Cheiranthus, Centaurea, Chrysan-
themum {annual), Coreopsis, Cosmos, Dianthus, Eschscholtzia, Lathyrus,
Unum, Matthiola, Nigella, Papaver {annual), Salpiglossis, Schizanthus,
Trapaeolum, Verbena, Zinnia.

{A) Seeds which require moist storage, preferably cold: Acer {some species},
Aesculus, Carpinus, Castanea, Corylus, Citrus, Fagus, juglans, Quercus.
{B) Seeds which keep themselves dry: open storage suffices: Acacia, Albizzia,
Eleagnus, Eucalyptus, Rhus, Robinia.
{C) Most other tree seeds store well at 0-1 0°C, preferably dry {ideally
4-6 per cent water content}.

is cheaper but more dangerous to use.

Seeds vary in their storage capacity. All seeds should be dried in a
warm dry atmosphere before storage and special care must be taken of
those kinds which have a low or medium storage capacity (Table 4.1).
When dry they preferably should be packeted and sealed in containers
with airtight lids, and kept in a room with a temperature which is not
excessively high or fluctuating. Under these conditions seeds of high
storage capacity will remain viable for at least five to six years, but those
of low capacity may lose their viability after a year or less. It is preferable
to keep the dried seeds in a refrigerator or deep freeze, and under these
conditions even low storage capacity types which normally last no more
than a year will keep for up to five or six years. Many tree and shrub
seeds will remain viable for a year or more only when stored in a deep
freeze. However, some large seeds of non-winter hardy species, such as
French and runner beans, may be damaged by too low temperatures.
54 Plant Breeding

Some seeds exhibit dormancy and their germination and subsequent

development is delayed even when sown under apparently favourable
conditions. This does not apply to most crops which are maintained by
seed rather than vegetative propagation, such as the majority of vegetables
and annual or biennial ornamentals, although it may do so to a small
degree in some, such as lettuce, carrot and sweet pea. Seed dormancy can
be a problem to the breeder of perennial crops by lengthening the time
taken to raise a generation of new plants. It is a major obstacle in the
breeding of bearded iris, causes some delay in apple and rose breeding and
can be a nuisance in the breeding of many other crops.
Seed dormancy may be due to impermeability of the seed coat, to a
physiological condition of the embryo, or to both. In some plants, notably
the tree peony (Paeonia suffruticosa) and several lily species, the epicotyl
of the seedling itself may become dormant. Seed coat impermeability is
common in tropical Leguminosae but occurs to a lesser degree in some
temperate genera such as Lathyrus. It occurs also in many Rosaceae, in
which it is associated with physiological dormancy.
Seed coat dormancy sometimes may, in small quantities of seed, be
overcome by abrasion from gentle rubbing between sheets of sandpaper,
as with sweet pea. Soaking in ethanol is effective with some species and
immersion in concentrated sulphuric acid which is dangerous for the
operator is frequently an effective treatment. Raspberry seeds undergo a
short dormant period, and to reduce this so that they can be sown to
produce a generation of seedlings the year following hybridisation, may be
treated for 20 minutes in concentrated sulphuric acid, then for one week
in 1 per cent calcium hypochlorite and an excess of calcium hydroxide,
followed by moist chilling for six weeks (Jennings and Tulloch, 1965). A
similar treatment is effective with many other Rosaceae but the majority
require longer low temperature treatments and are U?Ually stratified. This
is essentially moist low temperature (about freezing or just above) treat-
ment for 2-4 months. Seeds are generally mixed with moist sand and this
is effective for roses, and many rosaceous fruit trees.
Seeds without impermeable seed coats but subject to physiological
dormancy often need fluctuating temperature treatment, but sometimes
leaching with water or treatment with 0.01 M potassium nitrate or 0.5-3.0
per cent thiorea may be effective. In extreme cases such as the bearded
iris, embryo culture may be used.


We have examined the normal course of events which follows when a com-
patible pollination is made by transferring viable pollen to the stigma of
a plant with a visible embryo sac; but the processes leading to fertilisation
and seed and seedling development frequently may be blocked at some
stage and then the breeder is denied the gene combination he planned to
Fertilisation and Seed Development 55

obtain. Interruption can occur at many stages. Blockages occurring between

pollination and the release of sperms in the embryo sac, are generally
referred to under 'incompatibility' and this term will be used here for this
specific meaning, though it sometimes is used also to describe certaio
post-fertilisation failures.
Many plant species have a gene-controlled natural incompatibility
system which prevents or deters inbreeding by self-fertilisation or fertilisa-
tion between siblings. Such systems have been evolved by natural selection
because inbreeding is usually disadvantageous to species in the wild as it
frequently leads to a reduction in vigour and other undesirable character-
istics which weaken the stock. The plant breeder may wish to impose
inbreeding temporarily on a species in which outbreeding is the usual
course because this is a useful technique for improving genetic uniformity.
It may also be desirable to select for improved or complete self compat·
ibility without loss of vigour, and this has been achieved for some vege·
tatively propagated crops such as the raspberry, which depends on seed
s'et to produce a crop of fruit and yet is usually grown as a monoculture
and therefore not exposed to foreign pollen. Cultivated red raspberry
cultivars (Rubus idaeus) are self fertile whereas most wild forms of this
species have varying degrees of self incompatibility. This situation has
not been fully achieved with other fruits like apples and pears which
usually cannot be grown in monoculture and need some mixing of com-
patible cultivars in an orchard. Nevertheless some pear cultivars such as
'Conference', which is self incompatible, will produce fruit partheno-
carpically without fertilisation.
An incompatibility system may be a gene-controlled mechanical
device to prevent self pollination by insects, as with the dimorphic flowers

Figure 4.4 Dimorphic flowers of Primula. a, Pin-eyed; b, thrum-eyed.

56 Plant Breeding

of Primula (Figure 4.4) and Forsythia. However, when an 'illegitimate'

pollination is made by hand self-pollinating a thrum-eyed or a pin-eyed
flower, seed often is not produced because the plant also has a gene-
controlled physiological system which regulates pollen tube growth in
certain pollen-pistil combinations. Such pollen-pistil incompatibility
systems are also common in many species which do not have dimorphic
flowers. There are two types, the gametophytic and the sporophytic
systems in both of which the pathway to fertilisation is controlled by
different alleles of an incompatibility gene generally called S alleles.
These alleles can be crudely likened to keys to fit the S lock. The wrong
combination of lock and key prevents the pollen tube from delivering
its contents to the embryo sac. When any of the alleles controlling the
behaviour of the pollen are the same as those of the pistil the combina-
tion is incompatible. In most plants only one incompatibility gene is
involved but in grasses the system is complicated by two genes and there
are thus two locks to open.

Figure 4.5 Diagram illustrating gametophytic (G) and sporophytic (S) incompatibil-
ity systems. 5 1 -5 4 denote incompatibility alleles.
Fertilisation and Seed Development 57

4.8.1 The gametophytic incompatibility system

Here the reaction of the pollen is controlled by the interaction of the S
alleles of the pollen itself with those of the pistil of the pollinated plant
(Figure 4.5). Blockage of the pathway to fertilisation occurs by excessively
slow growth of the pollen tube and its cessation of growth before reaching
the embryo sac. This system is also associated with pollen grains having
only one generative nucleus at the time of dehiscence from the anthers. It
is probably the most widespread of the two systems and is known to occur
in Leguminosae, Onagraceae, Papaveraceae, Rosaceae, Solanaceae, Liliaceae
and in horticultural crops such as apple, pear, cherry, tomato, Papaver,
Petunia, Li/ium, Antirrhinum and Nemesia.

4.8.2 The sporophytic incompatibility system

Behaviour of the pollen is controlled by the S alleles of the pollen bearing
plant and not by those of the pollen itself, thus all the pollen from a plant
behaves in the same way in its incompatibility reactions (Figure 4.5). The
pollen as a rule does not germinate on the stigma of an incompatible plant
and if it does so the pollen tubes rarely penetrate the stigma. This system
is usually associated with pollen grains having two generative nuclei when
the pollen is shed from the anther. It occurs in Compositae, Cruciferae
and Rubiaceae and is important in the breeding of cabbages and other
forms of Brassica oleracea, radish, Cosmos, lberis and ?rimula. In ?rimula
it is associated with the dimorphic flowers and the pollen has only one
generative nucleus when it is shed.
Incompatibility, theoretically, occurs when the behaviour of the
pollen is controlled by any S allele which is the same as that of the pistil
as in Figure 4.5. However, this is an idealised situation and apparently
incompatible crosses can sometimes lead to fertilisation and seed produc-
tion because blockage is incomplete. This is especially true of the sporo-
phytic system in species like Brassica oleracea where there are as many as
40 different S alleles, some of which are dominant over others (Thompson
and Taylor, 1966) and some of which interact to induce so-called mutual
weakening and allow pollen tube growth. This apparent 'breakdown' of
the incompatibility system is very troublesome in the breeding of F 1
Brassica crops, as we shall see later.

4.8.3 Incompatibility in inter-species crosses

Another type of pollen-pistil incompatibility is that found when crosses
are made between different species. In these cases incompatibility may be
governed by S alleles if the species have common genes for incompatibility,
but it is often controlled by many other additional factors. The main
feature of this type of inter-specific incompatibility which is reviewed by
Lewis and Crowe (1957) is that it is frequently possible to make the cross
only one way. For example, pear 9 x apple; french bean 9 x runner bean;
58 Plant Breeding

radish 9 x cabbage hybrids may be obtained, but the reciprocal crosses are
more difficult or impossible to achieve. As a rule, when one of the parents
is self compatible it is advisable to use it as the female parent.


In Brassica o/eracea, the incompatibility blockage does not operate until

the flowers have opened. Thus it is possible, by prising open flower buds
and pollinating them prematurely, to obtain selfings and incompatible
crosses which cannot be achieved when open flowers are pollinated. The
blockage with Brassica and other plants with a sporophytic system occurs
at the stigma, and if this is cut off and self pollen applied to the stump
some seed will be produced, although usually not as much as when the
flowers are bud pollinated, but this is not effective with many species.
When a Brassica stigma is damaged mechanically or by momentarily touch-
ing it, prior to pollination, with a soldering iron maintained at 80°C, self-
fertilisation may be effected (Roggen and van Dijk, 1976). Petunias pro-
duce stigma exudate and incompatible combinations may sometimes
permit fertilisation and seed set if this exudate is washed off and replaced
by that from a compatible combination. Also, the removal of the entire
stigma and its replacement by laying on one cut from a compatible plant
may help with some species to obtain combinations which cannot other-
wise be achieved.
Excessively slow pollen tube growth in the style is sometimes a cause
of failure in wide crosses and when selfing plants with a gametophytic
incompatibility system. Various methods of reducing the distance that the
pollen tubes need to travel to reach the embryo sac have been used to
overcome this difficulty. Cutting off the style to the base of the ovary and
pollinating the stump enabled the hybrid of sweet pea Lathyrus odoratus
x L. hirsutus to be achieved (Davis, 1957). The injection of a suspension
of pollen in sterile water into the hollow style of Lilium and into the
ovary of Paeonia and Papaver has given fertilisation on an experimental
scale, but it does not seem to have been used regularly as a tool for the
breeder to obtain new hybrids. A more complex technique is to transplant
both ovules and pollen together under sterile conditions to a nutrient
medium and obtain fertilisation and seed production in vitro. This has
been achieved with Papaver somniferum (Kanta eta/., 1960) and attempt-
ed for several species by Guzowska (1971) but again it does not seem to
be used by breeders, even though it may have interesting possibilities.
Various environmental conditions may help to overcome incompat-
ibility. High carbon dioxide content of the surrounding atmosphere and
high temperatures tend to favour self fertilisation in Brassica o/eracea
forms which are partially self compatible, whereas relatively low tempera-
tures may favour some wide crosses. High atmospheric humidity may
have a similar effect.
Fertilisation and Seed Development 59

Attempts to obtain a chosen wide cross should not be given up lightly

if they do not succeed first time. A large number of pollinations should
be made using as many different varieties or genotypes of the two parents
as can be obtained and the crosses made both ways. In the early nine-
teenth century M. Debras made extensive pollinations for some years
between Lilium henryi and L. sargentiae before he obtained the first
'Aurelianense' hybrid of this very successful combination.


Many wide crosses may fail, not because the pollen tube is unable to
reach the ovary, but because the sperm and egg cells are unable to fuse.
For example, when tomato plants are pollinated with the snapdragon, the
pollen tubes will grow down to the ovary but fertilisation docs not occur
and no seeds are formed. There is little that can be done to overcome this
type of barrier except to persist in making the cross, although of course a
tomato-snapdragon cross is too wide to hope for success.
Even when a successful double fertilisation takes place there are still
sometimes obstacles which prevent the production of viable hybrid plants.
Such cases are often associated with a disruption of the endosperm which
fails to develop, dies at an early stage of seed development, or is in some
way antagonistic to the development of the embryo. For example the
lilies L. /ankongense x L. davidii and L. pyrenaicum x L. szovitzianum
produce seeds with very small embryos and lacking endosperm, and which
will not grow when sown on soil. The cross L. auratum x L. speciosum
develops apparently normal seed with full-sized embryos and plenty of
endosperm but when the dried seed is soaked the endosperm often pro-
duces a substance which is toxic to the embryo and kills or damages it so
that it is unable to develop into an autonomous plant.
The breeder may overcome such difficulties arising from the endo-
sperm by using an embryo culture technique. Although embryos some-
times have been excised from seeds and grown in soil, modern embryo
culture involves the removal of the embryo aseptically and its culture on a
sterile nutrient medium. This need not be a complex operation requiring
very specialised laboratory facilities and is well within the capabilities of
the part-time or amateur breeder. In spite of this, embryo culture has been
used in the breeding of only a few horticultural plants; the main recorded
successes are with beans, peach, potato, tomato, iris and lily.
Before it has dehisced, a fruit is a sterile package and all that is usually
necessary to obtain sterile embryos is to dip the fruit in industrial alcohol
before opening it to remove the seeds and embryos from the seeds. How-
ever, if seeds already free from the fruit are used, they should first be
soaked for 10 minutes to 2 hours in a 5 per cent stabilised calcium hypo-
chlorite solution and then washed in sterile water. In extreme cases a more
effective surface sterilant such as mercuric chloride may be necessary.
60 Plant Breeding

Excision of the embryos and their transfer to containers of nutrient

medium is best done in a room or cabinet with air filtered to extract
particles greater than 4 ~m in diameter. If this facility is not available the
operation may be done under a sheet of glass in a clean room with little
air movement, spraying under the glass with industrial alcohol shortly
before excising. From time to time the scalpel and other instruments used
in the operation should be dipped in industrial alcohol, which should be
allowed to evaporate from the instruments before re-using.
Proembryos taken shortly after fertilisation are often difficult to
culture, requiring complex liquid media, and sometimes more than one
change of medium, but many proper embryos will grow on one percent
agar with sugar and a simple salts solution. Examples of a simple and a
rather more complex medium are given in Table 4.2.
Some media can now be purchased weighed and ready to make up by
adding distilled water and these are especially valuable to the worker with
limited laboratory facilities. Different accessory substances sometimes

Table 4.2

Some media used for embryo culture (mg per litre of water)*

Emswel/er Tukey White

and Uhring (7934) (7 963)

KN03 200 136 80

Ca(N03b 800 200
KH2P04 200
NaH 2 P04 17
Ca3(P04b 170
Na2S04 200
MgS04 400 170 360
CaS04 170
KCI 680 65
FeP04.2H20 170
Fe 2 (S04)3 5 2.5
H3B03 1.5
Kl 0.8
MnS04 4.5
ZnS04 1.5
CuS04.5H 2 0 0.01
Mo03 0.001
Glycine 3
Nicotinic acid 0.5
Thiamine 0.1
Pyridoxine 0.1
20 g of sucrose and 5-10 g agar are added to each litre of water.

*Method of preparation is not straightforward and original references

should be consulted before making up media.
Fertilisation and Seed Development 61

have been added to media, including endosperm extracts such as coconut

milk, liquid extract from immature horse-chestnut seeds, or yeast extract;
also growth substances such as indole-acetic acid, naphthaleneacetic acid,
kinetin, adenine sulphate and gibberellins; also thiamine, ascorbic acid,
inosotol and nicotinic acid. It is advisable, however, to make first attempts
with simple media and to complicate them further only if the initial results
are unsatisfactory.
Embryos are usually cultured individually in glass bottles on slopes of
nutrient agar. The bottles must be sterilised before transferring the embryos
and this can be done conveniently in batches of 50-100 in a domestic
pressure cooker for 15-20 minutes at 15 pounds per square inch. It is
helpful, but not always necessary, to use a stereomicroscope as an aid
when excising small embryos, but a simple magnifying lens may prove
adequate with large seeds. As a rule, the embryos grow best when laid on
the surface of the agar, but cyclamen embryos grew better when imbed-
ded just below the surface (Gorter, 1955).
The conditions required for the optimum growth of the cultured
embryos depend on the species but adequate temperature and light levels
are often important, especially during the later stages of growth. When the
embryos have developed into plants which are too large for these bottles,
they should be transferred to soil compost and this stage is critical to their
survival; high humidity and adequate light are generally important during
this period and the use of a mist propagator may be advantageous.
Embryo culture may also be helpful to overcome seed dormancy. The
factors which prevent germination almost invariably lie in parts of the seed
other than the embryo itself. When the embryos from dormant seeds are
isolated on sterile nutrient media they usually begin growth without delay.
Embryo culture is not always an economic technique to overcome seed
dormancy but it can be valuable to help breeders to reduce the time
required to produce a new generation. It has been especially useful in the
breeding of bearded iris (Lenz, 1955).


This is an exciting new development which may lead to a technique

enabling breeders to achieve hitherto unobtainable genetic combinations.
In principle, it involves the removal of the cell walls from two different
genotypes and the fusion of these naked cells or protoplasts to produce
plants of novel genetic combinations. Such a technique would bypass
pollen-pistil incompatibilities and also endosperm antagonism because the
new plantlets would not depend on the endosperm for nourishment. It
might also get round any obstacles which occur about the time that the
sperm cell enters the embryo sac. A hybrid produced in this way would
differ from the same combinations produced sexually, in that the cyto-
plasm would originate as a combination between that of both partners.
62 Plant Breeding

As we have seen, in sexual reproduction the male cytoplasm is not usually

taken into the hybrid except in very small amounts.
The technique requires fairly sophisticated laboratory facilities and,
unlike embryo culture, it is probably beyond the capacity of the average
plant breeder. Cells, often from the mesophyll of leaves, are treated with
an enzymt: to dissolve the cell walls. The two kinds of protoplasts to be
fused are then mixed and sodium nitrate or polyethylene glycol added to
induce fusion. The protoplasts are then centrifuged, washed and distribut-
ed on a Petri dish with nutrient agar. These produce callus tissue which,
with the application of growth substances, can be induced to differentiate
shoots for isolation as new plants.
It is not difficult to obtain protoplasts; the problems lie in the induc-
tion of sufficient numbers of hybrid fusions, their screening from non-
hybrid tissue and the establishing of autonomous plants. Nicotiana and
Petunia have most frequently been used in the experiments and protoplast
fusion between these two genera has been achieved, but as yet no hybrid
plant has been obtained from this intergeneric combination. However,
several hybrids between species within the genus Nicotiana have been
produced by protoplast fusion. Sometimes these have abnormal chromo-
some numbers and this in itself may be of use to the breeder. It must be
stressed that protoplast fusion is at present mainly of academic interest
and is not yet a standard technique for the breeder, but it has such excit-
ing and far-reaching possibilities that breeders will need to keep in touch
with developments.

The progenies of some plants closely resemble their mother parent, even
when seed set appears to have resulted from deliberate pollination with a
different and distinct male parent. This phenomenon, in-which the embryo
of the seed is not the product of fertilisation, is called apomixis. Strictly
speaking the term also embraces the development of plantlets at sites
where flowers would normally form and includes the production of vivi-
parous plants as in the tree onion or Egyptian onion, which is a form of
the comon onion (Allium cepa} that produces bulbils instead of flowers
on the inflorescence. This phenomenon occasionally occurs also in some
forms of the leek (Allium porrum) and is made use of by flower-show
enthusiasts to maintain clones of their most successful plants.
Apomixis through seed production, which should more precisely be
called agamospermy, can arise by several different pathways. The processes
may be complex and are not completely understood in some instances,
but three types of seed apomicts can be recognised.
(1) In a few cases seed develops directly from the nucellus, no embryo
sac is formed, and the sexual process is completely bypassed. This occurs
frequently in Hosta and some species of Rubus, Allium and Opuntia, and
Fertilisation and Seed Development 63

can be a barrier to hybridisation in those genera. The new seedlings produc-

ed are genetically identical with the mother plant. In some circumstances
apomixis has been used to advantage by horticulturists; for example, the
propagation by apomictic seedlings of apple root-stocks which are difficult
to propagate vegetatively in the normal way.
(2) A normal embryo sac is formed and the egg cell develops into an
embryo without fertilisation. The embryo and subsequent seedling will
then have half the number of chromosomes of the seed parent. In most
cases pollination and sometimes fertilisation of the polar nuclei are neces-
sary to induce the unreduced egg cell to start dividing. This type of
apomixis can be used to advantage to obtain true breeding progenies; a
notable example is in barley breeding (Kasha and Kao, 1970).
(3) The most frequent type of apomixis through seed production is
when an embryo sac is formed directly from a diploid cell without reduc-
tion division or when reduction division occurs but there are anomalies in
the events which follow so that some of the products fuse to produce
diploid cells. In the first case the egg cell has the same genetic constitution
as the mother parent and in the second there is some segregation similar to
that which would occur from self fertilisation because there may have
been an opportunity for chromosome crossing-over or pairing between
unlike chromosomes may have occurred. In many cases pollination is
necessary before seed is produced and this type of apomixis is then refer-
red to as pseudogamy or false fertilisation. It is not always necessary to
have fertile pollen, frequently that from other species or pollen killed by
X-ray treatment is adequate to trigger off the process.
Non-segregating apomictic seeds are the usual mode of reproduction
of most forms of the wild dandelion (Taraxacum officina/e) and occur also
in Antennaria alpina. In most other cases, as in some species of Citrus,
Erigeron, Lilium, Malus, Potentilla, Rosa, Rubus and Rudbeckia, some
segregation of the progeny occurs.
From a practical point of view the breeder attempting to produce
hybrids is faced with three types of seed apomixis:
(1) Some Hosta species, for example, very rarely produce hybrid seed
even when plants are pollinated with chosen male parents. It may be
useless repeating such crosses in the same way but sometimes the reciprocal
cross will give hybrids.
(2) Lilium regale regularly produces hybrid seed when pollinated with
the closely related L. sargentiae, but when pollen of most other species is
used the progeny is nearly always apomictic L. regale. In this situation the
breeder should not claim a hybrid until there is clear morphological or
cytological evidence (see Section 7. 7) to confirm the belief. If the situation
is known then it is wise to use only the pollen of the 'apomictic' species
in future crosses.
(3) Some Rubus species and most plants producing apomictic seeds
also produce some functional embryo sacs. In these cases deliberate cross
64 Plant Breeding

pollinations produce varying proportions of hybrids against the resultant

seedlings and the problem then is to decide which are true hybrids and
which arc apomicts, especially if the apomicts segregate for slight differ-
ences from the seed parent. In known instances of this type it is advisable
whenever possible to use a pollen parent with a dominant character not
occurring in the female parent so that hybrids can be recognised with
certainty by morphological differences. For example, apomictic seedlings
of the cut-leaved blackberry (Rubus laciniatus) all have the typical foliage
character, whereas sexual hybrids from this species do not and can be
recognised at an early stage of growth.

BARSON, D. M. and BALLINGER, R. ). (1944). Extraction of tomato, cucumber
and marrow seed with hydrochloric acid, Agriculture, 51,178-184
CRANE M. B. and MARKS, G. E. (1952). Pear-apple hybrids, Nature, Lond., 170,
DAVIS, A. I. S. ( 195 7). Successful crossing in the genus Lathyrus through stylar
amputation, Nature, Lond.,180,612
EMSWELLER, S. A. and UHRING, ). (1962). Lilium speciosum X L. auratum,
North Am. Lily Soc. Lily Ybk, 15,7-15
GORTER, C. ). (1955). In vitro culture of Cyclamen embryos, Proc. K. Ned. Akad.
Wet., 58, 377-385
GUZOWSKA, I. (1971 ). In vitro pollination of ovules and stigmas in several species,
Genet. pol., 12, 261-266
HESLOP HARRISON,). and HESLOP HARRISON, I. (1970). Evaluation of pollen
viability by enzymatically induced fluorescence; intracellular hydrolysis of
fluorescin diacetate, Stain Techno!., 45, 115-120
HEYDECKER, W. (1974). Small-scale seed storage,}. R. hort. Soc., 99,216-220
JENNINGS, D. L. and TULLOCH, B. M. M. (1965). Studies on factors which pro-
mote germination of raspberry seeds, j. exp. Bot., 16, 329-340
JOHANSEN, D. A. (1950). Plant Embryology, Chronica Botanica, Waltham, Mass.,
KANTA, K., RANGASWAMY, N. S. and MAHESHWARI, P. (1962). Test-tube
fertilisation in a flowering plant, Nature, Lond., 194, 1214-121 7
KASHA, K. ). and KAO, K. N. (1970). High frequency haploid production in barley
(Hordeum vulgare L.), Nature, Lond., 225, 874-875
LENZ, L. W. (1955). Studies in Iris embryo culture. 1: Germination of embryos of
subsection Hexapogon Be nth (Sect. Regal is Senan Dykes), Aliso, 3, 1 7 3-18 2
LEWIS, D. and CROWE, K. (1958). Unilateral interspecific incompatibility in flower-
ing plants, Heredity, 12, 233-256
MARTIN, F. W. (1959). Staining and observing pollen tubes in the style by means of
fluorescence,Stain Tech., 34,125-128
NORTON, I. D. (1966). Testing of plum pollen viability with tetrazolium salts, Proc.
Am. Soc. hort. Sci., 89,132-134
ROGGEN, H. and van DI)K, A. ). (1976). Thermally aided pollination: a new
method of breaking self-incompatibility in Brassica oleracea, L., Euphytica, 25,
THOMPSON, K. F. and TAYLOR, I. P. (1966). Non-linear dominance relationships
between S alleles, Heredity, 21, 345-362
TUKEY, H. B. (1934). Artificial culture methods for isolated embryos of deciduous
fruits, Proc. Am. Soc. hort. Sci., 32, 313-322
Fertilisation and Seed Development 65

WHITE, P. R. (1963). The Cultivation of Animal and Plant Cells (2nd edition),
Ronald Press, New York, 228 pp

FRANKEL, R. and GALUN, E. (1977). Pollination Mechanisms, Reproduction and
Plant Breeding, Springer·Verlag, Berlin, 281 pp
de NETTANCOURT, D. (1977). Incompatibility in Angiosperms, Springer-Verlag,
Berlin, 230 pp
RAGHAV AN, V. ( 1976). Experimental Embryogenesis in Vascular Plants, Academic
Press, London, 603 pp


We have seen in Chapter 1 that with a knowledge of the mendel ian principle
of units of inheritance and a chequerboard diagram it is possible to work
out expected segregations. The technique can be used by the breeder to
forecast the numbers of individuals of different types in a family from a
deliberate cross and also, in the reverse way, by observing the numbers
segregating, to form an opinion on the gene or genes governing the inherit-
ance of the factor or factors of interest. In this chapter we shall expand
on what was described earlier.
The monohybrid situation is fairly simple. Suppose that flower
colour is governed by a single dominant gene A for red and a for white,
then homozygous AA red x homozygous aa white will give all red (Aa)
F 1 progeny. If we were not aware that red is dominant and crossed hetero-
zygous Aa red with white, which is always homozygous aa, then we
would get 1:1 red:white individuals in the progeny. The same situation
arises of course when the F 1 is deliberately backcrossed to the homozygous
recessive parent.
When the F 1 (all Aa) from AA x aa is selfed, then in the F2 genera-
tion we get a segregation of 3:1 (one AA, two Aa and one aa). The 1:1
backcross to recessive and 3:1 F2 are the hallmarks of the single dominant
gene situation. When A is not fully dominant then Aa would be inter-
mediate light red and AA x aa would give all light red F 1 and a 1 :2:1 ratio
of red AA, light red Aa and white aa in F2 •


The dihybrid situation, dealing with two genes, is fairly straightforward

when there is no linkage and the genes do not interact with one another.
Let us consider a case in which we are interested in two hypothetical genes
A and B. It will then be possible for the reader to substitute genes for a
specific character in place of A and B. When the homozygotes AABB and
aabb are crossed, all the F 1 individuals are AaBb as is also the case when
AAbb is crossed with aaBB. The F 1 AaBb produces four types of gametes
in equal numbers AA, Ab, aB, ab; this gives a chequerboard diagram for
the F2 as in Figure 5.7.

Segregation and Combining Ability 67
.t.B Ab oB ob


Ab AABb AAbb AoBb Aobb

oB AoBB AoBb ooBB ooBb

ob AoBb Aobb ooBb oobb

Figure S.l Chequerboard diagram for F 2 segregation from the cross AABB X aobb

From the 4 x 4 = 16 combinations there are nine different genotypes

AA88 one Aa88 two aa88 one
AA8b two Aa8b four aa8b two
AAbb one Aabb two aabb one
In the common situation, as described in Chapter 1, when A and 8 are
completely dominant, it is not possible to distinguish individuals with Aa
from those with AA and/or 8b from 88 and we have a 9:3:3:1 ratio for
the 16 combinations. However, if only A is dominant, individuals with
Aa cannot be distinguished from those with AA but the 8b individuals are
phenotypically different from 88 so we have a 6:3:3:2:1:1 ratio for
the 16 combinations. If neither A or 8 is dominant, the situation is
When the Aa8b F 1 in which each of the genes shows full dominance
is backcrossed to the dominant AA88 parent, all the progeny are pheno-
typically like the F 1 , but when it is backcrossed to the recessive aabb a
1:1:1:1 ratio is obtained, namely double dominant, dominant A only,
dominant 8 only, fully recessive. Crossing the F 1 to AAbb and aa88 types
gives 1 :1 ratios with double dominant and dominant A or dominant 8
68 Plant Breeding

according to which of the two types is used in the cross.

The segregations discussed above hold only when the genes do not
interact, but frequently two genes may influence the same morphological
or physiological character and the typical 9:3:3:1 ratio of the F 1 dihybrid
F 2 is then modified. There are six main types of gene interaction which
can be worked out from basic principles by studying the 16 combinations
of the chequerboard diagram given in Figure 5.1.

5.2.1 Complementary genes (9:7 ratio)

The character controlled by two genes is only expressed when the geno-
type has at least one dominant allele of both A and B. It is not expressed
with the genotypes AAbb, Aabb, aaBb, aabb.

5.2.2 Additive genes (9:6:1 ratio)

Both genes in dominant alleles give more intense expression of the charac-
ter than when only one is present. Segregation is similar to above, but
only aabb gives no expression at all.

5.2.3 Suppressor genes (13:3 ratio)

Gene A is dominant but its expression is inhibited when 8 is dominant.
Thus, only the one AAbb and the two Aabb are expressed. Of the other
13 the aabb is homozygous recessive aa and therefore not expressed, and
expression of A in all the other combinations is suppressed by the 8 allele.

5.2.4 Dupiicate genes (15:1 ratio)

Both genes have similar expression, AAbb as aaBB and Aabb as aaBb. All
have the same phenotype, only the full recessive aabb has no expression.

5.2.5 Epistatic dominant genes (12:3:1 ratio)

Epistasis is a dominance of one factor over another, when the character is
controlled by two genes rather than alleles of a single gene. Gene A effect
dominates 8 effect, which is only expressed when no dominant A allele
is present. The 8 gene can then only be expressed in the one aaBB, two
aaBb and one aabb, but the latter combination is homozygous recessive
so 8 is not expressed.

5.2.6 Epistatic recessive genes (9:3:4 ratio)

Gene A effect dominates 8 effect but 8 intensifies A. There is no expres-
sion with aaBB, aaBb or aabb. Of the remaining 12, expression is intensified
in AABB, AABb, AaBB and AaBb but not in the other combinations.


Lethal genes are genes which in the homozygous state have such a dele-
terious effect on the individual that it cannot survive. Their effect may
influence segregation ratios. The lethal condition is usually recessive and
Segregation and Combining Ability 69

in extreme cases the embryo fails to develop so that the homozygous

recessive individuals are never seen in segregating families; only the hetero-
zygotes and homozygous dominants can grow. For a lethal gene L the II
types would die and only U and LL forms survive. Thus, from U x LL
all individuals would survive but from L/ x U the II would die giving a
segregation of 2L/:1 LL which would all be dominant phenotypes instead
of the usual 3:1 ratio. In rare cases where the lethal gene is incompletely
dominant and only the LL genotypes die then the segregation would be
2U: 11/ with two-thirds of the plants being heterozygous debilitated types.
Sometimes genes are described as semilethal when they have a serious
deleterious or crippling effect but do not kill individuals in the homozygous
dominant condition. Fully dominant lethal genes cannot survive as not
only the LL but also the U forms die and the L allele is eliminated.
Probably the most common type of lethal condition is the chlorophyll-
deficient albino. Pale-coloured or yellow individuals can be found amongst
seedlings of families from many different species. These may die at an
early age, continue to grow very slowly, or recover and then grow norm-
ally, according to their genetic constitution. Sometimes the deleterious
alleles are only expressed in certain cytoplasms.


Occasionally pollen tubes of different genotypes may grow down the style
at different rates. The faster growing ones may then reach the ovules first
so that the potential hybrid combinations from the slower growing pollen
genotypes are reduced in number and the segregation ratio affected. This
condition, known as certation, has been observed in Datura, Gossypium
(cotton), Nicotiana, Me/andrium, Oenothera and Zea (corn). The type of
situation which can arise is that whereas A a x aa may give the normal 1:1
segregation, AA x Aa should also segregate 1:1 but in fact gives very few
Aa heterozygotes. This is because the haploid a pollen is able to grow at
the same rate as the A pollen in Aa styles, but its growth is slowed down in
AA styles whereas that of A pollen is not. Relatively few of the a gametes
reach an egg cell which has not already been fertilised by an A gamete so
that relatively few Aa zygotes are formed.


Having recorded certain characters of individuals in a segregating family,
the breeder may wish to know the genetic basis for the segregation so that
results of future crosses can be forecasted. If the crop is one like tomatoes
or peas which have been studied extensively then the mode of inheritance
may already be known and the results can be attributed to a certain gene
or genes, but for many crops the breeder will be working in the dark. On
examining the figures it well may be possible to -;uggest that a certain
character is controlled by a single dominant gene, giving a 3:1 segregation,
70 Plant Breeding

or that there is a more complicated situation of a dihybrid segregation of

two interacting genes. To test whether the results are a good fit-that is
whether the hypothesis falls within the accepted degree of probability
and is not entirely due to chance-a chi squared (x 2 ) estimate should be
difference between number observcd)2
2 ( and number expected
X =
number expected
To illustrate this an example is taken of segregation for glossy and
normal leaved Brussels sprout plants. In an F 2 from a normal x glossy
cross the plants segregated 206 and 77 for the two categories respectively,
suggesting a 3:1 ratio indicating that the glossy condition was controlled
by a single recessive gene. We can then work out x2 as follows:-
Class Number Number Deviation x2
observed expected
Normal 206 212.25 -6.25 0.184
Glossy 77 70.75 +6.25 0.552
Total 283 283.00 0.00 0.736
The numbers expected are calculated as 3/4 x 283 for normal and 1/4 x
283 for glossy. The total 2 can then be tested to find whether it deviates
significantly from the hypothesis that the segregation is 3:1. To do this we
need also the degree of freedom which is one less than the total number of
classes tested. In this case there arc two classes, normal and glossy, and
there is only one degree of freedom. With a 9:3:3:1 segregation we would
have four classes with four separate estimates of x2 and there could be
4- 1 = 3 degrees of freedom.

Table 5.1

t values for P = 0.05 (from Fisher and Yates, 1974}

Degrees of p = 0.05 Degrees of p = 0.05

freedom freedom

1 3.841 11 19.675
2 5.991 12 21.026
3 7.815 13 22.362
4 9.488 14 23.685
5 11.070 15 24.996
6 12.592 16 26.296
7 14.067 17 27.587
8 15.507 18 28.869
9 16.919 19 30.144
10 18.307 20 31.410
Segregation and Combining Ability 71

In Table 5. 7 the probability value of P = 0.05 is given for various

degrees of freedom and if the x2 is less than the given figure this indicates
that the figures do not deviate significantly from the hypothesis and that
they are a good fit and not due solely to chance.
We see that a 2 of 0. 736 for one degree of freedom is less than 3.841
so we are justified in assuming that the segregation is 3:1 and therefore
that the glossy condition is a single gene recessive condition. We might
have thought that the figures could fit a 13:3 ratio but x2 calculated for
this expectation works out to 13.293 also with one degree of freedom.
This figure is greater than 3.841 and is therefore not a good fit. If 2 isx
calculated for different ratios it is the lowest estimate of x2 which indicates
the hypothesis that best fits the observed figures.
We have seen an example of 2 calculated from one family only, but
x2 values for several families can be added together to obtain a total value
with an equivalent total number of degrees of freedom. The results from
five separate backcross glossy x normal sprout families are given below
(the first family being the same as the one given in the earlier example).

Family Normal Glossy x2
1 206 77 0.736
2 187 88 7.185
3 188 63 0.010
4 218 66 0.469
5 172 67 1.172
Total 9.572
Now we have a total x2 of 9.572 with five degrees of freedom which is less
than 11.070 and therefore a good fit to the 3:i ratio. We can also obtain
an estimate of 2 from the sum of the total observed normals and glossies
for all families, 971 and 361 respectively. This gives 3.139 with one degree
of freedom since it concerns only two classes, normal and glossy, and
being less than 3.841 is also a good fit. Subtracting these two estimates
and their degrees of freedom gives a x2 for heterogeneity, that is a portion
concerned with the disagreement among the groups. In this case, x2 for
heterogeneity is 6.433 with four degrees of freedom and less than 9.488. It
therefore shows that all the families agree in general with the hypothesis,
in spite of the fact that the individual 2 from family 2 does not alone fit
the hypothesis of 3:1 ratio.
In discussing segregation ratios in this chapter we have assumed that
the genes are inherited independently but linkage, like certation and lethal
factors, can alter these ratios. It is possible to use x2 calculations to detect
linkage and also to calculate intensity by techniques described by Mather
(1951 ). However, it is not often that the practical breeder will need to
72 Plant Breeding

use such calculations, although they can be helpful in some cases. For
example, let us suppose that we have male sterility controlled by a single
recessive gene, the expression of which is not influenced by the cytoplasm,
and that we want to produce only male sterile flowering plants. We cannot
self-fertilise such plants because they produce no pollen, so we can only
perpetuate them from seed by intercrossing the heterozygotes, which are
fertile, or by pollinating the male sterility homozygotes with pollen from
the fertile heterozygotes. This would give us segregations of 1 :3 and 1 :1
male sterile and male fertile plants respectively. If we could find a closely
linked foliage character then it would be possible to select many or most
of the male fertile plants before they flowered. Calculations on linkage
intensity would be very helpful in this situation to decide whether the
selection was worthwhile, though this could of course be judged in an
empirical way by repeatedly making the crosses and observing the results.


When a cross is made to recover a certain genotype, or to obtain a new

gene combination, it is frequently helpful to know how many plants one
should grow to have a reasonable chance of achieving one's objective.
Table 5.2 gives the number of plants which have to be grown to be 95 or
99 per cent sure of obtaining at least one of the required genotype. For
example, to obtain at least one double recessive in a dihybrid F 2 it would
be necessary to grow at least 46 individual seedlings for a 95 per cent
chance and 71 for a 99 per cent chance and for a triple recessive in a tri-
hybrid 191 and 296 respectively.

Table 5.2
Number of plants to grow to obtain at least one
plant of the required genotype

Ratio of required Level of probability

genotype to total 95% 99%

1:2 5 7
1:3 8 12
1:4 11 16
1:8 22 35
1:9 25 39
1:16 46 71
1:27 79 122
1:32 95 146
1:64 191 296


Frequently the breeder will be dealing with segregations which are much
more complex than the simple mono- or di-hybrid situations we have
Segregation and Combining Ability 73

been discussing. The characters of interest may have polygenic inheritance,

that is they may be controlled by more than two genes and sometimes
many genes. There may be multiple allele control-not merely two alterna-
tive alleles to a gene but many. The plants may be polyploids. This is not
to say that the breeder is never faced with the simpler segregations; many
features of major importance such as disease resistance, flower colour,
male sterility and the presence of thorns or hairs are usually controlled
by one or two genes only. But the inheritance of features such as yield,
fruit quality and vigour is usually complex and crosses do not give a
simple or clearly defined segregation ratio.
The polygenic situation with three or more genes can be worked out
by the chequerboard diagram but becomes very complex when inter-
actions are taken into account. In practical situations it is often desirable
to know how often the fully recessive condition or the fully dominant
condition may occur with three or more genes. The situation for independ-
ent genes, each sharing full dominance, is shown in Table 5.3.
Table 5.3

Basic segregations for 7-7 0 genes

Number of Number of Possible Number of Number of

genes different combinations dominant genetically
gametes in F, phenotypes different
in F, in F 2 classes

1 2 4 2 3
2 4 16 4 9
3 8 64 8 27
4 16 256 16 81
5 32 1,024 32 243
6 64 4,096 64 729
7 128 16,284 128 2,187
8 256 65,536 256 6,561
9 512 262,144 512 19,683
10 1,024 1,048,576 1,024 59,049

n 2n (2n)' 2n 3n

The possible combinations in the F2 generation corresponds to the number

of squares in the chequerboard diagram, and the full recessive occurs as
only one of these combinations, namely 3:1, 9:3:3:1, 27:9:9:9:3:3:3:1
when considering one, two and three genes respectively. If for example we
are considering five genes, then 32 full dominant and only one full recess-
sive would occur for every 1,024 plants grown in the F2 • To obtain full
recessives it would be profitable to self some of the F2 individuals which
showed the fewest dominant characters; these would then segregate a
higher proportion of recessives in the F3 generation than in the F2 . Thus
we see that inbreeding by selfing helps to select recessive types.
74 Plant Breeding

Complex segregations can arise with polyploids where there are more
than two of each chromosome type. In this situation it is possible for a
plant to carry more than two alleles of each gene and we can understand
the complication of gene dominance and interaction in such situations.


When plants are inbred through several generations of selfing or by sib

crossing (that is brother/sister mating) the progenies become more uniform
but frequently they suffer inbreeding depression. The inbred generations
are less vigorous and, from a horticultural point of view, less productive
than the original parent materia •. There may be more than one explanation
for this condition but frequently it is related to the accumulated segrega-
tion of deleterious or even semilethal genes resulting from inbreeding.
These do not have the same effect in the original parents because they are
in the heterozygous condition. The degree of inbreeding depression varies
according to the species; peas and French beans show very little loss of
vigour on inbreeding and, at the other extreme, Brussels sprouts rapidly
lose vigour after two or three generations of inbreeding. The loss of general
vigour may be accompanied by impaired reproductive fertility resulting in
poor fruit and seed set.
When plants are crossed with unrelated or distantly related individuals
the resulting progeny frequently shows heterosis, commonly referred to as
hybrid vigour. The hybrid individuals are more vigorous and frequently
earlier-maturing than either of the parents, a feature which was noted even
in 1790 by Kolreuter (see Section 1.2). However, very wide crosses, as
between different species or even different genera, sometimes may be
weak growing; the result depends very much on the combination. Nowa-
days there is much interest in heterosis for the production of hybrid
cultivars of many seed-propagated horticultural crops. These are not only
more productive, they are also more uniform than the older conventional
cultivars, and they will be discussed in more de.tail in Chapter 8.


When breeding for yield potential, or in high ploidy crops such as the
octaploid strawberry or tetraploid potato even for simpler characters such
as fruit or tuber skin-colour, the choice of parent material is not a clear-cut
decision because of the complexity of the inheritance. It is obviously help-
ful to the breeder if the plants to be used as parents can be assessed for
their capacity to transmit certain characters to their progenies. In most
cases there will be no one potential parent which is greatly superior to the
other; the choice is partly subjective even when a numerical assessment
is used.
Segregation and Combining Ability 75

The potential parents are first assessed for the characters of interest,
if necessary by subjective scoring on a basis of 1-3, 1-5, or 1-10. In these
cases it is customary to give the highest score for the most desirable
character. The plants are then pair-crossed if practicable in all possible
combinations to give a dial/el cross, coloquially referred to as a 'diallel'.
Often it is impossible to obtain a complete diallel, but a range of crosses
giving an incomplete diallel is also useful. The mathematical analysis of
the results is somewhat complex and it is advisable to obtain advice and
help from a biometrician before the crosses are planned and to have access
to a computer, especially for calculations on an incomplete diallel. Those
who wish to study the subject will find the mathematical aspect described
by Mather and Jinks (1971 ).
The results will give a measurement of combining ability-that is the
relative ability to transmit specific characters in crosses. There are two
aspects-general combining ability which applies to all crosses, and specific
combining ability which refers to certain specific crosses only. In most
cases the parents will show a degree of general combining ability; cases of
specific combining ability are less common.


Dioecious species produce two kinds of plants, male and female. As one
might expect, the families produced by crossing these two types are
approximately 50 per cent males and 50 per cent females. In most dio-
ecious species the plants have specialised sex chromosomes which carry
only those genes controlling sex expression. Maleness or femaleness is then
inherited independently of other characters. Occasionally the chromosomes
may carry a few other genes controlling characters which are always associ-
ated with either the male or female and said to be sex linked characters.
The usual situation with dioecious flowering plants, such as asparagus
and spinach, is that there are two types of sex chromosome, namely X
and Y. Homozygous XX plants are female and those which are hetero-
zygous XY are male. Crossing XX and YY plants will always segregate
1:1, as with a monohybrid backcross. Sometimes modifying genes are
present and these may induce the production of some hermaphrodite
flowers which allows for selfing, then XX females give all female progenies
and XY males produce 3:1 male:female plants, including some homo-
zygous YY males as with so-called 'super male' asparagus plants.
The heterozygous male condition is by far the most common in angio-
sperms but the situation is reversed in the wood strawberry Fragaria
elatior where XY plants are female. A similar XY sex control occurs with
animals, and in mammals as with plants, the XY is male; but in birds,
butterflies and fishes it is the female which is heterozygous.
76 Plant Breeding

Polyploidy can affect sex expression in dioecious plants, especially

when there are unequal complements of X and XY chromosomes as in
triploids and pentaploids. However, these situations are largely of academic
interest and unlikely to be encountered by the practical breeder.

FISHER, R. A., and YATES, F. (1974} Statistical Tables for Biological, Agricultural
and Medical Research, (6th Edition}, Longman Group, London (Previously
published by Oliver and Boyd, Edinburgh}
MATHER, K., and )INKS,). L. (1971}. Biometrical Genetics, Chapman and Hall,
London, 382 pp
MATHER, K. (1951}. The Measurement of Linkage in Heredity, Methuen, London,
149 pp

The structure of chromosomes and the genes they carry is very consistent
otherwise there would not be continuity in inheritance. However, changes
to chromosomes do occasionally occur. Sometimes individual genes change
to give new alleles and there may be larger alterations in structure. Chromo-
somes may break and pieces may be lost (deletion), pieces may break off
and be rejoined in the reverse order (inversion), or a portion of one
chromosome may be transferred to another chromosome (translocation).
The way such changes can take place and the pairing of altered chromo-
somes is illustrated in Figure 6.1. Inversions and translocations in which
the positions of chromosome portions are altered, but in which the gene
complement remains the same, can still have a heritable effect because
the chemical compounds produced by genes to influence development
interact with those of their neighbours. Linkages are also altered. These
changes, whether they involve individual genes or whole chromosomes,
introduce new heritable characters known as mutations. Many drastic
chromosome mutations are lethal and the cells containing them cannot
survive, but smaller changes often give the individuals carrying them an
opportunity to compete with more normal progeny and sometimes to
gain ascendency and to develop as new forms better adapted to the
When a gene mutates to a new allele, the change is usually recessive,
although occasionally it may be dominant like the Ll gene in raspberry
(Jennings, 1961 ). The mutant has larger fruit and considerably enlarged
calyx segments than the normal recessive form. When crossed with normal
forms the F 1 progenies always carry a proportion of plants with large
calyces and many of these are large fruited individuals, although fruit size
may not be increased with some genetic combinations because of reduced
fertility which results in poor drupelet set.
Every observant gardener knows that a cultivar of rose or dahlia or
other vegetatively propagated plant sometimes may produce a branch
with flowers of a different colour or double flowers. These new forms,
known in horticultural circles as 'sports', are mutations and often single
gene changes. When the flowers on the mutated branch are used in hybridi-
sation the progenies usually segregdte differently from those from similar
combinations made by crosses with flowers of the normal form. Many
cultivars of trees and shrubs with deeply indented laminas or weeping or

78 Plant Breeding

a a

D d~ b


a a a a
b b b b

I c c d
d c

a a a f
b b c

T g d

Figure 6.1 Dia gra m illu str ati ng

ho w ch rom oso me s
D, del eti on ; I, inv ers ma y un der go str uct
ion ; T, tra nsl oca tio ura l cha ng es.
Mutations 79

twisted branches have originated also in this way. Sometimes isolated

somatic cells mutate to give patterned areas of different petal or leaf colour
and these new chimceras can be maintained by careful vegetative propagation.
The effects of mutations that give rise to a change in flower colour
are fairly readily observed but mutations of minor genes which may have
an additive effect, for example on leaf shape or yield, are often undetected
by the plantsman. Their effects can accumulate and cultivars maintained
vegetatively as clones over a long period may be found to differ slightly
from one another. However, these differences are often masked by virus
infection and can only be observed when virus-free stocks are available.


Mutations provide a valuable source of variation of plant material from

which the breeder can make selections. Natural mutation rates may be
very slow and it is obviously advantageous if they can be speeded up to
give more variation. The most effective way to achieve this is to subject
plant material to ionising radiation such as X rays and gamma rays, but
ultraviolet light and chemical mutagens have been used successfully. The
mutagenic effects of X rays were first described by Muller (1927) and the
effects of radiation in general have been described by Catcheside (1948).
From a practical point of view X rays are probably the easiest avail-
able type of radiation to the plant breeder. Their use requires treatment
of material in a standard deep-therapy X-ray appartus by a specially
trained operator, but the cost of this is generally small compared with
gamma-ray treatment. Gamma rays are short-wave X rays generally obtain-
ed by radiation from a cobalt-60 source. This usually requires a gamma
field, a circular area of cultivated land surrounded with a thick radiation-
proof concrete wall and a cobalt-60 rod at the centre arranged so that it
can be raised from and lowered into a lead-lined radiation-proof container
without exposing the operator. Plants grown in the gamma field receive
different quantities of irradiation according to the square of their distance
from the source. Chemical mutagens are much less effective than radiation
treatments though they are cheap and simple to apply. The most com-
monly tried chemical is ethyl methane sulphonate (EMS).
The optimal dose of X rays depends on the species and the type of
material being treated, but 1,000-5,000 rad is most likely to be effective
with pieces of stem, root and tuber. Higher doses may be optional with
seeds. Rate of application is generally thought to be less important than
total rad treatment and about 300 rad per min is usual. However, recent
work suggests that lower application rates may reduce the number of
lethal mutations. Most induced mutations are recessive and since they
often involve a change to one chromosome only of a pair they may not be
observed directly. However, when selfed seed is produced from irradiated
plants, some seedlings homozygous for the mutated character will segre-
80 Plant Breeding

gate in the so-called X 1 generation. Nevertheless, some tissues show

mutations directly without the need to grow on seedlings especially when
the mutation is in a series of alleles without pronounced dominance.
Radiation treatment does not, of course, affect every cell of the
material in the same way, so that only parts of the plant may be mutated
and a sectorial chim.era is then produced. Tissues which form adventitious
shoots arising from single cells are especially suitable for radiation treat-
ment as entire shoots may then grow from one mutated cell. Broertjes
(1968) described over 350 species which form adventitious buds and
discussed their potential for mutation breeding. Polyploids are especially
suitable for treatment as damage to one chromosome of a set of three
or more is less likely to prove a serious disadvantage to a plant than when
there are only two replicates of each chromosome.
The results of induced mutation in plant breeding have fallen short of
the early enthusiastic claims and to date only about 100 cultivars have
been produced in this way. Most important economically are the short-
strawed macaroni wheats. Of special interest to horticulturists are induced
fruit colour changes in apples (Bishop, 1959), compact mutants in apple
and pear (Visser eta/. 1971 ), upright habit in black currant (Bauer, 1974)
and flower colour changes in several ornamentals (Broertjes eta/. 1968).

6.2.1 X-ray induction of flower colour mutants

This technique is well proven and effective for several vegetatively propa-
gated florists' crops and has been used successfully with Achimenes,
Alstromeria, Begonia, Chrysanthemum, Dahlia, Kalanchoe, Saintpaulia
and Streptocarpus. The effect of irradiation is generally to cause a muta-
tion of the gene or genes controlling flower colour to more recessive alleles.
It has therefore been suggested that the breeder should concentrate his
efforts to obtain the desired flower form and plant habit with flowers of
the most dominant colour type. When this has been achieved, a series of
cultivars of different flower colours can be produced by radiation, all
with the other desirable characters. The technique for treating chrysanthe-
mum cuttings is described by Broertjes (1966) and well illustrated with
coloured plates of the mutants. With this species the form of the petals
also was sometimes altered.
A recent development is the use of dithiothreitol to prevent extreme
and often lethal chromosome breakages and thus to increase the numbers
of treated plants with less drastic and potentially more useful mutations
The plant material is soaked in a 0.5 per cent aqueous solution for two
hours before irradiation (Broertjes, 1976).


It is well established that the natural mutation rate increases in ageing seed
of many species. The situation is reviewed by d'Amato and Hoffmann-
Ostenhof (1956) who quote effects in several plants of horticultural
Mutations 81

interest including Allium species, Antirrhinum majus, Lactuca sativa and

Pisum sativum. In Pisum sativum species a five- to six-fold increase in
mutations was reported for four-year-old seed. The increase in most other
species was lower; for example, the normal mutation rate in Antirrhinum
of one per cent was increased to 1.6-5.3 per cent for six to nine-year-old
seed. The cause of increased mutation in ageing seed is not fully under-
stood but it seems likely that it may be associated with the production by
the seed itself of certain metabolic products which are natural mutagens.

d'AMATO, F. and HOFFMANN-OSTENHOF, 0. (1956). Metabolism and spon-
taneous mutations in plants, Adv. Genet., 8, 1-28
BAUER, R. (1974). Westra, an X-ray induced erect-growing black currant variety,
and its use in breeding, in Polyploidy and Induced Mutation, International
Atomic Energy Agency, Vienna, 13-20
BROERT]ES, C. (1966). Mutation breeding of chrysanthemums, Euphytica, 15,
BROERT]ES, C., HACCIUS, B., and WEIDLICH, S. (1968). Adventitious bud
formation on isolated leaves and its significance for mutation breeding, Euphy-
tica, 17,321-344
BROERT]ES, C. (1976). Mutation breeding in vegetatively propagated floricultural
crops, Acta. hart., 63,187-195
BISHOP, C. ]. (1959). Radiation-induced fruit colour mutants in apples, Can./. Gen.
Cyt., 1,118-123
CATCHESIDE, D. G. (1948). Genetic effects of radiations, Adv. Genet., 2, 271-358
JENNINGS, D. L. (1961). Mutation for larger fruit in the raspberry, Nature, Land.,
MULLER, H.]. (1927). Artificial transmutation of the gene, Science, 66,84-87
VISSER, T., VERHAEGH, ]. ]. and de VRIES, D.P. (1971). Pre-selection of com-
pact mutants induced by X-ray treatment in apple and pear, Euphytica, 20,

HAGBERG, A. and .li.KERBERG, E. (1962). Mutations and Polyploidy in Plant
Breeding, Heinemann, London, 149 pp


Plant varieties which have originated under cultivation are called cultivars
(singular abbreviation cv.). They are of two principal types: (1) those
which are mainly propagated vegetatively by division, cuttings, grafting,
budding and nowadays sometimes also by rapid in vitro techniques, and
(2) those which are mainly propagated by seed. Most perennial horticul-
tural plants fall into the first category and annuals and biennials into the
second. However, a few perennial species such as Anemone coronaria,
Cyclamen and runner bean, all of which produce storage organs and can
be grown as perennials, are almost invariably seed propagated. The usual
reasons for resorting to seed propagation of perennials are the difficulty
of rapid vegetative propagation, the short-lived nature of plants of the
particular species, and disease problems. A few perennial ornamentals,
including Begonia, Delphinium, Lilium and certain Lupinus, Pelargonium,
Primula and Viola species are regularly propagated both by seed and
vegetatively. To a lesser extent this also applies to certain vegetable species
such as asparagus, chicory and seakale.
Each new vegetatively propagated cultivar comprises a clone of
genetically identical individuals, but in time small undetected mutations
may alter the genetic constitution of batches of the material so that several
different clones become embraced under the same cultivar name. Batches
of plants of a clone may also become differentially affected by viruses
which cause disease and affect performance, and it is not always possible
to attribute batch differences solely to disease or to accumulation of
The approach to the breeding of vegetatively propagated plants is
somewhat different to that required for those which are seed propagated.
In certain respects it is easier to achieve success with the former group
because once a good genetic combination has been obtained it can be
propagated immediately as a new cultivar, although sterility or incom-
patibility problems may have to be overcome when the desired end product
is a fruit or a seed such as a nut. With seed propagated plants it is necessary
to 'fix' the cultivar so that it breeds true, at least within certain prescribed
limits. As we shall discuss later, this may be a lengthy and difficult process.
Vegetatively Propagated Cultivars 83


Vegetatively propagated cultivars nevertheless have their special problems;

many of them arc especially troubled by disease. Vegetative propagation
often carries over disease to the new plants whereas seed propagation
usually screens out pathogens, most of which are not seed-borne. Virus
diseases especially are often difficult to eliminate from clones and many
good named clonal cultivars have disappeared through a build-up of
deleterious viruses or mycoplasmas. Nowadays it is possible, in many
cases, to eliminate viruses from clones by techniques which will be describ-
ed later.
Another troublesome aspect of diseases within clonal cultivars relates
to their uniform genetic constitution. A slight mutation of a disease
organism can result in the production of a highly virulent strain which
may even kill all the plants. Most seed-propagated cultivars are hetero-
geneous in many characters, including resistance to pathogens, and
individuals within a crop react differently to attack so that high propor-
tion of the plants escapes serious damage. However, the tendency nowa-
days is to use breeding techniques to produce seed propagated cultivars
which are much more homogeneous and this advantage over clonal propa-
gated cultivars is becoming less pronounced.
Unlike annuals and biennials, many perennial species take more than
one or two years to flower from seed. Some bulb crops such as daffodils
and tulips require six to eight years of growth before they flower, and
many fruit trees even longer. It may then be very important to the breeder,
for economic reasons, to shorten the generation time, especially when
the plant units are trees or bushes which each occupy a considerable area
of land or glasshouse space. Some techniques for hastening flowering are
described briefly in Section 3.1 0. It is also desirable to obtain an early
assessment of the flower or fruit qualities so that a large proportion of
the newly raised plants can be eliminated at an early stage to avoid the
cost of growing them to maturity. In some cases seedling characters give
an early indication of features of the mature plants. Raspberries with
thornless canes are desirable because thorns provide lodging for pathogenic
fungi and render the plants more liable to disease. Thornless types can be
recognised shortly after the emergence of seedlings by the absence of the
typical glands on the cotyledons.
Another problem in the breeding of some perennial crops is the slow
rate of multiplication of new cultivars or of disease-free stocks of establish-
ed cultivars. A few crops multiply vegetatively fairly rapidly by natural
means, such as runners (strawberry) or tubers (potato). Others can be
multiplied at a satisfactory rate by long-established horticultural tech-
niques such as stem cuttings (black currant, chrysanthemum, dahlia), root
cuttings (raspberry, sea kale), budding and grafting (tree fruits, roses,
rhododendrons). However, there are several crops like orchids and certain
bulb-forming species whose clones can be multiplied vegetatively only
84 Plant Breeding

very slowly by standard horticultural techniques, and benefit from the

application of recently developed manipulative treatments for rapid
propagation. These techniques which also can, in some circumstances, be
economically desirable for the relatively easily propagated species such
as the strawberry are described under the individual crops, and the more
complex in vitro techniques are described in Section 9.9.
We shall now consider the techniques and objectives in the breeding
of the more important temperate horticultural crop species. It is only
possible here to give an outline of procedures and any breeder wishing to
work on a specific crop should study the literature in some detail before


The cost of hand-harvesting soft fruit crops is 40-60 per cent of the total
cost of production and it is therefore not surprising that during the last
decade much effort has been put into devising ways to reduce costs by
machine harvesting. Suitable machines have been developed, and are used
extensively, for commercial harvesting of black currants and blueberries.
Means for mechanical harvesting of most other fruit crops need improve-
ment before they become standard practice, and the development of
suitable machines and the breeding of cultivars adapted to machine harvest-
ing arc interdependent. Breeders of new cultivars of soft fruits for com-
mercial production must keep ih touch with the latest developments in the
techniques for machine harvesting.

7.3.1 Blackberry and hybrid berries

Blackberries or brambles are, like raspberries, members ofthe genus Rubus,
but they differ from them in that the torus or 'plug' remains in the ripe
fruit-a feature which usually makes it easier to harvest fruit mechanically.
There are some 100-130 species of blackberry but the group is complicat-
ed by the frequent occurrence of localised populations of plants spreading
by apomictic seed production. Systematists disagree as to whether these
groups are separate species and subspecies or merely clones of a particular
genotype. Apomixis also makes it difficult for the breeder who may not
be sure whether seedlings from planned crosses are genuine hybrids. The
group includes species with a wide range of ploidy levels from diploids to
dodecaploids (12x ).
The main objective in blackberry breeding is to obtain types which
produce thorn-free canes with an easily manageable upright growth habit.
Hardiness and early fruit ripening are important but unfortunately some
of the available plant material with other useful characters lack these
Genes for thornlessness are dominant in 'Austin Thornless' (8x) and
recessive in 'Merton Thornless' (4x ). Thornlessness in the latter is associat-
ed with cotyledons lacking glands so that seedlings can be separated for
Vegetatively Propagated Cultivars 85

this character at a very early stage of growth and this is especially helpful
to the breeder since F2 families usually give tetraploid segregation of only
one thornless to 35 thorny. The 'Merton Thornless' source has been used
in the development of the cultivars 'Smoothstem' and 'Thornfrec'.
Some thornless mutants are chim<eras in which the gene for the thorn-
less character is confined to the outer cell layers so these types do not
transmit the character to their progeny.
Crosses between the blackberry and raspberry groups of Rubus have
produced some useful hybrid berries, notably the loganberry, whose exact
origin is obscure. Its hybrids with some blackberries arc fertile but those
with raspberries sterile. 'Phenomenal' is a loganberry x blackberry hybrid
and from this the 'Youngbcrry' was produced. The 'Boysenberry' is
similar but of unknown parentage. British hybrid berries of this type
include the 'Vcitchberry' and the recently introduced 'Tayberry'.
Blackberries are mainly propagated by tip layering or by leaf bud
cuttings and these techniques arc relatively expensive. The 'Tayberry'
can be propagated like raspberries from root cuttings and this capacity
would be a useful characteristic to breed into other blackberry and hybrid
berry types. Although this group has its difficulties for the breeder, it
seems of offer considerable scope for further improvements.

7.3.2 Black currant

Most black currant cultivars grown in Britain are forms of Ribes nigrum, a
species native to Europe and parts of Asia. During the last 20 years several
other species (all diploids, 2n = 16}, notably R. bracteosum, dikuscha,
hudsonianum, ussuriense, have been crossed with R. nigrum to give more
or less fertile hybrids. Other (2n = 16} species which hybridise with the
black currant, but give sterile or only partially fertile progeny, include the
gooseberry (R. grossu/aria}. red currant (R. rubrum} and flowering currant
(R. sanguineum}.
One of the major problems of commercial black currant production,
and one which prevents the crop becoming more popular, is irreglar crop-
ping. Flowers are damaged by late frosts and, even when this does not
occur, seed set may be poor so that development is hindered and a high
proportion of fruit is shed before it ripens. Ribes bracteosum, which is late
flowering, has been crossed with black currant and some of the resultant
progenies escape spring frost damage, but the fruits tend to lack the
characteristic black currant flavour. Most late flowering selections of
R. nigrum itself do not suffer from this disadvantage and arc being used
successfully in breeding. Some R. nigrum x hudsonianum hybrids seem
to have the capacity for continued fruit development without early
abscission, even when few seeds are set, and they have proved valuable
in breeding to combat the problem known to fruit growers as 'running
off' -that is, successive fruit shedding. However, since most wild forms of
Ribes are self incompatible it is necessary to select for self compatibility
86 Plant Breeding

amongst the hybrid populations with all species crosses. Most existing
cultivars are highly self compatible-an essential feature when crops of a
single cultivar are grown.
Mechanical harvesting requires bushes with a fairly upright habit, but
selecting for this characteristic tends to be accompanied by a reduction in
yield potential. However, a fastigiate growing cultivar 'Westra' obtained
as an X-ray mutant from 'Westwick Choice' {see Section 6.2), when used
as a parent confers upright habit without a pronounced loss in yield.
Black currants are sometimes devastated by American gooseberry
mildew and leaf spot fungi, reversion virus and black currant gall mite. All
these troubles can be controlled by modern spray regimens but it would
be preferable for economic and environmental reasons to control them by
using resistant cultivars.
Ribes sanguineum is fully resistant to American gooseberry mildew
caused by Sphaerotheca mors-uvae, but hybrids between it and black
currant are sterile and breeders have found it difficult to make use of this
source of resistance. A fair degree of resistance has been obtained from
some Scandinavian cultivars such as 'Brodtorp' which are said to be
heterozygous for a dominant resistant gene M. Laboratory and glasshouse
tests for susceptibility are not wholly reliable and resistance is usually
assessed by field performance.
Resistance to leaf spot caused by the fungus Pseudopeziza ribes, that
induces premature leaf fall, has been obtained from the Russian wild forms
and hybrids of Ribes nigrum sibiricum and R. dikuscha.
The black currant gall mite Cercidophyopsis ribis, formerly known by
the name big bud mite which clearly describes the symptoms it induces, is
the only known vector of the agent which causes reversion disease that can
seriously depress yield. The gooseberry is immune to the mite but hybrids
between it and black currant are sterile. Fertility can be restored by
colchicine-induced polyploidy and this material is being used as a source
for breeding resistant black currants by back-crossing. ·Recently some
Russian cultivars, especially those of R. nigrum sibiricum parentage
(Anderson, 1971 ), have been found to confer mite resistance; they are
fully fertile with existing European cultivars.
Most commercial black currant crops are used for juice production
and it is important that new cultivars should be capable of producing
fruits which have the colour, flavour and vitamin C content which make
them acceptable for this purpose. The assessment and selection for these
qualities is an important aspect of modern black current breeding projects.
Black currant pollen does not remain viable for long but can be
stored under dry conditions for one or two months. Seed is extracted
easily in a blender and if sown soon after extraction it germinates readily.
Seedlings fruit first when about three years old but cannot be fully assessed
before six years.
Black currants are usually propagated from hardwood cuttings about
20 em long. Propagation of a new selection can be done more rapidly by
Vegetatively Propagated Cultivars 87

softwood cuttings in, May-July or single-bud cuttings in slight heat in

February to give respectively some 200-400 and 500-1,000 new plants
from a mature bush.

7.3.3 Blueberries
These are species of the genus Vaccinium which also includes cranberries.
There are two main types of blueberry, the 'high bush' which grows at
least 1 metre tall, and the 'low bush' which is a dwarf shrub similar in
habit to the native British bilberry or blaeberry ( Vaccinium myrtil!is).
Several cultivars of high bush blueberry have been bred and are grown
extensively in the USA, (see Eck and Childers, 1966) but most low bush
blueberry crops are stands of wild plants managed in their natural habitat.
Very little breeding work has been done with blueberries in Europe,
although hybridising American species with V. myrtillis has been achieved
in Germany.
American high bush blueberry cultivars have received some attention
as a crop in Britain during the last decade. The fruit matures later in the
season than most other soft fruits, can be harvested mechanically 01.••:
carries well without damage. However, the plants require acid soils, take
six to eight years to reach full cropping capacity, and are slow and expens-
ive to propagate. Earlier fruit ripening would be a useful breeding objective
for British conditions so that the crop could be grown in acid marginal
soils, which are mostly found in the colder areas of the country.
Many wild Vaccinium species are known and several are interfertile.
American species alone range from diploids to tetraploids and hexaploids
with a basic chromosome number of 12. The genus and closely allied
genera of Ericaceae occur also in Asia and would form an interesting group
for crossability studies with a view to breeding for European conditions.

7.3.4 Gooseberry
European gooseberry cultivars are derived from the European endemic
Ribes g!ossu!aria, but American types are mainly hybrids between this
species and R. hirtel!um. The main objectives in gooseberry breeding are
spinelessness and resistance to American gooseberry mildew, but some
other characters such as resistance to the sawfly Nematus ribesii have
been included in breeding projects.
Species and varieties of the gooseberry (Eugrossu!aria) group of
Ribes differ in their degree of spininess and some like R. hirtellum and
R. rotundifo!ium have very small spines, but none is completely spine-free.
Inheritance of spinelessness is complex and crosses between Eugrossularia
types have so far failed to produce spine-free cultivars. Crosses have been
achieved between gooseberry and spineless members of the currant group
of Ribes, notably black currant and R. sanguineum, and selections made
at tetrapolid level, since the diploid hybrids are sterile. In spite of these
efforts, no truly spine-free gooseberry cultivar has yet been introduced.
88 Plant Breeding

Most North American gooseberry species, notably R. hirtellum and

R. divaricatum, have a high degree of resistance to mildew and have been
used to produce resistant cultivars; sharp segregations are found in F 1
progenies suggesting simple major gene control. However, the European
gooseberry has much larger fruits than those of most other species and it
is difficult to retain this characteristic in progenies from interspecific
hybrids. Fruit size is important in Europe where the crop is usually grown
commercially for harvesting early as immature green fruit.
Sawfly resistant cultivars have not yet been achieved but a degree of
resistance is obtained from crosses between gooseberry and R. nigrum
and R. sanguineum.

7.3.5 Raspberry
Most European raspberry cultivars ongmate entirely from the native
Rubus idaeus, but some modern firm-fruited cultivars are derived from
hybrids between this and the American black raspberry R. occidenta/is.
The major objectives in raspberry breeding are suitability for
mechanical harvesting, and resistance to fungal diseases of the canes
and to virus diseases. Less important objectives include resistance to
aphids and to raspberry beetle.
The mechanised harvesting of raspberries is not yet a routine operation
on commercial crops in Britain, but extensive research has shown that
fruit for pulping can be harvested by machine. Fruits of commonly grown
cultivars. can be shaken from the canes by machine but they tend to break
apart on the collecting plates. Firm-fruited cultivars are required and this
is being achieved by selection from red x black raspberry cultivars. Both
are diploids (2n = 14) and intercross readily to give fertile hybrids, but the
progenies tend to lack winter hardiness and are rather susceptible to
viruses. Backcrossing the hybrid to red raspberry is necessary to overcome
these defects and to obtain fruits with the colour and flavour of the true
red raspberry. Mechanical harvesting also causes superficial damage to new
canes which will produce the crop for the following year. This damage
allows entry of fungi causing cane diseases and a .reduction in yield.
Inherent resistance to these pathogens is being sought.
Fruit should not only be firm, it must also have an acceptable rasp-
berry flavour, neither too sweet nor too acid, and the colour should be a
bright red without excessive purple tints or a dull appearance due to many
small hairs on the surface of the druplets. Cultivars with yellow or amber
coloured fruit are occasionally grown in gardens; this character is control-
led by a single gene t, and minor genes influence the intensity. Large fruit
size is associated with high overall yield and case of hand-picking, and is
preferred for private gardens. A major dominant gene L 1 (see Section 6.1)
confers large fruit size but the character is modified by minor genes.
Resistance to several diseases is assisted by spine-free, hairy canes, the
accepted explanation being that these features help to prevent entry by
Vegetatively Propagated Cultivars 89

germinating fungal spores. Each of these characters is controlled by a

single major gene; s spinelessness (pleiotropic with eglandular cotyledons,
see Section 1.4) and H hairiness. A degree of resistance to most of the
damaging virus diseases is obtained in Europe by resistance to the rasp-
berry aphid, Amphorophora rubi, of which four strains are known. Several
dominant genes confer resistance, notably A 7 to aphid strains 1 and 3,
A2 to aphid strain 2 and A8 to all four strains. Immunity to the less
common soil-borne diseases caused by arabis mosaic virus, raspberry ring-
spot virus and tobacco black ring virus is conferred by the dominant genes
lam, lrr, ltb respectively all of which are found within cultivars of R. idaeus.
Several species in addition toR. occidenta/is have been used in crosses
with R. idaeus. Major dominant genes for aphid resistance have been
obtained from R. coreanus, and there is.some indication that resistance to
raspberry beetle may be transferred from R. phoenicolasius.
Air-dry raspberry pollen will remain viable for several years when
stored at 10-20 per cent relative humidity and -5°C. Raspberry plants
pollinated in the glasshouse in early summer produce seeds which can be
sown in autumn and will give fruiting plants within 18 months. Raspberry
cultivars are generally propagated by division of the plants grown in special
'spawn beds', which gives a multiplication rate of 10-30 times. More rapid
propagation of new cultivars and virus free nuclear stocks can be achieved
by root cuttings. A mature raspberry cane established under glass in spring
will give a large root system. If the roots are cut into 2 em long root
cuttings in autumn and the shoots which arise from them taken as stem
cuttings under mist propagation, in 18 months at least, 1 ,000 new clone
plants can be obtained from the original plant.

7.3.6 Red and white currants

These have both been developed from the European indigenous species
Ribes sativum, R. petraeum and R. rubrum, which readily intercross with
one another to produce fertile hybrids but are relatively infertile with
other Ribes species, except R. multiflorum. Red fruit colour is dominant
to white or pale colours but modifying genes affect segregation ratios.
Little work has been done on breeding of new red and white curants,
which are not as popular as black currants or gooseberries. Larger and
better flavoured fruit would be desirable objectives. Spring frost damage
and 'running off' are not serious troubles as with black currant, but
branches may be broken in strong winds. The main diseases are American
gooseberry mildew and leaf spot, and available cultivars differ in degree of

7.3.7 Strawberry
All large fruited cultivated strawberries are derived from hybrids between
the two North American species Fragaria chiloensis and F. virginiana
sometimes referred to as F. x ananassa. Both these species and their hybrids
90 Plant Breeding

are octaploids, but small fruited diploid, tetraploid and hexaploid species
occur in the wild and some are occasionally cultivated. The wood straw-
berry (F. vesca) which is endemic to Britain and many other parts of the
Northern Hemisphere, is a diploid (2n = 14). It is cultivated as a crop on
the island of Corfu, and cultivars of this species are sometimes offered for
garden cultivation in Britain. At least one of these, 'Baron Solemacher', is
a non-runnering type raised from seed. The hexaploid Hautbois strawberry
F. moschata (syn. F. elatior) is occasionally grown commercially, as for
example the cultivar 'Profumata de Tortona' in Italy. The fruit is scented
and plants tend to fruit over a long period. Other diploid and tetraploid
Asiatic species are sometimes cultivated in their land of origin but not in
Improvements in yield and fruit quality are the main objectives
of most strawberry breeding, but disease resistance and suitability for
mechanical harvesting may be equally important. The most troublesome
fungal diseases in Europe are red core and verticillium wilt. Resistance
to virus diseases is less important with strawberry than with other soft
fruit crops as virus spread can fairly easily be controlled in commercial
crops by propagation techniques and certification schemes. Proneness to
June yellows disease (Section 1.12) must also be avoided.
Strawberries are the most widespread of the soft fruits and it is not
surprising that there are more breeders working with them than with most
other crops of this group. Day-length and temperatures affect the runnering
and flower bud initiation so that cultivars bred at one site are often only
suitable for cultivation in areas of similar climatic conditions. For example,
Californian cultivars are productive in Southern Italy, but are less useful
for Northern Europe.
Being octaploid, segregation ratios are never clear-cut, and strawberry
breeding is largely a matter of raising and selecting a large number of seed-
lings-each breeder handling of the order of 10,000 annually. The breeder
eventually learns which clones are the most useful parents through careful
observation and well planned diallel crosses. As a rule, general combining
ability is more important than specific combining ability to the strawberry
breeder (Section 5.9) selecting clones as parents. Clones which transmit
undesirable traits to their progeny, such as susceptibility to june yellows
disease, should never be used. Crossing clones generally gives more promis-
ing progenies than those arising from selftng. All useful characters have so
far been found within the chi/oensis-virginiana complex, sometimes going
back to new collections of the original species which have a very wide
habitat range. It would, however, be useful to examine the possible con-
tribution of other species to a breeding programme and this has been
started (Jones, 1966). The strawberry has been crossed with Po ten til/a
fruticosa and Potentilla palustris but the hybrids are largely sterile (Ellis,
Selecting for resistance to red core disease caused by Phytophthora
fragariae is done by growing seedlings in a site where a wide range of
Vegetatively Propagated Cultivars 91

strains of the pathogen is established in the soil. Most plants contract the
pathogen but some show a degree of 'field resistance' and are able to grow
and crop satisfactorily even when infected. The fungus is confined to the
roots and clean plants for distribution arc obtained by rooting runners in
pots of sterilised soil under hygienic conditions. A similar method is used
for resistance to Verticillium a/bo-atrum, but a technique for inoculating
rooted plants for one hour in a conidial suspension and then planting in
soil has been used with this pathogen. Red core disease is a problem in the
cooler parts of the USA and Britain, and to a lesser extent in other parts
of Europe whereas Verticillium is most troublesome in areas with a warm
Mechanical harvesting is a once-over operation with strawberries so
that for optimal yields the fruits should all ripen simultaneously. There are
three processes to the operation: {1) cutting the crop, which requires that
the fruit should be held clear of the ground otherwise it is damaged or
left behind; {2) singling the fruits, which involves cutting so that each
individual fruit with its stalk is separated from the main stalk which bears
it, and {3) removing the stalk and calyx from the fruit by pulling out
the 'plug' or by cutting it off the fruit, which is possible only with fruits
of a certain shape. These requirements are illustrated in Figure 7.1. All
these features have been found within existing cultivars, but some attempts
have been made to introduce strong fruiting stems to hold fruit clear of
the ground by hybridising with F. vesca using artificially induced poly-
ploid forms of this species.
Fruit qualities required in cultivars for commercial production are
firm flesh and skin which is not easily damaged in transport to market or
to the processor {see Section 9.2.1 ), and bright red fruit as distinct from
that which is dark brownish red. Flavour should not be objectionable or
too pronounced. Some continental and American processors require red-
fleshed fruits.
Strawberry flowers must be emasculated one to three days before
anthesis to prevent self-fertilisation. Pollen can be stored for short periods.
Seed extraction is relatively easy (see Section 4. 7) and seed remains viable
for a long time when stored dry in a refrigerator but seed germination
may be irregular.
Propagating by runners gives some 20-30 plants from one individual
but a rate of propagation exceeding 1,000 per annum may be obtained
with some cultivars by in vitro techniques {see Section 9.7).


It takes a minimum of 15-20 years to produce a new tree fruit cultivar

by hybridisation and, because the individual plant units are relatively large,
a sizeable area of land has to be locked up in the testing process. Mutation
breeding plays a larger role with tree fruits than with many other crops
and this gives somewhat quicker results than hybridisation but the process
is nevertheless relatively slow.
92 Plant Breeding

Figure 7.1 Some morphological features of strawberry types suitable (unshaded)

and unsuitable •(shaded) for mechanical harvesting. 1, Weak ped-
uncle, fruit touches ground and is damaged by cutter blades. 2, Strong
upright peduncle. 3, Peduncles of fruit too short making mechanical
'singulating' of fruit difficult. 4, Recessed calyx; fruit damaged by knives
when calyx removed by cutting. 5, Fruit with short neck and reflexed
calyx making calyx removal by knives less difficult.

7.4.1 Apple
The cultivated apple is generally referred to as Malus pumila though its
precise origin is unknown and other species such as M. silvestris are thought
to have been used in its evolution. Most apple cultivars are diploids (2n ==
34) but about 10 per cent are triploids (2n == 51) including 'Baldwin',
'Gravenstein' and 'Blenheim Orange'. Some are chimceras with one or two
layers of diploid over tetraploid tissue and a few such as 'Paragon' and
'Stay man' are hexaploid-triploid chimceras.
All apple cultivars are, to some extent, self-incompatible and rarely
set more than 10 per cent fruit with their own pollen so that orchards
must contain at least two varieties. Relatively few cultivar combinations
are incompatible but triploids are generally poor pollen producers.
The main breeding objectives are cultivars which give high yields of
fruit of acceptable quality for specific areas of production; late flowering,
Vegetatively Propagated Cultivars 93

to escape frost damage, is important. Most damaging apple diseases and

pests can be controlled adequately by spray programmes but it would be
advantageous to avoid these if possible, and resistance to apple scab,
mildew, apple canker and woolly aphid are important features of some
breeding programmes. Virus infection of rootstocks is a serious problem
and attempts have been made to produce rootstock cultivars which can
be raised from apomictic seed. These would not show the wide variation
of type associated with those from sexually produced seeds but, like
them, they probably would not transmit viruses. However, this work
seems to have been temporarily overshadowed by new rapid techniques
for propagating virus-free clonal root stocks.
Pollen for hybridisation is obtained by collecting unopened flowers
in the 'balloon stage'. The anthers are rubbed off and allowed to dry.
Pollen collected in this way will keep for several weeks but may be stored
for a year in deep freeze. Because of incompatibilities, selfing is rarely
effective and selfed seedlings are generally weak growing. Flowers for cros-
sing should be emasculated at the balloon stage, and when this is done
outside it is convenient to remove stamens, petals and sepals. Insects will
not then visit the flower so that bagging is unnecessary, but fruit set is not
appreciably affected. Seeds should be collected from fruits as soon as they
are ripe and stratified {see Section 4.7).
Apple scab caused by the fungus Venturia inaequalis is the most
troublesome and ubiquitous apple disease. There are many strains of the
pathogen and some cultivars like 'White Transparent' are resistant to
several of them whereas 'Lord Lambourne' seems to be susceptible to
most. A single dominant resistant gene Vf has been identified in Malus
floribunda and extensively used in breeding, especially in the United
States. Apples are assessed for scab resistance in the seedling stage when
the first true leaves are showing between the cotyledons. They are sprayed
with a spore suspension of mixed inoculum and kept in a humid atmos-
phere at 18-20°C. Only very young leaves can be infected this way and it
has been shown that leaf infection is correlated with fruit infection.
Cultivars differ in their degree of polygenic resistance to mildew
caused by Podosphaera leucotricha and single dominant gene immunity
has been recorded in Malus robusta and M. zumi. Cultivars also differ in
their degree of resistance to canker caused by Nee tria galligena; some cider
apples and the Mailing rootstocks Ml and M12 are highly resistant. Seed-
lings can be assessed at an early stage of growth. The cultivar 'Northern
Spy' is highly resistant to woolly aphid (Eriosoma lanigerum) and has been
successfully used to breed resistant rootstocks.

7.4.2 Cherry
Most cultivars are diploid forms of the sweet cherry Prunus avium but
some triploid and tetraploid forms are known. The sour cherries, which
include principally the Morello, are tetraploid cultivars of P. cerasus.
94 Plant Breeding

Duke cherries are tetraploid and thought to be hybrids of P. avium x

cerasus. Mazzard rootstocks are forms of sweet cherry whereas the St Lucie
(Mahaleb) rootstocks are derived from P. maha/eb (2n = 16). Several other
cherry species have been used in hybridisation and also with, for example,
the sand cherry (P. besseyi), which has been crossed with apricot, peach
and plum species, but these are not important in Europe.
One of the main objectives in sweet cherry breeding is resistance to
fruit cracking. The firm fruited bigarrcau types are preferred as the fruit
travels well but it tends to crack more easily than the soft fruited gean or
heart types. Large dark coloured fruits are preferred and in general dark
fruit colour is dominant to light colour. Since nearly all sour cherries are
used for processing, fruit size is less important than overall yield with this
Because nearly all sweet cherries are highly self-incompatible it is
usually unnecessary to emasculate them before making deliberate crosses,
although the new Canadian cultivar 'Stella' is self-compatible. Sour cherries
are self-compatible and the flowers must be emasculated. Seeds from
early-maturing types often germinate poorly and embryo culture has
been used to overcome this difficulty. Seeds of later maturing types have
to be stratified and gibberellin soaks have sometimes helped in germination.
Some seedlings fruit when five years old.

7.4.3 Peach and nectarine

Both of these arc derived from Prunus perstca, nectarines being a smooth-
skinned form homozygous for the recessive gg. Hybrids have been obtain-
ed between peaches and several other Prunus species including the almond,
apricot, sweet cherry and sloe, but none of these hybrids has contributed
towards new cultivars.
Lateness of flowering for frost escape and earliness of ripening are
two of the important breeding objectives. Resistance to peach leaf curl,
peach canker and crown gall are also important but really good sources
of resistance have yet to be found. Peaches arc also grown as ornamental
flowering trees and the inheritance of petal colour and doubleness has
been studied by Lammerts (1945).
The seed of early cultivars is immature at the time the fruit ripens.
This is of little consequence for the edible qualities of the fruit but makes
it difficult for the breeder to obtain seedlings from such types. Embryo
culture has been used successfully to overcome this problem and the peach
is one of the important crops which has benefited most from this technique
in breeding.
Most characters are polygcnetically inherited as in the majority of
fruit crops, but some simply inherited characters arc known including large
flowers (L) dominant to small, pink flowers (R) dominant to red, single
flowers (D) dominant to double, white fruit flesh ( Y) dominant to yellow,
freestone flesh (F) dominant to clingstone, and pubescent fruit skin of
peach (G) dominant to smooth nectarine type.
Vegetatively Propagated Cultivars 95

Peach seed is usually stratified and it takes about five years before
seedlings produce their first fruits. This time can be reduced by removing
part of the testa before seed sowing. However, resultant seedlings often
'rosette' and form short shoots terminated by a bud instead of growing
in length. The terminal buds can be budded to a stock, several on the same
tree. In this way fruiting may be secured in the second or third season but
the technique is time-consuming and expensive and there is a risk of virus
infection from the rootstock.

7.4.4 Pear
Pyrus communis is the main cultivated pear of Europe but the snow pear
P. nivalis is grown on a small scale for the production of perry. In parts of
Asia P. pyrifolia and P. ussuriensis are cultivated and hybrids between
these and P. communis, especially the former, are grown in North America.
Most cultivated forms are diploids but a few polyploid cultivars occur.
Unlike apples and cherries, most pears are self-compatible, although the
majority give better and more reliable crops when cross-fertilised. Some
like 'Conference' regularly produce parthenocz:rpic fruit without even self
pollination but crop better when pollinated.
High quality fruit combined with high yields and cold resistance are
the major breeding objectives with pears. Resistance is sought to fire blight
caused by the bacterium Erwinia amy/ovora, which is also a serious disease
of many other woody plants including ornamentals, especially Rosaceae.
In the United States P. ussuriensis has been successfully used in hybridisa-
tion as a source of resistance, whereas in Britain escape from infection is
sought by producing cultivars which do not have a tendency to produce
late flowers. Infection occurs mainly through flowers but infectious
inoculum is not usually present at the normal time of flowering in Britain.
Pear pollen stores well when dry and can be kept for over a year in
deep freeze. Unlike apples, the flowers to be pollinated must be emasculat-
ed in the balloon stage in order to avoid self fertilisation and sometimes a
pair of notched scissors is used to remove petals and stamen in the one
operation. The seed has to be stratified and seedlings may take six years
or longer before they fruit. for the first time. Little can be done to hasten
first flowering but growing at high levels of fertility and avoiding pruning
and overcrowding are advantageous.
Root stocks have an influence on vigour, hardiness and disease resist-
ance of orchard trees. Several species notably Pyrus betulaefolia, callery-
ana, commumis, pyrifolia, pashia and the quince (Cydonia oblonga), haw-
thorn (Crataegus oxyacantha), june berry (Amelanchier canadensis) and
rowan (Sorbus aucuparia) have been used for pears. However, little breed-
ing work has been done with pear root stocks and this aspect seems to
deserve more attention.

1.4.5 Plum
About 15 different species of plum are cultivated, but the majority of
cultivars are forms of Prunus domestica, which is a hexaploid (2n = 48). It
96 Plant Breeding

has been suggested that P. domestica is not a wild species but that it
originated from chromosome doubling of a cross between the diploid P.
cerasifera (the myrobalan) and tetraploid P. spinosa (the blackthorn or
sloe). The damson plumP. insititita is also a hexaploid. Two other import-
ant species from China are the diploids P. sa/icina, which has been used
in hybridisation, and the apricot plum P. simonii, of which there are
several cultivars, although these are not grown in Britain. Self incom-
patibility is common in plums and the situation of cross compatibility is
sometimes complex in that a cross will give fruit when made one way but
not the other. However, some plums such as 'Belle de Louvain', 'Monarch',
'Coe's Golden Drop', 'Early Transparent Gage' and 'Victoria' are self-
The principal objectives are adaptability to specific areas, early
ripening and resistance to canker caused by Pseudomonas syringae. Im-
proved fruit quality and suitability for special requirements such as drying
cor prunes and the production of plum brandy are also taken into account.
Plum pollen can be stored like that of other rosaceous fruits and has
been known to remain viable for four and a half years at 2-8°C and 50
per cent humidity. Flowers to be pollinated are usually emasculated,
although this is not strictly necessary for self-incompatible types. Crossings
between types of different ploidy levels often give poor seed set or none
at all and aneuploids are sometimes produced. In crosses between diploid
and hexaploid types the diploids give more seeds than the hexaploids used
as the female parent and have produced hybrids which were tetraploid,
pentaploid, hexaploid, heptaploid and octaploid. Apomixis is a common
occurrence in interspecific hybrids.
Early ripening forms produce seed of low germinable capacity, as with
the peach, and the breeder may have to resort to embryo culture, although
this has successfully been done using peat and sand instead of a sterile agar
medium. Ripe seeds of other types should be cracked before they have
dried and then be stored in damp peat in polythene bags at 4-5°C. As soon
as the seed germinates it must be planted and kept growing. In this way
seedlings will flower in three or four years.
Resistance or tolerance to canker is sought from crosses with P.
salicina but no highly resistant cultivar is yet available.


As it is not the fruit or seed of these plants which are the end product of
the crop, it is unnecessary for cultivars to be fertile. Indeed, in some cases
fertility is undesirable as it may reduce the yield. The majority of potatoes
and Jerusalem artichokes do not flower when grown outside in northern
Europe, although it is important to the breeder that flowering should
occur under some conditions so that the desired crosses can be made.
Several perennial vegetables are minor crops and cannot be dealt with
in detail. Other perennial vegetables such as asparagus and chicory are
Vegetatively Propagated Cultivars 97

mainly propagated by seed and breeding of them will be described in

Chapter 8.

7.5.1 Potato
Only the tuberous southern American species of the large genus of Solanum
have been used in breeding the potato. Most cultivars are tetraploids
(2n = 48) assigned to Solanum tuberosum, but it is generally accepted that
this species should be split into two subspecies S. tuberosum tuberosum
and S. tuberosum andigena. The so-called 'andigena' types from the Andes
are also tetraploid and are considered to be the primitive type. They
require short days for tubering whereas the 'tuberosum' types will tuber in
long days. Some of the old Canary Island potatoes are andigena types and
include a triploid cultivar. A few other species have contributed towards
the breeding of the modern potato, notably the hexaploid Mexican S.
demissum (2n = 72), the tetraploid MexicanS. stoloniferum (2n = 48) and
the tetraploid South AmericanS. acaule (2n = 48).
The main objectives in potato breeding have included increased yield
and resistance to diseases and pests such as blight, wart disease, scab and
some viruses. Tuber quality is also important and recently selection has
been made for quality for specialised industries, such as suitability for
potato crisp production which requires high dry matter and low sugar
content, and for fermentation to produce industrial alcohol.
Most potato breeding involves hybridisation within the S. tuberosum
group. The crop has been very well worked by breeders in many parts of
the world and a very large number of seedlings have to be raised and tested
to find any which appear to have advantages over existing cultivars.
Although wild potato species flower profusely, many tetraploid
cultivars bloom sparsely and some, such as 'King Edward' and the Dutch
cultivar 'Bintje', do not flower in the field. Furthermore, flower buds tenel
to fall off prematurely. The two main techniques used to overcome these
difficulties are grafting on to tomato, which is effective for 'King Edward',
and the technique known as 'growing on a brick'. Tubers are planted
literally on a brick or tile and covered with sand or compost. As soon as
the roots have grown below the brick the covering is washed away from
the surface and the small tubers removed as they form. Long days will
also induce flowering.
Cultivars are mainly cross- and self-fertile, although selfing leads to
weak progenies. Many diploid types and wild species have strong incom-
patibility systems and cannot be selfed. Even when pollination has been
effected the flowers may drop off prematurely. In some cases this can be
overcome by pollinating stems of cut flowers taken from the field and
kept alive by standing in water or by spraying with 2-4D two or three
days after pollination.
Dry potato pollen can be kept viable for about a month in a refrigera-
tor and for considerably longer in deep freeze. Pollen sterility is a major
98 Plant Breeding

drawback in hybridisation and many desirable crosses cannot be achieved.

However, many pollen sterile cultivars have a high ovule fertility and
desired crosses should be attempted in both directions. In all circumstances
stringent measures must be taken to prevent virus infection of the breeding
material during the actual breeding process and the early stages of testing.
The main source of blight resistance, an extreme hypersensitivity
reaction, is 5. demissum. Crosses with tetraploid cultivars give mainly
pentaploid seedlings and these have to be backcrossed to 5. tuberosum to
regain tetraploids. The situation is complicated by the occurrence of
different races of the fungal pathogen Phytophthora infestans and control
of the disease is generally incomplete. Unfortunately good resistance is
associated with late maturation.
Resistance to wart disease is found within 5. tuberosum and is thought
to be controlled either by one dominant gene X or two complementary
genes Y and Z. Breeding for control of this disease has been notably
successful. Resistance to potato virus X is conferred by two dominant
genes Nx and N6 found in 5. tuberosum and has also been obtained from
5. acau!e. Resistance to viruses Y and C has been located in 5. sto!oniferum.
Somatic mutations in leaf shape, tuber colour, sprout colour and
flower form and colour occur from time to time in potato cultivars, some-
times as solid mutants but more commonly as chimceras. These have to be
rigorously removed in the propagation cycles.


In the past, amateur enthusiasts have played the major part in the breeding
of perennial ornamentals. However, during the last decade the considerable
economic importance of cut flowers and pot plants has been fully apprec-
iated and state-aided breeding of these crops is being done now on a
moderate scale. The main objectives are cultivars suitable as cut flowers
and for forcing out of season to obtain an all the year round supply.
Improvements in existing types are sought but the introduction of com-
pletely new types, including new genera or wide interspecific crosses, can
make outstanding contributions.
There are many species of ornamental plants with breeding potential
but only a few of the more important genera can be considered here.

7.6.1 Carnation
The border carnation is derived from the diploid (2n = 30) Dianthus
caryophyllus, which still grows wild in the Atlas Mountains. The yellow
colour probably comes from hybridisation with the Balkan species D.
knappei (2n = 30).
Perpetual flowering carnations originated in about 1750, probably as
a result of hybridisation of the border carnation type with the very flori-
ferous annual D. chinensis. Present-day cultivars are hexaploid (2n = 90).
Vegetatively Propagated Cultivars 99

Flower doubleness is due to petaloidy of the stamens and the extremely

double types produce practically no pollen; indeed, lack of pollen is often
a setback to breeding. Pollen production is improved by growing the
plants in a starved condition and at a relatively high temperature (23°C),
(Kho and Baer, 1973). However, high temperatures tend to reduce female
fertility. Pollen can be stored for one to two weeks when dry and seed
remains viable for several years.

7.6.2 Chrysanthemum
The cultivated chrysanthemum can be traced to 500 BC. Earliest forms
were probably all yellow·flowered types derived from C. indicum (tetra-
ploid 2n = 36). Hybridisation with C. morifolium (hexaploid 2n = 54)
introduced other colours. Cultivars have a fairly high rate of natural
mutation and many new forms have originated this way rather than from
seed. Chromosome numbers of cultivars are often irregular and range from
51-68. Radiation has been used successfully in the last decade to speed
up mutation and thus to produce new cultivars {see Section 6.2.1 ).

7.6.3 Dahlia
Dahlia cultivars are said to have been mainly derived from D. variabilis
which is an octaploid (2n = 64) and may itself be a form cultivated by the
Aztecs. Most species are tetraploids (2n = 32). D. pinnata with semi-double
flowers, and D. coccinea have probably also been involved in the breeding
of present-day Dahlia cultivars, and a 'rogue' plant in a batch imported
from Mexico on 1872 that became called D. popenovii gave rise to cactus
types. D. merckii, which has an extra two chromosomes (2n = 34) has
also been used in breeding.
Dahlia flowers are usually strongly self-incompatible and do not need
to be emasculated before hybridisation. Four major genes are known for
flower colour and their interaction is described in some detail by Crane
and Lawrence (1934). In recent years new mutant forms have been obtain-
ed by irradiation.

7.6.4 Delphinium
Most delphinium species are diploids (2n = 16) but the cultivated border
delphinium derived primarily from D. elatum, and probably also D. forma-
sum and D. grandiflorum, is tetraploid (2n = 32). The so-called Belladonna
types are sterile hexaploids (2n = 48) (Mehlquist and Gage, 1964) of
similar parentage. The cultivar 'Pink Sensation' is a fertile tetraploid of the
red flowered American species D. nudicaule and D. elatum. Recent work
with these species and with D. cardinale and D. zalil has produced some
very interesting scarlet flowered hybrids. The projects, which also include
mildew resistance and scented flowers, is described by Legro (1964).
Delphinium seed often has poor viability and should be sown soon
after ripening.
100 Plant Breeding

7.6.5 Freesia
Commerical cultivars are derived mainly from the white- or yellow-flowered
Tritonia (syn. Freesia) refracta and pink-flowered T. armstrongii. Although
some diploid (2n = 22) cultivars are grown, many of the largest flowered
modern types arc tetraploid (2n = 44); triploid and aneuploid (42, 43 and
45) cultivars are known. The diploids are fertile and can be raised from
seed but other forms are mainly very poor seed producers and have to be
vegetatively propagated so that disease is becoming a problem with them.
Tetraploids do produce some seed from which the breeder can raise new
cultivars but certain cultivar-crosses are incompatible, although the nature
of the incompatibility is not known (Sparnaaij eta/. 1968).

7.6.6 Fuchsia
There are some 100 species of Fuchsia from Central and South America
and a few from New Zealand. Many are readily cross-fertile and mainly
diploid (2n = 22) or tetraploid. Modern cultivars, mainly derived from
F. magel/anica, F. macrosternina and F. fu/gens, are mostly high poly-
plaids including octaploids and decaploids (2n = 11 0). Many species have
not yet been used by breeders and there is plenty of scope for further
introductions into the cultivar gene pool.

7.6.7 Gladiolus
There are several hardy European species of Gladiolus but present-day
cultivars are all derived from South African species. 'Colvillei' types are
thought to be hybrids of the two diploid species (2n = 30) G. cardinalis
and G. tristis, although both diploid and triploid cultivars exist. Large-
flowered types are probably hybrids of G. psittacinus and G. oppositifolius,
but other species such as G. purpureo-auratus, G. papi/ia and G. primulinus
have been used by breeders. Virus infection of clones is becoming a major
problem with Gladiolus.

7.6.8 Hyacinth
Cultivars are all derived from the diploid (2n = 16) wild Hyacinthus orient-
a/is of the Middle East and parts of Europe, which looks similar to the
cultivated Roman hyacinth. Most are polyploids and aneuploids.

7.6.9 Iris
Florists' bulbous iris are derived mainly from European species found wild
in Iberia. The English iris/. xiphioides (2n = 42) does not cross with other
species. The Spanish iris /. xiphium is diploid (2n = 34) and the Dutch
irises are mainly sterile hybrids of this species with/. filifolia and/. juncea.
The cultivar 'Wedgewood' is of special interest as it can be forced earlier
than other types. It is probably a hybrid of I. xiphium and the North
African species /. tingitana. Unfortunately it is sterile, but natural mutants
have given the cultivars 'White Wedgewood' and 'Dominator'. Irradiation
Vegetatively Propagated Cultivars 101

of the bulbs shortly after lifting has induced mutants, although these were
not observable until the second season, probably because the flower was
already formed at the time of treatment. The cultivar 'Prof. Blaauw' is
of similar parentage to 'Wedgwood' and also sterile but cannot be forced.
It seems that the earliness of 'Wedge wood' is due to a fortunate chance
gene combination. Virus infection is a problem with bulbous iris but
healthy stocks have been obtained by mcristem culture.
Pogon or bearded iris cultivars are complex hybrids of a number of
species, especially I. chamaeiris, I. pal/ida and I. variegata. They are often
tetraploids but higher polyploids and aneuploids occur. Recently the
spectacular Onococyclus species I. gatesii has been crossed with Pogon
irises to give the cultivar 'William Mohr' (2n = 22). This produces a few
seeds when crossed with other cultivars but no fertile pollen and has been
used in further hybridisation.
Seed of the bearded iris is slow to germinate and often difficult to
raise, and embryo culture has been extensively used in the United States
to aid germination to produce new cultivars (see Section 4.1 0). Dry pollen
stores well and can be kept at least 12 months.

7.6.10 Lily
All of the 80 or so true Lilium species arc diploid (2n = 24) with occasional
additional chromosomes. However, the tiger lily is a triploid (2n = 36) in
which one set of chromosomes differs from the other two and it is there-
fore of hybrid origin, in spite of the fact that it is referred to the species
L. tigrinum. The commercially most important group of cultivars includ-
ing the Asiatic hybrids, Mid Century hybrids, Fiesta hybrids and Umbel-
latum hybrids derive from the tiger lily and some eight species from
southern Asia including L. davidii, concolor, dauricum and cernuum all
of which are fairly cross-fertile. Trumpet lily hybrids are mainly derived
from crosses of L. sargentiae, L. brownii, L. regale <ind L. sulphureum, and
Oriental hybrids from crosses between L. speciosum and others, mainly
L. auratum.
Lily flowers are often strongly self-incompatible though they need
emasculating before pollinating to be certain that self fertilisation has not
occurred. Some species, notably L. regale, are apomictic when pollinated
with other species and one cannot be certain of hybridisation in wide
crosses without careful verification. Both pollen and seed can be kept well
in deep freeze but lose vitality rather rapidly when stored at ambient
temperatures. Seed of some species produce a small bulb below ground in
the first growing season without producing leaves until a year after sowing
but the seed of most Asiatic hybrids produces a leaf shortly after sowing.

7.6.11 Narcissus
Species are native only to south-west Europe, with the exception of N.
tazetta which is found in Asia. N. tazetta and the few allied species have a
102 Plant Breeding

basic chromosome number of 11 and are usually found wild as diploids

(2n -= 22). In all othe.- species the basic number is 7 with the majority of
species as diploids (2n = 14), although tetraploid forms occur wild occas-
ionally and many of the N. cyc!amineus wild forms are higher polyploids.
Many cultivars are tetraploids or triploids. Most diploid wild forms can be
readily intercrossed but the 7 and 11 chromosome number groups, when
intercrossed, give sterile hybrids notably the 'Poetaz' cultivars.
Seedling Narcissus hybrids do not flower until they are six to eight
years old and natural vegetative reproduction is slow, so that breeders are
greatly interested in new techniques for rapid vegetative propagation
(Section 9.7). Most daffodils have been bred by amateurs to win prizes at
flower shows and are not suitable for commercial cultivation for forcing as
cut flowers. More emphasis is now being put on such characteristics as
yield of flowers, suitability for forcing and longevity of the flowers.

7.6.12 Paeony
Most herbaceous paeony cultivars are diploids or tetraploids derived from
the diploid Chinese Paeonia /actiflora (syn. P. a/biflora) (2n -= 10). Two
yellow-flowered species from the Caucasus are of special interest for
crossing with this species but one, P. mlokosewitchii, does not hybridise
and the other P. wittmaniana a tetraploid (2n = 20), produces sterile
triploid hybrids with it. There are a number of European species, most of
which readily cross with one another, and some of the old garden cultivars
are hybrids of P. peregrina. These have only recently been crossed with the
P. /actiflora group.
The tree paeony is P. suffruticosa and cultivars are often hybrids with
P. !utea and P. de/aveyi, but this species does not seem to hybridise with
the herbaceous group.

7.6.13 Pelargonium
The 'zonal pelargoniums' or geraniums are said to have been derived
mainly from P. inquinans x P. zonate but other species may be involved,
especially P. scandens. They do not cross with the 'regal pelargoniums',
which is a complex group of hybrids of the sub-genus Pelargium, but they
hybridise with the ivy-leaved geranium P. pe!tatum.
Some leaf markings are chim<eras and can only be maintained by
vegetative propagation but there are genes for leaf zoning, mainly from
P. scandens, and certain cultivars are especially useful parents for these
characters. Traditionally, geranium cultivars have been clones propagated
by cuttings, but recently there has been much interest in F 1 seed propagat-
ed cultivars. The reason for this change in emphasis is the increasing
problems arising from virus infection of stocks.

7.6.14 Rhododendron
Both the deciduous azaleas and the evergreen rhododendrons are members
of the genus Rhododendron.
Vegetatively Propagated Cultivars 103

The so-called hardy hybrid rhododendrons were derived mainly from

R. arboreum, campanulatum, catawbiense, caucasicum, maximum and
ponticum. The first mentioned has the most showy red flowers but is less
hardy than the other five. Later the tender R. griffithianum with large
scented flowers was hybridised with this group to give such popular culti-
vars as 'Pink Pearl'. Recently many other species have been crossed with
older cultivars, especially R. griersonianum, repens, thomsonii and recently
R. yakusimanum. Some of the details of this work arc described by
Street (1954) and a comprehensive list of hybrids is given by the Royal
Horticultural Society (1964).
The scaly small-leaved '!epidote' species do not cross easily, if at all,
with the non-scaly larger-leaved 'elepidote' species and hybridisation
between deciduous and evergreen species is uncommon, though some
so-called 'Azaleodendron' cultivars exist.
The early original Ghent azaleas were hybrids of R. ca/endulaeeum
x nudiflorum but several other species, especially R. speciosum, viscosum,
luteum were later involved. 'Mollis hybrids' derive from R. mol/is x
japonicum. Recently the Ghent and Mollis groups have been hybridised
together, giving the Knaphill and Exbury strains of azalea.
It would be helpful in rhododendron breeding to induce polyploidy
artificially to regain fertility of hybrids, but this does not seem to be
easy to achieve.

7.6.15 Rose
Although there are several indigenous European rose species, modern rose
cultivars are largely derived from Asia. Rosa gallica, said to be the red rose
of Lancaster, was one of the first cultivated species and, crossed with
R. phoenicea, gave the damask rose. The musk rose R. moschata was also
used in early hybrids and later roses, notably the cabbage rose in which all
stamens are petaloid, and the moss rose, owe some of their character to
the native dog rose R. canina. Later the China rose R. chinensis hybridised
into the group gave a longerflowering period and the 'Hybrid Tea' rose and
'Hybrid Perpetual' were born. The pernettiana roses were hybrids with the
above group and the Austrian briar R. foetida, and these provided the
main gene pool for the modern floribunda types.
Most modern cultivars are tetraploids or higher polyploids and apo-
mixis is not uncommon within the genus. Although the seed often requires
stratification, the raising of seedlings to flowering stage does not usually
take more than 18 months. A recent appraisal of rose breeding was given
by Rowley (1966).

7.6.16 Tuhp
Our tulip cultivars are mainly derived from hybrids between unidentified
south-west Asian species, but during the last 30 years there has been
considerable activity in hybridising T. eich/eri, fosteriana, greigii and T.
104 Plant Breeding

kaufmanniana. Most species and cultivars are diploid (2n = 24) but a few
polyploid wild forms are known and some cultivars, including 'Apeldoorn'
'lnglescombe Yellow' al}d 'Keizerkroon' are triploids (2n = 36) and others,
such as 'Riant' and 'Sunburst' are tetraploid (2n = 48). Several cultivars
have originated as sports, and those with 'broken' flowers owe their
character to virus infection.
The small flowered species do not cross readily with the larger flower-
ed cultivars and seed production from cultivars is often poor, especially
with the triploids. It would be useful sometimes to obtain seed from
sterile plants derived from wide crosses. This might be done if the chromo-
some number could be doubled, but tulip bulbs and seedlings are not
suitable subjects for colchicine treatment. This problem has been partly
overcome by nitrous oxide treatment (see Section 2.2).
Tulip seedlings do not flower until they are five to eight years old and
an early method of screening useful types would be helpful. Work in
Holland has suggested that early sprouting of seedling bulbs is correlated
with capacity for early forcing (Van Eijk and Toxopeus, 1968).


Seed produced from a deliberate cross pollinatior may not be of hybrid
origin even when every precaution has been taken to prevent selfing.
Apomixis may have intervened and the seed will then be entirely maternal
in origin. This can be very disappointing to the breeder who is under a
moral obligation to refrain from claiming a wide cross hybrid unless there
is good evidence to support that claim.
Sometimes hybrids can be recognised at an early stage of seedling
development by morphological characters such as leaf shape and colour
which do not occur in seedlings of the mother parent when selfed or sib
crossed. Usually hybrids can be recognised at flowering by other morpho-
logical characters such as flower colour. However, the differences may
not be so clearly defined that one can be certain of hybridity and further-
more, one may not wish to wait for evidence if the plants take several
years to flower from seed.
Certain evidence of hybridity can sometimes be obtained at an early
stage of seedling development by an examination of the chromosomes
Root tip squashes are prepared as for chromosome counts (see Section 2.3)
and a search made for chromosome types which differ from any found
in the mother parent. In Figure 7.2 we see the typical chromosome set
of Lilium henryi and the somatic chromosome set of a suspected wide
hybrid with that species and another. The arrow indicates a long chromo-
some with two constrictions, which is not found in the seed parent and
therefore confirms that the examined plant is a true hybrid. This technique
is useful only with good preparations and is difficult with some species
which do not have distinctive chromosomes. In searching for differences
it is useful to look at the longer chromosomes and the positions and
numbers of constrictions.
Vegetatively Propagated Cultivars 105

Figure 7.2 Karyotype (haploid chromosome set) of Lilium henryi (top). and diploid
chromosome set of L. henryi X 'Asiatic cultivar' (bottom). Arrow indi-
cates a long chromosome with two constrictions of a type not found in
L. henryi, thus confirming hybridity.

Biochemical techniques, especially paper chromatography of leaf

extracts have also been used successfully to verify hybridity as with Pyrus
species (Catlin and Olsson, 1968).
106 Plant Breeding

ANDERSON, M. M. (1971). Resistance to gall mite (Phytoptus ribis Nal.) in the
Eucoreosma section of Ribes, Euphytica, 20, 422-426
CATLIN, P. B. and OLSSON, E. A. (1968). ldentific(ltion of interspecific hybrids of
Pyrus after paper chromatography of leaf extracts, Proc. Am. Soc. hart, Sci.,
CRANE, M. B. and LAWRENCE, W. j. C. (1934). The Genetics of Garden plants,
Macmillan, 236 pp
ECK, P. and CHILDERS, N. F. (1966). Blueberry Culture, Rutgers University Press,
New Brunswick, 378 pp
ELLIS, j. R. (1962). Fragaria-Potentilla intergenetic hybridisation and evolution in
Fragaria, Proc. Linn. Soc. London., 173, 99-106
JONES, 1. K. (1966). Evolution and breeding potential in strawberries, Scient. Hart.,
KHO, Y. 0. and BAER, j. (1973). The effect of temperature on pollen production in
carnations, Euphytica, 22,467-470
LAMMERTS, W. E. (1945). The breeding of ornamental edible peaches for mild
climates. I: Inheritance of tree and flower characters, Am. j. Bot., 32, 53-61
LEGRO, R. A. H. (1964). Species hybrids and delphinium breeding, Proc. Int. Hart.
Congress (1962), 4, Duclot, Gembloux, 50-53
MEHLQUIST, G. A. L. and GAGE, M.A. (1964). The origin of Delphinium X bela-
donna Hort., Proc. Int. Hart. Congress (1962), 4, Duclot, Gembloux, 41-49
Rhododendron Handbook, Part 2, Rhododendron hybrids, Spottiswoode and
Ballantyne, London, 436 pp
ROWLEY, G. D. {1966). The experimental approach to rose breeding, Scient. Hart.,
SPARNAAIJ. L. D., KHO, Y. 0. and BAER, J. (1968). Investigations on seed pro-
duction in tetraploid freesias, Euphytica, 17, 289-297
STREET, F. (1954). Hardy Rhododendrons, Collins, London, 192 pp
VAN EIJK, j. P. and TOXOPEUS, S. j. {1968). The possibilities of early selection for
forcing ability and productivity in tulips, Euphytica, 17, 177-183


Seed propagated cultivars must be fertile and capable of being maintained
through generations of seed production without radical changes of type.
Those produced in what may be called the conventional way must be
sufficiently homozygous to breed true within certain narrowly prescribed
limits. The so-called 'hybrid varieties', however, do not themselves breed
true and have to be reconstructed from true breeding parents each time a
batch of seed is produced.
Most seed-propagated cultivars, with the exception of French beans
and lettuce, do not suffer unduly from deterioration due to virus con-
tamination, as is common with vegetatively propagated cultivars. However,
they are subject to genetic deterioration resulting from segregation,
because for practical reasons it is nearly always impossible to maintain
them as completely homozygous units. To ameliorate this genetic shift,
stocks have to be frequently reselected and the process of maintenance
and propagation through successive generations of seed production then
becomes an extension of the breeding and an important aspect which the
breeder of new cultivars cannot ignore.
Unlike vegetatively propagated cultivars, those which are seed propa-
gated are mostly diploids and often derived from a single species. This is
because polyploidy and complex interspecific hybridisation tend to lead
to infertility, which makes economic seed production difficult. There are
of course exceptions, such as the tetraploid leek and the interspecific
hybrid swede.


Seed propagated crops fall into two fairly well defined categories-those
in which the plants are largely maintained by (1) self- and (2) cross-
Self-fertilisers obviously must be self-fertile and they must not suffer
appreciably from inbreeding depression. Their flowers may be constructed
in such a way that self pollination nearly always occurs, and out crossing
is rare under natural conditions as in peas, French beans, China aster
(Col/istephus) and African marigold (togetes). At the other extreme the
flowers may be capable of both self- and cross-fertilisation, as with cauli-
flower, and there are intermediate types such as broad bean, lettuce and
108 Plant Breeding

runner bean which are largely self-pollinated, but a small amount of out
crossing can occur. By contrast, cross-fertilisers usually have an incom-
patibility system which prevents or inhibits self-fertilisation and, as a rule,
their vigour is reduced by inbreeding depression.

8.2.1 Breeding self-pollinating crops

This is a relatively simple process. Seeds are collected from the most
promising individual plants and grown on for selection. Seeds are again
collected from the most desirable plants amongst the new seedlings and
the process repeated. The technique is known as line breeding or pedigree
breeding. If the crop is one like pea or French bean, which does not easily
cross-pollinate, it is unnecessary to isolate the plants. However, if cross-
fertilisation is likely to occur, each plant or an individual inflorescence or
flower from each plant must be isolated from pollinating agents at flower-
ing time under insect-free, or sometimes wind-free conditions.
It may be desirable to introduce new characters into the material by
enforced cross-fertilisation and then in the F 2 generation there will be
segregation of a heterogenous mixture of different types. In order to
regain stability of the material and obtain new true breeding types, it is
necessary to inbreed again by about six generations to regain the desired
degree of homozygosity. It may be possible to speed up this process by
forcing the plants in a glasshouse to obtain more than one crop of seed
a year, but it is then usually impossible to select under the conditions in
which the crop will normally be grown.
Sometimes it is desirable to maintain the majority of the character-
istics of a batch of material but to introduce into it one character, such as
disease resistance, from an otherwise undesirable type. This is done by a
programme of back crossing in which the F 1 plants and subsequent
generations are crossed to the most desirable parent instead of being
allowed to self fertilise and are selected in each generation for the 'new'
character. Six or eight generations of back crossing arc usually necessary
to achieve a desirable true breeding type with the new character added.
Some self-pollinating crops, notably peas, beans and lettuce, can
carry viruses through seeds, and precautions are necessary to reduce this
risk during the seed production stages.

8.2.2 Breeding cross-pollinating crops

This is more complex than with self-pollinating crops because a degree of
heterozygosity usually has to be maintained to prevent excessive loss of
vigour from inbreeding depression. A balance has to be achieved and
frequent reselection is usually necessary to prevent excessive hetero-
zygosity or a shift in gene balance towards an undesirable character, for
example premature bolting.
Sometimes it is desirable to treat the material temporarily as a self-
pollinating crop using pedigree breeding and to enforce self-fertilisation of
Seed Propagated Cultivars 109

individual plants to improve uniformity, and as it were to screen out

undesirable recessive characters, but usually this can be done only through
three or fewer generations before severe inbreeding depression becomes
Because selfing is difficult to achieve in many cross-pollinating crops,
due to the self incompatibility systems, they are often improved by mass
selection. This involves the selecting of a number of similar looking plants
(usually 10-50) of the 'improved' type, transplanting them close together
but isolated from other plants with which they are likely to hybridise, and
saving the seed. It may be necessary to repeat this process for several
generations. A cruder method, formerly used extensively, is to mark the
selected plants in a seed crop, leave them in situ and save the seed separately
from the rest of the crop. In this way the plants suffer less disturbance and
the yield is higher than if they arc transplanted. However, the method
gives no control over the pollen parents which may have been undesirable
plants and improvement requires more generations of selection and may
never be achieved. Another technique sometimes used is simply to rogue
out the undesirable types before flowering, leaving the rest to seed-a
technique known as negative mass selection.
In mass selection the plants are usually chosen solely on their appear-
ance. A more refined technique is to base their selection on the progenies
they give, by progeny testing. Seeds from each individual selected plant is
kept separate and part only is sown to test the progenies. The resulting
progeny plants arc compared and it is noted which of the selected parent
plants gave the best progenies. The reserve seeds only from the parents
which gave the best progenies are then sown and mass selection repeated
on the resultant plants. Various combinations of alternating generations of
mass selection and line breeding can be practised and the selected plants
may be deliberately pair crossed before progeny testing. A preliminary
stage of hybridisation followed by successive back-crossing may also be
practised, as with self-pollinating crops.

8.2.3 Hybrid cultivars

When selfing is enforced on out pollinating plants the progenies become
more uniform but suffer from inbreeding depression. If the resultant weak
inbred lines arc intercrossed the progenies are often uniform and exhibit
hybrid vigour (Section 5.8). Two such inbred lines when crossed together
produce an F 1 and this technique is being used increasingly to produce
new F 1 cultivars of several vegetable crops (Table 8. 7) and ornamentals
(sec Section 8.5).
Research is also in progress on the possibility of producing F 1 culti-
vars of other crops such as beetroot, celery, endive, leek and radish.
With a few highly priced crups, crossing between the two parent lines
is done by hand, especially when the yield of seed per individual hand
operation is high, as in tomato, cucumber and snapdragon. For most crops
110 Plant Breeding

Table 8.1

Numbers ofF, cultivars of some veqetable crops in the UK only and in all EEC
countries {after Wills and North, 1978)

Number of Percentage F, hybrids

cultivars 19 76 1973 1976
(EEC) (UK) (EEC)
Brussels sprout 113 26 65
Cabbage 128 10 40
Cucumber and gherkin 127 67 69
Carrot 57 3 11
Marrow and pumpkin 44 12 25
Onion 96 5 8
Spinach 86 9 20
Tomato 209 45 65
Turnip 55 0 2

crossing is 'controlled' by utilising self-incompatibility, male sterility,

dioecy or monoecy (Table 8.2). These systems will be discussed in more
detail under individual crops.

Table 8.2

Systems ofF, seed production

System Crops

A Genetic/cytoplasmic
(i) Male sterility Carrot
(ii) Self-incompatibility Ageratum
(sporophytic or gametophytic) Bellis
Brussels sprout
(iii) Dioecy Spinach
(iv) Monoecy Sweetcorn
B Mechanical or manual Snapdragon
C Chemical (male suppression or gametocidal) Cucurbits
Seed Propagated Cultivars 111

The maintenance ofF 1 cultivars often runs into two main difficulties.
The 'hybrid' seed crop may contain a proportion of seed of one or both of
the parent lines due to incomplete control of intercrossing between them.
If this rises above five or ten per cent the whole crop may have to be
destroyed. The other problem which can arise is a decreased fertility of the
parent lines due to continued inbreeding so that seed yields are considerably
reduced. Both these difficulties contribute to the relatively high price of
F 1 seed compared with conventionally produced seed. In spite of this
the superior performance of F 1 as compared with conventional cultivars
nearly always outweighs the disadvantages of extra costs. It is interesting
to note, however, that improved performance is not the only reason for
the introduction of F 1 cultivars. japanese scedsmen who pioneered them
in many vegetable and ornamental crops did so initially because they
afforded a measure of protection against pirating from their competitors.
Nowadays this aspect is of less importance to the breeder as adequate
protection is given in several countries by Plant Breeders' Rights {see
Section 9.5).
For some agricultural crops double cross cultivars are used in which
each of the parent lines is itself obtained by crossing two other lines-a
sort of F 1 between two F 1 s. This technique is cheaper than F 1 s because
of the reduced requirement for hand labour but does not give such a
genetically uniform end product as a typical F 1 • Three-way crosses arc
also used in which one parent is an F 1 and the other an inbred line. As we
shall see later, onions arc an example of this type. So-called synthetic
varieties are those in which several carefully selected inbred parent lines
are seeded together as an F 1 but with more than two parent lines. There
has been much discussion about the merits of synthetic varieties recently;
they are often productive but they do not give such uniform crops as F 1
cultivars and uniformity is usually as important as productivity, especially
for the freezing and canning industries, or when mechanical harvesting is


The distinction between self and out pollinating crops is not absolute and
breeders are constantly examining self pollinators to find if there is an
advantage in treating them as cross pollinators. Twenty years ago most
tomato cultivars were inbreds produced by selfing but nowadays the
majority are F 1 cultivars and Vicia beans and cauliflowers are currently
being examined with the same objective in view. Cauliflowers are a special
case and although most cultivars of this vegetable are still produced by
self-pollination techniques they arc forms of Brassica oleracea, which is
largely cross-pollinating and will be dealt with in Section 8.4.

8.3.1 Broad bean

The broad bean is said to be a form of Vicia faba which has been cultivat-
ed since antiquity, although the truly wild type has not been identified.
112 Plant Breeding

The agricultural field, horse or tick bean, which is especially hardy and has
relatively small seeds, is also a form of the same species and hybridises
with the broad bean. Most cultivars of Vicia faba have seeds with green
testas which tend to "become brown as they mature but one group, exem-
plified by the cultivar Threefold White', has white testas, pale coloured,
instead of dark, hilums and white flowers without the purplish blotch on
the wing petals as in most forms. These characters arc all controlled by a
single recessive gene and cultivars of this type are used almost exclusively
for canning because the seeds do not discolour the canning liquid.
The flowers of most forms of Vicia faba usually do not become self-
pollinated unless they are visited by insects. Pollen is shed in the keel and
cannot reach the stigma except by a brushing action of hairs on the style
brought about by the weight of an insect alighting on the flower to cause
'tripping'. However, forms are known which will self-pollinate without the
need for tripping. As a rule continued self-fertilisation of Vicia faba does
not lead to a pronounced loss of vigour but recent work shows that for
some forms heterosis can occur and that it may be advantageous to treat
the crop as a cross fertiliser. Plants may frequently behave as though they
are self incompatible because balanced lethal genes prevent self seed
development (Rowlands, 1964).
Very little work has been done on the breeding of improved broad
beans in recent years but there is at present a considerable interest and
activity in the breeding of field beans as a useful pulse protein source for
northern Europe. This work will no doubt produce new forms of interest
to horticulturists.

8.3.2 French bean

Phaseolus vulgaris is a native of South America and a relatively new,
although very important, introduction to Europe. From a horticultural
point of view four distinct forms can be recognised. (1) The type grown
for the dry seed and often referred to as the navy bean in America: there
is considerable interest in Britain in the breeding of a commercially viable
white-seeded form of this type to be used as prepared 'baked beans', but
as yet this has not been achieved. (2) Pole bean, a tall climbing form
occasionally grown commercially in glasshouses for high quality crops and
rarely in British gardens, as exemplified by the cultivar 'Climbing Blue
Lake'. (3) Dwarf string bean, typified by old cultivars such as 'The Prince',
'Masterpiece' and 'Canadian Wonder': the pods must be picked at an early
stage otherwise they become tough. (4) Snap-pod bush bean or stringless
green bean. Nearly all cultivars used for processing are of this type though
they mostly do not have such a good flavour as the string beans. White-
seeded types are preferred by processors, especially for canning and much
effort has been put into the breeding of improved forms of this type
during the last two decades. There are also several novelty types occasion-
ally grown on gardens with yellow, purple or red flecked pods and with
seeds of interesting shapes and colouration. All types are cross fertile.
Seed Propagated Cultivars 113

The flower ot Phaseolus has a coiled, spring-like style sheathed by the

fused filaments of the anthers and the keel. It is not easy to emasculate
without damaging the style and hand cross-pollinating requires practice
and skill before it is effective. Keeping the emasculated flowers in a humid
atmosphere for some hours after pollination helps to effect fertilisation.
The French bean P. vulgaris, when used as the female parent, will
cross with the runner bean P. coccineus. The hybrids are usually very
infertile when selfed but can be back-crossed toP. vulgaris and selected for
fertility. Germination is epigeal in P. vulgaris, hypogeal in P. coccineus and
the hybrid intermediate in this character. The cross has been made by
several breeders ostensibly to transfer the greater cold tolerance of runner
bean to the French bean but very few, if any, hybrids have given rise to
commercially acceptable cultivars.
A great deal is known about the genes and linkage groups of P. vul-
garis, but many of the characters studied arc of relatively little direct
interest to the practical breeder. There are at least eight genes controlling
seed colour and one P is basic to coloured seed so that all white-seeded
forms are the homozygous recessive pp and breed true. Black seed is
epistatic to all other colours. Pod stringlessness tends usually to be domin-
ant to stringiness but inheritance is not always clear cut, and green pod
colour is dominant to the yellow colour of wax-pod types. Some three
genes govern height: L tall dominant to short, A indeterminate dominant
to determinate habit, and T twining dominant to non-twining. Thus pole
beans carry all three dominants LA T and bush beans are typically 1/aatt.
The main objectives in breeding bush snap-pod beans for European
conditions are yield, earliness, pod quality for processing by freezing and
canning, suitability for mechanical harvesting (mainly upright habit and
concentrated ripening) and resistance to the diseases halo blight, anthrac-
nose and seed-borne viruses. Earliness has been sought by seedling tolerance
to cold soil conditions to permit earlier sowing as well as early maturation.
Processing quality is determined by sophisticated processing tests which
arc usually only available to the processors themselves. Susceptibility to
halo blight caused by the bacterium Pseudomonas phaseolicola and to
anthracnose caused by the fungus Col/etotrichum lindemuthianum can
be assessed by soaking the seeds in infective suspensions and observing
the progress of the diseases on small plants. There are several strains of
Co/letotrichum and most cultivars arc tolerant or resistant to some but
not all of them; however, a type 'Cornell 49-242' has a major gene for
immunity to all races. There are three main seed-borne viruses, Phaseolus
virus I, Phaseolus virus 2 and Nicotiana virus II. The symptoms vary
according to the genotype and some cultivars such as 'Processor' and
'Masterpiece' are resistant or tolerant to most of the symptoms; no culti-
vars are completely immune to all the viruses or symptomless when
infected. When planning their facilities breeders need to take account of
the fact that French bean seed is bulky to store and its viability is reduced
when stored at low temperatures.
114 Plant Breeding

8.3.3 Lettuce
Cultivated lettuce is described botanically as Lactuca sativa and was
probably developed from the ubiquitous wild lettuce L. scoria/a with
which it crosses readily to give fertile hybrids with regular pairing of the
chromosomes at meiosis, suggesting that the two forms are closely related.
It also hybridises with L. sa/igna and L. a/taica. Cultivated forms include
butter-head, crisp-head, cos, loose leaf or bunching, all of which readily
inter-cross with one another.
Lettuce flowers are small and difficult to emasculate by hand but this
can be done by the technique described in Section 3.4 (and see Figure 3.2)
or a selective gametocide (Section 3.7). After pollinating, the flowers
should be bagged to prevent cross-contamination by insects.
The main objectives in lettuce breeding arc suitability for growing
under glass in short days, resistance to downy mildew fungus (Bremia
lactucae) and to lettuce mosaic virus and beet western yellows virus.
Many British cultivars have white seeds but black and yellowish-brown
seeded forms occur. Seed colour is controlled by two genes, W and Y.
Genotypes WWYY have black seeds, wwYY yellow, and WWyy, wwyy
white seeds. Non-heading K is dominant to heading; lobed leaves (such as
in L. scoria/a) are dominant to entire leaves of the heading and cos types,
probably with two complementary genes UT and U2. The inheritance of
some characters of lettuce were reviewed by Lindqvist (1960).
There are at least seven strains of Bremia /actucae and many cultivars
are resistant to the weaker but a few to the more aggressive strain; 'Mildura'
is resistant only to the mild strain. Some American cultivars such as
'Calmar' and 'Fransisco' segregate for resistance to the aggressive strain
and have been used by breeders. The species L. scariola, L. sa/igna and
L. virosa have some resistance to mildew. The best source of resistance to
lettuce mosaic virus is a South American cultivar 'Gallega' (Yonder Pahlen
and Crnko, 1965), which is symptom-less when infected and docs not
produce infected seed. There is no known source of resistance to beet
western yellows virus but cultivars show some difference in the time taken
to develop disease symptoms.
Autotetraploid forms of some cultivars have been produced (Eenink
and Alvarez, 1965). They grow very vigorously but develop looser heads
and mature later than diploids; they are also rather infertile and produce
very small amounts of seed and arc not therefore commercially viable.
Gibberellic acid has been used to induce early seeding in lettuce and to aid
seed stalk development in seed crops of tight heading cultivars (Harrington,

8.3.4 Pea
The cultivated garden pea is generally included in the species Pisum
sativum and the field or agricultural pea in P. sativum arvense or as a
separate species P. arvense. These two forms differ mainly in one major
gene A which in the dominant condition gives coloured flowers, brownish
Seed Propagated Cu/tivars 115

seed coat and anthyocyanin coloration at the base of the stipules. Horti-
cultural cultivars are almost all recessive aa with white flowers and no dark
coioration of the stipules. However, some cultivars of sugar peas and old
Dutch cultivars of peas grown for their dry seeds for human consumption
carry the dominant A gene. Agricultural and horticultural forms readily
hybridise with each other when intercrossed by hand.
The pea has been intensively studied by geneticists following Mendel's
example. Over 300 genes are known for the species and there is consider-
able information also on linkage groups (Figure 1.5). Many of the genes
are for detailed seed-coat colour and for abnormalities of little interest to
the horticultural plant breeder, but the following major genes (dominant
character given first) are likely to be met in breeding work:-
(1) Seed
I green/yellow cotyledon
R round/wrinkled seed surface
(2) Pod
Bt blunt/acute pod apex
Cp curved/straight pod
Gp green/yellow pod
hard/soft (sugar pea) pod
~} both dominants required for normal
non-sugar pea type
N thick/thin pod wall (in sugar pea)
(3) Habit and foliage characters
Af leaves/tendrils only
Fa normal/fascinated stem
Fn One or two/three or more flowers per peduncle
Le tall/dwarf
Lf late/early flowering
St normal/reduced stipules
Tl tendril/no tendril to leaf
Uni normal/unfoliate leaf
Wb waxiness/no wax
Horticultural cultivars can be broadly categorised into four groups:
(1) grown for human consumption as dried seeds, (2) tall cultivars grown
primarily in gardens for fresh consumption, (3) dwarf cultivars grown for
processing by canning, freezing and dehydration and some also in gardens,
(4) sugar peas in which the whole immature pod is edible; they are largely
grown in gardens but on a small scale commercially for fresh consumption
in parts of Europe. Dwarf cultivars for processing are by far the most
important in Europe in terms of the area grown. Breeding is mainly for
improved yields and to extend the harvest season, but suitability for
116 Plant Breeding

processing is pre-eminent and can only be assessed satisfactorily by proces-

sing tests which arc generally solely available to the industry. Dark-green-
seeded types are required for freezing and dehydration but pale-seeded
cultivars are preferred for canning in Britain where some added colouring
matter is allowed.
Dry seed for human consumption is an important crop in some
countries but in Northern Europe wet weather in late summer makes it
difficult to harvest crops in good condition. Earlier maturing cultivars of
dry seed peas have created a renewed interest in this crop in Britain but it
is still a minor one. Interesting work is being done in England on afafstst
TIT/ types (Snoad, 1974) which have tendrils but no leaves and do not
lodge readily so that the plants hold less water and there is consequently
less damage to the drying seeds than in typical leafy types.
Very little recent work has been done on taller growing peas but there
is some interest in sugar peas. The latter are genetically ppvv or pp VV
types, the best beingpp vv NN genotypes.

8.3.5 Runner bean

This bean, Phaseolus coccineus (syn. P. multiflorus), is rarely grown as a
vegetable crop outside Britain. It outcrosses more readily than French
bean and although the flowers are usually self-pollinated they are so
constructed that this docs not readily occur unless they are, like the broad
bean, visited by insects. In addition to the tall climbing red-flowered
forms, there are recessive bush· and white-flowered types and some culti-
vars have pods which are relatively stringless.


From the breeder's point of view this is probably the most difficult group
of plants. Prior to 1960 most cultivars were developed by relatively simple
mass selection, but since then there has been much research into techniques
which take advantage or heterosis to give higher yields. The individual
crops differ considerably in their response to improved techniques.

8.4.1 Asparagus
About 100 species of Asparagus are known and a few are collected wild
for human consumption in Mediterranean areas; only A. officina/is is
cultivated. This Liliaceous species is dioecious (see Section 3.3), the male
plants arc usually more productive, and the 'spears' ~hey produce are of
better quality, than the females.
Plants can be propagated vegetatively, although less rapidly than by
seed. One way of improving the seed strain, therefore, is to test cross
individual plants and to maintain vegetatively only those clones which give
the best progenies when pair crossed. Seed cultivars produced by this
technique give approximately equal numbers of male (XY) and female
Seed Propagated Cultivars 117

(XX) plants but the females are usually rogued from the crop grown to
produce marketable spears because their spear quality and yield is inferior
to that of the males. Super mal~ ( YY) plants crossed with normal females
(XX) would theoretically give all male (XY) types from seed and roguing
would be unnecessary. Super male plants have been obtained by selfing
certain XY type plants which occasionally produce a few female flowers.
The progenies yield proportionately one YY, two XY, one XX and the
YY plants are recognised by their capacity to produce only male (XY)
plants when crossed with females; they are maintained vegetatively.
Asparagus occasionally produces seeds with twin embryos and one of
the twins is often a haploid (Marks, 1972). Since the haploids have devel-
oped from egg cells without fertilisation they are always the X type from
the XX female. Doubling the haploids by colchicine gives females which
are homozygous for all characters and can be used for crossing with
selected males to give more homozygous F 1 progenies than is possible
with heterozygous females.
In vitro techniques are used for culturing haploid embryos and have
been tested for rapid multiplication of clones. It seems likely that plants
vegetatively propagated in this way could be of considerable importance
to breeders in the near future.

8.4.2 Beetroot
Red beetroot, sugar beet mangold, swiss chard and spinach beet are all
forms of Beta vulgaris and readily intercross. Some wild sub-species have
been used in breeding experiments, but not with horticultural forms. At
least two genes govern root colour and red coloration is dominant to
yellow or white, but the intensity of red coloration is influenced by minor
Beetroot will set seed with its own pollen and pedigree selection can
be practised but soon leads to an undesirable reduction in yield through
inbreeding depression. As a rule, this crop is improved by mass selection or
by more sophisticated techniques utilising progeny testing. F 1 cultivars of
red beet are not yet used commercially though cytoplasmic male sterility
has been recorded in Beta vulgaris.
Flowers of beetroot are primarily wind pollinated and pair crosses can
be made by enclosing flowering shoots from two plants together in a
pollinating bag. Shaking the bag occasionally will effect pollination but
both self and cross seed will be produced and hybrids can only be dis-
tinguished from plants resulting from selfing if a suitable genetic marker
is available.
The main objectives in breeding red beet are high yields of dark-red
uniformly-coloured roots of uniform shape, and lack of proneness to bolt-
ing. Recently there has been some interest in transferring the monogerm
(single-seeded fruit) character from sugar beet to red beet to facilitate
singling of the plants. Assessment of root colour is made visually on
118 Plant Breedina

mature roots by removing a wedge of flesh with a sharp knife or a plug

with a cork borer. Selected sampled roots can be grown on to produce
seed. Growing in relatively infertile soils accentuates colour differences
and makes it easier to choose the darkest coloured roots. Premature bolt-
ing occurs most readily in long days and at relatively low temperatures.
Early sowing and growing at northerly latitudes encourages bolting and
can aid selection for resistance to this defect.

8.4.3 Brassica crops

Cabbage, Brussels sprout, cauliflower, broccoli, calabrese, kohl rabi and
most agricultural and horticultural kales are all forms of Brassica oleracea
and readily intercross. Summer cauliflower, and to a lesser extent early
round-headed summer cabbage, are self-fertile and do not suffer excessively
from inbreeding but plants of nearly all other forms are self-incompatible
and selfed progenies exhibit considerable inbreeding depression. To effect
selfing it is usually necessary to bud pollinate by hand to overcome the
natural incompatibility system.
Brassica oleracea will hybridise with difficulty to turnip and radish
but the hybrids are mainly sterile unless chromosome doubling is induced.
The first of these combinations is similar to the swede and the second,
called Raphanobrassica, is of considerable academic interest but has not
yielded any useful horticultural cultivars.
Mass selection and pedigree breeding is still used for early cauliflower
and summer cabbage. Mature cauliflower plants selected for breeding are
often difficult to seed because relatively few of the flowers develop and
survive rotting by pathogens. This has been overcome by vegetatively
propagating the plants from cuttings of curd portions, each with a piece of
leaf, and by grafting of curd portions (Crisp and Lewthwaite, 1974) on
seedling cauliflower plants. Most forms of B. oleracea can be maintained
as perennials by vegetative propagation from stem, leaf-bud and sometimes
root cuttings when it is desirable to keep a certain genotype as a clone for
breeding purposes. Hygienic conditions must be maintained to prevent
rotting, the temperature must be kept above about 15°C to maintain the
plants in the vegetatjve phase and precautions should be taken to prevent
infection by cauliflower mosaic, turnip mosaic and broccoli necrotic
yellows virus all of which are aphid transmitted.
Most modern Brussels sprouts are F 1 cultivars and the technique is
now being used for all other forms of Brassica o/eracea including, recently,
summer cauliflower. Crossing between the two parent lines is achieved by
having each homozygous for different 5 alleles. The parent lines are then
self-incompatible and, theoretically, produce no seed with their own
pollen when grown together as an F 1 seed crop, so that all of the seed
results from hybridisation between the two lines. In practice, the parents
do produce some selfed seed which gives rise to 'sib' plants. The propor-
tion of sibs should be below 5 per cent of the F 1 seed; when it rises above
Seed Propagated Cultivars 119

10 per cent the seed is usually considered to be unsaleable. High tempera-

ture, shortage of pollinating insects, and several unknown factors affecting
the seed crop can give rise to a high proportion of sibs. Sibbing is also
affected by the genetic constitution of the parents. Plants homozygous for
the so-called 'weak' incompatibility alleles such as 52, 55 and 575 are
especially likely to produce sibs and research is at present in progress to
identify 'strong' incompatibility alleles and to incorporate them into such
important horticultural forms as brussels sprout and cabbage. Before a
crop of F1 seed can be sold, it has to be tested for percentage sibs. This
is usually done by a 'growing-on test' in a glasshouse during the winter
following the harvesting of the seed but is usually a lengthy and rather
inaccurate test because sibs cannot always be recognised with certainty,
especially at an early stage of growth. Recently a laboratory test has been
devised whereby extracts of crushed seeds or of two- or three-day-old
seedlings are examined by electrophoresis for their enzyme content. This
gives a rapid and certain result but cannot be used for some 10 per cent
of F1 genotype combinations and is therefore not applicable with a few
F1 cultivars (Wills eta/., 1978).
F1 parent lines are produced by enforced breeding through some six
generations by hand bud pollination and it is usually desirable to select
them in each inbreeding generation on the basis of their performance when
crossed with an unrelated line. They must of course be homozygous for a
known 5 allele and this can be assessed by test crossing to a collection of
brassicas of known incompatibility status-a service provided by some
research institutes.
Several major genes are known for B. oleracea but most characters of
agronomic importance are polygenic. White flower, which is found in some
cauliflowers, is dominant to yellow, male sterility (abortive pollen) is
recessive to fertility; there are several genes for 'glossy' leaves lacking the
typical waxy bloom and nearly all are recessive to the normal condition;
inheritance of anthocyanin coloration is more complex but purple colora-
tion is mainly dominant to green.
The breeding objectives for all forms of horticultural brassicas usually
encompass increased yield, uniformity and lack of propensity to premature
bolting. Some of the objectives for specific crops include (1) Brussels
sprout-firm, medium-sized sprouts with few rough outer leaves and not
subject to internal browning (a physiological disorder of complex inherit-
ance); (2) cabbage-firm heads with short internal stems which do not split
when overmature, and late forms not subject to frost damage or rotting
of the outer leaves; (3) cauliflower-white curds without bracts in the curd
and not subject to riciness, or lack of buttoning (premature development
of very small poor quality curds); (4) calabrese-spears held well above the
foliage; (5) kohl rabi-lack of splitting.
Clubroot caused by the fungus Plasmodiophora brassicae is a serious
disease of brassica crops in some areas; there are several strains of the
120 Plant Breeding

fungus and no cultivar is resistant to all of them. Breeding for resistance

within Brassica o/eracea is a lengthy process but significant progress has
been made in recent years. There is a possibility of using some forms of
radish as a source of resistance to clubroot in B. oleracea hybrids and this
has been achieved in forage brassicas. Downy mildew caused by Erisyphe
polygoni has become troublesome during the last decade, especially on
Brussels sprout, and cultivars show differences in susceptibility so that it
is possible to breed for resistance. There seem to be few genetic differences
in susceptibility to other pathogens in B. o/eracea.
As a rule it is not necessary for breeders to store brassica pollen. Seeds
retain their viability for at least six years when kept dry at room tempera-
ture and very much longer when stored in deep-freeze.

8.4.4 Carrot
The wild carrot Daucus carota is widespread and common in Britain and
other parts of Europe. It hybridises readily with cultivated forms and can
cause genetic deterioration by pollen contamination of seed crops though
its peak flowering period does not coincide with them. The white root of
the wild form is dominant to the orange-red flesh of cultivated types and
white-rooted plants in commercial crops probably originate from hybridi-
sation of the seed crop with wild carrot.
Carrots suffer inbreeding depression but individual genotypes differ
in the degree of inbreeding which can be imposed on them without serious
loss of vigour. Plants can be selfed and crossed by the technique described
in Section 3.5. Most cultivars have been bred by simple mass selection
techniques but in recent years there has been much interest in the produc-
tion of F 1 cultivars. Control of crossing between parent lines is obtained
by male-sterility, of which there are two types. In the 'petaloid MS' form,
petal-like structures take the place of stamens and in some forms the true
petals are especially small and purplish-coloured instead of white. The
'brown anther MS' form has more or less normal anthers but produces no
fertile pollen. Petaloid male sterility is controlled by S cytoplasm as
compared with N cytoplasm in conjunction with two genes, one recessive
a and the other dominant B (Banga et of., 1964}. All genotypes with aa,
BB, or Bb and S cytoplasm are male sterile. To produce an F 1 cultivar a
fertile parent is crossed to a male-sterile parent, for example aabb S or
AABB S. The male sterile line is maintained by crossing two lines which
are genetically similar in all respects, except that one is an aa S or BB S
and the other respectively aa N or BB N type. Three lines are required to
maintain the hybrid. The situation is similar for the brown-anther type of
male-sterility, although in both cases more genes which restore fertility
may eventually be found.
Objectives in carrot breeding are improved yield and uniformity of
root shape and size, uniform root colour, lack of bolting and splitting.
Uniformitv of root size is complicated by the fact that size is closely
Seed Propagated Cultivars 121

related to time of emergence of the seedlings and this in turn influenced

by the stage of development of the embryo within the dry seed. Some of
the 'seeds' {single seeded fruits) have immature embryos, according to
their stage of development at the time of harvesting the seed crop. After-
ripening of the seed before sowing can improve uniformity of seedling
emergence but it may also be possible to improve embryo uniformity by
breeding selection. Premature bolting is encouraged by cool weather and
selection against this defect is best done in northerly parts of Europe.
Carrot seeds lose viability fairly quickly and need to be stored with

8.4.5 Celery
Celery and celeriac are forms of wild celery (Apium graveolens), a relatively
uncommon plant of wet areas of Europe. From a breeder's point of view
they are similar in many respects to carrot and flies can be used to effect
crossing with that crop (Honma, 1959).
Objectives for celery breeding include non-bolting, lack of pithiness,
and for celeriac, non-bolting also, and 'bulbs' with few basal roots and
lacking hollow cavities. Celery 'seed' is relatively small and individual
plants give high seed yields.

8.4.6 Cucumber
The cucumber Cucumis sativus belongs to the same genus as the musk
melon C. melo, but the two species do not hybridise.
Most modern cucumber cultivars are F 1 and these are relatively simple
to achieve without any risk of admixture by sibs. The plants are mono-
ecious so that emasculation is unnecessary when the plants are grown
under glass with insects excluded, and hand-pollination is economically
feasible because each individual pollination operation produces a fruit
with a large number of seeds. Insects can be used to effect crossing if the
male flowers are removed from plants of the parent destined to act as
the seed parent. Several chemical substances have been tested to bring
about pollen abortion or non-production of male flowers but are seldom
used to produce commercial seed crops. A genotype which produces only
female flowers is known. It can be maintained by selfing from male
flowers induced by spraying with gibberellin {Peterson, 1960), or silver
At least 20 genes are known for cucumber including S spiny/nearly
spine free fruits, A spines/no spines or fruits, 8 black/white spines, G dull/
glossy skin, Te tough/tender skins, Bt bitter/non-bitter taste, I determin-
ate/indeterminate growth habit, T tall/short plant height. Freedom from
bitterness is an important breeding objective and a technique to achieve
this was described by Andeweg and de Bruyn (1959).
122 Plant Breeding

8.4.7 Leek
The leek (Allium porrum) is thought to have been derived from the sand
leek (A. ame!oprasum) which grows wild in southern Europe, N. Africa
and the Middle East. Nearly all cultivars are tetraploids but wild A. ampe/o-
prasum may be diploid (2n = 16), tetraploid and rarely hexaploid.
Most cultivars have been bred by mass selection. Pedigree selection is
not favoured because the leek, although it is a tetraploid, suffers rather
severely from inbreeding depression. Attempts have been made to utilise
cytoplasmic male sterility to produce F1 cultivars but none is yet available
In some areas of Britain leek rust caused by Puccinia porri can be very
troublesome. Cultivars differ in their degree of susceptibility and breeding
for resistance would be a useful objective.

8.4.8 Marrow
The type of marrow grown in Britain is a form of the species Cucurbita
pepo which also embraces many kinds of squash, ornamental gourd and
some types of pumpkin. Some cultivars of two similar species C. maxima
and C. moschata are also known as squashes and pumpkin. Both of these
can be crossed with difficulty to C. pepo, especially with the aid of embryo
culture, but the hybrids are very infertile or completely sterile.
The marrow is monoecious and it is easy to produce F1 cultivars by
removing the male flowers from the line grown as seed parent and allowing
insects to effect cross pollination. A monogenic recessive condition for
male-sterility is known in C. pepo; staminate flowers are produced but the
pollen is not viable.
Green fruit skin colour is recessive to white or yellow and the warty
surface of some squashes is dominant to the smooth skin of the marrow.
The white striped fruit coloration is dominant to yellow striping.
Marrow chromosomes are very small and difficult to count in root tip
squashes. Groups of root cells may be tetraploid and give the impression
that the cultivar is tetraploid whereas as a rule the greater part of the root
system and the whole of the above-ground parts of the plant are diploid.

8.4.9 Melon
The species grown in Britain is the muskmelon (Cucumis melo). It differs
in several respects from the water melon (Citrullus vulgaris) and does not
hybridise with it. Like other Cucurbitaceae the melons are monoecious.
Mass selection and line breeding have been used for muskmelon and
there is now considerable interest in F 1 cultivars. These can be produced
by hand pollination and removal by hand of the male flowers of one of
the parents. Experiments have shown that it is also possible to prevent the
development of male flowers by treatment with ethrel (see Section 3.7).
Tetraploids have excellent fruit flesh quality but are later flowering
and less productive than diploids. Triploids are productive and have high
Seed Propagated Cultivars 123

fruit quality but are not as easy to produce in muskmelon as in water-

melon. The cross of tetraploid x diploid is only possible when the tetra-
ploid is used as the female parent and even then seed set is very poor
with muskmelon. Triploids are almost entirely sterile and fruits develop
parthenocarpically, which is a distinct advantage since they contain no
Interspecific hybridisation of C. melo with C. metiliferus has been
used in the United States to transfer disease resistance.

8.4.1 0 Onion
The bulbing onion grown in Europe is Allium cepa, differing from the
non-bulbing Welsh onion and the Japanese bunching onion which are
forms of A. fistu!osum. These two species can be crossed but the diploid
F 1 is sterile. Colchicine treatment to double chromosome numbers pro-
duces fertile amphidiploids and A. x cepa-fistu!osum cultivars have been
released in the United States as very vigorous bunching onion types. The
shallot, usually referred to as A. asca!onicum, is fully fertile with the onion
and is probably merely a form of A. cepa which proliferates into a cluster
of bulbs.
Onions show considerable inbreeding depression and the pedigree
breeding method is not effective with this crop. Techniques involving
alternate inbreeding and outbreeding generations are preferable and the
F 1 technique is used to produce many modern cultivars which are not
only more uniform but often mature earlier than conventional types.
Cytoplasmic male sterility, first made use of to produce F 1 cultivars of
onion, is the classic example of this technique.
The first F 1 cultivars were produced in the United States by maintain-
ing the male-sterile line vegetatively from small viviparous bulbils which
developed in the seed head. This system was abandoned because of the
difficulty of keeping the stock free from yellow dwarf virus. Present F1
cultivars require three parent lines generally referred to as A, B and C.
These are: the female A line (S ms ms) which is male sterile and produces
the hybrid seed, the n~tainer B line (N ms ms) which is genetically similar
to A but produces pollen, the male C line (N Ms Ms) which is genetically
dissimilar to the other two and chosen for its capacity to give a good
hybrid with the A line. S and N refer to sterile and normal types of cyto-
plasm. The B and C lines are produced by growing batches for seed in
insect-proof cages or as well isolated crops in the open. Line A can only
be produced by crossing with B. Blowflies are used for pollination of small
batches of the parents (see Section 3.5) but the natural insect fauna is
relied on for outdoor seed crops. Starting a search for male sterility from
basic material would- be a lengthy process but the newcomer to the breed-
ing of this crop can obtain the S cytoplasm and the recessive ms gene from
current F 1 cultivars which are fertile SmsMs forms and can be selfed to
give )4 sterile Smsms types. These cannot be used directly as they are very
124 Plant Breeding

heterozygous for other horticultural characteristics and several generations

of breeding are required to improve homozygosity.
High yield, uniformity, earliness- and long storage life are the main
objectives in onion breeding. Resistance to neck rot caused by Botrytis
a/Iii is important and there are cultivar differences, the most pungent being
the most resistant; pungency is itself an important feature of cultivars
grown primarily for dehydration. Resistance to the smut fungus (Urocystis
cepu/ae} would be helpful but is not found in A. cepa; however, A. fistula-
sum is highly resistant and some A. cepa x fistufosum hybrids are resistant
but non-bulbing.
There are at least two genes for bulb coloration in onion. I inhibits
all coloration whereas ii allows expression of the gene W which is thought
to have at least three alleles: W for red pigment, Wy for yellow and the
recessive ww for white. Thus Wii is red, Wli has a red neck only, Wyii is
yellow and WI/, w/1, wli, wii are white.
The genus Allium contains some 500 species and the potential gene
pool for the onion has not been fully exploited. It includes also chives
(A. schoenoprasum}, garlic (A. sativum}, leek, Welsh onion and several
native British species of wild garlic.

8.4.11 Parsnip
This is a yellow-flowered umbelliferous plant derived from the wild
parsnip Pastinaca sativa. Little breeding work has been done with this
crop, except to produce cultivars resistant to canker. Like the carrot,
parsnip flowers are protandrous and crossing can be controlled in a similar
way. Parsnip roots are fully winter-hardy and when selected they can be
replanted to flower and seed the following year, unlike mature carrot roots
which need more careful treatment and are usually kept in pots under glass
for the winter.

8.4.12 Radish
The cultivated forms of Raphanus sativus probably derive from the wild
radish R. raphanistrum, which can be found as a weed of arable land in
Britain. In addition to the relatively small-rooted and quick maturing types
grown in Britain, there are maincrop cultivars with large long roots having
white or rough dark grey or black skins grown in japan and parts of
eastern Europe, and others grown for their foliage as a fodder crop or for
human consumption as leaf salad.
The radish is a crucifer and, from a breeder's point of view, similar in
several respects to Brassica crops. It has an incompatibility system and
suffers from inbreeding depression, though self-fertilisation can occur.
Cytoplasmic male-sterility is known in radish and it would seem advan-
tageous to produce F 1 cultivars. However, seeding population rates for a
radish crop are high and the individual seeds are much larger than those of
B. oleracea so that seed costs are an important part of root production.
Seed Propagated Cultivars 125

Thus, F 1 seed is likely to be too expensive for commercial root production

and breeders have examined the possibility of producing F2 cultivars, but
none is yet available commercially (Bonnet, 1975).
Several genes control root colour but in practice white is recessive to
coloured roots and crosses between white and coloured roots give mainly
coloured progenies. The black and corky skinned characters are dominant
or epistatic to all other colours and to smooth skin. Some studies indicate
that long root is dominant to short round root, but several genes may
control this character.

8.4.13 Spinach
True spinach Spinacia oleracea should not be confused with New Zealand
spinach (Tetragonia expansa) or spinach beet which is a form of Beta
Cultivars of true spinach are sometimes classified into winter and
summer types. The latter were once traditionally prickly seeded types in
which the pseudocarp (spinach 'seed' is a one-seeded fruit) carried one to
four spiny projections. This character, controlled by a single dominant
gene, is not linked with hardiness and most winter cultivars have round
seed which is easier to sow than prickly seed.
Spinach is dioecious and male plants are often but not always less
'leafy' than females. However, plants frequently produce some flowers
which are partially or completely hermaphrodite (Figure 8.1). The situa-
tion is explained when sex is primarily determined by the sex chromosome
in which XX is female and XY male but with two other closely linked
modifying sex genes A and G. XXAAGG, XXaagg, XXAaGg plants are
female; XYAAGG, XYaagg, XYAGg male; other combinations are herma-
Until recently spinach cultivars were bred by mass selection but self-
ing can be practised to improve uniformity by growing hermaphrodite
individuals well-isolated from each other in the field. Isolating the plants
in bags or cages usually gives poor seed sets. F 1 cultivars can be produced
by growing selected parent lines in alternate rows and hand roguing all but
the male plants from one line and all but the females from the other.
However, this is an expensive method. Sneep (1957) described a technique
in which 'super females' XXaaGG types obtained by selfing hermaphrodites
are used. The following three lines are required: a population of XXaaGG
super female and XYaaGG hermaphrodite types which reproduce approxi-
mately equal numbers of each type, (2) a hermaphrodite XXAAgg type
which breeds true for sex character, and (3) an unrelated pollen line which
segregates for male, female and hermaphrodite types. Seed of (1) is grown
alongside that of (2) and hermaphrodite types rogued from (1 ). The
remaining super females will produce seed with the constitution XXAaGg,
which is all female. When this is sown as a seed crop with (3) the seeds
from it will be all hybrid and hand-roguing is not necessary in this final
126 Plant Breeding

Figure 8.1 Flowers of a, male; b, hermaphrodite and c, female plants of spinach

(Spinacia oleracea). {After Sneep, 1957.)

large scale stage of seed production. If a round-seeded form of the seed

parent and a prickly-seeded form of pollen parent is used, the seed of the
two can be mixed when the seed crop is sown. Each parent will produce
seed morphologically chracteristic of its kind since prickliness is a feature
of the maternal tissue and not a product of fertilisation. Round- and
prickly-seeded types can easily and accurately be separated by machine.
Spinach produces flowers in long days and selection for late bolting
is best done in northerly latitudes. Two generations of seed can be obtain-
ed in a glasshouse by growing under artificial long days at about 20°C.
The main objectives in spinach breeding are uniformity, delayed bolt-
ing, unproved leafiness of bolting male types, hardiness and resistance to
downy mildew caused by Perenospora effusa and to cucumber mosaic
virus. Resistance to both is found in cultivated types and is dominant. In
producing an F 1 only one parent need carry dominant homozygous
resistance genes.

8.4.14 Swede turnip

As we have seen, the swede (Brassica napus) is an amphidiploid (2n = 38)
which originated from a cross between B. campestris (2n = 20) and B.
Seed Propagated Cultivars 127

oleracea (2n = 18) and is of similar genetic constitution to rape and rape
kale. The combination has been resynthesised by breeders. Very little
work has been done by horticultural breeders, and the 'garden' swedes are
largely a byproduct of breeding agricultural cultivars. The swede is self·
fertile but cultivars intercross with each other and with non-bulbing
forms. The swede has a higher degree of resistance to clubroot than the
true turnip.

8.4.15 Turnip
Generally referred to as Brassica rapa, the wrnip and Chinese cabbage are
forms of Brassica campestris. The turnip will cross with swede, but the
hybrids are mainly sterile, and it is interfertile with Chinese cabbage. It has
an incompatibility system as in B. o/eracea, although selfing can take place
fairly readily in the open-flower stage with many genotypes. The violet-
red colour of the root is dominant or epistatic to white and yellow is
recessive to white.

8.4.16 Tomato
The tomato is typically Lycopersicum esculentum but several other South
American species, notably the small-fruited red currant tomato L. pimpin-
el/ifo/ium and also L. hirsutum and L. peruvianum, have been used by
breeders. All these species are diploid (2n = 24}.
In the typical tomato flower a cone of stamens envelops the stigma
and when the pollen is shed it effects self-fertilisation. This inbreeding
does not lead to severe loss of vigour and until the 1950s nearly all tomato
cultivars were self pollinated. Now most cultivars grown in Britain are
F1 s, which are more uniform and higher yielding than the older types.
Emasculation and crossing is usually by hand, an expet1sive operation, but
each fruit bears 100 or more seeds and the glasshouse tomato industry can
absorb fairly high seed costs. Attempts have been made to reduce F 1 seed
production costs for outdoor crops by using male-sterile types. Several
genes for male-sterility are known, including the so-called 'John Baer' type
of functional male-sterility in which fertile pollen is produced but the
anthers do not dehisce . without artificial aid. A condition known as
'exerted-style', in which the style is longer than the cone of stamens and
the stigma is held clear of the anther so that self-fertilisation occurs less
readily, is of interest to breeders. It cannot always be relied on to exclude
selfing but it makes emasculation easier. However, F 1 cultivars should not
themselves have a pronounced exerted style as pollination may not be so
easily effected under glass and fruit set may be poor, though some culti-
vars set fruit parthenocarpically if they are not fertilised.
Like peas and Phaseolus beans, the tomato is a crop which has receiv-
ed much attention from breeders. There are international associations for
each of these crops for breeders to communicate up-to-date advances in
techniques and to share ideas and breeding material. The genetics of
128 Plant Breeding

tomatoes is better known than that of most vegetable crops and the
following major genes for morphological characters are of special interest
to breeders.
(1) Foliage
c Normal cut leaf/'potato leaf'
(2} Habit
D tall/dwarf
Sp normal/determinate, 'self-pruning'
5 simple/much-branched compound, inflorescence
(3} Fruit
F normal fasciated shape of 'Marmande' type
N normal/nipple-tipped
0 normal/elongated shape
P smooth/hairy
R red/yellow fleck
T red/tangerine-coloured flesh
U dark base when ripe/non-'greenback' 'Stonor' type
Y yellow/transparent skin
Plants with RY have red, Ry pink, rY dark yellow and ry light yellow
to white-coloured fruits.
Disease resistance is important in tomato cultivars and several genes
are available: Ve for resistance to Verticil/ium, Tm-7, Tm-2 and Tm-2 2 to
tomato mosaic virus of which there are at least five isolates and Cf to
Cladosporium fulvum. Genetic control is known also for some resistance
to Fusarium oxysporum, Phytophthora infestans and to corky root caused
by Pyrenochaeta lycopersici. Many of these genes have been obtained from
L. pimpinellifolium and some from L. hirsutum, L. peruvianum and L.
glandolusum. Disease resistance genes are mostly dominant and can be
easily incorporated in F 1 cultivars by having one parent homozygous for
Tomato seed retains its viability for several years even when stored at
room temperature but the pollen is short-lived and easily damaged by
temperatures below freezing.


It is not possible to deal with these individually in any detail but the
importance of F 1 cultivars will be discussed. At present commercial F1 s
are offered in some 30 species in the following genera: Ageratum, Antir-
rhinum, Aqui!egia, Begonia, Bellis, Calceolaria, Chrysanthemum, Coleus,
Cyclamen, Dianthus, Hibiscus, Impatiens, Nicotiana, Papaver, Pelargonium,
Petunia, Portulaca, Primula, Saintpaulia, Salpiglossis, Sinningia (Gloxinia),
Tagetes, Viola and Zinnia. The cost of F1 seed compared to that of a
conventional cultivar may vary from two to three times higher for Begonia
to 25 times higher for Zinnia. Advantages of F 1 seed are improved vigour
Seed Propagated Cultivars 129

with three to four times the number of flowers per plant in some Bellis F 1 ,
improved uniformity and utilisation of fewer seeds of types which have
to be pricked oat and germinate uniformly, and greater earliness for
forcing-up to four weeks with some Primula and Calceolaria hybrids.
Pollination to produce the F 1 is often done by hand but incompati-
bility systems in Bellis and Ageratum allow for crossing by bees. The
heterostyle type of incompatibility of Primula cannot easily be used with
insects because flowering occurs early in the year when they are not
usually available.
Monogenic male-sterility occurs in Tagetes erecta, Zinnia and also in
Antirrhinum where a functional male-sterile type is known. However, as
true breeding male-sterile lines cannot be produced, roguing of plants at
flowering time is necessary and this may be expensive if heated glasshouse
space has to be allocated to plants which will eventually be discarded.
Cytoplasmically controlled male-sterility is available in Petunia, tuberous-
rooted Begonia and Aquilegia. In Petunia, however, the condition is
associated with flower abortion.


Seed-propagated cultivars are not static, for each time they are propagated
slight genetic changes can, and usually do, occur. To counteract these
changes it is necessary to apply selection pressure to each seed crop,
though this may be negative selection-roguing to remove off-types. Thus
seed production is an extension of breeding. Indeed there is not always a
clear distinction between the two and seedsmen sometimes make a deliber-
ate attempt to 'improve' the stock by selection during propagation rather
than maintain it without obvious change.
There are three distinct stages in seed production of conventional
cultivars to provide: (1) 'nuclear', (2) 'stock' and (3) 'commercial' seed.
Nuclear seed is produced only in very small quantities by the breeder or
maintainer of the cultivar and under the strictest precautions to ensure
that it is true to type. Sometimes it is produced from vegetatively main-
tained clonal plants.
Stock seed is produced from nuclear seed or from small quantities of
an earlier batch of stock seed known to be of high standard. As a rule, the
plants grown to produce stock seed are sown and treated in much the same
way as they would be to produce a market crop, so that their performance
can be monitored at all stages of growth and any deviant types discarded.
This requirement may introduce practical problems, especially for biennial
crops which have to be over-wintered in the mature state. Large mature
carrots do not survive well in the open and are sometimes lifted in the
autumn and grown to produce stock seed under glass. Mature cabbage
heads are often so solid that the main seed stalk cannot penetrate ami the
heads have to be cut crosswise on the upper surface several weeks before
130 Plant Breeding

the seed stem elongates to relieve the tension-an operation called 'racing'.
With the less hardy cabbage types and with spring cabbage the heads are
cut leaving a stalk from which flowering side shoots can grow-a technique
known to seed growers as 'stump seeding'. In parts of northern Europe
mature storing type cabbage are lifted and replanted in a trench so that the
top of the head only is above ground level.
Stock seed is usually grown by the seedsman or by the official body
responsible for maintaining the cultivar.

8.6.1 Commercial seed production

The large-scale production of seed for sale to commercial growers and in
packets for gardeners is usually carried out by specialised seed growers on
contract to a seed firm which provides the stock seed to raise the crop. As
a rule the site of the crop is decided in consultation between the seed
firm and the seed grower, and the contract usually states that the entire
crop of seed will be sold to the firm at an agreed price per kilogram me and
none retained by the grower for other purposes. Representatives of the
seed firm generally inspect the crop during the growing season and rogue
out any off-types. Sometimes surplus seed from a previous commercial
seed crop may be used for sowing instead of stock seed, especially with
peas and beans, but this is considered to be bad practice with most crops.
Relatively little seed of horticultural crops is grown now in northern
Europe; seed plants are raised mostly in warmer drier climates as in the
Mediterranean region, California and Idaho in the USA, parts of Africa
and Australia. However some commercial seed crops, especially of
ornamentals and cauliflower, are produced under glass in northern Europe.
Seed crops require frequent inspection and careful attention to details,
and are usually grown by experienced specialists. Seed from a good crop
of one hectare of Brussels sprouts may be worth £50,000 on the retail
market, but poor crops and complete crop failures due to uncontrollable
circumstances are common.
It is important to isolate seed crops of most species from one another
to prevent genetic contamination by cross fertilisation. In the Netherlands,
minimum isolation distances are prescribed by the Dutch Seed Association
(NAK-G) and an abridged version of the requirements is given in Table 8.3.
With commercial seed crops the plants are grown to obtain maximum
seed yield. For this reason biennials are sown later in the year than is
customary for market crops so that the plants are smaller at the onset of
winter and withstand cold weather with relatively little damage. Small
plants of this kind, which are often transplanted, are generally referred to
as 'stecklings '. It is not so easy to rogue off-types from these small plants
but this is less important than at the stock seed stage.
When mature, most horticultural seed crops are cut and stooked to
ripen before threshing, but direct combine harvesting is possible with some
crops. As a rule, the crop is threshed from the stooks using a combine
Seed Propagated Cultivars 131

Table 8.3

Minimal isolation distances required by Dutch Seed Association

(adapted from Bango, 1953)

Red beet, sugar beet and mangold 600 m
Different cultivars of red beet 200-400 m
Brassica oleracea
Different kinds of B. olerocea 600 m
Different cultivars except cauliflower 400 m
Different cauliflower cultivars 100m
Broad bean
Threefold white and other cultivars 100m
Other different cultivars 25-50 m
Different types 400 m
Different cultivars within a type 100m
Ridge cucumber and gherkin 400 m
Different cultivars 200m
Red cultivars and others 600 m
White cultivars and others 600 m
Different cultivars 200-400 m
Fodder and horticultural cultivars 600 m
Different types of horticultural cultivars 200m
Different cultivars within a type 50 m
Different cultivars 300m
Agricultural and horticultural cultivars 400 m
Different horticultural cultivars 200m
Turnip, turnip rape and Chinese cabbage 400 m

10m Cal/istephus, Matthiola
50 m Chrysanthemum leucanthemum, Dianthus
10-100 m Viola, Calendula
50-200m Clarkia, Godetia, Lobelia, Petunia, Phlox drummondi, Verbena
100m Gaillardia, Tagetes
100-400 m Antirrhinum, Begonia, Campanula medium, Centaurea cyanus,
Cheiranthus, Digitalis, Nemesia, Salvia splendens

harvester as a stationary thresher, carefully adjusting the drum speed and

concave setting to suit the seed. Radish seed is difficult to thresh and some
breeding work has been attempted to improve cultivars in this respect
(Banga et a!., 1965). French bean seed is easily damaged by incorrect
threshing and the germination may be seriously impaired.
132 Plant Breeding

Some species need individual treatment. Tomato and cucurbit fruits

are usually crushed and the pulp extracted by acid (see Section 4. 7).
Onion and leek heads are cut individually and dried on sheets or in bags
in an airy shed before threshing; leek is difficult to thresh unless the heads
are well dried. Seed heads of most ornamentals are picked individually as
they ripen. A few, such as Viola, eject the seed when ripe and have to be
left in a covered container otherwise seed will be lost.

ANDEWEG, j. M. and DE BRUYN, ). W. (1959). Breeding non-bitter cucumbers,
Euphytica, 8, 13-20
BANGA, 0. (1953). lnleiding tot de plantenveredeling, W. E. J. Tjeenk Willink,
Zwolle, 635 pp
BANGA, 0., PETIET, j. and VAN BENNEKOM, j. L. (1964). Genetical analysis of
male-sterility in carrots, Euphytica, 13,75-93
BANGA, 0., PETIET, j. and VAN BENNEKOM, j. L. (1965). Selecting radish for
more easily threshable siliquae, Euphytica, 14, 49-58
BONNET, A. (1975). Introduction et utilisation d'une sttlrlilte male cytoplasmique
dans des varithes precoces europeenes de radis (Raphanus sativus L. ), Ann.
Ametior. Plantes, 25,381-397
CRISP, P. and LEWTHWAITE, j. j. (1974). Curd grafting as an aid to cauliflower
breeding, Euphytica, 23, 114-120
EENINK, A. H. and ALVAREZ, J. M. (1975). Indirect selection for tetraploids in
lettuce (Lactuca sativa, L.), Euphytica, 24,661-668
HARRINGTON, j. F. (1960). The use of gibberellic acid to induce bolting and
increase seed yield of tight-heading lettuce, Proc. Am. Soc. hort. Sci., 75,
HONMA, S. (1959). A method of celery hybridisation, Proc. Am. Soc. hort. Sci., 73,
LINDQVIST, K. (1960).1nheritance studies in lettuce, Hereditas, 46,387-370
MARKS, G. E. (1972). Selecting asparagus plants as sources of haploids, Euphytica,
PETERSON, C. E. (1970). A gynoecious inbred line of cucumbers, Quarterly Bufl.
Michigan Agr. Exp. Sta., 43, 40-42
ROWLANDS, D. G. (1964). Fertility studies in the broad bean (Vicia faba L.),
Heredity, 19,271-277
SNEEP, j. (1957). De stand van de veredeling bij spinazie, Meded. lnst. Vered Tuin-
bouwg., 13,1-128
SNOAD, B. (1974). A preliminary assessment of 'leafless peas', Euphytica, 23,257-
VON DER PAHLEN, A. and CRNKO, J. (1965). El virus del mosaico de Ia Iechuga
(Mamor lactucae Holmes) en Mendoza y Buenos Aires, Rev. Invest. Agro-
pecuarias, 11, 25-31
WILLS, A. B., FYFE, S. K. and WISEMAN, E. M. (1978). Testing F 1 hybrids of
Brassica oleracea for sibs by seed isoenzyme analysis, Ann. appl. Bioi., to be
WILLS, A. B. and NORTH, C. (1978). Problems of hybrid seed production Acta
Hort., 83, 31-36

FOOD AND AGRICULTURE ORGANISATION (1961). Agricultural and Horti-
cultural Seeds; Their Production, Control and Distribution, Panetto and Petrelli,
Rome, 531 pp
JANICK, j. and MOORE, j. N. (1975). Advances in Fruit Breeding, Purdue Univer-
sity Press, West Lafayette, Indiana, 623 pp


The selection of potential newcultivars is usually the most time-consuming

aspect of a breeding project. Only when a limited number of new geno-
types can be produced, as with some difficult wide crosses or mutation
treatments, does selection play a minor role. As a rule the larger the
number of new individuals produced, or families ·in the case of seed
propagated plants, the greater will be the chance of selecting promising
new cultivars. Especially large numbers of new individuals have to be
raised for species which have already received much attention from breeders
if there is to be a reasonable chance of producing a superior new cultivar.
Because of the time required for selection, and the land area needed to
raise the volume of material, most amateurs are usually at a disadvantage
compared to professional breeders working with such crops as Brussels
sprouts, potatoes and strawberries. However, there is considerable scope
for amateur breeders with ornamentals and some of the lesser known
fruits and vegetables.
The breeding of horticultural crops is still largely a personal activity.
Even when a team of workers is employed on a project and the objectives
are clearly defined, it is important to have a leader to stamp his or her
individual mark on the work-like a conductor of an orchestra. For this
reason a project should not expand beyond the capabilities of the leader
of the team to supervise the selection. In practice this means that for
max1mum efficiency with most crop plants a breeder has to raise 5,000
to 10,000 new individuals for selection each year. With fewer than that
the chances of obtaining a successful new cultivar are low, and above that
the team leader may not be able to cope with the selection. If a greater
effort is needed then it is probably advisable to set up a second independ-
ent breeding team to complement the first rather than expand the existing
Selection of the end product of the breeding work, as distinct from
selection during back-crossing or inbreeding to produce more breeding
material, falls broadly into two stages. The first stage is often done by the
team leader alone and is essentially based on a visual assessment of individ-
ual seedlings or small families to choose material for the next stage. The
second stage of selection is based on carefully planned replicated experi-

134 Plant Breeding

ments with clone plants or moderately-sized families to measure yield and

horticultural quality by objective assessments, if possible. This may take
several years with perennials. Before the first stage of selection, there
may be a preHminary elimination of individuals or whole families suscep-
tible to specified disease pathogens or carrying some undesirable trait
which can be recognised at an early stage. After the second stage the
breeder may decide to submit some of the material to further trials at
more than one centre and to rigorous commercial scrutiny before finally
deciding to enter it for independent trials in comparison with standard
cultivars and probably also with comparable new material from com-
petitors (see Section 9.3}.


Before starting on the second stage of breeders' selection, a 'selection
index' of desirable plant characters should be drawn up and each plant
or family is then assessed numerically for these features. For cultivars
grown as commercial crops these will probably include yield, relative time
of maturity, uniformity (in the case of seed propagated crops}, plant
habit (especially in relation to ease of harvesting}, quality of marketable
product, disease and pest resistance, cold, heat or drought tolerance and
in some cases tolerance to pesticides and herbicides, and suitability for
A figure for yield is obtained by weighing and counting the produce,
but it may not always be meaningful. For those crops with a clearly
defined end point such as peas grown for consumption of the dry seed,
number of flowers per bulb, fresh weight of fruits like tomato, strawberry
and plum which change colour when they ripen, the yield figures will be
fairly meaningful for the particular trial. With crops that have a less well
defined end point, such as peas grown for canning or freezing, yield is defin-
ed as the weight of crop at a certain stage of maturity, usually measured
in the case of these crops by an instrument called a tenderometer (Section
9.2.1 ). If the crop is harvested two or three days too early or too late a
false view of the relative yield will be obtained. Even when an accurate
assessment of yield in a particular trial has been obtained it may not give
a very clear indication .of yield potential. Recent work by the National
Vegetable Research Station, England, for example, has drawn attention to
the considerable variation in relative yields of carrot cultivars in different
sites and seasons. The breeder can do no more than put forward those
selections which have yielded well in carefully planned trials in relation to
known standards and trust that more extensive trialling will support the
When it is impossible, or too time-consuming, to assess a character
objectively, then a subjective assessment on a numerical basis from 1 to 5
or 1 to 10 should be used. If several observers make i ndpendent assess-
ments, the figures can be analysed statistically. It is usually most con-
Selection, Introduction and Maintenance of New Cultivars 135

venient to arrange for the higher scoring for the most desirable aspect of
the recorded feature. Flavour is often recorded in this way and to obtain
meaningful results the participants should be tested first to find whether
they have a normal taste range. Some individuals are 'blind' to certain
tastes. Also, tasting tests should be carried out in dim, coloured lighting
as assessors of taste are easily prejudiced by visual appearance. It should
be made clear to the participants whether they are judging for personal
preference or some specific feature such as sourness or bitterness. Many
other characters including colour, plant habit and degree of disease resist-
ance can be assessed subjectively on a numerical basis.

9.2.1 Objective assessments, instrumentation

Whenever possible objective assessments are made of individual plants,
fruits and flowers by measuring sizes, fresh or dry matter weights, and of
sugar, acid or vitamin content by chemical tests. Instruments have been
designed specifically to measure certain features, especially the firmness or
consistency of the flesh of fruits and vegetables, and fruit skin toughness.
These work on two principles: (1) the degree of distortion when a pre-
determined force is applied to the product, and (2) the force required to
cause a probe to penetrate skin or flesh. The first method has been used to
assess the firmness of onions (Ang et a/., 1960) and Brussels sprouts
(North and Frith, 1959). Various laboratory-made and commercial
instruments, including the L.E.E. Kramer shear press and the highly
sophisticated lnstron machine, have been used to measure the flesh firm-
ness and skin toughness of apple (Yoisey and MacDonald, 1964), straw-
berry (Ourecky and Bourne, 1968), and tomato (Hutchings eta!., 1962).
The tenderometer is a shear press type of instrument used routinely to
measure the consistency and thus the degree of maturity of green pea seed.
Skin cracking can be a problem with tomatoes grown in the open, and
a relatively simple instrument, designed to measure the force required to
push a probe through the fruit skin, enabled two workers to measure skin
strength of 200 fruits per hour (Yoisey and Lyall, 1965). The assessments
were closely correlated with skin cracking in the field. These instruments
and techniques could probably be used to measure skin and flesh texture
on a wide range of other fruits and vegetables. Similar instruments have
been used to measure ease of harvesting by recording the force required to
sever fruit from the stalk, as with cucumber (Hutchings eta/., 1962) and
Instruments such as the tintometer are used for colour assessments
but the measurements are often partly subjective since they depend on
the operator matching the colour of the plant product, or of a liquid
extract from it, against standard colour samples. Two independent opera-
tors will not always agree on the precise matching, and a similar problem
also arises when a colour chart such as tha.t published by the Royal Horti-
cultural Society, London, England, is used. Nevertheless, these aids are a
136 Plant Breeding

considerable help as colour is difficult to describe verbally and personal

assessments without comparative standards can be unintelligible to other
observers. Truly objective colour measuring instruments, such as the colori-
meter, measure transmission or reflection of light on three different
wavebands and it is sometimes difficult for the breeder to interpret the
figures in a meaningful way. However, readings from these instruments
can measure, for example, the amount of blue as compared with red or
yellow light reflected from a fruit surface, and the overall density of
colour. Raspberry and strawberry fruits which are bright red are preferred
to those which have a purplish tinge, and black currants which produce
a very dark coloured juice are usually considered the most suitable for
juice extraction for processing.
Instruments may also be used to subject material to a standard treat-
ment before making a subjective assessment. For example, strawberry
cultivars differ in the extent 'to which their fruits are damaged during
transport and their shelf life is reduced. An instrument to inflict a standard
shaking regime on fruits (Gooding, 1976) can help when assessing cultivars
for resistance to damage in transit.


Once the breeder's selection is completed, the potential new cultivars are
usually submitted, when possible, for comparison with standard and other
new cultivars in independent trials. It may be at least two years before a
recommendation can be made, but preliminary recommendations are
sometimes available within a year.
Breeders' trials of vegetables in Britain are carried out by the National
Institute of Agricultural Botany (NIAB), Cambridge, although the trials
themselves are grown at several centres. These trials at present are designed
with few replicates of each batch of material submitted and their main
value to the breeder is the comparison with other potential new cultivars,
which would not otherwise be available, rather than a detailed breakdown
of performance. The material is usually submitted under a breeder's code
number and performance is not publicised except to the participants.
NlAB also carry out more comprehensive trials of vegetable cultivars,
including potatoes, and publicise details of performance, listing recom-
mended cultivars, but only those already in commerce are accepted for
comparison. Trials of vegetables used for processing in the UK, notably
peas, beans, Brussels sprout, calabrese, carrot and sweet corn are organised
by the Processor and Grower's Research Organisation (PGRO) centred at
Yaxley, near Peterborough. These trials, and some of those by NIAB,
depend on the co-operation of the Campden Food Preservation Research
Association for assessments of processing quality. Small scale trials of
some vegetable types have also been carried out from time to time by the
Royal Horticultural Society (RHS).
Selection, Introduction and Maintenance of New Cultivars 137

Trials of new cultivars of soft and tree fruits are organised in Britain
by the National Fruit Trials Committee (N FT) of the Ministry of Agricul-
ture, Fisheries and Food, mainly on the trial grounds at Brogdale in Kent,
but also at several other centres representative of British fruit production
areas, including Scotland. These trials also include foreign cultivars and
they are an important aid to British plant breeders in deciding which
material to release as new cultivars. In general, there are two stages of NFT
trials: preliminary trials conducted mainly at Brogdale in which a decision
is made to recommend release, further trial or discarding: and second-
stage trials conducted at several representative centres in which a decision
is made on those cultivars designated from the preliminary trials as requir-
ing further trial.
Comparative cultivar trials of a wide range of ornamentals are held
from time to time at the RHS gardens at Wisley, Ripley, Surrey, and certifi-
cates awarded to the most promising kinds, notably the RHS Award of
It is not obligatory for a breeder to submit material for the trial
system described above, and a charge may be made for submission to some,
as with the NIAB Breeders' trials.


Cultivars of the majority of vegetable types cannot be offered for sale

legally in Britain unless they have been accepted on the National List. New
cultivars must be submitted for trial by the Plant Variety Rights Office
(PVRO) and are required to pass tests for distinctness, uniformity and
stability before they can be accepted on the List. The applicant for a new
cultivar has to agree to provide nuclear seeds or to nominate an agent to
do so when called on. The purpose of the Official List is to protect con-
sumers and breeders from incorrectly named seed stocks and from a
bewildering assortment of synonyms. Similar systems are in operation in
other European countries, including Denmark, France, Holland and West
Germany. The EEC publishes a 'Common catalogue of Varieties of Vege-
table Species' and, as far as possible, nomenclature is the same for all
member countries, though some synonyms are permitted, especially to
overcome linguistic difficulties. A charge is made for entrance to the
trials, which usually take two years to complete. Similar regulations exist
for cultivars of most agricultural crops and for many of these species,
including potatoes, the trials also assess cultivar performance. Cultivars
which have poor performance, such as extreme susceptibility to disease
pathogens or very low yield, are not acceptable for inclusion in the List,
even though they may be distinct, uniform and stable. The extension of
such statutory performance trials to some vegetable species is under
discussion. Although the National List protects consumers, it puts diffi-
culties in the way of the amateur wishing to obtain unusual cultivars to
138 Plant Breeding

gratify his whims. Societies which distribute seeds to members without

exposing them for sale to the public could overcome this difficulty.
There is no National List of cultivars of fruits or ornamentals in
Britain except for the salad fruits such as cucumber, melon and tomato.
New cultivars of some horticultural crops can be covered by Plant
Breeders' Rights whereby the breeder or his or her nominated agent is
entitled to a levy on sales of the material, much as the patent scheme
covers inventions. The United Kingdom schemes at present include culti-
vars of apple, beans (French and runner), black currant, carnation, cherry
rootstocks, chrysanthemum, conifers, Cymbidium, dahlia, delphinium,
fuchsia, herbaceous perennials, Li/ium, lettuce, Pelargonium, pea, pear,
plum, potato, raspberry, Rhododendron, rhubarb, rose, strawberry, swede
turnip, tomato, ornamental trees, shrubs and woody climbers. Before new
cultivars are covered by 'Rights' they must be submitted for trial, and with
eligible vegetable types this is usually .done at the same time as they are
entered in trials for the Official List. A cultivar is not accepted if it has
previously been offered for sale and it must fulfil requirements of distinct-
ness, uniformity and stability. Once a cultivar is entered for Rights it may
be covered by a Protective Direction until confirmation is received of full
cover by Rights, in much the same way as application for patent rights is
covered by 'Patents pending'. Applications for entry of material for Rights
in the United Kingdom and requests for additional types of plants to be
covered by Rights are made to the Plant Variety Rights Office at Cambridge.
Plant Variety Rights are available in some other European countries,
notably Holland and West Germany. The United States does not operate
a Rights Scheme but cultivars can be patented there in much the same way
as inventions, and in France a scheme similar to a Trade Mark is available
for new cultivars.


It is inadvisable to publicise the name of a new cultivar until shortly before

its release, because confusion can rise if an objection is raised and the
name disallowed. This is especially important if application is made for
Rights or for inclusion in the National List, when a code of rules applies
to the naming of cultivars. The following are unacceptable.
(1) Names of more than three words.
(2) Figures and letters.
(3) Latinised names.
(4) Names already used for a cultivar of the same genus or names
closely resembling them.
(5) Inclusion of such words as 'variety', 'form', 'hybrid', 'cross' in the
(6) A name indicating that the cultivar concerned is related or derived
from another cultivar when this is not the case.
Selection, Introduction and Maintenance of New Cultivars 139

(7) The definite article or some titles such as "The", 'Mr', 'Miss', or an
abbreviation such as Mt for Mountain.
(8) Excessively long words or those difficult to pronounce.
(9) Names implying a trade mark such as the continued use of
'Bolgsville' as a prefix or suffix for a series of cultivars, though
such an addition to names could be used if it is registered as a trade
Proposed names have to be exhibited in the Plant Varieties and Seeds
Gazette of the PG RO and objections can be raised before they become
accepted. Only one name can be submitted at a time for each cultivar and,
if this is unacceptable, another has to be submitted for approval.
These rules for nomenclature are not obligatory for fruits and orna-
mentals when no application is made for Rights, but it is advisable to keep
as near to them as possible. Some societies such as the Royal Horticultural
Society maintain a register of cultivar names which can be helpful to a
breeder trying to choose one not previously used for the type of plant in
question, and the breeder has some moral obligation to make use of such
lists when applicable.
The decision to release a new cultivar is the responsibility of the
breeder or the breeder's agent; the official variety trials at present merely
help the breeder to make a decision and are not binding. However, failure
to meet the requirements of distinctness, uniformity and stability may
prevent a vegetable cultivar from being accepted on the National List and
thus preclude its commercialisation. This does not apply to fruits and
ornamentals, though Rights may be refused.


Many stocks of vegetatively propagated plants have become virus infected

and it can be difficult to find a source of some cultivars that is free from
known viruses or mycoplasmas. It is therefore in the breeder's interest to
ensure that some scheme is available whereby stocks which are free from
the most debilitating viruses are fed into the propagation system from
time to time. Badly infected stocks can destroy the reputation of an
otherwise excellent cultivar. Furthermore, stocks may become infected
during the early stages of trials because it is uneconomic or even undesir-
able to prevent infection. The breeder then will wish to 'clean up' the
stock of a new cultivar before it is released, and techniques for doing this
become part of the breeder's routine work.
Treatments with chemicals, including Malachite Green and Quin-
hydrone, have had a limited success in freeing material from virus under
experimental conditions but are not used as routine treatment. The most
useful techniques are heat treatment and meristem tip culture. Heat
treatments in air, usually at 35-38°C for 3-8 weeks, have successfully
freed some hosts from some viruses, and are used as a routine for rasp-
140 Plant Breeding

berry. The precise treatment is critical and advice from a plant virologist
should be sought before starting. During the last few years, in vitro meri-
stem tip culture has been very successful in freeing crops from some viruses
including strawberry, many bulbous species, carnation, chrysanthemum,
dahlia, rhubarb and potato. A portion of the meristem tip and first leaf
primordia, or an even larger cutting with certain genera such as Dahlia, is
transferred under sterile conditions to a bottle containing a nutrient and
sugar mixture, sometimes with auxins. As a general rule growth, and
especially root growth, is best when a liquid medium is used and the tissue
supported on a filter paper 'table' or 'bridge', although solid media contain-
ing agar are sometimes used (Hollings, 1965). Not all the plants produced
in this way are virus-free and virus testing is sometimes the most expensive
part of the operation. In Britain, the Nuclear Stock (Ornamentals) Organ-
isation and some biochemical companies provide a service to free plant
stocks from viruses, for which a charge is made. This can be helpful to a
breeder who does not have the facilities, time or expertise to devote to
this work.

9.6.1 Certification schemes

Independent crop certification schemes are an important aid to the main-
tenance of healthy true-to-type stocks of some vegetatively propagated
cultivars. The Ministry of Agriculture, Fisheries and Food operates schemes
in England and Wales for seed potatoes, black currant, raspberry, straw-
berry, fruit trees (including finished trees, mother trees and rootstocks)
and carnation. The Department of Agriculture and Fisheries for Scotland
has schemes for seed potatoes, black currant, raspberry, strawberry and
flower bulbs (at present narcissus and tulip).


The rate of vegetative propagation by normal horticultural techniques such

as division, cuttings, layers, budding and grafting may be slow for some
species and more rapid propagation could be helpful to the breeder wish-
ing to introduce a new cultivar or a disease-free stock of an existing one.
It has been known for many years that manipulative treatments of some
ornamental bulbous species can result in a greater rate of multiplication,
as in the use of isolated bulb-scales of lilies or the cutting or scooping out
of the baseplate of hyacinth bulbs. During the last few years it has been
found that similar treatments can aid in the propagation of many other
bulbous plants. Narcissus bulbs can be cut transversely into small pieces,
each with portions of two scales and a part of the baseplate (Eaton, 1973).
This technique described as 'twin-scaling', will give some 50-100 new
flowering bulbs from an individual specimen within three years, during
which time it would probably have developed no more than four or five
offset bulbs by natural division.
Selection, Introduction and Maintenance of New Cultivars 141

In vitro techniques, in which plant tissues are grown in glass containers

under sterile conditions on media containing sugar, nutrients and some-
times other chemical aids to growth, have excited much interest in recent
years, and their use for some plants is becoming part of the breeder's
routine. When meristem tips of orchids are grown in a manner used to
free the plants from virus, the tips themselves often proliferate in culture.
The new meristems can be transferred to further bottles of nutrient and
the operation repeated to produce a large number of clone plants in a
relatively short time. This technique has made it possible to commercial-
ise new clone cultivars of orchids which would otherwise have taken years
to propagate, and it has revolutionised the commercial orchid industry.
Meristem tips of most other plants do not proliferate in the same way
as orchids because of the special dominance resulting from auxin produc-
tion, although useful results have been obtained with bulbous species such
as gladiolus and hyacinth. By culturing meristems on a medium low in
auxin and rich in cytokinin, such as 6-benzylaminopurine, apical domin-
ance may be overcome and the tips then proliferate as with orchids. This
technique has been successful with strawberry (Boxus, 1974) and Gerbera,
and many thousands of new plants have been obtained from one individual
within a year. Recently it has been found that meristem tips of apple,
plum and cherry stocks can be induced to proliferate by the addition of
phloroglucinol to the medium and a 6,000-fold increase in apple obtained
in 8 months (jones eta!., 1977). It is probable that the addition of this
chemical to the medium may be helpful in the propagation of a wider
range of woody plants.
These new techniques, many of which are described by Murashige
(1974), have their problems and are at present applicable only to a limited
range of species. It may be necessary to transfer tissue to different media
to obtain autonomous plants; tissue of bulbous species may become
dormant and require a chilling treatment to induce active growth; there
may be serious losses of material after transference from in vitro culture
to soil; mutants may arise more frequently than when conventional
propagation techniques are used and the techniques are unsuitable for
chima:ras. In any case the new plantlets usually require growing on with
care under glass before they can be transferred to the open. Nevertheless,
these rapid in vitro propagation techniques offer exciting new prospects
for the breeder of vegetatively propagated cultivars and will undoubtedly
be used on a large scale in future. They may have a place also in the
breeding of seed propagated cultivars, as in the clonal propagation of
parents for F 1 hybrids.


We have seen that the modern trend is towards more uniform cultivars of
seed propagated plants and that Official Lists, and the publicising of
142 Plant Breeding

Recommended Varieties, are tending to restrict the number of cultivars

in commerce. Both these measures lead to the discarding of old cultivars
and land races. Much of this older material carried genes, gene alleles and
gene combinations which had originated or been maintained under the
protection of cultivation during periods of thousands of years, and they
may now be lost. This so-called gene erosion means the destruction of
much useful genetic material available to breeders. A pathogen may
mutate to a new virulent form for which no resistance is known amongst
current commercial cultivars, and a useful source to look for new resist-
ance may exist amongst old ones.
It is generally agreed amongst breeders that an attempt must be made
to maintain old cultivars, land races and also species threatened with
extinction. The work involved in this conservation may be costly and may
seem to be routine and insufficiently attractive to scientists wanting to
develop a career. The problem is to find organisations and individuals
willing to finance and carry out such work.
Several cultivar collections and seed banks do exist already for a
limited range of crop plants and it is to be hoped that this aspect of
conservation will be given its due prominence in future planning.

ANG, J. K., ISENBERG, F. M. and HARTMAN, J.D. (1960). Measurement of firm·
ness in onion bulbs with a shear press and a potentiometric recorder, Proc. Am.
Soc. hort. Sci., 15, 500-509
BOXUS, P. (1974}. The production of strawberry plants by in vitro micro·propaga·
tion, ]. hort. Sci., 49, 209-210
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Abutilon 18 Apricot 22, 94
Accessory chromosome 30 Aquilegia 4, 34, 38, 128
Aceto-orcein stain 27 Arabis mosaic virus 89
A chimenes 80 Artichoke 22
Additive gene 68 Ascorbic acid 61
Adenine sulphate 61 Asclepias 44
Aesculus 34 Asparagus 22, 29, 34, 53, 75, 116
Agamospermy 62 Atriplex 34
Ageratum 53,110,128 Autopolyploid 21
Albino 69 Azalea 102, 103
Allele 4, 15
Allium 38, 62, 81, 123
Allopolyploid 21
Allotetraploid 21 B chromosomes 30
Almond 22, 94 Back crossing 15, 16, 85, 108
Aloe 32 Beans 51, 53, 59, 107, 111, 112,
Alstromeria 80 136, 138
Amelanchier 95 Beet 22,33,37,38,41,42,51,53,
American gooseberry mildew 86, 117,131
87 Begonia 82, 128, 131
Amphidiploid 24, 126 Bellis 53, 110, 128
Amphorophora rubi 89 Big bud mite 86
Anaphase 7, 8, 25 Blackberry 22, 64, 84, 85
Androecium 33 Black currant 22, 51, 83, 84, 85, 86,
Anemone 82 89, 136, 138, 140
Aneuploid 21, 30, I 00 gall mite 86
Angiosperm 44, 46 Blow flies as pollinators 38, 123
Aniline blue 49 Blueberry 22, 51, 84, 87
Antennaria 63 Bolting 5, 118, 120
Anther culture 29 Brassica 22, 35, 49, 58, 118, 126,
Antholyza 32 127
Anthracnose of beans 113 Brassica oleracea 13, 14, 15, 39, 42,
Antipodal cells 46 57, 58, 118, 119, 131
Antirrhinum 38, 40, 53, 57, 81, incompatibility system 57
128, 131 'Breaking' of flowers 18
Apomixis 17, 62, 63, 84, 96,104 Bremia lactucae 114
Apple 2, 22, 42, 45, 5 I, 52, 54, 92, Broad bean 22, 53,111,112,131
93, 135, 138 Brown anther 39, 120
canker 93 Brussels sprouts 5, 13, 22, 53, 110,
mildew 93 135, 136
rootstocks 63, 93, 141 Bud pollination 58, 119
scab 93 Bumble bees as pollinators 32, 38


Cabbage 12, 13, 18, 22, 51,100, Cistron 19

118,119 Citrus 63
Calabrese 53, 118, 119 Cladosporum .fulvum 128
Calceolaria 128 Clarkia 30
Calendula 53, 131 Cleistogamous flowers 33
Callose 49 Clematis 40
Caltha 34 Cloned cultivars 82
Camerarius, Rudolph Jacob 3 Clubroot 119,120,127
Campanula 34, 131 Cobalt-60 79
Capsicum 38 Coconut 51
Carnation 3, 98, 138, 140 Coconut milk 61
Carrot 32, 34, 38, 39, 42, 51, 53, Colchicine 24, 25, 86
110,120,121,1 31,134 Colchicum 24
Cauliflower 13, 18, 22, 53, 111, 118, Coleus 19
119 Colour chart 13 5
mosaic 118 Colorimeter 136
Celeriac 121 Combining ability 74, 75
Celery 22, 51, 53, 121 Commercial seed 129, 130
Cellophane bags 35 Common Catalogue of Varieties
Centaurea 34, 53, 131 137
Centromere 6 Complementary gene 5, 68
'Cepa' 40 Complex segregations 72, 74
Ceratostigma 46 Cosmos 57
Cercidiophyopsis ribis 86 Cranberry 87
Certation 69 Crategomespilus 19
Certification schemes 140 Crataegus 95
Chemical mutagens 79 Crepis 26
Chequerboard diagram 13, 66, 67, Cross pollinators 108, 116
73 Crossing over 10, 11, 63
Cherry 22, 57, 93,94 Crown gall 94
Chervil 22 Cucumber 22, 29, 34, 40, 52, 53,
Chi squared test (statistics of gene 109, 121, 131, 135
segregation) 70, 71 Cucumis 40, 121, 122
Chiasma 11 Cucurbita 40, 45, 122
Chicory 22, 40, 42, 53 Cultivar 1
Chimaera 18, 19, 79, 80, 85, 92, Cyclamen 61, 82, 128
102 Cydonia 95
Chinese cabbage 127 Cytoplasmic inheritance 17, 48
Chives 124- Cytoplasmic male sterility 38, 39,
Chlorophyll-deficient seedlings 69 123
Chloroplasts 18
Chromatids 6, 10
Chromosome 5-13, 21, 22, 23, 27, Daffodil 21 , 102
28,30, 80 Dahlia 3, 21, 40, 53, 77, 80, 83, 99,
counting 27 138, 140
doubling 24, 26 Damson 96
map 10, 11 Dandelion 63
numbers 22 Darwin, Charles 3
Chrysanthemum 21, 32, 40, 53, 80, Datura 69
83, 92, 128, 131 Decaploid 21, 100

Degrees of freedom 70 Fertility I 07

Deletion mutants 77, 78 F 1 cultivars I 09, 110, 111, 120,
Delphinium 34, 53, 82, 99, 138 121,122,123,127,128
Deoxyribonucleic acid (DNA) 19 F 1 generation 15
Diallel cross 7 5, 90 F 2 generation 15, 66
Dianthus 3, 53, 98,128,131 Field bean 39, 112
Dihybrid segregation 15, 66 Fig 22, 33, 52
Dimorphic flowers 55, 56 Fire blight 95
Dioecious 34, 75 Fixative for root tip squashes 27
Dioecy 110 Flower colour mutants 80
Diploid 21, 28, 30,63 Flowering, control 41, 97
Dithiothreitol 80 Food Preservation Research
Dodecaploid 21 Association 136
Dominant genes 4 Fragaria 52, 75, 89, 90
Dormancy of seeds 54 Freesia I 00
Double cross cultivar Ill French bean 17, 22, 34, I 07, 112,
Double fertilisation 48 113,138
Double flowers 40,41 French bean X runner bean hybrid
Double stock 40 57' 113
Dryas 34 Fritil/aria 46, 51
Duplicate gene 63 Fruit, abcission 51, 85
Dutch Seed Association 130, 131 development 51
Dwarfing rootstocks 42 set 51
Fuchsia 32, 100, 138
Fuchs, Leonard I
Egg cel146 Functional male sterility 3 9, I 27,
Emasculation 3 5, 91, 9 5 129
Embryo culture 59, 60, 61, 94, 96
Embryo sac 46, 58, 61
Endive 22, 53 Garlic 22, I 24
Endosperm 46, 48, 49,51 Gamete 8, 44, 4 7
Epicoty1 dormancy 54 Gametophytic incompatibility 56,
Epistatic gene 5 57
dominant gene 68 Gene 4
recessive gene 68 chemical structure 19
Erigeron 63 erosion 142
Eriosoma lanigerum 93 pools 141
Erisyplze polygoni 120 General combining ability 75
Erwinia amylovora 95 Generative nuclei 44
Erytlzronium 46 Genetic marker 3 7
'Ethrel' 40 Genotype 5
Ethyl methane sulphonate 79 Geranium-see Pelargonium
Gibberellic acid 42, 114
Gladiolus 21, 100
Fairchild, Thomas 3 Glasshouses 41
False variegation 19 Gloxinia 128
False fruits 51 Goodness of fit (statistics of gene
Fennel 22, 39 segregation) 69
Fertilisation 47, 48,58 Gooseberry 22, 86, 87, 88
in vitro 58 Gossypium 69

Graft hybrid 19 Jerusalem artichoke 96

Gynaecium 33 June yellows 18
Gymnosperms 44

Kalanchoe 80
Halo blight 113 Kales 4, 13, 29, 53, 118
Hand pollination 34, 121, 127 Kinetin 61
Hamamelis 48 Kniphophia 32
Haploid 21, 28, 29, 117 Kohlrabi 13, 53, 118, 119
Hastening flowering 42, 83 Kolreuter, Joseph Gottleib 3, 74
Hawkweed 4
Hawthorn 19, 95
Hazel48 Laburnum 19
Helianthus 38 Lactuca 81, 114
Helleborus 34 Lamium 19, 33
Heptaploid 21 Lathyrus 53, 58
Heterogeneity 71 Laurel 30
Heterosis 74 Leafless pea 116
Heterozygosity 13 Leek 21, 22, 30, 38, 53, 62, 107,
Hexaploid 21, 90, 95, 99, 122 122
Hibiscus 32, 128 Leek rust 122
Hieracium 4 Lethal gene 68, 77
Hive bees as pollinators 38 Lettuce 22, 34, 35, 36, 39, 42, 107,
Homozygosity 13, 108 108,114,138
Homozygous 13, 15 mosaic 114
Horse chestnut 51, 61 pollination 35, 36
Hosta 18, 62,63 Ligustrum 18
House flies as pollinators 37 Lilium 24,35,41,49,54, 58,59,
Hybrid cultivars 17, 74, 109 63, 82, 101, 104, 105, 138
Hybrid seed crop 111 Lily see Lilium
Hybrid vigour 74, 109 Line breeding 108
Hyacinth 21, 100, 140 Linkage 11, 17, 66, 72
Hydrangea 18, 34 Linkage group 11, 13
Hypostatic gene 5 Locus 19
Loganberry 85
Long daylength treatment 42
Jberis 53, 57 Lonicera 52
Impatiens 33, 52, 128 Lupin-see Lupin us
Inbreeding depression 74, 108, 120 Lupinus 51, 53,82
Incompatibility 23, 54, 55, 56, 58, Lycopersicum 127, 128
techniques for overcoming 58
Incomplete dominance 4 Magnolia 34
Inositol 61 Maintenance of cultivars 133
Insect-proof containers 41 Male sterile 38, 72, 123
Inter-specific cross 3 9 Male sterility 17, 38, 39, 110, 120,
incompatibility 57 123
Introduction of new cultivars 133 Malus 22, 63, 92, 93
Inversion mutants 77, 78 Marigold 5 I
Iris 54, 59, 100, 101 Marrow 22, 34, 52, 53, 110, 122
Isolation distances 130, 131 Mass selection I 09, I 20

Matthiola 40, 53, 131 Octaploid 21, 22, 90, l 00

Mechanical harvesting 84, 86, 91, Ocnothera 69
92,111,113 Onion 22, 38, 53, 62, 110, 123,
Medlar 19 124, 131
Meiosis 6, 8, 9, ll, 23, 28 Ophrys 32
Melon 22, 34, 53, 122 Opuntia 62
Mendel, Gregor Johann 3 Oxalis 33
Mendelism 3
Meristem tip culture I 0 l, 139, 140
Metaphase 6-9 Paeony 54, 58, 102
Mimosa 44 Paeonia -see Paeony
Mitosis 6, 7,121 Pansy ll 0
Monoecious 34, I 21 Papaver 53, 58, 128
Monoecy 110 Papillae of stigma 4 7
Monogerm beet 117 Para-dichlorobenzene 2 7
Monohybrid segregation 13, 14, 66 Parnassia 32
Monosomes 30 Parsley 22, 53
Mulberry 21, 2 2 Parsnip 22, 32, 53, 124
Multiple alleles 73 Parthenocarpic fruit 52, 55, 95
Mutation 77, 79, 80,81 Pea3,4,23,42, 107,114,115,
artificial induction 79, 99 116, 134, 135, 138
of ageing seed 80, 81 Peach 22, 59, 94, 95
Myrobalan 96 leaf curl 94
canker 94
Pear 21 , 2 2, 55, 9 5, 13 8
Naming of cultivars 138 rootstocks 9 5
Narcissus 21, 25, 101, 102, 140 Pear-apple hybrid 52, 57
National Fruit Trials 13 7 Pedigree breeding l 08
National List 13 7 Pelargonium 18, 82, 102, 1 28, 138
Natural pollination 32 Penstemon 33, 53
Nectarine 22, 94 Pentaploid 21, 98
Nectria galligena 93 Pepper 23
Negative mass selection 109 Perianth 33
Nematus ribesii 87 Periclinal chimaera 18
Nemesia 131 Petunia 38, 53, 58, 62, 110, 128,
New Zealand spinach 125 131
Nicotiana 26, 32, 62, 69, 128 Phaseolus 17, 34, 112, 113, 116
Nicotinic acid 61 Phenotype 5
Nitrous oxide 26, 27 Phythopthora fragariae 90
Non-endospermic seeds 51 infestans 98, 128
Nucellus 46 Pistil 33
Nuclear seed 129 Plant Breeders' Rights Ill, 138
Nuclear Stock (Ornamentals) Plant Variety Rights Office 137
Organisation 140 P/asmodiophora brassicae ll 9
Nucleotide 19 Plastid 18, 19
Nullisome 30 Platanthera 32
Pleiotropic 4, 89
Plum 21, 22,45, 95, 96,138
Oak 48 Podosphaera /eucotricha 93
Objective assessment of plant Polar nuclei 45, 46, 48, 63
character 134, 135 Pole bean 112

Pollen 3 5, 3 7, 4 7, 48 Recessive genes 4

collection 37, 45 Reciprocal cross 1 7
germination tests 45,49 Red core disease 90, 91
morphology 45 Red currant 22, 89
mother cells 44 Reduction division 6
storage 45, 86, 89, 96, 97,99 Rhododendron 3, 44, 83, 102, 103,
tube 45, 47, 48,49 138
viability test 45 Rhubarb 23
Pollen-pistil incompatibility 56 Ribes 52, 85, 86, 88, 89
Pollinia 44 Root tip squashes 27
Polygenic 5, 73 Rose (including Rosa) 3, 17, 21, 47,
inheritance 73, 94 63, 103, 138
Polyploid 21, 24, 30, 45, 73, 80 Rowan 95
Polyploidy 30 Royal Horticultural Society 136
Portugese cabbage 13 Rubus52,62,84,85,88,89
Post-fertilisation failures 55 Runner bean 17, 23, 34, 82, 116,
Potato 23, 29, 83, 97, 98, 136, 138, 138
Potentilla 63, 90
S-alleles 56, 57, 118, 119
Primula 4, 21, 55, 56, 82, 128
Saintpaulia 80, 128
Processors and Growers Research
Salad crops 111
Organisation 136
Salpiglossis 53, 128
Pro-embryo 49, 60
Salsify 53
Progeny size 72
Sea kale 23, 83
Prophase 6-8
Second division of meiosis 11
Protandrous 34, 35, 37
Sectoral chimaera 80
Protogynous 34
Seed development 51
Protoplast fusion 61, 6 2
dormancy 54, 61
Prunus 22, 30, 93, 94, 95, 96
extraction 52, 86, 132
Pseudogamy 63
Pseudomonas syringae 96 production 129, 130
propagated cultivars 107
Pseudopeziza ribis 86
storage 53
Pumpkin 112
Pyrus 95 Segregation 13, 14, 15, 66, 67,68
forecasting 66
ratios 15, 68
Quercus 48 Selection I 09, 13 3
Quince 95 index 134
Selective gametocide 39, 40, 114
Radish 22, 39, 53, 57, 110,124, 131 Self fertilisation 55, 107
Radish X cabbage hybrid 58 Self incompatibility 55, 92, 94, 96,
Ranunculus 51 110
Raphanus 38 Self pollination 39, 107, 108, 111
Raphanobrassica 17, 118 Sex-linked character 75
Rapid propagation techniques 140 Sex ratio 75
Raspberry 2, 22, 23, 54, 55, 77, 83, Shear press 13 5
88, 89, 136, 138, 139, 140 Sibs 119
aphid 89 Silver nitrate 121
beetle 89 Sloe 96
black 88 Snap-pod bean 112
ringspot virus 89 Sodium dichloroacetate 40

Sodium 2, 3-dichloroisobutyrate 40 Three-way cross 111

Solanum 97 (see also potato) Threshing seed crops 130
Somatic cell 6 Thymus 34
Sorbus 95 Tin to meter 13 5
Spawn bed 89 Tobacco black ring virus 89
Specific combining ability 75 Tomato 23, 39, 52, 53, 57, 109,
Spermatophyta 44 110, 127, 128, 134, 135, 138
Sphaerotheca mors-uvae 86 seed extraction 52
Spinach 23, 33, 34,41, 42, 51, 53, Transformation 19
7 5, II 0, 12 5, 126, 131 Translocation mutants 77, 78
Spinach beet 117 Trials 136, 137
Spindle 6 Trihybrid 72
Spiraea 18 Triploid 21, 22, 27, 92, 100, 101,
Sporophytic incompatibility 23, 56, 102
57 Tripping of bean flower 11 2
Sports (mutant branches) 77 Trisome 30
Squashes 23, 122 Tritonia 100
Staminode 33 True breeding 13, 108
Steckling 130 Tulip 18, 21, 26, 27, 47, 83, 103,
Stigma exudate 47, 58 104
Stock seed 129 Turnip 22, 42, 51, 53, 110, 127,
Stomata of polyploids 26 131
Strawberry 21, 22, 34, 51, 75, 83, mosaic virus 118
89,90,91,138 Twin-scaling Narcissus bulbs 140
Strelitzia 3 2
Streptocarpus 80 Umbelliferae 32, 37
String bean 112
Stump seeding 130 Vaccinium 22, 52, 87
Style 48 Variegation 18, 19
Sugar peas 115, 116 Vegetatively propagated cultivars
Super female spinach 125 82,83
Super male asparagus 117 Vegetative nucleus 44
Suppressor gene 68 Venturia inequalis 93
Swede 1, 23, 53, 107, 126, 127, 131 Verifying hybridity 104
Sweet corn 53,110 Vernalisation 42
Sweet pea 58 Verticillium albo-atrum 91
Swiss chard 117 Verticillium wilt 91
Symphoricarpus 32 Viburnum 34
Synergids 46,47 Vi cia Ill, 112
Synthetic varieties 111 Vinca 18
Viola 33, 52, 53, 82
Violet 33
Tagetes 45, 53, 107, 110, 128,131 Virus-free stocks 79, 93, 139
Tare-leaved rogue of peas 48
Tasting tests 135
Telophase 7, 8, II Wallflower 18
Tenderometer 13 5 Watercress 23
Tepal33 Watermelon 23
Tetraploid 21, 22, 23, 26, 102,122 Wax-pod bean 23, 122
Tetrad 8, 44, 46 White currant 89
Tetrazolium 45 Woolly aphid 93

X rays 79, 80, 86

Yeast extract 61
Yucca 32

Zea 69
Zinnia 53, 128, 129
Zygote 48