Lecture No: 1

Definition, Aim, Objectives and Scope of Plant Breeding

Definition :

Plant breeding can be defined as an art, a science, and technology of improving the
genetic make up of plants in relation to their economic use for the man kind.
or
Plant breeding is the art and science of improving the heredity of plants for the benefit
of mankind.
or
Plant breeding deals with the genetic improvement of crop plants also known as
science of crop improvement.
or
Science of changing and improving the heredity of plants

Aim :

Plant breeding aims to improve the characteristics of plants so that they become more
desirable agronomically and economically. The specific objectives may vary greatly
depending on the crop under consideration.

Objectives of Plant Breeding :

1. Higher yield : The ultimate aim of plant breeding is to improve the yield of
economic produce. It may be grain yield, fodder yield, fibre yield, tuber yield, cane
yield or oil yield depending upon the crop species. Improvement in yield can be
achieved either by evolving high yielding varieties or hybrids.

2. Improved quality: Quality of produce is another important objective in plant

breeding. The quality characters vary from crop to crop. Eg. grain size, colour,
milling and backing quality in wheat. Cooking quality in rice, malting quality in
barley, size, colour and size of fruits, nutritive and keeping quality in vegetables,
protein content in pulses, oil content in oilseeds, fibre length, strength and fineness in
cotton.

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3. Abiotic resistance : Crop plants also suffer from abiotic factors such as drought, soil
salinity, extreme temperatures, heat, wind, cold and frost, breeder has to develop
resistant varieties for such environmental conditions.

4. B iotic resistance : Crop plants are attacked by various diseases and insects, resulting
in considerable yield losses. Genetic resistance is the cheapest and the best method of
minimizing such losses. Resistant varieties are developed through the use of resistant
donor parents available in the gene pool.

5. Change in maturity Duration / Earliness : Earliness is the most desirable character

which has several advantages. It requires less crop management period, less
insecticidal sprays, permits new crop rotations and often extends the crop area.
Development of wheat varieties suitable for late planting has permitted rice-wheat
rotation. Thus breeding for early maturing crop varieties, or varieties suitable for
different dates of planting may be an important objective. Maturity has been reduced
from 270 days to 170 days in cotton, from 270 days to 120 days in pigeonpea, from
360 days to 270 days in sugarcane.

6. Determinate Growth : Development of varieties with determinate growth is

desirable in crops like Mung, Pigeon Pea (Cajanus cajan ), Cotton (Gossypium sp.),
etc.

7. Dormancy : In some crops, seeds germinate even before harvesting in the standing
crop if there are rains at the time of maturity, e.g., Greengram, Blackgram, Barley
and Pea, etc. A period of dormancy has to be introduced in these crops to check loss
due to germinatio n. In some other cases, however, it may be desirable to remove
dormancy.

8. Desirable Agronomic Characteristics: It includes plant height, branching, tillering

capacity, growth habit, erect or trailing habit etc., is often desirable. For example,
dwarf ness in cereals is generally associated with lodging resistance and better
fertilizer response. Tallness, high tillering and profuse branching are desirable
characters in fodder crops.

9. Elimination of Toxic Substances : It is essential to develop varieties free from toxic

compounds in some crops to make them safe for human consumption. For example,
removal of neurotoxin in Khesari (Lathyruys sativus) which leads to paralysis of
lower limbs, erucic acid from Brassica which is harmful for human health, and

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 gossypol from the seed of cotton is necessary to make them fit for human
consumption. Removal of such toxic substances would increase the nutritional value
of these crops.

10. Non-shattering characteristics: The shattering of pods is serious problem in green

gram. Hence resistance to shattering is an important objective in green gram.

11. Synchronous Maturity : It refers to maturity of a crop species at one time. The
character is highly desirable in crops like Greengram, Cowpea, and Cotton where
several pickings are required for crop harvest.

12. Photo and Thermo insensitivity: Development of varieties insensitive to light and
temperature helps in crossing the cultivation boundaries of crop plants. Photo and
thermo-insensitive varieties of wheat and rice has permitted their cultivation in new
areas. Rice is now cultivated in Punjab, while wheat is a major rabi crop in West
Bengal.

13. Wider adaptability : Adaptability refers to suitability of a variety for general

cultivation over a wide range of environmental conditions. Adaptability is an
important objective in plant breeding because it helps in stabilizing the crop
production over regions and seasons.

14. Varieties for New Seasons : Traditionally Maize is a kharif crop. But scientists are
now able to grow Maize as rabi and zaid crops. Similarly, mung is grown as a
summer crop in addition to the main kharif crop.

Scope of plant breeding (Future Prospects)

From times immemorial, the plant breeding has been helping the mankind. With
knowledge of classical genetics, number of varieties have been evolved in different crop
plants. In order to combat the global alarm created by population explosion, the food front
has to be strengthened which is serious challenge to those scientists concerned with
agriculture. Advances in molecular biology have sharpened the tools of the breeders, and
brighten the prospects of confidence to serve the humanity. The application of biotechnology
to field crop has already led to the field testing of genetically modified crop plants.
Genetically engineered Rice, Maize, Soybean, Cotton, Oilseeds Rape, Sugar Beet and Alfalfa
cultivars are expected to be commercialized before the close of 20th century. Genes from
varied organisms may be expected to boost the performance of crops especially with regard

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 to their resistance to biotic and abiotic stresses. In addition, crop plants are likely to be
cultivated for recovery of valuable compounds like pharmaceuticals produced by genes
introduced into them through genetic engineering. It may be pointed out that in Europe
hirudin, an anti-thrombin protein is already being produced from transgenic Brassica napus.

Undesirable effects
Plant breeding has several useful applications in the improvement of crop plants.
However, it has five main undesirable effects on crop plants.

1. Reduction in Diversity : Modern improved varieties are more uniform than land
races. Thus plant breeding leads to reduction in diversity. The uniform varieties are
more prone to the new races of pathogen than land races which have high genetic
diversity.

2. Narrow genetic base : Uniform varieties have narrow genetic base. Such varieties
generally have poor adaptability.

3. Danger of Uniformity : Most of the improved varieties have some common parents
in the pedigree which may cause danger of uniformity.

4. Undesirable combinations : Sometimes, plant breeding leads to undesirable

combinations. The examples of man made crops having undesirable combination of
characters are Raphanobrassica and Pomato.

5. Increased susceptibility to minor diseases and pests : Due to emphasis on breeding

for resistance to major diseases and insect pests often resulted in an increased
susceptibility to minor diseases and pests. These have gained importance and, in some
cases, produced severe epidemics. The epidemic caused by Botrytis cinerea (grey
mold) in chickpea during 1980-82 Punjab, Haryana. The severe infection by Karnal
bunt (Tilletia sp.) on some wheat varieties, infestation of mealy bugs in Bt cotton.

Lecture No. 2
History and development of plant breeding

- The process of br inging a wild species under human management is referred to as

domestication
- Domestication may be the most basic method of plant breeding
- Domestication continuous today and is likely to continue for some time in future

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- Ex : In case of timber trees medicinal plants, microbes
- During the long period of historic cultivation natural selection has definitely acted on
the domesticated species.
- Movement of man from one place to another brought about the movement of his
cultivated plant species
- 700 BC - Babylonians and Assyrians pollinated date palm artificially
- 17th century - several varieties of heading lettuce were developed in France
- 1717 - Thomas Fair Child - produced the first artificial hybrid, popularly known
as Fair Child’s mule, by using carnation with sweet William
- 1727 - The first plant breeding company was established in France by the vilmorins.
- 1760-1766 - Joseph koelreuter, a German, made extensive crosses in tobacco.
- 1759-1835 – Knight was perhaps the first man to use artificial hybridization to
develop several new fruit varieties.
- Le couteur and Shireff used individual plant selections and progeny test to develop
some useful cereal varieties
- 1873 - the work of Patrick Shireff was first published.
- He concluded that only the variation heritable nature responded to selections, and that
there variation arose through ‘natural sports’ (= mutation) and by ‘natural
hybridization’
(= recombination during meiosis in the hybrids so produced).
- 1856 - Vilmorin developed the progeny test and used this method successfully in the
improvement of sugar beets.
- 1900 - Nilson-Ehle, his associates developed the individual plant selection method in
Sweden.
- 1903 - Johannsen proposed the pureline theory that provided the genetic basis for
individual plant selection.
- The science of genetics began with the rediscovery of Gregor Johan Mendel’s paper
in 1900 by Hugo de veris, Tshermark and Correns which was originally published in
1866.
- The modern plant breeding methods have their bases in the genetic and cytogenetic
principles.
- Numerous workers who determined the various modes of inheritance have contributed
to the development and understanding of plant breeding.

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 - The discovery of chromosomes as carriers of genes has led to the development of
specialized plant breeding methods for chromosome engineering.
- The totipotency of plant somatic and gametic cells allows regeneration of complete
plants from single cells. This, coupled with the development of recombinant DNA
technology, has enabled the transfer of desirable genes from any organism into plants.
Crop varieties developed in this manner are already in cultivation in several countries.

History of plant breeding in India

- 1871 – The Government of India created the Department of Agriculture

- 1905 – The Imperial Agricultural Research Institute was establish in Pusa, Bihar
- 1934 – The buildings of the institute damaged in earthquake
- 1936 – Shifted to New Delhi
- 1946 – Name was changed Indian Agricultural Research Institute
- 1901-05 – Agricultural Colleges were established at Kanpur, Pune, Sabour, Llyalpur,
Coimbatore
- 1929 – Imperial council of Agricultural Research was established
- 1946 – Name was change to Indian Council Agricultural Research
- 1921 – Indian Central Cotton Committee was established – Notable researches on
breeding and cultivation of cotton. Eg : 70 improved varieties of cotton
- 1956 – Project for intensification of regional research on cotton, oilseeds and millets
(PIRRCOM) was initiated to intensify research on these crops – located at 17
different centres through out the country
- 1957 – All India Coordinated maize improvement project was started with objective
of exploiting heterosis
- 1961 - The first hybrid maize varieties released by the project
- ICAR initiated coordinated projects for improvement of the other crops
- 1960 – First Agricultural University established at Pantnagar, Nainital, U.P.
Scientific contributions of eminent scientists

Name of the Scientists Contributions

Allard and Bradshaw - G x E interaction
Recurrent Selection for SCA - Hull
Recurrent Selection for GCA - Jenkins
Dominance hypothesis - Davenport

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Gene for gene hypothesis - Flor
Pureline concept - Johannsen
Backcross method - Harlan and Pope
Double cross scheme - Jones
Cytoplasmic Genetic Male sterility - Jones and Davis
Ear to row method - Hopkins
Colchicine - Blackslee and Nebel
Single Seed Descent Method - Goulden
Self incompatibility - Lewis
Vertifolia effect - Van Der Plank
Centres of diversity, Law of homologus series - Vavilov
Grater initial capital hypothesis - Ashby
Progeny test - Vilmorin
First artificial hybrid - Thomas Fairchild
Triticale - Rimpau
Mutation - Hugo de Vries
Sprophytic System of self incompatibility - Hughes and Babcock
Bulk method - Nilsson & Ehle
Raphano brassica - Karpenchenko
Heterosis - Shull
Male sterility - Jones and Davis
Father of hybrid rice - Yuan Long Ping
Self incompatibility classification - Lewis
Mechanism of insect resistance - Painter
Modified bulk method - Atkins
Components of genetic variance classification - Fischer
Male sterility in maize - Rhoades
Microcentre - Harlan
Chemical mutagen - Aurbach
Multiline concept - Jenson
Green revolution in India - M.S. Swaminathan
Semidwarf rice varieties at IRRI - T.T. Chang
Forage breeder - G.W. Burton

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 Forage breeder - T.J. Jenkin
Soyabean breeder - E.E. Hartwig

Some Indian Plant Breeding

T.S. Venkatraman - An eminent sugarcane breeder, he transferred thick stem and

high sugar contents from tropical noble cane to North Indian
Canes. This process is known as noblization of sugarcane.

B.P. Pal - An eminent Wheat breeder,developed superior disease

resistant N.P. varieties of wheat.

M.S. Swaminathan - Responsible for green revolution in India, developed high

yielding varieties of Wheat and Rice

Pushkarnath - Famous potato breeder

N.G.P. Rao - An eminent sorghum breeder

K. Ramaiah - A renowned rice breeder

Ram Dhan Singh - Famous wheat breeder

D.S. Athwal - Famous pearlmillet breeder

Bosisen - An eminent maize breeder

Dharampal Singh - An eminent oil-seed breeder

C.T. Patel - Famous cotton breeder who developed world’s first cotton
hybrid in 1970

V. Santhanam - Famous cotton breeder

Lecture No: 3
MODES OF REPRODUCTION
Mode of reproduction determines the genetic constitution of crop plants, that is,
whether the plants are normally homozygous or heterozygous. This, in turn, determines the
goal of a breeding programme. If the crop plants are naturally homozygous, e.g., as in self-
pollinators like wheat, a homozygous line would be desirable as a variety. But if the plants
are heterozygous naturally, e.g., as in cross-pollinators like Maize, a heterozygous population

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has to be developed as a variety. Consequently, the breeding methods have to be vastly
different for the two groups of crop plants. A knowledge of the mode of reproduction of crop
plants is also important for making artificial hybrids. Production of hybrids between diverse
and desirable parents is the basis for almost all the modern breeding programmes.
MODES OF REPRODUCTION
The modes of reproduction in crop plants may be broadly grouped into two
categories, asexual and sexual.

Asexual Reproduction
A sexual reproduction does not involve fusion of male and female gamentes. New
plants may develop from vegetative parts of the plant (vegetative reprodu ction) or may arise
from embryos that develop without fertilization (apomixis).
Vegetative Reproduction
In nature, a new plant develops from a portion of the plant body. This may occur
through modified underground and sub-acrial stems, and through bulbills.
Underground Stems
The underground modifications of stem generally serve as storage organs and contain
many buds. These buds develop into shoots and produce plants after rooting. Examples of
such modifications are given below.
Tuber : Potato
Bulb : Onion, Garlic
Rhizome : Ginger, turmeric
Corm : Bunda, arwi

Sub-aerial Stems
These modifications include runner, stolon, sucker etc.,. Sub-aerial stems are used for
the propagation of mint, date plam etc.

Bulbils
Bulbils are modified flowers that develop into plants directly without formation of
seeds. These are vegetative bodies; their development does not involve fertilization and seed
formation. The lower flowers in the inflorescence of garlic naturally develop into bulbils.
Scientists are trying to induce bulbil development in plantation crops by culturing young
inflorescence on tissue culture media ; it has been successfully done in the case of cardamom.
Artificial Vegetative Reproduction
It is commonly used for the propagation of many crop spec ies, although it may not
occur naturally in those species. Stem cuttings are commercially used for the propagation of

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sugarcane, grapes, roses, etc. Layering, budding, grafting and gootee are in common use for
the propagation of fruit trees and ornamental shrubs. Techniques are available for vegetative
multiplication through tissue culture in case of many plant species, and attempts are being
made to develop the techniques for many others. In many of these species sexual
reproduction occurs naturally but for certain reasons vegetative reproduction is more
desirable.

Significance of Vegetative Reproduction

Vegetatively reproducing species offer unique possibilities in breeding. A desirable
plant may be used as a variety directly regardless of whether it is homozygous or
heterozygous. Further, mutant buds, branches or seedlings, if desirable, can be multiplied
and directly used as varieties.

Apomixis
In apomixis, seeds are formed but the embryos develop without fertilization.
Consequently, the plants resulting from them are identical in genotype to the parent plant. In
apomictic species, sexual reproduction is either suppressed or absent. When sexual
reproduction does occur, the apomixis is termed as facultative. But when sexual reproduction
is absent, it is referred to as obligate. Many crop species show apomixis, but it is generally
facultative. The details of apomictic reproduction vary so widely that a confusing
terminology has resulted. A simplified classification of apomixis is given below.
Adventive Embryony
In this case, embryos develop directly from vegerative cells of the ovule, such as
nucellus, integument, and chalaza. Development of embryo does not involve production fo
embryo sac. Adventive embryony occurs in mango, citrus, etc.

Apospory
Some vegetative cells of the ovule develop into unreduced embryo sacs after meiosis.
The embryo may develop from egg cell or some other cell of this embryo sac. Apospory
occurs in some species of Hieraceum, Malus, Crepis, Ranunculus, etc.
Diplospo ry
Embryo sac is produced from the megaspore, which may be haploid or, more
generally, diploid. Generally the meiosis is so modified that the megaspore remains diploid.
Diplospory leads to parthenogenesis or apogamy.
Parthenogenesis
The embryo develops from embryo sac without pollination. It is of two types
Gonial parthenogenesis – embryos develop from egg cell,

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Somatic parthenogenesis – embryos develop from any cell of the embryo sac other than the
egg cell.
Apogamy
In apogamy, synergids or antipodal cells develop into an embryo. Like
parthenogenesis, apogamy may be haploid or diploid depending upon the haploid or diploid
state of the embryo sac. Diploid apogamy occurs in Antennaria, Alchemilla, Allium and
many other plant species.
Significance of Apomixis
Apomixis is a nuisance when the breeder desires to obtain sexual progeny, i.e., selfs
or hybrids. But it is of great help when the breeder desires to maintain varieties. thus in
breeding of apomictic species, the breeder has to avoid apomictic progeny when he is making
crosses or producing inbred lines. But once a desirable genotype has been selected, it can be
multiplied and maintained through apomictic progeny. This would keep the genotype of a
variety intact. Asexually reproducing crop species are highly heterozygous and show severe
inbreeding depression. Therefore, breeding methods in such species must avoid inbreeding.

SEXUAL REPRODUCTION
Sexual reproduction involves fusion of male and female gametes to form a zygote, which
develops in to an embryo. In crop plants, male and female gametes are produced in specialised
structures known as flowers.

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Flower
A flower usually consists of sepals, petals (or their modifications), stamens and/or pistil. A
flower containing both stamens and pistil is a perfect or hermaphrodite flower. If it contains
stamens, but not pistil, it is known as staminate, while a pistillate flower contains pistil, but
not stamens. Staminate and pistillate flowers occur on the same plant in a monoecious
species, such as maize, Colocasia, castor (Ricinus communis), coconut, etc. But in dioecious
species, staminate and pistillate flowers occur on different plants, e.g., papaya, date palm
(Phoenix dactylifera), pistachio (Pistacia vera), etc. In "crop plants, meiotic division of
specific cells in stamen and pistil yields microspores and megaspores, respectively. This is
followed by mitotic division of the spore nuclei to produce gametes; the male and female
gametes are produced in microspores and megaspores, respectively.

Spo rogenesis
Productions of microspores and megaspores is known as sporogenesis. Microspores are
produced in anthers (microsporogenesis), while megs.spores are produced in ovules
(megasporogenesis).

Microsporogenesis. Each anther has four pollen sacs, which contain numerous pollen mother
cells (PMCs). Each PMC undergoes meiosis to produce four haploid cells or microspores.
This process is known as microsporogenesis (Fig. 4.1). The microspores mature into pollen
grains mainly by a thickening of their walls.

Megasporogenesis. Megasporogenesis occurs in ovules, which are present inside the ovary. A
single cell in each ovule differentiates into a megaspore mother cell. The megaspore mother
cell undergoes meiosis to produce four haploid megaspores. Three of the megaspores
degenerate leaving one functional megaspore per ovule (Fig. 4.2). This completes
megasporogenesis.

Gametogenesis
The production of male and female gametes in the microspores and the megaspores,
respectively, is known as gametogenesis.
Microgametogenesis. This refers to the production of male gamete or sperm. During the
maturation of pollen, the microspore nucleus divides mitotically to produce a generative and
a vegetative or tube nucleus. The pollen is generally released in this binucleate stage. When
the pollen lands onto the stigma of a flower, it is known as pollination. Shortly after

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pollination, the pollen germinates. The pollen tube enters the stigma and grows through the
style. The generative nucleus now undergoes a mitotic division to produce two male gametes
or sperms. The pollen, along with the pollen tube, is known as microgametophyte. The pollen
tube finally enters the ovule through a small pore, micropyle, and discharges the two sperms
into the embryo sac.

Microsporogenesis and microgametogenesis (a generalized scheme)

Megagametogenesis. The nucleus of a functional megaspore divides mitotically to produce

four or more nuclei. The exact number of nuclei and their arrangement vary considerably
from one species to another. In most of the crop plants, megaspore nucleus undergoes three
mitotic divisions to produce eight nuclei. Three of these nuclei move to one pole and produce
a central egg cell and two synergid cells; one synergid is situated on either side of the egg
cell. Another three nuclei migrate to the opposite pole to give rise to antipodal cells. The two
nuclei remaining in the centre, the polar nuclei, fuse to form a secondary nucleus. The
megaspore thus develops into a mature megagametophyte or embryo sac. The development
of embryo sac from a megaspore is known as megagametogenesis. The embryo sac generally
contains one egg cell, two synergids, three antipodal cells (all haploid), and one diploid
secondary nucleus.

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 Megasporogenesis and megagametogenesis (generalized scheme)

Significance of Sexual Reproduction

Sexual reproduction makes it possible to combine genes from two parents into a
single hybrid plant. Recombination of these genes produces a large number of genotypes.
This is an essential step in creating variation thr ough hybridization. Almost the entire plant
breeding is based on sexual reproduction. Even in asexually reproducing species, sexual
reproduction, if it occurs, is used to advantage, e.g., in sugarcane, potato, sweet potato etc.

Lecture No. 4
MODES OF POLLINATION
Pollination refers to the transfer of pollen grains from anthers to stigmas. Pollen from an
anther may fall on to the stigma of the same flower leading to self-pollination or outogamy.
When pollen from flowers of one plant are transmitted to the stigmas of flowers of another
plant, it is known as cross-pollination or allogamy. A third situation, geitonogamy, results
when pollen from a flower of one plant falls on the stigmas of other flowers of the same
plant, e.g., in Maize. The genetic consequences of geitonogamy are the same as those of
autogamy.

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Self-pollination
Many cultivated plant species reproduce by self-pollination. Self-pollination species
are believed to have originated from cross-pollinated ancestors. These species, as a rule,
must have hermaphrodite flowers. But in most of these species, self -pollination is not
complete and cross -pollination may occur up to 5%. The degree of cross-pollination in self-
pollinated species is affected by several factors, e.g., variety environmenta l conditions like
temperature and humidity, and location.
Mechanisms promoting self-pollination The various mechanisms that promote self-
pollination are generally more efficient than those promoting cross -pollination. These
mechanisms are listed below.
1. Cleistogamy. In this case, flowers do not open at all. This ensures complete self-
pollination since foreign pollen cannot reach the stigma of a closed flower.
Cleistogamy occurs in some varieties of wheat, oats, barley and in a number of other
grasses.
2. In some species, the flowers open, but only after pollination has taken place. This
occurs in many cereals, such as, wheat, barley, rice and oats. Since the flower does
open, some cross-pollination may occur.
3. In crops like tomato and brinjal, the stigmas are closely surrounded by anthers.
Pollination generally occurs after the flowers open. But the position of anthers in
relation to stigmas ensures self-pollination.
4. In some species, flowers open but the stamens and the sigma are hidden by other
floral organs. In several legumes, e.g., pea, mung, urd, Soybean and gram the
stamens and the stigma are enclosed by the two petals forming a keel.
5. In a few species, stigmas become receptive and elongate through staminal columns.
This ensures predominant self -pollination.

Genetic Consequences of Self-Pollination

Self-pollination leads to a very rapid increase in homozygosity. Therefore,
populations of self-pollinated species are highly homozygous, self-pollinated species do not
show inbreeding depression, but may exhibit considerable heterosis. Therefore, the aim of
breeding methods generally is to develop homozygous varieties.

Cross-Pollination
In cross-pollinating species, the transfer of pollen from a flower to the stigmas of the
others may be brought about by wind (anemophily). Many of the crop plants are naturally

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cross-pollinated (Table 3.1). In many species, a small amount (up to 5-10 percent) of selfing
may occur.
Mechanisms promoting cross pollination There are several mechanism that facilitate cross-
pollination; these mechanisms are described briefly.
1. Dicliny : Dicliny or unisexuality is a condition in which the flowers are either
staminate (male) or pistillate (female).
a) Monoecy. Staminate and pistillate flowers occur in the same plant, either in
the same inflorescene, e.g., Castor, mango and coconut, or in separate
inflorescences, chestnut, strawberries, rubber, grapes and cassava.
b) Dioecy. The male and female flowers are present on different plants, i.e., the
plants in such species are either male or female, e.g., papaya, date, hemp,
asparagus, and spinach. In general, the sex is governed by a single gene, e.g.,
asparagus and papaya. In some cases, there are hermaphrodite plants in
addition to males and females, and a number of intermediate forms may also
occur.
2. Stamens and pistils of hermaphrodite flowers may mature at different times
facilitating cross -pollination.
a) Protogyny. In crop species like bajra, pistils mature before stamens.
b) Protandry. in crops like Maize and sugarbeets, stamens mature before pistils.
3. In Lucerne or alfalfa, stigmas are covered with a waxy film. The stigma does not
become receptive until this waxy film is broken. The waxy membrane is broken by
the visit of honey bees which also effect cross-pollination.
4. A combination of two or more of the above mechanisms may occur in some specie.
This improves the efficiency of the system in promoting cross-pollination. For
example, Maize exhibits both monoecy and protandry.
5. Self-Incompatibility. It refers to the failure of pollen from a flower to fertilize the
same flower or other flowers on the same plant. Self-incompatibility is of two types :
sporophytic and gametophytic. In both the cases, flowers do not set seed on selfing.
Self-incompatibility is common in several species of Brassica, some species of
Nicotiana, radish, rye and many grasses. It is highly effective in preventing self-
pollination.
6. Male Sterility. Male sterility refers to the absence of functional pollen grains in
otherwise hermaphrodite flowers. Male sterility is not common is natural
populations. But it is of great value in experimental populations, particularly in the

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 production of hybrid seed. Male sterility is of two types : genetic and Cytoplasmic.
Cytoplasmic male sterility is termed Cytoplasmic-genetic when restorer genes are
known. In view of the importance of self-incompatibility and male sterility, a more
detailed discussion on them follows later.
Genetic Consequences of Cross-Pollination. Cross-pollination preserves and promotes
heterozygosity in a population. Cross-pollinated species are highly heterozygous and show
mild to severe inbreeding depression and a considerable amount of heterosis. The breeding
methods in such species aim at improving the crop species without reducing heterozygosity
to an appreciable degree. Usually, hybrid or synthetic varieties are the aim of breeder
wherever the seed production of such varieties is economically feasible.

Often Cross-Pollinated Species

In many crop plants (Table 3.1), cross -pollination often exc eeds 5 per cent and may
reach 30 per cent. Such species are generally known as often cross-pollinated species, e.g.,
Jowar, Cotton, arhar, safflower etc. The genetic architecture of such crops is intermediate
between self-pollinated and cross-pollinated species. Con-sequently, in such species
breeding methods suitable for both of them may be profitably applied. But often hybrid
varieties are superior to others.
Lecture No: 5
METHODS OF PLANT BREEDING

Various approaches (viz., selection, hybridization, mutation, etc) that are used for
genetic improvement of crop plants are referred to as plant breeding methods or plant
breeding procedures or plant breeding techniques. The choice of breeding methods mainly
depends on the mode of pollination, mode of reproduction, gene action and breeding
objective of crop species. Plant breeding methods are generally classified on the basis of
application of crop improvement (general methods, special methods and population
improvement approaches) and hybridization (methods involving hybridization and methods
not involving hybridization).
Various breeding procedures that are more commonly used for the genetic
improvement of various crop plants are known as general breeding methods. Such breeding
methods include introduction, selection (pure line selection, mass selection, progeny
selection), hybridization (pedigree, bulk and backcross methods), heterosis breeding,
synthetic and composite breeding. On the other hand, those breeding procedures that are
rarely used for improvement of crop plants are referred to as special breeding methods. Such

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methods include: mutation breeding, polyploidy breeding, wide crossing or distant
hybridization and biotechnology. Four breeding approaches, viz., recurrent selection,
disruptive matin g and selection, diallel selective mating system and biparental mating are
used mainly for population improvement.
Classification of Plant Breeding Methods
Basis of classification and Breeding methods included
Types of methods
A. Application in crop improvement
(1) General Methods Plant introduction, Pure line selection, mass
selection, progeny selection, pedigree
method, bulk method, back cross method,
SSD, clonal selection, heterosis breeding,
synthetics and composites.
(2) Special Methods Mutation breeding, Polyploidy breeding,
transgenic breeding, molecular breeding.
(3) Population Improvement Recurrent selection, disruptive selection,
diallel selective approaches mating system,
biparental mating.
B. Hybridization
(1) Methods involving hybridization Pedigree, bulk, backcross and SSD Methods:
heterosis breeding, and population
improvement approaches and molecular
breeding (marker aided selection).
(2) Methods not involving hybridization Plant Introduction, pureline selection, mass
selection, progeny selection, clonal selection,
mutation breeding and transgenic breeding.

There are some differences in the breeding methods used for self pollinated and cross
pollinated species. Self pollinated species are homozygous, hence we can start hybridization
directly. Cross pollinated species, on the other hand, are highly heterozygous. Hence we can
not start hybridization directly. First we have to develop inbred lines by selfing or inbreeding
and then only hybridization can be taken up. We have to exploit homozygosity in self
pollinated crops and heterozygosity in cross pollinated species. Asexually propagated species
such as sugarcane, potato, sweet potato, etc., are highly heterozygous. Hence, F1 hybrids in

21
such crops exhibit segregation and selection can be practiced in F 1 generation. The superior
clones are identified and further multiplied. The maintenance or conservation of hybrid
vigour is easy in such crops because of asexually propagation.
Methods of Breeding Autogamous species
Plant breeding methods that are used for genetic improvement of self pollinated or
autogamous species include:
1. Plant Introduction
2. Pureline selection
3. Mass selection
4. Pedigree method
5. Bulk method
6. Single seed descent method
7. Backcross method
8. Heterosis breeding
9. Mutation breeding
10. Polyploidy breeding
11. Distant hybridization
12. Transgenic breeding.
Four breeding approaches, viz., recurrent selection, disruptive selection, diallele selective
mating, and biparental mating are used for population improvement.
Methods or Breeding Allogamous species
Breeding methods that are used for genetic improvement of cross pollinated or
allogamous species include
(1) Plant introduction
(2) Mass and progeny selection
(3) Backcross method
(4) Heterosis breeding
(5) Synthetic breeding
(6) Composite breeding
(7) Polyploidy breeding
(8) Distant hybridization
(9) Transgenic breeding

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 Mutation breeding is rarely used in allogamous species. Three breeding approaches
viz., recurrent selection, disruptive mating and biparental meting are used for population
improvement.
Methods of Breeding Asexually Propagated Species
Important breeding methods applicable to asexually propagated species are
(1) Plant Introduction
(2) Clonal selection
(3) Mass selection
(4) Heterosis breeding
(5) Mutation breeding
(6) Polyploidy breeding
(7) Distant hybridization
(8) Transgenic breedin g.
Mass selection is rarely used in asexually propagated species.
Brief account of breeding methods
Plant introduction is applicable to all three groups of crop plants, viz., self pollinated,
cross pollinated and asexually propagated species. It is an old est and rapid method of crop
improvement. The introduced material may be used in three ways viz.,
(1) Directly as a variety
(2) As a variety after selection
(3) As a parent in the hybridization for development of variety or hybrid
Pureline selection is applicable to self pollinated species. It is also used sometimes in
cross pollinated species for development of inbred lines. A single best pure line is released as
a variety. Thus a pureline variety is homozygous and homogeneous population.
Mass selection is common in cross pollinated species and rare in self pollinated and
asexually propagates species. In self pollinated crops, a mass selected variety is a mixture of
several purelines. Thus it is a homozygous but heterogeneous population. In cross pollinated
species, a mass selected variety is a mixture of several hetero and homozygotes. Thus, it is a
heterozygous and heterogeneous population
Progeny selection is used in cross pollinated species. A variety developed by this
method is heterozygous and heterogeneous population because it consists of several hetero
and homozygotes.
Pedigree method is applicable to both self and cross pollinated species. In self
pollinated crops progeny of a single best homozygote is released as a variety. Thus a variety
developed by this method has a homozygous and homogeneous population. In cross

23
pollinated species, it is used for developed of inbred lines. Bulk nad single seed descent
methods are used in self pollinated species. Progeny of a single best homozygote is released
as a varie ty by these methods. Thus, varieties developed by these methods are homozygous
and homogeneous.
Backcross method is applicable in all three groups of crop species. This method is
used for transfer of oligogenic characters from a donor source to a well adapted variety. This
method is also used for development of multilines, Isogenic lines and transfer of male
sterility. This method is more effective in transferring oligogenic characters than polygenic
traits. The end product of backcross method is similar to parent variety expect for the
character which has to be transferred from the donor source.
Multiline varieties are developed in self pollinated species. They are mixture of
several Isogenic lines, closely related lines or unrelated lines. Thus a multiline variety is a
homozygous but heterogeneous population.
Clonal selection is used in asexually propagated species. In this method progeny of a
single best clone is released as a variety. Such variety has heterozygous but homogeneous
population.
Heterosis breeding is used in/all the three groups. However, it is common in cross
pollinated and asexually propagated species and rare in self pollinated species. A hybrid
variety has homogeneous but heterozygous population. Synthetic and composite varieties are
developed in cross pollinated species. Such varieties consist of several homozygotes and
heterozygotes and thus constitute a heterogeneous population.
Mutation breeding is common in self pollinated and asexually propagated species and
rare in cross pollinated species. A mutant variety differs from parent variety in one or few
characters. A mutant differs from a segregant in two main ways. Firstly, the frequency of
segregants is very high and that of mutant is extremely low (0.1%). Secondly, mutant differs
from parent variety in one or few characters, where as a segregant differs from parent
material in several characters.
Polyploidy breeding is common in asexually propagated species and rare in self and
cross pollinated species. A polyploidy variety differs from parent variety in chromosome
numbers and exhibit gigant morphological characters.
Distant hybridization is used in all the three types of crop species. However, this
method is used for transferring some desirable genes from wild species to the cultivated ones.
Generally, backcross method is used for transfer of oligogenic characters and pedigree
method for transfer of polygenic characters.

24
 Transgenic breeding is applicable to all three types of crop species. This method is
used to solve specific problems which can not be solved by conventional breeding
techniques. This method will serve as a tool and can not be used as a substitute for
conventional breeding methods.
Recurrent selection is common in cross pollinated species and rare in other two
groups. It is used for accumulating favourable genes in a population i.e., for population
improvement. Other approaches which are used for population improvement include
disruptive mating, diallel selective mating (DSM) and biparental mating. DSM is used in self
pollinated species and other two techniques can be used both in self and cross pollinated
species.

Lecture No: 6
Plant Introduction
Plant introduction consists of taking a genotype or a group of genotypes of plants into
new environments where they were not being grown before. Introduction may involve new
varieties of a crop already grown in the area, wild relatives of the crop species or a totally
new crop species. Mostly materials are introduced from other countries or continents. But
movement of crop varietie s from one environment into another within a country is also
introduction. Some examples of within the country introduction are popularization of grape
cultivation in Haryana, Introduction of wheat in West Bengal, Rice in Punjab etc.
Primary Introduction : When the introduced variety is well suited to the new environment,
it is released fro commercial cultivation without any alteration in the original genotype, this
constitutes primary introduction. Primary introduction is less common, particularly in
countries having well organized crop improvement programmes. Introduction of semi dwarf
wheat varieties Sonora 64, Lerma Roja and of semi dwarf rice varieties Taichung Native 1
(TN-1), IR-8 and IR-36 are some examples of primary introductions.
Secondary Introduction: The introduced variety may be subjected to selection to isolate a
superior variety. Alternatively, it may be hybridized with local varieties to transfer one or few
characters from this variety to the local ones these processes are known as secondary
introduction. Secondary introduction is much more common than primary introduction.
Examples of secondary introduction are Kalyan Sona and sonalika wheat varieties selected
from material introduced from CIMMYT, Mexico.
History of Plant Introduction : crop plants have traveled ino many new areas from their
centres of origin. This movement of plants occurred with the movement of man. Most of
these introductions occurred very early in the histoy. For example, mung mustard, pear, apple
and walnut were introduced from the Central Asian Center of origin into various parts of
India. Similarly sesame, Jowar, arhar, Asian Cotton and finger millet originated in Africa and
traveled to India in the prehistoric period. From this it is clear that plant wealth of various
nations is to a large extent the result of plant introductions.
For several centuries A.D. the agencies of plant Introduction wre invaders, settlers,
traders, travelles, explorers and naturalists. The plant introduction were made either
knowingly or unknowingly. Muslim invaders introduced in India cherries and grapes from

25
Afghanistan by 1300 A.S. In the 16th century A.D. Portugues introduced Maize, groundnut,
chillies, potato, sweet potato, guava pineapple, papaya, cashewnut and Tobacco. East India
Company brought tea, litchi, and loquat from China. Cabbage, cauliflower and other
vegetables from the Mediterranean; annatto and mahogany from West Indies in the last
quarter of 18th century.
During 19th century, a number of botanic gardens played an important role in plant
introduction. The Calcutta botanic gardens was established in 1781. the Kew botanic gardens,
England arranged introduction of quinine and rubber trees from South America into India.
During and after the last part of 19th century various agricultural and horticulture research
stations were established in the country. These stations introduced horticulture and
agriculture plants independent of each other. There was no co-ordination among these
agencies regarding their introduction activities.
Plant Introduction Agencies in India
A centralized plant introduction agency was initiated in 1946 at the Indian
Agricultural Research Institute (IARI), New Delhi. The agency began as a plant introduction
scheme in the Division of Botany and was funded by ICAR. In 1956, during the second five
year plan, the scheme was expanded as the Plant Introduction and Exploration Organisation.
Subsequently in 1961, it was made an independent division in IARI, the Division of Plant
Introduction. The division was re organized as National Bureau of Plant Genetic Resources
(NBPGR) in 1976. the nature of activities and the functions of the bureau have remained the
same, but the scope and scale of its activities have increased considerably. The bureau is
responsible for the introduction and maintenance of germplasm of agricultural and
horticultural plants.
In addition to the National Bureau of Plant Genetic Resources, there are some other
agencies concerned with plant introduction. Forest Research Institute, Dehradun, has a plant
introduction organization which looks after the introduction, maintenance and testing of
germplasm of forest trees. The Botanical Survey of India was established in 1890 ; it was
responsible for the introduction, testing and maintenance of plant materials of botanical and
medicinal interest. But at present, introduction and improvement of medicinal plants is being
looked after by NBPGR. The Central Research Institute for various crops, e.g., tea, coffee,
sugarcane, potato, Tobacco, rice etc., introduce, test and maintain plant materials of their
interest. But their activities are coordinated by the NBPGR, which has the ultimate
responsibility for introduction activities. Plant material may also be introduced by individual
scientists, universities and other research organizations. But all the introductions in India
must be routed through the NBPGR, New Delhi.
The National Bureau of Plant Genetic Resources. The bureau has it headquarters at IARI,
New Delhi. It has four substations for the testing of introduced plant materials. These
substations represent the various climatic zones of India, they are listed below.
1. Simla. It is situated in Himachal Pradesh and represents the temperate zone ;
approximately 2,300 m above sea level.
2. Jodhpur, Rajasthan. It represents the arid zone
3. Kanya Kumari, Tamil Nadu. It represents the tropical zone
4. Akola, Maharashtra. It represents the mixed climatic zone. It was recently shifted
from Amravati.

26
 In addition, a new substation has recently been established at Shillong for collection of
germplasm from North-east India. This part of the country has a large genetic variability
for several crop species, e.g., rice, citrus, Maize etc.
The bureau functions as the central agency for the export and introduction of
germplasm of economic importance. The bureau is assisted in its activities by the various
Central Research Institutes of ICAR. The activities of the bureau are summarized below.
1. It introduces the required germplasm from its counterparts or other agencies in other
countries.
2. It arranges explorations inside and outside the country to collect valuable germplasm.
3. It is responsible for the inspection and quarantine of all the introduced plant materials.
4. Testing, multiplication and maintenance of germplasm obtained through various
sources. This may be done by the bureau itself at one of its substations or by one of
the concerned Central Institutes of ICAR.
5. To supply, on request, germplasm to various scientists or institutions. The germplasm
may be supplied ex-stock or may by procured from outside in case it is not available
in the country.
6. Maintenance of records of plant name, variety name, propagating material, special
characteristics, source, date and other relevant information about the materials
received.
7. To supply germplasm to its counterparts or other agencies in other countries.
8. To publish its exchange and collection lists. An Introduction News Letter containing
such lists is being published by the Food and Agriculture Organisation (FAO) since
1957 at irregular intervals. NBPGR has also published some lists, and is in the
process of publishing some other catalogues.
9. To set up natural gene sanctuaries of plants where genetic resources are endangered.
10. Improvement of certain plants like medicinal and aromatic plants.
Procedure of Plant Introduction
Introduction consists of the following steps : Procurement, quarantine, cataloguing,
evaluation, multiplication and distribution.
1. Procurement : Any individual or institution can introduce germplasm in India. But all the
introductions must be routed through the NBPGR, New Delhi. There are two routes for plant
introduction. In first route the individual or the institution makes a direct request to an
individual or institution abroad, who has the desired germplasm, to send it through the
NBPGR, New Delhi. In second procedure the individual or institute submits his germplasm
requirements to the NBPGR with a request for their import.
2. Quarantine : Quarantine means to keep materials in isolation to prevent the spread of
diseases etc. All the introduced plant propagules are thoroughly inspected for contamination
with weeds, diseases and insect pests. Materials that are suspected to be contaminated are

27
fumigated or are given other treatments to get rid of the contamination. If necessary, the
materials are grown in isolation for observation of diseases, insect pests and weeds. The
entire process is known as quarantine and the rules prescribing them are known as quarantine
rules.
3. Cataloguing : When an introduction is received, it is given an entry number. Further,
information regarding name of the species, variety, place of origin, adaptation and its various
characteristics are recorded. The plant materials are classified into three groups.
1. Exotic collections are given the prefix ‘EC’
2. Indigenous collections are designated as ‘IC’ and
3. Indigenous wild collections are marked as ‘IW’
4. Evaluation : To assess the potential of new introductions, their performance is evaluated at
different substations of the Bureau. In case of those crops for which Central Research
Institutes are functioning, e.g., rice, sugarcane, potato, Tobacco etc., the introduced materials
are evaluated and maintained by these institutes. The resistance to diseases and pests is
evaluated under environments favouring heavy attacks by them.
Acclimatization : Generally, the introduced varieties perform poorly because they are often
not adapted to the new environment. Sometimes, the performance of a variety in the new
environment improves with the number of generations grown there. The process that leads to
the adaptation of a variety to a new environment is known as acclimatization.
Acclimatization isbrought about by a faster multiplication of those genotypes (present in the
original population) that are better adapted to the new environment. Thus acclimatization is
essentially natural selection. Variability must be present in the original population for
acclimatization to occur. Therefore, land varieties are likely to get acclimatized, while
purelines are not likely to.
The extent of acclimatization is determined by (1) the mode of pollination, (2) the
range of genetic variability present in the original population, and (3) the duration of life-
cycle of the crop. Cross-pollination leads to a far greater gene recombinations than self-
pollination. As a result cross-pollination is much more helpful in acclimatisation than self-
pollination.
5. Multiplication and Distribution : Promising introductions or selections from the
introductions may be increased and released as varieties after the necessary trials. most of the
introductions, however, are characterized for desirable traits and are maintained for future
use. Such materials are used in crossing programmes and are readily supplied by the bureau
on request.
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.

28
To Serve as New Varieties. Sometimes introductions are directly released as superior
commercial varieties. The Maxican semidwarf wheat varieties Sonora 64 and Lerma Rojo,
semidrawf 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 entirely new crop plants.
2. It provides superior varieties either directly or after selection & hybridization.

29
 3. Introduction and exploration are the only feasible means of collecting germplasm and
to protect variability from genetic erosion.
4. It is very quick & economical method of crop improvement, particularly when the
introductions are released as varieties either directly or after a simple selection.
5. Plants may be introduced in new disease free areas to protect them from damage, e.g.,
coffee and rubber.

Demerits of Plant Introduction

The disadvantages of plant introduction are associated with the introduction of weeds,
diseases and pests.
Germplasm Collections
The sum total of hereditary material or genes present in a species is known as the
germplasm of that species. Therefore, a germplasm collection is the collection of a large
number of genotypes of a crop species and its wild relatives. Germplasm collections are also
known as gene banks (or world over the world). Furthe r, germplasm collections furnish the
richest source of variability. Crop improvement would ultimately depend upon the
availability of this variability to be utilized in breeding programmes.
With the modernization of agriculture, large tracts of land have been put under
pureline varieties of self-pollinated crops and hybrid varieties of cross-pollinated species.
This has led to a gradual disappearance of local or land varieties (‘desi’ varieties) and open-
pollinated varieties -both reservoirs of considerable variability. Cultivation and grazing are
gradually destroying many wild species and their breeding grounds. Wild relatives of crops
may be eliminated by introduced species of weedy nature or even by the cultivated forms
derived from them. The gradual loss of variability in the cultivated forms and in their wild
relatives is referred to as genetic erosion. This variability arose in nature over an extremely
long period of time and, if lost, would not be reproduced during a short period.
Most of the countries are greatly concerned about genetic erosion. The establishment
of IBPGR to coordinate germplasm conservation activities throughout the world reflects this
concern. Germplasm collections are being made and maintained to conserve as many
genotype as possible. The germplasm collections contain land varieties, various wild forms,
primitive races, exotic collections and highly evolved varieties. Some of the important
germplasm collections are listed below.
1. Institute of Plant Industry, Leningard. It has 1,60,000 entries of crop plants.
2. Royal Botanic Gardens, Kew, England, It has over 45,000 entries.

30
 3. Bellsville, U.S.A., maintains germplasm collections of small grain crops.
4. World collections of some of the crops are maintained at the following places.
i) Sugarcane. Canal Point, Florida, U.S.A. and Sugarcane Breeding Institute,
Coimbatore (2,800 entries).
ii) Groundnut. Bambey, Senegal (Africa).
iii) Potato. Cambridge, U.K. and Wisconsin, U.S.A.
iv) Annual New World Cottons. Near Tashkent, U.S.S.R.
v) Coffee. Ethiopis (Africa).
vi) Sweet Potatoes. New Zealand
5. The National Bureau of Plant Genetic Resources, New Delhi, is maintaining large
collections of Sorghum, Pennisetum, wheat, barley, oats, rice, Maize and other
agricultural and horticultural crops. For example, groundnut collection is maintained
at Junagarh, Cotton at Nagpur, Potato at Simla, Tobacco at Rajahmundhry, tuber
crops (other than potato) at Trivandrum etc. The Cotton collection maintained at
Central Institute for Cotton Research (CICR, Nagpur) are as follo ws ; Gossypium
hirsutum-4,100 entries ; G. barbadense-300 entries ; G. arboreum-1755 entries ; G.
herbaceum-393 entries (1991).
6. IRRI, Philippines, is maintaining 42,000 rice strains and varieties. More than 15,000
entries are maintained at CRRI, Cuttack.
7. The various International Institutes are building up and maintaining collections of
many species.
Seeds of most species lose viability quickly. Consequently, germplasm collections have
to be grown every few years. (1) Growing, harvesting and storing lar ge collections is a costly
affair requiring much time, labour, land and money. (2) There is also risk of errors in
labeling. (3) The genotypic constitution of entries may also change, particularly when they
are grown in environments considerably different from that to which they are adapted. This
is particularly true in case of cross-pollinated species and for local varieties of self-pollinated
species. These difficulties may be considerably reduced by cold storage of seeds. Seeds of
most of the plant species can be stored for 10 years or more at low temperatures and low
humidity. Thus the entries could be grown every 10 years or so instead of every one or two
years. Cold storage facilities are being utilized at Fort Collins, U.S.A and at IRRI,
Phillippines, NBPGR has developed cold storage facilities for germplasm maintenance this is
known as National Germplasm Repository.
Gene Sanctuaries. It has been proposed that within the centres of origin areas of the greatest
diversity should be demarcated and protected from human disturbances. In such areas, the
evolutionary potential of the local populations and the environment would be preserved. This
would not only preserve variability in these populations, but would also allow evolution to

31
continue and create new types. NBPGR proposes to establish gene sanctuaries in Meghalaya
for Citrus, and in the North-Eastern Region for Musa, Citrus, Oryza, Saccharum and
Mangifera.
Thus a gene sanctuary may be defined as an area of diversity protected from human
interference. A gene sanctuary conserves the germplasm in-situ, within the environment
where it naturally grows. This not only conserves the germplasm with very little labour and
expense, but also permits evolution to proceed on its natural course. This allows the
appearance of new gene combinations and new alleles not present in the preexisting
population.
Exploration: Explorations are trips for the purpose of collection of various forms of crop
plants and their related species. Explorations generally cover those areas that are likely to
show the greatest diversity of forms. The centres of origin are such areas and are often
visited by various exploration teams. In addition to wild forms, land races and open-
pollinated varieties are also collected. Exploration is the primary source of all the germplasm
maintained in germplasm collections.
Lecture No. 7
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 espistasis, there are just three basic principles of selection (Walker,
1969) :

32
 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 parent-
progeny 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 others : As a consequence of its past evolutionary history and breeding
struc ture, 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 characterics 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.
Selection intensity I) It is the amount of selection applied expressed as the proportion of the
population favoured (selected). The selection intensity is inversely proportional to the
percentage proportion selected (PS), as reflectedin Table 1.

33
Table 1 : Relationship between selection intensity and proportion of
population selected
Selection intensity 2.64 2.42 2.06 1.76 1.40 1.16 0.97 0.80 0.34 0
(i)
% selected (PS) 1% 2% 5% 10% 20% 30% 40% 50% 80% 100%

Thus, larger the size of I, more stringent is the selection pressure (hence low fraction
is selected) and vice versa. Then, no selection means all the members of a population are
allowed to reproduce (I=0, PS=100%), and zero selection means the whole population is
rejected (PS=0). However, in real selection experiments, as the desired alleles become
preponderant after each cycle of selection, I is also changed.
The I is of the greatest consequence is bringing about changes in the gene frequency
under selection. However, since the latter does not mean undue loss of desirable alleles, or
undue load of population size, the choice of an arbitrary value of I may be hazardous in a
plant breeding programme. The small size of I (i.e. low selection pressure) may cause a large
population size to be handled in the next generation, which will unnecessarily be taxing on
time and resources. On the other hand, a large size of I (high selection pressure) might cause
allelic erosion due to genetic drift (i.e. changes in gene frequencies due to sampling error, or
small sample size under selection in a finite population not due to genetic causes). Therefore,
an appropriate level of I should be chosen based upon the range of variability present in the
population subjected to selection.
The I=2.06 to 1.76 (i.e. around 5-10 per cent of individuals selected) has generally
been found appropriate in plant populations. However, the limit of selection intensity is set
by two factors : (i) population size, and (ii) inbreeding. Under natural selection, selection
intensity is expressed as the relative number of offsprings produced by different genotypes,
and is termed as selection coefficient.
Selection differential : Difference between the mean of the population and mean of the
selected individuals. Expressed in terms of standard deviation and is designated as ‘S’
Selection differential (S) S is the average superiority of selected individuals over the mean of
population of their origin. It is considered in the same parental generation before selection is
made. An arbitrary culling level, k(i.e. I) is fixed for a trait and individuals beyond that level
are selected. The average of all such selected individuals can be designated by X. then the
mean of selected individuals (X) exceeds the parental population mean by the measure of S.

34
That is S=X-µ. Therefore, wider the phenotypic variability (i.e. phenotypic variance, and
phenotypic standard deviation, that measures variability), greater is the possibility of S
being large.
Heritability : In crop improvement, only the genetic component of variation is important
since only this component is transmitted to the next generation. The ratio of genetic variance
to the total variance i.e. phenotypic variance is known as heritability.
H = Vg / Vp
Vp = Vg+Ve Where Vp = phenotypic variance
Vg = genotypic variance
Ve + error variance of environmental variance
Heritability estimated from the above formula is known as the broad sense
heritability. This is valid when homozygous lines are studied. But when segregating
generations are studied genotypic variance consists of (a) additive variance (b) dominance
variance (c) and variance due to epistasis.
Dominance variance is important when we are dealing with hybrids i.e. F1
generations. In self pollinated crops we release varieties only after making them homozygous
lines. Hence additive variance is more important in such cases. The proportion of additive
genetic variance to the total variance is known as narrow sence heritability.
If heritability is very high for any character it can be improved. Improvement of
characters with low heritability is very difficult.
Genetic Advance : Genetic advance is the difference between the mean of the selected
plants in the original population and the mean of the progeny raised from the selected plants
in the next generation. It can be predicted by the following formula.
Genetic advance(GA) = s P * H * K
K = selection intensity 2.06 when 5% of the population is selected
P = phenotypic standard deviation of the character in the population
H = heritability in broadsense

Lecture No. 8
MASS SELECTION
It is the earliest method of selection. Man has always practiced mass selection
consciously or unconciously from the time of domestication. In its most basic form mass
selection consists of selecting individuals on the basis of phenotypic superiority and mixing
the seeds for using as planting material for next season.

35
 Procedure for evolving variety by mass selection
First year : Large number of phenotypically similar plants having desirable characters are
selected. The number may vary from few hundred to few thousand. The seeds from the
selected plants are composited to raise the next generation.
Second year : composited seed planted in a preliminary field trial along with standard
checks. The variety from which the selection was made should also be included as check.
Phenotypic characterlistics of the variety are critically examined and evaluated.
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.

Modification of mass selection

Mass selection is used for improing a local variety. Large number of plants are
selected (I year) and individual plant progenies are raised (II year). Inferior, segregating
progenies are reflected. Uniform, superior rows are selected and the seed is bulked.
Preliminary yield trials are conducted in third year. Fourth to seventh year multilocation tests
are conducted and seed is multiplied in eight year and distributed in ninth year. Many other
modifications also are followed depending on the availability of time and purpose for ehich it
is used.
Merits of Mass selection :
1. Can be practiced both in self and cross pollinated crops
2. The varieties developed through mass selection are more widely adopted than pure
lines.
3. It retains considerable variability and hence further improvement is possible in future
by selection
4. Helps in preservation of land races
5. Useful for purification of pureline varieties
6. Improvement of characters governed by few genes with high heritability is possible.
7. Less time consuming and less expensive.

Demerits of mass selection

1. Varieties are not uniform
2. Since no progeny test is done, the genotype of the selected plant is not known

36
 3. Since selection is based on phenotype and no control over pollination the
improvement brought about is not permanent. Hence, the process of mass selection
has to be repeated not and then.
4. Characters which are governed by large number of genes with low heritability can not
be improved.
5. It can not create any new genotype but utilizes existing genetic variability.
Achievements
Mass selection must have been used by pre historic man to develop present day
cultivated cross from their wild parents. It was also used extensively before pureline
selection came into existence.
Cotton : Dharwad American Cotton
Groundnut : TMV-1 & TMV-2
Bajra : pusa moti, Baja puri, Jamnagar gaint, AF3
Sorghum : R.S. 1
Rice : SLO 13, MTU-15
Potato : K122

Lecture No 9
JOHANNSEN’S PURE LINE THEORY

The pure line theory

A pureline is a progeny of a single homozygous plant of a self-pollinated species. All
the plants of a pureline have the same genotype. The phenotypic differences within a
pureline is due to environment. Therefore variation within a pureline is not heritable. Hence
selection in a pureline is not effective.
The concept of pureline was proposed by Johannsen in 1903 on the basis of his
studies with princess variety of beans (Phaseolus vulgaris). From a commercial seed lot he
selected seeds of different sizes and grew them separately. The progenies differed in seed
size. Progenies from larger seeds produced larger seeds than those obtained from smaller
seeds. This clearly showed that the variation in seed size in the commercial seed lot of
princess variety had a genetic base. As a result selection for seed size was effective.
Johannsen further studied 19 lines, each line was a progeny of a single seed from the original
lot. He discovered that each line showed a characteris mean seed weght, rangin from 640mg

37
in Line No 1 to 350 mg in line No 19. the seed size within a line showed some variation,
which was much smaller than that in the original commercial seed lot. Johannsen postulated
that the original seed lot was a mixture of purelines. Thus each of the 19 lines represented a
pureline, and the variation in seed size within each of the urelines had no genetic basis and
was entirely due to environment.
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 Johannsens’ 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

38
 variation within a pureline is purely environmental and, as a result, selection within a
pureline is ineffective.

The two main conclusions of Johannsen’s experiment are

1. Selection is effective in population since it contains a mixture of several genotypes.
2. Selection is ineffective in a pureline, since it is a progeny of single, self fertilized
homozygous individual.

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

Characters 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

The 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 beat 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.

39
Genetic basis of pure line
Self pollination increases homozygosity with a corresponding decrease in
heterozygosity. The effect of homozygosity and heterozygosity may be illustrated by taking
an individual heterozygous for (Aa) a single gene as follows

No of Frequency % Frequency %
generations AA Aa aa Homo- Hetero -
of selfing zygosity zygosity
1 0 100 0 0 100
2 25 50 25 50 50
3 37.5 25 37.5 75 25
4 43.75 12.5 43.75 87.5 12.5
5 46.875 6.25 46.875 93.73 6.25
6 48.437 3.125 48.437 96.874 3.125
7 49.218 1.562 49.218 98.436 1.562
8 49.608 0.781 49.608 99.216 0.781
9 49.803 0.39 49.803 99.606 0.39

Proportion of completely homozygous plants in the population

[(2m – 1) / 2m]n
m = No. of generations of self pollination
n = No. of genes segregating
Suppose an individual heterozygous for a single gene (Aa) and the successive
generations derived from it are subjected to self-pollination.
Every generation of self -pollination will reduce the frequency of heterozygote Aa to
50 per cent of that in the previous generation.
There is a corresponding increase in the frequency of the two homozygotes AA and
aa. As a result, after 10 generations of selfing, virtually all the plants in the population would
be homozygous, i.e., AA and a.
On the other hand, the frequency of heterozygote Aa would be only 0.097 per cent,
which is negligible. It is assumed here that the three genotypes AA, Aa and aa have equal
survival. If there is unequal survival, it may increase or decrease the rate at which
homozygosity is achieved. If Aa is favoured, the rate of increase in homozygosity would be
lower than expected.

40
 But if Aa is selected against, hom ozygosity would increase at a faster rate than
expected.

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

General procedure for evolving a variety by pureline selection

The pureline selection has three steps.
1. Selection of individual plants from a local variety or some other mixed population.
2. Visual evaluation of individual plant progenies and
3. Yield trials
1. Selection
First year : A large number of plants (200-3000) which are superior than the rest are
selected from a local variety or mixed population and harvested separately (in some
cases individual heads or stems may be selected). The number of plants to be selected
depends upon the breeder’s discretion but should be as large as possible in view of the
available time, land, funds, labour etc. It is advisable to select for easily observable
characters such as flowering, maturity, disease resistance, plant height etc.
II. Evaluation :
Second year: Progenies of individual plants selected in 1st year are grown separately
with proper spacing (plant to row or head to row). The progenies are evaluated by
taking elaborate date on visual characters such as plant height, duration, grain type,
ear characters besides yield. The number of progenies should be reduced as much as
possible. Disease epiphytotics may be created to test the progenies for disease
resistance, poor, weak, diseased, insect attacked and segregating progenies are
rejected. The superior progenies are harvested separately. If necessary the process
may be repeated for one or more years.
III. Yield trials :

41
 Third year : The selected progenies, now called as cultures are grown in replicated
trial for critical evaluation of yield etc. The best local variety is used as a check and
should be grown at regular intervals, after every 15 or 20 cultures for comparision.
This is known as preliminary yield trial.
Superior cultures based on observable characters and yield are selected. The
number is drastically reduced.
Fourth & Fifth years : The superior cultures are tested against the local checks in
yield trials. Observations are recorded on many characters like diseases resistance,
days to flower, days to maturity, height of the plant ear characters, test weight and
yield. The data is subjected to statistical analysis to identify really superior cultures.
If necessary the trials may be extended for one more year or season. Inferior culture
are rejected and a few (4-5) promising cultures are selected.
Sixth, Seventh and Eighth years: The promising cultures selected are evaluated at
several locations along with strains or cultures of other breeders and local checks.
One or two promising cultures are selected.
Ninth year: The best progeny identified earlier is multiplied, named and released as a
variety for official release of any variety (approval from the variety releasing
committee of the state or central is necessary).

Advantage of pureline selection

1. The purelines are extremely uniform since all the plants in the variety will
have the same genotype.
2. Attractive and liked by the farmers and consumers.
3. Purelines are stable and long test for many years.
4. Due to its extreme uniformity the variety can be easily identified in seed
certification programmes.

Limitations or disadvantages of pureline selection

1. New genotypes are not created by pureline selection
2. Improvement is limited to the isolation of the best genotype present in
population. No more improvement is possible after isolation of the best
available genotype in the population.
3. Selection of purelines require great skill and familiarity with the crop.
4. Difficult to detect small differences that exist between cultures
5. The breeder has to devote more time

42
 6. Pure lines have limited adaptability hence can be recommended for cultivation
in limited area only.
Achievements :
Several varieties developed by pureline selection were released in many crops.
Some examples are given below
Rice : Mtu-1, Mtu-3, Mtu-7, Bcp-1, Adt-1, 3, 5, and 10
Sorghum : G 1 & 2, M 1 & 2, OO 1, 4 & 5,
Groundnut : TMV 3, 4, 7, 8 and Kadiri 71-1
Redgram : TM-1, ST-1
Chillies : G1 & G2
Ragi : AKP 1 to 7

43
 Differences between Mass and Pureline selections

S. No. Mass selection Pureline selection

1 Used both in self and cross pollinated crops Practiced in self pollinated crops only
2 Large number of plants are selected Comparatively less number of plants
are selected
3 The produce of the selected plants is mixed Produce of individual plants is kept
and sown as such in next year separate and progeny rows are raised
next year
4 No control of pollination Pollination is controlled
5 Variety developed is heterozygous and not Variety is homozygous homogeneous
uniform and uniform
6 Due to heterozygosity the variety deteriorates Due to homozygosity the variety lasts
quickly long
7 The method has to be repeated once in 2-3 No need to repeat
years to purify the variety
8 Wider adaptability due to heterozygosity Narrow adaptability due to
homozygosity
9 No knowledge of science is required. It is Knowledge of science and genetics is
more an art. required
10 Selection within a variety is effective Selection with in a pureline variety is
not effective
11 The variety is relatively difficult to identify It is relatively easy to identify in seed
certification programmes.

Lecture No 10
GENETIC BASIS OF PURE LINE

Self pollination increases homozygosity with a corresponding decrease in

heterozygosity. The effect of homozygosity and heterozygosity may be illustrated by taking
an individual heterozygous for (Aa) a single gene as follows

44
No of Frequency % Frequency %
generations AA Aa aa Homo- Hetero -
of selfing zygosity zygosity
1 0 100 0 0 100
2 25 50 25 50 50
3 37.5 25 37.5 75 25
4 43.75 12.5 43.75 87.5 12.5
5 46.875 6.25 46.875 93.73 6.25
6 48.437 3.125 48.437 96.874 3.125
7 49.218 1.562 49.218 98.436 1.562
8 49.608 0.781 49.608 99.216 0.781
9 49.803 0.39 49.803 99.606 0.39

Proportion of completely homozygous plants in the population

[(2m – 1) / 2m]n
m = No. of generations of self pollination
n = No. of genes segregating
Suppose an individual heterozygous for a single gene (Aa) and the successive
generations derived from it are subjected to self-pollination.
Every generation of self -pollination will reduce the frequency of heterozygote Aa to
50 per cent of that in the previous generation.
There is a corresponding increase in the frequency of the two homozygotes AA and
aa. As a result, after 10 generations of selfing, virtually all the plants in the population would
be homozygous, i.e., AA and a.
On the other hand, the frequency of heterozygote Aa would be only 0.097 per cent,
which is negligible. It is assumed here that the three genotypes AA, Aa and aa have equal
survival. If there is unequal survival, it may increase or decrease the rate at which
homozygosity is achie ved. If Aa is favoured, the rate of increase in homozygosity would be
lower than expected.
But if Aa is selected against, homozygosity would increase at a faster rate than
expected.

45
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.
4. Local varieties
5. Old pureline varieties and
6. Introduced varieties

General procedure for evolving a variety by pureline selection

The pureline selection has three steps.
4. Selection of individual plants from a local variety or some other mixed population.
5. Visual evaluation of individual plant progenies and
6. Yield trials
2. Selection
First year : A large number of plants (200-3000) which are superior than the rest are
selected from a local variety or mixed population and harvested separately (in some
cases individual heads or stems may be selected). The number of plants to be selected
depends upon the breeder’s discretion but should be as large as possible in view of the
available time, land, funds, labour etc. It is advisable to select for easily observable
characters such as flowering, maturity, disease resistance, plant height etc.
II. Evaluation :
Second year: Progenies of individual plants selected in 1st year are grown separately
with proper spacing (plant to row or head to row). The progenies are evaluated by
taking elaborate date on visual characters such as plant height, duration, grain type,
ear characters besides yield. The number of progenies should be reduced as much as
possible. Disease epiphytotics may be created to test the progenies for disease
resistance, poor, weak, diseased, insect attacked and segregating progenies are
rejected. The superior progenies are harvested separately. If necessary the process
may be repeated for one or more years.
III. Yield trials :
Third year : The selected progenies, now called as cultures are grown in replicated
trial for critical evaluation of yield etc. The best local variety is used as a check and

46
 should be grown at regular intervals, after every 15 or 20 cultures for comparision.
This is known as preliminary yield trial.
Superior cultures based on observable characters and yield are selected. The
number is drastically reduced.
Fourth & Fifth years : The superior cultures are tested against the local checks in
yield trials. Observations are recorded on many characters like diseases resistance,
days to flower, days to maturity, height of the plant ear characters, test weight and
yield. The data is subjected to statistical analysis to identify really superior cultures.
If necessary the trials may be extended for one more year or season. Inferior culture
are rejected and a few (4-5) promising cultures are selected.
Sixth, Seventh and Eighth years: The promising cultures selected are evaluated at
several locations along with strains or cultures of other breeders and local checks.
One or two promising cultures are selected.
Ninth year: The best progeny identified earlier is multiplied, named and released as a
variety for official release of any variety (approval from the variety releasing
committee of the state or central is necessary).

Advantage of pureline selection

1. The purelines are extremely uniform since all the plants in the variety will
have the same genotype.
2. Attractive and liked by the farmers and consumers.
3. Purelines are stable and long test for many years.
4. Due to its extreme uniformity the variety can be easily identified in seed
certification programmes.

Limitations or disadvantages of pureline selection

7. New genotypes are not created by pureline selection
8. Improvement is limited to the isolation of the best genotype present in
population. No more improvement is possible after isolation of the best
available genotype in the population.
9. Selection of purelines require great skill and familiarity with the crop.
10. Difficult to detect small differences that exist between cultures
11. The breeder has to devote more time
12. Pure lines have limited adaptability hence can be recommended for cultivation
in limited area only.

47
 Achievements :
Several varieties developed by pureline selection were released in many crops.
Some examples are given below
Rice : Mtu-1, Mtu-3, Mtu-7, Bcp-1, Adt-1, 3, 5, and 10
Sorghum : G 1 & 2, M 1 & 2, OO 1, 4 & 5,
Groundnut : TMV 3, 4, 7, 8 and Kadiri 71-1
Redgram : TM-1, ST-1
Chillies : G1 & G2
Ragi : AKP 1 to 7

Differences between Mass and Pureline selections

S. No. Mass selection Pureline selection
1 Used both in self and cross pollinated crops Practiced in self pollinated crops only
2 Large number of plants are selected Comparatively less number of plants
are selected
3 The produce of the selected plants is mixed Produce of individual plants is kept
and sown as such in next year separate and progeny rows are raised
next year
4 No control of pollination Pollination is controlled
5 Variety developed is heterozygous and not Variety is homozygous homogeneous
uniform and uniform
6 Due to heterozygosity the variety deteriorates Due to homozygosity the variety lasts
quickly long
7 The method has to be repeated once in 2-3 No need to repeat
years to purify the variety
8 Wider adaptability due to heterozygosity Narrow adaptability due to
homozygosity
9 No knowledge of science is required. It is Knowledge of science and genetics is
more an art. required
10 Selection within a variety is effective Selection with in a pureline variety is
not effective
11 The variety is relatively difficult to identify It is relatively easy to identify in seed
certification programmes.

48
Lecture No 11
BIOMETRICS – COMPONENTS OF GENETIC VARIATION

Biometry or biomatrics is the science thatdeals with the application of statistical procedures
to the study of biological problems.
Biometrical genetics or Quantitative genetics is that branch of genetics, which attempts to
unravel the inheritance of quantitative traits using statistical concepts and procedure.
INTRODUCTION :
The two basic requirements of Plant Breeding are the presence of genetic variation
and exploitation of this variation through selection. The selection of plants from a population
is almost always based on their phenotype. Phenotype has both heritable and nonheritable
components. The heritable component is due to the genes present in plants, that is genotype.
The non-heritable component consists of the effects of environment. The value of progeny
obtained from a selected plant, therefore, would largely depend upon therelative contributions
by the heritable and the non-heritable components to its phenotype., clearly, the breeder
should be thoroughly familiar with the laws of inheritance and the relative importance of the
genotype and the environment in determining the concerned phenotype.
Qualitative and Quantitative characters :
Some characters are little affected by other genes, i.e. the genetic back ground, or the
environment. Such characters are generally governed by one or few genes with large, easily
detecta ble effects, such genes are known as oligogenes. The characters produced by
oligogenes show distinct classes and are known as qualitative characters or olygogenic
traits. On the other hand, the development of many characters is very much affected by the
genetic background and, more particularly, by the environment. These characters are
governed by several genes with small individual effects; these genes one known as
polygenes. The characters produced by polygenes are referred to as quantitative
characte rs, because they do not show clear -cut classes and have to be studied by
measurement. They are also called Polygenic traits since they are governed by polygenes.
The inheritance of both qualitative and quantitative characters follows the laws of Mendel.
But the effects of individual genes in the two cases are totally different in magnitude
consequently, the techniques used to study the two types of characters are also different.

Role of the environment in Quantitative Inheritance

Quantitative characters are considerably affected by environment. The main result of this
effect is that the relationship between genotype and phenotype is partially or completely
hidden i.e. the phenotype doesnot reveal the genotype.

49
For eg: Phenotype = Genotype+ Environment P=G+E
If environment = 0 then phenotype = Genotype. However the effect of environment is
seldem zero. So phenotype is the joint action of Genotype and Environment.
In crop improvement, the breeder selects plants on the basis of their phenotype.
The effectiveness of selection depend on the proportion of phenotype due to the genotype.
Therefore, it is important to know the extent to which environment influences different
quantitative characters.
To estimate the effect of environment on a character, large No. of strain / genotypes are
grown in a replicated trial and the data is subjected to analysis of variance as per the
experimental design used.
The genotype x Environmental interaction signifies that the relative performance of various
genotype in effected by the environment. For eg: Performance of genotype ‘A’ may be
superior to the genotype ‘B’ in one environment but in another environment inferior to that of
‘B’
If G X E interaction is absent, genotype ‘A’ will be superior ‘B’ in all the environments.

BIOMETRICAL TECHNIQUES IN PLANT BREEDING

The biometrical techniques are useful to the plant breeders in the following 4
different ways.
1. Assessment of genetic variability present in the population. – It can be assesed by
Range, variance, standard deviatio n, coefficient of variation, D2 statistics, metro
glyph analysis
2. In the selection of elite genotypes from mixed populations – correlation, path and
discriminant function analysis)
3. Selection of parents and breeding procedures – diallel , partial diallel, line x tester,
generation means, triallel by parental cross and triple test cross analysis
4. Determining varietal adaptation – Stability analysis
Finley and Wilkinson (1963)
Eberhart and Russel (1966)
Perkins and Jinks (1968)
Freeman and Parkins (1971)
5. AMMI – Model – Additive main effects and multiplicative interaction
Components of Genetic variance :
Phenotypic variance = Genotypic variance + Environment variance
Vp = Vg +Ve

50
Fisher (1918) divided the genotypic variance into three components :
1. Additive
2. Dominance
3. Epistasis
Later Hayman and Mather partitioned the epistatic component into three types of
interactions – vig.,
Additive x additive ; additive x dominance and dominance x dominance
1. Additive variance : Arises of from difference between two homozygotes for a gene
i.e. AA and aa. It is generally represented ‘d’
2. Dominance variance or Intra allelic interaction: It is due to the deviation of
heterozygote (Aa) phenotype from the average of phenotypic values of the two
homozygotes (AA and aa) it is represented by ‘h’.
3. Epistasis variance or Inter allelic Interaction : Results from an interaction between
two or more genes represented by ‘e’.
Difference between additive and dominance variance
It refers to difference between homozygotes It refers to deviation of Aa from the mean of
(AA/aa) AA and aa
Genes show lack of dominance Genes show incomplete, complete or over
dominance
Associated with homozygosity and is more in Associated with heterozygosity and is more
inbreders in outbreeders
It is fixable It is non-fixable
Selection is very effective as it is fixable Selection is ineffective as it is non-fixable
it is the chief cause of transgressive It is the chief cause of heterosis
segregation

Different types of gene actions involving two genes

Component of Specific Description General
genetic variance symbol symbol
Additive da The different between AA and aa phynotypic d
values
db The different between BB and bb phynotypic
values
Dominance ha The deviation of Aa phenotype from the average h
of AA and aa phenotypes
hb The deviation of Bb phenotype from the averave
of BB and bb phenotypes
Epistasis da x db Additive x Additive effect due to interaction i
between AA & BB
da x hb Additive x dominance interaction due to j
and interactions between and AA and Bb and between
ha x db Aa and BB respectively
ha x hb Dominance x dominance interaction due to l
interation between Aa and Bb

51
Lecture No 12
HYBRIDIZATION : TECHNIQUES AND CONSEQUENCES

The mating or crossing of two plants or lines of dissimilar genotype is known as

hybridization. In plants, crossing is done by placing pollen grains from one genotype, the
male parent, on to the stigma of flowers of the other genotype, the female parent. It is
essential to prevent self-pollination as well as chance cross-pollination in the flowers of the
female parent. At the same time, it must be ensured that the pollen from desired male parent
reaches the stigma of female flowers for successful fertilization. The seeds as well as the
progeny resulting from the hybridization are known as hybrid or F1. The progeny of F1,
obtained by selfing or intermating of F1 plants, and the subsequent generations are termed as
segregating generations. The term cross is often used to denote the products of hybridization,
i.e. the F1 as well as the segregating generations.

OBJECTIVES OF HYBRIDIZATION
The chief objective of hybridization is to create genetic variation. When two
genotypically different plants are crossed, the genes from both the parents are brought
together in F1. segregation and recombination produce many new gene combinations in F2
and the later generations, i.e. the segregating generations. The degree of variation produced
in the segregating generations would, therefore, depend on the number of heterozygous genes
in the F1. This sill, in turn, depend upon the number of the genes for which the two parents
differ. If the two parents are closely related, they are likely to differ for a few genes only.
But if they are not related, or are distantly related, they may differ for several, even a few
hundred, genes. However, it is not likely that the two parents will ever differ for all their
genes. Therefore, when it is said that the F1 is 100 per cent heterozygous, it has reference
only to those genes for which the two parents differ.
The aim of hybridization may be the transfer of one or few qualitative characters, the
improvement in one or more quantitative characters, or use the of 1 as a hybrid variety. These
objectives are briefly discussed below.
Combination Breeding : The main aim of combination breeding is the transfer of one or
more characters into a single variety from other varieties. these characters may be governed
by oligogenes or polygenes. The intensity of the character in the new variety is either

52
comparable to or, more generally, lower that in the parent variety from which it was
transferred. In this approach, increase in the yield of a variety is obtained by correcting the
weaknesses in the yield contributing traits, e.g., tiller number , grains per spike, test weight is
that for disease resistance. The backcross method of breeding was designed for combination
breeding, and often pedigree method also fulfils the same purpose. In combination breeding,
the genetic divergence between parents is not the major consideration. What is important is
that one of the parents must have in a sufficient intensity the character(s) under transfer,
while the other parent is generally a popular variety.
Transgressive Breeding : Transgressive breeding aims at improving yield or its
contributing characters through transgressive segregation. Transgressive segregation is the
production of plants in an F2 generation that are superior to both the parents for one or more
characters. Such plants are produced by an accumulation of plus or favourable genes from
both the parents as a must combine well with each other, and should preferably be genetically
diverse, i.e., quite different. This way, each parent is expected to contribute different plus
genes which when brought together by recombination give rise transgressive segregant. As a
result, the intensity of character in the transgressive segregant, i.e., the new variety, is greater
than that in either of the parents. The pedigree method of breeding and its modifications,
particularly the population approach, are designed for the production of transgressive
segregants.
Hybrid Varieties : In most self-pollinated crops, F1 is more vigorous and higher yielding
than the parents. Wherever it is commercially feasible, F1 may be used directly as a variety.
In such cases, it is important that the two parents should produce an outstanding F 1.
TYPES OF HYBRIDIZATION
The plants or lines involved in hybridization may belong to the same variety, different
varieties of the same species, different species of the same genus or species from different
genera. Based on the taxonomic relationship of the two parents, hybridization may be
classified into two broad groups :
1. Intervarietal and
2. Distant hybridization
Intervarietal Hybridization : The parents involved in hybridization belong to the same
species ; they may be two strains, varieties or races of the same spicies. It is also known as
intraspecific hyrbidization. In crop improvement programmes, intervarietal hybridization is
the most commonly used. In fact, it is so common that it may often appear to be the only
form of hybridization used in crop improvement. an example would be crossing of two

53
varieties of wheat, rice or some other crop. The intervarietal crosses may be simple or
complex depending upon the number of parents involved.
Simple Cross : In a simple cross, two parents are crossed to produce the F1. The F 1 is selfed
to produce F2 or is used in a backcross programme, e.g.,
A X B à F1 (A X B)
Complex Cross : more than two parents are crossed to produce the hybrid, which is then
used to produce F2 or is used in a backcross. Such a cross is also known as convergent cross
because this crossing programme aims at converging, i.e., bringing together, genes from
several parents into a single hybrid. A few examples of convergent cross are described in
Fig. 7.1. As
Three Parents (A, B, C)
AXB

F1 (A X B) X C

Complex hybrid (A X B) X C
FOUR Parents (A, B, C, D)
AXB CXD

F1 (A X B) X (C X D)

Complex hybrid (A X B) X (C X D)
Eight Parents (A, B, C, D, E, F, G, H)
AXB CXD EXF GXH

F1 (A X B) X (C X D) (E X F) X (G X H)

[(A X B) X (C X D)] X [(E X F) X (G X H)]

Complex hybrid [(A X B) X (C X D)] X [ (E X F) X (G X H)]

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 Fig 7.1. Complex crosses involving 3, 4 and 8 parents.

Crop improvement progresses, the crop varieties would accumulate more and more
favourable genes. This would lead to greater similarities between even unrelated varieties. In
view of this, it may be expected that in future complex crosses would become more and more
important. In breeding of highly improved self-pollinated crops like wheat and rice, complex
crosses are a common practice today. Complex crosses would become routine in near future
in the improvement of other self-pollinated crops with the progress in the level of their
improvement.
Distant Hybridization : Distant hybridization includes crosses between different species of
the same genus or of different genera. When two species of the same genus are crossed, it is
known as interspecific hybridization; but when they belong to two different genera, it is
termed as intergeneric hybridiztion. Generally, the objective of such crosses is to transfer one
or few simply inherited characters like disease resistance to a crop species. Sometimes,
interspecific hybridization may be used for developing a new variety, e.g., Clintonoat variety
was developed from a cross between Avena sativa x A. byza ntina (both hexaploid oat
species), and CO 31 rice variety was developed from the cross Oryza sativa var. indica x O.
perennis. Almost al the present-day sugarcance varieties have been developed from complex
crosses between Saccharum officinarum (noble canes), S. barberi (Indian canes) and other
Saccharum species, e.g., S. spontaneum (Kans.). The improvement in fiber length of Indian
Cotton (Gossypium arboreum) has been brought about by crossing it with American
cultivated Cotton ; many improved varietie s have resulted from such crosses. Intergeneric
hybridization may also be used to develop a new crop species, e.g., Triticale from a cross
between Triticum sp. And Secale cereale (rye). Wild species often provide genes which are
not present in the cultivated species. For example, many of the genes for rust resistance in
wheat are derived from related wild species. Distant hybridization is likely to become
increasingly important in the correction of specific defects of crop species. In many cases,
wild species may contribute valuable ‘yield genes’ as well to the cultivated species.
Pre -requisites for hybridization
Breeder should have clear knowledge about the following before taking up
hybridization.
1. Requirements of the tract
2. Local conditions i.e. soil, climate, Agronomic practices and market requirements
3. Existing varieties of crops both local and introduced
4. Facilities like funds, land, labour and equipment

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 5. Plant material i.e. germ plasm
6. Objectives : Well set objectives and planning
Hybridization proce dure or steps involved in hybridization
Details of the following steps have to be covered in
Practical classes
1. Choice or selection of parents
2. Evaluation of parents i.e. by selfing and studying the progeny
3. Emasculation
4. Crossing or pollination
5. Bagging & Labelling
6. Harvesting of F1 seed
7. Raising F 1 generation
From F2 onwards the generations are known as segregating generations and they may be
handled either by pedigree method of Bulk method or backcross method for evolving new
varieties.

Lecture No 13
HANDLING OF SEGREGATING GENERATIONS
Pedigree Method
In the pedigree method, individual plants are selected from F 2 and subsequent
generations, and their progenies are tested. During the entire operation a record ofall parent
off spring relationships is kept. This is known as pedigree record. Individual plant selection
is continued till the progenies show no segregation. At this stage the selection is done among
the progenies, multilocation tests are conducted and released as varieties.
The pedigree may be defined as a description of the ancestors of an individual and it
generally goes back to some distant ancestors. It is useful to know the relationship of two
individuals and useful for selection of parents and prediction of outcome of the cross.
Procedure of pedigree method
1 st year : cross is made between the parents possessing desirable characters.
2 nd year : Sow the F1 seed giving wide spacing so that each F1 plant produces more seeds.
Raise as many F1 plants as possible to produce large number of F2 seeds. Harvest in bulk.
3 rd year : Grow 2000-10000 plants of F2 giving wide spacing for full expression of the
characters in F2 generation plants. Grow parents for comparision. Depending upon the
facilities and objectives of the programme about 100-500 superior plants are selected. The

56
value of selection depend on the skill of the breeder. He has to judge which F2 plant will
produce superior progeny for characters under consideration. The breeder develops this skill
through close study of the crop for many generations. The selection in F2 is done for simply
inherited characters like head type disease resistance etc. and selection for characters
governed by many genes like yield will be reserved for later generations. The selected plants
are harvested separately and given serial numbers and description entered in pedigree
registers.
4 th year : Progeny rows of F3 i.e. seeds of one selection plant in one row are space planted
along with parents and checks. From superior progeny rows, individual plants with desirable
characters are selected (about 50-100 families and about 5 plants in each family and
harvested separately). Diseased, lodging and undersirable progenies are discarded.
5 th year : F 4 plants raised again as head to row. Desirable plants are selected from desirable
rows and harvested separately.
6 th year : F 5 plants raised in 3 row plots i.e. seeds of each selected plant sown in 3 rows. By
this time many families might have become reasonly homozygous. For comparision check
variety is grown for every 3 or 5 block. Progenies are evaluated for yield and the inferior
ones are rejected. The number should be reduced to 25-50. superior plants from superior
progenies are selected. Plants from each progeny are bulked.
7 th year : F 6 individual plant progenies are grown in multi-row plots and evaluated. Inferior
progenies are rejected and superior progenies are selected. Plants of each progeny are
harvested in bulk. Diseased and inferior plants from the progenies are removed.
8 th year : F 7 preliminary yield trial with 3 or more replications ar conducted to identify
superior lines. The progenies are evaluated for many characters including yield. Standard
commercial varieties must be included as checks. Two to five outstanding lines are selected
and advanced to coordinated yield trials.
9 th, 10 th & 11 th year : selected lines are tested in several localities for 2 or 3 years for
adaptation tests. Lines are evaluated for all characters mainly yield and disease resistance. A
line that is superior to commercial variety in yield and other characters is selected. 11th and
12 th year : Selected superior lines is named, multiplied and released as a new variety.
Number of year can be reduced if generations are advanced during off seasons either in green
house or under irrigated conditions.
Several modifications for the above described pedigree method are followed by
breeders depending upon the crop, time and availability of funds and facilities like labour,
land etc.

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Early generation tests :
The objective of these test is to find out superior crosses and superior progenies in
early generations i.e. in F2 and F3. we need not advance all the crossed and all selected
progenies in each cross upto F8. much labour, time and cost would be saved by this early
generation testing. A more reliable information about the potential crosses and progenies
may be obtained by conducting replicated tests (preferably in more location) and evaluating
them for yield and other characters in F2 or F3 itself. A desirable cross or progeny should
have high mean yield, high genetic variance and high expected genetic advance under
selection. Other crosses and progenies are rejected in the beginning i.e. F2 and F3 generations
itself.
F2 progeny testing : Another modification for pedigree method. In F2 make as many single
plants selections as possible. From F3 to F6 advance the progenies in bulk making selections
of the progenies as a wholw and discarding the inferior progenies. Thus each of the progeny
is derived from the single plant selected in F2 generation. In F6 make single plant selections
in each of the progeny. Compare the yields of the single plants with progenies from which
they are selected. Select superior single plant progenies and advance to preliminary yield
trials, multilocation tests etc. There are two advantages 1. No. of crosses can be handled
simultaneously 2. Natural selection operates from F 3 to F 6 since they are advanced in bulk.
Mass pedigree method : This is another modified pedigree method. Crosses are made and
further generations grown in bulk or as mass until suitable season occurs for making desirable
selections against drought, insect and diseases etc. The population will be exposed to the
natural conditions of vagaries. From the remaining population individual plants are selected
and harvested progenies are evaluated for yield and other characters in preliminary yield
trials and further generations are proceeded as in pedigree method till release of variety. The
advantages of both bulk and pedigree methods can be obtained and large number of crosses
can be handled at a time. The disadvantage is that it takes a bit longer time.
Merits of pedigree method :
1. It gives maximum opportunity for the breeder to use his skill and judgement in
selection of plants
2. It is well suited for the improvement of characters which can be easily identified and
are simply inherited.
3. Transgressive segregation for yield and other quantitative characters may be
recovered.
4. Information about the inheritance of characters and pe digree of lines can be obtained.

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 5. Inferior plants and progenies are eliminated in early generations.
6. It takes less time than bulk method to develop new variety.
Demerits of pedigree method :
1. Valuable genotypes may be lost in early generations, if sufficient skill and knowledge
are lacking in the breeder, at the time of selection.
2. No opportunity for natural selection
3. Difficult to handle many crosses
4. Maintenance of records, selections, growing progeny rows etc are time consuming
and laborious.
Achievements : Large number of varieties have been developed by pedigree method in
many crops.
A few examples are
Wheat – NP-52, 120,125, 700 and 800 series
Rice – ADT – 25, Jaya, Padma
Cotton – Lakshmi, Digvijay, Sorghum – Co 18, RS 610 etc., Tobacco – NP 222
Sorghum – Co 18, Rs 610, Tobacco – NP 222
Lecture No. 14
BULK METHOD
The bulk method was first proposed by Nilsson Ehle in 1908 at Svalof. This method
is also known as mass method ‘or’ Population method of breeding
v Isolation of Homozygus lines
v Waiting for the opportunity for selection
v Opportunity for natural selection.
v F2 and subsequent generations are harvested in mass as bulk to raise the next
generation.
v At the end of the bulking period (after attaining homozygosity) individual plants are
selected and evaluated similar manner as pedigree method of breeding.

THE PROCEDURE FOR BULK METHOD

The exact procedure for the bulk method would vary depending upon the objective of
breeder. The following procedure is described for the isolation of homozygous lines. The
breeder may introduce various modifications in the scheme to suit his needs.
Hybridization : Parents are selected according to the objective of the breeding programme.
A simple or a complex cross is then made depending upon the number of parents involved.

59
F1 Generation : F 1 is space-planted and harvested in bulk. The number of F1 plants should
be as large as possible ; usually more than 20 plants should be grown.
F2-F 6 Generations : F2 to F6 generations are planted at commercial seed rates and spacings.
These generations are harvested in bulk. During this period, environmental factors, disease
and pest outbreaks would change, the frequencies of different genotypes in the population.
Artificial selection is generally not done. The population size should be as large as possible,
preferably 30,000-50,000 plants in each generation.
F7 Generations : About 30-50 thousand plants are space-planted. 1000 to 5000 plants with
superior phenotypes are selected and their seeds harvested separately. Selection is based on
the phenotype of plants, grain characteristics, disease reaction, etc.
F8 Generation : Individual plant progenies are grown in single or multi-row plots. Most of
the progenies would be reasonably homozygous and are harvested in bulk. Weak and inferior
progenies are rejected on the basis of visual evaluation. Only 100-300 plant progenies with
desirable characteristics are saved.
Some progenies which show segregation are generally rejected unless they are of
great promise. In promising pr ogenies, individual plants may be selected ; preliminary yield
trial will be delayed for one year in such cases.
F9 Generation : Preliminary yield trial is conducted by using standard commercial varieties
as checks. The progenies which are superior than the check are advanced. Quality test may
be conducted to further reject undesirable progenies. The progenies are evaluated for height,
lodging resistance, maturity date, disease resistance and other important characteristics of the
crop species.
F10-F13 Generations : Replicated yield trials are conducted over several locations using
standard commercial varieties as checks. The lines are evaluated for important characteristics
in addition to yield, disease resistance and quality. If a line is superior to the standard
varieties in yield trials, it would be released as a new variety.
F14 Generation : Seed of the released variety is increased for distribution to the cultivators.

MERITS OF BULK METHOD

1. The bulk method is simple, convenient and less expensive.
2. Since, each F2 plant is equally represented till F6, no chance of elimination of good
genotypes in early generations.

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 3. Artificial or natural disease epiphytics, winter killing high temperature etc. eliminates
undesirable types and increases the frequency of desirable type. Thus isolation of
desirable types becomes easier.
4. Progenies select from long term bulks are superior than the selection from F2 or short
term bulk.
5. Since, little work and attention is needed in F2 and subsequent generation more no. of
crosses can be handled.
6. No pedigree records which saves time
7. Since large population are grown, transgressive segregants are more likely to appear
and increase due to natural selection. Hence, there is a greater chance to isolate good
segregants than pedigree method.
DEMERITS OF BULK METHOD
1. The major disadvantage of bulk method is that it takes a much longer time to develop
a new variety. Natural selection becomes important only after F8 or F10, and bulking
may have to be done upto F20 or more. Thus the time required is considerably longer,
and most breeders do not use the bulk method simply for this reason.
2. In short-term bulks, natural selection has little effect on the genetic composition of
populations. But short -term bulks are useful for the isolation of homozygous lines
and for specific objectives as in Harlan’s mass -pedigree method.
3. It provides little opportunity for the breeder to exercise his skill or judgement in
selection. But in the modified bulk method, the breeder has ample opportunity for
practicing selection in the early segregating generations.
4. A large number of progenies have to be selected at the end of the bulking period.
5. Information on the inheritance of characters cannot be obtained which is often
available from the pedigree method.
6. In some cases, at least, natural selection may act against the agronomically desirable
types.
Comparison between bulk and pedigree method.
S. No. PEDIGREE METHOD S. No. BULK METHOD
1 Most widely used Breeding method 1 Used only to a limited extent
2 Individual plants are selected in F2 2 F 2 and subsequent generations are
and subsequent generations and grown in bulk
individual plant progenies are grown
3 Artificial selection ; artificial disease 2 Mainly natural selection. In certain
epidemics etc. are an integral part of cases artificial selection may be
the method essential

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 3 Natural selection does not play any 3 N.S. determines the composition of
role the pop n at the end of the bulking
period
4 Pedigree Records have to be 4 No pedigree records are maintained
maintained which is often time
consuming and laborious
5 Generally its taken 12-13 years to 5 Takes more than 15 years.
release a new variety
6 Requires close attention of breeder 6 It is quite simple and does not
from F2 onwards require much attention
7 Planting (spacing) the segregating 7 The bulk populations are generally
generations are space planted to planted at commercial planting
permits effective individual plant rates
selection
8 Population size is small in 8 The population size is large
comparison to bulk

Much improvement in crop plants could not be done through this method reason
being.
1. Long time required for Natural Selector
2. Lack of opportunity for the breeder to use his skills
3. Lack of facilities to raise large population
Achievements of bulk method:
The method has been used to a limited extent is Barley breeding in U.S.A. and more
than 50 varieties were developed. They are : ARIVAL, BEECHER, GLACIER, and GEM.
Originated from a cross : Atlas x Vaughn. The bulk was maintained for 7 to 8 months.

SINGLE-SEED-DESCENT METHOD
Another modification of the bulk method is the single-seed-descent method, which is
becoming increasingly popular. In this method, a single seed from each of the one to two
thousand F2 plants is bulked to raise the F3 generation. Similarly, in F3 and the subsequent
generations one random seed is selected from every plant present in the population and
planted in bulk to raise the next generation. This procedure is followed till F5 or F6 when the
plants would have become nearly homozygous. In F5 or F6, a large number (1 to 5 hundred)
of individual plants are selected and individual plant progenies are grown in the next
generation. Selection is done mainly among the progenies, and the number of progenies is
sufficiently reduced to permit replicated trial in the next generation. Individual plants may
be selected only from outstanding families not showing segregation. Thus preliminary yield
trials and quality tests begin in F7 or F8 and coordinated yield trials in F8 or F 9.

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 The objective of single-seed-descent method is to rapidly advance the generations of
crosses ; at the end of the scheme, a random sample of homozygous or near homozygous
genotypes/lines is obtained. F2 and the subsequent generations are grown at very high plant
densities as vigour of individual plants is not important. In each year, 2-3 generations may be
raised using off-season nurseries and greenhouse facilities. The important features of this
scheme are : (1) lack of selection, natural or artificial, till F5 or F6 till the population is
reasonably homozygous , and (2) raising of F3 and later generations from a bulk of one seed
from each F2 and the subsequent generation plant in order to ensure that each F2 plant is
represented in the population. As a result of the speed and economy, the single-seed-descent
scheme is becoming increasingly popular with the breeders.
The single-seed-descent scheme (1) advances the generation with the maximum
possible speed in a conventional breeding method; (2) requires very little space, effort and
labour ; (3) Makes the best use of greenhouse and off -season nursery facilities; and (4)
ensures that the plants retained in the end population are random sample from the F2
population. However, (1) it does not permit any form of selection (which is implied in the
scheme) during the segregating generations; and (2) in each successive generation, the
population size becomes progressively smaller due to poor germination and death of plants
due to diseases, insect pests and accidents. In some crops, e.g., pulses, plant loss may be one
of the most serious problems of the scheme.

Lecture No. 15
BACK CROSS

Breeders of early 20th century engaged in the development of disease resistant

varieties observed that pureline selections with genes for resistance from intra-or inter-
specific hybridization were inferior to the generally acceptance superior parent in yield or
quality characteristic. To overcome this problem, (Harlan and Pope (1922) suggested the
back cross method by which an undesirable allele at a particular locus is replaced by the
desirable allele in otherwise elite variety. In other words, B.C. procedure conserves all good
characteristics of a popular adapted variety and incorporates a desirable character from
another variety.
Back cross : A cross between a hybrid (F 1 or a segregating generation) and one of its parents
is known as backcross.

63
Back cross method : In the B.C. method, the hybrid and the progenies in the subsequent
generations are repeatedly back crossed to one of their parents.
Objective : To improve or correct one or two specific defects of a high yielding variety,
which is well adapted to the area and has other desirable characteristics.
Recipient parent : Well adapted, high yielding variety, lacking one or two characters and
hence receives these genes from other variety.
Donor parent : The variety which donates one or two useful genes.
Recurrent parent : Since the recipient parent is repeatedly used in the backcross
programme, it is also known as the recurrent parent.
Non-recurrent parent : The donor parent, on the other hand, is known as the non-recurrent
parent because it is used only once in the breeding programme (for producing the F1 hybrid).

REQUIREMENTS OF A BACK CROSS PROGRAMME

1. Existence of a good recurrent parent variety which requires improvement is some
qualitatively inherited character or a quantitative character with high heritability.
2. A suitable donor parent must be available possessing the character or characters to be
transferred in a highly in tense form.
3. High expressivity of the character under transfer through several back crosses in the
genetic back ground of the recurrent parent.
4. The character to be transferred must have high heritability-preferably determined by
one or few genes.
5. Simple testing technique for detecting the presence of the character under transfer.
6. Recovery of the recurrent genotype in a reasonable number of back cross generations.
Applications of Back Cross method
B.C. method in applicable to both S.P. & C.P. crops.
1. Inter varietal transfer of simply inherited characters : characters governed by one or
two major genes – Eg. disease resistance, used color.
Linkage drag : Failure of transfer of simply inherited characters like disease resistance
by B.C. method due to a tight linkage between the gene being transferred and some
other undesirable gene.
2. Inter varietal transfer of Quantitative characters : Quantitative characters with high
heritability can be transferred.
Eg. Early ness, Pl. height seed size, seed shape.

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 3. Inter specific transfer of simply inherited characters : Mostly disease resistance from
related species into a cultivated species.
Eg. 1. Leaf and stem rust resistance from Triticum timopheevii
T. monococcum, Aegilops speltoides and rye (S. cereale ) to T. aestivum
2. Black arm resistance from several Gossypium species to G. hirsutum
4. Transfer of cytoplasm : Back Cross methodin used to transfer cytoplasm from one
variety or species to another. This is especial desirable in cases of Cytoplasmic or
Cytoplasmic-genetic male sterility.
E. Transfer of T. timopheevii cytoplasm to T. aesticem
5. Transressive segregation : Back cross method may be modified to obtain transgressive
segregants. It may be modified is one of the following two ways.
I. The F1 may be Back crossed only 1 or at most 2 times to the recurrent parent
leaving much heterozygosity for transgressive segregants to appear.
II. To or more recurrent parents may be used in the back cross programme to
accumulate genes from them in to the back cross progeny. Such a modification of the
back cross would produce a new variety that would not be exactly like any one of the
recurrent parents.
6. Production of Isogenic lines : Isogenic lines are identical in their genotype, except for
one gene. Such lines are useful in studying the effects of individual genes on yield
and other characteristics. Isogenic lines and easily produced using the back cross
method.
7. Germplasm conversion : Conversion of photosensitive germplasm lines (using as
recurre nt parent) to photo insensitive line (using a photo insensitive line as a donar or
non-recurrent parent.
Transfer of a Dominant Gene
Let us suppose that a high yielding and widely adapted variety A is susceptible to
stem rust. Another variety B is resistant to stem rust, and that resistance to stem rust is
dominant to susceptibility. A generalized scheme of the backcross programme for the
transfer of rust resistance from variety B to variety A is given below.
Hybridization : Variety A is crossed to varie ty B. Generally, variety A should be used as
the female parent. This would facilitate the identification of selfed plants, if any.
F1 Generation : F1 plants are backcrossed to variety A. Since all the F1 plants will be
heterozygous for rust resistance, selection for rust resistance is not necessary.

65
First Backcross Generation (BC1) : half of the plants would be resistant and the remaining
half would be susceptible to stem rust. Rust resistant plants are selected and backcrossed to
variety A. BC 1 plants resistant to rust may be selected for their resemblance to variety A as
well.
BC2-BC5 Generations : In each backcross generation, segregation would occur for rust
resistance. Rust resistant plants are selected and backcrossed to the recurrent parent A.
Selection for the plant type of variety A may be practiced, particularly in BC2 and BC3.
BC6- Generation : On an average, the plants will have 98.4 per cent genes from variety A.
Rust resistant plants are selected and selfed; their seeds are harvested separately.
BC6 F 2 Generation : Individual plant progenies are grown. Progenies homozygous for rust
resistance and similar to the plant type of variety A are harvested in bulk. Several similar
progenies are mixed to constitute the new variety.
Yield Tests : The new variety is tested in a replicated yield trial along with the variety A as a
check. Plant type, date of flowering, date of maturity, quality etc. are critically evaluated.
Ordinarily, the new variety would be identical to the variety A in perfor mance. Detailed
yield tests are, therefore, generally not required and the variety may directly be released for
cultivation.
Transfer of a Recessive Gene
When rust resistance is due to a recessive gene, all the backcrosses cannot be made
one after the ot her. After the first backcross, and after every two backcrosses, F2 must be
grown to identify rust resistant plants. The F1 and the backcross progenies are not inoculated
with rust because they would be susceptible to rust. Only the F2 is tested for rus t resistance.
A generalized scheme for the transfer of a recessive gene for rust resistance is given below.
Hybridization : The recurrent parent is crossed with the rust resistant donor parent. The
recurrent parent is generally used as the female parent.
F1 Generation : F1 plants are backcrossed to the recurrent parent.
BC1 Generation : Since rust resistance is recessive, all the plants will be rust susceptible.
Therefore, there is no test for rust resistance. All the plants are self-pollinated.
BC1 F2 Generation : Plants are inoculated with rust spores. Rust resistant plants are selected
and backcrossed with the recurrent parent. Selection is done for the plant type and other
characteristics of the variety A.
BC2 Generation :There is no rust resistanc e test. Plants are selected for their resemblance to
the recurrent parent A, and backcrossed with the recurrent parent.

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 BC3 Generation : There is no disease test. The plants are self-pollinated to raise F2.
selection is usually done for the plant type of variety A.
BC3F2 Generation : Plants are inoculated with stem rust. Rust resistant plants resembling
variety A are selected and backcrossed to variety A. Selection for plant type of A is generally
effective.
BC4 Generation : There is no rust resistance test. Plants are back-crossed to variety A.
BC5 Generation : There is no rust test. Plants are self -pollinated to raise F2 generation.
BC5F2 Generation : Plants are subjected to rust epidemic. A rigid selection is done for rust
resistance and for the characteristics of variety A. Selfed seeds from the selected plants are
harvested separately.
BC5F3 Generation : individual plant progenies are grown and subjected to rust epiphytotic.
A rigid selection is done for resistance to stem rust and for the characteristics of variety A.
Seeds from several similar rust resistant homogeneous progenies are mixed to constitute the
new variety.
Yield Tests : It is the same as in the case of transfer of a dominant gene.

Lecture No: 16
BACK CROSS

Transfer of Two or Mo re Characters Into a Single Recurrent Parent

When two of more characters are to be transferred into the same variety, one of the
following three approaches may be used.
Simultaneous Transfer : Genes for the different characteristics may be transferred
simultaneously in the same backcross programme. The characters to be transferred are
bought together into the hybrid by successively crossing each of the non-current parents to
the recurrent parent or the hybrid thus produced. But in such a case, a larger backcross
population would be needed than in the case of transfer of a single character. Further, the
breeding programme may be delayed because the conditions necessary for the selection of all
the characters may not occur every year. Sometimes, the two genes under transfer may be
linked. In such a case, the transfer becomes very easy, and selection for only one gene may
be necessary. some examples of such a favourable linkage are; between the genes Lr 24 and
Sr 24, Lr 19 and Sr 25, and Lr 26 and Sr 31.
Stepwise Transfer : The recurrent parent is first improved for one character. The improved
recurrent parent is then used as the recurrent parent in a backcross programme for the transfer
of the second character. If additional characters are to be transferred, they are transferred one

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at a time in a stepwise fashion. This approach takes much longer time for the transfer of two
or more characters.
Simultaneous But Separate Transfers : Each character is transferred to the same recurrent
parent in simultane ous but separate backcross programmes. The resulting improved versions
from the different programmes are then crossed together. Homozygous lines for the
characters being transferred are then selected from the segregating generations using the
pedigree method. This approach appears to be the most suitable of the three strategies.
Merits
1. The genotype of new variety is nearly identical with that of the recurrent parent,
except for the genes transferred. Thus the outcome of a backcross programme is
known beforehand and it can be reproduced any time in the future.
2. It is not necessary to test the variety developed by the backcross method in extensive
yield tests because the performance of the recurrent parent is already known. This
may save upto 5 years’ time and a considerable expense.
3. The backcross programme is not dependent upon the environment, except for that
needed for the selection of the character under transfer. Therefore, off -season
nurseries and green-houses can be used to grow 2-3 generations each year. This would
drastically reduce the time required for developing the new variety.
4. Much smaller populations are needed in the backcross method than in the case of
pedigree method.
5. Defects, such as susceptibility to disease, of a well-adapted variety can be removed
without affecting its performance and adaptability. Such a variety is often preferred
by the farmers and the industries to an entirely new variety because they know the
recurrent variety well.
6. This is the only method for inter-specific gene tra nsfers, and for the transfer of
cytoplasm.
7. It may be modified so that transgressive segregation may occur for quantitative
characters.
Demerits
1. The new variety generally cannot be superior to the recurrent parent, except for the
character that is transferred.
2. Undesirable genes closely linked with the gene being transferred may also be
transmitted to the new variety.

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 3. Hybridization has to be done for each backcross. This often difficult, time taking and
costly.
4. By the time the backcross programme improves it, the recurrent parent may have been
replaced by other varieties superior in yielding ability and other characteristics.

Achievements
1. Two cotton varieties 170-C o-2 and 134 – Co 2m were developed
2. Kalyana sona susceptible to leaf rust. Resistant has been transferred from several
diverse sources i.e., Robin, K1, Blue bird, Tobari, Frecor and HS-19
3. Tift 23A is susceptible to downy mildew. The line backcrossed with MS-521A, MS-
541 A, MS-570A resistant hybrids were produced.

Comparison between backcross and pedigree methods

Pedigree method Backcross method

F1 and the subsequent generations are F 1 and the subsequent generations are
allowed to self-pollinate backcrossed to the recurrent parent

The new variety developed by this method is The new variety is identical with the
different from the parents in agronomic and recurrent parent, except for the character
other characteristics under transfer

The new variety has to be extensively tested Usually extensive testing is not necessary
before release before release

The method aims at improving the yielding The method aims at improving specific
ability and other characteristics of the variety defects of a well adapted, popular variety

It is useful in improving both qualitative and It is useful for the transfer of both
quantitative characters quantitative and qualitative characters
provided they have high heritability

It is not suitable for genes transfer from It is the only useful method for gene transfers
related species and for producing substitution from related species and for producing
of addition lines addition and substitution lines

Hybridization is limited to the production of Hybridization with the recurrent parent is

the F1 generations necessary for producing every backcross
generation

The F1 and the subsequent generations are The backcross generations are small and

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much larger than those in the backcross usually consist of 20-100 plants in each
method generation

The procedure is the same for both dominant The procedures for the transfer of dominant
and recessive genes and recessive genes are different

Multiline variety are mixtures of several purelines of similar height, flowering and maturity
dates, seed colour and agronomic characters of each of which has a different gen for
resistance to the given disease.
Characteristics of a good Multiline
1. Its genetic diversity for vertical resistance genes for the concerned disease
2. The vertical resistance genes should be strong enough
3. It should have normal resistance to othe r diseases
4. Components of multiline should be uniform for agronomic and other features.
5. It should have yield advantage
Development of multiline varieties
A multiline variety is usually created by mixing the seeds of several lines that are
similar in appearance but have different genes for resistance to a given disease. There are two
main steps in the development of multilines:
1. Development of component lines
2. Evaluation and grouping of the components.
Development of component lines
The resistance genes are incorporated in an elite variety or line to produce as many
near-isogenic lines as there are distinct R genes. This is done through a conventional
backcross programme (5-6 backcrosses), a limited backcrossing (2-3 backcrosses, followed
by pedigree selection) or by making double or multiple crosses. The lines obtained from the
last two approaches are likely to differ for agronomic and other features as well; therefore, a
detailed evaluation of such lines is essential.
Evaluation and grouping of the components
The number of component lines should be large, 15-20 according to Borlaug (1959),
if durability of resistance is desired. But if a reduced level of disease is the objective, a rather
small number of component lines would be adequate.
Achievements
Multiline variety appears to be a useful approach to control disease like rusts where
new races are continuously produced. In India, four multiline varieties have been released in

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wheat. Kalyan Sona and Sonalika, one of the most popular varieties during the late sixties
were used as the recurrent parent to produce these varieties. Variety ‘KSML3’ consists of 8
lines having rust resistance genes from Robin, Ghanate, K1, Rend, Gabato, Blue Bird, Tobari,
etc. Multiline ‘MLKS11’ is also a mixture of 8 lines ; the resistance was derived from E6254,
E6056, E5868, Frecor, HS19, E4894, etc. The third variety, KML 7406, has 9 lines deriving
rust resistance from different sources. In addition, Sonalika Multiline-1 was released for
cultivation in Punjab state (six component lines).

Lecture No. 17
SELF-INCOMPATIBILITY
More than 300 species belonging to 20 families of angiosperms show self-
incompatibility. Self -incompatible pollen grains fail to germinate on the stigma of the flower
that produced them. If some pollen grains do germinate, pollen tubes fail to enter the stigma.
In many species, the pollen tubes enter the style, but they grow too slowly to effect
fertilization before the flower drops. Sometimes, fertilization is effected, but the embryo
degenerates at a very early stage. Self-incompatibility appears to be a biochemical reaction,
but the precise nature of these reactions is not clearly understood. The genetic control of
incompatibility reactions is relatively simple. Lewis (1954) has suggested various
classifications of self-incompatibility ; a relatively simple classification is as follows ; 1.
heteromorphic system, 2. homomorphic system, (2a) gametophytic control, and (2b)
sporophytic control.
Heteromorphic System.
In this system, flowers of different incompatibility groups are different in
morphology. For example, in Primula there are two types of flowers, pin and thrum. Pin
flowers have long styles and short stamens, while thrum flowers have short styles and long
stamens. This situation is referred to as distyly. Tristyly is known in some plant species, e.g.
Lythrum ; in such cases, the style of a flower may be either short, long or of medium length.
In the case of distyly, the only compatible mating is between pin and thrum flowers. This
characteristic is governed by a single gene s ; Ss produces thrum, while ss produces pin
flowers. The incompatibility reaction of pollen is determined by the genotype of the plant
producing them. Allele S is dominant over s. The incompatibility system, therefore, is
heteromorphic -sporophytic. The pollen grains produced by pin flowers, would all be s in
genotype as well as incompatibility reaction. The pollen produced in thrum flowers would be
of two types genotypically, S and s, but all of them would be S phenotypically. The mating

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