Breeding Methods

The following are the methods of breeding self-pollinated crop plants.

1. Introduction
2. Selection a) Pure line selection b) Mass selection
3. Hybridization and selection i) Inter varietal a) Pedigree Method b) Bulk Method. c)
Single Seed Descent Method. d) Modified Bulk Method e) Mass - Pedigree Method.
ii) Interspecific hybridization
4. Back cross method
5. Multiline varieties
6. Population approach
7. Hybrids.
8. Mutation breeding
9. Polyploidy breeding
10. Innovative techniques

Plant introduction
Taking a genotype or a group of genotypes in to a new place or environment where they
were not grown previously. Thus, introduction may involve new varieties of a crop already
grown in that area, a wild relative of the crop species or totally a new crop species for that
area.
E.g. a) Introduction of lRRl rice varieties.
b) Introduction of sunflower wild species from Russia
c) Introduction of oilpalm in to Tamil Nadu.
Plant introduction may be of two types. 1. Primary Introduction and 2. Secondary
Introduction
1. Primary Introduction
When the introduced crop or variety is well suited to the new environment, it is directly
grown or cultivated without any alteration in the original genotype. This is known as primary
introduction. E.g. IR. 8, IR 20, IR 34, IR 50 rice varieties; oil palm varieties introduced from
Malaysia and Mashuri rice from Malaysia.
2. Secondary Introduction
The introduced variety may be subjected to selection to isolate a superior variety or it may
be used in hybridization programme to transfer some useful traits. This is known as
secondary Introduction. e.g. In soybean EC 39821 introduced from Taiwan is subjected to
selection and variety Co 1 was developed. In rice ASD 4 is crossed with IR 20 to get Co 44
which is suited for late planting.
Objectives of Plant Introduction
1. To introduce new plant species there by creating ways to build up new industries.
E.g. Oil palm
2. To introduce high yielding varieties to increase food production. E.g. Rice and wheat.
3. To enrich the germplasm collection. E.g. Sorghum, Groundnut.
4. To get new sources of resistance against both biotic and abiotic stresses. E.g. NCAC
accessions to have rust resistance in groundnut. Dasal rice variety for saline
resistance.
5. Aesthetic value – ornamentals are introduced for aesthetic value.
Plant Introduction Agencies
Most of the introductions occurred very early in the history. In earlier days the agencies were
invaders travellers, traders, explorers, pilgrims and naturalists Muslim invaders introduced
in India cherries and grapes. Portuguese introduced maize, ground nut, chillies, potato, sweet
potato, guava, pine apple, papaya and cashew nut. East India Company brought tea. Later
Botanic gardens played a major role in plant Introduction.
A centralized plant introduction agency was initiated in 1946 at IARI, New Delhi. During
1976 National Bureau of Plant Genetic Resources (NBPGR) was started. The bureau is
responsible for introduction and maintenance of germplasm of agricultural and horticultural
plants.
Similarly, Forest Research Institute, Dehradun has a plant introduction organization, which
looks after introduction, maintenance and testing of germplasm of forest trees. Besides
NBPGR the Central Research Institutes of various crops also maintain working germplasm.
All the introductions in India must be routed through NBPGR, New Delhi. The bureau
functions as the central agency for export and introduction of germplasm.
At International level International Board of Plant Genetic Resources (IBPGR) with
headquarter at Rome, Italy is responsible for plant introduction between countries.
Procedure for plant Introduction
The scientist / University will submit the requirement to NBPGR. If the introduction is to be
from other countries, NBPGR will address IBPGR for effecting supply. The IBPGR will
assign collect the material from the source and quarantine them, pack them issue
phytosanitary certificate suitably based on the material and send it to NBPGR. The NBPGR
will assign number for the material, keep part of the seed for germplasm and send the rest to
the scientist. There are certain restrictions in plant introduction. Nendran banana from Tamil
Nadu should be not be sent out of state because of bunchy top disease. Similarly, we cannot
import Cocoa from Africa, Ceylon, West Indies, Sugarcane from Australia, Sunflower from
Argentina.
Functions of NBPGR
1. Introduction maintenance and distribution of germplasm
2. Provide information about the germplasm through regular publications.
3. Conduct training courses to the scientist with regard to introduction and maintenance of
germplasm.
4. Conduct exploratory surveys for the collection of germplasm.
5. To set up Natural gene sanctuaries.
Purpose of plant introduction
The main purpose of plant introduction is to improve the plant wealth of the country. The
chief objectives of plant introduction may be grouped as follows:
To obtain an entirely new crop plant
Plant introductions may provide an entirely new crop species. Many of our important crops,
e.g., Maize, potato, tomato, Tobacco, etc., are introductions. Some recently introduced crops
are Soybean, gobhi sarson, oil palm etc.
To serve as new varieties
Sometimes introductions are directly released as superior commercial varieties. The
Maxican semidwarf wheat varieties Sonora 64 and Lerma Rojo, semidwarf rice varieties TN
1, IR-8 and IR-36 are more recent examples of this type.
To be used in crop improvement
Often the introduced material is used for hybridization with local varieties to develop
improved varieties. Pusa Ruby tomato was derived from a cross between Meeruty and
Sioux, an introduction from U.S.A.
To save the crop from diseases and pests
Sometimes a crop is introduced into a new area to protect it from diseases and pests. Coffee
was introduced in South America from Africa to prevent losses from leaf rust. Hevea rubber,
on the other hand, was brought to Malaya from South America to protect it from a leaf
disease.
For scientific studies
Collections of plants have been used for studies on biosystematics, evolution and origin of
plant species. N.I. Vavilov developed the concept of centres of origin and that of
homologous series in variation from the study of a vast collection of plant types.
For aesthetic value
Ornamentals, shrubs and lawn grasses are introduced to satisfy the finer sensibilities of man.
These plants are used for decoration and are of great value in social life.
Varieties selected from introductions
Many varieties have been developed through selection from introductions. Two varieties of
wheat, Kalyan Sona and Sonalika, were selected from introductions from CIMMYT,
Mexico.
Varieties Developed through Hybridization
Introductions have contributed immensely to the development of crop varieties through
hybridization. All the semidwarf wheat varieties are derived from crosses with Mexican
semi-dwarf wheats. All but few semidwarf rice varieties possess the dwarfing gene from
Dee-geo-woo-gen through either TN1 or IR 8. Thus, almost all these semi-dwarf wheat and
rice varieties have been developed from crosses involving introductions. All the sugarcane
varieties have been derived from the introduced noble canes.
Other examples of varieties developed through hybridization with introductions are Pusa
Ruby tomato obtained from a cross between Meeruti and Sioux; Pusa Early Dwarf Tomato
derived from the cross Meeruti x Red Cloud; Pusa Kesar carrot, Pusa Kanchan turnip etc.
Merits of plant introduction
1. It provides new crop varieties, which are high yielding and can be used directly
2. It provides new plant species.
3. Provides parent materials for genetic improvement of economic crops.
4. Enriching the existing germplasm and increasing the variability.
5. Introduction may protect certain plant species in to newer area will save them from
diseases. E.g. Coffee and Rubber.
Demerits
1. Introduction of new weed unknowingly. E.g. Argemone mexicana, Eichornia and
Parthenium
2. Introduction of new diseases: Late blight of potato from Europe and Bunchy top of banana
from Sri Lanka
3. New pests: Potato tuber moth came from Italy
4. Ornamentals becoming weeds: Lantana camara
5. Introduction may cause ecological imbalance. E.g. Eucalyptus.
Acclimatization
When superior cultivars from neighbouring or distant regions are introduced in a new area,
they generally fail initially to produce a phenotypic expression similar to that in their place
of origin. But later on they pickup and give optimal phenotypic performance, in other words
they become acclimatized to the new ecological sphere. Thus, acclimatization is the ability
of crop variety to become adapted to new climatic and edaphic conditions. The process of
acclimatization follows an increase in the frequency of those genotypes that are better
adapted to the new environment. Factors affecting acclimatization are:
i. Mode of pollination
ii. Amount of variability present in original population
iii. Life cycle of crop plant and
iv. Mutation
SELECTION
Selection is basic to any crop improvement. Isolation of desirable plant types from the
population is known as selection. It is one of the two fundamental steps of any breeding
programme viz., 1. creation of variation and 2. Selection. There are two agencies involved
in carrying out selection: one is Nature itself (Natural selection) and the other is man artificial
selection. Though both may complement each other in some cases, they are mostly opposite
in direction since their aims are different under the two conditions (nature and
domestication). The effectiveness of selection primarily depends upon the degree to which
phenotype reflects the genotype. Before domestication, crop species were subjected to
natural selection. The basic for natural selection was adaptation to the prevailing
environment. After domestication man has knowingly or unknowingly practiced some
selection. Thus, crop species under domestication were exposed to both natural and artificial
selection i.e. selection by man. For a long period, natural selection played an important role
than selection by man. But in modern plant breeding methods natural selection is of little
importance and artificial selection plays an important role.
Basic Principles of Selection: Notwithstanding the highly complex genetic situation imposed
by linkage and epistasis, there are just three basic principles of selection (Walker, 1969):
1. Selection operates on existing variability: The main function of the selection exercise is
to discriminate between individuals. This is possible only when sufficient variation is
present in the material subjected to selection pressure. Thus, selection acts on the existing
variation it cannot create new variation.
2. Selection acts only through heritable differences: only the selected individuals are
permitted to contribute to the next genetion / progenies. Therefore, should there be greater
influence of non-heritable agencies on the individuals selected, the parentprogeny
correlation will be greatly vitiated. Hence the variation among individuals to be selected
must be genetic in nature, since it is the genetic variation that tends to close the gap between
phenotype and genotype. Environmental variability cannot be of any use under selection.
3. Selection works because some individuals are favoured in reproduction at the expense of
other : As a consequence of its past evolutionary history and breeding structure, a population
or a crop consists of highly genetically variable individuals with regards to such diverse
phenomena as differential viability, differential maturity, differences in mating tendencies,
fecundity, and duration of reproductive capacity. Hence some individuals tend to become
superior to others for some or other traits desirable under domestication. These superior
individuals are retained for reproduction while others discarded under selection.
Selection has two basic characteristics viz. 1. Selection is effective for heritable differences
only,
2. Selection does not create any new variation. It only utilizes the variation already present
in a population.
The two basic requirements for select on to operate are:
1. Variation must be present in the population.
2. The variation should be heritable.
Selection intensity: Parcentage of plants selected, to be advanced to next generation, from
a population.
Mass Selection
Large number of plants having similar phenotype are selected and their seeds are mixed
together to constitute a new variety. Thus, the population obtained from selected plants will
be more uniform than the original population. However, they are genotypically different.
Steps:
First year: From the base population select phenotypically similar plants which may be 200
- 2000. Harvest the selected plants as a bulk.
Second year: The bulk seed is divided into smaller lots and grown in preliminary yield trial
along with control variety. Dissimilar phenotypes are rejected. High yielding plots are
selected.
Third to sixth year: The variety is evaluated in coordinated yield trials at several locations.
It is evaluated in an initial evaluation (IET) trial for one year. If found superior it is promoted
to main yield trials for 2 or 3 years.
Seventh year: If the variety is proved superior in main yield trials it is multiplied and released
after giving a suitable name.

Schematic representation of Mass selection

Merits of mass selection
1. Since large number of plants are selected, the variety developed has wider adaptability
as compared to variety developed through Pureline selection.
2. Often extensive yield trials are not necessary. This reduces the time and cost for
developing new variety.
3. Mass selection retail considerable genetic variability in the new variety. Therefore,
another round of mass selection after few years would be effective in improving variety
further.
 4. It is less demanding method so that breeders can devote more time to other breeding
programmes.
Demerits of mass selection
1. The varieties developed through mass selection show variation and are not as uniform
as Pureline varieties. Therefore, such varieties are less liked than pureline varieties.
2. Improvement through mass selection is less than that could be achieved through
pureline selection, because some progenies will be poorer in performance as compared
to the best progeny/pureline present in new variety.
3. In the absence of progeny test it is difficult to find out whether selected plants are
homozygous as there may be some degree of cross pollination in self-pollinated crops,
some of the selected plants may be heterozygous.
4. Varieties developed through mass selection are more difficult to identify than pureline
varieties in seed certification programmes.
5. Mass selection utilizes variability already present in a variety or population, and it can
not generate variability.
Pureline theory given by Johannsen (1903)
 Pureline is the progeny of a single self-fertilized homozygous plant.
 The concept of Pureline was proposed by Johannsen on the basis of his studies with
beans (Phaseolus vulgaris) variety called Princess. He obtained the seeds from the
market and observed that the lot consisted of a mixture of larger as well as smaller size
seeds. Thus, there was variation in seed size. Johannsen selected seeds of different sizes
and grown them individually. Progenies of larger seeds produced larger seeds and
progenies from smaller seeds produced small seeds only.
 This clearly showed that there is variation in seed size in the commercial lot and it has
a genetic basis. He studied nineteen lines altogether. He concluded that the market lot
of the beans is a mixture of purelines. He also concluded whatever variation observed
with in a line is due to environment only.
 Confirmatory evidence was obtained in three ways. In line 13 which is having 450 mg
seed weight he divided the seeds on weight basis. He divided the line into seeds having
200, 300, 400 and 500 mg weights and studied the progenies.
 Ultimately, he got lines having weight ranging from 458 to 475. Thus, the variation
observed is purely due to environment. The second evidence was that selection with in
a Pureline is ineffective. From a Pureline having 840 mg selection was made for large
as well as small seeds.
  After six generations of selection the line for large seed as well as for small seed gave
progenies having 680-690 mg. did not change further. Thus, it was proved that section
within a Pureline is ineffective. In third evidence when parent - offspring regression
was worked in line thirteen found be to zero indicating that variation observed is non
heritable and it is due to environment only.

Confirmatory evidence was obtained in three ways

In the first case, he classified the seed from each pureline into 100 mg classes, and grew them
separately. The mean seed weight of progenies from different seed weight class of a single pure
line were comparable with each other, and with that of the parent pureline. For example, line
no 13 had seed size classes of 200, 300, 400, and 500 mg. The mean seed weights of the
progenies derived from these seed weight classes were 475, 450, 451 and 458 mg respectively.
The second line of evidence came from selection within a pureline. From each pureline, the
largest and the smallest seeds were selected to raise the next generation. In the subsequent
generations, large seeds were selected in the progenies obtained from large seeds while in these
from small seeds selection was done fro small seeds. Six generations of selection was
ineffective in increasing or decreasing the seed size. For example, after 6 generations of
selection, the mean seed weight in Line No 1 was 690 and 680 mg in the progenies selected for
small and large seeds respectively. Thus, selection within a pureline was ineffective.
The third approach was to estimate parent offspring correlation. The value of parent offspring
correlation within line no 13 was – 0.018+ 0.038, that is, zero, while it was 0.336 + 0.008 in
the original seed lot of the Princess which is highly significant. The parent-offspring
correlation will be zero when the variation is nonheritable, while it will be significantly greater
than zero when the variation has a genetic basis, i.e., is heritable.
These observations reveal that the variation for seed size in the original seed lot of Princess
had a genetic basis and was heritable. But the variation within the purelines obtained from the
single seeds selected from this seed lot was purely due to the environment and, therefore, non-
heritable.
The two main conclusions from the Johannsen’s experiment are:
1. A self-fertilized population consists of a mixture of several homozygous genotypes.
Variation in such a population has a genetic component, and therefore selection is effective.
2. Each individual plant progeny selected from a self-fertilized population consists of
homozygous plants of identical genotype. Such a progeny is known as pureline. The variation
within a pureline is purely environmental and, as a result, selection within a pureline is
ineffective.
Origin of variation in pure lines
Pure lines show genetic variation after some time because of the following reasons:
1. Mechanical Mixture: During cultivation, harvesting threshing and storage, other genotypes
may get mixed up.
2. Natural hybridization: Through pure lines are produced in self-pollinated crops, some
amount of natural cross pollination occurs in them also can be avoided by isolation and rouging.
3. Mutation: occur spontaneously in nature at random
Characteristics of purelines
1. All the plants within a pureline have the same genotype
2. The variation within a pureline is environmental and non-heritable
3. Purelines are stable
Progeny test
Evaluation of the worth of plants on the basis of performance of their progenies is known as
progeny test. This was developed by Louis de Vilmorin and so it is also known as the Vilmorin
Isolation principle. Vilmorin worked on sugar beet plants. The progeny test serves two
valuable function:
1. Determines the breeding behaviour of a plant i.e. whether it is homozygous or heterozygous.
2. Whether the character for which the plant was selected is heritable i.e. is due to genotype or
not. Selections have to be based or phenotype and so it is necessary to know the genotype of
the selected plant.
Pureline selection
Pureline selection has been the most commonly used method of improvement of self-pollinated
crops. Almost all the present day varieties of self-pollinated crops are purelines. Pureline
selection has several applications in improvement of self-pollinated crops. It is used to
improve:
1. Local varieties
2. Old pureline varieties and,
3. Introduced varieties
Procedure of Pureline selection
First year:
 An old variety or landrace is used for Pureline selection. Population they selected for
pureline selection is homozygous. Single plant is selected and harvested separately
superior plants must be selected from the mixed population. About 1000-2000 plants are
selected depending on the available resources.
Second year:
 The individual progenies are grown separately with proper spacing the top 15-20
progenies are selected and they are bulked. Poor, defective, weak and segregating
progenies are discarded. Selection should be based on simply inherited character like
plant type, Plant height, grain type, flowering and maturity duration disease resistance
this process may be repeated
Third year:
 Seed of the individual plant progenies are not enough to conduct a replication trail. So,
they are grown in unreplicated trial with check. Here yield of progenies are taken as a
criteria for selection.
Fourth year:
 Replicated yield trials are conducted using the best available check variety. This may be
repeated for 2-3 year. All the observations are recorded
Fifth to Eighth year:
 Promising strains are evaluated at several locations along with strains or check. The best
progeny / strain is released as a new variety and its seed multiplication in initiated for
distributed to the farmer.

Advantages:
 Maximum possible improvement over original variety.
 Pureline varieties are extremely uniform and therefore, more preferred by farmers and
consumers
 Due to uniformity, the variety is easily identified in seed certification programmes.
Disadvantages:
 This method can isolate only superior genotypes, it cannot create new genotypes. And
not applicable in cross pollinated crop.
  Poor adaptability due to narrow genetic base vulnerable for new diseases and pests.
 Pureline selection requires more time, space and expensive yield trials than mass
selection.
 Improvement is dependent on genetic variation present in the original population.
 The breeder has to devote more time to pureline selection than mass selection.

Comparison between pureline and mass selection

Pure line selection Mass selection
1 The new variety is a pureline The new variety is a mixture of purelines.
2 The new variety is highly uniform. In fact, The variety has genetic variation of
the variation within a pureline variety is quantitative characters, although it is
purely environmental. relatively uniform in general appearance.
3 The selected plants are subjected to Progeny test is generally not carried out.
progeny test.
4 The variety is generally the best pureline The variety is inferior to the best pureline
present in the original population. The because most of the purelines included in
pureline selection brings about the it will be inferior to the best pureline.
greatest improvement over the original
variety.
5 Generally, a pureline variety is expected to Usually the variety has a wider adaptation
have narrower adaptation and lower and greater stability than a pureline
stability in performance than a mixture of variety.
purelines.
6 The plants are selected for the desirability. The selected plants have to be similar in
It is not necessary they should have a phenotype since their seeds are mixed to
similar phenotype. make up the new variety.
7 It is more demanding because careful If a large number of plants are selected,
progeny tests and yield trials have to be expensive yield trials are not necessary.
conducted. Thus, it is less demanding on the breeder.

Hybridization
 Natural variability in self-pollinated population is exhausted during selection, for
further improvements new genetic variability has to be created by crossing two different
 pure lines. Hybridization means the mating or crossing of two plants or lines of
dissimilar genotypes.
 The seeds as well as the progeny resulting from the hybridization are known as hybrid
or F1. The progeny of F1 obtained by self or inter mating of F1 plants and the
subsequent generations are called segregating generations. Today hybridization is the
most common method of crop improvement and the majority of the crop varieties have
originated from hybridization.
 One of the objectives of hybridization is to create genetic variation. Two genotypically
different plants are crossed together to obtain F1 generation. F1 is advanced to generate
F2 generation. The degree of genetic variation in F2 and subsequent generation depend
on number of heterozygous genes in F1.

Aims of hybridization
1. To transfer of one or few qualitative characters.
2. Improvement in one or more quantitative character.
3. F1 Hybrid as variety.
I. Combination breeding:
This method is used for the transfer of one or more character into
or single variety from another variety. Eg: improving the yield by correcting the defect.
i.e. disease resistance. The other parent selected for hybridization must have a sufficient
intensity of a character under transfer.
II. Transgressive breeding:
It aims at improving yield or its contributing character through
transgressive segregation. It refers to the appearance of such plants in F2 generation that
are superior to both the parents for one or more character. It is due to accumulation of
plus or favourable genes from the parents as a consequence of recombination. The
parents used for crossing must combine very well and are genetically diverse. So,
pedigree breeding followed by population approach are designed for production of
transgressive segregants.
III. Hybrid varieties:
In self-pollinated crops F1 is more vigorous and high yielding than
the parents. Two parents should combine well to produce outstanding F1 hybrid.
Types of hybridization
 Inter-varietal Hybridization / Intra specific: Parents involved in hybridization belong to
the same species. They may be two strains, varieties or races.
 Varietal crosses may be simple crosses or complex crosses
a.) Simple crosses: Two parents are crossed to produce F1 (A x B)
b.) Complex crosses: More than two parents are crossed to produce the hybrid (A x B) x C
x F1
Procedure of hybridization:
The breeder has clear cut objective in developing the variety. He has to select the variety
accordingly.
1. Choice of parents: One of the parents involved in crosses should be a well-adapted and
proven variety in the area. The other variety should be having the character that are absent in
this variety. Combining ability of the parents serves as useful guides in the selection of parents,
which produce superior F1 and F2.
2. Evaluation of parents: Parents are evaluated for their combining ability.
3. Emasculation: The removal of stamens/anther without affecting the female reproductive
organs, hand emasculation is mostly followed.
4. Bagging: Immediately after emasculation the flowers are enclosed in suitable bags to prevent
cross pollination.
5. Tagging: The emasculated flowers are tied with a thread. The information on date of
emasculation, date of pollination, names of female and male parents are recorded in the tag
with pencil. The name of the female parent is written first then male parent.
6. Pollination: Mature fertile and viable pollen from the male parent should be placed on
receptive stigma of emasculated flowers to bring about fertilization. Pollen grain is collected,
allowed for dehiscence and pollination is carried out with camel hair brush.
7. Harvesting and storing of F1 seeds: The crossed heads/pods should be harvested and
threshed. The seeds should be dried and properly stored to protect them from storage pests.
8. Rising the F1 generation: Identify the selfed seeds in the F1 generation by using dominant
marker gene. Larger F1 population is desirable, because both the genes are present in
heterozygous condition.
9. Selfing: To avoid cross pollination and to ensure self-pollination. In often cross-pollinated
crops, they are bagged to prevent cross pollination.
Distant Hybridization
When crosses are made between two different species or between two different genera, they
are generally termed as distant hybridization (or) wide hybridization
History
Thomas Fairchild 1717 was the first man to do distant hybridization. He produced a hybrid
between two species of Dianthus Dianthus caryophyllus (Carnation) x D. barbatus (Sweet
william)
Inter generic hybrid produced by Karpechenko, a Russian Scientist in 1928. Raphano brassica
is the amphidiploid from a cross between Radish (Raphanus sativus) and cabbage (Brassica
oleracea). Triticale was produced by Rimpau in 1890 itself. Triticale is an amphidiploid
obtained from cross between wheat and rye. Another example is Saccharum noblisation
involving three species.
Hybrids in self-pollinated crops - problems and prospects
Exploitation of heterosis through F1 hybrids has hitherto been the prerogative of cross-
pollinated crops, chiefly due to their breeding systems favouring allogamy. However,
possibilities of working for such a proposition have recently been realized in self-pollinated
corps also. Indeed, exploitation of hybrid vigour in autogamous/ self-pollinated crops is easy
and less time consuming as homozygous inbreds are already available. There is practically no
difference with regard to hybrid breeding between self and cross-pollinated crops. But the
prospects of hybrids in self-pollinated crops are dependent on three major considerations:
1. How high a heterotic effect can be gained under optimal production conditions.
2. In fact, a breeder's main concern is the magnitude rather than the frequency of occurrence of
heterosis in crops. Thus, the consideration is whether or not it is possible to obtain economically
viable heterosis.
3. How much of the yield surplus due to high heterosis can offset the extra seed cost? In major
self-pollinated crops like wheat, barley, rice, etc., the seed rate per unit area is exorbitant and
hence the hybrid seed requirement is also more.
4. How efficient and effective is the mechanism of cross-pollination in self-pollinated crops?
By nature, self-pollinated crops are shy pollinators with very poor pollen manoeuvrability (or
movability to effect allogamy). Therefore, the efficiency (degree of allogamy) with which cross
pollination can take place on a commercial scale is the true determinant of the success of a
hybrid programme in self-pollinated crops.
5. Among self-pollinated crops, F1 hybrids have been graduated into the farmer's field in rice,
barely, tomato, Sorghum (often-cross-pollinated) and wheat.
Pedigree method of handling segregating generations
 In pedigree method individual plants are selected from F2 and their progenies are tested in
subsequent generations. A record of the entire parent off spring relationship is maintained
 and known as pedigree record. The pedigree may be defined as a description of the ancestor
of an individual and it generally goes back to some distant ancestor. So, each progeny in
every generation can be traced back to the F2 plant from which it is originated.
 This method used for selection from segregating population of crosses in self-pollinated
crops. It is used for combination or transgressive breeding.
Procedure:
1. Hybridization: The selected parents are crossed to produce a simple / complex cross (F1
seed)
2. F1 generation: F1 seeds are space planted to each produces maximum number of F2 seed.
15-30 F1plants are sufficient to produce good F2 populations.
3. F2 generation: 200-10000 plants are space planted and 100-500 plants are selected and
their seeds are harvested separately. He should select as many as F2 plants as he can handle
efficiently. The selection depends on skill of the breeder and his ability to judge to select
F2 which produce good progeny.
4. F3 generation: Individual plant progeny are space planted. Individual plant with desirable
characters from superior progenies is selected.
5. F4 generation: Individual plants progenies are space planted desirable pants are selected
undesirable progenies are rejected. Progenies are compared visually and more plants are
selected from superior progenies. Selection of desirable plants from superior
progenies selection is practiced within / between family.
6. F5 generation: Many families have reached homozygous and may be harvested in bulk.
The breeder has to assess the yielding potential of progenies, 25-100 progenies are
advanced and tested in preliminary yield trial.
7. F6 generation: Multi row plots and evaluated visually progenies harvested bulk and they
have become homozygous.
8. F7 generation: Preliminary yield trail with replication to identify the superior progenies.
Progenies are evaluated for other component character 2-5 outstanding lines superior to
check are advanced to multi location testing.
9. F8 –F10 generation: Replicated yield trial at several locations. They are tested for yield as
well as for resistance.
10. F11 generation: Seed multiplication and release.
Merits:
1. Maximum opportunity for the breeder to use his skill and judgment for the selection of
plants in segregating generation.
2. It provides information about the inheritance of qualitative character from the pedigree
record.
3. Chances of recovering transgressive segregants is more.
4. Plants and progenies with visible defects and weaknesses are eliminated at an early stage.
5. Well suited for the improvement of easily identified and simply inherited characters.
Demerits:
1. Maintenance of accurate pedigree record is tedious and takes up valuable time
 2. Selection of progenies in every generation laborious, time consuming. Difficult to handle
many crosses.
3. No opportunity for natural selection.
4. Possibility of losing the valuable genotype is early segregating generation.
5. The success of this method largely depends on skill of the breeder.
Bulk method
Bulk method was first used by Nilsson Ehle in 1908. F2 and the subsequent generations are
harvested as bulks to raise the next generation. At the end of bulking period individual plants
are selected and evaluated in a similar manner as in the pedigree method. The duration of
bulking may vary from 7-30 generation artificial selection may seldom be practiced
1. Isolation of homozygous lines
It is used for the isolation of homozygous lines with a minimum of effort and expense. The
population is carried to F6-F7 as Bulk, where it reaches homozygosity. Individual plants are
selected and evaluated to derive pure line. So preliminary yield trails are conducted to derive
homozygous lines.
2. Waiting for the opportunity for selection:
Selection for resistance to disease, lodging and cold depends upon the presence of suitable
environmental conditions favouring epidemic. Waiting till such environment do occur so the
segregating generations are carried as bulk in such environment. Individual plants are
selected and handled as in pedigree method. The duration of bulking depends upon the
occurrence of the concerned environment. This is known as mass pedigree method of Harlan.
3. Opportunity for natural selection:
Maintenance of bulk is inexpensive and without much efforts. Some bulk populations are
carried up to F20 to F30 to provide an opportunity for natural selection to act on their
composition. Up to F6 generation the population is heterozygous and after F7 generation
natural selection to act on homozygous plants and would change the frequency of
homozygous genotypes present in the population. It is assumed that natural selection would
favour higher yielding genotypes and eliminate poorer genotypes.
Procedure of bulk method:
1. Hybridization: Parents are selected and crossed
2. F1 generation: F1 is space planted more than 200 F1 plants
3. F2-F6 Generation: Planted at commercial seed rate, spacing and harvested as bulk, during
this period. Frequency of population changes due to outbreak of disease or pest.
4. F7 generation: 50000 plants are space planted about 1000-5000 plants with phenotype is
selected and the seeds are harvested separately.
5. F8 generation: Individual plant progenies are single/multi row plants, since progenies are
homozygous and harvested in bulk weak and inferior progenies are rejected and 100-300
individual plant progenies with desirable characters.
6. F9 generation: Preliminary yield trial with standard check, yield and quality parameter is
taken for selection.
7. F10---F12 generations: Replicated yield trails are conducted. Yield and its component
characters are evaluated along with the check. Superior progenies are released as variety
8. F13 generation: Seed multiplication of the newly released variety and distribution to
farmers.
Merits
1. Simple, convenient and inexpensive method
2. Natural selection is likely to increase the frequency of superior genotypes in the population.
Therefore, progenies selected from long term bulks are likely to be superior to those selected
from F2 or short bulks.
3. Little work and attention are required in F2 and subsequent generations, and no pedigree
record is to be kept. This save time and labour, and the breeder can concentrate more on
other breeding projects.
4. Since large populations are grown, chances of getting transgressive segregants are more.
5. Individual plant selection is done when population has become homozygous. Therefore,
selection is expected to be more effective than in F2 and F3 generations.
6. Particularly suited to small grain crops grown in high crop densities.
7. Natural selection is expected to improve characters like adaptation to prevailing
environment which are otherwise difficult to assess and select for.
Demerits
1. The major disadvantage of bulk method is longer time taken to develop a new variety.
Natural selection becomes important only after F10 generation and bulking may have to be
done up to F20 or more which is considerably longer than the time taken in pedigree method
2. In short term bulks, natural selection has little effect on the genetic composition of
populations. But short-term bulks are useful for isolation of homozygous lines.
3. It provides little opportunity to breeders to exercise their skill in selection.
4. A large number of progenies have to be handled at the end of the bulking period.
5. Information on inheritance of characters cannot be obtained
6. Off season and green house facilities cannot be used to advance the generation since
environment at such locations may be markedly different from that in target location.

Single seed decent method

 Single seed descent method This method is a modification of bulk method
 Single seed from each F2 plants is bulked to raise the F3 generation. Similarly, F3, F4, F5
generation when the plants are homozygous plant progenies are advanced to next
generation. Selection is done mainly among the progenies and number of progenies is
sufficiently reduced to permit replicated trail. Individual plants may be selected from
 outstanding families showing segregation. So preliminary yield trial and quality tests begin
in F7 to F8.
Objectives
1. Rapidly advance of generation of crosses.
2. F2 and subsequent generation are grown with a very high plant density.
3. F2 plant is represented equally in the end population.
4. Off season nursery/green house facilities are utilized.
5. Maximum possible speed.
6. Require very little space/effort/ labour.
7. Do not permit any form of selection during the segregating generation.
8. In each successive generation the population size become small due to poor generation
and death of plants due to disease/pest.
 Pedigree Method Bulk Method
Individuals plants are selected in F2 and F2 and the subsequent generation are
Segregation generation and individual plant maintained as bulk
progenies are grown
Artificial selection, Artificial disease Artificial selection, artificial disease
epidemics are an integral part epidemic can be created to assist
natural selection.
No role of Natural section Natural selection determines the
composition of population.
Pedigree record have to be maintained which No pedigree records are maintained
is laborious and time consuming
It takes 12 years to develop new variety More than 12 years bulk population > 10
years for effective natural selection
Widely used method Limited use
Needs close attention from F2 onwards It is simple convenient.
Segregating generation are space planted to Bulk population are planted at commercial
permit individual plant selection planting rate
Population size is small Larger population are grown and the
natural selection expected to the increase
the chances recovery of transgressive
segregants

Back cross method

A Crossing between a F1 hybrid or its segregating generation with one of its parents is known
as Back cross. The hybrid and its progenies in the subsequent generations are repeatedly
back crossed to one of their parents. As a result, the genotype of back cross progeny becomes
increasingly similar to that parent to whom the back crosses are made. At the end of 6-8 back
crosses, the progeny would be almost identical with the parent involved in back crossing.
Objective:
1. To improve one or two specific defects of a high yielding variety and a well-adapted
variety with desirable character.
2. The characters lacking in this variety are transferred to it from a donor parent without
changing the genotype of this variety except for the genes being transformed.
Requirements of back cross breeding
1. Suitable recurrent parent must be available which lacks in one or two characteristics.
2. A suitable donor parent must be available; the character must be highly intense.
 3. The character(s) to be transferred must have high heritability and preferably, should be
governed by one or few genes
4. A sufficient number of back crosses should be made so that genotype of recurrent parent
is recovered in full. Ordinarily, 6-7 backcrosses are sufficient for the purpose.
Applications of back cross breeding
1. Inter varietal transferring of simply inherited traits. Characters governed by one or two genes
like disease resistance are successful.
2. Inter varietal transfer of quantitative characters and highly heritable quantitative characters
like earliness, plant height, seed size and seed shape are transferred.
3. Inter specific transfer of simply inherited characters: Disease resistance is transferred from
related species to cultivated species. Inter specific transfer of genes are easy when the
chromosome of the two species pair regularly.
4. Transferring of cytoplasm: wild species cytoplasmic are transferred to cultivated species
transfer of male sterility. The variety or species from which the cytoplasm is to be transferred
is used as the female parent. The parent to which the cytoplasm is to be transferred is used
as the male parent in the original cross and back cross. After 6-8 back crosses the progeny
would have the nuclear genotype of the recurrent parent and the cytoplasm from the donor
parent.
5. Transgressive segregation: F1 is back crossed to one or two times to the recurrent parent
leaving much heterozygosity for transgressive segregation to appear. In the second
modification two or more recurrent parent may be used in the back-cross progeny to
accumulate genes from them into the back cross. Progeny of the new variety is not exactly
like any one of the recurrent parents.
6. Production of isogenic lines: Isogenic lines are identical in their genotype except for one
gene
7. Germplasm conversion: When valuable germplasm cannot be utilized in breeding
programmes and may be used as recurrent parent in separate back cross programme these
lines are called converted lines.
Procedure for transfer to dominant gene
E.g. High yielding and widely adapted wheat variety A is susceptible to stem rust another
variety B is resistance to stem rust. Stem rust is dominant to susceptibility.
1. Hybridization: Variety A is crossed to variety B. Generally, variety A should be used as
female parent. This would help in identification of selfed plants.
2. BC1 generation: F1 plants are back crossed to variety A. Since all the F1 are heterozygous
for rust resistance, selection for rust resistance is not necessary.
3. First BC1 generation: Half of the plants in BC1 generation are resistant and the remaining
half would be susceptible to stem rust. Rust resistant plants are selected and back crossed to
variety A.
4. BC2 to BC 5 generation: Segregation would occur for rust resistance. Rust resistant plants
are selected and back crossed to variety A.
5. BC6 generation: BC6 plants will have 99 percent genes from variety A Rust resistant plants
are selected and selfed, their seeds are harvested separately
6. BC6F2 generation: Individual plants progeny from the selfed seeds of the selected plants
are grown. Rust resistant plant similar to the plant type of variety A are selected and they are
selfed. Seeds are harvested separately.
7. BC6 F3 generation: Individual plants progeny are grown. Progenies homozygous for rust
resistant and similar to plant type of variety Aare harvested in bulk. Several similar progenies
are usually mixed to constitute the new variety.
8. Yield tests and seed multiplication: New variety is tested in replicated yield trails along
with variety A as check. Ordinarily, the new variety would be identical to variety A in
performance, detailed yield trials are therefore, generally not required.
Procedure for transfer to recessive gene
1. Hybridization: Variety A, recurrent parent is crossed as female to variety B, donor parent
for rust resistance.
2. BC1 generation: F1 plants are back crossed to variety A. Since all the F1 are heterozygous
for rust resistance, selection for rust resistance is not necessary.
3. First BC1 generation: Since rust resistance is recessive, all plants will be susceptible to
rust. Therefore, there is no test for rust resistance and all plants are self -pollinated.
4. BC1F2 generation: Plants are inoculated with rust spores and rust resistant plants are
selected and back crossed with recurrent parent. Selection will be done for plant type like
recurrent parent A.
5. BC2 generation: There is no rust resistance test. Plants are selected for resemblance to
recurrent parent A and back crossed with recurrent parent A.
6. BC3 generation: There is no disease test. The plants are self pollinated to raise F2.
Selection is done for plant type of recurrent parent A.
7. BC3F2 generation: Plants are inoculated with rust spores. Rust resistant plants resembling
recurrent parent A are selected and back crossed to recurrent parent A.
8. BC4 generation: No rust resistance test and plants are back crossed to recurrent parent A.
9. BC5 generation: There is no rust resistance test. Plants are self-pollinated to raise F2
generation.
10. BC5F2 generation: Plants are inoculated with rust spores and rigid selection is done for
resistance and for the characteristics of recurrent parent A. Selfed seeds from selected plants
are harvested separately.
11. BC5F3 generation: Individual plant’s progeny from the selfed seeds of the selected plants
are grown and subjected to rust epiphytotic. A rigid selection is done for rust resistant plant
similar to the plant type of variety A. Seeds from several similar rust resistant homogenous
progenies are usually mixed to constitute the new variety.
12. Yield tests and seed multiplication: New variety is tested in replicated yield trails along
with variety A as check. Ordinarily, the new variety would be identical to variety A in
performance, detailed yield trials are therefore, generally not required.
Merits:
1. Back cross method retains all desirable character of a popular adapted varieties and
replaces undesirable allele at particular locus
2. Useful for the transfer of disease resistance and incorporation of quality traits into a variety
3. This is used for the development of isogenic lines,
4. Extensive tests are not required 2-3 generation can be raised in off season nurseries
greenhouses; it would save time.
5. This is the only method for the inter specific gene transfer and transfer of cytoplasm.
6. Male sterility and fertility restoration genes can be transferred to various back ground.
Demerits:
1. New variety cannot be superior to recurrent parent except for the character transferred
2. It involves lot of crossing work. 6-8 back cross is often difficult and time consuming.
3. Sometime undesirable gene linked with desirable also may be transferred.
4. By the time the back cross programme the recurrent parent may have been replaced by
other varieties superior in yield and other character.
Multiline Varieties
Generally, pureline varieties are highly adapted to a limited area, but poorly adapted to wider
regions. Further, their performance is not stable from year to year because of changes in
weather and other environmental factors. Purelines often have only one or a few major genes
for disease resistance, such as, rust resistance, which make them resistant to some races of
the pathogen. New races are continuously produced in many pathogens, which may
overcome the resistance present in the pureline varieties. For example, Kalyan Sona wheat
(T. aestivum) originally resistant to brown rust (leaf rust), soon became susceptible to new
races of the pathogen. To overcome these limitations, particularly the breakdown of
resistance to disease, it was suggested to develop multiline varieties.
Multiline varieties are mixtures of several purelines of similar height, flowering and maturity
dates, seed colour and agronomic characteristics, but having different genes for disease
resistance. The purelines constituting a multiline variety must be compatible, i.e., they
should not reduce the yielding ability of each other when grown in mixture.
In 1954, Borlaug suggested that several purelines with different resistance genes should be
developed through back cross programmes using one recurrent parent. This is done by
transferring disease resistance genes from several donor parents carrying different resistant
genes to a single recurrent parent. Each donor parent is used in a separate backcross
programme so that each line has different resistant gene or genes. Five to ten of these lines
may be mixed depending upon the races of the pathogen prevalent in the area. If a line or
lines become susceptible, they would be replaced by resistant lines. New lines would be
developed when new sources of resistance become available. The breeder should keep
several resistant lines in store for future use in the replacement of susceptible lines of
multiline verities.
Merits of Multiline varieties
1. All the lines are almost identical to the recurrent parent in agronomic characteristics,
quality etc. Therefore, the disadvantages of the pureline mixtures are not present in the
multiline varieties.
2. Only one or a few lines of the mixture would become susceptible of the pathogen in
anyone season. Therefore, the loss to the cultivator would be relatively low.
3. The susceptible line would constitute only a small proportion of the plants in the field.
Therefore, only a small proportion of the plants would be infected by the pathogen.
Consequently, the disease would spread more slowly than when the entire population was
susceptible. This would reduce the damage to the susceptible line as well.
Demerits of Multiline Varieties
 1. The fanner has to change the seed of multiline varieties every few years depending upon
the change in the races of the pathogen.
2. There is a possibility that a new race may attack all lines of a multiline variety.
Achievements
Multiline variety appears to be a useful approach to control diseases like rusts where new
races are continuously produced. In India, three multiline varieties have been released in
wheat (T. aestivum). Kalyan Sona, one of the most popular varieties in the late sixties, was
used as the recurrent parent to produce these varieties. Variety 'KSML 3' consists of 8 lines
having rust resistance genes from Robin, Ghanate, Kl, Rend, Gabato, Blue Brid, Tobari etc.
Multiline 'MLKS 11' is also a mixture of 8 lines; the resistance is derived from E 6254, E
6056, E 5868, Frecor, HS 19, E 4894 etc. The third variety, KML 7406 has 9 lines deriving
rust resistance from different sources.
Dirty Multiline
This term is used when a multiline is having one or two susceptible lines also. The idea of
including susceptible lines is to prevent race formation.
Hardy Weinberg Law
 Cross pollinated crops are highly heterozygous due to the free inter mating among them
so these are random mating populations. Because each individual of the population has
equal opportunity of mating with any other individual. It is also known as
mendelian/panmictic population. A Mendelian population may be thought of having a
gene pool consisting of all gametes produced by the population. So, gene pool may be
defined as the sum total of all genes present in the population. A population consists of all
such individuals that share the same gene pool has an opportunity to inter mate with each
other and contribute to the next generation of the population.
 Each generation of a Mendelian population may be considered to arise from a random
sample of gametes from the gene pool of previous generation. Hence, it is not easy to
follow the inheritance of a gene in a Mendelian population. It cannot be estimated by using
the techniques of classical genetics. So, to understand the genetic makeup of such
population a population genetics has been developed.
 This law is independently developed by Hardy (1908) in England and Weinberg (1909) in
Germany. The law states that the gene and genotype frequencies in a Mendelian population
remain constant from generation after generation if there is
no selection, mutation, migration or random drift.
 The frequencies of these genotypes for a locus with two alleles A and a would be P 2(AA),
2pq (Aa)and q2(aa)
Where, p= Frequency of ‘A’ allele in the population.
 q= Corresponding frequency ‘a’ allele in the population the sum of p+q is equal=1
Such a population would be at equilibrium since the genotypic frequencies would be stable,
that is, would not change from one generation to the next. This equilibrium is known as
Hardy Weinberg equilibrium.
Migration: Migration is the movement of individual into a population from a different
population. Migration may introduce new alleles into the population or may change the
frequency of existing allele. The amount of change in gene frequency ‘q’ will primarily
depend upon two factors.
a. Ratio of migrant individuals to those of the original population.
b. The Magnitude of difference between the values of q in the population and in the migrants.
So, in plant breeding migration is by inter varietal crosses or poly crosses.
Mutation: mutation is a sudden heritable change in an organism and is generally due to a
structural change in a gene. It may produce a new allele not present in the population or may
change the frequency of existence allele that 10-6 mutation is detected. So, such effects in
breeding population may be ignored.
Random drift or genetic drift: It is a random change in gene frequency due to sampling error.
Random drift is more in small population than larger. Ultimately, the frequency of one of the
alleles becomes zero and that of the other allele becomes one. The allele with the frequency
one is fixed in the population because there would be no change in the frequency. So, all genes
become homozygous. The genetic drift can be reduced by handling large population.
Inbreeding: Mating between individuals sharing a common parent in their
ancestry inbreeding reduces the proportion of the heterozygosity and increase the frequency of
homozygosity by the rate of decrease in heterozygosity is equal to ½ N (N- Number of plants
in the population) per generation in monoecious or hermaphrodite species. In dioecious species
and monoecious species where self-pollination is prevented the decrease in heterozygosity is
low. In small population, even with strict random mating / strict cross pollination the frequency
of homozygotes increases while that of heterozygotes decreases due to inbreeding
Selection: The selection in a random mating population is highly effective in increasing or
decreasing the frequency of allele, but it is unable to either fix or eliminate them. However, in
combination with a system of inbreeding, selection is highly efficient in the fixation and
elimination of an allele.
Inbreeding
It is mating between individuals related by descent or having common ancestry. The highest
degree of inbreeding is obtained by selfing. Inbreeding depression refers to decrease in
fitness and vigour due to inbreeding. The degree of inbreeding is measured by
the inbreeding coefficient.
History of inbreeding:
In breeding depression has been recognised by man for a long time. Knowing the
consequences of inbreeding many societies have prohibited marriages between closely
related individuals. Darwin in 1876 published a book “cross and self-fertilization in
vegetable kingdom” in which he concluded that progenies obtained from self-fertilization
were weaker in maize. Detailed and precise information on inbreeding in maize was
published by East in 1908 and Shull in 1909.
The different effects of inbreeding are:
1. Appearance of Lethal and Sublethal Alleles: IB results in appearance of lethal;
sublethal and sub vital characters. e.g.: Chlorophyll deficiencies, rootless seedlings, flower
deformities – They do not survive, they lost in population.
2. Reduction in vigour: General reduction in vigour size of various plant parts.
3. Reduction in Reproductive ability: Reproductive ability of population decreases
rapidly. Many lines reproduce purely that they cannot be maintained.
4. Separation of the population into distinct lines: Population rapidly separates into
distinct lines i.e. due to increase in homozygosity. This leads to random fixation of alleles in
different lines. Therefore, lines differ in genotype and phenotype. It leads to increase in the
variance of the population.
5. Increase in homozygosity: Each line becomes homozygous. Therefore, variation within
a line decreases rapidly. After 7-8 generations of selfing the line becomes more than 99%
homozygous. These are the inbreds which have to be maintained by selfing.
6. Reduction in yield: Inbreeding leads to loss in yield. The inbreds that survive and
maintained have much less yield than the open pollinated variety from which they have been
developed.
Degrees of inbreeding depression
Various plant species exhibit different degrees of inbreeding depression. The depression
may be from very high to nil. Based on degree of depression, the plant species can be
grouped into 4 broad categories.
 1. High inbreeding depression: E.g. Lucerne, Carrot. Inbreeding leads to severe
depression and exhibit lethal effects. After 3 or 4 generations of selfing it is hard to maintain
lines.
2. Moderate inbreeding depression: E.g. Maize, Jowar, Bajra. Though lethal effects are
there, lines can be separated and maintained.
3. Low inbreeding depression: E.g. Cucurbits, Sunflower. Only a small degree
of inbreeding depression is observed.
4. No inbreeding depression: The self-pollinated crops do not show inbreeding depression.
Heterosis
The term heterosis was first used by Shull in 1914. Heterosis may be defined as the
superiority of an F1 hybrid over both of its parents in terms of yield or some other character.
Generally, heterosis is manifested as an increase in vigour, size, growth rate, yield or some
other characteristic.
History
Koelreuter is first reported hybrid vigour in tobacco produced artificial hybrids. In 1876,
Darwin concluded that hybrids form unrelated plant type were highly vigorous. Most of our
present knowledge on heterosis comes from the work on maize. Crossing inbred lines rather
than open pollinated varieties produces the commercial maize hybrids. Hybridization
between inbreds developed from the same variety or from closely related varieties produced
only a small degree of heterosis.
 But a vast majority of the cases of heterosis are cases of superiority of hybrids over their
parents. Hybrid vigour describes only superiority of hybrids over the parents. The few cases
where F1 hybrids are inferior to their parents may also be regarded as cases of hybrid vigour
in the negative direction.
Heterosis is the superiority of a hybrid over its parents.
1. Increased yield: Heterosis is generally expressed as an increase in the yield of hybrids.
The yield may be measured in terms of grain, fruit, seed, leaf, tubers or the whole plant.
2. Increased reproductive ability: More number of flowers/fruits/seeds. Increase in Size and
General Vigour: The hybrids are generally more vigorous, i.e., healthier and faster
growing and larger in size than their parents.
3. Better quality: In many cases, hybrids show improved quality. For example, many hybrids
in onion show better keeping quality, but not yield, than open-pollinated varieties.
4. Earlier flowering and maturity: In many cases hybrids are earlier in flowering and
maturity than the parents. But earliness is highly desirable in many situations particularly
in vegetables.
5. Greater resistance to disease and pest: Some hybrids are known to exhibit a greater
resistance to insect or diseases than their parents.
6. Greater adaptability: Hybrids are generally more adapted to environmental changes than
inbreds.
7. Faster growth rate: In some cases, hybrids show a faster growth rate than their parents.
But the total plant size of the hybrids may be comparable to that of parents. In such cases,
a faster growth rate is not associated with a larger size.
8. Increase in the number of a plant part: In some cases there is an increase in the number of
nodes, leaves and other plant parts, but the total plant size may not be larger.
Estimation of heterosis
1. Average heterosis:
It is the heterosis where F1 is superior to mid parent value. In other words, superior to
average of two parents.
F1 - MP
--------- x 100
MP
Where F1 = Mean of hybrid
MP = Mid parental value.
(P1 + P2)
MP = ----------
2
where P1 = Parent 1; P2 = Parent 2
This type of heterosis is of no use in agriculture since the superiority is below the better
parent value
2. Heterobeltiosis:
Superiority of F1 over the better parent.
F1 - BP
--------- x 100
BP
Where BP = Mean of Better Parent.
3. Economic heterosis:
Superiority of the F1 compared to the high yielding commercial variety in a particular crop.
F1 - CV
--------- x 100
CV
Where CV = Mean of Commercial Variety.
4. Negative heterosis:
Performance of F1 inferior to better parent / mid parent value. - e.g. Duration.
Breeding Methods for Cross Pollinated Crops
Populations of cross-pollinated crops are highly heterozygous. When inbreeding is practiced,
they show severe inbreeding depression. So, to avoid inbreeding depression and its undesirable
effects, the breeding methods in the crop is designed in such a way that there will be a minimum
inbreeding. To isolate plants with superior genotypes. Individual plants are highly
heterozygous. The progeny from such plants are highly heterogeneous and usually different
from the parent plant due to segregation and recombination. This would increase the frequency
of desirable genes combinations. As a result, the phenotype of the population would be
favorably changed. In cross pollinated species genotype of the individual plants is of little
importance but it is the frequency of desirable genes or alleles in the population as a whole that
determines the value of a population.

The breeding methods commonly used in cross pollinated crops may be broadly grouped into
two categories.
I. Population improvement
2. Hybrids
3. Synthetics and Composites
Population Improvement
a) Intra-population improvement
In cross pollinated crops, population improvement is used to enhance the frequency of desirable
alleles in a population. It may be intra or interpopulation improvement. Requirement of intra-
population improvement:
 Presence of genetic variation for different traits
 Presence of additive genetic variation with partial dominance in population
 High heritability of traits
 Absence of any undesirable linkages
 There should not be negative association between desirable traits
 Population under improvement should be grown to fairly large plant population size in
selection blocks.
 Improvement is expected to be high for characters which can be easily and precisely
rated particularly before flowering.
Here mass selection or its modification are used to increase the frequency of desirable alleles
thus improving the characteristics of the population.
 1. Mass selection without progeny testing: Plants are selected on the basis of their
phenotype and no progeny test carried out
2. Mass selection with progeny testing: Initial selection on the basis of phenotype but
final selection on the basis of progeny test. This includes the ear-to -row method and
recurrent selection
Mass selection
This is similar to the one, which is practiced, in self-pollinated crops. A number of plants are
selected based on their phenotype and open pollinated seed from them are bulked together to
raise the next generation. The selection cycle is repeated one or more times to increase the
frequency of favourable alleles. Such a selection is known as phenotypic recurrent selection.
Mass selection is a form of maternal selection as there is no control on pollination.

Merits
i) Simple and less time consuming
ii) Highly effective for character that are easily heritable. Eg. Plant height, duration.
iii) It will have high adaptability because the base population is locally adapted one.
Demerits
1. Selection is based on phenotype only which is influenced by environment
2. The selected plants are pollinated both by superior and inferior pollens present in the
population.
3. High intensity of selection may lead reduction in population there by leading to inbreeding.
To overcome these defects modified mass selection is proposed they are

a) Detasseling: This is practiced in maize. The inferior plants will be detasseled there by inferior
pollen from base population is eliminated.

b) Panmixis: From the selected plants pollen will be collected and mixed together. This will be
used to pollinate the selected plants. This ensures full control on pollen source.

c) Stratified mass selection: Here the field from which plants are to be selected will be divided
into smaller units or plots having 40 to 50 plants / plot. From each plot equal number of plants
will be selected. The seeds from selected plants will be harvested and bulked to raise the next
generation, by dividing the field into smaller plots, the environmental variation is minimized.
This method is followed to improve maize crop. It is also known as Grid method of mass
selection

B) Family selection
Selection based on means of the individual plant progenies or families.
I) Half sib family selection: Half sibs are those, which have one parent in common. Here only
superior progenies are planted and allowed to open pollinate.
1. Ear to row method: It is the simplest form of progeny selection. Which is extensively used
in maize. This method was developed by Hopkins (1908).
a) A number of plants are selected on the basis of their phenotype. They are allowed to open
pollinate and seeds are harvested on single plant basis.
b) A single row of say 50 plants i.e. progeny row is raised from seeds harvested on single
plant basis. The progeny rows are evaluated for desirable characters and superior progenies are
identified.
c) Several phenotypically superior plants are selected from progeny rows. There is no control
on pollination and plants are permitted to open pollinate. Though this scheme in simple, there
is no control over pollination of selected plants. Inferior pollen may pollinate the plants in the
progeny row. To overcome this defect, the following method is suggested:
a) At the time of harvest of selected plants from base population on single plant basis, part of
the seed is reserved.
b) While raising progeny rows, after reserving part of the seeds, the rest are sown in smaller
progeny rows.
c) Study the performance of progenies in rows and identify the best ones.
d) After identifying the best progenies, the reserve seeds of the best progenies may be raised in
progeny rows.
e) The progenies will be allowed for open pollination and best ones are selected.
There are number of other modifications made in the ear to row selection. For example,
i. The selected progenies may be selfed instead of open pollination
ii. The selected plants may be crossed to a tester parent. The tester parent may be an open
pollinated variety, or inbred
iii. The progeny test may be conducted in replicated trial.
II) Full sib family selection: Full sibs are those which are produced by mating between
selected plants in pairs. Here the progenies will have a common ancestry. The crossed
progenies are tested.
AxB BxA
III) Inbred or selfed family selection: Families produced by selfing.
a) S1 family selection: Families produced by one generation of selfing. These are used for
evaluation and superior families are intermated (Simple recurrent selection).
b) S2 family selection: Families obtained by two generations of selfing and used for evaluation.
Superior families are intermated.
Merits of Family selection
1. Selection based on progeny test and not on phenotype of individual plants.
2. In breeding can be avoided if care is taken raising a larger population for selection.
 3. Selection scheme is simple.
Demerits
1. No control over pollen source. Selection is based only on maternal parent only.
2. Compared to mass selection, the cycle requires 2-3 years which is time consuming.

b) Inter -population improvement

Suggested by Hayes and Garber (1919) and independently by East and Jones (1920) and
breeding schemes developed during 1940s
Recurrent selection
This is one of the breeding methods followed for the improvement of cross-pollinated crop.
The different Recurrent selection schemes are variations of progeny selection, the difference
lying in the manner of obtaining the progeny for evaluation, and in making the all possible inter
crosses among the selected lines in place of open pollination.
Here single plants are selected based on their phenotype or by progeny testing. The selected
single plants are selfed. In the next generation they are intermated (cross in all possible
combinations) to produce population for next cycle of selection. The recurrent selection
schemes are modified forms of progeny selection programmes. The main difference between
progeny selection and recurrent selection:
i) The manner in which progenies are obtained for evaluation.
ii) Instead of open pollination, making all possible inter crosses among the selected lines.
The recurrent selection schemes are of 4 different types:
ii. Simple Recurrent Selection
iii. Recurrent Selection for GCA
iv. Recurrent Selection for SCA
v. Reciprocal Recurrent Selection
1. Simple recurrent selection
In this method a number of desirable plants are selected and self-pollinated. Separate progeny
rows are grown from the selected plants in next generation. The progenies are intercrossed in
all possible combination by hand. Equal amount of seed from each cross is mixed to raise next
generation. This completes original selection cycle. From this, several desirable plants are
selected and self-pollinated. Progeny rows are grown and inter crosses made. Equal amount of
seeds are composited to raise next generation. This forms the first recurrent selection cycle.
First Year:
i) Several superior plants are selected.
ii) Selected plants selfed.
iii) Harvest the single plants.
iv) Seeds are evaluated, superior plants are identified.
Second Year:
i) Progeny rows raised
ii) Inter crosses are made in all combination by hand.
iii) Equal amount of seed bulked from each cross.
Third Year:
i) Composited seeds raised
ii)Repeat the operation as in first year
Fourth Year:
Repeat as in second year.
i) Recurrent selection is effective in increasing the frequency of desirable genes in the
population
ii) Most suited for characters having high heritability.
iii) Inbreeding is kept at minimum.
Eg: oil content, protein content and high heritability traits are effective for increasing the
frequency of desirable genes in the selected populations

Fig: Simple Recurrent Selection schematic representation

Merits of recurrent selection
i) Recurrent selection is effective in increasing the frequency of desirable genes in the
population ii) Most suited for characters having high heritability
iii) Inbreeding is kept at minimum.

2. Recurrent selection for general combining ability

It is the modification of the early testing suggested by the Jenkins, 1935. In this case the
progenies selected for progeny testing are obtained by crossing the selected plants to a tester
parent with broad genetic base. A tester parent is a common parent mated to a number of lines.
Such a set of crosses is used to estimate the combining ability of the selected lines. A tester
with broad genetic base means an open pollinated variety, a synthetic variety or segregating
generation of a multiple cross. The gametes of the tester would be variable the difference
between population and tester is primarily due to general combining ability (GCA). Recurrent
selection for GCA can be used for two basically different purposes:
1. It may be used to improve the yielding ability and the agronomic characteristics of a
population. In this case the end product will be a synthetic variety.
2. It may be used to concentrate genes for superior GCA. Here the end product will be superior
inbreds. Such inbreds can be developed after a few cycles of RSGCA
The steps involved in recurrent selection for GCA are outlined below:
First year
A number of phenotypically outstanding plants are selected from source population. Each
selected plant is selfed and crossed (as male) to a number of plants selected from tester with
broad genetic base. Tester with broad genetic base implies a population that has a large genetic
variation. e.g. an open pollinated variety/synthetic/segregating population. The selfed seeds are
harvested separately and saved for planting in third year. The test crossed (seed from plant x
tester cross) from each selected plant is harvested separately and used for replicated yield trial
in second year.
Second year
A replicated yield trial is conducted using plant x tester seeds, and plants producing superior
test cross progeny are identified.
Third year
Selfed seeds (from first year) from those plants producing superior test cross progenies are
planted in separate progeny rows in a crossing block. The progenies are crossed in all possible
combinations, and equal amounts of seeds from all intercrosses are composited to obtain the
next generation. This completes the original selection cycle.
Fourth year
The composited seed (from all the inetrcrosses) is planted and used as the source population
for first cycle of recurrent selection. All operations of first year are carried out.
Fifth year
Operations of second year are repeated.
Sixth year
Operations of third year are repeated. This completes the First recurrent selection cycle.
Seventh year
The second recurrent cycle may be initiated, and in this manner, several cycles of selection
may be carried out.

Fig: Schematic representation of Recurrent Selection for GCA

3) Recurrent Selection for Specific Combining Ability
Recurrent selection for SCA was first proposed by Hull in 1945. This is similar to RSGCA
except, that in the case of Tester. Here the tester will be an INBRED instead of open pollinated
variety. The other operations are similar to RSGCA. The objective of RSSCA is to isolate from
population such lines that will combine well with an inbred. These lines are expected to give
best hybrids in heterosis breeding.
4. Reciprocal recurrent selection
Proposed by Comstock (1949), Robinson and Harvey. The objective is to improve two different
populations in their ability to combine well with each other. In this method we can make
selection for both GCA and SCA. Basically, two populations A and B are used. Each serve as
a tester for the other.
First year
1. Several plants selected in population A and B.
2. Selected plants are self-pollinated
3. Selected plant from A is test crossed with plants in B and Vice versa. Seeds from testcross
of each selected plant is harvested separately.
Second year
1. Separate yield trials conducted from test cross progenies of A and B
2. Superior progenies identified
Third year
1. Selfed seed from plants producing superior test cross progenies planted.
2. All possible inter crosses made 3. Seeds harvested and composited
Fourth year
Populations A and B are planted from the composited seed produced in the third year, and
operations of first year are repeated.
Fifth year
Operations of the second year are repeated
Sixth year
Operations of third year are repeated. This completes the first recurrent selection cycle. The
population may be subjected to further selection cycles, if desired, by repeating the operations
of first recurrent selection cycle.
Use of RRS
1. Two populations are developed by this method
2. They may be intermated to produce a superior population with broad genetic base. This is
similar to a varietal cross but in this case the populations have been subjected to selection for
combining ability (GCA and SCA)
3. Inbreds may be developed from populations A and B. These inbreds may be crossed to
produce a single cross or double cross hybrids.
4. Production of synthetic variety
Fig: Schematic representation of Reciprocal Recurrent Selection
2. Hybrids
They are the first generation from crosses between two pure lines, inbreds, open pollinated
varieties of other populations that are genetically not similar. Pure line hybrids: Tomato.
Inbred hybrids: Maize, bajra.
Kinds of hybrids
1. Single cross hybrids (A x B)
Crossing two inbreeds or pure lines.
2. Three-way cross hybrid (A x B) x C
A cross between a single cross hybrid and an inbred.
3. Double cross hybrid (A x B) x (C x D)
cross between two Fls.
4. Double Top Cross hybrid
Double cross hybrid crossed with open pollinated variety.
Operation in production of hybrids:
In production of hybrids inbreds are preferred rather than open pollinated varieties for the
following reasons:
1. Inbreds can be maintained without a change in the genotype. Whereas open pollinated
variety cannot be maintained pure. They may alter genotypically due to natural selection etc.
2. The hybrids derived from inbreds will be uniform where as it may not be in case, of open
pollinated variety.
3. The inbreds are homogenous and their performance can be predicted whereas open
pollinated variety are heterogenous and their prediction in performance cannot be made.
Development of inbreds
1. By inbreeding, selfing etc.
2. Development of inbreds from haploids - rice, sorghum, maize.
Evaluation of inbreds
a) Phenotypic evaluation
Based on phenotypic performance. Highly suitable for characters with high heritability.
b) Top cross test
Top cross test provides a reliable estimate of GCA. The selected inbreds will be crossed to a
tester parent with wide genetic base i.e. open pollinated variety. The cross progenies will be
evaluated in replicated progeny rows. Based on results better inbreds can be selected.
c) Single cross evaluation
The developed inbreds can be crossed and the single crosses can be estimated in replicated
trial. Outstanding hybrids tested over years in different locations, then released.
d) Prediction of double cross performance
'The predicted performance of any double cross is the average performance of the four non
parental single crosses involving the four parental inbreds. Inbreds: A, B, C, D.
6 possible single crosses = A x B, A x C, A x D, B x C, B x D, C x D.
From these 3 double crosses produced = (A x B) x (C x D), A x C) x (B x D), (A x D) x (B x
C)
The performance of these anyone double cross can be predicted from performance of the four
single crosses not involved in producing that particular hybrid.
(A x B) x (C x D) = [(A x C) + (A x D) + (B x C) + (B x D)]/4
Production of Hybrids
Methods
I. Hand emasculation and dusting - Cotton, Tomato, Chillies, Bhendi
2. Use of male sterile lines
a) Cytoplasmic male sterility – ornamentals
b) Genic male sterility - Redgram, Castor.
c) Cytoplasmic - genic male sterility Jowar, Bajra, Rice
3. Use of self in compatibility
By planting cross compatible lines hybrids are produced. Here both are hybrids. E.g. Brassicas.
Success of hybrids
a) Easy hand emasculation
b) Abundant seed set to compensate cost of hand emasculation.
c) Stable male sterile lines.
d) Effective restorers.
e) Effective pollen dispersal.
3. Synthetic Varieties
A synthetic variety is produced by crossing in all combinations a number of inbreds (4-6) that
combine well with each other. The inbreds are tested for their GCA. Once synthesised, a
synthetic is maintained by open pollination. The lines that make up a synthetic may be usually
inbred line but open pollinated variety, or other population tested for general combining ability
are also be used.
Synthetic varieties are common in grasses, clover, maize and sugar beets. The normal
procedure is equal amounts of seeds from parental lines (Syn 0) is mixed and planted in
isolation. Open pollination is allowed. The progeny obtained is Syn 1. This is distributed as
synthetic variety or it may be grown in isolation for one more season and Syn 2 is distributed.
Merits
1. Less costly compared to hybrids.
2. Farmer can maintain his synthetic variety for more seasons which is not possible in
hybrids.
3. Because of wider genetic base the synthetics are more stable over years and environments.
4. Seed production is more skilled operation in hybrids where as it is not so in synthetics.
Demerits
1. Performance is little bit lower compared to hybrids because synthetics exploit only
GCA while hybrids exploit both GCA and SCA.
2. The performance may not be good when lines having low GCA are used.
4. Composite varieties
It is produced by mixing seeds of phenotypically outstanding lines and encouraging open
pollination to produce crosses in all possible combinations among mixed lines. The lines used
to produce a composite are rarely tested for combining' ability. So, the yield of composite
varieties cannot be predicted easily. Like synthetics, composites are commercial varieties and
are maintained by open pollination.
 Synthetic variety Composite variety
Parental components are generally inbreds It is not so in composite. The lines are not
tested for their GCA tested for their GCA.
No of parental lines are limited to 4 - 6 No such limit
inbreds
Synthetic produced with inbreds can be It is not possible to reconstitute composite
reconstituted variety
Yield performance can be predicted Yield performance cannot be predicted

Combining ability
Ability of a strain to produce superior progeny when crossed with other strains.
General combining ability (GCA)
Average performance of a strain in a series of cross combinations. The GCA is estimated from
the performance of F1S from the crosses. The tester will have a broad genetic base.
Specific combining ability (SCA)
Deviation in performance of a cross combination from that predicted on the basis of general
combining ability of the parents involved in the cross. The testing will be on inbred.
Breeding methods for vegetatively propagated crops
Some agricultural crops and a large number of horticultural crops are asexually propagated.
Some common asexually propagated crops are sugarcane (S. officinarum), potato (S.
tuberosum), sweet potato (I. batatas), Colocasia (Taro), Arum, Dioscorea (yams), Mentha,
ginger (Zingiber sp.), turmeric (C. domestica), banana (Musa paradisiaca), etc., almost all the
fruit trees, e.g., mango (Mangifera indica), citrus (Citrus spp.), apples (P. malus), pears (P.
communis), peaches (P. persica), litchi (Litchi chinensis), loquat (Eriobotrya japonica), etc:,
and many ornamentals and grasses. Many of these crops show reduced flowering and seed set,
'e.g., sugarcane, potato, sweet potato, banana, etc., and some varieties of these crops do not
flower at all. But many of these crops flower regularly and show satisfactory seed set.
However, they are propagated asexually to avoid the ill effects of segregation and
recombination, both being the inevitable consequences of sexual reproduction.
Segregation and recombination produce new gene combinations due to which the progeny
differ from their parents in genotype and phenotype. Asexual reproduction, on the other hand,
produces progeny exactly identical to their parents in genotype because the progeny is derived
from vegetative cells through mitosis.
The advantage of asexual reproduction is immediately clear. It preserves the genotype of an
individual indefinitely. It must be noted that this does not depend on the homozygosity of the
genotype of an individual. Any genotype is preserved and. maintained through asexual
reproduction. In contrast self-pollination preserves and maintains only homozygous genotypes
giving rise to purelines.
Characteristics of Asexually Propagated Crops
 A great majority of them are, perennial, e.g., sugarcane, fruit trees, etc. The annual crops are
mostly tuber crops, e.g., potato, cassava (M. utilissima), sweet potato, etc.
 Many of them show reduced flowering-and seed set. Many varieties do not flower at all.
Only the crops grown for fruit, particularly where good fruit set depends upon seed formation,
show regular flowering and satisfactory seed set.
 They are invariably cross-pollinated.
 These crops are highly heterozygous and show severe inbreeding depression.
 A vast majority of asexually propagated crops are either polyploids, eg., sugarcane, potato,
sweet potato, etc., or have polyploid species or varieties.
 Many species are interspecific hybrid, eg., Banana (M. paradisiaca), sugarcane, Rubus, etc.
 These crops consist of a large number of clones, that is, progeny derived from a single plant
through asexual reproduction. Thus, each variety of an asexually propagated crop is a clone.
Clone
A clone is group of plants produced from a single through asexual reproduction. Thus,
asexually propagated crops consist of large number of clones, and they are often known as
clonal crops. All the members of a clone have the same genotype as the parent plant. As a
result, they are identical with each other in genotype. Consequently, the phenotypic differences
within a clone do not have a genetic basis and are purely due to the environmental effects. A
selection within a clone is thus useless. The various characteristics of a clone are summarised
below.
Identical Genotype
All the individuals belongings to a single clone are identical in genotype. This is so because a
clone is obtained through asexual reproduction, which involves mitotic cell division only.
Genetic variation in the progeny of a plant is produced chiefly by segregation and
recombination, which occur during meiosis only. Thus, the genotype of a clone is maintained
indefinitely without any change.
Lack of genetic variation
The phenotypic variation present within a clone is due to the environment only. This is so
because all the individuals belonging to a single clone have the same genotype. The phenotype
of a clone is due to the effects of genotype (G), the environment (E) and the genotype X
'environment interaction (G x E) the population mean (µ). Thus, the phenotype (P) of a clone
may be expressed as follows:
P = µ + G + E + GE
Thus, the phenotypic differences among clones would be partly due to E and GE components.
Hence the efficiency of selection among clones, as among purelines, would depend upon the
precision with which the E and GE components of phenotype are estimated.
Immortality
Theoretically, clones are immortal i.e., a clone can be maintained indefinitely through asexual
reproduction. But clones usually degenerate due to viral or bacterial infection. A clone may
become extinct due to its susceptibility to diseases or insect pests. Further, genetic variation
may arise within a clone changing its characteristics.
Severe Inbreeding Depression
Generally, clones are highly heterozygous and show severe loss in vigor due to inbreeding.
Clonal Selection
The phenotypic value of a plant or clone is due to the effects of its genotype (G), the
environment (E) and genotype x environment (G x E) interaction. Of these, only the G effects
are heritable. The environmental and interaction effects are non-heritable and cannot be
selected for. Therefore, a selection for quantitative characters based on observations on single
plants is highly unreliable. In fact, plants selected in this way may be no better than a random
sample.
Further, a selection for characters like yielding ability, etc. on the basis of unreplicated clonal
plots would often be misleading and unreliable. Therefore, the value of a clone can be reliably
estimated only through replicated yield trials. However, selection for highly heritable
characteristics, such as plant height, days to flowering, color, disease resistance, etc., are easy
and effective even on the basis of individual plants or single plots. Clearly, these situations are
the same as those in the case of sexually reproducing crops.
Selection Procedure
In view of these considerations, in the earlier stages of clonal selection, when selection is based
on single plants or single plots, the emphasis is on the elimination of weak and undesirable
plants or clones. The breeder cannot reasonably hope to identify superior' genotypes at this
stage. In the later stages, when replicated trials are the basis of selection, the emphasis is to
identify and select the superior clones. The various steps involved in clonal selection are briefly
described below and are depicted in Fig:
1. First Year
From a mixed variable population, few hundreds to few thousand desirable plants are selected.
A rigid selection can be done for simply inherited characters with high heritability. Plants with
obvious weaknesses are eliminated.
2. Second Year
Clones from the selected plants are grown separately, generally, without replication This is
done in view of the limited supply of the propagating material for each clone, and because of
the large number of clones involved. The characteristics of clones will be clearer now than in
the previous generation when the observations were based on individual plants. The number of
clones is drastically reduced and inferior clones eliminated. The selection is based on visual
observations and on the basis of clonal characteristics. Fifty to one hundred clones may be
selected on the basis of clonal characteristics.
3. Third Year
Replicated preliminary yield trial is conducted. Suitable, checks included for comparison. Few
superior performing clones with desirable characteristics selected for multilocation trials. At
this stage, selection for quality is also done. If necessary, separate disease nurseries may be
planted to evaluate the disease resistance of selected clones.
4. Fourth to Sixth Years
Replicated yield trials are conducted at several locations along with a suitable check. The
yielding ability, quality and disease resistance, etc. of the clones are rigidly evaluated. The best
clone that is superior to the check in one or more characteristics is identified for release as a
new variety.
5. Seventh Year
The superior clone is multiplied released as a new variety.
Merits of Clonal Selection
 It is the only method of selection applicable to clonal crops. It avoids inbreeding depression,
and preserves the gene combinations present in the clones.
 Clonal selection, without any substantial modification, can be combined with hybridization
to generate the variability necessary for selection.
 The selection scheme is useful in maintaining the purity of clones.
Demerits of Clonal Selection
 This selection method utilizes the natural variability already present in the population; it has
not been devised to generate variability.
 Sexual reproduction is a prerequisite for the creation of variability through hybridization
Clonal Hybridization
Clonal crops are generally improved by crossing two or more desirable clones, followed by
selection in the F1 progeny and in the subsequent clonal generations. Once the F1 has been
produced, the breeding procedure is essentially the same as clonal selection. The improvement
through hybridization involves the following three steps:
1. Selection of parents,
2. Production of F1 progeny, and
3. Selection of superior cones.
Hybridization can be used only in such crops, which can reproduce sexually. In case of those
crops where sexual reproduction is lacking, mutagenesis or biotechnological approaches can
be applied.
Selection of Parents
Selection of the parents to be used in hybridization is very important since the value of F1
progeny would depend upon the parents used for producing the F1. Parents are generally
selected on the basis of their known performance both as varieties and as parents in
hybridization programmes. The performance of a strain in hybridization programmes depends
on its prepotency and general combining ability. It would be highly desirable to know the
relative values of CGA and SCA in the crop to be improved. If GCA is more important, a small
number of parents with good should be used in hybridization programmes. On the other hand,
when SCA is more important, a large number of parents should be used to produce a large
number of F1 families in an effort to find some outstanding crosses.
A recent suggestion is to partially inbreed the parents to be used in hybridization programmes.
Clonal crops show severe inbreeding depression, but it is expected that one generation of
selfing or 2-3 generations of sib-mating may not reduce vigour and fertility too severely.
Inbreeding may enable the breeder to identify plants that would have a greater concentration
of desirable alleles. These plants may be more prepotent as parents than the highly
heterozygous clones. The practice is gaining some favour with plant breeders.
Production of F1 progeny
Generally, clonal crops are cross-pollinated and they may show self-incompatibility. The
selected parents may be used to produce single crosses involving two parents or an equivalent
of a polycross involving more than two parents.
Selection among F1 Families
When the breeding value of parents is not known, and the relative contributions of GCA and
SCA is not available, a large number of crosses have to be made in order to ensure that at least
some of the crosses would produce outstanding progeny in F1. This is particularly true in a
species where crop improvement has not been done or has been done at a small scale. In such
cases, it would be cumbersome to evaluate a large number of F1 progeny in detail. To avoid
this, generally small samples of several F1 populations are grown. The general worth of
individual F1 populations is estimated visually. The presence of outstanding individuals in the
F1, populations is also noted, and inferior F1’s are eliminated. Promising F1’s with outstanding
individuals are then grown at a much larger scale for selection. The procedure is designed to
save time, space and labour by planting only small populations of a large number of crosses at
the preliminary stage.
Selection within F1 Families
The selection procedure within F1 populations is essentially the same as that in the case of
clonal selection. The various steps involved in the breeding of clonal crops through
hybridization are briefly described below. From second year onward, these should be read
along with the steps described in clonal selection.
First Year
Clones to be used as parents are grown and crosses are made to produce F1 progeny.
Second Year
Sexual progeny from the cross, i.e., seedlings obtained from seeds, are grown. Undesirable
plants are eliminated. Few hundred to few thousand desirable plants are selected.
Third Year
Clones from the selected individual plants are grown separately. Poor and inferior clones are
eliminated. Up to 200 superior clones may be selected for preliminary yield trial.
Fourth Year
A replicated preliminary yield trial is conducted in which suitable checks are included for
comparison. Few outstanding clones are selected for trials at several locations.
Fifth to seventh year
Replicated yield trials are conducted at several locations. Suitable checks are included for
comparison. One or a few outstanding clones are identified and released as new varieties.
Eighth year
The clones released as varieties are multiplied and distributed among farmers.
MUTATION BREEDING
The term mutation was coined by Hugo De Vries in 1900 for the first time and the word is
derived from the latin word ‘MUTARE’ means to change. Mutation is the sudden heritable
change other than the Mendelian segregation and gene recombination in an organism.
Mutation may be the result of a change in a gene, a change in chromosome that involves several
genes or a change in plasma gene. Mutations produced by changes in the base sequence of
genes are known as gene or point mutations some mutations may be produced by changes in
chromosome structure or even in chromosome number they are termed as chromosomal
mutation. There are three types of mutations based on genetic basis of heritable change:
1. Gene mutations: These are produced by change in the base sequence of genes. The change
may be due to base substitutions, deletion or addition.
2. Chromosomal mutation: These arise due to change in chromosome number that may leads
to polyploidy or aneuploidy or change in chromosome structure that result in deletions
duplication, inversion and translocation.
3. Cytoplasmic or plasma gene mutation: These are due to change in the base sequence of
plasma genes. The plasma genes are present in mitochondria or chloroplast. Here the mutant
character occurs in buds or somatic tissues which are used for propagation in clonal crops.
Classification of mutations:
A. Based on origin, the mutations are classified as spontaneous and induced mutations.
1. Spontaneous mutations: Mutations occur in natural populations at a low rate (10-6) but
different genes may show different mutation rates. Here the different genes show different
mutation rate. For example: in maize R-locus mutates at the frequency of 4.92 x 10-4 i.e. (1 in
20000 population), when as Su locus at 2.4 x 10 -6 (1 in 25 lakhs). The Wx locus considered to
be highly stable. The difference in mutation rate may be due to
a) Genetic back ground i.e. presence of mutator genes
b) Genes themselves
c) Environment
2. Induced mutation: Mutations may be artificially induced by treatment with certain physical
or chemical agents. Available evidence indicates that induced mutation rarely produces new
alleles they produce alleles which are already known to occur spontaneously. Induced
mutations are comparable to spontaneous mutations in their effects and in the variability they
produce. Induced mutation occurs at a relatively higher frequency so that it is practical to work
with them.
B. Based on magnitude of phenotypic effects mutation as classified as
Macro mutations: Oligogenic mutation – Large phenotypic effect and recognizable on
individual plant basis and can be seen easily in M2 generations. e.g. Ancon breed in sheep, pod
maize to cob maize
Micro mutations: Polygenic mutations – Small phenotypic effect which cannot be recognized
on individual plant basis but can be recognize only in a group of plants. Selection should be
done in M3 or later generations.
Characteristic features of mutations
1. Mutations are generally recessive but dominant mutations also occur
2. Mutations are generally harmful to the organism. Most of the mutations have deleterious
effects but small proportion (0.1%) of them are beneficial.
3. Mutations are random i.e. they may occur in any gene. However, some genes show high
mutation rates than the others.
4. Mutations are recurrent
5. Induced mutations commonly show pleiotropy often due to mutation in closely linked genes.
Procedure for irradiation: The plant material may be treated in any of the following source.
1. Seeds, 2. Seedlings, 3. Flowers, 4. Cuttings
1. Seeds: Seeds are used after soaking to get greater frequency of induced mutations than air
dried.
2. Seedlings: At any stage of life cycle can be subjected to radiation but usually seedlings
neither too young nor too old are irradiated due to their convenience in handling in pots
transportation from nursery easily.
3. Flowers: Meiotic cells have been found more sensitive than the mitotic cells and therefore
plants are irradiated in the flowering stage in order to affect the developing gametes.
4. Cuttings: In case of fruit tree when they are propagated by clones – the desirable cuttings are
exposed to irradiation.
Selection of the variety for mutagen treatment
The variety selected for mutagenesis should be the best available in the crop.
Dose of the Mutagen
An optimum dose of the mutagen should be used. An optimum dose is the one which produces
the maximum frequency of mutations and causes the minimum killing. Many workers feel that
a dose close to LD50 should be optimum. LD50 is that dose of a mutagen, which would kill
50% of the treated individuals.
Mutation Breeding for oligogenic traits
The handling procedure described here is based on the selection for a recessive mutant allele
of an oligogene.
1. M1 generation: Several hundred seeds are treated with a mutagen and are space planted. In
general, the number of treated seeds is so adjusted as to give rise to 500 fertile M 1 plants at the
harvest. Care should be taken to avoid outcrossing; this can be achieved either by planting the
M1 population in isolation or by bagging the inflorescence of M 1 plants or even the whole M1
plants. M1 plants will be chimeras for the mutations present in heterozygous state. About 20
to 25 seeds from each M1 spike are harvested separately to raise the M2 progeny rows.
2. M2 generation: About 2,000 progeny rows are grown. Careful and regular observations are
made on the M2 rows. But only distinct mutations are detected in M2 because the observations
are based on single plants. All the plants in M2 rows suspected of containing new mutations
are harvested separately to raise individual plant progenies in M3. if the mutant is distinct, it is
selected for multiplication and testing. However, most of the mutations will be useless for crop
improvement. Only 1-3 per cent of M2 rows may be expected to have beneficial mutations.
3. M3 generation: Progeny rows from individual selected plants are grown in M3. Poor and
inferior mutant rows are eliminated. If the mutant progenies are homogeneous, two or more
M3 progenies containing the same mutation may be bulked. Mutant M 3 rows are harvested in
bulk for a preliminary yield trial in M 4.
4. M4 generation: A preliminary yield trial is conducted with a suitable check, and promising
mutant lines are selected for replicated multilocation trials.
5. M5-M7 generations: Replicated multilocation yield trials are conducted. The out-standing
line may be released as a new variety. The low yielding mutant lines, however, should be
retained for use in hybridization programmes.
Mutation breeding for polygenic traits: Mutagenesis does produce genetic variation in
polygenic traits; this variation is usually as much as 50% of that generated in F2 generation, but
sometimes it may be as much as or even greater than the latter.
1. M1 and M2 generations: M1 and M2 generations are grown in the same way as in the case of
oligogenic traits. In M2 generation, vigorous, fertile and normal looking plants that do not
exhibit a mutant phenotype are selected and their seeds are harvested separately to raise
individual plant progeny rows in M3.
2. M3 generation: Progeny rows from individual selected plants are grown. Careful
observations are made on M3 rows for small deviations in phenotype from the parent variety.
Inferior rows are discarded. Few rows may be homogeneous and would be harvested in bulk.
Selection in done in M3 rows showing segregation; a majority of M3 rows would show
segregation. Intensive and careful evaluation of a large number of M3 progeny rows allows
identification of mutants with altered quantitative traits, e.g., partial or horizontal disease
resistance. Such mutants occur in high frequencies that approach 1% or even high, so that their
isolation becomes quite cost effective.
3. M4 generation: Bulked seed from homogeneous M3 rows may be planted in a preliminary
yield trial with a suitable check; superior progenies are selected for replicated multilocation
yield trials. Individual plant progenies from M 3 are critically observed. Progenies showing
segregation may be subjected to selection only if they are promising. Superior homogeneous
progenies are harvested in bulk for preliminary yield tests in M5.
4. M5-M8 generations: Preliminary yield trials and / or multi-location trials are conducted
depending upon the stage when the progenies become homogeneous. Outstanding progenies
may be released as new varieties.
Applications of Mutation Breeding
Mutation breeding has been used for improving both oligogenic as well as polygenic
characters. Mutagenesis has been used to improve morphological and physiological characters
including yielding ability. Various applications of mutation breeding are:
1. Induction of desirable mutant alleles which may not be available in the germplasm
2. It is useful in improving specific characteristics of a well-adapted high yielding variety.
3. Mutagenesis has been successfully used to improve various quantitative characters including
yield.
4. F1 hybrids from intervarietal crosses may be treated with mutagens in order to increase
genetic variability by inducing mutation and to facilitate recombination of linked genes.
5. Irradiation of interspecific (distant) hybrids has been done to produce translocations.
Advantages:
1. Mutation create inexhaustible variation.
2. When no improvement is possible this method has to be adopted.
Limitations:
1. Frequency of desirable mutations is very low about 0.1 percent. To detect the desirable one
in M2 considerable time, labour & other resources are to be employed.
2. To screen large population, efficient quick and inexpensive selection techniques are needed.
3. Desirable mutations may be associated with undesirable side effects due to other mutations
thus extending the mutation breeding programme.
4. Detection of recessive mutations in polyploids and clones is difficult and larger doses of
mutagen have to be applied and larger populations are to be grown.
Polyploidy Breeding
The somatic chromosome number of any species, whether diploid or polyploidy, is designated
as 2n, and the chromosome number of gametes is denoted as n. An individual carrying the
gametic chromosome number, n, is known as haploid. A monoploid, on the other hand, has
the basic chromosome number, x. In a diploid species, n = x, one x constitutes a genome or
chromosome complement. The different chromosomes of a single genome are distinct from
each other in morphology and or gene content and homology; members of a single genome do
not show a tendency of pairing with each other. Thus, a diploid species has two, a triploid has
3 and a tetraploid has 4 genomes and so on.
In euploids, the chromosome number is an exact multiple of the basic or genomic number.
Euploidy is more commonly known as polyploidy.
When all the genomes present in a polyploidy species are identical, it is known as
autopolyploid and the situation is termed as autopolyploidy.
In the case of allopolyploids, two or more distinct genomes are present.
Euploids may have 3(triploid), 4(tetraploid), 5 (pentaploid), or more genomes making up their
somatic chromosome number.
In case of autopolyploidy, they are known as autotriploid, autotertaploid, autopentaploid, and
so on, while in the case of allopolyploidy they are termed as allotriploid, allotetraploid,
allopentaploid, etc.
Amphidiploid is an allopolyploid that has two copies of each genome present in it and, as a
consequence, behaves as a diploid during meiosis.
A segmental allopolyploid contains two or more genomes, which are identical with each other,
except for some minor differences.
Autopolyploids
Origin and production of doubled chromosome numbers:
1. Spontaneous: chromosome doubling occurs occasionally in somatic tissues and unreduced
gametes are produced in low frequencies.
2. Production of adventitious buds: decapitation in some plants leads to callus development at
the cut ends of the stem. Such a callus has some polyploid cells and some of the shoot buds
regenerated from the callus may be polyploid. In Solanaceae 6-36% of adventitious buds are
tetraploids.
3. Treatment with physical agents: Heat or cold treatment, centrifugation, x -ray or gamma ray
irradiation may produce polyploids. Exposing the plants or ears of maize to a temperature of
38-45oC at the time of the first division of zygote produce 2 -5 % tetraploid progenies.
4. Regeneration in vitro: polyploidy is a common feature of the cells cultured in-vitro.
5. Colchicine treatment: Colchicine treatment is the most effective and the most widely used
treatment for chromosome doubling.
Autopolyploidy
In autopolyploidy, triploidy, tetraploidy and higher levels of ploidy are included.
Morphological and cytological features of auto polyploids:
The general features are summarised below:
1. Polyploids have larger cell size than diploids. Guard cells of stomata are larger and the
number of stomata per unit area is less in polyploids than diploids.
2. Pollen grains of polyploids are generally larger than those of the corresponding diploids.
3. Polyploids are generally slower in growth and later in flowering.
4. Polyploids usually have larger and thicker leaves, and larger flowers and fruits which are
usually less in number than in diploids.
5. Polyploids generally show reduced fertility due to irregularities during meiosis and due to
genotypic imbalance leading to physiological disturbances.’
6. In many cases autopolyploidy leads to increased vigour and vegetative growth.
7. Different species have different levels of optimum ploidy. For sugar beet the optimum level
is 3x, sweet potato 6x while for timothy grass it is between 8 -10x.
8. Autopolyploids generally have a lower dry matter content than diploids.
Application of autopolyploidy in crop improvement
Triploids
Triploids are produced by hybridization between tetraploid and diploid strains. They are
generally highly sterile, except in a few cases. This feature is useful in the production of
seedless watermelons. In certain species, they may be more vigorous than the normal diploids,
e.g., in sugar beets. These two examples are described in some detail. Seedless watermelons
are produced by crossing tetraploid (4x, used as female) and diploid (2x, used as male) lines,
since the reciprocal cross (2x x 4x) is not successful. The triploid plants do not produce true
seeds; almost all the seeds are small, white rudimentary structures like cucumber (Cucumis
sativus) seeds. But few normal size seeds may occur which are generally empty. For good seed
setting pollination is essential. For this purpose, diploid lines are planted in the ratio 1 diploid:
5 triploid plants. There are several problems viz. genetic instability of 4x lines, irregular fruit
shape, a tendency towards hollowness of fruits, production of empty seeds and the labour
involved in trioploid seed production.
1. Triploid sugar beets: Among root crops triploid sugar beets apparently represent the
optimum level of polyploidy because 3n plants have longer roots than diploid and also yield
more sugar per unit area.
2. Tetraploid rye: The advantage of tetraploid over its diploid counterpart are large kernel size,
superior ability to emerge under adverse condition and higher protein content. Tetraploid rye
varieties have been released for cultivation. e.g. Double steel, Tetra petkus.
Limitations of autopolyploidy:
1. Larger size autopolyploid generally contain more water and produce less dry matter content
than diploids
2. High sterility with poor seed setting is observed
3. Due to complex segregation, progress through selection is slow
4. Monoploids and triploids cannot be maintained except through clonal propagation
5. The varieties cannot be produced at will
6. Effects of autopolyploidy cannot be predicted.
Allopolyploidy: Allopolyploids have genomes from two or more species production of
allopolyploids has attracted considerable attention; the aim almost always was creation of new
species. Some success has been evident from the emergence of triticale. Raphano brassica
and allopolyploids of forage grasses.
Morphological and cytological features of allopolyploids
1. Allopolyploids combine the morphological and physiological characteristics of the parent
species but it is very difficult to predict the precise combination of characters that would appear
in the new species.
2. Many allopolyploids are apomictic eg: Tulips, Solanum
3. The chromosome pairing in the new species depends upon the similarities between the
chromosomes of the parental species. Chromosomes with such similarities are known as
homoeologous chromosomes. After chromosome doubling, the allopolyploid would have two
homelegous chromosomes for each chromosome present in the F 1 hybrid, comparable to the
diploid species. Such allopolyploid is referred as amphidiploid or Allotetraploid.
4. Fertility of Allopolyploids can be improved by hybridization and selection.
Application of allopolyploidy in crop improvement:
1. Utilization as a Bridging species: Amphidiploids serve as a bridge in transfer of characters
from one species to a related species, generally from a wild species to cultivated species. An
example of use of an amphidiploid as a bridging species in the use of synthetic N. digluta or
transfer of resistance to tobacco mosaic virus from N. glutinosa to N. tabacum. The F1 hybrid
from the cross N. tabacum x N. glutinosa is sterile. Chromosome doubling of the F1 hybrid
produces the synthetic allohexaploid N. digluta which is reasonably fertile. N. digluta is
backcrossed to the recipient species (N. tabacum) to produce a pentaploid having complete
somatic chromosome complement of N. tabacum and one genome of N. glutinosa. The
pentaploid is sufficiently fertile to be backcrossed to N. tabaccum. In the progeny N. tabacum
like plants resistant to tobacco mosaic are selected and cytologically analysed.
2. Creation of new crop species: Triticales, Raphanobrassica
3. Widening the genetic base of existing allopolyploids: The genetic base of some natural
allopolyploids may be narrow, and it may be useful to introduce variability in such cases by
producing the allopolyploids afresh from their parental species. B. napus is a case in point; the
genetic variability of this species is narrow and the only recourse available is to synthesize new
allopolyploid B. napus to widen its genetic base. This is being done by crossing B. campestris
(n=10, AA) with B. oleracea (n=9, CC), the parental diploid species, to produce the
amphidiploid B. napus (n=19, AACC). The two species, B. campestris and B. oleracea, have
to be crossed as autotetraploids; the cross is very difficult and embryo culture has to be used;
somatic hybridization is being used to get around these problems.
Limitations of Allopolyploidy
1. The effects of allopolyploidy cannot be predicted. The allopolyploids have some features
from both the parental species, but these features may be the undesirable ones, e.g.,
Raphanobrassica, or the desirable ones, e.g., Triticale.
2. Newly synthesized allopolyploids have many defects, e.g., low fertility, cytogenetic and
genetic instability, other undesirable features etc.
3. The synthetic allopolyploids have to be improved through extensive breeding at the
polyploidy level. This involves considerable time, labour and other resources.
4. Only a small proportion of allopolyploids are promising; a vast majority of them are
valueless for agricultural purposes.
Wheat Breeding
Wheat - Triticum spp. (x =7)
Wheat is the most important cereal in the world, giving about one-third of the total
production, followed closely by rice. In temperate regions it is the major source of food. The
chief use of wheat is, the flour for making bread. Basic chromosome no x=7, chromosome
number: Diploid: 2n = 14 Tetraploid: 2n = 28 Hexaploid: 2n = 42
Place of origin:
Diploid: Asia minor
Tetraploid: Abyssinia, North Africas
Hexaploid: Central Asia
Classification:
Ploidy level Species Common name Genome
Diploid (2n=14) T. boeticum Wild einkorn AA
T. monococum Einkorn AA
Tetraploid (2n=28) T. dicoccoides Wild Emmer AABB
T. dicoccum Emmer AABB
T. durum Macaroni wheat
T. persicum Persian wheat AABB
T. turgidum Rivet wheat AABB
T. polonicum Polish wheat AABB
T. timopheevi - AABB
Hexaploid (2n= 42) T. aestivum Common or bread wheat AABBDD
T. compactum Club wheat AABBDD
T. sphaerococcum Dwarf wheat AABBDD
T. spelta Spelt wheat AABBDD
T. macha Macha wheat AABBDD
There are fourteen species of wheat according to Vavilov.
Fig:1 Origin of diploid, tetraploid and hexaploid wheat
Related Species of Triticum:
1.T. boeoticum: forms with one to two seeded spikelets occur. The brittle ears shatter at
maturity into individual spikelets armed with awns which provide an effective means of seed
dispersal.
2.T. monococcum: Primitive diploid form domesticated, evolved from T. boeoticum by
mutation and selection.
3. Aegilops speltoides: (2n=14; B genome). It is naturally cross-pollinating. It is the
recognized donor of the B genome.
4.T. dicoccoides: It is an amphidiplod form resulting from the hybridization of T. boeoticum
and Ae. speltoides.
5.T. dicoccum: The spikes are dense, bearded and laterally compressed, the spikelets are two
grained and the grains are retained within the glumes after threshing (speltoid). It is the oldest
of the cultivated wheat.
6.T. durum: Free thrashing wheat with naked grains, important of the tetraploid wheats.
Grains contain high gluten.
7. Ae. squarrosa: (2n=14; D genome) It is the source of D genome in the cultivated hexaploid
wheat, high adaptability.
8. T. spelta: Hexaploid species, considered an amphidiploid from hybridization between T.
dicoccoides and Ae. squarosa.
The most important of all the hexaploid wheat is the common bread wheat, T. aestivum
grown in all parts of the tropics and sub tropics. This hexaploid wheat from which most
modern wheats have been developed. It exhibits an extremely wide range of morphological
and physiological variation and ecological adaptation.
Breeding objectives
1. High yield: High yield depends on
a) The number of heads / unit area
b) The number of grains /head.
c) The average weight of grain
While breeding for high yielding varieties all the above three components must be looked
into. Omitting any one of them may not yield results. Further while breeding for high yield
it is necessary to combine into a variety a favourable combination of genes influencing all
yield process.
2. Breeding non- lodging varieties: This is achieved after the identification of dwarfing gene
in Japanese variety Norin 10. Most of our dwarf wheats are two gene dwarfs. E.g. Sonora
63, Sonora 64, Kalyan Sona. Emphasis is now on triple gene dwarfs.
3. Breeding for disease resistance: Rust is the major disease. Both stem rust and leaf rust are
important ones. There are different races of rust. So, while breeding for rust resistance
horizontal resistance is to be looked into. Back cross method of breeding and development
of multi lines are the methods.
4. Breeding for insect resistance: Hessian fly is the major pest. Resistance in most varieties
is through antibiosis.
5. Breeding for quality. Different wheat varieties vary greatly in their chemical composition
which is considerably influenced by environment. The varieties of hard wheat or bread wheat
which have higher gluten content. The soft wheat contains lesser gluten content which is
suitable for cake making, pastries. The durum wheats are unsuited for either cakes or bread
but they are suitable for making macaroni. So, depending upon the use the quality breeding
objective is to be fixed.
Methods of breeding:
1. Introduction: Semi dwarf wheat introduced from Mexico, Sonora 63, Sonora 64, Mayo
64, Lerma Roja 64
2. Pure line selection: Earlier varieties like P4, P6, P12 evolved at Pusa Institute are result
of pure line selection from local population.
3. Hybridisation and selection
a) Inter varietal: A number of successful derivatives were developed at IARI New Delhi and
Punjab. NP 809 - New Pusa multiple cross derivative. However, all these varieties were
lodging and poor yielder when compared to other countries. Hence the wheat hybridization
programme was changed by Dr. M.S. Swaminathan during 1963. Dr. N. E. Borlaug was
invited to our country and he suggested for introduction of semi dwarf varieties from
Mexico. As a result, four commercial spring wheat varieties viz., Sonara 63, Sonara 64,
Mayo 64 and Lerma Roja 64 were introduced. However, they had red kernel hard wheats.
These were utilised in our breeding programme and amber colour wheat varieties like Kalyan
Sona, Safed Lerma, Sharbati Sonora were released, these are double gene dwarfs.
b) Inter specific crosses: To get resistance against Hessian fly and also for rust resistance.
c) Back cross method of breeding: Rust resistance in Chinese spring wheat was introgressed
from Thatcher.
4. Hybrid wheat: At Kansas Agri. Expt. Station USA, male sterile lines were identified by
crossing T. timophevi x T. aestivum Bison variety
By repeated back crossing, a male sterile line resembling Bison variety was developed. At
present USA and Canada are doing work on this.
5. Mutation breeding: Dr. M. S. Swaminathan did extensive work on this with gamma
rays. Sharbati Sonora with increased protein content was evolved.
6. Development of multiline varieties: Borlaug developed multiline varieties against rust.
MLKS 15 was developed at IARI. Multiline variety is a mixture of pure lines which are
phenotypically similar but genotypically dissimilar. Each line is produced by separate back
cross method of breeding. Each line having resistance against a particular race of a disease.
7. Wheat genetic engineering: Genetic engineering opened up new opportunities for plant
breeders by enabling them to incorporate genes isolated from organisms outside the gene
pools to which they usually have access. Despite huge promise and potential of transgenic
wheat, the approach to transgenic wheat commercialization by private sector multinational
seed companies appears to be cautious.
8. Molecular markers and wheat breeding: Resistance to Fusarium head blight is
governed by QTL Fhb1 and near-isogenic lines carrying a resistant allele at QTL Fhb1 have
been developed using the resistant cultivar “Sumai3,” which showed a 23% reduction in
disease severity. A marker linked to the “Sumai3” allele at Fhb1 showed a large phenotypic
effect on resistance across different populations (Pumphrey et. al., 2007) and was used in
wheat breeding programs across the world (Haberle et al., 2009).
International Programme
International Centre for Maize and Wheat Improvement (CIMMYT) ia an international non-
profit, scientific research and training organization at Mexico and offices at 20 different
locations all over the world operates a global programme on maize, wheat and triticale
improvement, investigates economic issues related to these crops and supports national
programmes in developing countries.
National Programme
All India Coordinated Wheat Improvement Project established in 1965, was upgraded to
Project Directorate in 1978. The headquarter of Directorate of Wheat Research (DWR) was
at IARI, New Delhi till 1991 and afterwards shifted to Karnal, now known as Indian Institute
of Wheat and Barley Research. The project undertakes integrated, multidisciplinary and
multilocational research programmes to increase overall productivity of wheat at national
level. This is sought to be achieved by developing high yielding varieties with crop
production and protection technologies.
Barley Breeding
Barley (Hordeum vulgare L.) is one of the oldest of the cultivated cereals and is widely
grown in many climates of the world. Barley ranks fourth among the cereals worldwide in
the production of grain, after wheat, corn, and rice. The major production areas are northern
and western Europe and the USSR, where seasons are too short or too dry to grow feed
grains such as corn or sorghum. Barley is tolerant of alkali, drought, or frost. Barley grows
in the arid climates of the Sahara, the high plateaus of Tibet, and the tropical plains of India,
but it produces best where fertility is favourable and the spring seasons are long and cool. It
does not mature well in hot humid weather. Barley has developed a diversity of head and
seed types, disease resistance, and quality characteristics. Many improvements have been
made in the cultivated varieties of barley by breeding. Barley is mostly used for feed and
malt industry as it can not be used for bread making because of low gluten content.
Origin
The Near East is considered as the origin of barley. Barley together with emmer wheat was
the first cereal to be domesticated in the Middle East, at least 9000 years ago. The first
archeological material of barley was two rowed barley which is closely resemble with some
races of wild barley, i.e. Hordeum spontaneum. This wild species crosses readily with
cultivated barley and is the progenitor species of H. vulgare (Helback, 1966; Harlan, 1975).
Taxonomy
The genus Hordeum contains 28 species with a basic chromosome number of x=7.
Depending upon the level of ploidy, the species are grouped as diploid (2n=2x=14),
tetraploid (2n=2x=28) and hexaploid (2n=2x=42).
The diploid species of barley:
Cultivated species: H. vulgare
Wild species: H. chilense, H. comosum, H. muticum and H. pusillum
Tetraploid barley species: H. capense and H. secalinum (all wild spp.)
Hexaploid species: H. arizonicum and H. lechleri (all wild spp.)
A species named as H. bulbosum exists both as diploid and polyploid and used to produce
haploid barley embryos.
H. vulgare is only cultivated species with two phenotypic forms, viz. two rowed and six
rowed barley based on ear morphology. Earlier these two forms were grouped into two
separate species but now these has been grouped into single species H. vulgare because these
two have same chromosome number, intercross freely and produce fertile hybrids.
Cytology
There are seven pairs of chromosomes in cultivated barley whereas in wild species,
chromosome no is 7, 14 and 21 pairs. Chromosomes are relatively large, easily distinguished
at metaphase of mitosis. Chromosome 1 is the longest and chromosome 5 is the shorted
chromosome, chromosome 6 and 7 are satellite chromosomes. Chromosome 6 has larger
satellite and 7 has shorter satellite. The chromosomes are median to submedian (Nilan,
1974).
Kasha and Kao (1970) reported the use of wild cross-pollinated barley species H. bulbosum
for producing haploids in common barley. The crosses are made between H. vulgare and H.
bulbosum and shortly after fertilization the embryo is removed ans cultured on artificial
medium in a test tube. The chromosomes of wild species are selectively eliminated, resulting
thereby in a haploid embryo which is sterile unless diploid chromosome number is restored
by treating the haploid seedlings with a 0.1 percent colchicine solution for five hours in the
two to three leaf stage.
Three complete sets of primary trisomics (2n+1) have been developed in barley (Nilan,
1974). These trisomics offer one of the most effective methods of associating genes with
their respective chromosomes.
Breeding objectives
 Yield improvement: To enhance the production, breeding for high yield is the first
and foremost objective of barley breeders.
 Increased adaptability: Barley does not perform well in hot and humid weather. So,
to increase adaptability, barley varieties are to be developed for hot and humid weather
conditions.
 Resistance to yellow rust, aphid and nematode: Breeding for biotic stresses put major
focus on developing varieties resistant to major diseases and pests of barley. Due to various
races of yellow rust, rust resistant varieties should be developed to uncreas the production.
Corn leaf aphid is the major pest of barley and resulted in yield reduction upto 25 per cent.
Due to the development of hull less varieties of barley, this problem is expected to aggravate
further s hull less varieties are damaged more as compared to hulled varieties.
 Improvement in nutritional quality: Developing varieties with high lysine and crude
protein content is one of the breeding objective of barley breeders. The correlations between
crude protein-grain yield and lysine- grain yield were negative.
 Improvement in traits related to malt industry: Barley is preferred gran for malt
production due to tightly adhering lemma and pericarp which protects coleoptile during
malting process which prevents damage to tender shoot, hulls form natural filter and aid in
separating soluble materials from undissolved particles in the production of wort, due to firm
texture grains can be handled with less damage at high moisture level and conversion of
starch to dextrins and fermentable sugars is more efficient due to the combined presence of
alpha and beta amylases.
Methods of breeding
1. Introduction:
Direct Introduction: Clipper
Secondary Introduction (Selection in other collections): Dolma, Sonu (HBL 87), LSB2
2. Pure line selection in local material: Earlier varieties like NP13, NP21, NP100, T4, T5,
K12, K14, C50, C84, CN292, CN294 are result of pure line selection from local population.
3. Hybridisation and selection
a) Inter varietal: i.) A number of successful derivatives were developed by intercrossing
within indigenous germplasm (tall types). BR32, DL157, K18, K19, K24, K70, K71, K125,
K141, K169, K225
ii.) Tall type varieties developed through intercrossing between indigenous and exotic
germplasm include BG25, BG105, BG108, C144, C155, C164, DL100, DL260, DL349,
DL353, NP103, NP104, NP105, NP106, NP 109
iii.) Semidwarf varieties developed by crossing indigenous and exotic germplasm. e.g.
DL70, DL85, DL88, DL171, P100, P103, P267, RD118
iv.) Semidwarf varieties were developed by crossing within exotic germplasm. e.g. BH87
b) Back cross method: Backcrossing had historical significance as its first application was
in breeding of smooth awned varieties of barley.
5. Mutation breeding: Brley responds favourably to mutagenic treatments due to its diploid
nature and low chromosome number. The old classical experiment of Stadler on induction
of mutation was on barley. Use of mutations in barley is now limited to resistance to diseases
and simply inherited traits like earliness and dwarfness etc. Barley varieties developed
through mutation breeding included PL56, DL253and RDB1
6. Molecular markers and barley breeding: In barley, MAS has been employed for the
management of stripe rust caused by Puccinia striiformis Westend. f. sp. hordei, an
important disease of barley worldwide. Several qualitative and quantitative genes conferring
resistant against barley stripe rust have been reported by many workers. Three QTLs (QTL4,
QTL5, and QTL7) were identified on chromosomes 4, 5, and 7, respectively. Castro et al.
(2003) pyramided these three QTLs and studied their effect on resistance against the disease
at the seedling stage. The parents used in the study were Orca, Harrington, and D1-72. Orca,
a barley cultivar derived from the cross between Calicuchima-sib and Bowman, was the
source of QTL4 and QTL7. Harrington is a two-row malting barley cultivar. D1-72 is a line
derived from the cross between Shyri and Galena population that has a resistance allele at
QTL5 tracing to Shyri. MAS was performed for resistance alleles at QTL4 and QTL7 in the
BC1 generation (HarringtonOrca). Four BC 1 plants with Orca alleles at marker loci flanking
QTL4 and QTL7 were crossed with D1-72. DH lines were derived from the F 1 plants of
these crosses. Phenotyping of the parents and the DH lines was done using three races of the
fungus: PSH-13, PSH-14, and PSH-71. Genotypic screening for the polymorphism was done
using 14 SSR markers. The absence of resistance alleles at both QTL4 and QTL5 was
associated with the susceptible phenotype, which validated the effect and location of two
QTLs, QTL4 and QTL5, each tracing from a different parent. However, QTL7 had no effect
on barley stripe resistance at the seedling stage.
International Programme
Barley is mandated crop of the International Centre for Agricultural Research in the Dry
Areas (ICARDA), located at Aleppo, Syria. It was established in 1976 with purpose to
increase and stabilize food production in developing countries of temperate zones having
arid and semi-arid climate.
National Programme
Till mid-sixties, barley improvement programme in India was carried out by individual
states, organizations and the scientists independently with little chance of mutual
cooperation and the evaluation of varieties of one state into other states. In order to rectify
these limitations, ICAR launched All India Coordinated Barley Improvement Project in
1967. It is mandated crop of Indian Institute of Wheat and Barley.
Chickpea breeding
Chickpea (Cicer arietinum L.) is cultivated in almost all parts of the world covering Asia,
Africa, Europe, Australia, North America and South America continents. Chickpea, a
member of Fabaceae, is a self- pollinated true diploid (2 n = 2 x = 16) with genome size of
738 Mbp (Varshney et al. 2013). It is an ancient cool season food legume crop cultivated
by man and has been found in Middle Eastern archaeological sites dated 7500–6800 BC
(Zohary and Hopf, 2000). Its cultivation is mainly concentrated in semiarid environments
(Saxena, 1990). India ranks first in the world’s production and area by contributing around
70.7 % to the world’s total production (FAOSTAT, 2011). It is one of the most important
food legume plants in sustainable agriculture system because of its low production cost,
wider adaptation, ability to fix atmospheric nitrogen and fit in various crop rotations (Singh,
1997) and presence of prolific tap root system. It also helps in enhancing the soil quality for
subsequent cereal crop cultivation by adding organic matter for the maintenance of soil
health and ecosystem.
It is a rich source of quality protein (20–22 %), has the highest nutritional compositions and
free from anti-nutritive components compared to any other dry edible grain legumes, and
thus, it is considered a functional food or nutraceutical. Besides proteins, it is rich in fibre
and minerals (phosphorus, calcium, magnesium, iron and zinc), and its lipid fraction is high
in unsaturated fatty acids (Williams and Singh, 1987). It has no anti-nutritional factors
(Mallikarjuna et al., 2007) and contains higher amounts of carotenoids like β-carotene than
genetically engineered ‘golden rice’ (Abbo et al. 2005). This plant holds a good repute in
‘Ayurvedic’ and ‘Unani’ systems of medicine. In India, acid exudates from the leaves were
used medicinally for aphrodisiac, bronchitis, cholera, constipation, diarrhoea, dysentery,
snakebite, sunstroke and warts. It also has the property to act as hypo-cholesteremic agent;
germinating chickpea is believed to reduce the blood cholesterol level. Sprouted seeds are
eaten as a vegetable or salad. Young leaves and stems and green pods are eaten like
vegetables. Leaves yield an indigo-like dye. The dried seeds may be used in soups or after
grinding as flour. Grain husks, stems and leaves may be used in livestock feed. In the USA
and Europe, chickpeas are marketed dried, canned or in various vegetable mixtures. Mashed
chickpea mixed with oils and spices (hummus) is a popular hors d’oeuvre in the
Mediterranean Middle East.
Origin
Vavilov (1926) supported the idea of Southwest Asia and the Mediterranean region being
the primary centres of origin, with Ethiopia as the secondary centre. van der Maesen (1987)
suggested that Anatolia in Turkey was the area where chickpea was believed to have
originated because three wild species of Cicer closely related to chickpea are found here
namely C. bijugum K. H. Rech, C. echinospermum P. H. Davis, C. reticulatum Lad. C.
reticulatum can be regarded as probable progenitor of chickpea or may be presumed to have
common encestor with chickpea.
Taxonomy
The Cicer genus belongs to family Leguminosae, subfamily Papilionaceae and tribe
Cicereae Alef. The Cicer genus currently comprises 43 species, out of which 9 are annual
and 34 are perennial species (Muehlbauer et al. 1994). Most of these species are found in
West Asia and North Africa, covering Turkey in the north to Ethiopia in the south and
Pakistan in the east to Morocco in the west. Of the 9 annual Cicer species, C. arietinum is
the only cultivated species. The eight other annual Cicer species are C. reticulatum, C.
echinospermum, C. pinnatifidum, C. judaicum, C. bijugum, C. cuneatum, C. chorassanicum
and C. yamashitae.
The wild annual progenitor of chickpea has been identified as C. reticulatum L. (Ladizinsky
and Adler, 1976), and the perennial progenitor is proposed as C. anatolicum (Tayyar and
Waines, 1996). The Cicer species, including cultivated and wild, have been classified into
four sections based on their geographical distribution, life cycle and morphological
characteristics (van der Maesen, 1987).
Monocicer section: The 8 annual species, namely, C. arietinum, C. reticulatum, C.
echinospermum, C. pinnatifi dum, C. bijugum, C. judaicum, C. yamashitae and C. cuneatum,
were grouped in this section. These species are annual having small flowers with firm, erect
to prostrate stems.
Chamaecicer section: This section includes annual or perennial shrubby species with thin
creeping branches. e.g. C. chorassanicum and C. incisum (perennial species) in,
Polycicer section: 23 perennial, rather large flowered species with imperripinnate leavesor
rachis ended in tendril and
Acanthocicer section: The remaining 7 woody perennial species with large floers and
persistent spiny leaf rachis and spiny calyx teeth.
Based on seed size and colour, cultivated chickpea are of two types (Cubero, 1987):
 Macrosperma (kabuli type): Large seeded (100 seed weight >25g), round or ramhead,
cream coloured. The plant is medium to tall in height, with large leaflets and white
flowers, and contain no anthocyanin.
 Microsperma (desi type): Seeds of this type are small and angular in shape. Seed colour
varies from cream, black, brown, yellow to green. There ate 2-3 ovules per pod but on an
average 1-2 seeds per pod produced. The Plants are short with small leafletsand purplish
flowers and contain anthocyanin.
Two types of chickpea cultivars are recognized globally – kabuli and desi. The kabuli types
are generally grown in the Mediterranean region including Southern Europe, Western Asia
and Northern Africa, and the desi types are grown mainly in Ethiopia and Indian
subcontinent.
Cytology
Chickpea has 2n=16. All Cicer species except C. pungens, C. montbretii, C. songaricum and
C. anatolicum have 2n=16. The diploid chromosome number was reported to be 14 in C.
pungens, both 14 and 24 in C. montbretii, and 14 and 16 in C. songaricum and C. anatolicum.
Gupta and Bahl (1983) reported that chickpea has a pair of very long chromosome, distinctly
satellite and sub metacentric. Six pairs of metacentric and submetacentric chromosomes and
a pair of very short metacentric chromosomes.

Genepool
van der Maesen et al. (2007) proposed recent classification in which primary gene pool
consists of cultivated species and landraces. The secondary gene pool consists of the
progenitor species, C. reticulatum and C. echinospermum, the species that are crossable with
C. arietinum but with reduced fertility of the resulting hybrids and progenies; nevertheless,
both are cross-compatible with the cultigen and do not need in vitro interventions to produce
hybrids. The tertiary gene pool consists of all the annual and perennial Cicer species that
are not crossable with cultivated species.
Breeding objectives
Increased seed yield: Developing high yielding varieties is the main objective of crop
improvement programme. The yield potential of chickpea is almost static for the last decade.
To increase productivity varieties with higher yield potential are required to be developed.
Increase biomass, tall, erect and compact cultivars: Increased biomass is resulted in
increased yield due to more photosynthesis. Similarly, with the development of compact and
erect varieties, planting density can be increased and such varieties are also suitable for
mechanical harvesting.
Resistance to diseases: Fusarium wilt is the most devastating disease of chickpea and yield
losses upto 90% has been reported in susceptible cultivars. So, developing varieties resistant
to Fusarium wilt is major objective of chickpea breeding programme.
Another disease Ascochyta blight is major disease of chickpea prevalent if north west India.
Developing Ascochyta blight resistant varieties to enhance the productivity is one of the
objectives of chickpea improvement programmes. Botrytis grey mold ia an important foliar
disease of chickpea and three types of root rot diseases affect chickpea in dry areas.
Resistance to insects: Gram pod borer is the major insect pest of chickpea globally due to
high mobility, polyphagy, high reproductive rate and wide host range. Yield losses due to
this pest ranges from 18% to 24%. Sources of resistance are lacking in cultivated chicpea
germplasm. Resistance sources are available in germplasm belonging to tertiary genepool
i.e. in C. bijugum which can be used provided cross ability barriers overcome.
Breeding for abiotic stress tolerance: Drought and salinity are major abiotic stresses in
chickpea cultivation. developing drought and salinity tolerant varieties is important objective
of chickpea breeding programmes.
Breeding methods: Being highly self-pollinated crop, the breeding method used in
developing improved varieties will be same as used in other self-pollinated crops like wheat,
barley etc.
Commercial cultivars
In the 1970s, most of the commercial varieties were developed through selection from
landraces. The major emphasis was given on increasing yield potential.
During the 1980s, the focus was laid on breeding for disease resistance. Consequently,
several varieties (Avarodhi, JG 315, Pusa 209, GNG 16, Pusa 212, Pusa 240, Pusa 244, Pusa
256, Pusa 413, ICCV 10, ICCV 37, Phule G 5, Phule G 12, etc.) resistant/tolerant to
Fusarium wilt were developed and released for their cultivation in different regions of the
country (Chaturvedi et al. 2003).
Similarly, genotypes tolerant to Ascochyta blight, namely, C235, Gaurav, H 75-35, BG 261,
PBG 1, GNG 146 and PBG 5, were developed for North West Plain Zone (Delhi, Punjab,
Haryana, North Western Rajasthan and Western UP) and G 543 and PBG 5 for the state of
Punjab (Sandhu et al. 2004).
However, during the 1990s, the major thrust was given on breeding for multiple disease
resistance and high-input responsive varieties. Sources for drought tolerance (RSG 44, RSG
963, RSG 888, ICC 4958, ICCV 10, Vijay, GL 769, GPF 2, PDG 3, PDG 4, Phule G 5), cold
tolerance (ICCV 88506, ICCV 88503, Phule G 96006, ICC 8923, PDG 84-10, GL 28008,
GL 28028) and salt tolerance (CSG 88101, CSG8962) were identified for their use in
breeding programme. As a result, multiple disease-resistant varieties, namely, Bharati, Pusa
372, Pusa 362, BG 391, KWR 108 and GNG 1581 against wilt and root rot and GNG 469
against Ascochyta blight and root rot, and high-input responsive variety like DCP 92-3 were
released for cultivation. Rice fallows (about 11.0 m ha) in Eastern India (eastern UP, Bihar,
West Bengal, Orissa, Jharkhand and Assam) and Central India (eastern MP and
Chhattisgarh) provide opportunities for horizontal expansion of area under chickpea.
The development of short-duration varieties like ICCV 2, JG 74, Vijay, JG 11, JG 16, JAKI
9218 and KAK 2 was the major catalyst for the expansion of chickpea area in Southern and
Central India. In spite of reduction in duration, the yield potential of these early maturing
varieties remains almost unaffected, thus improving per day the productivity of the crop.
Presently emphasis has been laid on the development of extra-large seeded kabuli chickpea
varieties (>50 g/100 seed weight).
A major breakthrough has been witnessed in developing large seeded kabuli varieties with
high- yield potential such as KAK 2, BG 1003, BG 1053, JGK 1, Phule G 95311, IPCK
2002-29, IPCK 2004-29, L 555 and HK 05-169 (Chaturvedi et al. 2010). Similarly,
prominent large seeded desi varieties, viz. BG 256, Phule G 5, BGM 391, K 850, Radhey,
Gujarat Gram 2 and L 556, were also developed.
International Programme
Two international crop research institutes viz., ICRISAT (International Crop Research
Institute for Semi-Arid Tropics) at Patancheru, India and ICARDA (International Centre for
Agricultural Research in Dry Areas) at Aleppo, Syria are engaged in research on chickpea
improvement. Desi type chickpea comes under ICRISAT and kabuli type is under both
ICRISAT and ICARDA.
National Programme
All India Coordinated Pulses Improvement Project including chickpea was started in 1967-
68 which was upgraded to Project Directorate in 1977 and to Directorate of Pulsed Research
(DPR), at Kanpur in 1984. Now DPR is known as Indian institute of Pulses Research (IIPR)
w.e.f. 1993. An independent all India coordinated project on chickpea was started in 1993.
Lentil breeding
Among the cool season legume crops, lentil production has displayed the greatest increase,
while cropping of many other grain legumes is actually in decline. This increase is most
likely due to a convenient fast cooking coupled to a saving of fuel and time – dehulled lentils
cook even faster than milled rice. Worldwide lentil consumption has augmented more than
twice the rate of human population growth, with lentil consumption over the past 40 years
having increased more than any other food crop. Therefore, continued genetic improvement
of lentil is essential in all production regions; nevertheless, without access to and use of
diverse germplasm, prospects for yield genetic gain in lentil may be difficult to accomplish.
Wild Lens accessions represent less than 1 % of the world germplasm collection of this
genus, this small fraction likely representing the greatest untapped pool of genetic variation
in lentil. Germplasm evaluation together with a successful and targeted hybridization allows
for an efficient breeding and selection of varieties adapted to specific environments.
Origin
The oldest remains of wild lentils have been found in Mureybit (Syria) dated around 10,000
BP, while those of the cultigens have been dated around 9,000 BP and were discovered in
aceramic Neolithic layers located in the Near East (see Cubero et al. 2009 for archaeological
data). Given the coexistence of the wild and domesticated forms which is not found
elsewhere, coupled with archaeological evidence, the Fertile Crescent is the most likely
candidate to be the center of origin of the cultivated lentil. Ladizinsky (1999) also suggested
the Near East as the center of origin based on the polymorphism found in wild accessions of
ssp. orientalis and the monomorphism of culinaris . Yadun et al. ( 2000 ) suggested a lentil
domestication site close to or overlapping the area where einkorn and emmer wheats had
been domesticated in the Fertile Crescent, concretely, the Karacadag/ Diyarbakir region as
the domestication center of lentil.
Furthermore, two centers of diversity for L. culinaris ssp. orientalis have been described:
(1) Southeast Turkey and Northwest Syria and (2) West and North Jordan and southern Syria
(Ferguson et al. 1998).
Compared to L. orientalis, cultivated lentils have a greater stem and rachis length, more
leaflets per leaf, a greater leaf area, an increased number of flowers per peduncle, as well as
an increased number of pods and seeds. In addition, peduncles of the cultivated forms are
generally shorter or equal in length to the rachis when compared to the wild forms. Aside
from all of the above, pod indehiscence of the cultivated forms in contrast to the dehiscent
mode of the wild species, together with the erect growth habit of the cultivated lentil
compared to the wild procumbent and prostrate plant structure, typifies the most noticeable
differences between the domesticated and wild relatives, being traits of great economic value
and with many implications on yield and ease of harvesting. All of the above mentioned
characters are strongly associated with increased yield levels, in a similar way as observed
for the other domesticated food legumes. Lentil cultigens show a higher frequency of white
flowers compared to the dominant purple-colored inflorescences of the wild relatives, most
probably a character associated to a better culinary quality and fixed by indirect selection of
lighter-colored seed coats (Fratini et al. 2011).
Taxonomy
Taxonomic analyses based on morphological and/or biochemical markers ranged from four
species in 1979, namely, L. culinaris, L. orientalis , L. nigricans, and L. ervoides, to two
species in 1984: L. culinaris (with subspecies culinaris (the cultigen), orientalis , and
odemensis ) and L. nigricans (with ssp. nigricans and ervoides ); again to four species in
1993, i.e., L. culinaris (ssp. culinaris and orientalis), L. odemensis, L. nigricans, and L.
ervoides; and to the previous four species plus the addition of L. tomentosus and L. lamottei
to comprise a total of six species in 1997.
The number of six species is now widely accepted: L. culinaris (ssp. culinaris, the cultigen
and ssp. orientalis (Boiss.) Ponert, the wild ancestor of the cultivated forms),
L. odemensis (Godr.) Ladiz., L. tomentosus Ladiz., L. nigricans (Bieb.) Godr., L. ervoides
(Bring.) Grande., and L. lamottei Czfr. The geographical distribution of the Lens species
which are all self-pollinated is de`1scribed by Cubero et al. (2009).
Cultivated lentil (L. culinaris ssp. culinaris): The cultivars are grouped into two inter
grading clusters- small seeded lentil (microsperma) with small pods and seeds diameter 3-
6mm and large seeded lentils (macrosperma) with large pods and seeds size of 6-9mm.
Wild species: are delicate, small flowered annuals distributed over south west Asia and
Mediterranean basin.
Cytology
All Lens species possess the same chromosome number (2 n=14) and share similar
karyotypes, which consists of three pairs of metacentric and submetacentric chromosomes,
three pairs of acrocentric chromosomes, and a pair of metacentric chromosomes with the
nucleolar organizing region (NOR) secondary constriction site proximal to the centromere
location (Ladizinsky 1993), although some karyotype variants have been described (Balyan
et al., 2002).
Gene pools
Besides of the forms of the cultigen (L. culinaris ssp. culinaris), L. culinaris ssp. orientalis
obviously belongs to the primary gene pool and L. odemensis to the secondary, although
success of L. odemensis crosses with the cultigen may or may not require embryo rescue
depending on the specific accessions used. The tertiary gene pool is composed of L.
tomentosus, L. lamottei, L. nigricans, and L. ervoides, but these can become part of the
secondary gene pool by means of embryo rescue. In any case, further hybridization studies
are needed to truly establish whether L. tomentosus and L. lamottei belong definitely to the
secondary or tertiary gene pool (Cubero et al. 2009).
Breeding Objectives
 High yield: Plant height is strongly associated to yield index. Plants of erect and a very tall
habit tend to lodge as they approach maturity; thus, erect varieties of medium height should
be expected to be more lodging resistant, which results in higher yields and also to a more
amenable mechanical harvesting. Flower number per peduncle in lentil may retain
significance in relation to productivity potential.
 Bold seed size, high protein and less cooking time
Seed size, protein content and cooking time are the most important quality parameters of
lentil. There is negative correlation between yield and protein content and a positive
correlation between seed size and cooking time. Seed size can be used to predict cooking
quality and close relationship between seed size and cooking time eliminates the need for
screening early generation lentil genotypes for cooking time (Erskine et al., 1985).
According to Sharma (2011), in most crops, especially cereals, protein content is invariably
negatively correlated to seed size; however, lentil is possibly an exception and protein
content has been claimed to be positively (although mildly) correlated to seed size.
 Early maturity: Earliness is a desirable trait ensuring completion of the crop cycle in a
relatively short period of time, thereby making more efficient use of resources as well as
avoiding losses due to high temperatures during crop maturation.
 Resistance to diseases: Ascochyta blight, rust and wilt
Ascochyta blight, rust and wilt are the serious diseases of lentil in India. A number of
resistance sources for these diseases have been identified and multiple disease resistant
varieties have been developed. Pant L406 is resistant to wilt and rust.
 Resistance to insects: Pod borer, aphids and cutworms
Pod borers, cut worms and aphids are occasionally serious pest of lentil. A few lines are
reported to be free from pod borer infestation are P202, P248, P262, P772, P803, P927.
 Resistance to shattering
Reduced shattering is a trait of great economic value as pod dehiscence can cause
significant losses before or during harvest; the trait is considered to have played a major
role in lentil domestication.
 Tolerance to drought: Lentil has been traditionally grown in semiarid regions under rainfed
conditions; thus, it combines a high degree of drought resistance and a low water
requirement; in fact, excessive water supply is damaging to the crop.
Breeding methods
Breeding methods used for developing improved varieties of lentil are similar to those of
other self-pollinated crops.
Selection: Breeding history of released varieties showed that majority of these varieties are
selection from germplasm and it is evident that Pureline selection among locally adapted
germplasm lines has been very successful.
Hybridization: Varietal hybridization followed by application of pedigree, bulk methods or
their modifications are now increasingly being used. The trends are in favour of bulk and
SSD methods where segregating populations are handled in bulk till F5/F6 generations and
single plant selections are made in F5/F6 to be grown as individual plant progeny row
followed by further selection among rows. The bulk harvest of selected progeny rows (each
row to be separately bulked) are evaluated in yield trails.
Interspecific hybridization
Artificial cross-pollination in a highly self- pollinated crop species, such as lentil, is
important to increase genetic variability. Wide crosses to yield interspecifi c hybrids allow
for the introgression of important alleles of agricultural interest from wild species to
cultivars, as, for instance, the resistance or tolerance to abiotic and biotic stresses (Erskine
et al. 1994; Ocampo et al. 2000; Davis et al. 2007; Cubero et al. 2009; Pérez de la Vega
et al. 2012). Lentil is generally described as a strictly self-pollinated species holding
cleistogamous flowers (Wilson 1972; Kumar and Singh 1998); nonetheless, recent studies
point out that natural outcrossing rates of lentil can be relatively high (Horneburg 2006).
In the case of interspecific hybridization, crossing efficiencies are much lower than
intraspecific ones, for instance, out of a total of 1,707 pollinations, six interspecific hybrids
with L. odemensis, two with L. nigricans, and one with L. ervoides were recovered using
embryo rescue (Fratini and Ruiz, 2006). Therefore, a large number of manual pollinations
are needed to recover each single interspecific hybrid.
With regard to inter-subspecific hybrids of lentil, it has been reported that the domesticated
lentil is readily crossable with subspecies orientalis (Ladizinsky 1979; Muehlbauer et al.
1989; Vandenberg and Slinkard 1989; Vaillancourt and Slinkard 1992; Fratini et al. 2004),
although the fertility of the hybrids depends on the chromosome arrangement of the wild
parent (Ladizinsky 1979; Ladizinsky et al. 1984).
Interspecific embryos between cultivated lentils and either L. ervoides or L. nigricans abort
(Abbo and Ladizinsky 1991, 1994; van Oss et al. 1997; Fratini and Ruiz 2006) and embryo
rescue techniques are necessary to recover hybrids (Cohen et al. 1984; Ladizinsky et al.
1985; Fratini and Ruiz 2006; Fiala et al. 2009). Nonetheless, gibberellic acid (GA 3)
application after pollination aided to develop viable pods and interspecific Lens hybrids were
obtained without the need of in vitro embryo culture (Ahmad et al. 1995).
Mutation Breeding: Mutation breeding can also be applied in lentil. The n- nitroso-n-methyl
urea (NMU) was almost three time more effective and efficiency was 1.5 to 2 times higher
than gamma rays (Sharma and Sharma, 1979).
Molecular markers
Lentil is a relatively minor crop compared to common bean, pea, and chickpea; as a result,
genomic information regarding lentil is still limited by a relatively large genome size
together with scarce information available on gene sequences, constituting a major obstacle
to undertake genomic studies in lentil. So far for lentil, there exist no descriptions of a
bacterial artificial chromosome (BAC) library, BAC-end sequence, or a physical map (Ford
et al. 2007, 2009; Varshney et al. 2009; Pérez de la Vega et al. 2012). In comparison to
other crop species, the number of Lens data indexed continues to be scarce, although this
situation is rapidly changing, in particular for nucleotide-expressed sequence tags (ESTs)
(Kaur et al. 2011; Bhadauria et al. 2013; Sharpe et al. 2013; Verma et al. 2013).
Lastly, the development of a deep and diverse transcriptome resource for lentil using next-
generation sequencing technology allowed to generate data in multiple-cultivated (L.
culinaris) and wild ( L. ervoides ) genotypes, which together with the use of a bioinformatics
workflow enabled for the identification of a large collection of SNPs and SSR markers for
the subsequent development of a genotyping platform that was used to establish the first
comprehensive genetic map of the L. culinaris genome, comprising seven linkage groups
corresponding to the number of chromosome pairs of lentil (Sharpe et al. 2013).
International Centre
International Centre for Agricultural Research in Dry Areas (ICARDA) is actively engaged
in research on lentils. With a conservation capacity of more than one thousand accessions,
the International Centre for Agricultural Research in the Dry Areas (ICARDA;
http://www.icarda. org), the Indian Agricultural Research Institute (http://www.iari.res.in),
the N I Vavilov Research Institute of Plant Industry ( http://www.vir.nw.ru), the National
Bureau of Plant Genetic Resources, India, and finally the USDA collection at the Regional
Plant Introduction Station, Pullman, Washington (http://www.ars.usda.gov) stand out.
Another national germplasm bank holding an important collection of lentils is the Spanish
Plant Genetic Resource Center (http://www.inia.es) with more than 600 accessions. A
complete picture of the genetic resources available can be obtained at the germplasm
collection directory web page of Bioversity International
(http://www.bioversityinternational.org/), which integrates information regarding most of
the world germplasm collections.
National Programme
All India Coordinated Pulses Improvement Project including lentil was started in 1967-68
which was upgraded to Project Directorate in 1977 and to Directorate of Pulsed Research
(DPR), at Kanpur in 1984. Now DPR is known as Indian institute of Pulses Research (IIPR)
w.e.f. 1993. An all India coordinated project on MULLaRP (mungbean, urdbean, lentil,
lathyrus, rajmash and field pea) was started in 1993.
MUSTARD and RAPE SEED
Brassica spp. (2n = 16, 18, 20, 22, 36, 38 and 48)
Oilseed Brassicas, rapeseed and mustard are the third most important edible oil source in
world after soybean and palm and in India Brassicas rank second after groundnut. Because
of their ability to germinate and grow at low temperature, the oilseed Brassicas can be grown
in cooler areas and at higher elevations. Cultivated Brassicas can be broadly divided in to
two distinct types viz. Vegetable type: Cabbage, Cauliflower, turnip
Oil seed type - Rape seed and mustard.
Origin
Primary centre of origin of B. campestris is near the Himalayan region. The secondary
centres of origin are located in the European -Mediterranean are and Asia (Downey and
Roebbelen, 1989).
Regarding the origin if B. juncea, Prakash and Hinata (1980) reported that species originated
in middle east where B. compestris and B. nigra might have first come into contact.
However, Haminway (1976) argued that B. juncea has probably arisen by hybridization
between different B. compestris and B. nigra genotypes at several different times and
localities resulting in secondary centres of origin in China, north eastern India and Caucasus.
Taxonomy:
Belongs to family Brassicaceae or Cruciferae. The genus Brassica contains more than 3000
species of which 40 are of economic importance. Harberd (1972) examined 85 species of
Brassica and grouped species of the genus into cytodemes. These cytodemes are composed
of different species with the same chromosome number and which are cross fertile and other
having species with different chromosome number and cross infertile. According to him,
most important agricultural species are four diploids, three allopolyploids, each belong to a
separate cytodeme. Four diploids are:
1. B. nigra - Black mustard
2. B. oleracea - Cabbage
3. B. campestris - Rape seed.
4. B. tourne frotii - Wild turnip
Three allopolyploids
1. B. napus - Rape seed of Europe (ghobi sarson)
2. B. juncea - Indian mustard
3. B. carinata – Ethiopian mustard (veg / oil seed)
The genetical relationship between the oilseed brassicas are diagramaticaly represented as
follows:
B. napus will cross readily with B. campestris but with extreme difficulty in case of B.
oleracea.
Rape seed
1.Brassica compestris: Chromosome no 2n=20. Also known as Indian Rape Seed. It is self-
sterile in nature. Important oil seed crop of North India. three cultivated types include:
i.) B. campestris var. Brown sarson
ii.) B. campestris var. Yellow sarson
iii.) B. campestris var. toria
1. B. napus: Chrmosome no. =38. Also known as European Rape Seed. It is self-fertile.
Mustard
1. B. nigra: Chromosome no.= 16. Known as Black mustard and native of Eurasia. 28%
fixed oil. Used as medicine pungent due to glucoside sinigrin.
2. B. alba: Chromosome no. =24. Also known as White mustard. Young seedling used as
salad, yellowish seed contains 30 % oil.
3. B. juncea: Chromosome no.= 36. Known as Indian mustard or Rai, seeds contain 35% oil.
Leaves used as herb contains sinigrin.
Cytology
Cytological analysis of chromosome pairing in the progeny of interspecific crosses has
clealy established that three species with higher chromosome number are amphidiploids
derived from diploid species as follows:
B. oleracea (CC, n=9) x B. campestris (AA, n=10) – B. napus (AACC, n=19)
B. nigra (BB, n=8) x B. campestris (AA, n=10) - B. juncea (AABB, n=18)
B. nigra (BB, n=8) x B. oleracea (CC, n=9) – B. carinata (BBCC, n=17)
This relationship has further been confirmed by artificially synthesizing the amphidiploid
species from their diploid parents. Roebbelen (1960) carried out detailed chromosomal
analysis of diploid species and concluded that only six chromosomes were distinctly
different and remaining being homologous with one or another of the basic six. This
suggested early evolution from a common progenitor species with basic chromosome
number of x=6. The diploid Brassica species with n=8,9,10 chromosome number resulted
from secondary balanced polyploidy.
Breeding objectives
1. Seed yield: Yield is the end product of many biological processes which are under control
of complex polygenic systems. An ideal plant type is having increased branch number,
pods per plant, seeds per pod and seed size. Further yield increase could result from
increase in biomass and harvest index. Increased biomass can result from reduced photo
respiration and increased light saturated rate of photosynthesis.
2. Early maturity: For use in various multiple cropping sequence.
3. Resistance to abiotic factors Frost resistance is needed to prevent yield losses. Winter
hardiness is very important.
4. Resistance to biotic stress: Powdery mildew, Black leg, Sclerotinia rot, alternaria blight
ae major diseases affecting yield. Mustard aphid – major pest and so far no resistance
source has been identified. The variety RH813 and JM1 have been developed for white
rust resistance.
5. Herbicide resistance: Resistance to herbicides viz. Atrazine, simabine etc. A few sources
of resistance are available.
6. Shattering resistance: Major problem in B. napus, highly shattering whereas B. juncea
exhibited tolerance. Introgressive breeding done.
7. Increased oil content and quality: High oil content upto 45% in yellow seed varieties. For
industrial purpose more Erucic acid varieties are required. Development of low erucic
acid cultivars for edible purpose. Reduced linolenic acid content is also desirable. First
“00”hybrid of B. napus (gobhi sarson), Hyola-401 has been developed.
8. Meal quality: Meal having less Glucosinolate content.
9. Breeding for drought tolerance: In India more than 50% of rapeseed and mustard is grown
under residual moisture supply conditions. Therefore, it would be desirable to breed
varieties for such situations.
10.Breeding for frost tolerance: Frost resistant variety RH781 has been commercialised.
In Brassicas the following ideotype is suggested for moisture stress situations:
Roots: Well developed, deep root system with thick long hairs
Seedling growth: Rapid development of hypocotyl and epicotyl
Leaf characters: Rapid build up of leaf area index at early stage, smaller, thicker, hairy,
waxy, erect or moderately droopy leaves with higher photosynthetic and low dark
respiration rate, rapid cell elongation and maintenance of high leaf water potential.
Plant characters: Intermediate plant height, profused primary branches, no tertiary
branches, compact branches, higher siliquae density, high root shoot ratio, ability to
survive internal water stress and to recover growth on hydration, long siliquae, more
 number of seeds/siliquae with comparatively bold seed size, resistant to major diseases
and pests
Flowering: Synchrony in flowering
Duration: Short- medium duration depending upon species and location specific rainfall

Breeding methods
Breeding methods of self-pollinated crops are used in case of B. juncea which is self-
compatible. Pureline selection, pedigree, bulk, SSD, backcross methods and doubled haploid
breeding are used by Brasssica breeders. Breeding methods of cross-pollinated crops are
used in B. campestris (brown sarson, yellow sarson and toria) for improvement. Mass
selection, recurrent selection, synthetic varieties and hybrid varieties are developed.
Hybrid varieties in rapeseed is dependent on cytoplasmic- genetic male sterility where three-
line breeding comes in use.
Production of hybrids using sporophytic self-incompatibility has now been used in vegetable
crops like cabbage, radish and other B. campestris vegetables. However, its application in
oilseed form of self-incompatible B. campestris has not been successful because of high
inbreeding depression by bud selfing and inbreds are difficult to maintain by continuous
selfing.
1. Introduction: Regina variety introduced from Sweden
2. Simple selection: Seeta, Krishna, Kranti varieties were developed through selection.
3. Hybridization and selection
i.) Intervarietal ii.) Interspecific
a) Bulk method b) Pedigree method c) single seed descent
4. Back cross method
5. Population improvement: Recurrent selection, mass selection
6. Heterosis breeding: Heterosis can be exploited using CMS lines.
7. Mutation breeding: RLM 198 variety developed through mutation breeding at PAU
Ludhiana by gamma radiation treatment of mustard cultivar RL18.
8. Tissue culture technique for production of homozygous diploids. First somaclonal variety
Pusa Jai Kisan developed through tissue culture in 1993.
9. Embryo rescue technique for inter specific crosses.
10. Polyploidy breeding: Highly competitive cultivars at least for production of vegetative
matter may be bred from autotetraploids of B. campestris and B. oleracea. Natural
allopolyploids species B. napus, B. juncea and B. carinata can be resynthesized from
 crosses between the parental diploid species. Amphidiploids involving leafy types as one
of the parents, has a large number of primary and secondary branches and more pods.
Similarly, amphidiploids from oleiferous types had more oil.
11. Interspecific somatic hybrids: Interspecific protoplast fusion has been used in Brassica
to transfer disease resistance and other desirable traits from wild to cultivated species.
Blackleg resistance has been transferred from B. tournefortii to B. napus through
protoplast fusion. B. napus and B. oleracea protoplasts have been fused to transfer
resistance to bacterium Xanthomonas campestris pv. campestris from B. napus to B.
oleracea. Resultant somatic hybrids backcrossed with B. oleracea to develop high
resistant progenies.
12. Recombinant DNA technology: In Brassicas, transgenic plants have been produced for
oil quality, seed storage protein, low glucosinolates, herbicide tolerance, fungal disease
resistance, virus resistance, pest resistance and other traits. Conola transgenics have been
commercially released in USA, Canada and Chile.
13. Molecular markers in Brassicas: Different molecular markers have been extensively used
in Brassicas for identification and characterization of transferred chromosomes from
different sources of Brassica species.
National Programme
Directorate of Oilseeds Research (DOR), Hyderabad is national organization with
responsibility to plan, coordinate and execute the research programmes to augment the
production and productivity of oilseeds.
National Research Centre on Rapeseed- mustard (NRC), Bharatpur
NRC has following mandates
 National repository of rapeseed and mustard genetic resources and information
 Basic, strategic and applies research to improve the productivity and quality of oil and meal
 Development of viable agro-production and protection technologies
 To extend technical expertise and consultancies
 Establishment of linkages and promotion of cooperation with national and international
agencies to achieve objectives
Sunflower breeding
Sun flower (Helianthus annuus)
Place of origin: North America.
Classification: The genus comprises nearly 67 species - all native to America. Of these two
are cultivated:
a) H. annuus – diploid, 2n = 34, oil seed crop.
b) H. tuberosus – Hexaploid, 2n = 102, known as Jerusalem artichoke and is cultivated for
tuber.
Wild species: H. hirsutus, H. rigidus moderately resistant to Alternaria.
Putative parent: Weed sunflower gave rise to cultivated one. The weed sunflower was
modified by introgression with H. petiolaris.
Cultivars of sunflower:
a) Giant types: 6 - 14 feet tall, late maturing, large heads 12 - 30” in diameter, seeds large,
white or grey or with black stripes. Oil content is very low. E.g. Mamoth Russian.
b) Semi dwarf varieties: Medium tall - 4 ½ to 6 feet, early maturing, heads 7 - 9” in diameter.
Seeds are smaller, black, grey or striped. High oil content 35%. E.g. Jupiter, Pole star.
c) Dwarf types: 2 to 4½ feet tall, early maturing. Head size 5½ - 6½” in diameter. Small
seeds, high oil content 37%. e.g. Sunrise, Morden, Co1, Co2
Cytology
The genus Helianthus has bacic chromosome number of 17 and includes diploid, tetraploid
and hexploid species. The 17 chromosome number is divided into four groups based on
position of centromere and presence of satellite or absence of satellites. First group has two
pairs of satellite chromosomes with sub-median centromere, the second group has five pairs
of chromosomes with median to sub-median centromere. The third group has six pairs with
sub-median centromeres. The fourth group has four pairs with sub- median to sub- terminal
centromeres.
Breeding objectives
1. To develop short duration varieties suitable for dry land and irrigated conditions: In
dryland, it is successful in black soils only. In red soil under rainfed conditions it is not
successful.
2. Breeding varieties with high oil content: Ranges 38 to 48%. Complex character yield and
oil content are negatively correlated. To increase oil content the shell must be thin.
3. Breeding for self-fertile lines: Protoandry and self incompatability mechanism operates in
sunflower. Hence hand pollination is necessary. To avoid this, self-fertile lines can be
developed.
4. Breeding for disease resistance. Powdery mildew, rust, charcoal rot, Alternaria are major
diseases of sunflower. Wild species like H. hirsuta are moderately resistance to Alternaria.
5. Resistant to pests: Heliothis, Grass hoppers, Jassids are major pests of sunflower.
Breeding Methods:
1. Introduction: Morden variety introduced from Canada.
2. Mass selection: EC 68414 from Russia. Variety Co1 was mass selection from Morden.
Useful for characters which are highly heritable. For e.g. plant height, disease resistance.
3. Hybridization and selection
a) Intervarietal : Variety Co2 was derivative of multiple cross Co4 - (Dwarf x Surya)
b) Interspecific : Wild species of North American origin and best Soviet varieties were
crossed and number of varieties were evolved. E.g. Progress, Novelty, Jubilee 60 which are
resistant to Verticillium wilt also
4. Mutation: Variety Co3 (Mutant from Co2 through gamma rays)
5. Head to row and remnant seed method: Developed by Pustovoit in Russia. By this method
oil content is increased. In this method the following are the steps:
a) From open pollinated type a large no (10,000 to 12,000) plants are selected based on Head
size.
b) The selected lines are analysed for oil content and high oil content lines are isolated (1000
plants).
c) Part of the seed reserved and the part is sown in progeny rows along with check to estimate
yield.
d) Second season testing is also done. The best lines are identified.
e) The remnant seed of elite plants which give high yield were raised in isolation and
multiplied for crossing inter se next season.
f) The multiplied lines also tested for oil content and high yielding high oil content lines
were raised in isolation and crossed inter se.
6. Population improvement: By mass selection, recurrent selection and use of male sterile
lines population can be improved and utilised for breeding.
7. Heterosis breeding: Development of inbred lines and crossing them to harness heterosis
was first done as early as 1920 in Russia. During 1970 cytoplasmic genetic male sterility
was identified in wild types and obsolete cultivars. Now this system is being extensively
used for production of hybrids. First hybrid BSH 1 CMS 234 A x RHA 274 BSH 2 BSH 8.
A number of CGMS lines were bred by Government as well as private seed growers and are
utilised now. Male sterility can also be inducted by GA 100 ppm.
Steps 1. Development of inbreds.
2. Evaluation of inbreds for combining ability.
3. Conversion of inbreds into CGMS lines and R lines.
4. Production of hybrids.
Varietal renovation: In sun flower the varieties released are renovated annually to produce
super elite (Breeder seed) and Elite Seed (Foundation seed).
8. Molecular markers in sunflower breeding: Molecular markers have been successfully used
in sunflower breeding to study the genetic variability of inbred lines, for identification of
interspecific hybrids and MAS to identify disease and pest resistant plants during resistance
breeding programmes.
National Programme
Breeding research for sunflower improvement were initiated in 1972 with establishment of all
India coordinated research programme on sunflower with headquarters at University of
Agricultural Sciences, Bangalore. ICAR launched project to promote research and
development efforts on hybrid sunflower from 1989.
Carrot Breeding
Origin of Carrot:
The genus Daucus has many wild forms that grow mostly in the Mediterranean region and
south-west Asia. Afghanistan is believed to be the primary centre of genetic diversity. There
are evidences that purple carrot together with a yellow variant spread from Afghanistan to the
Mediterranean region as early as the tenth or eleventh century.
Wild carrot, Daucus carota subsp. carota L., (a.k.a. Queen Anne’s Lace). Wild carrot is
abundant in temperate regions across the globe and is widely distributed across much of the
United States where it is often found along roadsides, abandoned fields, and pastures. Wild
carrot is the progenitor of the cultivated carrot, D. carota subsp. sativus, and the two
subspecies are sexually compatible. The cultivated carrot was likely domesticated in Central
Asia roughly 1,100 yrs ago and is grown worldwide from both open pollinated and hybrid seed.
The white and orange carrots are probably mutations of the yellow form. The domestic carrot
readily crosses with widely adapted wild carrot known as Queen Anne’s Lace.
Floral Biology and Pollination of Carrot:
The inflorescence of carrot is a compound umbel. A primary umbel can have over 1000 flowers
at maturity, whereas secondary, tertiary and quaternary umbels bear fewer flowers. Floral
development is centripetal i.e. the flowers to dehisce first are on the outer edges of the outer
umbellets. Carrot is protandrous.
After straightening of filament, the pollen is shed and stamens quickly fall. After this, the petals
open fully and the style elongates. The style is divided into two parts. The petals of petaloid
plants are persistent unlike those of brown-anther, male sterile plants.
Flowers are epigynous. There are five small sepals, five petals, five stamens and two carpels.
Emasculation is laborious and time consuming. As soon as the first bud in an umbel opens, the
whole umbel of the female parent is bagged in a muslin/cloth bag.
The flowers are removed daily until peak flowering has reached. Anthers are removed from the
early opening outer flowers in the outer whorl of umbellets until sufficient flowers are
emasculated.
Unopened central florets in the emasculated umbellets and all late flowering umbellets are
removed. Thus, only the emasculated flowers are left on the female inflorescence inside the
bag. A pollen bearing umbel from previously protected male plant is inserted into the bag of
the female parent along with some house-flies to ensure pollination. Daily for a few days in the
morning, the male umbel is gently rubbed against the emasculated umbel to enhance artificial
cross-pollination.
Sometimes, 1-2 flowering umbels of both the parents are enclosed in the same cloth cage along
with some house-flies. Seed from each parent is sown in adjacent rows. The hybrids and the
parents could be identified (not always) and necessary roguing done to remove the selfed
plants.
Simon has described another alternative commonly used in Europe for intercrossing male
fertile plants. A single isolated umbel will not develop seeds even though pollen is present in
the flowers as this is protanadrous. This umbel can serve as the seed parent in a cross if, one
week after anthesis, the flowers of such an isolated umbel are sprinkled with water to flush out
pollen.
fter it dries, pollen from the intended pollen parent can be introduced with a brush and the seed
parent umbel again placed in isolation. Seeds thus produced are nearly always hybrids.
Sometimes two parents to be crossed are covered by a plastic or cloth screen pollination cage.
Flies are released in the pollination cage to move pollen or pollen is moved by hand or brush.
In this system, selfed and crossed seeds are harvested together. The selfed and crossed
progenies need to be identified by phenotypic or molecular markers or by hybrid vigour when
inbreds are crossed.
Qualitative Genes in Carrot:
As per compilation by Simon (1984) and Peterson and Simon (1986) only 20 genes (Table
22.1) have been described.

Breeding Goals of Carrot:

i. High root yield
ii. Good eating quality
iii. Scarlet/orange colour roots
iv. High carotene content in roots
v. Uniformity in root shape and size
vi Thick flesh roots
vii. Thin and self coloured core in roots
viii. Broad shouldered, cylindrical, uniformly tapering or stump rooted carrot with non-
branching habit
ix. Early rooting
x. Cracking free roots (major gene Ck known for root cracking)
xi. Smooth, shiny root
xii. High sugar and dry matter in roots
xiii. Slow bolting habit
xiv. Heat tolerance
xv. Crown or upper surface (shoulder) free from green colour, flat or slightly up-lifted rather
than concave or shrunken
xvi. Resistance to: Alternaria blight (Alternaria dauci), Cercospora leaf blight (Cercospora
carotae)
Varietal Groups of Carrot:
Simon (2008) have described several varietal groups based on root colour, root length, colour
intensity, core colour and size, root tip shape and days to edible root maturity. Those same
distinctions are still used to help categorize carrot varietal groups even today.
The major root types used today include varietal groups as:
European – Nantes, Chantenay, Danvers, Paris Market, Flakkee, Berlicum, Amsterdam
Forcing
Asian – Kuroda
North American – Imperator
South American – Brasilia
Numerous cultivars have been named for all of these root types. The typical carrot root shapes
are illustrated in Fig. 22.1.
European Nantes types (cylindrical roots, orange in colour) and Japanese Kuroda type (conical
roots, deep orange in colour) are common in India and Asia at large. A dark orange selection
from Gosun by Mr. Kuroda in 1950s resulted in the Kuroda-Gosun cultivar that has become
grown widely, often simply referred to as Kuroda in India. Most of private sector seed
companies are marketing Kuroda carrot on a large scale.
Genetic Resources of Carrot:
As for most vegetables, carrot genetic resources are in the form of open-pollinated cultivars.
Centres on carrot germplasm accessions are as follow:
1. USDA – ARS (http://www(dot)ars = 723 accessions of Daucus carota.
2. European cooperative programme for Crop Genetic Resources – ECP/GR = 5037 accessions
of D. carota
3. National Centre for Vegetable Crops research- Carrot Breeding Collection (CNPH),
Eurpresa Brasileira de Pesquisa Agropecularia-Brazil = 200 accessions.
4. BAZ – Inst, of Horticultural Crops, Germany = 5 species, 25 subspecies, 30 wild relatives.
5. National Gene Bank – Rural Development Administration, Korea = 695 accessions
Selection Criteria of Carrot:
Colour and Quality:
Visual examination of roots, cross section of roots and longitudinal section of roots is effective.
Same colour should extend from crown to down tap root. The colour of xylem, phloem and
vascular cambium should match as far as possible.
Sugar and Flavour:
A thin cross-sectional slice could be cut and tested. The roots with harsh flavour are eliminated.
Total sugars which contribute to sweetness and are an important component of general
preference can be estimated by a refractometer. Selection for high soluble solids and dry matter
is also possible by specific gravity. High dry matter is useful for processors. Percentage dry
weight is easily determined by weighing a fresh sample, drying and reweighing.
Non-bolting:
Bolting may cause serious losses in yield and quality, hence it is important to apply selection
pressure for non-bolting.
Disease Resistance:
Susceptible cultivars are generally planted between rows of breeding lines and the spreader
row plants are inoculated to ensure the spread of disease. This practice is applicable for both
Alternaria leaf blight and Cercospora leaf spot.
Breeding Methods of Carrot:
Open-pollinated Varieties and Synthetics:
In the recent past, mass selection and pedigree selection within the different populations have
been the most important breeding methods. Open-pollinated varieties are adapted to different
situations. Nevertheless, all efforts to breed varieties with high level of uniformity have been
limited by genetic factors originating from breeding methods used for open-pollinated
varieties.
One reason for this is the heterozygosity of the open- pollinated varieties, while another is the
inbreeding depression resulting from Radom selfing of plants within the populations.
Inbreeding depression is considerable as quoted by Stein and Nothnagel (1995) in a review
(Table 22.2).
All open-pollinated varieties suffer from inbreeding depression and a limited degree of
uniformity, and hybrid breeding of carrots has now been started by the carrot breeders
intensively to improve uniformity.
In view of severe inbreeding depression in carrot, selection for resistance to inbreeding
depression is a major consideration in carrot where self-pollination is essential to breeding
progress. In this situation, it is better to develop new inbreds from intercrosses between two
good inbreds.
In case where extreme uniformity seems to be unnecessary, e.g. for juice or pulp production or
for regions with weakly developed agriculture, breeding of synthetics would also be
worthwhile.
As an outcrossing diploid with significant inbreeding depression, adequate population size is
vital to maintain population vigor in development of open-pollinated cultivars.
The development of most open-pollinated carrot cultivars begins with a pre-existing open-
pollinated cultivar and selection is exercised for one or more traits. Population improvement
for carrot typically starts with inter crosses among plants of open-pollinated varieties followed
by phenotypic mass selection for root shape, smoothness, length and desirable root colour in a
particular local production area.
Where facilities are available, selection should also be done for disease resistance (Alternaria
leaf blight), root quality, cavity spot, nematodes, flavour, carotene content and processing
quality. Plants with premature bolting, excessive root cracking and poor plant vigour are always
removed.
Several of the more successful open-pollinated carrot cultivars of North America began with
an intercross between two cultivars, like the breeding of ‘Imperator’. Other examples include
‘Waltham Hicolor’ from ‘Hutchinson’ x ‘Turkey Red Carrot’ and ‘Gold Pak’ from ‘Long
Imperator’ x ‘Nantes’.
Hybrid Breeding:
Thompson (1961) and Hanschke and Gabelman (1963) were the first scientists to detect and
analyze male sterility. The first carrot hybrids were sold in early 1960s in the USA. Today
more than 100 hybrid varieties exist worldwide. The percentage of hybrids is 60-90% in Europe
for early and late varieties and in USA, the value has reached to almost 100%.
Hybrid breeding in carrot is generally based on two systems of cytoplasmic male sterility
(CMS) with different genetic backgrounds and origin. These are brown anther type and the
petaloid type. The brown anther type (ba) is present in all cultivated orange coloured open
pollinated varieties.
The phenotype is characterized by deformed, brown coloured anthers without functional pollen
caused by a genetic block in meiosis. According to O. Banga and his colleagues (1964) the
type ba results from an interaction of the Sa cytoplasm with two independent nuclear genes
(homozygous aa or dominant B).
The two complementary genes E and D operate with their dominant alleles as restorer genes.
Due to this complex inheritance, many test crosses are necessary for the development of a
suitable maintainer.
The cytoplasm of the petaloid type (pt) is derived from a wild form of Daucus carota L. and
has been introduced into many open-pollinated varieties of cultivated carrot. The ‘pt’ type is
characterized by a transformation of the anthers into petals or petal like structures which are
unable to produce functional pollen.
For the inheritance of this type, an interaction between Sp-cytoplasm and two independent
dominant genes (M1; M2) was postulated by T.S. Morelock in 1974. A maintainer for this type
cannot be detected in the F) because of the dominant state of the M genes. Backcrosses should
be performed by use of the male-fertile genotype (Sp) m1m1, m2m2 as a tester in order to find
dominant alleles.
As a matter of fact, petaloid male sterility was discovered in North American wild carrot by
Munger in 1953. It has also been found in other North American wild carrots. Petaloidy is a
homeostatic mutation where a second whorl of petals exists in place of anthers.
Petaloid CMS is the most widely used form of male sterility for production of commercial
carrot hybrids in North America today. It is stable over a wide range of environments
throughout flowering and seed production, although in some genetic backgrounds petaloidy
breaks down and late season umbels can be fertile.
The incorporation of CMS into carrot inbreds for production of hybrid cultivars is similar to
that for other crops. Generally, this process begins at the F2 or F3 generation with the
intercrossing of individual selected fertile plants to a male sterile plant. In the next generation
progeny from the male sterile are examined for male fertility in the process of backcrossing for
sterile line development.
Presence of male fertile plants indicates the presence of restorer alleles from the original fertile
parent. It has been easy to identify and select male sterility maintaining carrot inbreds from a
wide range of germplasm backgrounds but the incidence of restorers varies widely.
A third CMS system has been detected in an alloplasmic form of orange-coloured carrot
originating from a cross between the wild carrot D. carota gummifer Hook. fil. and the
cultivated carrot D C. sativus Foffm by T. Nothangel in 1992.
This type of male sterility called ‘gum’ type, is characterized by a total reduction of anthers
and petals. Recent results on the genetic mechanisms suggest that an interaction of the
‘gummifer’ cytoplasm with a recessive allele (gugu) in the nucleus is responsible for the
expression of this type of male sterility.
The two CMS systems ‘ba’ and ‘pt’ generally suffer from instability of male sterility under
specific conditions. The instability is mainly influenced by high temperature. Nevertheless,
observations over many years have revealed that other factors such as dry conditions, growing
time or long day conditions operate provocatively.
A strict selection scheme is therefore necessary because carrot is partially andromonoecial, i.e.
in umbels of higher order, male flowers can be produced. Umbels of the 5—7th order must be
examined carefully.
The development of the male sterile (ms) and maintainer lines is very laborious process due to
the dominant state of male sterility. Crossing, backcrossing, selfing and testing of the progenies
in the following two generations, including isolation of the positive progenies, are
characteristic steps in breeding processes.
The breeder is forced to eliminate male fertile plants or phenotypes with partial male fertility
within the ms lines developed, and to develop new lines with low inbreeding depression. Such
lines can be found but they rarely have good combining ability.
An excellent uniformity of the hybrid varieties demands a high degree of homozygosity in the
ms and maintainer lines but selection of lines with low inbreeding depression reduces the
probability of finding good parents for hybrid varieties. To overcome this problem, the
development of three-way hybrids has been recommended. In this way, the production of
hybrid seed is realized on ms F1 lines (Fig. 22.1)
For this method, an original maintainer and a second line (exchange maintainer) are necessary.
The universal maintainers which can maintain all ms lines due to complete homozygous state
of the maintainer genes is desirable for such a system.
A universal maintainer must also possess the specific characters of the ms line and a good
combining ability. Exchange maintainers have therefore been propagated which were selected
following a diallel with all lines of one type.
These lines must have the same genetic state as the specific ms line, a state which can be found
relatively frequently. Thus, the best line for performance and uniformity can be selected.
Recently, exchange maintainer lines for the most important ms lines have become available
and can be used for seed production of new lines. Pollinator lines which are necessary for
hybrid seed production are derived from op-varieties or special breeding lines by using a top-
cross system for the testing of general combining ability.

.
Applications of Biotechnology of Carrot:
Genome size is relatively small. Variation in molecular genetic markers is quite extensive, and
both genetic transformation and regeneration of carrot are readily achieved. Therefore, carrot
is a good candidate for biotechnological applications.
The carrot genetic linkage map includes around 1000 isozymes, RFLPs, AFLPs, RAPDs, and
other molecular markers. From 10-25% of genomic clones tested are estimated to be in high
copy numbers, while 20-30% of the RFLP probes and RAPD primers, and 40% of AFLP bands
were polymorphic. The incidence of molecular marker polymorphism and map size to date
makes marker-assisted selection a viable option.
Reports of success have not yet been published but markers linked to nematode resistance,
CMS restorers, carotene content, and sugar content are being sought. One significant limitation
to the application of molecular markers to carrot improvement is the need for radioactive labels,
which cannot to be used in many laboratories. Because of this, efforts are underway to convert
AFLPs to PCR-based co-dominant markers and to develop microsatellite markers.
One of the most intensively studied traits of carrots today continues to be CMS with a particular
emphasis on molecular evaluation. Mitochondrial restriction fragment pattern and protein
product differences between fertile, petaloid, and brown anther cytoplasm’s have been
compared and analysis of variation for a few specific genes, such as atpA have been reported.
Seven monogenic traits have been mapped for carrot. These are yel, cola, Rs, Mj-1, Y, Y 2 and
P1. QTLs have been mapped for carrot total carotenoids and five component carotenoids
namely, phytoene, alpha-carotene, beta-carotene, zeta-carotene and lycopene and majority of
the structural genes of the carotenoid pathway is placed into this map (Just et. al., 2007). MAS
has been reported for two genes: MJ-1 and Rs.
With the pioneering work of F. C. Steward and co-workers in 1964, totipotency of plant cells
was first demonstrated using carrot. Carrot has, in fact, proven to be a model organism for plant
tissue culture, transformation, and regeneration.
Taking advantage of carrot’s facile manipulation in cell culture, carrot- Daucus capillifolius
and carrot-bishop’s weed (Aegopodium podagraria) protoplast fusions were among the first
examples of successfully regenerated somatic hybrids.
Maize transposable elements have been successfully introduced into carrot cell cultures and
found to be mobile. Two interesting experiments with carrot cell cultures demonstrate evidence
for easy regeneration of carrot callus after one year of being air-dried and stored at room
temperature.
Transgenics with genes for disease resistance and enhanced root color have been field-tested.
With more genes of potential agricultural application cloned and improved, public perception
of transgenics, generation, and testing of carrot transgenics will increase.
Now there are reports that carrot transformation is quite efficient. Carrot transformants have
been developed for hepatitis B vaccine and other unique plant products but like most other
vegetables, commercial release of the carrot transgenics has not materialized. However, it is to
be recognized that herbicide and fungal resistance could have lot of advantages in crop
production.
Research and Trends in Carrot Seed Production:
The individual carrot flowers, in common with most other species in Umbelliferae, are borne
on terminal branches in compound umbels. There is a distinct order of flowering which relates
to umbel position. The first umbel to flower is the primary (sometimes referred to as the ‘king’
umbel) which is terminal to the main stalk.
Branches from the main stalk form secondary umbels, and subsequent branches from these
form tertiary umbels. Quaternary branches and umbels may also be formed.
The modern methods of carrot root crop production for the fresh market and processing require
high quality carrots with the minimum of variation between individual roots derived from the
same seed lot. The relationships between seed weight and endosperm characteristics and effect
of seed position on seed quality have been reviewed by George (1999).
The variation in weight of individual seeds and weight of their embryo can contribute to
subsequent variability in seedling size. Seeds from primary umbels are of better quality in terms
of uniformity and producing heavy seedlings. Therefore, it is better to have high plant densities
as there will be less branching and therefore a high proportion of primary to secondary umbels
than at lower plant densities.
Further overall seed quality is improved if manual harvesting is restricted to primary umbels
only so that seeds of shorter maturity span are harvested. This is usually needed to propagate
basic seeds.
For quality seed production in early stages, root-to-seed production is followed where fully
evaluated and selected roots are planted for seed production. Root to seed production is used
for nucleus/breeder/foundation seed production. It is also quite common for commercial carrot
seed production as this method provides an opportunity to the breeder and the seed producer
for selection of roots with desired quality and phenotype.
The production location should be free of wild carrots. Limited size plot can be covered by
pollinating honey-bee hives. There could be geographical isolation by at least a few kilometers
and pollination be accomplished by honeybees or naturally occurring insect pollinators. Seed
to seed production though possible should be avoided.
Cultivar Description of Carrot:
There are open-pollinated and hybrid cultivars. The following outline is based on UPOV (1990)
and as described by George (1999).
1. Leaf:
Length (including petiole): very short, short, medium, long or very long
Division: very fine, fine, medium, coarse or very coarse
Intensity of green colour: light, medium or dark
Anthocyanin coloration of petiole: absent or present
2. Carrot roots:
Length: very short, short, medium, long or very long
Width: narrow, medium or broad
Ratio width/length: very small, small, medium, large or very large
Shape of longitudinal section: circular, obovate, obtriangular or narrowly oblong
Shape of shoulder: flat, flat to rounded, rounded, rounded to conical or conical
Tip: blunt, slightly pointed or pointed
External colour: white, yellow, orange or red
Intensity of external colour: light, medium or dark
Extent of green colour of skin of shoulder: absent or very small, small, medium, large or very
large
Diameter of core relative to total diameter: very small, small, medium, large or very large
Colour of core: white, yellow, orange or red
Intensity of colour of core: light, medium or dark
Colour of cortex: white, yellow, orange or red
Intensity of colour of cortex: light, medium or dark
Colour of core compared with colour of cortex: lighter, same or darker
Green coloration of interior of top (longitudinal section): absent or very weak, weak, medium,
strong or very strong
Time of maturity: very early, early, medium, late or very late
Isolation Distance:
Breeder/foundation seed – 1600 m
Certified seed – 1000 m
Seed Production Methods:
1. Seed – to – seed
2. Root – to – seed
Seed Yield:
1. 600-1000 kg/ha
2. 1000 seed weight – Approximately 0.8 g
3. SMR – 50
Varieties of Carrot:
Pusa Kesar:
This has been bred at Indian Agricultural Research Institute, New Delhi and is an old release
of 1963. It was evolved from a cross of Local Red and Nantes. The roots are scarlet in colour.
It is rich in carotene (38 mg/100 g edible portion). Roots stay longer in field without bolting.
Seed production in north Indian plains is successful.
Pusa Meghali:
This is a tropical or Asiatic type cultivar. It has been developed at Indian Agricultural Research
Institute, New Delhi from a cross between Pusa Kesar and Nantes and was released by the
Institute in 1985.
It has short top, smooth roots with orange flesh and self-coloured core. It is richer in carotene
content (11571 IU/100 g) than Pusa Kesar (7753 IU/100 g) and produces seeds in plains. It is
suitable for early sowing and takes 110-120 days to attain harvest maturity and yields 260-280
q/ha.
Nantes:
This variety belongs to European or temperate types. Its seed production is possible only in the
hilly areas. It is an introduction by IARI Regional Station, Katrain. The roots are half long (12-
15 cm), slim, well-shaped, cylindrical with stumped end forming a small thin tail.
The cortex and core are deep orange. It ranks good in quality, but the top is brittle making
pulling difficult. Keeping quality is poor. It is suitable for cultivation in cooler months. It takes
110-120 days for root formation.
Pusa Yamdagni (Selection 5):
This has been developed at IARI, Regional Station, Katrain from an inter-varietal cross
between EC 9981 and Nantes to combine earliness and size of root of the former and self-
coloured core character of the latter. It has performed well both in the hills and plains. It is
early in maturity. It has been released by the central variety release committee.
Kuroda:
It is an old variety developed in Japan by Mr. Kuroda but is very popular in Indian seed
companies for large scale sales. It has long, sweet, tender orange colour roots with wide
shoulders. Roots are tapering to a blunt point. Roots are smooth, uniform, and conical in shape.
Roots have better storability.
Pea breeding
Peas (Pisum sativum L.) are consumed as fresh vegetable or dry seeds in most of the countries.
In India it is grown as winter vegetable in plains and summer vegetable in the hills.
Origin of Pea
The geographical region comprising of Central Asia, the Near East, Abyssinia, and the
Mediterranean is considered as centre of origin based on genetic diversity. According to Blixt
(1970), the Mediterranean is the primary centre of diversity with secondary centres in Ethiopia
and the Near East.
The genus Pisum comprises of only a small number of taxa. All taxa within Pisum are diploid
(2n = 2x = 14) and the majority are fully inter-crossable with a few being more difficult, but
possible. Pisum humile syn. syriacum is considered a possible candidate as progenitor, as it
resembles closely to the cultivated form. There has been introgression to cultivated types from
P. humile and P. elatius.
The characters associated with domestication are as follows:

Botany of Pea
Pea is an annual herbaceous plant. It has a tap root system. Stems are slender, usually single,
and upright in growth. Leaves are pinnately compound with two to several leaflets. The rachis
terminates in a simple or branched tendril. There are large stipules at the base of leaf.
The plant may be single stemmed or many axillary stems may originate at the cotyledonary
node or any superior node, especially if the apical growing point is destroyed, leaflets of a pair
are opposite or slightly alternate. The lower leaflets are larger than the upper leaflets. The
margins of leaflets and stipules may be entire or serrated.
The inflorescence is raceme arising from the axil of a leaf. The lowest node at which flower
initiation occurs, is normally constant under a given set of conditions and is used in classifying
the varieties into early and late types.
Most early cultivars produce the first flower from nodes 5 to 11 and the late cultivars start
flowering at about nodes 13 to 15. Early cultivars are often single flowered or bear some single
and some double flowers. Late cultivars are usually double/triple flowered.
The flowers are typical papilionaceous with green calyx comprising of five united sepals, five
petals (one standard, two wings and two keels). The stamens are in diadelphous (9+1)
condition. Nine filaments are fused to form a staminal tube, while the tenth is free throughout
its length.
The gynoecium is monocarpellary, with ovules (up to 13) alternately attached to the two
placentas. Style normally bends at right angle to the ovary. Stigma is sticky. Pea is strictly self-
pollinated in nature. Stigma is receptive to pollen from several days prior to anthesis until 1
day or more after the flower wilts.
Pollen is viable from the time anthers dehisce till several days thereafter. For emasculation, the
flower bud chosen should have developed to the stage just before anther dehiscence, indicated
by extension of petals beyond sepals. Flowers can be emasculated at any time.
The first step in emasculation is to tear away with the forceps the tip of the sepal from in front
of the keel. The forefinger is positioned behind the flower and thumb in front and a light
pressure is applied. This spreads the standard and wings to expose the keel. The exposed keel
is slit-open by tips of forceps. Pressure can be applied by the thumb and finger on keel for
increased exposure of the pistil and stamens. The 10 stamens are pulled out.
Pollen can be obtained throughout the day, preferably from a freshly opened flower. For pollen
collection, it is more convenient to pick the male flowers, remove the standard and wings, pull
back the keel so that the style protrudes and use the pollen covered stylar brush as an applicator
to transfer the pollen to the stigma of the emasculated bud.
Older flowers and other flower buds not used in crossing are removed from the peduncle to
increase the pod set after crossing. Normally emasculation is done in afternoon followed by
pollination next forenoon/morning.
Cytology
The somatic chromosome number of peas is 14. The translocations and other chromosome
arrangements are common.The seven characters studied by Mendel have been mapped as
indicated below:
(i) The shape of mature seeds, smooth/wrinkled (R/r), in chromosome 7
(ii) Seed colour, yellow/green (I/i), in chromosome 1
(iii) Flower colour, purple/violet or white (A/a), in chromosome 1
(iv) Mature pods, smooth and expanded/wrinkled and indented (V/v) in chromosome 4
(v) Colour of unripe pods, green/yellow (Gp/gp), in chromosome 5
(vi) Inflorescence, axillary/terminal (Fa/fa), in chromosome 4
(vii) Plant height, tall/dwarf (Le/le) in chromosome 4
Thus, Mendel probably dealt with the genes a and i in chromosome l, le, fa and v in
chromosome 4, gp in chromosome 5, and r in chromosome 7. One may ask as to why he did
not run into the complication of linkage while formulating the law of independent assortment.
The answer is that with respect to a and i in chromosome 1 and fa in relation to le and v in
chromosome 4, these genes are so distant, that linkage is not realized. The only two genes
which could have shown linkage were le and v. As far as is known, Mendel did not study the
simultaneous segregation in these two.
Breeding Goals of Pea
1. High green pod yield
2. Long, attractive green pods with more seeds/pod (9-12 seeds)
3. Sweetness
4. High shelling percentage
5. Specific maturity (early, mid)
6. Suitable for freezing and canning
7. Resistant/tolerant to frost
8. Resistant to diseases, namely:
(i) Downy mildew (Peronospora viciae (Berk.) de Bary)
(ii) Powdery mildew (Erysiphe polygoni DC)
(iii) Rust (Uromyces viciae – fabes (Pers). Schroet, and U. pisi (Pers.) Wint.)
(iv) Wilt (Fusarium oxysporum Schl. f. sp. pisi (van Hall) Synd. & Hans.)
9. Resistance to insects, namely:
(i) Leaf miner
(ii) Aphids
(iii) Pod-borer
(iv) Pea stem fly
Varietal Groups:
The primary characters used for grouping of varieties include traits related to seed and pod
types, maturity groups and plant height.
A useful point of reference list of characters and descriptors used in the UPOV guidelines
is given in Table 15.2:
Genetic Resources of Pea
A large number of ex-situ germplasm collections have been reported around the world in public
domain as compiled by Ambrose (2008) and given in Table 15.3.

These centers have been actively collaborating with each other in PGR management of peas.
In absence of CGIAR funded international agricultural research center with global mandate on
peas, an international consortium for pea genetic resources (Pea GRIC) has been formed that
links key collections in Europe, USA, ICARDA and Australia.
In India, about 2000 pea germplasm accessions are conserved at NBGPGR, New Delhi, IIVR,
Varanasi and IIPR, Kanpur, Besides, a few state agricultural universities rich in vegetable pea
germplasm are G.B. Pant University of Agric. and Technology, Pantnagar, Punjab Agricultural
University, Ludhiana, Haryana Agriculture University, Hisar, JNKVV, Jabalpur and Indian
Agric.Res. Inst., New Delhi.
Important Donors for Pea Breeding Programme
Singh (1991, 1995) has compiled extensive information on genetics and breeding of peas
including listing of superior lines with multiple disease resistance in pulse crops. Kalloo (1993)
and Narsinghani and Tewari (1993) have also given detailed accounts of pea breeding.
A few examples of donors on peas are as follows:
Earliness: Asauji, Lucknow, Bonia, Hans, EC 3
More pods/plant: PL P 26, 50, 69, 179, 279, 496
Long pods: EC 109171, 109176, 109190, 109195
Bold pods: EC 4103, 6185, 95924
Powdery mildew: EC 326, 42959, 109190, 109196, T 10, P 185, P 288, PC 6578, B 4048, P
6587, P 6588, BHU 159, EC 42959, IC 4604, JP 501, VP 7906
Wilt: Early Perfection, Bonneville, PL 43, 124, 6101, Glacier
Rust: PJ 207508, 222117, EC 109188, EC 42959, IC 4604, PJ 207508, JP Batri Brown 3, JP
Batri Brown 4
Pea mosaic: American Wonder, Perfecion Canner’s Gem, Dwarf White Sugar, Little Marvel
Leaf miner: EC 16704, 21711, 25173
Pea stem fly (tolerant): Bonneville, Asaugi, Boach Sel., GC 141, IP 3 (Pant Uphar), Dwarf
Gray Sugar, T 10, T 163
Breeding Procedures of Pea
Peas are self-pollinated due to cleistogamy and accordingly, the common breeding procedures
applicable to self-pollinated crops viz. pedigree, bulk, single seed descent (SSD), back-cross
and mutation breeding are used in pea breeding.
Single seed descent method is now becoming common in peas. This is particularly, useful in
those situations where selected better lines are intercrossed. F1 plants are grown to produce 500
or more F2 seeds.
One seed is harvested from each F2 plant and the harvested seeds are bulked to plant F3. This
procedure continues till F5 in which phenotypically, superior individual plants are selected for
future plant to progeny planting and evaluation. A major advantage of this method is, that, it
can be carried out with less resources and the rapid advancement of generations is possible in
field and glass-house/off season-nursery.
While advancing the generations, selection for highly heritable traits is practiced frequently in
early generations, before lines are grown out as small plots in F4/F5 generation. Shuttle
breeding is also practised in peas where alternate generations (like F3 and F5) are grown at off-
site locations. For example, in India alternate generations can be grown in late kharif in Pune
and Nasik in Maharashtra and followed by winter season in northern plains.
In this way, 2 generations can be grown in a year. There is widespread use of SSD utilizing
glasshouses or plant growth chambers to speed-up early generations while also maintaining a
wider level of variability between lines before growing plant to progeny rows for field
evaluation and selection. Bulk selection is also used by some breeders.
In garden pea number of green pods/plant, green pod weight, pod length and number of
seeds/pod have been shown to be the major yield components affecting the green pod yield.
These yield components usually do not show component compensation effect and therefore,
simultaneous improvement for these characters should be possible.
Prospects of mutation breeding in peas have been discussed by Jaranowski and Micke (1985).
A dose of 10-15 krad of gamma rays is appropriate for seeds. A good criterion of effectiveness
of any mutagen is germination reduction, not exceeding 50% for gamma radiation, better only
30% for neutrons and certain chemical mutagens.
Stronger germination reduction may result for a high number of chromosomal aberrations and
this in turn will lead to high sterility. Among chemicals, EMS, NEU, EI, NMU and sodium
azide seem to be the most efficient mutagens for peas.
Chemical compounds are applied as water solutions and seeds are usually presoaked for 12-16
hours. Presoaking facilitates the penetration of the mutagen into the tissues. The optimum
temperature and duration are 21-24°C and 2-4 hours, respectively.
Recommended concentrations of certain chemical mutagens for pea seeds are:
EMS: 0.05 – 0.3%
NEU: 0.20 – 0.40 millimole
EI: 0.05 – 0.15%
NMH: 0.01 – 0.03%
DES: 0.03%
The solutions should be fresh prepared and buffered to pH 5-6. All the mutagenic compounds
are toxic and some are very carcinogenic and should be handled with extreme care. The
probability of obtaining a favourable variant/mutant (as in other breeding approaches) is
strongly related to the size of the plant population screened.
Treating only a few hundred seeds can hardly be expected to give positive results.
Approximately, 1000 surviving and fertile plants in M ; generation are certainly the minimum,
considering that even the effective mutagen treatments may lead to only one mutation per locus
in 100000 cells.
M1 generation should be grown under optimum conditions. Each M 1 plant should be harvested
individually for growing M2 progenies. If M1 generation is large, a single pod or even a single
seed/M1 plant may be harvested.
Starting with the M2 generation, the optimum methods are very similar to those used for
hybridization programme from F2 onwards. Most reliable is pedigree selection. Bulk handling
followed by pedigree method is also recommended.
However, it should be recognized that in spite of lot of efforts on mutation breeding in peas, in
Sweden, Italy, Germany, Poland and Bulgaria, hardly a few mutations have been released as
commercial cultivars. Thus, it is clear that mutants have provided a significant range of
variation, but that is widely represented in modem day pea breeding, materials.
As a matter of fact, applied mutation breeding is certainly less charming and is on decline,
although one can find numerous mutagenesis programmes in public sector research. In India,
this facility is easily available in Nuclear Agric. and Biotech Division of Bhaba Atomic
Research Centre, Trombay, Mumbai.
Ideotype breeding
A major step in producing a plant model more suited to the crop environment was made by B.
Snoad who introduced the ‘st’ gene for reduced stipule size and the ‘af’ gene, which substitutes
tendrils for leaflets. A detailed review on this has been written by Hedley and Ambrose (1981).
Plants with the genetic constitution ‘afaf stst’ have acquired the descriptive name of ‘leafless’.
The main advantage of the ‘leafless’ pea is its improved standing ability due to its greater
number of tendrils. The risk of lodging is reduced and the crop can be more easily harvested.
The improved canopy structure may also allow the crop to dry more rapidly with a reduced risk
of disease.
Comparisons between near-isogenic leafed and leafless lines grown as spaced pot plants,
however, have shown that the yield per plant of the mutant is reduced relative to that of the
leafed plant.
However, there are also reports where no significant difference in yield per plant between
comparable near-isogenic lines has been observed when grown in randomized plots. More
research is still needed regarding these plant types but it seems that they do not offer yield
advantage as such.
Resistance to diseases
It is convenient to plant spreader rows of a highly susceptible cultivar for field screening against
powdery mildew and rust. Plants can also be inoculated by dusting the powdery mildew spores
from freshly infected leaves. Similarly, for rust, spore suspensions prepared from infected
plants can be sprayed. Screening for wilt will be more effective in the ‘sick’ plots.
Considering the fact that powdery mildew is a serious disease of pea and for which new good
resistant donors are available, a typical backcross breeding approach as applicable to a
character governed by a single recessive gene (powdery mildew in pea is under a single
recessive gene) as outlined by Gritton (1986) has been shown in Fig. 15.4.

Molecular Markers in Pea

Since the development of the polymerase chain reaction (PCR) in generating random amplified
polymorphic DNA in 1990, this technique has been found valuable in the construction of
genetic maps in several species and in production of genetic markers linked to specific
phenotypic traits in particular using bulked segregant pools. RAPD technique became popular
because of its simplicity and ease of use.
Laucou et al. (1998) constructed a genetic linkage map of Pisum sativum L. based primarily
on RAPD markers that were carefully selected for their reproducibility and scored in a
population of 139 recombinant inbred lines (RILs). The mapping population was derived from
a cross between a protein-rich dry-seed cultivar ‘Terese’ and an increased branching mutant (K
586) obtained from the pea cultivar ‘Torsdag’.
The map currently comprises nine linkage groups with two groups comprising only 6 markers
(n = 7 in pea) and covers 1139 cM. This RAPD-based map has been aligned with the map based
on the (J1281 x J1399) RILs population that includes 355 markers in seven linkage groups
covering 1881 cM.
For this alignment 7 RFLPs, 23 RAPD markers, the morphological marker le and the PCR
marker corresponding to the gene Uni were used as common markers and scored in both
populations. Genes for which linked markers have been reported in the pea are listed in Table
15.4.

McClendon et al. (2002) identified DNA markers linked to fusarium wilt race 1 resistance in
pea. Eighty recombinant inbred lines (RILs) from the cross of Green Arrow (resistant) and PI
179449 (susceptible) were developed through single-seed descent, and screened for disease
reaction in race 1 infested field soil and the greenhouse using single-isolate inoculum.
The RILs segregated 38 resistant and 42 susceptible fitting the expected 1: 1 segregation ratio
for a single dominant gene (chi2 = 0.200). Bulk segregant analysis (BSA) was used to screen
64 amplified fragment length polymorphism (AFLP) primer pairs and previously mapped
random amplified’ polymorphic DNA (RAPD) primers to identify candidate markers. Eight
AFLP primer pairs and 15 RAPD primers were used to screen the RIL mapping population and
generate a linkage map.
One AFLP marker, ACG: CAT_222, was within 1.4 cM of the Fw gene. Two other markers,
AFLP marker ACC: CTG₋159 at 2.6 cM linked to the susceptible allele, and RAPD marker
Y15 1050 at 4.6 cM linked to the resistant allele, were also identified. The probability of
correctly identifying resistant lines to fusarium wilt race 1, with DNA marker ACG: CAT_222,
is 96%. These markers will be useful for marker assisted breeding in applied pea breeding
programs.
Marker assisted selection (MAS) is now being integrated in on-going conventional pea
breeding programmes. MAS is particularly, useful to speed-up selection for those traits that
express late in plant development. Such target traits include resistance for diseases and even
lodging and seed characters.
Isozyme marker alcohol dehydrogenase (Adh 1) has been shown to be linked with resistance
to pea enation virus (En.). Two recent examples related to disease resistance are development
of PCR markers designed from cDNA-AFLP fragments providing tight linkage to genes (subm-
1 and mo) conferring resistance to pea seed borne mosaic virus and an SSR marker suitable for
resistance to powdery mildew of peas as mentioned by Ambrose (2008). QTLs for lodging
resistance have been reported.
Integration of Biotechnology in Conventional Pea Breeding:
Transformation and regeneration protocols are now available in peas. The most common
method involves Agrobacterium tumefaciens mediated transformation. The major difficulty
lies in the fact that this transformation is genotype specific and only a small portion of cultivars
have responded to this technique.
Somaclonal variation arising from the regeneration of plants from callus, led to the use of
cotyledonary meristem from freshly imbibed seed as a source of tissue for successful
transformation. The use of this technology in the pea breeding is limited to proof of concept.
Partial resistance to alfalfa mosaic virus (AMV) has been reported as a consequence of
transformation with chimeric virus coat protein gene, a-amylase inhibitor (α-A 1) and the
promoter phytohemagglutin, both found in French-bean when transferred to pea, have shown
constitutive expression and resistance to pea weevil. The expression of inhibitor (α-amylase)
served to block the development of the larvae at an early stage and this resulted in less seed
damage and better seed quality.
This transgenic pea product could not reach to large scale field testing due to legal issues.
Transfer of herbicide resistance both as a reportable marker and a trait have also been reported,
but not carried through to commercial release.
While GM crops are on increase in many parts of world with global acreage of 134 million
hectares in 2009, the adverse reaction to GM crops in Europe and low rates of transfer have all
contributed to the pea breeding industry not engaging in the development and release of GM
peas till date.
Disease Resistance Breeding and Markers:
Pea powdery mildew, caused by Erysiphe pisi, is an air-borne disease with a worldwide
distribution, being particularly important in climates with warm dry days and cool nights.
Although varying levels of resistance to E. pisi have been observed in pea only three genes for
resistance named er1, er2 and Er3 have been described so far. Gene er1 is widely used in pea
breeding programmes and provides complete or incomplete resistance depending on the
locations.
Resistance conferred by this gene has been proved to be stable and is caused by a barrier to the
pathogen penetration. RFLP, RAPD/SCAR and SSR markers linked to the er1 have been
identified. Gene er2 is not used commercially. The gene confers a high level of resistance in
some locations, but is ineffective in others. The expression of er2 is influenced by temperature
and leaf age.
Gene er2 governed resistance is based mainly on post-penetration cell death complemented by
a reduction of percentage penetration success in mature leaves. AFLP, RAPD and SCAR
markers linked to er2 are available. Gene Er3 was recently identified in P. fulvum and has
successfully been introduced into the adapted P. sativum material by sexual crossing.
Resistance conferred by the gene Er3 is due to a high frequency of cell death that occurs both
as a rapid response to attempted infection and a delayed response that follows the colony
establishment. RAPD markers tightly linked to Er3 have been identified and converted into
SCARs.
Pea rust has become an important pathogen of pea particularly in regions with warm, humid
weather. Pea rust has been reported to be caused either by the fungus Uromyces viciac-fabae
(Syn. U. fabae) or U. pisi. U. viciae-fabae is the principal causal agent of pea rust in tropical
and subtropical regions such as India and China, where warm humid weather is suitable for the
appearance of both the uredial and the aecidial stage.
Several sources of incomplete resistance against U. viciae-fabae have been reported. A single
major gene (Ruf) has been reported as responsible for this partial resistance. Two RAPD
markers have been detected flanking the gene Ruf, but both markers were not close enough to
the gene to allow a dependable marker-assisted selection for rust resistance.
Only recently, pea germplasm collections have been screened to identify sources of resistance
to U. pisi both under field and growth chamber conditions. No complete resistance has been
identified so far. However, incomplete resistance has been observed in the collections.
All the identified accessions have displayed a compatible interaction (high infection type) both
in adult plants under field conditions and in seedlings under growth chamber conditions, but
with varying levels of disease reduction.
This resistance was not associated with host cell death. Preliminary results performed on F2:3
revealed two QTLs for resistance to U. pisi in the field and controlled conditions, respectively,
which seems to be the reason for high percentage of the phenotypic variance.
Aschochyta blight, caused by Mycosphaerella pinodes, the teleomorph of Ascochyta pinodes
(Berk & Blox) Jones, is a widespread pea disease. Ninteen QTLs associated with resistance
have been reported. More recently, 6 QTLs (mp1-mp6) have been associated with resistance
to M. pinodes in a cross of the cultivar Messire with P. sativum subsp. syriacum.
Resistance to fusarium wilt in peas caused by Fusarium oxysporum Schlect. f. sp. pisi race 1
(van Hall) Snyd. & Hans, is conferred by a single dominant gene Fw. The gene has been located
in the pea genome by analyzing progenies from crosses involving genetic markers across all
pea linkage groups.
Fw has been shown to be located on linkage group III, about 13 map units from Lap-1 and b
and 14 map units from Td. The relatively large distances between these markers and Fw
precludes the use of the linked markers in marker-assisted selection for wilt resistance.
DNA markers linked to recessive gene sbm-1 for resistance to pea seed-borne mosaic virus
(PSbMV) pathotype P-l have been identified. Markers linked to the dominantly inherited gene
En for resistance to pea enation mosaic virus (PEMV) have also been reported.
Snap Pea
The snap pea is a type of edible-podded pea that is conspecific to field and garden peas (P.
sativum L.). Edible-podded peas lack pod parchment or fibre. Most snap pea cultivars have
wrinkled seeds with green cotyledons, white flowers and short internodes. Snap pea cultivars
released by public and private breeders and seedsmen in 20th century in USA are as follows:
Sugar Stick, Round Podded Sugar, Sugar Snap, Sugar Bon, Sweet Snap, Early Snap, Honey
Pod, String-less Sugar Snap, Sugar Daddy, Sugar Gem. Sugar Pop, Sugar Boys. Mega. Super
Sugar Snap, Crystal. Sugar Lady, Sugar Star and Jessy, etc.
Among edible podded types. Oregon Sugar Podded (Mithi Phali) has been made popular at
PAU, Ludhiana in 1996 through introduction. Pods are light green and devoid of parchment
layer. Its average yield is about 100-110 q/ha and it is tolerant to powdery mildew under field
condition.
Vegetable Pea Varieties:
In vegetable peas, early introductions from Europe and USA were found quite successful and
popular in India. These included Arkel (early maturing, dwarf type, introduction from England
in 1970s) and Bonneville (main season, late maturing, tall type, introduction from USA in
1970s). These introductions were obtained at IARI, New Delhi and were released for
commercial cultivation after preliminary evaluation.
Arkel is still a very popular variety grown throughout the country. Arkel has 45-60 cm plant
height, long well filled, sickle shape green pods with 7-9 seeds/pod. The seeds are sweet and
become wrinkled at maturity. First picking is done in 60 days. It is highly susceptible to
powdery mildew and rust. It has double flowers at lower nodes and single onwards. Shelling
percentage is about 40%. The yield potential is 75-80 q/ha.
VL Ageti Matar (VL-7) is an early maturing variety developed at VPKAS, Almora in 1995.
Plants are dwarf with dark green foliage. Pods are light green, completely filled, two pods in a
bunch with 6-7 seeds/pod and 42% shelling. Its average yield is about 80-90 q/ha. Seeds are
wrinkled.
Kashi Nandini (VRP-5) is an early maturing variety developed at IIVR, Varanasi in 2006
through hybridization (P-1542 x VT-2-1) followed by pedigree selection. Plants are dwarf,
erect and come to flowering in 34 days after sowing. Pods are 8-9 cm long, well filled with 8-
9 seeds having 48% shelling. Its average yield is 110-120 q/ha. Seeds are wrinkled.
Matar Ageta-6 is an extra early maturing variety (45-50 days) developed at PAU, Ludhiana
in 1996 through hybridization (Massey Gem x Harabona) followed by pedigree selection.
Plants are dwarf, erect and bear 12-15 pods with 6 seeds/pod and 45% shelling.
First picking is done in 45-50 days after sowing and about 50% of green pod yield is obtained
in the very first picking. Plants are dwarf (40 cm), erect and green. Dry seeds are light green,
smooth with slight dimples. It is tolerant to high temperature and its yield potential is about 60-
65 q/ha. This variety is suitable for single harvest. Seeds are wrinkled.
Pant Sabji Matar 3 (Arkel x GC 141) developed at Pantnagar is similar to Arkel. however,
the pods are slightly longer and attractive. Pusa Pragati developed at IARI is an early maturing
cultivar with slightly straight green pods having 8-10 seeds/pod. First green pods are harvested
in 60 days. It has become popular due to longer pods and higher yield potential (100 q/ha).
Kashi Udai (VRP 6) is an early maturing cultivar developed through pedigree method of
breeding from the cross of Arkel x FC 1 at IIVR, Varanasi. Plant height is 58-62 cm and 50%
plants bear flowers at 35-37 days after sowing. Plants have dark green foliage and short
internodes with 8-10 pods/plant. Green pods are attractive, 9-10 cm long, filled with 8-9 bold
seeds. Shelling percentage is 48% and green pod yield potential is 100-110 q/ha.
Among main season/late varieties requiring 100 days for first green pod harvest, the popular
cultivars are Bonneville, Perfection New Line, Lincoln, Jawahar Matar-1, Jawahar Matar-4,
Arka Ajeet, Azad Pea 2, Azad Pea 5, etc. These varieties are not very popular and are on decline
from cultivation and the market.
Bonneville is an introduction from USA and made popular by IARI. Plants are medium tall
(60-70 cm) come to flowering in about 60 days having two pods per peduncle. Pods are light
green, straight having 6-7 seeds/ pod with 45% shelling and seeds are green, bold and wrinkled.
Its average yield is 100 q/ha and it is susceptible to powdery mildew. Green pods are harvested
in 100 days.
Arka Ajit (FC-I) is a variety of medium maturity group developed at IIHR, Bangalore in 2006
through multiple cross involving Bonneville, IIHR 209 and Freezer 656. Pods are 8-9 cm long
with green bold seeds having 55%, shelling. It takes 90 days from sowing to first picking. It
has resistance to powdery mildew and rust. Its average yield is about 95-100 q/ha.
Azad P-5 (KS-225) is late maturing variety developed at CSAUAT, Kanpur in 2006. Plant
growth is medium with straight pods full of grains and bearing may be extended up to March.
Its average yield potential is 95-105 q/ha and has resistance to powdery mildew.
Pant Sabji Matar-4 (Arkel x HFP 4) developed at Pantnagar is late maturing and is classified
as leafless type as leaflets are converted into tendrils.
Sugarcane breeding
Sugarcane is a large grass of the genus Saccharum, tribe Andropogoneae, family Poaceae.
Modern sugarcane (Saccharum spp.) cultivars are interspecific hybrids derived from a
hybridization process involving Saccharum officinarum (or “noble cane”) and Saccharum
spontaneum (wild cane), followed by a series of backcrosses to the noble parent (Daniels and
Roach, 1987). The earliest known historical record of sugarcane and sugar is from Indian
writings from 3000 to 3400 years ago. The generic name for sugarcane, Saccharum, originated
from the Indian Sanskrit term “sharkara” for the crude sugary product obtained from the honey
reeds. Dispersal of Indian sugarcane westward seems to have occurred during the first
millennium BC. Soldiers of Alexander the Great are known to have carried it to Europe from
India about 325 BC. Later, Greek and Roman writers were familiar with the concept of the
Indian honey reed and its “honey” (sugar) product. The early history of sugarcane is covered
by a number of authors, including Deer (1949) and Barnes (1964).
Origin
The origin of sugarcane is a complex question that is best discussed in relation to its taxonomy
and distribution in Southeast Asia, the Indonesian Archipelago, and New Guinea. Different
species likely originated in various locations with S. officinarum and Saccharum robustum in
New Guinea, Saccharum barberi in India, and Saccharum sinense in China. Dispersal of S.
officinarum over a period of thousands of years is believed to have occurred both into the
Pacific Ocean area, and along the island chain into Asia, whilst the thinner Indian canes were
developed and cultivated in the North India/South China region. The wild progenitor of S.
officinarum is S. robustum.
Sugarcane technologists consider six species to be important as progenitor species in the origin
of modern commercial hybrids (Saccharum spp.). These six species (listed below) are
distinguishable on the basis of sugar content, thickness of stalk, floral characteristics,
chromosome number and epidermal hairs. The first four in the list below are in cultivation,
while the last two (S. spontaneum and S. robustum) are wild species growing in southern Asia
and New Guinea.
S. officinarum L.: sweet, juicy, thick stalk garden cane, initially in New Guinea
S. barberi Jesw.: sweet, thin stalk Indian canes
S. sinense Roxb.: sweet, thin stalk Chinese canes
S. edule Hassk.: edible inflorescence garden cane, New Guinea, Melanesia
S. spontaneum L.: very thin, hardy wild canes, low sugar, New Guinea and southern Asia
S. robustum Brandes & Jeswiet ex Grassl: tall, harder, thick stalk wild canes, a little juice and
sugar, New Guinea and eastern Indonesia
Cytology
Saccharum spp. having basic chromosome number x= 6,8,10. S. officinarum has diploid
chromosome number of 2n=80 with minor deviation in chromosome number (aneuploids) are
also found. Triploids are rarely found in among selfed progenies of S. officinarum and are
indistinguishable from diploid progenitors. Diploid chromosome number in S. barberi and S.
sinense has been reported to vary in the five morphologically distinct groups as:
i. Sunnabile-2n=82 and 116
ii. Mungo- 2n=82
iii. Nargori- 2n=107 and 124
iv. Suretha- 2n=90 and 92
v. Pansahi-2n= 118
First four groups are considered as S. barberi and the Pansahi group is equivalent to S. sinense.
Bremer (1966) suggested that a subgenus in Saccharum with 2n=68,102 and 136 chromosomes
and having secondary basic chromosome number of x=17.
Breeding objectives
i. High cane yield
ii. Moderate to high sucrose content
iii. Early to full season maturity
iv. Resistance to diseases
A) Red rot
B) Smut
C) Wilt
D) Mosaic
E) Ratoon stunting disease
F) Grassy shoot disease
v. Resistance/tolerance to insect pests
a) Shoot borer
b) Cane borer
c) Pyrilla
d) Mealy bugs
f) Whiteflies
g) Termites
h) White grub
vi. Tolerance to abiotic stresses
a) Drought
b) Salinity
c) High temperature

Breeding methods of sugarcane are based on following consideration:

 The sugarcane plant is complex polyploid and heterozygous
 The flowering occurs under specific environmental conditions
 Self-incompatibility and male sterility are present
 Sugarcane propagated vegetatively by stem cuttings or sets

Breeding methods
Breeding methods used for developing improved varieties of sugarcane as similar to the
methods used in vegetatively propagated crops. Clonal selection and clonal hybridization
methods are used to develop imroved varieties in sugarcane. In hybridization three types of
crosses are made:
Biparental crosses: Crosses betweentwo known parental clones. This is easily acieved by
bringing together two parents in an isolated area or under lanterns.
Area crosses: In this system several male sterile female clones are pollinated by one male parent
in isolated area.
Melting pot crosses: or polycrosses are made by bringing together arrows of large number of
superior/ potential parental cultivars in an isolated area. Natural cross pollination allowed. This
procedure allows the evaluation of breeding behavour of large number of clones with minimum
expense.
Mutation breeding: Presently, mutation breeding in sugarcane aims at creating economic
mutants for higher cane yield, non-flowering and resistance to various dieases like red rot,
smut, downy mildew and various borers. Difficulty with mutation breeding in sugarcane is that
at least three time consuming vegetative generations are needed to get rid of chimerism and to
test for stable mutants.
National Programmes
The research on sugarcane in India is being carried out :
i. Under all India coordinated research project
 ii. At two central institutes, namely, Indian Institute of Sugarcane Research, Lucknow
and Suagrcane breeding Institute, Coimbatore
Limitations of Conventional Breeding and Rationale for Transgenic Approaches Conventional
breeding in sugarcane has several limitations that can be overcome by transgenic approaches.
Listed below are a few such examples.
1. Commercial sugarcane cultivars possess different proportions of chromosomes, complex
recombinational events, and varying chromosome sets (aneuploidy) (Sreenivasan et al., 1987).
This genomic complexity brings difficulties in applying conventional plant breeding for
cultivar improvement. In addition, conventional breeding is a multistage, laborious, and time-
consuming process requiring 10–14 years to develop a new cultivar. A single fault, such as
disease susceptibility in an otherwise elite cultivar, could cause the cultivar to be abandoned.
Conventional breeding approaches to correct such faults in an existing cultivar are impractical
in sugarcane, due to the genetic complexity of cultivars (Birch and Maretzki, 1993). The
capacity to introduce specific genes by transgenic approach, without major genetic
reassortment following crossing, could be used to rescue flawed cultivars (Birch and Maretzki,
1993). For example, the successful production of sugarcane plants resistant to leaf scald disease
was achieved by transgenic approach (Zhang et al., 1999).
2. Although breeding efforts in sugarcane have been successful in increasing cane production,
only limited success has been achieved recently in increasing sugar content. For example, there
has been no increase in sugar content over the last 40 years in Australian sugarcane (Bonnett
et al., 2004b). In the USA, Legendre (1995) reported that the average sucrose content of new
candidate varieties decreased 3.5% on the fifth cycle of recurrent selection, as compared to the
previous cycle, indicating that a limit has been reached for this trait. The QTL analysis of
interspecific F1 populations also indicated that modern sugarcane cultivars have a limited
(biased subset) population of genes controlling sugar content (Ming et al., 2001b). In contrast,
metabolic engineering of sugarcane through transgenic approaches could improve sugar
content. For example, transgenic sugarcane with doubled sugar content was achieved when
attempting to produce isomaltulose in sugarcane (Wu and Birch, 2007).
3. Production of novel products in sugarcane is not possible by conventional breeding. In
contrast, metabolic engineering through transgenic approaches could produce new products,
such as alternative sugars, biopolymers, pharmaceuticals, and high-value proteins. For
example, successful production of sorbitol (Chong et al., 2007), isomaltulose (Wu and Birch,
2007), p-hydroxybenzoic acid (pHBA) and biodegradable polymer (McQualter et al., 2005;
Petrasovits et al., 2007) has been achieved in transgenic sugarcane, which cannot be achieved
through conventional breeding.
4. Conventional breeding allows transfer of traits and genes only between sexually compatible
species. Hence, transfer of traits from noncompatible species is impossible. In contrast,
transgenic approaches allow insertion of novel genes from sexually noncompatible
plants/organisms, enable expression of native genes at different levels in specific tissues or
under novel developmental patterns of expression.
5. The number of traits to be considered when selecting for variety development is determined
by the degree of genetic linkage among those traits. If linkages are rare, several traits can be
selected simultaneously. In the case of sugarcane the extent of those linkages is still uncertain
(Ming et al., 2006). Recent advances in molecular marker-assisted selection and transformation
technologies can alleviate the problem. Thus, genetic transformation by modern molecular
techniques has the potential to enhance a host of traits including sugar, pest and disease
resistance, tolerance to drought and cold, vigor, plant architecture and fiber, and to produce
alternative products such as biopolymers and isomaltulose in sugarcane.