Professional Documents
Culture Documents
IN
HORTICULTURE
Science in Horticulture Series
General Editor: Professor L. Broadbent, University of Bath
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.
PLANT BREEDING
AND GENETICS
IN HORTICULTURE
C. North
M
© C. North 1979
Set I.B.M. by
REPRODUCTION DRAWINGS LTD, SUTTON, SURREY
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
Preface ix
THE MECHANISM OF INHERITANCE
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 CHROMOSOME NUMBER 21
2.1 Polyploids
2.2 Techniques for inducing polyploids
2.3 Chromosome counting
2.4 Haploids
2.5 Aneuploids
2.6 Accessory chromosomes
3 FLOWER FORM AND POLLINATION 32
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
Index 143
PREFACE
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
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.
1.3 MENDELISM
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
1.4 GENES
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-
ment.
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
differently.
1.5 CHROMOSOMES
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
1.6 MITOSIS
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
2
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
nucleus.
1.7 MEIOSIS
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.
2 3
' I
I I
\
~0 \
I
\
,I
J
J
J
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
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.
1.10 SEGREGATION
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
p X
1
Wh wh Fn fn
Fl
~
WhFn
// \~ Wh fn wh Fn wh fn
Wh
Fn
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.
1.12 VARIEGATION
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.
1.13 CHIMAERAS
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
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
inheritance.
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
units.
20 Plant Breeding
REFERENCES
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-
188
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,
361-372
FURTHER READING
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
CHROMOSOME NUMBER
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.
21
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.)
FRUITS
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
VEGETABLES
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
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
~
( A'\"1\
X~ I VJ . VI
"'\!/
\ W!}!)
'-___-) \______
\ METAPHASE ANAPHASE TELO PHASE
@,
I ./.
PROPHASE
I
,x~, ("~ (~
__)
COLCHICINE M ITOSI~
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,
1938.)
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.
2.4 HAPLOIDS
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
2.5 ANEUPLOIDS
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.
REFERENCES
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,
22,310-316
NITSCH, C. (1974}. La culture de pollen isole sur milieu synthetique C.r. hebd.,
Seanc. Acad. Sci., Paris, 278,1031-1034
Chromosome Number 31
FURTHER READING
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
FLOWER FORM AND POLLINATION
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
32
Flower Form and Pollination 33
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.
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
pansy.
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
s
'\
I
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.
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.
(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
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
space.
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.
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.
REFERENCES
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
FURTHER READING
FREE, ). B. (1970). Insect Pollination of Crops, Academic Press, London, 544 pp
HUDSON, ). P. (1957). Control of the Plant Environment, Butterworths, London,
240 pp
FERTILISATION AND
SEED DEVELOPMENT
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
44
Fertilisation and Seed Development 45
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
s
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.
4.4 FERTILISATION
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
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
phosphate.
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
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.
4.7 SEEDS
Table 4. 7
VEGETABLE SEEDS
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.
TREE SEEDS
{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}.
4.8 INCOMPATIBILITY
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
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
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.
4.10 EMBRYOCULTURE
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
Table 4.2
Some media used for embryo culture (mg per litre of water)*
4.12 APOMIXIS
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
REFERENCES
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,
1017
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.,
305
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,
643-646
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
FURTHER READING
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
SEGREGATION AND
COMBINING ABILITY
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 •
66
Segregation and Combining Ability 67
.t.B Ab oB ob
Figure S.l Chequerboard diagram for F 2 segregation from the cross AABB X aobb
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
5.4 CERTATION
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.
Table 5.1
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
Observed
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
x
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,
x
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.
Table 5.2
Number of plants to grow to obtain at least one
plant of the required genotype
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
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
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 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
REFERENCES
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
MUTATIONS
6.1 CHANGES IN CHROMOSOME AND GENE STRUCTURE
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
environment.
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
77
78 Plant Breeding
a a
b
D d~ b
c
c
e
d
a a a a
b b b b
I c c d
d c
e
e
a a a f
b b c
c
T g d
d
e
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
REFERENCES
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,
156-162
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.,
191,302-303
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,
153-207
FURTHER READING
HAGBERG, A. and .li.KERBERG, E. (1962). Mutations and Polyploidy in Plant
Breeding, Heinemann, London, 149 pp
VEGETATIVELY
PROPAGATED CULTIVARS
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.
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.
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
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
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
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
Britain.
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,
1962).
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
climate.
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).
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
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
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-
fertile.
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
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
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
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
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
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).
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.
REFERENCES
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.,
93,88-109
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.,
18,121-130
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
ROYAL HORTICULTURAL SOCIETY, RHODODENDRON GROUP (1964).
Rhododendron Handbook, Part 2, Rhododendron hybrids, Spottiswoode and
Ballantyne, London, 436 pp
ROWLEY, G. D. {1966). The experimental approach to rose breeding, Scient. Hart.,
18,131-135
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
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.
Table 8.1
Numbers ofF, cultivars of some veqetable crops in the UK only and in all EEC
countries {after Wills and North, 1978)
Table 8.2
System Crops
A Genetic/cytoplasmic
(i) Male sterility Carrot
Leek
Onion
Petunia
Radish
Tagetes
Tomato
(ii) Self-incompatibility Ageratum
(sporophytic or gametophytic) Bellis
Broccoli
Brussels sprout
Cabbage
(iii) Dioecy Spinach
(iv) Monoecy Sweetcorn
B Mechanical or manual Snapdragon
Sweetcorn
Tomato
Pansy
Cucurbits
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
used.
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.
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.
8.3.3 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,
1960).
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
Wa}
Wb waxiness/no wax
Wsp
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
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
polygenes.
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
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
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
nitrate.
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
commercially.
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
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
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
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
vulgaris.
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-
phrodite.
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
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
resistance.
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.
Table 8.3
VEGETABLES
Beet
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
Carrot
Different types 400 m
Different cultivars within a type 100m
Cucumber
Ridge cucumber and gherkin 400 m
Different cultivars 200m
Onion
Red cultivars and others 600 m
White cultivars and others 600 m
Different cultivars 200-400 m
Radish
Fodder and horticultural cultivars 600 m
Different types of horticultural cultivars 200m
Different cultivars within a type 50 m
Spinach
Different cultivars 300m
Swede
Agricultural and horticultural cultivars 400 m
Different horticultural cultivars 200m
Turnip
Turnip, turnip rape and Chinese cabbage 400 m
ORNAMENTALS
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
REFERENCES
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,
476-479
HONMA, S. (1959). A method of celery hybridisation, Proc. Am. Soc. hort. Sci., 73,
345-348
LINDQVIST, K. (1960).1nheritance studies in lettuce, Hereditas, 46,387-370
MARKS, G. E. (1972). Selecting asparagus plants as sources of haploids, Euphytica,
22,310-316
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-
265
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
published
WILLS, A. B. and NORTH, C. (1978). Problems of hybrid seed production Acta
Hort., 83, 31-36
FURTHER READING
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
SELECTION, INTRODUCTION AND
MAINTENANCE OF NEW CULTIVARS
9.1 SELECTION
133
134 Plant Breeding
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.
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
Merit.
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.
(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
mark.
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.
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.
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
REFERENCES
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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
EATON, H. J. (1973}. Narcissus propagation, Proc. int. Plant Propagators Soc., 23,
132-134
GOODING, H. J. (1976). Resistance to mechanical injury and assessment of shelf-life
in fruits of strawberry (Fragaria X ananassa), Hort. Res., 16,71-82
HOLLINGS, M. (1965). Disease control through virus-free stock, A. Rev. Phyto-
pathol., 3, 367-396
HUTCHINGS, I. J., JOHN, C. A., PREND, J. and WYATT, C. C. (1962), An instru·
ment for the measurement of the force required for the separation of cucumber
fruit from the peduncle and for the measurement of the firmness of tomato
fruit, Proc. Am. Soc. hort. Sci., 81, 487-492
JONES, 0. P., HOPGOOD, M. E. and O'FARRELL, D. (1977). Propagation in vitro
of M.26 apple rootstocks, j. hort. Sci., 52, 235-238
MURASHIGE, T. (1974). Plant propagation through tissue cultures, Ann. Rev.
Plant. Physiol., 25, 135-166
NORTH, C. and FRITH, L. H. (1959). An instrument for measuring the firmness of
Brussels sprouts,]. hort. Sci., 34, 183-188
OU RECKY, D. K. and BOURNE, M. C. (1968). Measurement of strawberry texture
with an instron machine, Proc. Am. Soc. hort. Sci., 93, 317-325
VOISEY, P. W. and LYALL, L. H. (1965). Methods of determining the strength of
tomato skins in relation to fruit cracking, Proc. Am. Soc. hort. Sci., 86, 597-
609
VOISEY, P. W. and MACDONALD, D. C. (1964). An instrument for measuring the
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563
INDEX
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
143
144
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
82,93
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
147
Zea 69
Zinnia 53, 128, 129
Zygote 48