UNIT -1

Introduction to plant breeding :

Historical development, concept, nature and role of plant breeding.

Major achievements and future prospects and genetics in relation to

plant breeding.

Domestication germplasm conservation and plant introduction :

Domestication, acclimatization and introduction, centres of

origin/diversity.
 PLANT BREEDING

“It is an art, science and technology which deals with genetic

improvement of crop plants in relation to their economic use for
mankind.”

Nature of Plant Breeding:-

1. Art
• In earlier days man depends on his skills and judgement in selecting better plants. He
knew nothing about the inheritance of characters, role of environment in producing
them and the basis of variation in various plant characters. His method of selection was
designed without the understanding of principles of inheritance. During primitive time
plant breeding was largely an art and very less science was involved in that.

• Plant breeding is an art because selection of superior plants requires human skills,
imagination and experience.
2. Science
• Plant breeding is considered as the current phase of crop evolution.
As the knowledge of genetics and other related science progresses
plant breeding become less art and more science. Especially,
discovery of Mendels̕ work in 1900 added a lot to the knowledge of
science.
• Plant breeding is a science because development of superior
varieties or hybrids involves genetic principles, sequential steps
(hybridization, selection, evaluation, multiplication, releases) and
experimentation.

3. Technology
• Product of all plant breeding activities, whether dependent on the
art or science, is improved variety, hybrids, synthetics and
composites. This product is utilized by farmers for commercial
cultivation.
• Therefore, plant breeding can be rightly viewed as a
technology since it generates a useful product
 ROLE OF PLANT BREEDING

1. Food: directly (food grains) or indirectly (meat

and milk)

2. Shelter: mainly in rural areas

3. Clothing: Cotton

4. Drugs : Rauwolfia serpentina, Withania somnifera

5. Fuel: Euphorbia, Jatropha, Maize

 HISTORY OF PLANT BREEDING

In broad sense history of plant breeding can be divided into 4 parts :

1. Pre Mendelian era:- before 1900
2. Mendelian era:- 1900 to 1920.
3. Post Mendelian era:- 1921 to 1950
4. Modern era:- after 1950
1. Pre Mendelian era:-
700 BC Assyrians and Babylonians - Hand pollination of date palm

1665 Robert Hooke (England) – 1st Described the cell and known as father of
cell-biology.

1676 Millington - Anthers function as male organs

1694 Camerarius (Germany) - First to demonstrate existence of sexes

in plants.

1717 Thomas Fairchild – produced the first artificial hybrid, popularly known as
Fairchild҆s mule.
1753 Linnaeus - Published "Species Plantarum".
1761-66: Koelreuter (Germany) - Demonstrated that hybrid offspring
received traits from both parents and were intermediate in most traits.

1779 Knight - Emphasized the practical aspects of hybrids

1801 Lamarck - Theory of evolution through inheritance of acquired

characters .

1819 Shirreff - Utilized pure line selection to develop a new oat cultivar

(released in 1824), and a new wheat cultivar (released in 1832)

1831 Brown - Discovered the eukaryotic cell nucleus
1837-38 Schleiden and Schwann - cell theory
1859-89 Darwin - Published "Origin of Species“ and noted inbreeding, sterility, and
differences in reciprocal crosses

1866 Mendel - Published "Experiments in plant hybridization"

1884 Strasburger - Demonstrated fertilization and showed the fusion of the 2 nuclei
to form the zygote

1899 Novaschin and Guignard - Discovered double fertilization of egg and

endosperm

1899 Hopkins - Described ear to row selection method.

2. Mendelian era:-
1900 Correns (Germany), DeVries (Holland) and Von Tschermak
(Austria) - rediscovered Mendel҆s laws of heredity.

Bateson- Introduced the terms "allelomorph", "homozygote",

"heterozygote ", "F1" and "F2"

1902 DeVries (Holland) - mutation theory of evolution

1903 Johannsen - proposed pure line theory of selection

1904 Hannig - Contributed to the idea of embryo culture

1906 Bateson - term "genetics“

1908 Nilsson- Ehle – given the explanation of multiple factor hypothesis

1908-09 Hardy (England) and Weinberg (Germany) –Hardy- Weinberg law.

1914 Shull - Introduced the term "heterosis "

Blakeslee - Discovered trisomics in Datura.
3. Post Mendelian era:-
1927 Muller - Reported artificial mutations in animals by X-rays

1928 Stadler - Described the mutagenic effects of X-rays in barley

1931 Stern, Creighton and McClintock - cytological proof of crossing-over

1933 Rhoades - cytoplasmic male sterility in maize

1944 Avery, MacLeod and McCarty – DNA is the hereditary material

1945 Hull - Proposed recurrent selection method

1946 Comstock et al. - Suggested reciprocal (half-sib) recurrent selection

1950 McClintock – Jumping genes

4. Modern era:-
1952 Jensen - 1st suggested the use of multilines in Oats.

1955 Benzer – Based on his work on rII locus of T4 Bacteriophage he gave the
subdivisions of genes in cistron, recon and muton.

1963 Vanderplank – developed the concept of vertical and horizontal

resistance.

1964 N.E.Borlaug – developed the high yielding dwarf varieties of Wheat

which resulted in green revolution.

1968 Donald – developed the concept of crop ideotype in wheat

1978 - Development of worlds 1st Rice hybrid (CMS based) for commercial
cultivation in China.

1983 – Development of 1st transgenic (genetically engineered) plant of

Tobacco in U.S.A.

1987 – Development of 1st transgenic cotton plant by Monsanto company in

U.S.A.

1994 – Flavr Savr tomato was introduced by Calgene company of California.

 HISTORY OF PLANT BREEDING IN INDIA

1871- Government of India created the Department of Agriculture.

1905- Imperial Agricultural Research Institute was established in Pusa (Bihar)
1921- The Indian Central cotton committee was established.
1929- Imperial Council of Agricultural Research was established in New
Delhi.
1936- Imperial Agricultural Research Institute shifted to its present location in
New Delhi.
1956- Project for Intensification of Regional Research on Cotton, Oilseeds and
Millets (PIRCOM)
1957- All India Coordinated Maize Improvement Project

1961- First Maize hybrid released (Ganga1, Ganga101, Deccan and

Ranjit)

1964- First Sorghum hybrid (CSH-1) was released.

1965- First Bajra hybrid (HB-1) was released.

1991 – 1st pigeon pea hybrid (ICPH-8) was released from ICRISAT,
Hyderabad
OBJECTIVES OF PLANT BREEDING

1. Higher Yield:
2. Improved Quality:
3. Abiotic resistance
4. Biotic resistance
5. Change in maturity Duration / Earliness
6. Determinate Growth
7. Dormancy :
8. Desirable Agronomic Characteristics:
9. Elimination of Toxic Substances :
10. Non-shattering characteristics:
11. Synchronous Maturity :
12. Photo and Thermo insensitivity:
13. Wider adaptability:
14. Varieties for New Seasons
1. Higher Yield:
 -The ultimate aim of plant breeding is to improve the yield of
“economic produce on economic part”.
 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.
i. Grain size, colour, milling and baking quality in wheat.
ii. Cooking quality in rice,
iii. Malting quality in barley,
iv. Colour and size of fruits,
v. Nutritive and keeping quality in vegetables,
vi. Protein content in pulses,
vii. Oil content in oilseeds,
viii. Fibre length, strength and fineness in cotton.
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.

Drought affected field

4. Biotic 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

Blast affected rice field

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
 Eg: 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 germination.

seed germination on mother plant

8. Desirable Agronomic Characteristics:
 It includes plant height, branching, tillering capacity,
growth habit, erect or trailing habit etc., is often
desirable.
 For example:
 dwarfness 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 – lentil (Lathyruys sativus)
which leads to paralysis of lower limbs
 Erucic acid from Brassica which is harmful for
human health
 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.

Mung
11.Synchronous Maturity :
 It refers to maturity of a crop species at one time.

 The character is highly desirable in crops like greengram,

cowpea, castor 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.
 ACHIEVEMENTS OF PLANT BREEDING

A) Production of dwarf and semi dwarf cereal varieties:

B) Nobilisation of sugarcane

C) Development of hybrid and synthetics

A) Production of Dwarf and Semi Dwarf Cereal Varieties:

Many dwarf and semi dwarf varieties are developed in crops like wheat
and rice.
Dr. Borlaug used NORIN-10 as a source of dwarfing gene, in wheat at
CIMMYT (Mexico).
Variety Kalyan Sona and Sonalika were selected from these materials.

Similarly the development of semi dwarf varieties of Rice has

revolutionized rice cultivation. These varieties were developed by
introducing the gene from japonica variety Dee-Geo-Woo-Gen.
Ex TN. 1 developed at Taiwan and IR-8 at IRRI, Philippines, both were
introduced in India in 1966.
B) Nobilisation of Sugarcane:

The Indian canes were of Saccharum barberi, largely grown in North

India. They were hardy but poor in yield and sugar content,
……while tropical noble cane of Saccharum officinarum had thicker
stem and higher sugar content but it performed badly in North India due
to low winter temperature.

C.A Barber and T.S Venkatraman at Sugarcane Breeding Institute,

Coimbatore, transferred thick stem, higher sugar content and other
desirable characters from the Noble cane to Indian cane is commonly
referred as Nobilisation on of Indian canes.
C) Development of Hybrid and Synthetic:

1. Maize: Ganga series of hybrid, Ganga safed-2, African Tall,

Manjari, Deccan etc.

2. Sorghum : CSH-1,2,3,4,5,6,7,8,910,12,14 and 15 R.

3. Bajara: WCC-75, PHB-10, ICTP-8203, Shraddha and Saburi.

4. Cotton: H-4, VaraLaxmi, Savitri, NH-44, Jaylaxmi, etc.

Indian Scientist:
1. Dr. M.S. Swaminathan – Father of green revolution in India. He developed the high
yielding dwarf varieties of Wheat and Mutation breeding
2. Maheshwari and Guha (1964)- Produced haploid plant in Vitro from pollen
grain.
3. C.A.Barber and T.S venkatraman – Nobilisation of sugarcane.
4. Dr. Athwal – Bajra breeder.
Developed world’s first grain pearl millet hybrid ‘Hybrid Bajra 1’ in 1965

5. K. Ramiah- Rice breeder

6. Dr. N.G.P. Rao – Sorghum breeder
Father of Hybrid Sorghum in India and developed 1st hybrid of sorghum (CSH-1).

7. Pushkarnath - Potato Breeder

Developed several high yielding varieties of potato.

8. Ram Dhan Singh- Wheat breeder

9. C. T Patel- Cotton breeder
World’s first interspecific cotton hybrid- Hybrid-4 and known as father of Hybrid cotton
 Scope of Plant Breeding

• 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.

• Since the population is increasing at an alarming rate, there is

need to strengthened the food production 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.
• Genes from varied organisms may be expected to boost
the performance of crops especially with regard 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
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 in Punjab and Haryana.

- The severe infection by Karnal bunt (Tilletia sp.) on some wheat

varieties, infestation of mealy bugs in Bt cotton
 DOMESTICATION

“Domestication is the process of bringing wild species under

human management.”

All crops grown today have

undergone extensive genetic
change from their wild ancestors.
 SELECTION UNDER DOMESTICATION

1)Natural Selection

Mechanism of Natural selection given by Nichlson 1962

i) Environmental selection

ii) Competition

2) Artificial Selection
PATTERNS OF EVOLUTION IN CROPS

1. Mendelian Variation
a) Macromutation
b) Micromutation

1. Interspecific Hybridization

2. Polyploidy
 Centres of plant domestication

Alphonse De Candolle was the first to suggest in 1886

that a crop plant originates from the area where its wild
progenitor occurs.

• French systematic botanist (taxonomist) proposed

that the crops of the world originated in 3 centers:
1). Mesoamerica
2). the Fertile Crescent of the Middle East
3). South East Asia
 CENTRES OF ORIGIN
• Center of origin is the region of the world where a crop is believed to
have originated.

1. PRIMARY CENTRES OF ORIGIN

2. SECONDARY CENTRES OF ORIGIN

Studies Conducted At Institute Of Plant Industry,

Leningrad (1916-1936)

Law of Homologous Series in Variation

 LIST OF CENTRES OF ORIGIN

Centre of Origin Primary Centre of Origin Secondary Centre of Origin

Barley , Triticum spp, jawar, bajara,
gram, lentil,pea,khesari, liseed,
safflower, seasamum, castor,
Abyssinian Centre coffee, onion, okra,etc Broad bean (Vicia faba)
Triticum spp, rye, alfalfa, carrot,
cabbage, oat, lettuce, apple, pyrus
spp, prunus spp, grape, alomond,
Asia Minor Centre ( Syn, near chestnut, pistachio nut, persain B. compestris, B. nigra, turnip,
East or Persian Centre) clover, etc. apricot, etc.
Maize, rajma, lina beans, melon,
pumkin, sweet potato, arrowrrot,
Central American Centre (Syn. chillies, G.hirsutum, papaya,
Mexican Centre) guava, avacado, etc.
Triticum aestivum , pea, mung,
linseed, sasamum, safflower,
hemp, G. herbaceum, radish,
muskmelon, carrot, onion, garlic,
Centra Asia Centre ( Syn, spinach, pear, almond, grape,
Afganistan Centre) apple, etc. Rye (Secale cereale)
 Centre of Origin Primary Centre of Origin Secondary Centre of Origin

Soybean , radish, bunda

(Colocasia sp), proso millet,
buckwheat, opium poppy, brinjal,
pear, peach, apricot, plum, orange, Maize, rajma, cowpea, turnip,
China Centre Chinese tea, etc seasamum, etc.

Rice, pigeon pea, chickpea,

cowpea, mung, brinjal, cucumber,
Hindustan Centre ( Divided Indian radish , noble canes,
into 1) Indo-burma and 2) G.arboreum, Mango, orange,
Siam-malaya- java Centre) coconut, banana, etc.

Triticum spp, barley, Avena spp,

lentil, pea, broad bean, lathyrus
spp, chickpea, clovers, Brassica
spp, onion, garlic, beets, lettuce,
asparagus, lavender, peppermint,
Mediterranean Centre etc.
South American Centre Potato, maize, pineapple,
(1. Peru, 2. Chile and 3. pumpkin, G.b barbadense,
Brazil-Paraguay Centres) tomato, tobacco
(1) Central American Centre (2) South American Centre
(3) Mediterranean, (4) Asia Minor Centre (5) Abyssinian Centre
(6) Central Asia, (7) Hindustan centre of origin (7A) Indo-Burma, (7B) Siam-
Malaya-Java, (8) China Centre
 CENTER OF DIVERSITY

Geographic region in which greatest variability of a crop occurs.

Types of Centers of Diversity of Crops

The centers of crop diversity of three types viz:

1) Primary centers of diversity
2) Secondary centers of diversity
3) Micro –Centers
1. Primary Centers of Diversity:

Primary centers are regions of vast genetic diversity of crop

plants. These are original homes of the crop plants which are
generally uncultivated areas likes mountains, hills, river
valleys, forests etc

Main features of these centers are given below:

1. They have wide genetic diversity.

2. Have large number of dominant genes.
3. Mostly have wild characters.
4. Natural selection operates.
2. Secondary Centers of Diversity:

• N.I. Vavilov (1926) suggested that valuable forms of crop plants

are found far away from their primary area of origin, which he
called secondary centers of diversity.

These are generally the cultivated areas and have following main
features.

1. Have lesser genetic diversity than primary centers.

2. Have large number of recessive genes.
3. Mostly have desirable characters.
4. Exhibit more crossing over
5. Both natural and artificial selections operate.
3. Microcenters:

In some case, small areas within the centers of diversity exhibit

tremendous genetic diversity of some crop plants. These areas are
referred to as micro-centres.

The main features of micro centers are given below:

1. They represent small areas within the centers of diversity.
2. Exhibit tremendous genetic diversity.
3. The rate of natural evolution is faster than larger areas.
4. They are important sites for the study of crop evolution.
GENE BANKS
Gene Banks:

Gene bank refers to a place where germplasm can be conserved in

living state. Gene banks are also known as germplasm banks.

The germplasm is stored in the form of seeds, pollen or in vitro

cultures, or in the case of a field gene bank, as plants growing in the
field.

Gene banks are mainly of two types, viz:

1) Seed gene banks, and
2) Field gene banks.
1. Seed Gene Bank:
 A place where germplasm is conserved in the form of seeds is called
seed gene bank.

 Seeds are very convenient for storage because they occupy smaller
space than whole plants.

 However, seeds of all crops can not be stored at low temperature in

the seed banks.

 The germplasm of only orthodox species (whose seed can be dried

to low moisture content without losing viability) can be conserved
in seed banks.
In the seed banks, there are three types of conservation, viz.

1) Short term:- Working collections are stored for short term (3-5
years) at 5-10 0C.

2) Medium term:- Active collection are stored for medium term (10-
15 years) at below 15 0C.

3) Long term. Base collections are conserved for long term (50 years
or more) at -18 or -20 0C.
Advantages of gene banks:
1. Large number of germplasm samples or entire variability can be
conserved in a very small space.
2. In seed banks, handling of germplasm is easy.
3. Germplasm is conserved under pathogen and insect free environment.

Disadvantages of gene banks:

1. Seeds of recalcitrant species can not be stored in seed banks.
2. Failure of power supply may lead to loss of viability and there by loss of
germplasm.
3. It requires periodical evaluation of seed viability. After some time
multiplication is essential to get new or fresh seeds of storage.
2. Field Gene Banks:
Field gene banks also called plant gene banks are areas of land in which
germplasm collections of growing plants are assembled.

This is also ex-situ conservation of germplasm. Those plant species that

have recalcitrant seeds or do not produce seeds readily are conserved in
field gene bank.

In field gene banks, germplasm is maintained in the form of plants as a

permanent living collection.

Eg. coconut, rubber, mango, cassava, yam, and cocoa.

Advantages field gene banks:

1. It provides opportunities for continuous evaluation for various economic

characters.
2. It can be directly utilized in the breeding programmes.

Disadvantages field gene banks:

1. Field gene banks can not cover the entire genetic diversity of a species. It
can cover only a fraction of the full range of diversity of a species.
2. The germplasm in field gene banks is exposed to pathogen and insects and
some times is damaged by natural disasters such as bushfires, cyclones,
floods , etc.
3. Maintenance of germplasm in the field gene banks is costly affair.
Gene Banks for Various Crops in India
Location of Gene
Crop Species Bank Name of Research Institute
Indian Institute of Wheat and Barley
Wheat Karnal Research (IIWBR)
Rice Cuttack National rice research institute ( NRRI)
Potato Shimla Central Potato Research Institute ( CPRI)
Cotton Nagpur Central Institute for Cotton Research ( CICR)
Pulses Kanpur Indian Institute for Pulses Research ( IIPR)
Oil seed crops Hyderabad Indian Institute of Oilseeds Research( IIOR)
Sorghum Hyderabad Indian Institute of Millets Research (IIMR)
Soybean Indore Indian Institute of Soybean Research
Groundnut Junagarh National Research Centre of Groundnut
Maize New Delhi Indian Agricultural Research Institute
Citrus Nagpur Central Citrus Research Institute
Sugarcane Coimbatore Sugarcane Breeding Institute
Indian grassland and Fodder Research
Forage Crop Jhansi Institute ( IGFRI)
Tobacco Rajahmundry Central Tobacco Research Institute ( CTRI)
International Gene Banks for Different Crops
Sr. Location of
No. Name of Institute Gene Bank Crops Maintained
International Rice Research
1 Institute (IRRI) Philippines Rice
International Wheat and Maize Maize, Wheat,
2 Improvement Centre ( CIMMYT) Mexico Triticale, Barley
International Centre for Tropical Cassava, beans, Rice
3 Agriculture ( CIAT) Columbia and Maize
Cowpea, Soybean,
International Institute for lima bean, cassava,
4 Tropical Agriculture ( IITA) Nigeria Sweet potato.
5 International Potato Centre Peru Potato
International Crop Research Sorghum, Pearl Millet,
Institute for Semiarid Tropics ( pigeon pea,
6 ICRISAT) India groundnut.
International Centre for
Agriculture Research in Dry land Durum Wheat, Barley,
7 Areas ( ICARDA) Syria beans.
PLANT INTRODUCTION
Plant Introduction
 It consists of taking a genotype or group of genotypes
into a new environment where they have not been
grown before.

 The introduction may involve introducing new varieties

of the crop already grown in the area, wild relatives of
the crop species or a totally new crop species for the
area.

 It can be done within the country or from other countries

of continents.
Types: Introduction can be categorized as:-
1. Primary Introduction
2. Secondary Introduction

1. Primary Introduction
 An introduction is said to be primary introduction when
the introduced variety is well suited for new
environment.
 It is released for commercial cultivation without any
alteration in original genotype.
Eg: introduction of semidwarf wheat varieties (Sonora 64,
Lerma Rojo) and semidwarf rice (TN1, IR 8, IR28, IR 36) in
India
2. Secondary Introduction
 The introduced variety may not be suitable for direct
cultivation and can be subjected to selection to isolate a
superior variety suitable for a particular area.

 The introduced variety may also be hybridized with local

variety to transfer one or two characters from it to local
variety.
 Eg: Kalyan Sona, Sonalika
PROCEDURE FOR PLANT INTRODUCTION

1. Procurement

2. Quarantine

3. Cataloguing

4. Evaluation

5. Multiplication

6. Distribution
1) Procurement of germplasm
 The new germplasm is procured through NBPGR, New Delhi.

 Scientists, individuals and institutions can submit their

requirement to Director, NBPGR, Pusa Complex New Delhi.

 If the bureau is unable to meet the request from its own stock
or from known source it attempts to procure them from the
counterparts in other countries.

 Generally the material is obtained through correspondence

as gifts or exchange of germplasm in consideration of past
gifts to the Bureau or in anticipation of future gifts.

 The plant part depending on the crop species e.g. seeds;

tubers; suckers, bulbs or cuttings etc. can be procured.
2. Quarantine
• It is to keep the material in isolation to prevent spreading of diseases
etc.

• All introduced material is thoroughly inspected for contamination

with diseases, weeds and insects. Plants that are suspected to be
contaminated are fumigated or are given other isolation for
treatments and observed for insect pests and disease.

• The entire process is known as quarantine and the rules which are
used for this are known as Quarantine rules.

• It is essential that all the material being introduced must be

accompanied by an authentic phytosanitary certificate.

• The plant material being introduced or exported must confirm to

certain quarantine regulations and quarantine control is exercised by
NBPGR at different points of entry.
3) Cataloging
 All the plant material which is introduced is given an
entry number and information regarding agency, place of
origin, adaptation etc and is well documented.

 Plant material is classified in three categories

1. Exotic collection (EC)
2. Indigenous collection(IC)
3. Indigenous wild (IW) collection
4) Evaluation
 The plant material is sent to sub stations of the bureau
and evaluated with respect to various characters to
assess the potential of new introductions.
5) Multiplication and distribution
 Plant material which is introduced is to be multiplied and
further tested at various locations.
 The suitability of cultivation in different regions of the country
should be assessed before using it as a commercial variety.
 Acclimatization
 It is the process that leads to the adaptation of a variety
to a new environment.
 Variability must be present in original population so that
natural selection could lead to acclimatization.
 The extent of acclimatization is determined by range of
genetic variability in original population and duration of
life cycle of crop.
 Cross pollination leads to far more gene recombination
than self pollination. The greater the initial variation the
more is acclimatization.
 Purpose of Plant Introduction
1. Entirely new crops
- (Maize, Potato, Soybean)
1. New varieties
-Introductions are directly released as superior commercial
varieties
-TN1, IR8 (Rice); Sonora 64 and Lerma rojo (Wheat)
3. Utilization in crop improvement programmes
4. Saving a crop from a disease or pest
5.Utilization in Scientific studies
6.Used for aesthetic value
 Merits of Plant Introduction

1. Provides entirely new crop plants and superior varieties

for different crops
2. Helps in germplasm collection
3. Quick and economic method of crop improvement
 Demerits of Plant Introduction
Plant introduction are associated with entry of weeds,
diseases and Pests into India with introduced materials
1. Weed
- Phylaris minor, Argemone maxicana
2. Diseases
-Late blight of Potato from Europe in 1883
3.Insect Pests
-Potato tuber moth from Italy in 1900
4. Ornamentals turned weeds
- Lantana camara
5. Threat to ecological balance
- Eucalyptus spp introduced from Australia cause rapid
depletion of subsoil water reserves
 AGENCIES IN INDIA

National Bureau of Plant Genetic Resources (NBPGR)

Forest Research Institute

Botanical Survey of India

NBPGR has its headquarters at IARI New Delhi

Genetics of qualitative and quantitative traits

Components of Genetic Variance

Dominance:- The allele that that express itself in the
presence of another allele is called as dominant.

Recessive: The one that unable to express itself is knows

as recessive.
Qualitative Traits:- The characters which are governed by one or
few genes called as Qualitative characters also called as
oligogenes.

Ex. Mendel law of segregation.

 Inheritance of qualitative characters

Eg.
Laws of Mendel-Law of Segregation

Two alleles of a gene present in the F1 do not contaminate

each other; they separate and pass into different gametes in
their original form producing two different types of gametes in
equal proportion.
Quantitative Traits:- The characters which are governed by several
genes called as Quantitative characters also called as polygenes.

EX. Multiple factor hypothesis.

 Inheritance of quantitative characters

Ex. Multiple factor hypothesis:-

This hypothesis was originally postulated by Yule in 1906, but

experimental evidence presented by Nilsson-Ehle in 1908.
In studies on the inheritance of seed colour in wheat
Thus the studies on Nilsson-Ehle demonstrated the existence of several genes
with small cumulative effects governing a single traits; such genes are commonly
known as polygenes (sometimes minor genes).
Sr. Particulars Qualitative Traits Quantitative Traits
no.
1 Gene involved One or few Several
2 Effect of individual Large & detectable Small and undetectable
gene
3 Variation Discontinuous Continuous
4 Grouping into distinct Possible Not possible
classes
5 Effect of environment Little High
6 Metric Measurement Not possible Possible
7 Analysis Based on frequencies Means, variance,
and ratios covariances
8 Stability High Low to medium
9 e.g., Colour, shape, surface Height, weight,
of plant parts width,length,duration
Components of Genetic Variance:

In crop improvements programme, only the genetic components of

variation are important because only this component is transmitted to
the next generation.

According to Fisher in 1918, components of genetic variance divided

into three components viz.
1) Additive,
2) Dominance
3) Epistatic
1) Additive Components:
It is the component arising from difference between the two
homozygotes for a gene, E. g AA and aa.

2) Dominance Component:
It is due to the deviation of heterozygote ( Aa) phenotype from the
average of phenotypic value of the two homozygotes (AA and aa). It is
also referred as intra allelic interaction.

3) Epistatic or Interaction Components:

It results from an interaction between two or more genes.
Later Hayman and Mather classified the epistatic components into
three types interaction viz.
1) Additive X additive
2) Additive X dominance,
3) Dominance X dominance.
Heritability

1. The ratio of genotypic variance to the phenotypic variance is

known as heritability.

2. It is expressed in per cent.

3. It is a good index of the transmission of characters from parents

to their offspring.

4. The estimates of heritability help the plant breeder in selection

of elite genotypes from diverse genetic populations.
 Types of Heritability

Depending upon the components of variance used as numerator in

the calculation, heritability is of two types,

1. Broad sense heritability

2. Narrow sense heritability

 Broad sense heritability

It is the ratio of genotypic variance to total or phenotypic variance.

It is calculated from total genetic variance which consists of

additive, dominance and epistatic variances.
Main features of broad sense heritability
h 2 (ns) = VA/VP
Since narrow sense heritability is estimated from additive genetic
variance it plays important role in the selection of elite genotypes
from the segregating populations, whereas broad sense
heritability estimates are more useful in selecting superior lines
from the homozygous materials.
 Genetic Advance

Improvement in the mean genotypic value of selected plants over the

parental population is known as genetic advance.

It is the measure of genetic gain under selection.

The success of genetic advance under selection depends on three main

factors:

1. Genetic Variability
2. Heritability
3. Selection Intensity
1. Genetic Variability

The magnitude of genetic variability present among plants or

families in the base population has important bearing on the
genetic advance.

The greater the genetic variability the higher is the genetic advance
and vice versa.
2. Heritability

The heritability of the character under consideration affects the

genetic advance.

The genetic advance is generally high with the characters having

high heritability and vice versa.
3. Selection Intensity

The proportion of plants or families selected for the study is called

as selection intensity, which plays important role in the success of
genetic advance.

The high selection intensity generally gives better results than low
selection intensity.
 Computation of genetic advance

Knowledge of genetic gain and selection differential is essential before

dealing with the computation of genetic advance.

Genetic gain:-
The difference between the mean phenotypic values of the progeny of
selected plants and the base or parental population is known as genetic
gain.
It is denoted by R.
R = XP – XO

Where,
XP – Mean phenotypic values of the progeny of selected plants
XO – Mean of base population
Selection Differential:-

The difference between the mean phenotypic value of selected plants and
mean phenotypic value of parental population.

This is a measure of the intensity of selection and is denoted by K.

Selection intensity (K):-

1% 2% 5% 10%
2.64 2.42 2.06 1.76

Genetic advance = K . h2 . P
K – Standardize selection differential
h2 – heritability
P – phenotypic standard deviation
 Maintenance of breeding records
Field trials are essential in crop breeding to evaluate the
progeny and comparison with other varieties. It is important that the
breeder, master the techniques to keep the records of all kind of data to
achieve the objective he has planned to achieve. The procedure that he
may use to keep the record will depend on the particular objective of
his experiment.

In field breeding, breeder has to maintain various records, like-

1. Accession Register: This will contain the details of the seeds/

planting material with regard to receipt date, source, their number,
number assigned at the receiving unit, short description of the planting
material, to whom sent for evaluation, date, feedback information
about the genotype, how to maintained etc.
Accession number given by the serial number followed by the year of
entry i.e. serial 145 in 1991. Then accession number will be 145191 or
91145. It will be mentioned as EC = Exotic collection IC = Indigenous
collection.
2. Standard form of a Field Note Book
Each field notebook should contain the following information.
A. For Yield Trial:
i) 1st Page: Number and title of the project, Season of raising
the crop and Unit under which the trial is being conducted

ii) 2nd page: A full plan of the field showing the location of the
trial with the approach path, North East directions should be
specified.

iii) 3rd Page: Plan of the experiment, Experiment details,

Name of the experiment, Season, Number of variants, Design
of the experiment, Replication, Size of the plot
(Block/Plot/Row etc.), Spacing (Between rows and within the
row in cm), Date of sowing/planting, Date of harvest, Name
of the Principal Investigator
iv) 4th page: Details of cultural practices followed for the plot/ field like
Date of ploughing, Date of layout of the trial, Manurial schedule
adopted, Irrigation schedules with date from life irrigation onwards,
Plant protection schedules followed, Details of intercultural operations
(hoeing, weeding, and earthing up etc.), Date of harvest, Duration of
processing till storage, Rainfall received during the crop growth,
General remarks on the seasonal condition prevailed and its effects on
crop growth including the occurrence of pests and disease.

v) 5th page: One page for each variant per replications allotted and the
following information has to be recorded in each page. Date of
germination, Date of gap filling, Initial stand on, Date of first flowering,
Date of general flowering, Date of harvest, Final stand, Wet weight of
grain, Wet weight of haulms/ straw etc., Dry weight of produce after
cleaning, Yield per ha in kg. The page will also have additional
information on observations about the variant, recorded by the breeder
in relation to the object of the project.
The fifth page will also contain the following information and
their modification depending upon the crop.
e.g. Rice (Date of ear head emergence in the main shoot,
Date of ear head emergence in tillers and Number of tillers)
Cotton (Number of sympodial branches, Number of
monopodial branches)

B. Generation study:
This field notebook will contain the following information.
a. Plan for the segregation generation
b. Details of the generation (Name of the generation study,
Number of crosses, Details of the cross, Length of row,
Spacing (cm), Date of sowing, Dates of harvest, and Name of
the Principal Investigator)
Modes of reproduction and selection in self
pollinated crops: modes of reproduction and apomixis
Modes of reproduction and pollination control and its
relevance to breeding :

1. Sexual reproduction
2. Asexual reproduction
3. Apomixis and its types
4. Different mechanism of pollination
5. Self Incompatibility
6. Male sterility
Mode of Reproduction: Production of a new generation of
individuals by sexual or asexual means.

Two types

1. Asexual reproduction
2. Sexual reproduction
Asexual reproduction

Asexual reproduction:- The development of new individuals without

fusion of male and female gametes is known as asexual reproduction.

New plants may develop from vegetative part of the plant (vegetative
reproduction) or may develop from embryos without fertilization
(apomixis).

1. Vegetative reproduction
2. Apomixis
A) Vegetative Reproduction:

In this new plants developed from a portion of the plant body.

This may be occurred thorough modified underground and sub-aerial

stems or through bulbils.

For example Rhizome – Ginger, Tuber-potato, Bulbs- onion, corm- arwi.

Sub-aerial stems- mint, bulbils- garlic
Similarly artificial vegetative methods stem cutting ( Sugarcane) and
Root cutting ( Lemon, citrus).
Layering, budding and grafting (mango) are common methods in
propagation of fruits and ornamental horticultural crops.
B. APOMIXIS

It is the type of asexual reproduction in which seeds

are formed and embryo developed without
fertilization.

In apomixis, sexual reproduction is either suppressed

or absent.
 Types of Apomixis

1. Adventitious Embryony:

2. Gametophytic Apomixis:

a) Apospory:

b) Diplospory:
i) Adventive Embryony:

Embryo directly developed from vegetative cells of

the ovule such as integuments and chalaza.
E.g, Mango, Citrus.
2. Gametophytic Apomixis: Embryos developed without
fertilization from egg cell or other cells of embryo sac.

a) Apospory:
Diploid embryo sac develop from vegetative cells of ovule.
Embryo may be developed from egg cell or other cell of embryo
sac.

b) Diplospory:
Embryo sac is developed from the megaspore, which may haploid
or diploid.
Sexual reproduction

Sexual reproduction:-Sexual reproduction involves fusion of male

and female gametes to form a zygote, which develops into an
embryo.

Flower:- In crop plants, male and female gametes are produced in

specialized structure known's as flower.
Perfect or Hermaphrodite flower:-

Staminate flower:-

Pistillate flower:-

Monoecious species:-

Dioecious species:-

Sporogenesis :-

Gametogenesis :-
Sporogenesis:- Production of microspore and megaspore is known's
as Sporogenesis.

Microspores are produced in anthers called microsporogenesis,

while megaspore are produced in ovules called megasporogenesis.

Gametogenesis:- The production of male and female gametes in

microspore and megaspore called gametogenesis.

Microgametogenesis:- Production of male gamete or sperm.

Megagametogenesis:- Development of embryo sac from a

megaspore.
Meiosis: Two successive spindle using division which reduce
the chromosome no. from diploid to haploid. Or “Two cell
divisions, occurring one after the other, following only one
round of DNA replication, shows chromosome pairing and
chromosome separation in the first division and chromatid
separation in the second division, which leads to the
formation of haploid daughter cells.”

Mitosis: The spindle using nuclear division which produce two

identical daughter cells from mother cell. Or “The cell division
in which each nuclear division is preceded by one round of
DNA replication and during anaphase sister chromatids
separate and pass to the opposite poles”
Double fertilization

Fertilization:- The fusion of one of the two sperms with the egg
cell, producing a diploid zygote, is known's as fertilization.

triple fusion:- The fusion of the remaining sperm with secondary

nucleus, leading to the formation of a triploid primary endosperm
nucleus, term as triple fusion.
Synergids contain Filiform apparatus
in its micropylar end which secretes a
chemotropically active substance that
directs the growth of the pollen tube.

Antipodals are nutritive in function.

Antipodals usually degenerate before
or soon after fertilization in most
plants and therefore considered
vestigial. But in some plants it
remains persistent, where it produces
substances that controls the growth
and development of endosperm.
Pollination

1. Self pollination or autogamy

2. Cross pollination or allogamy

3. Geitonogamy

4. Often cross-pollinated species

Pollination:-Pollination refers to the transfer of pollen grain from anthers to
stigmas.

1. Self pollination or auto gamy:- Pollen from an anther may fall on the
stigma of the same flower leading to self pollination or auto gamy.

2. Cross pollination or allogamy:- Some times pollen from an anther may

fall on the stigma of another flower of different plants leading to cross
pollination or allogamy.

3. Geitonogamy:- Some times pollen from an anther of one flower fall on

the stigma of the another flower of same plant leading to the
geitonogamy. Eg. Maize

4. Often cross-pollinated: This type of plants are self pollinated, however the
extent of cross pollination often exceed 5 to 30 %. Such species are generally
known as often cross pollinated species.
E.g. Jawar, Cotton, Safflower, Arhar, etc.
 Self pollination
Mechanism of self-pollination
1. Bisexuality: Male and female sexual organs present in the same
flower e.g Wheat, rice, groundnut, etc.

2. Homogamy: Male and female sexual organs mature at the same

time e.g wheat, groundnut, etc.

3. Cleistogamy: In this condition flowers does not open at all and

ensure complete self pollination e.g Rice, Wheat.

4. Chasmogamy: In some species, flower open but only after

pollination has taken place. E.g Barley, Oat.

5. In crop like Tomato and Brinjal stigma are closely surrounded by

anthers , hence pollination occurs after opening of flower but the
position of anther in relation to stigma ensure self – pollination.
6. Stamens and stigma hidden by floral organs
 Genetic Consequences of Self – Pollination

1. It leads to a very rapid increase in homozygosity; therefore self

pollinated species highly homozygous in nature.

2. Self pollinated species do not show inbreeding depression,

exhibit considerable heterosis.
 Self Pollinated Crop Species

Cereals and Millets: Vegetables:

1. Wheat (Triticum aestivum) 1. Tomato (Lycopersicon esculentum)
2. Rice (Oryza sativa) 2. Okra (Ablmoschus esculentus)
3. Barley (Hordeum vulgare) 3. Lettuce (Lactuca sativa)
4. Oats (Avena sativa) 4. Brinjal (Solanum melongana)
Legumes: 5. Chilies (Capsicum annum)
1. Pea (Pisum sativum) 6. Potato (Solanum tuberosum)
2. Groundnut (Arachis hypogea) Oil Seeds:
3. Gram (cicer arientinum) 1. Til (Seasamum indicum)
4. Mung (Vigna radiata) 2. Linseed (Limum ustitatissimum)
5. Urd (Vigna mungo) Fruit Trees:
6. Cowpea (Vigna anguiculata) 1. Apricot (Prunus armeniaca)
7. Soybean (Glycine max) 2. Nectarine (Prunus persica)
8. Lentil (Lens esculentus) 3. Citrus (Citrus sp)
 Cross Pollination

The transfer of pollen from a flower to the stigma of the other flower
of different flower plant.

In cross pollinated species pollination may be brought about by wind,

water insect or animals.

Wind (anemophily)
Water ( hydrophily)
Insect ( entomophily)
Animal ( Zoophily)
Mechanism of cross pollination

i) Dicliny (Unisexuality):
a) Monoecy:
b) Dioecy:
ii) Dichogamy:
a) Protogyny:
b) Protandry:
iii) Heterostyly:
iv) Herkogamy:
vii) Self –Incompatibility:
Viii) Male Sterility:
i) Dicliny (Unisexuality):
It is a condition in which flower is either staminate or pistillate.
a) Monoecy:
Staminate and pistillate flowers occur in the same plant either in the
same inflorescence. E.g Maize, Cucurbit, Strawberry, etc.
b) Dioecy:
The male and female flowers are present on different plants i.e. the in
such species are male or female. E.g. Papaya, date, palm, etc.
ii) Dichogamy:
Anther and stigma of hermaphrodite flower mature at different time,
facilitating cross pollination.
a) Protogyny:
Gynoceium matures earlier than the androceium E.g. Bajara.
b) Protandry:
Androceium matures earlier than gynoceium. E.g. Maize.
iii) Heterostyly:
Different length of style and filaments E.g Linseed, Buckwheat
iv) Herkogamy:
Presence of physical barrier. In lucerne or alfalfa stigma are covered by
waxy film and it does not become receptive unless this waxy film is
broken by honeybees.

vii) Self –Incompatibility:

It refers to the failure of pollen from a flower to fertilize the same flower
or other flowers on the same plants. e,g mustard , tobacco, sunflowers.

Viii) Male Sterility:

It refers to the absence of functional pollen grains in hermaphrodite
flower.
Genetic Consequences of Cross Pollination:

1) It preserves and promotes heterozygosity in population.

2) Cross pollinated species shows inbreeding depression and

considerable heterosis.
Cereals:
1. Maize (Zea mays)
Cross Pollinated Species
2. Rye (Secale cereale)
Legumes:
3. Bajra (Pennisetum americanum)
1. Alfalfa (Medicago sativa)
Vegetables:
2. Red clover (Trifolium pratense)
1. Cabbage (Brassica oleracea)
3. White clover (Trifolium repens)
2. Carrot (Daucus carota)
Crops:
3. Cauliflower (Brassica oleracea)
Sugarcane (Saccharum oficinarum)
4. Cucumber (Cucumis sativus)
Hemp (Canabis indica)
5. Onion (Allium cepa)
Fruits:
7. Radish (Raphaus sativus)
1. Apple (Pyrus malus)
9. Muskmelon (Cucurbita moschata)
2. Avocado (Persea americana)
10. Watermelon (Citrullus vulgaris)
3. Mango (Mangifera indica)
12. Sweet potato (lpomoea batatas)
4. Pear (Pyrus communis)
15. Broccoli (Brassica oleracea)
5. Blackberries (Rubus fruticous)
21. Garlic (Allium sativum)
11. Cherry (Prunus avium)
22. Coriander (Coriandrum sativum)
14. Coconut (Cocos nucifera)
Oil Seeds:
15. Grapes (Vitis vinifera)
2. Sunflower (Helianthus annus)
16. Papaya (Carica papaya)
3. Castor (Ricinus communis)
Self incompatibility
 Self incompatibility
• It refers to the failure of pollen to fertilize the same flower or
other flower of the same plant.

• Self incompatibility is an important out breeding mechanism,

which prevent auto gamy and produce allogamy.

• Its is results due to morphological, genetic, physiological and

biochemical causes

• Controlled by a single locus termed S, which has many

different alleles in the species population
 Self incompatibility may occurs due to:

1. Pollen grain fails to germinate on the stigma.

2. Pollen grain germinates, but pollen tube fails to enter inside the
stigma and style.

3. Sometimes pollen tube enters inside the style but growth is very
slow.

4. Pollen tube enters the ovule but there is no fertilization due to

degeneration of eggs cells.

5. Sometimes fertilization is effected but embryo degenerate at

very early stage.
 Classification
Lewis (1954) has suggested various classification of self – incompatibility.

• On the basis of – Flower morphology, Genes involved,

Site of expression of SI reaction and pollen cytology
• Flower morphology : Heteromorphic and Homomorphic
• Genes involved : Monoallelic, Diallelic, polyallelic
• Site of expression : Stigmatic, stylar, Ovarian
• Pollen cytology : Binucleate, Trinucleate
 Classification
Flower morphology : Heteromorphic and
Homomorphic
1) Heteromorphic System:
In this system, flowers of different incompatibility groups are different
in morphology.

E.g In Primula there are two types of flowers. Pin and thrum. Pin
flowers have long style and short stamens.
While thrum flowers have short styles and long stamens. This
situation is referred as distyly.

Tristyly is known in some plant species. E.g Lythrum in such case, the
style of a flower may be either short, long or of medium length.
 Distyly
• Found in Primula

Pin (ss) x Pin (ss) Incompatible

Thrum (Ss) x Pin (ss) 1 thrum , 1 pin
Pin (ss) x Thrum (Ss) 1 thrum , 1 pin
Thrum (Ss) x Thrum Incompatible
(Ss)
 Tristyly
• Lythrum
 2) Homomorphic System:

In the Homomorphic system incompatibility is not associated with

morphological differences among flowers.

Two types of homomorphic system

1. Gametophytic control
2. Sporophytic control
1. Gametophytic control

When the self incompatibility is controlled by the genetic

constitution of gametes, it is known as Gametophytic self
incompatibility.

Gametophytic incompatibility was first described by East and

Mangelsdor in 1925 in Nicotiana sanderae.

Eg. Potato, Tomato.

 Pollen grains are unable to germinate on pistil having similar allele as
that of pollen grain.

Gametophytic system permits recovery of male parent only in partially

fertile crosses.

In a single system there are three types of mating.

1) Fully Incompatible (S1 S2 X S1 S2)

2) Fully Compatible (S1 S2 X S3 S4)

3) Partially Compatible (S1 S2 X S1 S3)

 Pollen grain

Pistil

S1 S3, S2 S3 S1 S3, S1 S4, S2 S3, S2 S4

2. Sporophytic control

When the self incompatibility is governed by the genotype of

pollen producing plant, it is called as Sporophytic self
incompatibility.

Hughes and Babcock first reported it in 1950 in Crepis foetida and

by Gerstel in Parthenium argentatum.

Eg. Radish, Cabbage, sunflower

The sporophytic system contains a form of dominance in which S1 is
dominant over all the other alleles, S2 is dominant over all except S1,
and so on S1>S2>S3>S4.

Sporophytic system permits recovery of both male & female parent.

 Utilization in plant breeding
• The production of hybrids
• Combining desirable genes from different
sources

• Limitation:
- Difficult to produce inbred
- Breakage due high temp and humidity
Male Sterility
 Male Sterility

Definition:- Male sterility is defined as an absence or non-

function of pollen grain in plant or incapability of plants to
produce or release functional pollen grains.

Importance
The use of male sterility in hybrid seed production has a great
importance as it eliminate the process of mechanical
emasculation.
 Types of Male Sterility

The male sterility is of five types

1. Genetic male sterility (GMS) – governed by nuclear genes-

TGMS, PGMS
2. Cytoplasmic male sterility (CMS) – governed by cytoplasmic
genes
3. Cytoplasmic-Genetic male sterility (CGMS) – governed by both
nuclear and cytoplasmic genes
4. Transgenic male sterility – induced by the technique of genetic
engineering
5. Chemical induced male sterility – induced by the use of
chemical.
Phenotype expression of male sterility

1) Absence of male organs.

2) Failure to develop normal microsporogenous tissue- anther
3) Abnormal microsporogenesis deformed or inviable pollen
4) Abnormal pollen maturation
5) Non dehiscent anthers but viable pollen.
6) Barriers other than incompatibility preventing pollen from
reaching ovule
1) Genetic Male Sterility:

1. The pollen sterility, which is caused by nuclear genes, is termed as

genetic male sterility.

2. It is usually governed by a single recessive gene ms, but dominant

gene governing male sterility are also known Eg. Safflower.

3. The male sterility alleles may rise spontaneously or it can be

induced artificially and is found in several crops viz. Pigeon pea,
castor, tomato, barley, cotton, etc.

4. A male sterile line may be maintained by crossing it with

heterozygous male fertile plant, such a mating produces 1:1 male
sterile and male fertile plants.
 Genetic Male Sterility

Nuclear gene Nuclear gene

msms
ssssss S Cytoplasm X MSMS
ssssss S/F Cytoplasm

Male sterile Male fertile

Nuclear gene

F1 MSms Cytoplasm
ssssss S

Male fertile
Utilization in Plant Breeding:

1) Genetic male sterility is usually recessive and monogenic hence can

be used in hybrid seed production.

2) It is used in both seed propagated crops and vegetatively

propagated species.

1) The genotypes of msms and Msms lines are identical except for the
‘ms’ locus i.e. they are isogenic and are known as male sterile A)
Maintainer B) Line respectively.
Thermo sensitive genetic male sterility

Plants are sterile when temperature exceeds 32 0c/ 24 0c (day/night)

and become fertile when the temp. is below 24 0c/ 18 0c (day/night).

Photoperiod sensitive genetic male sterility

Within temperature, complete sterility is obtained in rice plants

grown under long-day condition (day length more than 13 hr 45 min),
but under short conditions almost normal fertility is obtained.
2) Cytoplasmic Male Sterility:

The pollen sterility which is controlled by cytoplasmic genes is known

as cytoplasmic male sterility (CMS).

Usually the cytoplasm of zygote comes primarily from the eggs cell
and due to this progeny of such male sterile plants would always be
male sterile.
 Cytoplasmic Male Sterility:

Nuclear gene Nuclear gene

rr
ssssss S Cytoplasm X ssssssrr F Cytoplasm

Male sterile Male fertile

Nuclear gene

F1 rr Cytoplasm
ssssss S

Male sterile
Transfer of cytoplasmic male sterility into new strain
CMS may be transferred easily to a given strain by using that strain as a
pollinator (recurrent parent) in the successive generation of backcross
programme.

After 6-7 backcrosses the nuclear genotype of male sterile line would be
almost identical to that of the recurrent pollinator strain.

The male sterile line is maintained by crossing it with pollinator strain

used as a recurrent parent in backcross, since the nuclear genotype of
the pollinator is identical with that of the new male sterile line.

Such a male fertile line is known as maintainer line or ‘B’ line and ‘male
sterile line is also known as ‘A ‘ line.
Utilization in Plant Breeding:

CMS has limited application.

It cannot be used for development of hybrid, where seed is the

economic product.

But it can be used for producing hybrid seed in certain ornamental

species or asexually propagated species like sugarcane, potato, and
forage crops.
3) Cytoplasmic Genetic Male Sterility:

When pollen sterility is controlled by both cytoplasmic and nuclear

genes is known as cytoplasmic and nuclear genes is known as
cytoplasmic genetic male sterility.

Jones and Davis first discovered this type of male sterility in 1944 in
onion.

The fertility restore gene ‘R’ is dominant and found in certain strains of
the species. This genes restores male fertility in the male sterile line,
hence is known as restores gene.
This system includes A, B, and R lines.

A line is a male sterile line,

B is similar to ‘A’ in all features but it is a male fertile and

R is restore line it restore the fertility in the F1 hybrid.

since B line is used to maintain the fertility and is also referred as

maintainer line.
 Nuclear gene Cytoplasm
Fertile Sterile
MSMS F F

MSms F F

msms F S

For sterility both cytoplasm and nuclear gene should be responsible.

If any one out of these, govern the expression for fertility then plant will
be fertile.
The most important line in CGMS is male sterile line used in hybrid
seed production.

And genetic constitution is msms called ‘’A’’ line.

Nuclear gene

msms
ssssss S
Cytoplasm

Male sterile line or ‘’A’’ line

 For seed production of ‘’A’’ line an isogenic line is required i.e its genetic
constitution is exactly same as ‘’A’’ line, but in place of sterile cytoplasm
it has fertile cytoplasm.

Its expression is male fertile called ‘’B’’ line or maintainer line or

isogenic line.

Nuclear gene

Nuclear gene
msms
ssssss S
Cytoplasm

msms
ssssss F
Cytoplasm

Male sterile line or ‘’A’’ line

Male fertile line or ‘’B’’ line

 Seed production of ‘’A’’ line

Nuclear gene Nuclear gene

msms
ssssss S Cytoplasm X msms
ssssss F Cytoplasm

Male sterile line or ‘’A’’ line Male fertile line or ‘’B’’ line

Nuclear gene

msms
ssssss S Cytoplasm

Male sterile line or ‘’A’’ line

How to maintain ‘’B’’ line?

Nuclear gene

msms
ssssss F Cytoplasm

Male fertile line or ‘’B’’ line

 Hybrid seed production

Nuclear gene Restorer gene

msms
ssssss S Cytoplasm X MSMS
ssssss S/F Cytoplasm

Male sterile line or ‘’A’’ line Male fertile, Restore (R) line

Nuclear gene

F1 MSms
ssssss S Cytoplasm

Male fertile
Utilization in Plant Breeding:

Cytoplasmic genetic male sterility is widely used for hybrid seed

production of both seed propagated species and vegetatively
propagated species.

It is used commercially to produce hybrid seed in maize, Bajara,

cotton, rice, sunflower, jowar, etc.
Transgenic Male Sterility

When the male sterility is induced by the techniques of genetic

engineering, it is called as transgenic male sterility.

It is heritable and basically comes under genetic male sterility.

In this system, the two kinds of genes are involved.

One gene causes male sterility (integrated with genome of A line) while
the other suppresses it (in R line).
Chemical Induced Male Sterility

The chemical which induces male sterility artificially is called as male

gametocide.
It is rapid method but the sterility is non-heritable.

In this system A, B and R lines are not maintained.

Some of the male gametocides used are gibberellins (rice, maize),

Sodium Methyl Arsenate (rice) and Maleic hydrazide (wheat, onion).
 Fundamentals of Plant Breeding

Plant breeding is an art and science of changing the genetics of plants for the benefit of
humankind. Plant breeding can be accomplished through many different techniques ranging
from simply selecting plants with desirable characteristics for propagation, to more complex
molecular techniques Plant breeding has been practiced for thousands of years, since near the
beginning of human civilization. It is now practiced worldwide by individuals such as
gardeners and farmers, or by professional plant breeders employed by organizations such
as government institutions, universities, crop-specific industry associations or research
centers.
International development agencies believe that breeding new crops is important for
ensuring food security by developing new varieties that are higher-yielding, resistant to pests
and diseases, drought-resistant or regionally adapted to different environments and growing
conditions.
Nature of Plant Breeding
Plant breeding is an art or science and is as old as agriculture, started since man learnt to
cultivate the plants. In earlier days, man depends on his skill and judgement in selecting
better plants. His knowledge about the plant was very limited. He knew nothing about the
inheritance of characters, role of environment in producing them and the basis of variation in
various plant characters. His method of selection was designed without the understanding of
the principle of inheritance. Therefore, during primitive time plant breeding was largely an
art and very less science was involved in that, but the present breeding methods are entirely
based on the scientific principles of plant sciences, particularly of genetics a cytogenetic.
Thus, plant breeding is purely science with very little art involved. Science is the knowledge
gathered through scientific method. The scientific method consists of observation,
formulation of hypothesis, experimentation and conclusion either to accept or reject the
hypothesis.
Plant breeding is considered as the current phase of crop evolution. As the knowledge of
genetics and other related science progresses plant breeding become less art and more
science. Especially discovery of Mendel‟s work in 1900, added lot to the knowledge of
science. Selection of desirable plant even today is an art it depends on the skill of a person
but alone skill is not enough, modern plant breeding is based on through understanding and
use of genetics principles. To be successful, a plant breeder must know each and everything
about the crop with he is working. He should have an understanding of principles of
difference disciplines viz. Genetics, Cytology, Morphology and Taxonomy, Plant Physiology,
Plant Pathology, Entomology, Agronomy, and Soil Science, Biochemistry, Statistics, and
Biometrics, Computer and Plant biotechnology. Thus, plant breeding is an art science and a
technology of developing genetically superior plants in terms of the economics utility for the
mankind.
Objectives of Plant Breeding:
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.
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. e.g. 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.
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. Biotic 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 desira ble 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 germination. 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 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.
Major achievements of Plant Breeding
Development of dwarf and semi dwarf cereal varieties
Many dwarf and semi dwarf varieties are developed in crop like wheat and Rice. Dr. Borlaug
used a Japanese variety NORIN-10 as a source of dwarfing gene, in wheat at CIMMYT
(Mexico). In 1963, ICAR has introduced some dwarf selection from CMMYT. Variety
Kalyansona and Sonalika were selected from these materials. In India, majority of the wheat
varieties grown are semi dwarf, and are resistant to water lodging, responsive to fertilizer
doses etc. Similarly, the development of semi dwarf varieties of rice has revolutionized rice
cultivation. These varieties were developed by introducing the gene Dee-Geo-Woo-Gene. e.g.
TN1 variety developed at Taiwan and IR-8 at IRRI Philippines, both were introduced in India
in 1966.
Noblisation of Indian Canes
The Indian canes were of Saccharum barberi, largely grown in North India. They were hardy
but poor in yield and sugar content, while tropical cane of Saccharum officinarum had thicker
stem and higher sugar content but, it performed badly in North India due to low winter
temperature. C.A. Barber and T.S Venkatraman at Sugarcane Breeding Institute, Coimbtore
transferred thick stem higher sugar content and other desirable characters from the noble cane
to Indian cane is commonly referred as nobilization on of Indian canes.
Development of Hybrid and Synthetic varieties in millets
Development of hybrid varieties in maize, sorghum and pearl millet enhanced the production
of these crops in the country. First maize hybrid was released in India in 1957.
a) Maize- Canga series of hybrid, Ganga safed-2, African Tall, Manjari, Deccan etc.
b) Sorghum -CSH-1,2,3,4,5,6,7,8,910,12,14 and 15 R.
c) Bajra-WCC-75, PHB-10, ICTP-8203, Shradha and Saburi.
Hybrid cotton
First cotton hybrid was developed in India in 1970 named H4 at Surat station of Gujrat
Agriculture University. After that many intervarietal and interspecific cotton hybrids were
developed which further increased the area under hybrid cotton and enhanced the production
of cotton in the country. e.g. H-4, Varalaxmi, Savitri, NH-44, Jaylaxmi, etc.
Molecular Breeding
Use of molecular biology techniques to supplement conventional plant breeding further
enhanced the efficiency of various breeding methods. Marker assisted selection or marker
aided selection (MAS) is a process whereby a marker (morphological, biochemical or one
based on DNA/RNA variation) is used for indirect selection of a genetic determinant or
determinants of a trait of interest (i.e. productivity, disease resistance, abiotic stress tolerance,
and/or quality). Many varieties in rice, maize and wheat were developed with help of MAS
and released for commercial cultivation in India. e.g. Improved Pusa Basmati1, Improved
Samba Mahsuri in rice, maize hybrid Vivek QPM9 etc.
Transgenic varieties
Recombinant DNA technology is most powerful tool for creating crops with novel desired
phenotypes or designer crops. Genetic modification of plants is achieved by adding a specific
gene or genes to a plant, or by knocking out a gene with RNAi, to produce a
desirable phenotype. The plants resulting from adding a gene are often referred to as
transgenic plants. Genetic modification can produce a plant with the desired trait or traits
faster than classical breeding because the majority of the plant's genome is not altered.
The majority of commercially released transgenic plants, are currently limited to plants that
have introduced resistance to insect pests and herbicides. Insect resistance is achieved
through incorporation of a gene from Bacillus thuringiensis (Bt) that encodes a protein that is
toxic to some insects. For example, the cotton bollworm, a common cotton pest, feeds on Bt
cotton it will ingest the toxin and die. Herbicides usually work by binding to certain plant
enzymes and inhibiting their action. The enzymes that the herbicide inhibits are known as the
herbicides target site. Herbicide resistance can be engineered into crops by expressing a
version of target site protein that is not inhibited by the herbicide. This is the method used to
produce glyphosate resistant crop plants Genetic modification of plants that can produce
pharmaceuticals (and industrial chemicals), sometimes called pharma crops, is a rather
radical new area of plant breeding.
Undesirable consequences
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/Genetic erosion: 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 manmade 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.
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 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.
Different phases of Plant Breeding
1. Domestication
The process of bringing 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.
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. This map shows the sites of domestication for a
number of crops. Places where crops were initially domesticated are called centers of origin
Plant breeding in certain situations may lead the domestication of wild
plants. Domestication of plants is an artificial selection process conducted by humans to
produce plants that have more desirable traits than wild plants, and which renders them
dependent on artificial (usually enhanced) environments for their continued existence. The
practice is estimated to date back 9,000-11,000 years. Many crops in present day cultivation
are the result of domestication in ancient times, about 5,000 years ago in the Old World and
3,000 years ago in the New World. In the Neolithic period, domestication took a minimum of
1,000 years and a maximum of 7,000 years. Today, all of our principal food crops come from
domesticated varieties.
A plant whose origin or selection is due primarily to intentional human activity is called
a cultigen, and a cultivated crop species that has evolved from wild populations due to
selective pressures from traditional farmers is called a landrace. Landraces, which can be the
result of natural forces or domestication, are plants (or animals) that are ideally suited to a
particular region or environment. An example are the landraces of rice, Oryza
sativa subspecies indica, which was developed in South Asia, and Oryza
sativa subspecies japonica, which was developed in China.
2. Selection
i.) Mass selection
ii.) Pureline selection
iii.) Recurrent selection
3. Hybridization
i.) Intervarietal/Intraspecific
ii.) Distant Hybridization
a) Interspecific/Intrageneric
b) Intergeneric
c) Somatic hybridization
4. Creation of novel genetic variations
i) Mutation
ii.) Polyploidy
iii.) Somaclonal variation
iv.) Gametoclonal variation
5. Genetic engineering/Recombinant DNA technology
Activities of Plant Breeding
The desired changes in the 'genotypes of crop species mi the consequent benefits to the
farmers are brought about by a series interrelated and largely interdependent activities. Major
activities of plant breeding are following;
1. Creation - variation
2. Selection
3. Evaluation
4. Release
5. Multiplication
6. Distribution of the new variety
Creation of variation: Genetic variation is a prerequisite for any improvement in a crop.
Therefore, in any breeding programme, this is always the first step unless variation pre-exists.
Genetic variation can be created by domestication, germplasm collections, plant introduction,
hybridization, mutation, polypolidy, somaclonal variation and genetic engineering.
Selection: The next step consists of identification and isolation of plants having the desirable
combinations of characters, and growing their progeny, this is called selection. Selection is
necessarily based on phenotype. The efficiency of this activity determines the success of a
breeding programme. Various breeding methods have been designed to increase the efficacy
of selection. Selection finally yields an improved line/ stream of population.
Evaluation: The newly selected lines/strains/population are tested for yield and other traits
and their performance is compared with the existing best varieties called checks. Evaluation
is a stepwise process, ordinarily conducted at several locations for three or more years under
the concerned. All India coordinated crop improvement project. If the new
line/strain/population is superior to the checks, it is released and notified as the new variety
and its seed can now be multiplied and, more importantly, certified by a seed certification
agency for quality.
Multiplication: This step concerns with the large scale production of certified seed of the
released and notified variety. Seed production is usually done by seed production agencies in
a step wise manner, and the seed is certified by a seed certification agency.
Distribution: Certified seed is ultimately sold to the farmers who use it for commercial crop
cultivation. This activity alone makes it possible to reap the economic benefits from the
above activities in form of: (i) an enhanced and (ii) Stable production of (iii) Superior
produce (iv) often at a lower cost.
MODE OF REPRODUCTION
Knowledge of the mode of reproduction and pollination is essential for a plant breeder,
because these aspects help in deciding the breeding procedures to be used for the genetic
improvement of a crop species. Choice of breeding procedure depends on the mode of
reproduction and pollination of a crop species.
Reproduction refers to the process by which living organisms give rise to the offspring of
similar kind (species). In crop plants, the mode of reproduction is of two types: viz. 1)
Asexual reproduction and 2) Sexual reproduction
I. Asexual reproduction
Multiplication of plants without the fusion of male and female gametes is known as asexual
reproduction. Asexual reproduction can occur either by vegetative plant parts or by
vegetative embryos which develop without sexual fusion (apomixis). Thus, asexual
reproduction is of two types: viz. a) vegetative reproduction and b) apomixis.
Vegetative reproduction refers to multiplication of plants by means of various vegetative
plant parts. Vegetative reproduction is again of two types: viz. i) natural vegetative
reproduction and ii) artificial vegetative reproduction.
Natural vegetative reproduction
In nature, multiplication of certain plants occurs by underground stems, sub aerial stems,
roots and bulbils. In some crop species, underground stems (a modified group of stems) give
rise to new plants. Underground stems are of four types: viz. rhizome, tuber, corm and
bulb. The examples of plants which reproduce by means of underground stems are given
below:
Rhizome: Turmeric (Curcuma domestica), Ginger (Zingiber officinale)
Tuber: Potato (Solanum tuberosum)
Corm: Arvi (Colocasia esculenta), Bunda (C. antiquorum)
Bulb: Garlic (Allium sativum), onion (A. cepa)

Rhizome: Turmeric Tuber: Potato Bulb: Onion

Sub aerial stems include runner, sucker, stolon, etc. These stems lead to vegetative
reproduction in mint (Mentha spp) rose, strawberry, banana, etc.
Bulbils are modified forms of flower. They develop into plants when fall on the
ground. Bulbils are found in garlic.
Artificial vegetative reproduction
Multiplication of plants by vegetative parts through artificial method is known as artificial
vegetative reproduction. Such reproduction occurs by cuttings of stem and roots, and by
layering and grafting. Examples of such reproduction are given below:
Stem cuttings: Sugarcane (Saccharum sp.) grapes (Vitis vinifera), roses, etc.
Root cuttings: Sweet potato, citrus, lemon, etc.
Layering and grafting are used in fruit and ornamental crops.
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
Apomixis refers to the development of seed without sexual fusion (fertilization). In apomixis
embryo develops without fertilization. Thus apomixis is an asexual means of
reproduction. Apomixis is found in many crop species. Reproduction in some species occurs
only by apomixis. This apomixis is termed as obligate apomixis. But in some species
sexual reproduction also occurs in addition to apomixis. Such apomixis is known
as facultative apomixis.
There are two types of apomixis: viz.
A) Adventive embryony
B) Gametophytic apomixis
a) Apospory
b) Diplospory
i) Pathenogenesis
ii) Peudogamy
Adventive embryony
The development of embryo directly from the diploid cells of ovule lying outside the
embryosac belonging to either nucellus or integuments is referred to as adventive embryony.
Gametophytic apomixis
In this form of apomixis, embryo develop without fertilization from egg cell or other cells of
embryo sacs. In recurrent apomixis, unreduced embryo sacs are produced by a process of
apomeiosis, which is a collective term used for various substitutes for meiosis that give rise to
unreduced embryo sacs. Apomeiosis is of two main types:
Apospory
Some vegetative cells of ovule develop into unreduced embryo sacs through a series of
mitotic divisions and without meiosis, e.g., in some species of Malus, orchids etc.
Diplospory
Embryo sac is produced from the megaspore, which may be haploid or, more generally,
diploid. In apomictic species, the meiosis is so modified that the megaspore remain diploid.
In all such cases, either meiosis is omitted altogether or restitution of unreduced chromosome
number occur during first meiotic division. The embryos in such embryo sacs may arise by
either i) Parthenogenesis ii) Pseudogamy
i) Parthenogenesis
Embryo develop from the embryo sac without pollination. Parthenogenesis is of following
two types:
Gonial parthenogenesis: Embryo develop from egg cell.
Somatic parthenogenesis: Embryo develop from some other cells of embryo sac.
ii) Pseudogamy
In such cases, pollination is necessary for embryo development, but fertilization of egg cell
does not take place. Fertilization of the secondary nucleus, however, does occur and is
necessary for endosperm development. Pseudogamy is also of two types:
Gonial pseudogamy: Embryo develop from egg cell
Somatic pseudogamy: Embryo develop from some other cells of embryo sac.
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.
II SEXUAL REPRODUCTION
Sexual reproduction involves fusion of male and female gametes to form a zygote, which
develops into an embryo. In crop plants, male and female gametes are produced in specialised
structures known as flowers.
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.
Sporogenesis
Productions of microspores and megaspores is known as sporogenesis. Microspores are
produced in anthers (microsporogenesis), while megaspores 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. 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 degenerates leaving
one functional megaspore per ovule. 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 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.
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.
Fertilization:
The fusion of one of the two sperms with the egg cell producing a diploid zygote is known as
fertilization. The fusion of the remaining sperm with the secondary nucleus leading to the
formation of a triploid primary endosperm nucleus is termed as triple fusion. The primary
endosperm nucleus after several mitotic divisions develops into mature endosperm, which
nourishes the developing embryo.

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 through 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.
Mode of pollination
The process by which pollen grains are transferred from anthers to stigma is referred as
pollination. Pollination is of two types: viz. 1) Autogamy or self-pollination and 2) Allogamy
or cross pollination.
I. Autogamy
Transfer of pollen grains from the anther to the stigma of same flower is known as autogamy
or self-pollination. Autogamy is the closest form of inbreeding. Autogamy leads
to homozygosity. Such species develop homozygous balance and do not exhibit significant
inbreeding depression.
Mechanism promoting self-pollination
1. Bisexuality. Presence of male and female organs in the same flower is known as
bisexuality. The presence of bisexual flowers is a must for self-pollination. All the self-
pollinated plants have hermaphrodite flowers.
2. Homogamy. Maturation of anthers and stigma of a flower at the same time is called
homogamy. As a rule, homogamy is essential for self-pollination.
3. Cleistogamy. When pollination and fertilization occur in unopened flower bud, it is known
as cleistogamy. It ensures self-pollination and prevents cross pollination. Cleistogamy has
been reported in some varieties of wheat, barley, oats and several other grass species.
4. Chasmogamy. Opening of flowers only after the completion of pollination is known as
chasmogamy. This also promotes self-pollination and is found in crops like wheat, barley,
rice and oats.
5. Position of Anthers. In some species, stigmas are surrounded by anthers in such a way
that self-pollination is ensured. Such situation is found in tomato and brinjal. In some
legumes, the stamens and stigma are enclosed by the petals in such a way that self-pollination
is ensured. Examples are greengram, blackgram, soybean, chickpea and pea.
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.
II. Allogamy
Transfer of pollen grains from the anther of one plant to the stigma of another plant is called
allogamy or cross pollination. This is the common form of out-breeding. Allogamy leads to
heterozygosity. Such species develop heterozygous balance and exhibit significant
inbreeding depression on selfing.
Mechanism promoting cross-pollination
1. Dicliny. It refers to unisexual flowers. This is of two types: viz. i) monoecy and ii)
dioecy. When male and female flowers are separate but present in the same plants, it is
known as monoecy. In some crops, the male and female flowers are present in the same
inflorescence such as in mango, castor and banana. In some cases, they are on separate
inflorescence as in maize. Other examples are cucurbits, grapes, strawberry, cassava and
rubber. When staminate and pistillate flowers are present on different plants, it is
called dioecy. It includes papaya, date palm, spinach, hemp and asparagus.
2. Dichogamy. (from the Greek dikho-apart and gamous-marriage) It refers to maturation of
anthers and stigma of the same flowers at different times. Dichogamy promotes cross
pollination even in the hermaphrodite species. Dichogamy is of two types: viz. i) protogyny
and ii) protandry. When pistil matures before anthers, it is called protogyny such as in pearl
millet. When anthers mature before pistil, it is known as protandry. It is found in maize,
sugarbeet and several other species.
3. Heterostyly. When styles and filaments in a flower are of different lengths, it is called
heterostyly. It promotes cross pollination, such as linseed.
4. Herkogamy. Hinderance to self-pollination due to some physical barriers such as
presence of hyline membrane around the anther is known as herkogamy. Such membrane
does not allow the dehiscence of pollen and prevents self-pollination such as in alfalfa.
5. Self-incompatibility: The inability of fertile pollens to fertilize the same flower is
referred to as self-incompatibility. It prevents self-pollination and promotes cross
pollination. Self-incompatibility is found in several crop species like Brassica, Radish,
Nicotiana, and many grass species. It is of two types sporophytic and gametophytic.
6. Male sterility: In some species, the pollen grains are not functional. Such condition is
known as male sterility. It prevents self-pollination and promotes cross pollination. It is of
three types: viz. genetic, cytoplasmic and cytoplasmic genetic. It is a useful tool in hybrid
seed production.
Study of floral biology and aforesaid mechanisms is essential for determining the mode of
pollination of various crop species. Moreover, if selfing has adverse effects on seed setting
and general vigour, it indicates that the species is cross pollinated. If selfing does not have
any adverse effect on these characters, it suggests that the species is self-pollinated.
The percentage of cross pollination can be determined by growing a seed mixture of two
different varieties together. The two varieties should have marker characters say green and
pigmented plants. The seeds are harvested from the recessive (green) variety and grown next
year in separate field. The proportion of pigmented plants in green variety will indicate the
percentage of outcrossing or cross pollination.
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
Often Cross-Pollinated Species
In many crop plants, cross-pollination often exceeds 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.
Significance of pollination
The mode of pollination plays an important role in plant breeding. It has impact on five
important aspects viz. 1) gene action, 2) genetic constitution, 3) adaptability, 4) genetic purity
and 5) transfer of genes.
Classification of crop plants based on mode of pollination and mode of reproduction
Mode of pollination and Examples of crop plants
reproduction
A. Autogamous Species
1. Seed Propagated Rice, Wheat, Barley, Oats, Chickpea, Pea, Cowpea, Lentil,
Green gram, Black gram, Soybean, Common bean, Moth
bean, Linseed, Sesame, Khesari, Sunhemp, Chillies, Brinjal,
Tomato, Okra, Peanut, etc.
2. Vegetatively Propagated Potato
B. Allogamous Species
1. Seed Propagated Corn, Pearlmillet, Rye, Alfalfa, Radish, Cabbage,
Sunflower, Sugarbeet, Castor, Red clover, White clover,
Safflower, Spinach, Onion, Garlic, Turnip, Squash,
Muskmelon, Watermelon, Cucumber, Pumpkin, Kenaf,
Oilpalm, Carrot, Coconut, Papaya, etc.
2. Vegetatively propagated Sugarcane, Coffee, Cocoa, Tea, Apple, Pears, Peaches,
Cherries, grapes, Almond Strawberries, Pine apple, Banana,
Cashew, Irish, Cassava, Taro, Rubber, etc.
C. Often Allogamous Species Sorghum, Cotton, Triticale, Pigeonpea, Tobacco.

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 affected, 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 between pin and thrum plants would produce Ss and ss progeny in equal
frequencies. This system is of little importance in crop plants; it occurs in sweet potato
and buckwheat.

Homomorphic System
In the homomorphic system, incompatibility is not associated with morphological
differences among flowers. The incompatibility reaction of pollen may be controlled by
the genotype of the plant on which it is produced or by its own genotype.
Gametophytic System
Gametophytic incompatibility was first described by East and Mangelsdorf in 1925 in
Nicotiana sanderae. The incompatibility reaction of pollen is determined by its own
genotype, and not by the genotype of the plant on which it is produced. Generally,
incompatibility reaction is determined by a single gene having multiple alleles, e.g.,
Trifolium, Nicotiana, Lycoperscion, Solanum, Petunia etc. Pollen tube grows very slowly
in the style containing the same S allele as the pollen, and fails to effect fertilization.
Therefore, all the plants are heterozygous at the S locus. In a single gene system, there are
three types of mating:
i) Fully incompatible, e.g., S1S2 x S1S2
ii) Fully compatible, e.g., S1S2 x S3S4
iii) Partially (i.e., 50% of the pollen) compatible, e.g., S1S2 x S2S3

Sporophytic System
In the sporophytic system also, the self-incompatibility is governed by a single gene, S,
with multiple alleles; more than 30 alleles are known in Brassica oleracea. In general, the
number of S alleles is considerably larger in the gametophytic than in the sporophytic
system. The incompatibility reaction of pollen is governed by the genotype of the plant on
which the pollen is produced, and not by the genotype of the pollen. It was first reported
by Hughes and Babcock in 1950 in Crepis foetida, and by Gerstel in Parthenium
argentatum (in the same year). In the sporophytic system, the S alleles may exhibit
dominance, individual action (codominance) or competition. In many cases, different S
alleles vary in their activity leading to varying degrees of self-incompatibility, e.g., B.
oleracea. Polygenes (modifying genes) are known to increase as well as decrease the
activities of S alleles both in the gametophytic as well as sporophytic systems.
Mechanism of Self-Incompatibility
The mechanism of self-incompatibility is quite complex and is poorly understood. The
various phenomena observed in self-incompatible matings are grouped into three broad
categories:
(1) pollen-stigma interaction,
(2) pollen tube-style interaction, and
(3) pollen tube-ovule interaction.
Pollen-Stigma Interaction
These interactions occur just after the pollen grains reach the stigma and generally prevent
pollen germination. At the time they reach stigma, pollen grains generally have two nuclei
in the gametophytic system, while they have three nuclei in the sporophytic system. This
was once considered to be the basis for the two incompatibility systems, but the available
evidence indicates otherwise. However, the structure of stigmatic surface appears to be
definitely involved in the differences between the two systems. In the gametophytic
system, the stigma surface is plumose having elongated receptive cells and is commonly
known as „wet‟ stigma. Incompatible pollen grains generally germinate on reaching the
stigma; the incompatibility reaction occurs at a later stage.
In the sporophytic system, the stigma is papillate and dry, and is covered with a hydrated
layer of proteins known as „pellicle‟. There is evidence that the pellicle is involved in
incompatibility reaction. Within few minutes of reaching the stigmatic surface, the pollen
releases an exine exudates which is either protein or glycoprotein in nature. This exudate
induces immediate callose formation in the papillae (which are in direct contact with the
pollen) of incompatible stigma. Often callose is also deposited on the young protruding
pollen tubes preventing any further germination of the pollen. Thus, in the sporophytic
system, stigma is the site of incompatibility reaction; once the pollen tube crosses the
stigmatic barrier, there is no further inhibition of pollen tube growth.
In the homomorphic sporophytic system, the incompatibility reaction of pollen is probably
due to the deposition of some compounds from anther tapetum on to the pollen exine.
Pollen Tube-Style Interaction
In most cases of the gametophytic system, pollen grains germinate and pollen tubes
penetrate the stigmatic surface. But in incompatible combinations, the growth of pollen
tubes is retarded within the stigma, e.g., in Oenothera, or a little later in the style, e.g., in
Petunia, Lycopersicon, Lilium etc. In the latter case, there is a cessation of protein and
polysaccharide synthesis in the pollen tubes, which leads to the degeneration of tube wall
and the bursting of pollen tube.
Pollen Tube-Ovule Interaction
In some cases, e.g., Theobromo cacao, pollen tubes reach the ovule and effect fertilization.
However, in incompatible combinations, embryos degenerate at an early stage of
development.
Exploitation of self-incompatibility in hybrid production
Self-incompatibility may be used in hybrid seed production. For this purpose,
(1) two self-incompatible, but cross-compatible, lines are interplanted; seed obtained from
both the lines would be hybrid seed.
(2) Alternatively, a self-incompatible line may be interplanted with a self-compatible line.
from this scheme, seed from only the self-incompatible line would be hybrid.
(3) Schemes for the production of double cross and triple cross hybrids have also been
proposed and their feasibility has been demonstrated in the case of brassicas. The
gametophytic system has been used, to a limited extent, for hybrid seed production in
clover, Trifolium (Leguminosae). In Solanaceae, the cultivated species are generally self-
fertile, and self-incompatibility is confined to wild species.
The sporophytic system has been exploited for hybrid seed production in brassicas
(Cruciferae), primarily by the Japanese seed companies. In Compositae, another
economically important family showing sporophytic self-incompatibility, the cultivated
varieties are generally self-fertile.
The use of self-incompatibility in hybrid seed production is hampered by several problems
mentioned below:
(1) Production and maintenance of inbred lines by hand pollination is tedium and costly.
(2) This raises the cost of hybrid seed.
(3) Continued selfing leads to a depression in self-incompatibility, and it unintentionally,
but unavoidably, selects for self-fertility.
(4) In the gametophytic system, continued inbreeding gives rise to new incompatibility
reactions, which may limit the usefulness of such inbreds as parents. (5) Environmental
factors, e.g., high temperature and high humidity etc., reduce or even totally overcome
self-incompatibility reaction leading to a high (30% or more) proportion of selfed seed.
(6) Bees often prefer to stay within a parental line, particularly when the parental lines
differ morphologically. This, in turn, increases the proportion of selfed seed.
7) Transfer of S alleles from one variety or, more particularly, species into another variety
or species is tedious and complicated. This has prevented the use of self-incompatibility in
hybrid seed production in Solanaceae and Compositae.
MALE STERILITY
Male sterility is characterized by non-functional pollen grains, while female gametes
function normally. It occurs in nature sporadically, perhaps due to mutation. Male sterility
is classified into three groups:
(1) Genetic,
(2) Cytoplasmic, and
(3) Cytoplasmic-genetic.
Genetic Male Sterility
Genetic male sterility is ordinarily governed by a single recessive gene, ms, but dominant
genes governing male sterility are also known, e.g., in safflower. Male sterility alleles
arise spontaneously or may be artificially induced. A male sterile line may be maintained
by crossing it with heterozygous male fertile plants. Such a mating produces 1:1 male
sterile and male fertile plants.
Utilization in Plant Breeding
Genetic male sterility may be used in hybrid seed production. The progeny from ms ms x
Ms ms crosses are used as female, and are interplanted with a homozygous male fertile (Ms
Ms) pollinator. The genotypes of ms ms and Ms ms lines are identical except for the ms
locus, i.e., they are isogenic; they are known as male sterile (A) and maintainer (B) lines,
respectively. The female line would, therefore, contain both male sterile and male fertile
plants; the latter must be identified and removed before pollen shedding. This is done by
identifying the male fertile plants in seedling stage either due to the pleiotropic effect of
the ms gene or due to the phenotypic effect of a closely-linked gene. Pollen dispersal from
the male (pollinator) line should be good for a satisfactory seed set in the female line.
however, generally pollen dispersal is poor and good, closely-linked markers are rare.
Rouging of male fertile plants from the female lines is costly as a result of which the cost
of hybrid seed is higher. Due to these difficulties, genetic male sterility has been exploited
commercially only in a few countries. In USA it is used in Castor. In India it was being
used in Redgram, but presently it is being used in safflower. Marker genes which are
linked to male sterility/fertility can be used to identify the male fertile plants before
flowering stage. For example, in Maize there is a gene, pigmented hypocotyl (P) and green
hypocotyl (P) which is closely linked with sterility locus
PS - Pigmented & Sterile
PF – Green & Fertile
At seedling stage all the green plants are to be removed and pigmented plants are retained,
as they are sterile.
Suggestions have been made for its use in several other crops, e.g., Cotton, barley, tomato,
sunflower, cucurbits etc., but it is not yet practically feasible.
Cytoplasmic Male Sterility
This type of male sterility is determined by the cytoplasm. Since the cytoplasm of a zygote
comes primarily from egg cell, the progeny of such male sterile plants would always be
male sterile. Nuclear genotype of male sterile line would be almost identical to that of the
recurrent pollinator strain. The male sterile line is maintained by crossing it with the
pollinator strain used as the recurrent parent in the backcross programme since its nuclear
genotype is identical with that of this new male sterile line. such a male fertile line is
known as the maintainer line or B line as it is used to maintain the male sterile line is also
known as the A line, there is considerable evidence that the gene or genes conditioning
Cytoplasmic male sterility, particularly in Maize, reside in mitochondria, and may be
located in a plasmic like elements.

Utilization in Plant Breeding

Cytoplasmic male sterility may be utilized for producing hybrid seed in certain ornamental
species, or in species where a vegetative part is of economic value. But in those crop
plants where seed is the economic part, it is of no use because the hybrid progeny would be
male sterile. Eg: Onion, Fodder Jowar, Cabbage, Palak etc.
Cytoplasmic-Genetic Male Sterility
This is a case of Cytoplasmic male sterility where a nuclear gene for restoring fertility in
the male sterile line is known. The fertility restorer gene, R, is dominant and is found in
certain strains of the species, or may be transferred from a related species, e.g., in wheat.
This gene restores male fertility in the male sterile line, hence it is known as restorer gene.

The cases of Cytoplasmic male sterility would be included in the Cytoplasmic-genetic

system as and when restorer genes for them would be discovered. It is likely that a restorer
gene would be found for all the cases of Cytoplasmic male sterility if a thorough search
were made. This system is known in Maize, Jowar, bajra, sunflower, rice, wheat, etc.
Plant would be male sterile in the presence of male sterile cytoplasm if the nuclear
genotype were rr, but would be male fertile if the nucleus were Rr or RR. New male sterile
lines may be developed following the same procedure as in the case of Cytoplasmic
system. For the production of hybrid seed, removal of anthers before fertilization is
essential to avoid selfing. Manually removing of anthers is very tedious and time-
consuming process in almost all the crops except in Maize and Castor which are
monoecious. The pre-requisites for successful hybrid seed production in large quantities
are:
1. Existence of male sterility or self-incompatibility through which hand emasculation
can be avoided.
2. Sufficient cross-pollination should be there to get good seed set.
Breeding Methods
The following are the methods of breeding self-pollinated crop plants.
1. Introduction
2. Selection a) Pure line selection b) Mass selection
3. Hybridization and selection i) Inter varietal a) Pedigree Method b) Bulk Method. c)
Single Seed Descent Method. d) Modified Bulk Method e) Mass - Pedigree Method.
ii) Interspecific hybridization
4. Back cross method
5. Multiline varieties
6. Population approach
7. Hybrids.
8. Mutation breeding
9. Polyploidy breeding
10. Innovative techniques

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

Schematic representation of Mass selection

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

Confirmatory evidence was obtained in three ways

In the first case, he classified the seed from each pureline into 100 mg classes, and grew them
separately. The mean seed weight of progenies from different seed weight class of a single
pure line were comparable with each other, and with that of the parent pureline. For example,
line no 13 had seed size classes of 200, 300, 400, and 500 mg. The mean seed weights of the
progenies derived from these seed weight classes were 475, 450, 451 and 458 mg
respectively.
The second line of evidence came from selection within a pureline. From each pureline, the
largest and the smallest seeds were selected to raise the next generation. In the subsequent
generations, large seeds were selected in the progenies obtained from large seeds while in
these from small seeds selection was done fro small seeds. Six generations of selection was
ineffective in increasing or decreasing the seed size. For example, after 6 generations of
selection, the mean seed weight in Line No 1 was 690 and 680 mg in the progenies selected
for small and large seeds respectively. Thus, selection within a pureline was ineffective.
The third approach was to estimate parent offspring correlation. The value of parent offspring
correlation within line no 13 was – 0.018+ 0.038, that is, zero, while it was 0.336 + 0.008 in
the original seed lot of the Princess which is highly significant. The parent-offspring
correlation will be zero when the variation is nonheritable, while it will be significantly
greater than zero when the variation has a genetic basis, i.e., is heritable.
These observations reveal that the variation for seed size in the original seed lot of Princess
had a genetic basis and was heritable. But the variation within the purelines obtained from
the single seeds selected from this seed lot was purely due to the environment and, therefore,
non-heritable.
The two main conclusions from the Johannsen’s experiment are:
1. A self-fertilized population consists of a mixture of several homozygous genotypes.
Variation in such a population has a genetic component, and therefore selection is effective.
2. Each individual plant progeny selected from a self-fertilized population consists of
homozygous plants of identical genotype. Such a progeny is known as pureline. The
variation within a pureline is purely environmental and, as a result, selection within a pureline
is ineffective.
Origin of variation in pure lines
Pure lines show genetic variation after some time because of the following reasons:
1. Mechanical Mixture: During cultivation, harvesting threshing and storage, other genotypes
may get mixed up.
2. Natural hybridization: Through pure lines are produced in self-pollinated crops, some
amount of natural cross pollination occurs in them also can be avoided by isolation and
rouging. 3. Mutation: occur spontaneously in nature at random
Characteristics of purelines
1. All the plants within a pureline have the same genotype
2. The variation within a pureline is environmental and non-heritable
3. Purelines are stable
Progeny test
Evaluation of the worth of plants on the basis of performance of their progenies is known as
progeny test. This was developed by Louis de Vilmorin and so it is also known as the
Vilmorin Isolation principle. Vilmorin worked on sugar beet plants. The progeny test serves
two valuable function:
1. Determines the breeding behaviour of a plant i.e. whether it is homozygous or
heterozygous.
2. Whether the character for which the plant was selected is heritable i.e. is due to genotype
or not. Selections have to be based or phenotype and so it is necessary to know the genotype
of the selected plant.
Pureline selection
Pureline selection has been the most commonly used method of improvement of self-
pollinated crops. Almost all the present day varieties of self-pollinated crops are purelines.
Pureline selection has several applications in improvement of self-pollinated crops. It is used
to improve:
1. Local varieties
2. Old pureline varieties and,
3. Introduced varieties
Procedure of Pureline selection
First year:
 An old variety or landrace is used for Pureline selection. Population they selected for

pureline selection is homozygous. Single plant is selected and harvested separately

superior plants must be selected from the mixed population. About 1000-2000 plants are
selected depending on the available resources.
Second year:
 The individual progenies are grown separately with proper spacing the top 15-20

progenies are selected and they are bulked. Poor, defective, weak and segregating
 progenies are discarded. Selection should be based on simply inherited character like
plant type, Plant height, grain type, flowering and maturity duration disease resistance
this process may be repeated
Third year:
 Seed of the individual plant progenies are not enough to conduct a replication trail. So,

they are grown in unreplicated trial with check. Here yield of progenies are taken as a
criteria for selection.
Fourth year:
 Replicated yield trials are conducted using the best available check variety. This may be

repeated for 2-3 year. All the observations are recorded

Fifth to Eighth year:
 Promising strains are evaluated at several locations along with strains or check. The best

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

Comparison between pureline and mass selection

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

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

Aims of hybridization
1. To transfer of one or few qualitative characters.
2. Improvement in one or more quantitative character.
3. F1 Hybrid as variety.
I. Combination breeding:
This method is used for the transfer of one or more character into
or single variety from another variety. Eg: improving the yield by correcting the
defect. i.e. disease resistance. The other parent selected for hybridization must have a
sufficient intensity of a character under transfer.
II. Transgressive breeding:
It aims at improving yield or its contributing character through
transgressive segregation. It refers to the appearance of such plants in F 2 generation
that are superior to both the parents for one or more character. It is due to
accumulation of plus or favourable genes from the parents as a consequence of
recombination. The parents used for crossing must combine very well and are
genetically diverse. So, pedigree breeding followed by population approach are
designed for production of transgressive segregants.
III. Hybrid varieties:
In self-pollinated crops F1 is more vigorous and high yielding than
the parents. Two parents should combine well to produce outstanding F 1 hybrid.
Types of hybridization
 Inter-varietal Hybridization / Intra specific: Parents involved in hybridization belong to

the same species. They may be two strains, varieties or races.

 Varietal crosses may be simple crosses or complex crosses
a.) Simple crosses: Two parents are crossed to produce F1 (A x B)
b.) Complex crosses: More than two parents are crossed to produce the hybrid (A x B) x C
x F1
Procedure of hybridization:
The breeder has clear cut objective in developing the variety. He has to select the variety
accordingly.
1. Choice of parents: One of the parents involved in crosses should be a well-adapted and
proven variety in the area. The other variety should be having the character that are absent in
this variety. Combining ability of the parents serves as useful guides in the selection of
parents, which produce superior F1 and F2.
2. Evaluation of parents: Parents are evaluated for their combining ability.
3. Emasculation: The removal of stamens/anther without affecting the female reproductive
organs, hand emasculation is mostly followed.
4. Bagging: Immediately after emasculation the flowers are enclosed in suitable bags to
prevent cross pollination.
5. Tagging: The emasculated flowers are tied with a thread. The information on date of
emasculation, date of pollination, names of female and male parents are recorded in the tag
with pencil. The name of the female parent is written first then male parent.
6. Pollination: Mature fertile and viable pollen from the male parent should be placed on
receptive stigma of emasculated flowers to bring about fertilization. Pollen grain is collected,
allowed for dehiscence and pollination is carried out with camel hair brush.
7. Harvesting and storing of F1 seeds: The crossed heads/pods should be harvested and
threshed. The seeds should be dried and properly stored to protect them from storage pests.
8. Rising the F1 generation: Identify the selfed seeds in the F1 generation by using dominant
marker gene. Larger F 1 population is desirable, because both the genes are present in
heterozygous condition.
9. Selfing: To avoid cross pollination and to ensure self-pollination. In often cross-pollinated
crops, they are bagged to prevent cross pollination.
Distant Hybridization
When crosses are made between two different species or between two different genera, they
are generally termed as distant hybridization (or) wide hybridization
History
Thomas Fairchild 1717 was the first man to do distant hybridization. He produced a hybrid
between two species of Dianthus Dianthus caryophyllus (Carnation) x D. barbatus (Sweet
william)
Inter generic hybrid produced by Karpechenko, a Russian Scientist in 1928. Raphano brassica
is the amphidiploid from a cross between Radish (Raphanus sativus) and cabbage (Brassica
oleracea). Triticale was produced by Rimpau in 1890 itself. Triticale is an amphidiploid
obtained from cross between wheat and rye. Another example is Saccharum noblisation
involving three species.
Hybrids in self-pollinated crops - problems and prospects
Exploitation of heterosis through F1 hybrids has hitherto been the prerogative of cross-
pollinated crops, chiefly due to their breeding systems favouring allogamy. However,
possibilities of working for such a proposition have recently been realized in self-pollinated
corps also. Indeed, exploitation of hybrid vigour in autogamous/ self-pollinated crops is easy
and less time consuming as homozygous inbreds are already available. There is practically no
difference with regard to hybrid breeding between self and cross-pollinated crops. But the
prospects of hybrids in self-pollinated crops are dependent on three major considerations:
1. How high a heterotic effect can be gained under optimal production conditions.
2. In fact, a breeder's main concern is the magnitude rather than the frequency of occurrence
of heterosis in crops. Thus, the consideration is whether or not it is possible to obtain
economically viable heterosis.
3. How much of the yield surplus due to high heterosis can offset the extra seed cost? In
major self-pollinated crops like wheat, barley, rice, etc., the seed rate per unit area is
exorbitant and hence the hybrid seed requirement is also more.
4. How efficient and effective is the mechanism of cross-pollination in self-pollinated crops?
By nature, self-pollinated crops are shy pollinators with very poor pollen manoeuvrability (or
movability to effect allogamy). Therefore, the efficiency (degree of allogamy) with which
cross pollination can take place on a commercial scale is the true determinant of the success
of a hybrid programme in self-pollinated crops.
5. Among self-pollinated crops, F1 hybrids have been graduated into the farmer's field in rice,
barely, tomato, Sorghum (often-cross-pollinated) and wheat.
Pedigree method of handling segregating generations
 In pedigree method individual plants are selected from F2 and their progenies are tested in

subsequent generations. A record of the entire parent off spring relationship is maintained
and known as pedigree record. The pedigree may be defined as a description of the
 ancestor of an individual and it generally goes back to some distant ancestor. So, each
progeny in every generation can be traced back to the F 2 plant from which it is originated.
 This method used for selection from segregating population of crosses in self-pollinated

crops. It is used for combination or transgressive breeding.

Procedure:
1. Hybridization: The selected parents are crossed to produce a simple / complex cross (F 1
seed)
2. F1 generation: F1 seeds are space planted to each produces maximum number of F 2 seed.
15-30 F1plants are sufficient to produce good F2 populations.
3. F2 generation: 200-10000 plants are space planted and 100-500 plants are selected and
their seeds are harvested separately. He should select as many as F 2 plants as he can
handle efficiently. The selection depends on skill of the breeder and his ability to judge to
select F2 which produce good progeny.
4. F3 generation: Individual plant progeny are space planted. Individual plant with
desirable characters from superior progenies is selected.
5. F4 generation: Individual plants progenies are space planted desirable pants are selected
undesirable progenies are rejected. Progenies are compared visually and more plants are
selected from superior progenies. Selection of desirable plants from superior
progenies selection is practiced within / between family.
6. F5 generation: Many families have reached homozygous and may be harvested in bulk.
The breeder has to assess the yielding potential of progenies, 25-100 progenies are
advanced and tested in preliminary yield trial.
7. F6 generation: Multi row plots and evaluated visually progenies harvested bulk and they
have become homozygous.
8. F7 generation: Preliminary yield trail with replication to identify the superior progenies.
Progenies are evaluated for other component character 2-5 outstanding lines superior to
check are advanced to multi location testing.
9. F8 –F10 generation: Replicated yield trial at several locations. They are tested for yield as
well as for resistance.
10. F11 generation: Seed multiplication and release.
Merits:
1. Maximum opportunity for the breeder to use his skill and judgment for the selection of
plants in segregating generation.
2. It provides information about the inheritance of qualitative character from the pedigree
record.
3. Chances of recovering transgressive segregants is more.
4. Plants and progenies with visible defects and weaknesses are eliminated at an early
stage.
5. Well suited for the improvement of easily identified and simply inherited characters.
Demerits:
1. Maintenance of accurate pedigree record is tedious and takes up valuable time
 2. Selection of progenies in every generation laborious, time consuming. Difficult to
handle many crosses.
3. No opportunity for natural selection.
4. Possibility of losing the valuable genotype is early segregating generation.
5. The success of this method largely depends on skill of the breeder.
Bulk method
Bulk method was first used by Nilsson Ehle in 1908. F2 and the subsequent generations are
harvested as bulks to raise the next generation. At the end of bulking period individual
plants are selected and evaluated in a similar manner as in the pedigree method. The
duration of bulking may vary from 7-30 generation artificial selection may seldom be
practiced
1. Isolation of homozygous lines
It is used for the isolation of homozygous lines with a minimum of effort and expense. The
population is carried to F6 -F7 as Bulk, where it reaches homozygosity. Individual plants are
selected and evaluated to derive pure line. So preliminary yield trails are conducted to
derive homozygous lines.
2. Waiting for the opportunity for selection:
Selection for resistance to disease, lodging and cold depends upon the presence of suitable
environmental conditions favouring epidemic. Waiting till such environment do occur so
the segregating generations are carried as bulk in such environment. Individual plants are
selected and handled as in pedigree method. The duration of bulking depends upon the
occurrence of the concerned environment. This is known as mass pedigree method of
Harlan.
3. Opportunity for natural selection:
Maintenance of bulk is inexpensive and without much efforts. Some bulk populations are
carried up to F20 to F30 to provide an opportunity for natural selection to act on their
composition. Up to F6 generation the population is heterozygous and after F7 generation
natural selection to act on homozygous plants and would change the frequency of
homozygous genotypes present in the population. It is assumed that
natural selection would favour higher yielding genotypes and eliminate poorer genotypes.
Procedure of bulk method:
1. Hybridization: Parents are selected and crossed
2. F1 generation: F1 is space planted more than 200 F1 plants
3. F2 -F6 Generation: Planted at commercial seed rate, spacing and harvested as bulk, during
this period. Frequency of population changes due to outbreak of disease or pest.
4. F7 generation: 50000 plants are space planted about 1000-5000 plants with phenotype is
selected and the seeds are harvested separately.
5. F8 generation: Individual plant progenies are single/multi row plants, since progenies are
homozygous and harvested in bulk weak and inferior progenies are rejected and 100-
300 individual plant progenies with desirable characters.
6. F9 generation: Preliminary yield trial with standard check, yield and quality parameter is
taken for selection.
7. F10---F12 generations: Replicated yield trails are conducted. Yield and its component
characters are evaluated along with the check. Superior progenies are released as variety
8. F13 generation: Seed multiplication of the newly released variety and distribution to
farmers.
Merits
1. Simple, convenient and inexpensive method
2. Natural selection is likely to increase the frequency of superior genotypes in the
population. Therefore, progenies selected from long term bulks are likely to be superior to
those selected from F2 or short bulks.
3. Little work and attention are required in F2 and subsequent generations, and no pedigree
record is to be kept. This save time and labour, and the breeder can concentrate more on
other breeding projects.
4. Since large populations are grown, chances of getting transgressive segregants are more.
5. Individual plant selection is done when population has become homozygous. Therefore,
selection is expected to be more effective than in F2 and F3 generations.
6. Particularly suited to small grain crops grown in high crop densities.
7. Natural selection is expected to improve characters like adaptation to prevailing
environment which are otherwise difficult to assess and select for.
Demerits
1. The major disadvantage of bulk method is longer time taken to develop a new variety.
Natural selection becomes important only after F10 generation and bulking may have to be
done up to F20 or more which is considerably longer than the time taken in pedigree
method
2. In short term bulks, natural selection has little effect on the genetic composition of
populations. But short-term bulks are useful for isolation of homozygous lines.
3. It provides little opportunity to breeders to exercise their skill in selection.
4. A large number of progenies have to be handled at the end of the bulking period.
5. Information on inheritance of characters cannot be obtained
6. Off season and green house facilities cannot be used to advance the generation since
environment at such locations may be markedly different from that in target location.

Single seed decent method

 Single seed descent method This method is a modification of bulk method
 Single seed from each F2 plants is bulked to raise the F3 generation. Similarly, F3, F4 , F5

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

Back cross method

A Crossing between a F1 hybrid or its segregating generation with one of its parents is
known as Back cross. The hybrid and its progenies in the subsequent generations are
repeatedly back crossed to one of their parents. As a result, the genotype of back cross
progeny becomes increasingly similar to that parent to whom the back crosses are made.
At the end of 6-8 back crosses, the progeny would be almost identical with the parent
involved in back crossing.
Objective:
1. To improve one or two specific defects of a high yielding variety and a well-adapted
variety with desirable character.
2. The characters lacking in this variety are transferred to it from a donor parent without
changing the genotype of this variety except for the genes being transformed.
Requirements of back cross breeding
1. Suitable recurrent parent must be available which lacks in one or two characteristics.
2. A suitable donor parent must be available; the character must be highly intense.
3. The character(s) to be transferred must have high heritability and preferably, should be
governed by one or few genes
4. A sufficient number of back crosses should be made so that genotype of recurrent parent
is recovered in full. Ordinarily, 6-7 backcrosses are sufficient for the purpose.
Applications of back cross breeding
1. Inter varietal transferring of simply inherited traits. Characters governed by one or two
genes like disease resistance are successful.
2. Inter varietal transfer of quantitative characters and highly heritable quantitative characters
like earliness, plant height, seed size and seed shape are transferred.
3. Inter specific transfer of simply inherited characters: Disease resistance is transferred from
related species to cultivated species. Inter specific transfer of genes are easy when the
chromosome of the two species pair regularly.
4. Transferring of cytoplasm: wild species cytoplasmic are transferred to cultivated species
transfer of male sterility. The variety or species from which the cytoplasm is to be
transferred is used as the female parent. The parent to which the cytoplasm is to be
transferred is used as the male parent in the original cross and back cross. After 6 -8 back
crosses the progeny would have the nuclear genotype of the recurrent parent and the
cytoplasm from the donor parent.
5. Transgressive segregation: F1 is back crossed to one or two times to the recurrent parent
leaving much heterozygosity for transgressive segregation to appear. In the second
modification two or more recurrent parent may be used in the back-cross progeny to
accumulate genes from them into the back cross. Progeny of the new variety is not exactly
like any one of the recurrent parents.
6. Production of isogenic lines: Isogenic lines are identical in their genotype except for one
gene
7. Germplasm conversion: When valuable germplasm cannot be utilized in breeding
programmes and may be used as recurrent parent in separate back cross programme these
lines are called converted lines.
Procedure for transfer to dominant gene
E.g. High yielding and widely adapted wheat variety A is susceptible to stem rust another
variety B is resistance to stem rust. Stem rust is dominant to susceptibility.
1. Hybridization: Variety A is crossed to variety B. Generally, variety A should be used as
female parent. This would help in identification of selfed plants.
2. BC1 generation: F1 plants are back crossed to variety A. Since all the F1 are
heterozygous for rust resistance, selection for rust resistance is not necessary.
3. First BC1 generation: Half of the plants in BC1 generation are resistant and the
remaining half would be susceptible to stem rust. Rust resistant plants are selected and
back crossed to variety A.
4. BC2 to BC 5 generation: Segregation would occur for rust resistance. Rust resistant
plants are selected and back crossed to variety A.
5. BC6 generation: BC6 plants will have 99 percent genes from variety A Rust resistant
plants are selected and selfed, their seeds are harvested separately
6. BC6 F2 generation: Individual plants progeny from the selfed seeds of the selected plants
are grown. Rust resistant plant similar to the plant type of variety A are selected and they
are selfed. Seeds are harvested separately.
7. BC6 F3 generation: Individual plants progeny are grown. Progenies homozygous for rust
resistant and similar to plant type of variety Aare harvested in bulk. Several similar
progenies are usually mixed to constitute the new variety.
Merits:
1. Back cross method retains all desirable character of a popular adapted varieties and
replaces undesirable allele at particular locus
2. Useful for the transfer of disease resistance and incorporation of quality traits into a
variety
3. This is used for the development of isogenic lines,
4. Extensive tests are not required 2-3 generation can be raised in off season nurseries
greenhouses; it would save time.
5. This is the only method for the inter specific gene transfer and transfer of cytoplasm.
6. Male sterility and fertility restoration genes can be transferred to various back ground.

Demerits:
1. New variety cannot be superior to recurrent parent except for the character transferred
2. It involves lot of crossing work. 6-8 back cross is often difficult and time consuming.
3. Sometime undesirable gene linked with desirable also may be transferred.
4. By the time the back cross programme the recurrent parent may have been replaced by
other varieties superior in yield and other character.
Multiline Varieties
Generally, pureline varieties are highly adapted to a limited area, but poorly adapted to
wider regions. Further, their performance is not stable from year to year because of
changes in weather and other environmental factors. Purelines often have only one or a few
major genes for disease resistance, such as, rust resistance, which make them resistant to
some races of the pathogen. New races are continuously produced in many pathogens,
which may overcome the resistance present in the pureline varieties. For example, Kalyan
Sona wheat (T. aestivum) originally resistant to brown rust (leaf rust), soon became
susceptible to new races of the pathogen. To overcome these limitations, particularly the
breakdown of resistance to disease, it was suggested to develop multiline varieties.
Multiline varieties are mixtures of several purelines of similar height, flowering and
maturity dates, seed colour and agronomic characteristics, but having different genes for
disease resistance. The purelines constituting a multiline variety must be compatible, i.e.,
they should not reduce the yielding ability of each other when grown in mixture.
In 1954, Borlaug suggested that several purelines with different resistance genes should be
developed through back cross programmes using one recurrent parent. This is done by
transferring disease resistance genes from several donor parents carrying different resistant
genes to a single recurrent parent. Each donor parent is used in a separate backcross
programme so that each line has different resistant gene or genes. Five to ten of these lines
may be mixed depending upon the races of the pathogen prevalent in the area. If a line or
lines become susceptible, they would be replaced by resistant lines. New lines would be
developed when new sources of resistance become available. The breeder should keep
several resistant lines in store for future use in the replacement of susceptible lines of
multiline verities.
Merits of Multiline varieties
1. All the lines are almost identical to the recurrent parent in agronomic characteristics,
quality etc. Therefore, the disadvantages of the pureline mixtures are not present in the
multiline varieties.
2. Only one or a few lines of the mixture would become susceptible of the pathogen in
anyone season. Therefore, the loss to the cultivator would be relatively low.
3. The susceptible line would constitute only a small proportion of the plants in the field.
Therefore, only a small proportion of the plants would be infected by the pathogen.
Consequently, the disease would spread more slowly than when the entire population was
susceptible. This would reduce the damage to the susceptible line as well.
 Demerits of Multiline Varieties
1. The fanner has to change the seed of multiline varieties every few years depending upon
the change in the races of the pathogen.
2. There is a possibility that a new race may attack all lines of a multiline variety.
Achievements
Multiline variety appears to be a useful approach to control diseases like rusts where new
races are continuously produced. In India, three multiline varieties have been released in
wheat (T. aestivum). Kalyan Sona, one of the most popular varieties in the late sixties, was
used as the recurrent parent to produce these varieties. Variety 'KSML 3' consists of 8 lines
having rust resistance genes from Robin, Ghanate, Kl, Rend, Gabato, Blue Brid, Tobari
etc. Multiline 'MLKS 11' is also a mixture of 8 lines; the resistance is derived from E 6254,
E 6056, E 5868, Frecor, HS 19, E 4894 etc. The third variety, KML 7406 has 9 lines
deriving rust resistance from different sources.
Dirty Multiline
This term is used when a multiline is having one or two susceptible lines also. The idea of
including susceptible lines is to prevent race formation.
Hardy Weinberg Law
 Cross pollinated crops are highly heterozygous due to the free inter mating among them

so these are random mating populations. Because each individual of the population has
equal opportunity of mating with any other individual. It is also known as
mendelian/panmictic population. A Mendelian population may be thought of having a
gene pool consisting of all gametes produced by the population. So, gene pool may be
defined as the sum total of all genes present in the population. A population consists of
all such individuals that share the same gene pool has an opportunity to inter mate with
each other and contribute to the next generation of the population.
 Each generation of a Mendelian population may be considered to arise from a random

sample of gametes from the gene pool of previous generation. Hence, it is not easy to
follow the inheritance of a gene in a Mendelian population. It cannot be estimated by
using the techniques of classical genetics. So, to understand the genetic makeup of such
population a population genetics has been developed.
 This law is independently developed by Hardy (1908) in England and Weinberg (1909) in

Germany. The law states that the gene and genotype frequencies in a Mendelian population
remain constant from generation after generation if there is
no selection, mutation, migration or random drift.
 2
 The frequencies of these genotypes for a locus with two alleles A and a would be P (AA),

2pq (Aa)and q2(aa)

Where, p= Frequency of „A‟ allele in the population.
 q= Corresponding frequency „a‟ allele in the population the sum of p+q is equal=1

Such a population would be at equilibrium since the genotypic frequencies would be

stable, that is, would not change from one generation to the next. This equilibrium is
known as Hardy Weinberg equilibrium.
Migration: Migration is the movement of individual into a population from a different
population. Migration may introduce new alleles into the population or may change the
frequency of existing allele. The amount of change in gene frequency „q‟ will primarily
depend upon two factors.
a. Ratio of migrant individuals to those of the original population.
b. The Magnitude of difference between the values of q in the population and in the
migrants.
So, in plant breeding migration is by inter varietal crosses or poly crosses.
Mutation: mutation is a sudden heritable change in an organism and is generally due to a
structural change in a gene. It may produce a new allele not present in the population or
may change the frequency of existence allele that 10 -6 mutation is detected. So, such
effects in breeding population may be ignored.
Random drift or genetic drift: It is a random change in gene frequency due to sampling
error. Random drift is more in small population than larger. Ultimately, the frequency of one
of the alleles becomes zero and that of the other allele becomes one. The allele with the
frequency one is fixed in the population because there would be no change in the frequency.
So, all genes become homozygous. The genetic drift can be reduced by handling large
population.
Inbreeding: Mating between individuals sharing a common parent in their
ancestry inbreeding reduces the proportion of the heterozygosity and increase the frequency
of homozygosity by the rate of decrease in heterozygosity is equal to ½ N (N- Number of
plants in the population) per generation in monoecious or hermaphrodite species. In dioecious
species and monoecious species where self-pollination is prevented the decrease in
heterozygosity is low. In small population, even with strict random mating / strict cross
pollination the frequency of homozygotes increases while that of heterozygotes decreases due
to inbreeding
Selection: The selection in a random mating population is highly effective in increasing or
decreasing the frequency of allele, but it is unable to either fix or eliminate them. However, in
combination with a system of inbreeding, selection is highly efficient in the fixation and
elimination of an allele.
Inbreeding
It is mating between individuals related by descent or having common ancestry. The
highest degree of inbreeding is obtained by selfing. Inbreeding depression refers to
decrease in fitness and vigour due to inbreeding. The degree of inbreeding is measured by
the inbreeding coefficient.
History of inbreeding:
In breeding depression has been recognised by man for a long time. Knowing the
consequences of inbreeding many societies have prohibited marriages between closely
related individuals. Darwin in 1876 published a book “cross and self-fertilization in
vegetable kingdom” in which he concluded that progenies obtained from self-fertilization
were weaker in maize. Detailed and precise information on inbreeding in maize was
published by East in 1908 and Shull in 1909.
The different effects of inbreeding are:
1. Appearance of Lethal and Sublethal Alleles: IB results in appearance of lethal;
sublethal and sub vital characters. e.g.: Chlorophyll deficiencies, rootless seedlings, flower
deformities – They do not survive, they lost in population.
2. Reduction in vigour: General reduction in vigour size of various plant parts.
3. Reduction in Reproductive ability: Reproductive ability of population decreases
rapidly. Many lines reproduce purely that they cannot be maintained.
4. Separation of the population into distinct lines: Population rapidly separates into
distinct lines i.e. due to increase in homozygosity. This leads to random fixation of alleles
in different lines. Therefore, lines differ in genotype and phenotype. It leads to increase in
the variance of the population.
5. Increase in homozygosity: Each line becomes homozygous. Therefore, variation within
a line decreases rapidly. After 7-8 generations of selfing the line becomes more than 99%
homozygous. These are the inbreds which have to be maintained by selfing.
6. Reduction in yield: Inbreeding leads to loss in yield. The inbreds that survive and
maintained have much less yield than the open pollinated variety from which they have
been developed.
Degrees of inbreeding depression
 Various plant species exhibit different degrees of inbreeding depression. The depression
may be from very high to nil. Based on degree of depression, the plant species can be
grouped into 4 broad categories.
1. High inbreeding depression: E.g. Lucerne, Carrot. Inbreeding leads to severe
depression and exhibit lethal effects. After 3 or 4 generations of selfing it is hard to
maintain lines.
2. Moderate inbreeding depression: E.g. Maize, Jowar, Bajra. Though lethal effects are
there, lines can be separated and maintained.
3. Low inbreeding depression: E.g. Cucurbits, Sunflower. Only a small degree
of inbreeding depression is observed.
4. No inbreeding depression: The self-pollinated crops do not
show inbreeding depression.
Heterosis
The term heterosis was first used by Shull in 1914. Heterosis may be defined as the
superiority of an F1 hybrid over both of its parents in terms of yield or some other
character. Generally, heterosis is manifested as an increase in vigour, size, growth rate,
yield or some other characteristic.
History
Koelreuter is first reported hybrid vigour in tobacco produced artificial hybrids. In 1876,
Darwin concluded that hybrids form unrelated plant type were highly vigorous. Most of
our present knowledge on heterosis comes from the work on maize. Crossing inbred lines
rather than open pollinated varieties produces the commercial maize hybrids.
Hybridization between inbreds developed from the same variety or from closely related
varieties produced only a small degree of heterosis.
 But a vast majority of the cases of heterosis are cases of superiority of hybrids over their
parents. Hybrid vigour describes only superiority of hybrids over the parents. The few
cases where F1 hybrids are inferior to their parents may also be regarded as cases of hybrid
vigour in the negative direction.
Heterosis is the superiority of a hybrid over its parents.
1. Increased yield: Heterosis is generally expressed as an increase in the yield of hybrids.
The yield may be measured in terms of grain, fruit, seed, leaf, tubers or the whole plant.
2. Increased reproductive ability: More number of flowers/fruits/seeds. Increase in Size
and General Vigour: The hybrids are generally more vigorous, i.e., healthier and faster
growing and larger in size than their parents.
3. Better quality: In many cases, hybrids show improved quality. For example, many
hybrids in onion show better keeping quality, but not yield, than open-pollinated
varieties.
4. Earlier flowering and maturity: In many cases hybrids are earlier in flowering and
maturity than the parents. But earliness is highly desirable in many situations
particularly in vegetables.
5. Greater resistance to disease and pest: Some hybrids are known to exhibit a greater
resistance to insect or diseases than their parents.
6. Greater adaptability: Hybrids are generally more adapted to environmental changes than
inbreds.
7. Faster growth rate: In some cases, hybrids show a faster growth rate than their parents.
But the total plant size of the hybrids may be comparable to that of parents. In such
cases, a faster growth rate is not associated with a larger size.
8. Increase in the number of a plant part: In some cases there is an increase in the number
of nodes, leaves and other plant parts, but the total plant size may not be larger.
Estimation of heterosis
1. Average heterosis:
It is the heterosis where F1 is superior to mid parent value. In other words, superior to
average of two parents.
F1 - MP
--------- x 100
MP
Where F1 = Mean of hybrid
MP = Mid parental value.
(P1 + P2)
MP = ----------
2
where P1 = Parent 1; P2 = Parent 2
This type of heterosis is of no use in agriculture since the superiority is below the better
parent value
2. Heterobeltiosis:
Superiority of F1 over the better parent.
F1 - BP
--------- x 100
BP
Where BP = Mean of Better Parent.
3. Economic heterosis:
Superiority of the F1 compared to the high yielding commercial variety in a particular crop.
F1 - CV
--------- x 100
CV
Where CV = Mean of Commercial Variety.
4. Negative heterosis:
Performance of F1 inferior to better parent / mid parent value. - e.g. Duration.

Breeding Methods for Cross Pollinated Crops

Populations of cross-pollinated crops are highly heterozygous. When inbreeding is practiced,
they show severe inbreeding depression. So, to avoid inbreeding depression and its
undesirable effects, the breeding methods in the crop is designed in such a way that there will
be a minimum inbreeding. The breeding methods commonly used in cross pollinated crops
may be broadly grouped into two categories.
I. Population improvement
2. Hybrids
3. Synthetics and Composites
Population Improvement
In cross pollinated crops, population improvement is used to enhance the frequency of
desirable alleles in a population.
Population improvement without progeny test: Mass selection,
Population improvement with progeny testing includes Recurrent selection
Mass selection
This is similar to the one, which is practiced, in self-pollinated crops. A number of plants are
selected based on their phenotype and open pollinated seed from them are bulked together to
raise the next generation. The selection cycle is repeated one or more times to increase the
frequency of favourable alleles. Such a selection is known as phenotypic recurrent selection.
Merits
i) Simple and less time consuming
ii) Highly effective for character that are easily heritable. Eg. Plant height, duration.
iii) It will have high adaptability because the base population is locally adapted one.
Demerits
1. Selection is based on phenotype only which is influenced by environment
2. The selected plants are pollinated both by superior and inferior pollens present in the
population.
3. High intensity of selection may lead reduction in population there by leading to inbreeding.
Recurrent selection
This is one of the breeding methods followed for the improvement of cross-pollinated crop.
Here single plants are selected based on their phenotype or by progeny testing. The selected
single plants are selfed. In the next generation they are intermated (cross in all possible
combinations) to produce population for next cycle of selection. The recurrent selection
schemes are modified forms of progeny selection programmes. The main difference between
progeny selection and recurrent selection:
i) The manner in which progenies are obtained for evaluation.
ii) Instead of open pollination, making all possible inter crosses among the selected lines.
The recurrent selection schemes are of 4 different types:
i. Simple Recurrent Selection
ii. Recurrent Selection for GCA
iii. Recurrent Selection for SCA
iv. Reciprocal Recurrent Selection
1. Simple recurrent selection
In this method a number of desirable plants are selected and self-pollinated. Separate progeny
rows are grown from the selected plants in next generation. The progenies are intercrossed in
all possible combination by hand. Equal amount of seed from each cross is mixed to raise
next generation. This completes original selection cycle. From this, several desirable plants
are selected and self-pollinated. Progeny rows are grown and inter crosses made. Equal
amount of seeds are composited to raise next generation. This forms the first recurrent
selection cycle.
First Year:
i) Several superior plants are selected.
ii) Selected plants selfed.
iii) Harvest the single plants.
iv) Seeds are evaluated, superior plants are identified.
Second Year:
i) Progeny rows raised
ii) Inter crosses are made in all combination by hand.
iii) Equal amount of seed bulked from each cross.
Third Year:
i) Composited seeds raised
ii)Repeat the operation as in first year
Fourth Year:
Repeat as in second year.
i) Recurrent selection is effective in increasing the frequency of desirable genes in the
population
ii) Most suited for characters having high heritability.
iii) Inbreeding is kept at minimum.
Eg: oil content, protein content and high heritability traits are effective for increasing the
frequency of desirable genes in the selected populations

Merits of recurrent selection

i) Recurrent selection is effective in increasing the frequency of desirable genes in the
population ii) Most suited for characters having high heritability
iii) Inbreeding is kept at minimum.
2. Hybrids
They are the first generation from crosses between two pure lines, inbreds, open pollinated
varieties of other populations that are genetically not similar. Pure line hybrids: Tomato.
Inbred hybrids: Maize, bajra.
Kinds of hybrids
1. Single cross hybrids (A x B)
Crossing two inbreeds or pure lines.
2. Three-way cross hybrid (A x B) x C
A cross between a single cross hybrid and an inbred.
3. Double cross hybrid (A x B) x (C x D)
cross between two Fls.
4. Double Top Cross hybrid
Double cross hybrid crossed with open pollinated variety.
Operation in production of hybrids:
In production of hybrids inbreds are preferred rather than open pollinated varieties for the
following reasons:
1. Inbreds can be maintained without a change in the genotype. Whereas open pollinated
variety cannot be maintained pure. They may alter genotypically due to natural selection etc.
2. The hybrids derived from inbreds will be uniform where as it may not be in case, of open
pollinated variety.
3. The inbreds are homogenous and their performance can be predicted whereas open
pollinated variety are heterogenous and their prediction in performance cannot be made.
Development of inbreds
1. By inbreeding, selfing etc.
2. Development of inbreds from haploids - rice, sorghum, maize.
Evaluation of inbreds
a) Phenotypic evaluation
Based on phenotypic performance. Highly suitable for characters with high heritability.
b) Top cross test
Top cross test provides a reliable estimate of GCA. The selected inbreds will be crossed to a
tester parent with wide genetic base i.e. open pollinated variety. The cross progenies will be
evaluated in replicated progeny rows. Based on results better inbreds can be selected.
c) Single cross evaluation
The developed inbreds can be crossed and the single crosses can be estimated in replicated
trial. Outstanding hybrids tested over years in different locations, then released.
d) Prediction of double cross performance
'The predicted performance of any double cross is the average performance of the four non
parental single crosses involving the four parental inbreds. Inbreds: A, B, C, D.
6 possible single crosses = A x B, A x C, A x D, B x C, B x D, C x D.
From these 3 double crosses produced = (A x B) x (C x D), A x C) x (B x D), (A x D) x (B x
C)
The performance of these anyone double cross can be predicted from performance of the four
single crosses not involved in producing that particular hybrid.
(A x B) x (C x D) = [(A x C) + (A x D) + (B x C) + (B x D)]/4
Production of Hybrids
Methods
I. Hand emasculation and dusting - Cotton, Tomato, Chillies, Bhendi
2. Use of male sterile lines
a) Cytoplasmic male sterility – ornamentals
b) Genic male sterility - Redgram, Castor.
c) Cytoplasmic - genic male sterility Jowar, Bajra, Rice
3. Use of self in compatibility
By planting cross compatible lines hybrids are produced. Here both are hybrids. E.g.
Brassicas.
Success of hybrids
a) Easy hand emasculation
b) Abundant seed set to compensate cost of hand emasculation.
c) Stable male sterile lines.
d) Effective restorers.
e) Effective pollen dispersal.
3. Synthetic Varieties
A synthetic variety is produced by crossing in all combinations a number of inbreds (4 -6) that
combine well with each other. The inbreds are tested for their GCA. Once synthesised, a
synthetic is maintained by open pollination. The lines that make up a synthetic may be
usually inbred line but open pollinated variety, or other population tested for general
combining ability are also be used.
Synthetic varieties are common in grasses, clover, maize and sugar beets. The normal
procedure is equal amounts of seeds from parental lines (Syn0) is mixed and planted in
isolation. Open pollination is allowed. The progeny obtained is Syn 1. This is distributed as
synthetic variety or it may be grown in isolation for one more season and Syn 2 is distributed.
Merits
1. Less costly compared to hybrids.
2. Farmer can maintain his synthetic variety for more seasons which is not possible in
hybrids.
3. Because of wider genetic base the synthetics are more stable over years and environments.
4. Seed production is more skilled operation in hybrids where as it is not so in synthetics.
Demerits
1. Performance is little bit lower compared to hybrids because synthetics exploit only
GCA while hybrids exploit both GCA and SCA.
2. The performance may not be good when lines having low GCA are used.
4. Composite varieties
It is produced by mixing seeds of phenotypically outstanding lines and encouraging open
pollination to produce crosses in all possible combinations among mixed lines. The lines used
to produce a composite are rarely tested for combining' ability. So, the yield of composite
varieties cannot be predicted easily. Like synthetics, composites are commercial varieties and
are maintained by open pollination.

Synthetic variety Composite variety

Parental components are generally inbreds It is not so in composite. The lines are not
tested for their GCA tested for their GCA.
No of parental lines are limited to 4 - 6 No such limit
inbreds
Synthetic produced with inbreds can be It is not possible to reconstitute composite
reconstituted variety
Yield performance can be predicted Yield performance cannot be predicted

Combining ability
Ability of a strain to produce superior progeny when crossed with other strains.
General combining ability (GCA)
Average performance of a strain in a series of cross combinations. The GCA is estimated
from the performance of F1 S from the crosses. The tester will have a broad genetic base.
Specific combining ability (SCA)
Deviation in performance of a cross combination from that predicted on the basis of general
combining ability of the parents involved in the cross. The testing will be on inbred.
Breeding methods for vegetatively propagated crops
Some agricultural crops and a large number of horticultural crops are asexually propagated.
Some common asexually propagated crops are sugarcane (S. officinarum), potato (S.
tuberosum), sweet potato (I. batatas), Colocasia (Taro), Arum, Dioscorea (yams), Mentha,
ginger (Zingiber sp.), turmeric (C. domestica), banana (Musa paradisiaca), etc., almost all
the fruit trees, e.g., mango (Mangifera indica), citrus (Citrus spp.), apples (P. malus), pears
(P.communis), peaches (P. persica), litchi (Litchi chinensis), loquat (Eriobotrya japonica),
etc:, and many ornamentals and grasses. Many of these crops show reduced flowering and
seed set, 'e.g., sugarcane, potato, sweet potato, banana, etc., and some varieties of these crops
do not flower at all. But many of these crops flower regularly and show satisfactory seed set.
However, they are propagated asexually to avoid the ill effects of segregation and
recombination, both being the inevitable consequences of sexual reproduction.
Segregation and recombination produce new gene combinations due to which the progeny
differ from their parents in genotype and phenotype. Asexual reproduction, on the other hand,
produces progeny exactly identical to their parents in genotype because the progeny is
derived from vegetative cells through mitosis.
The advantage of asexual reproduction is immediately clear. It preserves the genotype of an
individual indefinitely. It must be noted that this does not depend on the homozygosity of the
genotype of an individual. Any genotype is preserved and. maintained through asexual
reproduction. In contrast self-pollination preserves and maintains only homozygous
genotypes giving rise to purelines.
Characteristics of Asexually Propagated Crops
, perennial, e.g., sugarcane, fruit trees, etc. The annual crops
are mostly tuber crops, e.g., potato, cassava (M. utilissima), sweet potato, etc.
-and seed set. Many varieties do not flower at all.
Only the crops grown for fruit, particularly where good fruit set depends upon seed
formation, show regular flowering and satisfactory seed set.
-pollinated.

t majority of asexually propagated crops are either polyploids, eg., sugarcane, potato,
sweet potato, etc., or have polyploid species or varieties.
M. paradisiaca), sugarcane, Rubus,
s consist of a large number of clones, that is, progeny derived from a single
plant through asexual reproduction. Thus, each variety of an asexually propagated crop is a
clone.
Clone
A clone is group of plants produced from a single through asexual reproduction. Thus,
asexually propagated crops consist of large number of clones, and they are often known as
clonal crops. All the members of a clone have the same genotype as the parent plant. As a
result, they are identical with each other in genotype. Consequently, the phenotypic
differences within a clone do not have a genetic basis and are purely due to the environmental
effects. A selection within a clone is thus useless. The various characteristics of a clone are
summarised below.
Identical Genotype
All the individuals belongings to a single clone are identical in genotype. This is so because a
clone is obtained through asexual reproduction, which involves mitotic cell division only.
Genetic variation in the progeny of a plant is produced chiefly by segregation and
recombination, which occur during meiosis only. Thus, the genotype of a clone is maintained
indefinitely without any change.
Lack of genetic variation
The phenotypic variation present within a clone is due to the environment only. This is so
because all the individuals belonging to a single clone have the same genotype. The
phenotype of a clone is due to the effects of genotype (G), the environment (E) and the
genotype X 'environment interaction (G x E) the population mean (µ). Thus, the phenotype
(P) of a clone may be expressed as follows:
P = µ + G + E + GE
Thus, the phenotypic differences among clones would be partly due to E and GE components.
Hence the efficiency of selection among clones, as among purelines, would depend upon the
precision with which the E and GE components of phenotype are estimated.
Immortality
Theoretically, clones are immortal i.e., a clone can be maintained indefinitely through asexua l
reproduction. But clones usually degenerate due to viral or bacterial infection. A clone may
become extinct due to its susceptibility to diseases or insect pests. Further, genetic variation
may arise within a clone changing its characteristics.
Severe Inbreeding Depression
Generally, clones are highly heterozygous and show severe loss in vigor due to inbreeding.
Clonal Selection
The phenotypic value of a plant or clone is due to the effects of its genotype (G), the
environment (E) and genotype x environment (G x E) interaction. Of these, only the G effects
are heritable. The environmental and interaction effects are non-heritable and cannot be
selected for. Therefore, a selection for quantitative characters based on observations on single
plants is highly unreliable. In fact, plants selected in this way may be no better than a random
sample.
Further, a selection for characters like yielding ability, etc. on the basis of unreplicated clonal
plots would often be misleading and unreliable. Therefore, the value of a clone can be
reliably estimated only through replicated yield trials. However, selection for highly heritable
characteristics, such as plant height, days to flowering, color, disease resistance, etc., are easy
and effective even on the basis of individual plants or single plots. Clearly, these situations
are the same as those in the case of sexually reproducing crops.
Selection Procedure
In view of these considerations, in the earlier stages of clonal selection, when selection is
based on single plants or single plots, the emphasis is on the elimination of weak and
undesirable plants or clones. The breeder cannot reasonably hope to identify superior'
genotypes at this stage. In the later stages, when replicated trials are the basis of selection, the
emphasis is to identify and select the superior clones. The various steps involved in clonal
selection are briefly described below and are depicted in Fig:
1. First Year
From a mixed variable population, few hundreds to few thousand desirable plants are
selected. A rigid selection can be done for simply inherited characters with high heritability.
Plants with obvious weaknesses are eliminated.
2. Second Year
Clones from the selected plants are grown separately, generally, without replication This is
done in view of the limited supply of the propagating material for each clone, and because of
the large number of clones involved. The characteristics of clones will be clearer now than in
the previous generation when the observations were based on individual plants. The number
of clones is drastically reduced and inferior clones eliminated. The selection is based on
visual observations and on the basis of clonal characteristics. Fifty to one hundred clones may
be selected on the basis of clonal characteristics.
3. Third Year
Replicated preliminary yield trial is conducted. Suitable, checks included for comparison.
Few superior performing clones with desirable characteristics selected for multilocation
trials. At this stage, selection for quality is also done. If necessary, separate disease nurseries
may be planted to evaluate the disease resistance of selected clones.
4. Fourth to Sixth Years
Replicated yield trials are conducted at several locations along with a suitable check. The
yielding ability, quality and disease resistance, etc. of the clones are rigidly evaluated. The
best clone that is superior to the check in one or more characteristics is identified for release
as a new variety.
5. Seventh Year
The superior clone is multiplied released as a new variety.
Merits of Clonal Selection

depression, and preserves the gene combinations present in the clones.

to generate the variability necessary for selection.

lones.
Demerits of Clonal Selection

has not been devised to generate variability.

hrough hybridization
Clonal Hybridization
Clonal crops are generally improved by crossing two or more desirable clones, followed by
selection in the F 1 progeny and in the subsequent clonal generations. Once the F 1 has been
produced, the breeding procedure is essentially the same as clonal selection. The
improvement through hybridization involves the following three steps:
1. Selection of parents,
2. Production of F1 progeny, and
3. Selection of superior cones.
Hybridization can be used only in such crops, which can reproduce sexually. In case of those
crops where sexual reproduction is lacking, mutagenesis or biotechnological approaches can
be applied.
Selection of Parents
Selection of the parents to be used in hybridization is very important since the value of F 1
progeny would depend upon the parents used for producing the F 1. Parents are generally
selected on the basis of their known performance both as varieties and as parents in
hybridization programmes. The performance of a strain in hybridization programmes depends
on its prepotency and general combining ability. It would be highly desirable to know the
relative values of CGA and SCA in the crop to be improved. If GCA is more important, a
small number of parents with good should be used in hybridization programmes. On the other
hand, when SCA is more important, a large number of parents should be used to produce a
large number of F1 families in an effort to find some outstanding crosses.
A recent suggestion is to partially inbreed the parents to be used in hybridization
programmes. Clonal crops show severe inbreeding depression, but it is expected that one
generation of selfing or 2-3 generations of sib-mating may not reduce vigour and fertility too
severely. Inbreeding may enable the breeder to identify plants that would have a greater
concentration of desirable alleles. These plants may be more prepotent as parents than the
highly heterozygous clones. The practice is gaining some favour with plant breeders.
Production of F1 progeny
Generally, clonal crops are cross-pollinated and they may show self-incompatibility. The
selected parents may be used to produce single crosses involving two parents or an equivalent
of a polycross involving more than two parents.
Selection among F1 Families
When the breeding value of parents is not known, and the relative contributions of GCA and
SCA is not available, a large number of crosses have to be made in order to ensure that at
least some of the crosses would produce outstanding progeny in F 1. This is particularly true in
a species where crop improvement has not been done or has been done at a small scale. In
such cases, it would be cumbersome to evaluate a large number of F 1 progeny in detail. To
avoid this, generally small samples of several F1 populations are grown. The general worth of
individual F1 populations is estimated visually. The presence of outstanding individuals in the
F1, populations is also noted, and inferior F1‟s are eliminated. Promising F1‟s with
outstanding individuals are then grown at a much larger scale for selection. The procedure is
designed to save time, space and labour by planting only small populations of a large number
of crosses at the preliminary stage.
Selection within F1 Families
The selection procedure within F1 populations is essentially the same as that in the case of
clonal selection. The various steps involved in the breeding of clonal crops through
hybridization are briefly described below. From second year onward, these should be read
along with the steps described in clonal selection.
First Year
Clones to be used as parents are grown and crosses are made to produce F 1 progeny.
Second Year
Sexual progeny from the cross, i.e., seedlings obtained from seeds, are grown. Undesirable
plants are eliminated. Few hundred to few thousand desirable plants are selected.
Third Year
Clones from the selected individual plants are grown separately. Poor and inferior clones are
eliminated. Up to 200 superior clones may be selected for preliminary yield trial.
Fourth Year
A replicated preliminary yield trial is conducted in which suitable checks are included for
comparison. Few outstanding clones are selected for trials at several locations.
Fifth to seventh year
Replicated yield trials are conducted at several locations. Suitable checks are included for
comparison. One or a few outstanding clones are identified and released as new varieties.
Eighth year
The clones released as varieties are multiplied and distributed among farmers.
MUTATION BREEDING
The term mutation was coined by Hugo De Vries in 1900 for the first time and the word is
derived from the latin word „MUTARE‟ means to change. Mutation is the sudden heritable
change other than the Mendelian segregation and gene recombination in an organism.
Mutation may be the result of a change in a gene, a change in chromosome that involves
several genes or a change in plasma gene. Mutations produced by changes in the base
sequence of genes are known as gene or point mutations some mutations may be produced by
changes in chromosome structure or even in chromosome number they are termed as
chromosomal mutation. There are three types of mutations based on genetic basis of heritable
change:
1. Gene mutations: These are produced by change in the base sequence of genes. The change
may be due to base substitutions, deletion or addition.
2. Chromosomal mutation: These arise due to change in chromosome number that may leads
to polyploidy or aneuploidy or change in chromosome structure that result in deletions
duplication, inversion and translocation.
3. Cytoplasmic or plasma gene mutation: These are due to change in the base sequence of
plasma genes. The plasma genes are present in mitochondria or chloroplast. Here the mutant
character occurs in buds or somatic tissues which are used for propagation in clonal crops.
Classification of mutations:
A. Based on origin, the mutations are classified as spontaneous and induced mutations.
1. Spontaneous mutations: Mutations occur in natural populations at a low rate (10-6) but
different genes may show different mutation rates. Here the different genes show different
mutation rate. For example: in maize R-locus mutates at the frequency of 4.92 x 10-4 i.e. (1
in 20000 population), when as Su locus at 2.4 x 10 -6 (1 in 25 lakhs). The Wx locus
considered to be highly stable. The difference in mutation rate may be due to
a) Genetic back ground i.e. presence of mutator genes
b) Genes themselves
c) Environment
2. Induced mutation: Mutations may be artificially induced by treatment with certain
physical or chemical agents. Available evidence indicates that induced mutation rarely
produces new alleles they produce alleles which are already known to occur spontaneously.
Induced mutations are comparable to spontaneous mutations in their effects and in the
variability they produce. Induced mutation occurs at a relatively higher frequency so that it is
practical to work with them.
B. Based on magnitude of phenotypic effects mutation as classified as
Macro mutations: Oligogenic mutation – Large phenotypic effect and recognizable on
individual plant basis and can be seen easily in M2 generations. e.g. Ancon breed in sheep,
pod maize to cob maize
Micro mutations: Polygenic mutations – Small phenotypic effect which cannot be
recognized on individual plant basis but can be recognize only in a group of plants. Selection
should be done in M3 or later generations.
Characteristic features of mutations
1. Mutations are generally recessive but dominant mutations also occur
2. Mutations are generally harmful to the organism. Most of the mutations have deleterious
effects but small proportion (0.1%) of them are beneficial.
3. Mutations are random i.e. they may occur in any gene. However, some genes show high
mutation rates than the others.
4. Mutations are recurrent
5. Induced mutations commonly show pleiotropy often due to mutation in closely linked
genes. Procedure for irradiation: The plant material may be treated in any of the following
source. 1. Seeds, 2. Seedlings, 3. Flowers, 4. Cuttings
1. Seeds: Seeds are used after soaking to get greater frequency of induced mutations than air
dried.
2. Seedlings: At any stage of life cycle can be subjected to radiation but usually seedlings
neither too young nor too old are irradiated due to their convenience in handling in pots
transportation from nursery easily.
3. Flowers: Meiotic cells have been found more sensitive than the mitotic cells and therefore
plants are irradiated in the flowering stage in order to affect the developing gametes.
4. Cuttings: In case of fruit tree when they are propagated by clones – the desirable cuttings
are exposed to irradiation.
Selection of the variety for mutagen treatment
The variety selected for mutagenesis should be the best available in the crop.
Dose of the Mutagen
An optimum dose of the mutagen should be used. An optimum dose is the one which
produces the maximum frequency of mutations and causes the minimum killing. Many
workers feel that a dose close to LD 50 should be optimum. LD 50 is that dose of a mutagen,
which would kill 50% of the treated individuals.
Mutation Breeding for oligogenic traits
The handling procedure described here is based on the selection for a recessive mutant allele
of an oligogene.
1. M1 generation: Several hundred seeds are treated with a mutagen and are space planted. In
general, the number of treated seeds is so adjusted as to give rise to 500 fertile M1 plants at
the harvest. Care should be taken to avoid outcrossing; this can be achieved either by
planting the M1 population in isolation or by bagging the inflorescence of M1 plants or even
the whole M1 plants. M1 plants will be chimeras for the mutations present in heterozygous
state. About 20 to 25 seeds from each M1 spike are harvested separately to raise the M 2
progeny rows.
2. M2 generation: About 2,000 progeny rows are grown. Careful and regular observations are
made on the M2 rows. But only distinct mutations are detected in M 2 because the
observations are based on single plants. All the plants in M2 rows suspected of containing
new mutations are harvested separately to raise individual plant progenies in M 3. if the
mutant is distinct, it is selected for multiplication and testing. However, most of the
mutations will be useless for crop improvement. Only 1-3 per cent of M2 rows may be
expected to have beneficial mutations.
3. M3 generation: Progeny rows from individual selected plants are grown in M3. Poor and
inferior mutant rows are eliminated. If the mutant progenies are homogeneous, two or more
M3 progenies containing the same mutation may be bulked. Mutant M 3 rows are harvested in
bulk for a preliminary yield trial in M4.
4. M4 generation: A preliminary yield trial is conducted with a suitable check, and promising
mutant lines are selected for replicated multilocation trials.
5. M5 -M7 generations: Replicated multilocation yield trials are conducted. The out-standing
line may be released as a new variety. The low yielding mutant lines, however, should be
retained for use in hybridization programmes.
Mutation breeding for polygenic traits: Mutagenesis does produce genetic variation in
polygenic traits; this variation is usually as much as 50% of that generated in F 2 generation,
but sometimes it may be as much as or even greater than the latter.
1. M1 and M2 generations: M1 and M2 generations are grown in the same way as in the case
of oligogenic traits. In M2 generation, vigorous, fertile and normal looking plants that do not
exhibit a mutant phenotype are selected and their seeds are harvested separately to raise
individual plant progeny rows in M3.
2. M3 generation: Progeny rows from individual selected plants are grown. Careful
observations are made on M3 rows for small deviations in phenotype from the parent variety.
Inferior rows are discarded. Few rows may be homogeneous and would be harvested in bulk.
Selection in done in M3 rows showing segregation; a majority of M3 rows would show
segregation. Intensive and careful evaluation of a large number of M 3 progeny rows allows
identification of mutants with altered quantitative traits, e.g., partial or horizontal disease
resistance. Such mutants occur in high frequencies that approach 1% or even high, so that
their isolation becomes quite cost effective.
3. M4 generation: Bulked seed from homogeneous M3 rows may be planted in a preliminary
yield trial with a suitable check; superior progenies are selected for replicated multilocation
yield trials. Individual plant progenies from M3 are critically observed. Progenies showing
segregation may be subjected to selection only if they are promising. Superior homogeneous
progenies are harvested in bulk for preliminary yield tests in M5 .
4. M5 -M8 generations: Preliminary yield trials and / or multi-location trials are conducted
depending upon the stage when the progenies become homogeneous. Outstanding progenies
may be released as new varieties.
Applications of Mutation Breeding
Mutation breeding has been used for improving both oligogenic as well as polygenic
characters. Mutagenesis has been used to improve morphological and physiological
characters including yielding ability. Various applications of mutation breeding are:
1. Induction of desirable mutant alleles which may not be available in the germplasm
2. It is useful in improving specific characteristics of a well-adapted high yielding variety.
3. Mutagenesis has been successfully used to improve various quantitative characters
including yield.
4. F1 hybrids from intervarietal crosses may be treated with mutagens in order to increase
genetic variability by inducing mutation and to facilitate recombination of linked genes.
5. Irradiation of interspecific (distant) hybrids has been done to produce translocations.
Advantages:
1. Mutation create inexhaustible variation.
2. When no improvement is possible this method has to be adopted.
Limitations:
1. Frequency of desirable mutations is very low about 0.1 percent. To detect the desirable
one in M2 considerable time, labour & other resources are to be employed.
2. To screen large population, efficient quick and inexpensive selection techniques are
needed. 3. Desirable mutations may be associated with undesirable side effects due to other
mutations thus extending the mutation breeding programme.
4. Detection of recessive mutations in polyploids and clones is difficult and larger doses of
mutagen have to be applied and larger populations are to be grown.
Polyploidy Breeding
The somatic chromosome number of any species, whether diploid or polyploidy, is
designated as 2n, and the chromosome number of gametes is denoted as n. An individual
carrying the gametic chromosome number, n, is known as haploid. A monoploid, on the
other hand, has the basic chromosome number, x. In a diploid species, n = x, one x
constitutes a genome or chromosome complement. The different chromosomes of a single
genome are distinct from each other in morphology and or gene content and homology;
members of a single genome do not show a tendency of pairing with each other. Thus, a
diploid species has two, a triploid has 3 and a tetraploid has 4 genomes and so on.
In euploids, the chromosome number is an exact multiple of the basic or genomic number.
Euploidy is more commonly known as polyploidy.
When all the genomes present in a polyploidy species are identical, it is known as
autopolyploid and the situation is termed as autopolyploidy.
In the case of allopolyploids, two or more distinct genomes are present.
Euploids may have 3(triploid), 4(tetraploid), 5 (pentaploid), or more genomes making up
their somatic chromosome number.
In case of autopolyploidy, they are known as autotriploid, autotertaploid, autopentaploid, and
so on, while in the case of allopolyploidy they are termed as allotriploid, allotetraploid,
allopentaploid, etc.
Amphidiploid is an allopolyploid that has two copies of each genome present in it and, as a
consequence, behaves as a diploid during meiosis.
A segmental allopolyploid contains two or more genomes, which are identical with each
other, except for some minor differences.
Autopolyploids
Origin and production of doubled chromosome numbers:
1. Spontaneous: chromosome doubling occurs occasionally in somatic tissues and unreduced
gametes are produced in low frequencies.
2. Production of adventitious buds: decapitation in some plants leads to callus development at
the cut ends of the stem. Such a callus has some polyploid cells and some of the shoot buds
regenerated from the callus may be polyploid. In Solanaceae 6-36% of adventitious buds are
tetraploids.
3. Treatment with physical agents: Heat or cold treatment, centrifugation, x -ray or gamma
ray irradiation may produce polyploids. Exposing the plants or ears of maize to a temperature
of 38-45oC at the time of the first division of zygote produce 2 -5 % tetraploid progenies.
4. Regeneration in vitro: polyploidy is a common feature of the cells cultured in-vitro.
5. Colchicine treatment: Colchicine treatment is the most effective and the most widely used
treatment for chromosome doubling.
Autopolyploidy
In autopolyploidy, triploidy, tetraploidy and higher levels of ploidy are included.
Morphological and cytological features of auto polyploids:
The general features are summarised below:
1. Polyploids have larger cell size than diploids. Guard cells of stomata are larger and the
number of stomata per unit area is less in polyploids than diploids.
2. Pollen grains of polyploids are generally larger than those of the corresponding diploids.
3. Polyploids are generally slower in growth and later in flowering.
4. Polyploids usually have larger and thicker leaves, and larger flowers and fruits which are
usually less in number than in diploids.
5. Polyploids generally show reduced fertility due to irregularities during meiosis and due to
genotypic imbalance leading to physiological disturbances.‟
6. In many cases autopolyploidy leads to increased vigour and vegetative growth.
7. Different species have different levels of optimum ploidy. For sugar beet the optimum
level is 3x, sweet potato 6x while for timothy grass it is between 8 -10x.
8. Autopolyploids generally have a lower dry matter content than diploids.
Application of autopolyploidy in crop improvement
Triploids
Triploids are produced by hybridization between tetraploid and diploid strains. They are
generally highly sterile, except in a few cases. This feature is useful in the production of
seedless watermelons. In certain species, they may be more vigorous than the normal
diploids, e.g., in sugar beets. These two examples are described in some detail. Seedless
watermelons are produced by crossing tetraploid (4x, used as female) and diploid (2x, used as
male) lines, since the reciprocal cross (2x x 4x) is not successful. The triploid plants do not
produce true seeds; almost all the seeds are small, white rudimentary structures like cucumber
(Cucumis sativus) seeds. But few normal size seeds may occur which are generally empty.
For good seed setting pollination is essential. For this purpose, diploid lines are planted in the
ratio 1 diploid: 5 triploid plants. There are several problems viz. genetic instability of 4x lines,
irregular fruit shape, a tendency towards hollowness of fruits, production of empty seeds and
the labour involved in trioploid seed production.
1. Triploid sugar beets: Among root crops triploid sugar beets apparently represent the
optimum level of polyploidy because 3n plants have longer roots than diploid and also yield
more sugar per unit area.
2. Tetraploid rye: The advantage of tetraploid over its diploid counterpart are large kernel
size, superior ability to emerge under adverse condition and higher protein content.
Tetraploid rye varieties have been released for cultivation. e.g. Double steel, Tetra petkus.
Limitations of autopolyploidy:
1. Larger size autopolyploid generally contain more water and produce less dry matter
content than diploids
2. High sterility with poor seed setting is observed
3. Due to complex segregation, progress through selection is slow
4. Monoploids and triploids cannot be maintained except through clonal propagation
5. The varieties cannot be produced at will
6. Effects of autopolyploidy cannot be predicted.
Allopolyploidy: Allopolyploids have genomes from two or more species production of
allopolyploids has attracted considerable attention; the aim almost always was creation of
new species. Some success has been evident from the emergence of triticale. Raphano
brassica and allopolyploids of forage grasses.
Morphological and cytological features of allopolyploids
1. Allopolyploids combine the morphological and physiological characteristics of the parent
species but it is very difficult to predict the precise combination of characters that would
appear in the new species.
2. Many allopolyploids are apomictic eg: Tulips, Solanum
3. The chromosome pairing in the new species depends upon the similarities between the
chromosomes of the parental species. Chromosomes with such similarities are known as
homoeologous chromosomes. After chromosome doubling, the allopolyploid would have
two homelegous chromosomes for each chromosome present in the F 1 hybrid, comparable to
the diploid species. Such allopolyploid is referred as amphidiploid or Allotetraploid.
4. Fertility of Allopolyploids can be improved by hybridization and selection.
Application of allopolyploidy in crop improvement:
1. Utilization as a Bridging species: Amphidiploids serve as a bridge in transfer of characters
from one species to a related species, generally from a wild species to cultivated species. A n
example of use of an amphidiploid as a bridging species in the use of synthetic N. digluta or
transfer of resistance to tobacco mosaic virus from N. glutinosa to N. tabacum. The F1 hybrid
from the cross N. tabacum x N. glutinosa is sterile. Chromosome doubling of the F1 hybrid
produces the synthetic allohexaploid N. digluta which is reasonably fertile. N. digluta is
backcrossed to the recipient species (N. tabacum) to produce a pentaploid having complete
somatic chromosome complement of N. tabacum and one genome of N. glutinosa. The
pentaploid is sufficiently fertile to be backcrossed to N. tabaccum. In the progeny N.
tabacum like plants resistant to tobacco mosaic are selected and cytologically analysed.
2. Creation of new crop species: Triticales, Raphanobrassica
3. Widening the genetic base of existing allopolyploids: The genetic base of some natural
allopolyploids may be narrow, and it may be useful to introduce variability in such cases by
producing the allopolyploids afresh from their parental species. B. napus is a case in point;
the genetic variability of this species is narrow and the only recourse available is to
synthesize new allopolyploid B. napus to widen its genetic base. This is being done by
crossing B. campestris (n=10, AA) with B. oleracea (n=9, CC), the parental diploid species,
to produce the amphidiploid B. napus (n=19, AACC). The two species, B. campestris and B.
oleracea, have to be crossed as autotetraploids; the cross is very difficult and embryo culture
has to be used; somatic hybridization is being used to get around these problems.
Limitations of Allopolyploidy
1. The effects of allopolyploidy cannot be predicted. The allopolyploids have some features
from both the parental species, but these features may be the undesirable ones, e.g.,
Raphanobrassica, or the desirable ones, e.g., Triticale.
2. Newly synthesized allopolyploids have many defects, e.g., low fertility, cytogenetic and
genetic instability, other undesirable features etc.
3. The synthetic allopolyploids have to be improved through extensive breeding at the
polyploidy level. This involves considerable time, labour and other resources.
4. Only a small proportion of allopolyploids are promising; a vast majority of them are
valueless for agricultural purposes.

Wide hybridization or Distant hybridization

Introduction
When individuals from two distinct species of the same genera are crossed, it is known as
inter specific hybridization. e.g. Inter specific hybridization: Oryza sativa x O. perennis
When individuals being crossed belong to two different genera, it is referred to as inter
generic hybridization. e.g. Wheat x rye.
Hybridization between individuals from different species belonging to the same genus or two
different genera, is termed as distant hybridization or wide hybridization, and such crosses
are known as distant crosses or wide crosses.
History
The first distant hybridization; hybrid between carnation (Dianthus caryophyllus) and sweet
willian (Dianthus barbatus) by Thomas Fairchild in 1717 and the hybrid is called as
Fairchilds‟ mule.
1. Most of the interspecific hybrids were of no agricultural value.
2. Many interspecific hybrids particularly in case ornamentals, served as commercial
varieties. 3. An interesting inter generic hybrid Raphano brassica was an amphidiploid cross
between radish and cabbage but it was useless.
The first inter generic hybrid with a great potential was TRITICALE
Objectives:
1. To transfer some desirable character from wild relatives that are not available in cultivated
varieties. e.g. i) Many disease resistance and, insect resistance genes
ii. Wide adaptability: (i.e. drought-resistance, cold tolerance etc.)
iii. Quality improvement (e.g. Cotton (fibre), Tobacco (leaf)
iv. Yield improvement (e.g. Oats, Tobacco, Maize, Sugarcane)
v. Other characters (e.g. CMS, Earliness, dwarfness morphological characters)
2. Exploitation of luxuriance (heterosis) in vegetatively propagated / ornamental crops.
Prolonged vegetative period, Prolonged blooming period
3. Creation of Novel genotypes: New species or F 1 hybrids hither to non – existent in nature.
Barriers to the production of distant hybrids
1. Failure of zygote formation / Cross incompatibility
2. Failure of zygote development / Hybrid inviability
3. Failure of F1 seedling development / Hybrid sterility
A variety of mechanisms may be responsible for each of these three difficulties:
1. Failure of zygote formation / cross incompatibility: Inability of the functional pollens of
one species or genera to effect fertilization of the female gametes of another species or genera
is referred to as cross incompatibility. It may be due to –
i. Failure of fertilization, because the pollen may not germinate.
ii. Pollen tube is unable to reach to embryo sac and hence sperms are not available for
fertilization
iii. Pollen tube may burst in the style of another species. e.g. Datura.
iv. The style of the female parent may be longer than the usual length of the pollen tube
growth therefore the pollen does not reach the embryo sac. e.g. Zea mays and Tripsacum sp.
v. Pollen tubes of polyploidy species are usually thicker than those of diploid species.
vi. When a diploid is used as female and a polyploidy as male, the polyploidy pollen tube
grows at a slower rate in the diploid style than it would be in a polyploid style.
These barriers are known as pre-fertilization barriers.
Techniques to make wide crosses successful
1. Removal or scarification of stigma
2. Using short styled parent as female.
3. Using the diploid species as the male parent.
2. Failure of zygote Development / Hybrid inviability
The inability of a hybrid zygote to grow into a normal embryo under the usual conditions of
development is referred to as hybrid inviability. This may be due to:
i.) Lethal genes: some species carry a lethal gene, which causes death of the interspecific
hybrid zygote during early embryonic development. For e.g. Aegilops umbellulata carries a
lethal gene with 3 alleles against diploid wheats.
ii.) Genetic disharmony between the two parental genomes: The genetic imbalance between
the two parental species may cause the death of embryos. For e.g. in cotton - G. gossypioides
x other G. spp., Brassica – B. napus x B. oleracea
iii.) Chromosome elimination: In some cases of distant hybridization, chromosomes are
gradually eliminated from the zygote. This generally does not prevent embryo development,
but the resulting embryo and the F1 plants obtained from such crosses are not true
interspecific hybrids since they do not have the two parental genomes in full. Generally,
chromosomes from one are successively eliminated due to mitotic irregularities. For e.g.
Hordeum bulbosum x H. vulgare, Hordeum bulbosum x Triticum aestivum, Triticum
aestivum x Zea mays, chromosome of H. bulbosum and Zea mays are eliminated.
iv.) Incompatible cytoplasm: Embryo development may be blocked by an incompatibility
between cytoplasm of the species used as female and the genome of the species used as male.
Such an interaction, more generally, leads to hybrid weakness and male sterility in the
hybrids or may sometimes leads to failure of embryo developments.
v.) Endosperm Abortion: Seeds from a large number of distant crosses are not fully
developed and are Shrunken due to poorly developed endosperm. Such seeds show poor
germination, and may often fail to germinate. When the endosperm development is poor or is
blocked, the condition is generally known as endosperm abortion. For e. g.-
1. Triticum sp. x Secale sp. – Triticale. In this case the endosperm aborts at a much later
stage so that a small frequency of viable seed is obtained.
2. Hordeum bulbosum x Hordeum vulgare – the endosperm aborts at an early stage so that
viable seeds are not produced. In case of endosperm abortion - embryo rescue culture is
practiced.
3. Failure of Hybrid seedling development / Hybrid sterility
Some distant hybrids die during seedling development or even after initiation of flowering.
The mechanisms involved in the failure of seedling development most likely involve
complementary lethal genes. For e.g.-
1. In cotton-certain interspecific hybrids appear normal, but die in various stages of seedling
growth; some plants die at flowering.
2. Interspecific and intergeneric F1 hybrids of wheat show both chlorosis and necrosis;
Hybrid sterility: Hybrid sterility refers to the inability of a hybrid to produce viable off
spring. The main cause of hybrid sterility is lack of structural homology between the
chromosomes of two species.
Applications of wide hybridization in crop improvement
1. Alien addition lines: Carries one chromosome pair from a different species in addition to
somatic chromosome complement. For e.g. Disease resistance in Wheat, oats, tobacco
2. Alien substitution lines: has one chromosome pair from different species in place of the
chromosome pair of the recipient parent.
3. Introgression of genes: Transfer of small chromosome segments with desirable genes. e.g.
i. Disease resistance: In Cotton transfer of black arm disease resistance from G. arboreum to
G. barbadense
ii. Wider adaptation: Cold tolerance has been transferred from wild relatives to Wheat, onion,
potato, tomato and grape.
iii. Quality: Oil quality in oil palm was improved by genes from wild relatives.
iv. Changing the mode of reproduction: 1. Self-incompatibility: S.I. genes from B.
campestris to self-compatible B. napus for hybrid seed production.
v. Yield
vi. Other characters
4. Development of New crop species: e.g. Triticale
5. Utilization as New hybrid varieties: e.g. F1 hybrids in cotton Varalaxmi cotton (G.
hirsutum x G. barbadense)
Sugarcane: All the present day commercial varieties are complex interspecific hybrids
involving S. officinarum & S. spontaneum
Limitations of Distant Hybridization:
1. Incompatible Crosses
2. F1 Sterility
3. Problems in Creating New species
4. Lack of homoeology between Chromosomes of the Parental Species
5. Undesirable Linkages
6. Problems in the Transfer of Recessive Oligogenes and Quantitative Taits
7. Lack of Flowering in F 1
8. Problems in using Improved varieties in Distant Hybridization
9. Dormancy

Breeding for disease resistance

Disease is an abnormal condition in the plant produced by an organism
Host: Plant affected by disease.
Pathogen: Organism that produces the disease.
Damage due to disease
i) Reduces total Biomass leading to yield loss
ii) Stunted growth
iii) Sterility
Need for disease resistance breeding
i) To prevent yield loss
ii) High cost reduction
iii) Prevention of environmental pollution
Kinds of disease reaction:
i). Susceptible reaction: Disease reaction is profuse, if unchecked it may lead to total yield
loss.
ii) Immune reaction: Host does not show the symptoms of a disease
iii) Resistance reaction: Infection and establishment takes place but growth of the pathogen in
the host is restricted
iv). Tolerance: Host is attacked by the pathogen in the same manner as the susceptible variety
but there may not be yield loss.
Vertical and horizontal resistance
These terms were introduced by Van der Plank.
Vertical resistance: It is also known as race specific, pathotype specific or specific
resistance. Vertical resistance is generally determined by major genes and is characterised by
pathotypic specificity. Pathotype specificity denotes that the host carrying a gene for vertical
resistance is attacked only by that pathotype which is virulent towards the resistant gene, to
all other pathotypes the host will be resistant.
Only two types of disease reaction can be seen i.e. immune or susceptible reaction. When
virulent pathotype becomes frequent. There may be epidemics. Vertical resistance is not long
lasting.
Horizontal resistance: It is race non-specific, pathotype non-specific or general resistance.
Horizontal resistance is governed by polygenes, that is many genes with small effects and it is
pathotype non-specific. Horizontal resistance does not prevent the development of symptoms
but it slows down the rate of spread to the disease in the population. HR is more stable
compared to VR.
Mechanism of disease resistance:
a) Mechanical: Certain mechanical or anatomical features of host may prevent infection. e.g.
Closed flowering habit of wheat and barley prevents infection by spores of ovary infecting
fungi.
b) Hypersensitivity: Immediately after infection several host cells surrounding the point of
infection die. This leads to death of pathogen also. Phytoalexins present in plant body is
responsible for hypersensitivity reaction.
c) Antibiosis: Presence of some toxic substance. This is more correct for insect resistance.
E.g. Gossypol content in cotton.
d) Nutritional factors: The reduction in growth and spore formation may be due to nutritional
factors of the host.
Genetics of disease resistance:
a) Oligogenic resistance: Resistance is governed by one or few major genes and resistance is
generally dominant. The action of major genes may be altered by modifiers. Gene for gene
relationship Flor (1956) proposed this based on his work in linseed rust. According to this for
every resistance gene present in the host, the pathogen has a gene for virulence. Susceptible
reaction will result when the pathogen is able to match all the resistant genes with virulence
gene.
b) Polygenic inheritance: The genes show both additive and non - additive effects and there is
large environmental effects.
c) Cytoplasmic inheritance: T cytoplasm - Maize Tift 23A cytoplasm - Cumbu Susceptible to
disease. C and M cytoplasm of maize resistant to Helminthosporium. L 111A and 732, A
cytoplasm resistant to downy mildew in cumbu.
Methods of disease resistance breeding
1. Plant introduction: Resistant varieties from other can be directly introduced for
cultivation. e.g. IR 20 rice resistant to blast.
2. Selection: This may be from local land races or from introduced cultivars. e.g. Co 4 Gobi
Anaikomban resistance to blast. NCAC 17090 ground nut resistant against leaf spot.
3. Hybridisation and Selection:
a) Intervarietal hybridization - Co37 Rice resistant to blast
b) Inter specific hybridization - Powdery mildew resistance in Sesamum
c) Inter generic - Atylosia for root rot in red gram. Depending on gene action the selection
procedure may vary.
If the resistance is governed by polygenes, then pedigree method of selection is to be
followed.
If the resistance is governed by major genes linked with other undesirable characters we have
to go for back cross method of breeding. For dominant gene the back cross method is
different from recessive gene governed traits.
4. Mutation breeding: Co2. Ground nut tolerant to late leaf spot disease.
5. Polyploidy breeding: Nicotiana crosses for resistance against leaf spot.
6. Tissue culture method: Resistance reaction can be screened easily in test tubes and
resistant lines can be mass multiplied. e.g. Banana, Cardamom.
Screening techniques for disease resistance
Depending on mode of spread of disease the screening technique may differ. The screening
can be done both at screen or glass house level and field level. The different screening
techniques are as follows:
Soil borne diseases: Wilt and root rot are produced by soil borne fungi. In this case sick plot
technique is followed. Susceptible varieties can be grown and infected plants can be ploughed
in situ to maintain optimum condition for infection.
Air borne diseases: e.g. Rust, Smut, mildews, blights. For ground nut rust, infestor rows can
be sown 15 days earlier as border rows and the disease will infest the susceptible infestor
rows. After 15 days the varieties tested to be are to be sown. Spraying the spore suspension
from affected leaves will also increase the load.
Seed borne disease: Smut, bunt etc. Artificial inoculation can be done by soaking the seeds
in solution of pathogen under vacuum condition. Insect transmitted diseases.
Insect transmitted diseases: e.g. virus diseases, Red gram sterility mosaic virus.
Sap transmitted: Here the stapling technique is used. Leaves from affected plants can be
stapled to the entries to be tested. The insect feeding in susceptible leaf will transmit virus to
test entries.
Inbreeding
depression and
heterosis
– Cross pollinated species are highly heterozygous in nature

– So upon inbreeding they show loss in vigour and conversely show hybrid vigour
when two different plants are hybridized
Inbreeding depression

– Reduced biological fitness in a given population as a result of inbreeding or

breeding of related individuals.

– The highest degree of inbreeding is?????

Effects of inbreeding

1. Appearance of lethal and sublethal alleles

2. Reduction in vigour
3. Reduction in reproductive ability
4. Increase in homozygosity
5. Reduction in yield
6. Separation of population into distinct lines
Degree of Inbreeding depression

– High Inbreeding depression- eg alfalfa, carrot plants die

– Moderate Inbreeding depression- eg maize, bajra
– Low Inbreeding depression- eg onion, sunflower
– Lack of Inbreeding depression- all SP crops
Heterosis

– When two homozygous inbreeds (a true breeding line obtained by continuous

inbreeding) of genetically unlike constituents are crossed together, the resulting
hybrids obtained from the crossed seeds are usually robust, vigorous,
productive and taller than the either parents.

– This increased productivity or superiority over the parents is known as heterosis

or hybrid vigour.

– Heterosis can be defined as the superiority of F1 hybrid over both the

parents in terms of yield or some other character.
History

– Heterosis has been known since the art of hybridization came into existence.

– Koelreuter (1763) was the first to report hybrid vigour in the hybrids of tobacco,
Datura etc.

– Mendel (1865) observed this in pea crosses.

– Dr. G.H. ShuII (1914) proposed the term heterosis (Gr. heteros different and osis
= condition).
– More appropriate to term the developed superiority of the hybrids as hybrid vigour and
to refer to the mechanism by which the superiority is developed as heterosis.
– Heterosis is estimated in three different ways,
– 1) Mid parent heterosis
– 2) Better parent heterosis
– 3) Standard heterosis
Mid parent heterosis

– When the heterosis is estimated over the mid parent i.e. mean value or average
of the two parents
– It is also known as average heterosis It is also known as average heterosis or
relative heterosis
 Heterobeltiosis
2) Better
Parent Heterosis (Heterobeltiosis)
• The term heterobeltosis is used to describe the improvement of heterozygote
over the better parent of the cross.

• When the heterosis is estimated over the better parent is known as better parent
heterosis or heterobeltiosis
 Standard heterosis
It refers to the superiority of F1 over the standard commercial check variety. It is also
called as economic heterosis or useful heterosis
Heterosis
 Genetic basis of heterosis

– Three main theories to explain heterosis

– Dominance hypothesis
– Overdominance hypothesis
– Epistasis hypothesis
Dominance hypothesis

– The dominant hypothesis of heterosis was proposed by Davenport (1910),

Bruce (1910) and Keeble and Pellow (1910).
– According to dominance hypothesis, heterosis is the result of accumulation of
dominant genes in F1 from both the parents.
– Suppose the genetic constitution of female parent is AABBccdd and that of male
parent as aabbCCDD.
– A hybrid between these two parents will have four dominant genes (Aa Bb Cc
Dd) and exhibit superiority over both the parents having two dominant genes
each.
Objections

– i) If the dominance hypothesis is true, it should be possible to obtain pure

heterotic individuals in F2 which are homozygous for all the dominant genes.
However, such individuals are never obtained.

– (ii) If the heterosis is due to dominance, the F2 curve should be skewed towards
dominant genes, but the curve of F2 is found always smooth and symmetrical.
Over dominance hypothesis

– The over dominance hypothesis of heterosis was independently proposed in

1908 by Shull and East and it was supported by East (1936) and Shull (1945).
– According to this hypothesis, heterosis is due to superiority of heterozygote
over its both homozygous parents.
Epistasis

– In 1952 Gowen suggested that influence of one locus on the expression of

another may be the cause of heterosis
– Many cases showed it to be true
– But all heterotic crosses do not show epistasis and all crosses that show
epistasis they are not heterotic
Breeding in CP
plants
– The population improvement methods may be grouped into two classes.
– 1. Selection without progeny testing. Eg : Mass selection.
– 2. Selection with progeny testing
a. Progeny selection or ear to row method
b. Recurrent selection
a) Progeny selection:
– The simplest form of progeny
selection is the ear to row
method, extensively used in
(50-100)
maize.
– This method was developed
by Hopkins in 1908.
b) Recurrent Selection:
In cross pollinated populations, scheme of selection (on the basis of phenotype or progeny
test) followed by intermating (in all combinations) of the selected plant to produce
population for the next cycle of selection.

Improve the frequency

of desirable alleles for a
character in population

Note: The recurrent selection schemes were devised in relation to heterosis breeding.

The idea was to ensure the isolation of superior inbreds from the populations subjected to
recurrent selection for their ultimate utilization in the production of hybrid and synthetic
varieties.
 Main features

– Recurrent Selection is modified form of Progeny Selection.

– Application: It is more commonly used in cross pollinated crops

– Base population: Heterozygous population required

– End product: It can be used to produce inbreds, hybrid varieties and synthetic
varieties

– Impact: Improve the frequency of desirable alleles for a character in population

Recurrent Selection
1. Simple Recurrent Selection

2. Recurrent Selection for General Combining Ability

3. Recurrent Selection for Specific Combining Ability

4. Reciprocal Recurrent Selection

Procedure of Simple
Recurrent Selection

• A type of recurrent
selection which does
not include testers

• Condition of use
This method is useful only
for those characters which
have high heritability
 Simple Recurrent Selection
– In simple recurrent selection:
1. A number of plants with desirable phenotype are selected and self-pollinated.
2. In the second year, separate progeny rows are grown from selfed seeds of the selected plants.
3. The progenies are intercrossed in all possible combinations by hand, and equal amount of seed
from each cross is composited to produce the next generation. This completes the original
selection cycle.
4. For recurrent selection, several desirable plants are selected from the composited population
obtained from the original selection cycle; they are selected on the basis of phenotype and are self-
pollinated.
5. Next year, progeny rows are grown from the selfed seed and all possible intercrosses are made by
hand.
6. Equal seeds from all the intercrosses are composited to produce the next generation.
7. This constitutes the first recurrent selection cycle. The population may be subjected to one or
more recurrent selection cycles
 Recurrent Selection for General Combining Ability (RSGCA)

• A form of recurrent selection that is used to improve general combining ability

of a population for a character and includes heterozygous tester with a broad
genetic base

– A tester strain is the common parent mated to a number of lines, strains or plants

• Condition of use : This method is used for the improvement of those

characters which are governed by additive gene action

• Impact of this method: It improves the gca of a character

1. First Year. A number of phenotypically outstanding plants are selected from the source population. The source
population may be an open-pollinated variety, a synthetic or an advanced generation of a hybrid. Each selected
plant is selfed as well as crossed (as male) to a number of randomly selected plants from a tester with broad genetic
base. The selfed seeds are harvested separately and saved for planting in the third year. The test-cross progeny
(plant x tester progeny) from each selected plant is harvested separately and used for a replicated yield trial in the
second year.
2. Second Year. A replicated yield trial is conducted using the plant x tester progenies. The superior progenies are
identified.
3. Third Year. Selfed seeds (from the first year) from those plants that produced superior test-cross progenies
(identified on the basis of yield trial in the second year) are planted in separate progeny rows in a crossing black.
These progenies are intercrossed in all possible combinations. Equal amounts of seeds from all the intercrosses are
composited to obtain the next generation. This completes the original cycle of selection.
4. Fourth Year. The seed obtained from bulking of all the intercrosses is planted as the source population for the first
cycle of recurrent selection. Several plants are selected on the basis of their phenotype; each of them is selfed as
well as crossed (as male) to a number of random plants from the tester with wide genetic base.
5. Fifth Year. Operations of the second year are repeated.
6. Sixth Year. Operations of the third year are repeated. This completes the first recurrent selection cycle.
7. Seventh Year. The second recurrent selection cycle may be initiated as in the case of the first recurrent selection
cycle in the fourth year. In this manner several cycles of recurrent selection may be carried out.
 Recurrent Selection for Specific Combining Ability (RSSCA)

• A form of recurrent selection that is used to improve specific combining ability

of a population for a character using homozygous tester.

– The procedure for RSSCA is identical with that for GCA, except that in this case an inbred is used as a tester in the
place of an open-pollinated variety.

• Condition of use : This method is used for the improvement of those characters
which are governed by non- additive gene action
• Impact of this method: It improves the sca of a character
• The objective is to isolate from a population such lines that will combine well
with a given inbred.
1. First Year. Several plants are selected from the population and self-pollinated. The selected plants (used as males)
are also crossed to an outstanding inbred used as the tester (used as female).
2. Second Year. A replicated yield trial is planted using the testcross progeny. Outstanding progenies are identified
3. Third Year. Selfed seed from the plants that produced the outstanding progenies are planted in separate progeny-
rows in a crossing block. All possible intercrosses among these progenies are made by hand. Equal amounts of seed
from all the intercrosses are composited. This completes the original selection cycle.
4. Fourth Year. The composited intercross seed is planted and the operations of the first year are repeated.
5. Fifth Year. Operations of the second year are repeated.
6. Sixth Year. Operations of the third year are repeated. This completes the first recurrent selection cycle. The
population may be subjected to one or more recurrent selection cycles, if desired, by repeating the operations of
the first recurrent selection cycle (fourth to sixth years).
 Reciprocal Recurrent Selection (RRS)

• A form of recurrent selection that is used to improve both gca and sca of a
population for a character using two heterozygous tester.

• Condition of use : This method is used for the improvement of those characters
which are governed by additive and non- additive gene action
• Impact of this method: It improves the gca and sca of a character
• The objective of RRS is to improve two different populations in their ability to
combine well with each other
Heterosis can be fully exploited in the form of hybrids in cross
pollinated species

In cross-pollinated species, heterosis can also be exploited

partially in the form of synthetic and composite varieties.

Definition of Synthetic Variety:

A Variety which is produced by crossing in all combination a

number of inbred lines that combine well with each other.

Once synthesized, a synthetic is maintained by open-

pollination in isolation is referred as synthetic variety.

Hayes and Garber suggested the commercial utilization of

synthetic varieties in Maize in 1919.
Features of Synthetic Variety
1. Heterogeneous
2. Synthetic variety can be developed by using clones, inbreds or OPV i.e. open
pollinated variety
3. Cross pollination is must
4. Maintained by open pollination
5. Unlike composite variety, exact reconstitution of synthetic variety is possible
6. More adaptive to varying growing conditions as compared to hybrids
7. Less uniform as compared to hybrids
8. Less attractive as compared to hybrids
9. Show some amount of heterosis as compared to OPV
10. Have better disease resistance
Steps in Development of Synthetic Variety:
Development of synthetic variety consists of three major steps. i.e.

1) Evaluation of lines for GCA.

2) Production of synthetic variety,

3) Multiplication of synthetic variety.

Step 1. Evaluation of Lines for GCA

The lines that make up a synthetic variety may be inbred lines,

clones, open pollinated variety.

Inbred lines are evaluated for general combining ability

There are three different methods of evaluating, which is
produced by GCA.

1. Top cross
2. Polycross
3. Single cross
In top cross, the inbreds are crossed with a common tester and the
progeny are evaluated in replicated trials for general combining
ability of yield and yield contributing characters.

In Polycross, selected inbreds are allowed to intermated by open

pollination in isolation.

Single cross all possible single crosses are made among selected
inbreds. These crosses are evaluated for GCA of yield in replicated
trial using local variety as a check.

Thus, inbred lines with good GCA are identified and finally
selected for development of synthetic variety
Step 2. Production of A Synthetic variety

A synthetic variety may be produced in one of the following two

ways.

1. Equal amounts of seeds from the parental lines are mixed and
planted in isolation.
• Open-pollination is allowed
• The seed from this population is harvested in bulk; the
population raised from this seed is the Syn1 generation.

2. All possible crosses among the selected lines are made in

isolation.
Equal amounts of seed from each cross is composited to produce
the synthetic variety.
Step 3. Multiplication of Synthetic Varieties

Multiplied in isolation for one or more generations before its

distribution for cultivation.

The synthetic varieties are usually maintained by open-

pollinated.
 Merits and Demerits Synthetic Variety

Merits

•Less costly compared to hybrids

•Farmer can maintain his synthetic variety for more seasons
which is not possible in hybrids
•Because of wider genetic variety base, the synthetics are more
stable over years and environments.
•Seed production is more skilled operation in hybrids where as it
is not so in synthetics.
•Synthetic varieties have vast genetic variability which provides
them better protection from the infestation of new races of a
pathogen.
Demerits

•Performance is little bit lower compared to single or double

cross hybrids because synthetics exploit only GCA while hybrids
exploit both GCA and SCA.
•The performance of synthetic is adversely affected by lines with
relatively poorer GCA.
•The performance of synthetic varieties is less uniform and less
attractive.
•Synthetic varieties are utilized in cross pollinated species only,
Whereas hybrids can be developed in both cross & self
pollinated species
Achievements of Synthetic Varieties:

Synthetic varieties have been developed in cross- pollinated

crops like maize, pearl millet, sunflower, Sugarbeet, alfalfa,
lucerne, etc. in U.S.A.

In India synthetic varieties have been evolved in pearl millet

at ICRISAT and in Sugarbeet at pantnagar.
Ex. Sugarbeet- pant synthetic – 3, Cauliflower- synthetic -3,
bajara- ICMS-7703.
 Composite varieties

Composite varieties

In cross pollinated crops, the varieties produced by open

pollination among a number of outstanding strains usually not
tested for combining with each other.
Features of Composite Variety

1. Heterogeneous
2. Relevant to cross pollinated species only
3. Can be developed from open pollinated variety or any other
heterozygous variety
4. Farmer can use his own saved seed for 3 to 4 years, after that
seed should be replaced
5. There can be two or more constituent genotypes
6. Evaluation for general combining ability (gca) as in synthetic
variety production, is not carried out
7. Exact reconstitution of composite variety is not possible
Steps In Development of Composite Variety

1. Selection of parental material

2. Intermating of selected genotype.
3. Evaluation of crosses.
4. Mixing of parental material of superior crosses in equal
quantity
1. Selection Of Base Material
Open pollinated varieties or other heterozygous sources serve as the
base material.

Several open pollinated varieties with similar maturity duration ,

height , seed colour are selected for this purpose.

2. Intermating Selected Genotype

Selected genotypes are crossed in all possible combinations.

Total crosses to be made are equal to n(n-1)/2, where n is the no
of varieties selected for intermating.
3. Evaluation Of Crosses

All the single crosses are evaluated in replicated trials for yield performance
in F1,F2,F3 & F4 generations using standard hybrid or open pollinated
variety as a check.

Crosses exhibiting little or no inbreeding depression in F2,F3 &F4

generations are identified & selected for the development of composite
cultivar.

4. Mixing Parental Seed of Superior Crosses

Parental seeds (of the superior crosses) is mixed in equal quantity to

develop a composite variety.
Seed multiplication is carried out by open pollination in isolation for 1 or 2
generations.
 Procedure of developing composite variety

Year Main breeding activity

First Selection of base material, intermating in all possible
combination and harvesting crossed seed separately
second Evaluation of F1 crosses in replicated trial using hybrid or
open pollinated variety as check.
Third to fifthEvaluation of F2, F3 and F4 generations in replicated trial
using standard check, identification of crosses exhibiting
little or no inbreeding depression.
Sixth and Mixing parental seed of superior crosses in equal quantity
seventh to constitute composite variety, seed multiplication by
open pollination in isolation.
Eighth Release of new composite variety , distribution of seed to
the farmers for commercial cultivation.
Achievements: Composite varieties

1. In India, the first composite varieties were released in 1967: the

six maize composites were, Ambar, Jawahar, Kisan, Vikram, Sona
and Vijay.
2. Some of the recently released maize composites are: Co 1 (full
season, resistant to downy mildew), Renuka (very early), Kanchan
(very early). These composites yield as much as 90 per cent of the
best hybrid varieties.
3. Three opaque 2 composties, viz., Shakti, Ratan and Protina, have
been released in maize. They have twice the amount of lysine and
tryptophan as compared to the normal maize hybrids and
composites.
4. A composite variety, Composite I, has been evolved in Brassica
campestris var. toria.
Differences between Synthetic and Composite Cultivars

Particulars Synthetic cultivars Composite cultivars

Base Material Inbreds or clones Open pollinated varieties or

any other heterozygous
source
Component Two to eight Two to many
genotypes
Evaluation of Always done Usually not done
GCA
Seed After 4-5 years After 3-4 years
Replacement
Exact Possible Not possible
Reconstitution
 Development of inbred lines through conventional
and anther culture method

A) Conventional method

1. Continuous selfing

B) Tissue culture method

1. Anther culture method
 Haploid and doubled haploid

– Haploid: An individuals with the gametic chromosome number (n).

– Doubled haploids (DHs): A plant or line obtained by doubling the chromosome number of a
haploid plant (n).

The resulting individual is completely homozygous.

B) Tissue culture method
1. Anther culture 2. Ovary or ovule culture

What is Anther Culture?

Anther culture is a technique by which the developing anthers at a precise and critical stage are
excised from unopened flower bud and are cultured on a nutrient medium, where the
microspores within the cultured anther develop into callus tissue or embryoids that give rise to
haploid plantlets.
Callus:-

In plant cell biology a callus is a mass of

undifferentiated cells (usually cultivated on gel media that
develops during tissue culture of plant parts).

Proliferation:-

Cell proliferation is the process that results in an increase of the

number of cells.

Embryoid:-

Mass of plant or animal tissue that resembles an embryo.

Or Embryo like structures produced as a result of differentiation
processes such as embryogenesis and androgenesis.
 What is Hybrid?
Hybrids are the first generation (F1 ) from a cross between
two pure lines or inbred line that are genetically dissimilar.
Most of the commercial hybrids are F1s from two or more
pure lines (tomato, rice), inbred lines (maize, sunflower,
caster etc.), open pollinated varieties.
Pureline: Progeny of a single homozygous self-
pollinated plant.

Inbred line:- A nearly homozygous line developed

by continued inbreeding, usually selfing.
 Types of hybrids

1. Intraspecific hybrids
2. Interspecific hybrids
1. Intraspecific hybrid: A hybrid between
genetically different genotypes of the same
species.
Also called intervarietal hybrid.
These hybrid are always fertile.

Three types of Intraspecific hybrid

1. Single cross
2. Three way cross
3. Double cross
1. Single cross: when two inbred lines or pure lines are crossed to produce
the F1 hybrid it
is known as single cross.

A X B

P P
1 2

F1
Number of single crosses: n(n-1)/2
Where, n – Number of parents
2. Double cross :
when two single
crosses are crossed
the resulting hybrid
population is
known as double
cross.

Four inbred lines are

used in Double cross
The high productivity of F1 used as female.
High pollen production efficiency of F1 used as
male parent.

Number of Double crosses: n(n-1)(n-2)(n-3)/8

 Three way cross :
A AX B
It is a cross between a
single cross and an
inbred to give hybrid
population

F1 X C
Three inbred lines are
used in three way cross

F1
The single cross is used as female and inbred as
male

Number of Three way crosses: n(n-1)(n-2)/2

Interspecific hybrid: The F1 progeny between two
different species of the genus is known as Interspecific
hybrid.

Also called intrageneric hybrid

Such hybrids are fertile in few cases.
DISTANT HYBRIDIZATION FOR CROP IMPROVEMENT
Introduction

❑ Hybridization
Crossing between two genetically dissimilar parent is called
hybridization.

❑ Distant hybridization
Hybridization between individuals from different species belonging
the same genus (interspecific hybridization) or two different
genera of same family (intergeneric hybridization) is termed as
distant hybridization and such crosses are known as distant crosses
or wide crosses.
History

❑ Thomas Fairchild (1717):

The first authentic record of a distant hybridization for crop improvement

is the production of a hybrid between Carnation (Dianthus caryophyllus)
and Sweet willian (Dianthus barbatus).

❑ Karpechenko (1928):

An intergeneric hybrid, between Brassica (cabbages) and Raphanus

(radish). Raphanobrassica, was produced.

❑ Rimpu (1890):

Produce the first intergeneric hybrid triticale (Wheat X Rye) which have
greater potential than raphanobrassica.
 Inter- specific hybridization:

– Ex. Nerica, an upland rice for Africa

– Oryza sativa (Asian upland rice) : non-shattering , resistant to lodging,

high yield potential

– Oryza glaberrima(African rice): drought tolerant, disease resistant, weed-

suppressing

➢ Nerica rice combines the best of both species.

 Intergeneric crosses

– Triticale, a new cereal created in the lab.

– Triticale, a cross (intergeneric cross) between wheat and rye, was

produced by embryo rescue of the product of fertilization and a
chemically induced doubling of the chromosomes.
Intergeneric combinations of wheat and rye that produces hexaploid
and octaploid triticale
 Problems associated with wide crosses

❑ The major problems associated with wide crosses are:

– Cross Incompatibility

– Hybrid Inviability

– Hybrid Sterility

– Hybrid Breakdown
CROSS INCOMPATIBILITY

❑ This is the inability of the functional pollen grains of one species or genus to effect

fertilization in another species or genes.

– There are three main reasons of cross incompatibility viz.

I. Lack of pollen germination,

II. Insufficient growth of pollen tube to reach ovule and

III. Inability of male gamete to unite with the egg cell.

– These barriers are known as pre –fertilization barriers.

– This is overcome by employing different techniques like reciprocal crosses, bridge

crosses, using pollen mixtures, pistil manipulations, use of growth regulators etc.
HYBRID INVIABILITY

❑ This refers to the inviability of the hybrid zygote or embryo. In some cases,
zygote formation occurs, but further development of the zygote is arrested. In
some other cases, after the completion of the initial stages of development, the
embryo gets aborted.

– The reasons for this are:

1. Unfavorable interactions between the chromosomes of the two species

2. Unfavorable interaction of the endosperm with the embryo.

3. Disharmony between cytoplasm and nuclear genes

– Reciprocal crosses, application of growth hormones and embryo rescue are

the techniques that can be used to overcome this problem.
HYBRID STERILITY

❑ This refers to the inability of a hybrid to produce viable offspring.

This is more prominent in the case of intergeneric crosses.

❑ The major reason for hybrid sterility is the lack of structural

homology between the chromosomes of the two species.

– Sterility caused by structural differences between the chromosomes

of two species can be overcome by amphidiploidization using
colchicine.
 HYBRID BREAKDOWN

❑ Hybrid breakdown is a major problem in interspecific crosses.

– When F1 hybrid plants of an interspecific crosses are vigrous and fertile but
there F2 progeny is weak and sterile it is known as hybrid breakdown.

– So hybrid breakdown hinders the progress of interspecific gene transfer.

– This may be due to the structural difference of chromosomes or problems

in gene combinations.
 TECHNIQUES TO MAKE WIDE CROSSES SUCCESSFUL

1. SELECTION OF PLANTS
The most compatible parents available should be selected for the crosses.

2. RECIPROCAL CROSSES
Reciprocal cross may be attempted when one parental combination fails.
e.g. Mung x udid- cross compatible and Udid x mung-cross incompatible

3. MANIPULATION OF PLOIDY
Diploidization of solitary genomes to make them paired will be helpful to make the
cross fertile.
4. BRIDGE CROSSES
When two parents are incompatible, a third parent that is compatible with both the
parents can be used for bridge crosses and thus it becomes possible to perform cross
between the original parents.
e.g. Tobacco
-Nicotiana repanda x N.tabaccum– cross incompatible
-Nicotiana repanda x N.sylvestris- cross compatible
-Product x N.tabaccum- cross compatible

5. USE OF POLLEN MIXTURE

Unfavorable interaction between pollen and pistil in the case of wide crosses can be
overcome to some extent by using pollen mixture.
 Continue.......

6. MANIPULATION OF PISTIL
Decapitation of the style will sometimes prove helpful in overcoming incompatibility.

7. USE OF GROWTH REGULATORS

Pollen tube growth can be accelerated by using growth hormones like IAA, NAA, 2,4-D
and Gibberellic acid.

8. PROTOPLAST FUSION
When fusion of gametes fails, protoplast fusion of somatic cells can be attempted.

9. EMBRYO RESCUE
Hybrid zygotes formed by wide crosses may fail to grow in a number of cases. The zygotes
are taken out and grown in in vitro medium to overcome this problem.
Achievements

– Hybrid varieties:

➢ Upland cotton – MCU-2, MCU-5, Khandwa1, Khandwa2 etc are

derivatives of interspecific hybridization.

➢ Hybrid between Pearl millet x Napier gress- Hybrid Napier which is very
popular for its high fodder yield and fodder quality e.g. Jaywant and
Yashwant

➢ Interspecific hybrids in cotton- Varlaxmi, Savitri, DCH-32, NHB-12, DH-

7, DH-9 etc.

➢ Prabhani Kranti variety of bhindi.

Pre-breeding

– Pre-breeding refers to all activities designed to identify desirable

characteristics and /or genes from unadapted materials that cannot be used
directly in breeding populations, and to transfer these traits to an
intermediate set of materials that breeders can use further in producing new
varieties for farmers.
– This is also known as developmental breeding or germplasm enhancement.
– The lines developed through pre-breeding may not be suitable for commercial
cultivation but serves as carriers of genes from wild species and can easily be
utilized for the development of commercial cultivars.
Backcross
Method
– The hybrid and the progenies in subsequent
generations are repeatedly backcrossed to one of the
parents.
– As a result, the genotype of the backcross progeny
becomes increasingly similar to that of the recurrent
parent.
Pre-requisite for back cross
breeding
– 1. A suitable recurrent parent must be available which lacks in one or two
characteristics.
– 2. A suitable donor parent must be available
– 3. The character to be transferred must have high heritability and preferably it
should be determined by one or two genes.
– 4. A sufficient number of back crosses should be made so that the genotype of
recurrent parent is recovered in full
Application of back cross method

– This method is commonly used to transfer disease resistance from one variety to another. But
it is also useful for transfer of other characteristics.
– 1. Intervarietal transfer of simply inherited characters E.g. Disease resistance, seed coat colour
– 2. Intervarietal transfer of quantitative characters. E.g. Plant height, Seed size, Seed shape.
– 3. Interspecific transfer of simply inherited characters E.g. Transfer of disease resistance from
related species to cultivated species. E.g. Resistance to black arm disease in cotton from wild
tetraploid species into Gossypium hirsutum
– 4. Transfer of cytoplasm This is employed to transfer male sterility. The female parent will be
having the sterile cytoplasm and recurrent parent will be used as male parent.
– 5. Transgressive segregation Back cross method may be modified to produce
transgressive segregants. The F1 is backcrossed to recurrent parent for 2 to 3
times for getting transgressive segregants.
– 6. Production of near isogenic lines
– 7. Germplasm conversion E.g. Production of photo insensitive line from photo
Sensitive germplasm through back crossing. This was done in the case of
sorghum
Procedure for backcross method

– The Plan of backcross method would depend upon whether the gene being
transferred is recessive or dominant.
– The plan for transfer of a dominant gene is simpler than that for a recessive
gene.
 Transfer of
dominant
allele
First Season Hybridization between parent B donor
(Female) and Susceptible parent A recipient (male)
Second Season Raising the F l and backcrossed to
recurrent parent A.
Third season Growing the BClFl . It will be
segregating for 1 susceptible (rr): 1 resistant (Rr).
Rust resistant plants are backcrossed with recurrent
parent A. This is second backcross.
Fourth Season Raising BC2 F1 will again segregate
in the ratio of 1: 1. Third backcross is done with
resistant plants.
– Fifth Season to Eighth Season Raising backcross Fls and crossing resistant plants
with recurrent parent is continued up to 7th backcross
– Ninth season Raising BC7F1 and observing resistant lines. The plants resembling
parent A coupled with resistance is selected and harvested on single plant basis.
– Tenth Season Growing the progeny row and observing each row for resistance.
Best rows are selected and harvested
– Eleventh Season The row bulk is raised in yield trial along with check and the
best plots are selected.
– Twelfth season Selected plot seeds are multiplied and released as new variety.
– Steps
– I Season: Make a cross between donor parent A and recurrent parent B and Harvest the hybrid. The
donor parents A is resistant which is governed by recessive genes. The susceptibility is governed by
dominant gene in parent B.
– II Season: Grow the F1 which will be susceptible – Harvest them.
– III Season: Grow F2 which will be segregating in the ratio of I :2: I i.e. 3/4 susceptible and 1/4 resistant.
With the resistant lines (rr) make first backcross with parent A having dominant RR gene. Harvest BCI
F1
– IV Season: Grow BCIF1
– V season: Grow BC1 F2 which will be segregating as we saw in III season. Repeat the process of third
season. This will give BC2F1I
– VI Season : Grow BC2F1
– VII season: Grow BC2F2 them repeat the process of V Season. This will give BC3F1.
– VIII Season: Grow BC3F1
– IX Season: Grow BC3F2 and repeat the process of VII Season. Harvest BC4F1.
– X season: Grow BC4F1
– XI Season: Grow BC4F2 and repeat the process of IX Season. Harvest BC5F1.
– XII, XIII & XIV: Repeat the process and carry out backcross upto 7 time.
– XV Season: While studying BC7F2 select single plants having resistance and resembling
parent B.
– XVI Season: Study the progenies in progeny rows and select best progenies.
– XVII Season: Conduct yield trial and select best material. XVIII Season: Multiply the seeds
and distribute it as improved variety with resistance to disease.
 What is Multiline Variety – Definition?

Multiline variety is a mixture of several pure lines of similar phenotype (height, seed
color, flowering time, maturity time and various other agronomic characteristics) but
have different genes for disease resistance (means these are isogenic lines).
 Main features of multiline varieties are briefly presented below:

1. Application:
The multiline approach is applicable to self-pollinated species only. Multiline
cultivars are commercially used in self-pollinated crops like oat, wheat, soybean,
groundnut and many other crops.

2. Genetic Constitution:
Multiline cultivars are mixtures of several pure-lines. The pure-lines may be
isogenic lines, closely related lines or unrelated lines. Thus, multi-lines are
homozygous but heterogeneous populations or genetically diverse populations.
3. Adaptation:
Multi-lines are more adaptable to environmental variations than pure-lines by
virtue of their genetic diversity. In other words, multi-lines have more buffering
capacity to environmental changes than pure lines.

4. Disease Control:
The use of multiline cultivars is an effective way to minimize the yield losses due
to the attack of multiracial disease.

In a multiline cultivar, each component genotype has a resistant gene for a

different race of a disease.

All races of a disease will never appear at a time and all the genotypes of a
heterogeneous mixture are never attacked at a time.
5. Qualityof Produce:
The produce of multiline cultivars is generally less uniform and less attractive
than that of a pure-line, because it is a mixture of several pure lines.

6. Yield:
The yield of a multiline would be lesser than that of the most productive
cultivar of a pure-line under normal conditions.

But under adverse conditions, the yield of a multiline would be much higher
than that of most productive pure-line cultivar.

Because highly productive pure-line cultivars are more prone to biotic and
abiotic stresses due to narrow genetic base.
MULTILINE
BREEDING
Multilines:
First Suggested IN OATS BY JENSEN IN 1952
Multilines are developed for self-pollinating species.

These cultivars consist of a mixture of specially developed

genotypes called isolines (or near isogenic lines) because they differ
only in a single gene (or a defined set of genes).

Isolines are developed primarily for disease control, even though

these cultivars could, potentially, be developed to address other
environmental stresses.

Isolines are developed by using the techniques of backcrossing in

which the F1 is repeatedly crossed to one of the parents (recurrent
parent) that lacked the gene of interest (e.g., disease resistance).
1. Mixtures of isolines.

2. Mixture of closely related lines.

3. Mixture of unrelated or distinctly

different genotypes.
1. It has genetic diversity.

2. It should have normal resistance.

3. Components should be uniform

agrnomically.

4. It should have yield advantage.

1. Selection of recurrent parent.
2. Selection of donor parents.
3. Transfer of resistance.
4. Mixing of isolines.
1. Identical to recurrent parent
2. Loss to cultivator is low.
3. Spreading of diseases is slow
4. It reduces the risk of homogenising the
pathogen population globally.
5. Stabilises and optimizes production on
farm.
1. Seed is to be repaced every few years.
2. Production and mainatenance is time taking
job.
3. It has to be regularly reconstituted, and new
lines have to be continuously developed.
4. Seed certification poses difficulties.
5. It is less attractive, less uniform and costly
method.
 Multiline cultivars have been developed for
commercial cultivation in oats, wheat,
soybean and peanut in USA.
 In India three multiline varieties, viz.
KSML 3, MLKS 11 and KML 7404 have
been released in wheat from Punjab.
 PLANT HYBRIDIZATION
Genotype peculiarities of two or more different varieties or species of
plants could be brought together only by crossing them. The mating or
crossing of two plants or lines of dissimilar genotype is called
hybridization. In plant hybridization, one of the plants is taken as the
female plant and the other as the male plant. Pollen grains from the
male parent are made to pollinate the stigma of the flowers of the
female parent. The seeds obtained from such a cross are called F1 seeds
and the progeny raised from it is called F1 (First Filial) generation. The F1
is selfed to produce F2 and the subsequent generations like F3, F4 etc are
raised in the same way. These generations are called segregating
generations and they are handled differently based on the scope and
objectives of the breeding program.
 OBJECTIVES OF HYBRIDIZATION
The major objectives of hybridization are:
A. Combination Breeding
Combination breeding is the transfer of one or more characters from other
varieties to a particular variety. These characters may be oligogenic or
polygenic. In this way, genes for disease resistance, quality traits etc can be
transferred.
B. Transgressive Breeding
Transgressive breeding is based on transgressive variation of characters in
segregating generations like F2. A cross is made between two strains of plants
and the F2 is screened for transgressive variations.
Transgressive segregation is the segregation of characters beyond the parental
limits, in the segregating generations like F2.
C. Production of Hybrids
Hybrid (F1) plants show higher vigor and yield when compared to parents, in
some cases. This phenomenon is called hybrid vigor. F1 seeds can be raised in
bulk through hybridization and distributed directly for cultivation, especially in
cross-pollinating crops.
 TYPES OF HYBRIDIZATION
Based on the genetic difference between parents, hybridization can be
classified into:

1. Inter-varietal Hybridization
2. Distant Hybridization
INTER-VARIETAL HYBRIDIZATION
The cross between the members of the same species (intra-specific) is
called inter-varietal hybridization. In this type of hybridization, different
cross patterns can be used.

1. Simple Cross
In this case, two parents are used to produce an F1 hybrid.

2. Complex Crosses
In complex crosses, more than two parents are involved. Such
crosses can be called convergent crosses since they bring genes
from different sources together.
 DISTANT HYBRIDIZATION
• Hybridization between the members of different species or hybridization
beyond species level is called distant hybridization. Thus, it may be
interspecific (intra-generic) or inter-generic. When conventional methods
of hybridization fails, para-sexual methods are used in such cases.

• Para-sexual hybridization is the technique of fusing somatic protoplasts

when reproductive cells fail to fuse or fertilize.
 THE PROCESS OF HYBRIDIZATION
The major steps involved in the process of hybridization are:

1. Selection of Parents
2. Emasculation
3. Bagging
4. Tagging
5. Pollination
6. Harvesting F1 Seeds
7. Further handling of the plants
 SELECTION OF PARENTS
The choice of the parents depends on the objective of the cross. In
combination breeding, the genetic diversity of the parents is not
important. In the case of transgressive breeding, genetically diverse
plants are selected as parents. If the characteristics of the parents are
not completely known, they are evaluated for the agronomic features.
 EMASCULATION
In the case of crops with bisexual flowers, stamens of the flowers of the
female parents are removed or the pollen grains are killed. This process
is called emasculation. Mechanical, physiological or genetic methods of
emasculation are used, depending upon the nature of the crop and the
cross.
 MECHANICAL METHODS OF
EMASCULATION
Here, the anthers are removed from the flowers of the female parents.
Hand emasculation and suction method are generally used. For hand
emasculation, the flower buds are opened carefully before anthesis (First
opening of the flower) and the anthers are removed with the help of
forceps. Care should be taken so that the gynoecium of the flowers in
not damaged. In suction method, the petals are removed from the
flowers before anthesis, with the help of forceps. Then, a thin rubber or
glass tube attached to a suction hose is used to suck the anthers from
the flowers.
 PHYSIOLOGICAL METHODS OF
EMASCULATION
Here, the anthers are killed with the help of heat treatment, cold
treatment or alcohol treatment.
 GENETIC EMASCULATION
Genetic or cytoplasmic male sterility factors are introduced into the
female parents to make them sterile.
 BAGGING
The emasculated inflorescences of female plants are covered using
butter paper bags or cloth bags. However, in the case of cross-pollinated
crops, male plants may also be bagged if desired, so as to avoid pollen
mixture. The bags are removed 2-3 days after pollination.
 TAGGING
Emasculated flowers are tagged properly after bagging. Circular or
rectangular tags may be used. Details of the cross, date of emasculation,
date of pollination and the number of flowers emasculated must be
noted on the tag. Carbon pencil or permanent ink may be used for
tagging.
 POLLINATION
Mature, fertile and viable pollen grains are collected from the male
parent and dusted on the stigma of the female parent. Care should be
taken to see that the pollen grains are dusted at the optimum stage of
viability.
 HARVESTING F1 SEEDS
Crossed seeds are harvested carefully and stored to raise the F1
generation.
FURTHER HANDLING OF THE PLANTS
Further handling of the hybrids depends on the objective of the cross. In
the case of hybrid seed production, the F1 seeds are directly released to
farmers. In the case of combination breeding and transgressive breeding,
F2 is raised and the most appropriate solution program is used.
THE END
 Inbreeding and
Inbreeding Depression
INTRODUCTION
• Inbreeding is a form of mating system in sexual organism.
• It implies mating together of individual that are close to each
other by ancestral or pedigree relationship.
• When the individuals are closely related E. g Full sib was mating,
half sib mating.
• The highest degree of inbreeding is achieved by selfing.
• The chief effect of inbreeding is to increase homozygosity in
the progeny, which is proportionate to the degree of inbreeding.
• Cross – pollinated and asexually reproducing species are highly
heterozygous in nature.
INTRODUCTION CONT…
• These species show a severe reduction in fertility and vigour
due to inbreeding (inbreeding depression).
• It contrast to this hybridization between unrelated strains
leads to an increased vigour and fertility (hybrid vigour or
heterosis).
• These two aspects are of great significance in breeding of these
species.
• In fact heterosis and inbreeding depression may be considered
as the two opposite sides of the same coin.
Inbreeding Depression:
• It refers to decrease in fitness and vigour due to
inbreeding or it may be defined as the reduction or
loss in vigour and fertility as a result of inbreeding.
• The most revealing impact of inbreeding is the loss
of vigour and the physiological efficiency of an
organism characterised by reduction in size and
fecundity.
• For example selfing reduces heterozygosity, by a
factor ½ in each generation.
• In fact the dwgree of inbreeding in any generation
is equal to the degree of homozygosity in that
generation.
Inbreeding Depression Cont…
• Inbreeding depression results due to fixation of unfavourable
recessive genes in F2, while in heterosis the unfavourable
recessive genes of one line (parent) are covered by favourable
dominant genes of other parent.
• Man has recognised inbreeding depression for a long time.
• In many species marriage between closely related ancestries
have been prohibited.
• In hindu society perhaps presents the extreme example, where
marriages between individual related by ancestry is prohibited.
Effects of Inbreeding
 Inbreeding is due to a reduction in vigour and reproductive capacity that is
fertility.
 There is a general reduction in the size of various plant parts and in yield.
 The effects of inbreeding may be summarised as under.
a) Appearance of Lethal and Sublethal Alleles:
 Inbreeding to the appearance of lethal, sublethal and subvital characteristics.
 Such characteristics include chlorophyll deficiencies E.g Albino, chlorine rootles
seedlings , defects in flower structure etc. generally, plants carrying such
characteristics cannot maintained and are lost from the population.
b) Reduction in Vigour:
 There is a general reduction in the vigour of the population.
 Plants become shorter and weaker because of general reduction in the size of
various plant parts.
c) Reduction in Reproductive Ability:
 The reproductive ability of the population decreases rapidly.
 Many lines (plant progenies) reproduction poorly that they cannot be maintained.
Effects of Inbreeding cont…
d) Separation of the Population into Distinct Lines:
 The population rapidly separates into phenotypically distinct lines.
 This is because of an increase in homozygosity due to which there is random
fixation of various alleles of different lines.
 Therefore, the lines differ in their genotype and consequently in phenotype.
e) Increase in Homozygosity:
 Each line becomes increasingly homozygous following inbreeding.
 Consequently, the variation within a line decreases rapidly.
 Ultimately, after 7 to 8 generations of selfing, the lines become almost uniform.
 Since they approach complete homozygosity (> 99 percent homozygosity).
 The lines, which are almost homozygous due to continued inbreeding and are
maintained through close inbreeding, are known as inbred lines or inbreds.
i) Reduction in Yield:
 Inbreeding generally leads to a loss in yield.
 The inbred lines that is able to survive and be maintained yield much less than
the open pollinated varieties from which they were derived.
 In maize, the best – inbred lines yield about half as much as the open pollinated
varieties from which they were produced.
 In alfalfa and carrot, the reduction in yields is much greater, while in onions and
many cucurbits the reduction in yield is very small.
Degree of Inbreeding Depression
• The various plant species differ considerably in their responses
to inbreeding.
• Inbreeding depression may range from very high to very low or
may even be absent into the following four broad categories.

1) High inbreeding depression,

2) Moderate inbreeding depression,
3) Low inbreeding depression, and
4) Absence of inbreeding depression.
High Inbreeding Depression:
• Several plant species, Eg. alfalfa (M. sativa) carrot (D. carota) ,
hayfield, tarweed etc show very high inbreeding depression.

• A large proportion of plants produced by selfing shows lethal

characteristics and do not survive.

• The loss in vigour and fertility is so great that very few lines
can be maintained after 3 to 4 generation of inbreeding.

• The line shows greatly reduced yields, generally less than 25

percent of the yield of open – pollinated varieties.
Moderate Inbreeding Depression:
• Many crops species, such as maize, jowar, bajara etc. shows
moderate inbreeding depression.

• Many lethal and sublethal types appear in the selfed progeny,

but a substantial proportion of the population can be maintained
under self- pollination.

• There is appreciable reduction in fertility and many line

reproduce so poorly that they are lost.

• However, a large number of inbred lines can be obtained, which

yield upto 50 percent of the open- pollinated varieties.
Low Inbreeding Depression:
• Several crop plants, E. g onion (A. cepa), many cucurbits, rye (S.
cereale), sunflower (Hannus), hemp etc show only a small degree
of inbreeding depression.

• Only a small proportion of the plants show lethal or subvital

characteristics.

• The loss in vigour and fertility is small; rarely a line cannot be

maintained due to poor fertility.

• The reduction in yield due to inbreeding is small or absent.

• Some of the inbreds lines may yields as much as the open

pollinated varieties from which they were developed.
Lack of inbreeding Depression:
• The self- pollinated species do not show inbreeding depression
although they do not show heterosis.

• It is because their species reproduce by self – fertilization and

as a result, have developed homozygous balance.

• In cost of the cross- pollinated species exhibit heterozygous

balance.
THANK YOU
 CONTENT
1) Introduction
2) History of Distant Hybridization
3) Objective of Distant Hybridization
4) Features
5) Barriers associated with Distant Hybridization
6) Techniques for production of Distant Hybrid
7) Role of Wide crossing in crop improvement
8) Limitations of Distant Hybridization
9) Achievements
 HYBRIDIZATION
Hybridization:- Mating between two different strains.

Interspecific Hybridization:- When the individuals from two distinct species of the genus
are crosssed, it is known as interspecific hybridization. e.g. Oryza sativa x Oryza perennis.

Intergeneric Hybridization:- When the individuals being crossed belong to species from
two different genera it is referred as intergeneric hybridization. e.g. Triticum spp. X rye
(Secale cereale).

Distant Hybridization:- Hybridization between individuals from different species,

belonging to the same genus or to different genera is termed as distant hyridization and such
crosses are known as distant crosses or wide crosses.
 HISTORY
 Thomas Fairchild (1717):- He first authentic record of a
distant hybridization for crop improvement is the production
of a hybrid between Carnation (Dianthus caryophyllus) and
Sweet William (Dianthus barbatus).

Thomas Fairchild
Karpechenko (1928):- An interesting intergeneric hybrid,
Raphanobrassica, (R. sativus x B. oleracea) was produced.

Rimpu (1890):- Produce the first intergeneric hybrid

triticale which have greater potential than Raphanobrassica.

Karpechenko
 OBJECTIVES DISTANT HYBRIDIZATION

Wide crossing or distant hybridization has been used in the genetic

improvement of some crop plants. It is an effective means of transferring

desirable genes into cultivated plants from related species and genera.
INTER- SPECIFIC HYBRIDIZATION
 Ex. Nerica, an upland rice for Africa

 Oryza sativa (Asian upland rice) : non-shattering , resistant to lodging,

high yield potential

 Oryza glaberrima (African rice) : drought tolerant, disease resistant, weed

suppressing

 Nerica rice combines the best of both species.

African rice

Nerica rice

Asian rice
 INTERGENERIC CROSSES
 Triticale, a new cereal created in the lab.

 Triticale, a cross(intergeneric cross) between wheat and rye , was produced by

embryo rescue of the product of fertilization and a chemically induced doubling

of the chromosomes.

 Embryo rescue become necessary when fertile offspring is never produced by an

interspecific cross.
Wheat Rye

Triticale = Wheat x Rye

 Interspecific Combinations of Wheat and Rye
that produces hexaploid and octaploid triticale
 Main features of Interspecific or
Intergeneric hybridization
1. It is used when the desirable character is not found within the species of a crop.
2. It is an effective method of transferring desirable gene into cultivated plants from
their related cultivated or wild species.
3. It is more successful in vegetatively propagated species like sugarcane and
potato than in seed propagated species.
4. It gives rise to three types of crosses viz. a) fully fertile, b) Partially fertile and
c) Fully sterile in different crop species.
5. It leads to introgression which refer to transfer of some genes from one
species into genome of another species.
6.F1 hybrid between two genus are always sterile. The fertility has to be restored by
doubling of chromosome through colchicine treatment.
 Interspecific hybridization gives rise to
three types of crosses

a) Fully fertile
b) Partially fertile
c) Fully sterile in different crop species
 Fully fertile crosses
 Interspecific crosses are fully fertile between those species that have complete
chromosomal homology. Chromosome in such hybrids have normal pairing at
meiosis and result the F1 plants are fully fertile.

• Example- Cotton, Wheat, oat and Soybean.

 Cotton: There are four cultivated species of cotton Viz.,
• G. hirsutum and G. barbadense (Tetraploid,2n=52).
• G. arboreum and G. herbaceum (Diploid,2n=26).

 Triticum aestivum* Triticum compactum ( Where * F1 plants are Fully fertile)

 Partially fertile crosses
 Interspecific crosses are partially fertile between those species which differ in
chromosome number but have some chromosome in common.
• In such situation , F1 plants are partially fertile and partially sterile.
• Example- Wheat, Cotton, Tobacco.

Cotton: G. hirsutum (AADD) X G. thurberi (DD)

2n=52 | 2n=26
F1(ADD)
13II + 13I
Partially sterile or fertile, 2n=26.
 Fully sterile crosses
Interspecific crosses are fully sterile between those species which do not
have chromosomal homology. Such hybrids can be made self fertile by
doubling the chromosomes through colchicine treatment e.g. Tobacco,
wheat, cotton, brassica and Vigna species.

Cotton : G. arboreum (2n=26 ) X G. thurberi(2n=26)

|
F1 was sterile
Colchicine treatment- Fertile amphidiploid 2n=52.
 Problems associated with wide crosses
 The major problems associated with wide crossesare:
1) Cross Incompatibility.
2) Hybrid Inviability.
3) Hybrid Sterility.
4) Hybrid Breakdown.
 CROSS INCOMPATIBILITY
 This is the inability of the functional pollen grains of one species or genus to effect
fertilization in another species or genes.
 There are three main reasons of cross incompatibility viz.
i. Lack of pollen germination,
ii. Insufficient growth of pollen tube to reach ovule and
iii. Inability of male gamete to unite with the egg cell.
• These barriers are known as pre –fertilization barriers.

• This is overcome by employing different techniques like reciprocal crosses, bridge

crosses, using pollen mixtures, pistil manipulations, use of growth regulators etc.
 HYBRID INVIABILITY
 This refers to the Inviability of the hybrid zygote or embryo. In some cases, zygote
formation occurs, but further development of the zygote is arrested. In some other cases,
after the completion of the initial stages of development, the embryo gets aborted.
• The reasons for this are:
1. Unfavorable interactions between the chromosomes of the two species
2. Unfavorable interaction of the endosperm with the embryo.
3. Disharmony between cytoplasm and nuclear genes

• Reciprocal crosses, application of growth hormones and embryo rescue are the
techniques that can be used to overcome this problem.
 HYBRID STERILITY
 This refers to the inability of a hybrid to produce viable offspring. This is more
prominent in the case of intergeneric crosses. The major reason for hybrid sterility is the
lack of structural homology between the chromosomes of the two species.
 This may lead to meiotic abnormalities like chromosome scattering, chromosome
extension, lagging of chromosome in the anaphase, formation of anaphase bridge,
development of chromosome rings and chains, and irregular and unequal anaphase
separations.
 These irregularities may lead to aberrations in chromosome structure. Lack of
homology between chromosomes may also lead to incomplete pairing of
chromosomes.
 Sterility caused by structural differences between the chromosomes of two species can
be overcome by amphidiploidization using colchicine.
 HYBRID BREAKDOWN
 Hybrid breakdown is a major problem in interspecific crosses.

 When F1 hybrid plants of an interspecific crosses are vigorous and fertile but
there F2 progeny is weak and sterile it is known as hybrid breakdown.

 So hybrid breakdown hinders the progress of interspecific gene transfer.

 This may be due to the structural difference of chromosomes or problems

in gene combinations.
 EMBRYO RESCUE
 Embryo rescue :

 When embryos fails to develop due to endosperm degeneration, embryo

culture is used to recover hybrid plants; this is called hybrid rescue.

 e.g :- H. vulgare x Secale cereale.

 Embryo rescue generally used to overcome endosperm degeneration.
 Embryo culture.
A. Proembryo dissected 3 to 5 days after pollination
B. Proembryo culture on solid agar media
C. Plantlet developing from embryo
D. Plantlet transplanted into soil.
 Embryo rescue in barley

Hordeum vulgare (Barley) X Hordeum bulbosum (Wild relative)

2n = 14 2n = 2X = 14
Embryo Rescue

Haploid Barley 2n = X= 7 (H. bulbosum)

chromosomes eliminated

This technique was once more efficient than microspore culture in creating
haploid barley
 Wide crossing of wheat and rye requires embryo
rescue and chemical treatments to double the no. of
chromosomes triticale
Triticum durum (4X) x Secale cereale (2X)
AABB RR

ABR F1(3x): Embryo Rescue

Chromosome Doubling Hexaploid
Triticale
(6x) AABBRR
 Limitations of embryo rescue
 High cost of obtaining new plantlets .
 Sometimes deleterious mutations may be induced during the in vitro
phase.
 A sophisticated tissue culture laboratory and a dependable greenhouse
are essential for success.
 Specialized skill for carrying out the various operations are required.
 Differences between Interspecific and
Intergeneric Hybridization
Particular Interspecific Hybridization Intergeneric Hybridization

Parents involved Involves two different species of the same Involves two different genera of
gene. the same family.

Fertility Such hybrids vary from completely fertile Hybrids are always sterile.
to completely sterile

Seed Setting More than intergeneric crosses. Less than interspecific crosses.

Use in crop improvement More than intergeneric crosses. Less than interspecific crosses.

Release of Hybrid Variety Possible in some crops. Not Possible.

Evolution of new crops Not possible, but evolution of new species Sometime possible,
is sometimes possible. e.g., Triticale.
 Techniques to make wide crosses successful
 SELECTION OF PLANTS
 The most compatible parents available should be selected for the crosses.
 RECIPROCAL CROSSES
 Reciprocal cross may be attempted when one parental combination fails.
 e.g. Mung x udid- cross compatible and Udid x mung-cross incompatible

 MANIPULATION OF PLOIDY
 Diploidization of solitary genomes to make them paired will be helpful to make the cross fertile.

 BRIDGE CROSSES
 When two parents are incompatible, a third parent that is compatible with both the parents can be
used for bridge crosses and thus it becomes possible to perform cross between the original parents.
 e.g. Tobacco
 Nicotiana repanda x N.tabaccum– cross incompatible
 Nicotiana repanda x N.sylvestris- cross compatible
 Nicotiana syivestris x N.tabaccum- cross compatible Continue.......
 USE OF POLLEN MIXTURE
 Unfavorable interaction between pollen and pistil in the case of wide crosses can be
overcome to some extent by using pollen mixture.
 MANIPULATION OF PISTIL
 Decapitation of the style will sometimes prove helpful in overcoming incompatibility.
 USE OF GROWTH REGULATORS
 Pollen tube growth can be accelerated by using growth hormones like IAA, NAA, 2,4-D
and Gibberellic acid.
 PROTOPLAST FUSION
 When fusion of gametes fails, protoplast fusion of somatic cells can be attempted.
 EMBRYO RESCUE
 Hybrid zygotes formed by wide crosses may fail to grow in a number of cases. The zygotes
are taken out and grown in in vitro medium to overcome this problem.
 Role Of Wide Crosses In Crop Improvement
 Wide crosses are generally used to improve crop varieties for disease resistance, pest resistance, stress
resistance, quality, adaptation, yield etc. These crosses can even be used to develop new crop species.
Techniques like alien addition and alien substitution may also be effective.
(A)Disease and insect resistance
Crop Character transferred Species transferred from Species transferred to
Cotton Jassid resistance G. tomentosum G. hirsutum
Blackarm resistance G. arboreum G. barbadense
Okra Resistance to YMV Abelmoschus manihot A. esculenta
Groundnut Resistant to leaf chewing Arachis monticola A. hypogea
insect
Wheat Rust resistance Agropyron T. aestivum
Tobacco Resistant to mosaic virus N. repanda N. Tabaccum
Sugarcane Sereh disease resistant Saccharum spontaneum S. Officinarum
Potato Resistant to late blight and Solanum denissum S. tuberosum
leaf roll.
Continue.......
 (B) Improvement in quality
Crop Charactertransferred Species transferred from Species transferred to
Cotton Fibre length G. thurberi & G.hirsutum
Male sterility G. raimondii
G. harkenssii G.hirsutum
Potato Starch content Frost resistance Wild species Cultivated Spp.
Solanum acaule S. tuberosum
Tomato Carotenoid content Lycopersicon L.esculentum
Wild Spp.
Palm Oil quality Wild Spp. Cultivated Spp.
Rice, Oat & Rye Protein quality Wild Spp. Cultivated Spp.
Tobacco Leaf quality Nicotiana debneyi N. tabacum
Oat High oil content Avena sterilis A.sativa

(C) Improvement in yield: This also been achieved through the use of wild Spp. in some crops e.g. Oat,
Mung bean, Groundnut, Potato, Tobacco.
 ALIEN ADDITION LINES
 These lines carries one chromosome pair from a different species in addition to the
normal somatic chromosome complement of the parent species.
 When only one chromosome (not a pair of chromosome) from another species is
present, it is known as alien addition monosome.
 Alien addition have also been done in rice, sugar beet, cotton, brassicas.
 The main purpose of alien addition is the transfer of disease resistance from related wild
species. e.g. Transfer of mosaic resistance from Nicotiana glutinosa to N. tabacum.
 The alien addition lines have been developed in case of wheat, oats, tobacco and
several other species.
 Alien addition lines are of little agricultural importance since the alien chromosome
generally carries many undesirable genes. e.g. Reduced growth and short, broad leaves
in addition to mosaic resistance.
 ALIEN SUBSTITUTION LINES
 This line has one chromosome pair from a different species in place of the chromosome pair
of the recipient species.

 When a single chromosome (not a pair) from different species in place of a single
chromosome of the recipient species, known as alien-substitution monosome.
 Alien –substitution line have been developed in wheat, cotton, tobacco, oats, etc.

 In case of tobacco, mosaic resistance gene N was transferred from the N. glutinosa to N.
tabacum line had 23 pairs of N. tabacum chromosomes and one pair (chromosome H) of N.
glutinosa chromosomes.

 The alien substitution show more undesirable effects than alien additions and as a
consequence are of no direct use in agriculture.
 Limitations of Distant hybridization
1) Incompatible crosses
2) F1 sterility
3) Problems in creating new species
4) Lack of homoeology between chromosome of the parental species
5) Undesirable linkages
6) Problems in the transfer of recessive oligo genes and quantitative traits
7) Lack of flowering in F1
8) Problems in using improved varieties in distant hybridization
9) Dormancy
 Achievements
 Hybrid varieties:-
 Upland cotton – MCU-2, MCU-5, Khandwa1, Khandwa2 etc are derivatives of
interspecific hybridization.
 Hybrid between Pearl millet x Napier grass - Hybrid Napier which is very popular for
its high fodder yield and fodder quality e.g. Jaywant and Yashwant

 Interspecific hybrids in cotton- Varlaxmi, Savitri, DCH-32, NHB-12, DH- 7, DH-9

etc.

 Prabhani Kranti variety of bhindi.

Unit 4
 POLYPLOIDY HISTORY

Eduard Strasburger
▪ Founder of modern cytology
▪ First coined the term
‘polyploid’

Hugo DeVries (1901)

published “Mutation Theory”
Polyploid mutants in
Evening primerose played an
important role in his work
 POLYPLOIDY HISTORY

 De Vries did large-scale greenhouse experiments to

find mutants.
 The ‘gigas’ mutant was important as it turned out to
be a spontaneously derived tetraploid (4n=28 is typical
for Oenothera)
 POLYPLOIDY HISTORY

O. Winge (1917)
 First classic paper on polyploidy

 Distinguished two types of polyploids

 Autopolyploids - genomes identical

 Allopolyploids - genomes from two or more spp.

Kihara and Ono (1926) actually coined the terms

‘autopolyploidy’ and ‘allopolyploidy’
Terminology related to ploidy

The chromosome number is of three types as

 Basic: (x): Gametic chromosome no. of the true diploid
species
 Haploid: (n): Half of the somatic chromosome no. of
the species
 Diploid: (2n): Somatic chromosome no. of the species
 CLASSIFICATION OF HETEROPLOIDY

HETEROPLOIDY

ANEUPLOIDY EUPLOIDY

 Nullisomic (2n-2)  Haploidy (n)

 Monosoic (2n-1)  Polyploidy
 Trisomic (2n+1) (3n,4n,5n,6n & so on)
 Tetrasomic (2n+2)
 Haploid Vs Monoploid
 The monoploid chromosome set is the basic set of
chromosomes multiplied in a ploidy series (x)

 The haploid chromosome set is the set chromosomes

present in a gamete (n)

 Monoploid is always haploid and haploid may or may

not monoploid.

 In diploids, monoploids and haploid are equivalent

Pearlmillet (diploid) 2n=14=2x
n=x
Wheat (hexaploid): 2n=42=6x n>x
 POLYPLOIDY
Polyploidy occurs when all the chromosomes are
present in three or more copies.
Polyploidy and Plant Evolution

Polyploids, like the one on the

Most Crop Species are Polyploid left, are larger than their diploid
progenitors (strawberry on
right).
Nearly 35 per cent of flowering plants arose through polyploidization.
 Polyploid complex
Sugarcane

Bread Wheat

Maize

Banana

Pearl millet
 Polyploidy
Definition: Presence of more than two identical or distinct
genomes in an individual is known as polyploid and
condition of being polyploid is polyploidy.
Types of polyploidy
1. Autopolyploidy
a) Autotriploid:-
b) Autotetraploid:-
c) Autopentaploid:-
2. Allopolyploidy
a) Allotetraploid:-
b) Allohexaploid :-
c) Allooctaploid :-
1. Autopolyploidy: If all the genomes present in an individual are
identical, it is called as autopolyploidy.

a) Autotriploid:- Three copies of the same genome (3x)

b) Autotetraploid:- Four copies of the same genome (4x)
c) Autopentaploid:- Five copies of the same genome (5x)

2. Allopolyploidy: In the case of allopolyploids, two or more

distinct genomes are present.

a) Allotetraploid:- Two distinct genome, each genome has two copies

b) Allohexaploid :- Three distinct genome, each genome has two copies
c) Allooctaploid :- Four distinct genome, each genome has two copies
Autopolyploid
Origin and production of doubled chromosome numbers:

1. Spontaneous
2. Production of adventitious buds
3. Treatment with physical agents
4. Regeneration in vitro
5. Colchicine treatment
Autopolyploid
Origin and production of doubled chromosome numbers:

1. Spontaneous :
Chromosome doubling occurs occasionally in somatic tissues and
unreduced gametes are produced in low frequencies.
2. Production of adventitious buds : Decapitation (is the complete
separation of the head from the body) in some plants leads to callus
development at the cut ends of the stem.

Such a callus has some polyploid cells and some of the shoot buds
regenerated from the callus may be polyploid.

In solanaceae 6-36% of adventitious buds are tetraploid.

3. Treatment with physical agents: Heat or cold treatment,
centrifugation, x-ray or gamma ray irradiation may produce
polyplods.

Exposing the plants or ears of maize to a temperature of 38-45 oC

at the time of the first division of zygote produce 2-5 % tetraploid
progeny.
4. Regeneration in vitro: polyploidy is a common feature of the cells
cultured in-vitro. Some of plants regenerated from callus and
suspension culture may be polyploids.

5. Colchicine treatment: Colchicine treatment is the most effective and

the most widely used treatment for chromosome doubling.
A. Seedless watermelons are produced by crossing tetraploid (4x,
used as female) and diploid
• For good seed setting pollination is essential.
• For this purpose diploid lines are planted in the ratio 1 diploid : 5
tetraploid plants.
There are several problem viz.
• irregular fruit shape,
• a tendency toward hollowness of fruits,
• production of empty seeds and the labour involved in trioploid seed
production.
B. Triploid sugarbeets : Among root crops triploid sugar beets
apparently represent the optimum level of polyploidy because 3n
plants have longer roots than diploid and also yield more sugar per
unit area.

2. Tetraploid rye : the advantage of tetraploid over its diploid

counterpart are large kernel size, superior ability to emerge under
adverse condition and higher protein content.
• tetraploid rye varieties have been released for cultivation.
• Barseem var. pusa giant
 Limitations of autoployploidy:
1. Larger size autoployploiods generally contain more water and
produce less dry matter content than diploids.
2. High sterility with poor seed setting is observed.
3. Due to complex segregation, progress through selection is slow.
4. Monoploids and triploids cannot be maintained except through clonal
propagation.
5. New polyploids are always characterized by a few or more
undesirable features. E.g irregular fruit size in watermelons
6. Effects of autopolyploidy cannot be predicted.
 Allopolyploids
• ORIGIN AND PRODUCTION OF ALLOPOLYPLOIDS
• The present-day allopolyploids were most likely
produced by chromosome doubling in F1 hybrids
between two distinct species belonging to same genus or
to different genera.
• Chromosome doubling might have occurred in somatic
tissues due to an irregular mitotic cell division leading to
formation of allopolyploids. Alternatively, irregular
meiosis may lead to production of allopolyploid progeny.
 Morphological and cytological features of allopolyploids

1. Allopolyploids combine the morphological and physiological

characteristics of the parent species but it is very difficult to predict
the precise combination of characters that would appear in the new
species e.g., Raphanobrassica. But Triticale has combined
favourable features, hardiness of rye & yielding ability of
wheat.

2. Many allopolyploids are apomictic. Ex : Solanum sp.

3. The chromosome pairing in the new species depends upon the
similarities between the chromosomes of the parental species.
Chromosomes with such similarities are known as homoeologous
chromosomes.

After chromosome doubling, the allopolyploid would have two

homelegous chromosomes for each chromosome present in the F1
hybrid, comparable to the diploid species. Such allopolyploid is
referred as amphidiploid or Allotetraploid.

4. Fertility of Allopolyploids can be improved by hybridization and

selection.
A) Examples of autopolyploids
1) Autotriploids Banana : Seedless, larger fruits than
diploid
Apples : Propogate asexually by budding
or grafting
Sugerbeet : Higher sugar content than
diploid resistant to moulds
Watermelon : Melons are seedless or
rudimentary seeds.
2) Autotetraploids Rice : Larger seed and Higher protein
than diploid
Grape : Larger fruits, fewer seed per
fruit
Alfalfa : Better in yield
B) Examples of Allopolyploids
Crops and parental Number Resulting allopolyploid
(2n) and Species
Radish - Cabbage
Raphanus sativus 18 Raphanobrassica
Brassica olereacea 18 2n=36
Wheat
Triticum monococcum 14 A Triticum turgidum
Unknown species 14 B (2n=28) AABB
T. turgidum 28 AB Triticum aestivum
T. tauschi 14 D (2n=42) AABBDD
Tobacco
Nicotiana sylvestris 24 Nicotiana tomentosa
Nicotiana tomentosa 24 (2n=48)
N. paniculata 24 N. rustica
N. undulata 24 (2n=48)
Cotton
Gossypium herbaceum Var. 26 A G. hirsutum
abricanum
G. raimondii 26 D (2n=52) AADD
Brassica
Brassica Nigra 16 B B. corinata
Oleracea 18 C (2n=34)BBCC
B. compesiris 20 A B. napus
B. oleracea 18 C (2n=38) AACC
B. compestris 20 A B. junecea
B. nigra 16 B (2n=36) AABB
Triticale
Triticum turgidum 28 Triticale
Secale cereale 14 (2n=42)
Role of Allopolyploids in evolution
Evolution of Bread Wheat (Triticum aestivum)
EVOLUTION OF BRASSICA SPECIES

Nagaharu’s Brassica U tiangle

Evolution of Nicotiana tabacum
EVOLUTION OF COTTON
IMPACT OF EVOLUTION OF COTTON
Application of allopolyploids in crop improvement :

1. Utilization as a Bridging species :

Amphidiploids serve as a bridge in transfer of characters from one species

to a related species, generally from a wild species to cultivated species.

Example: use of synthetic Nicotiana digluta for the transfer of resistance

to tobacco mosaic virus from N. sylvestris to N.tabacum.
2. Creation of New crop species. Ex : Triticales,
Raphanobrassica

Triticales:- Triticum turgidum x secale cereale.

Triticales:- Triticum aestivum x secale cereale.
Raphanobrassica:- radish x cabbage
Intergeneric combinations of wheat and rye that produces
hexaploid and octaploid triticale
 ARTIFICAL ALLOPOLYPLOIDS
• Raphanobrassica : Developed by Russian scientist “Karpechenko” in
1928.
• It was developed by a cross between raddish and cabbage .
3. Widening the genetic base of existing Allopolyploids :

This is being done by crossing B.campestris (n=10, AA) with B.oleracea

(n=9, CC), the parental diploid species, to produce the amphidiploid
B.napus (n=19, AACC).
 Limitations of Allopolyploidy

1. The effects of allopolyploidy cannot be predicted. The allopolyploids

have some features from both the parental species, but these
features may be the undesirable ones.
e.g., Raphanobrassica

2. Newly synthesized allopolyploids have many defects,

e.g., low fertility
3. The synthetic allopolyploids have to be improved through
extensive breeding at the polyploidy level. This involves
considerable time, labour and other resources.

4. Only a small proportion of allopolyploids are promising; a vast

majority of them are valueless for agricultural purposes.
 Thank you

Happy Learnings !!!

1
1
UNIT - 4 Mutation Breeding
B
Dr. Chetariya Chana P.
Assistant Professor
Ph.D. (Agri.) Genetics and Plant Breeding
School of Agriculture
Lovely Professional University
Phagwara, Punjab
CONTENT
✓ History
✓ Terminology
✓ Characteristics of mutation
✓ Classification of mutation
✓ Molecular basis of mutation
✓ Classification based on origin
✓ Procedure for irradiation
✓ Applications of mutation breeding
✓ Advantages and Limitations
✓ Role in crop evolution and
✓ Achivements
HISTORY
➢ The mutation was first discovered by Wright,
1791 in Male lamb (sheep)
➢ The term mutation was introduced by Hugo de
Vries in 1900.

4
➢Mutation:
•It refers to sudden heritable changes in the characters of
organism.
➢Mutagen:
•The agents capable of inducing mutations are known as
mutagens.
➢Mutation breeding:
•Utilization of induced mutations for crop improvement is
known as mutation breeding.

5
1. Mutation is originated spontaneously or induced
artificially by radiations or chemicals.
2. Mutations are random events.
3. Mutations are mostly recurrent.
4. Spontaneous mutations occur at very low frequencies
(10-4 to 10-7) in nature.
5. The rate of induced mutations is markedly affected by
the environmental conditions.
6. Most mutant alleles are plelotropic in nature.

6
 CHARACTERISTICS OF MUTATION
7. Mutations occur in both forward and reverse directions.
8. The mutator genes and antimutator genes effect the rate
of some other genes of the genome.
9. It occurs in any tissue/cell and at any developmental
stage of organism.
10. Generally, mutant alleles are recessive to their wild type
or normal alleles.
11. Fine structure analyses of genes have shown that
mutations producing the same phenotypic effect may be
located at different sites within the gene.

7
PHYSICAL MUATGEN
Chemical mutagen:-
 X rays were first discovered by Roentgen in 1895. They
are sparsely ionizing and highly penetrating. They are
generated in X rays machines. X rays can break
chromosomes and produce all types of mutations in
nucleotides, viz. addition, deletion, inversion,
transposition, transitions and transversions.
 X rays were first used by Muller in 1927 for induction of
mutations in Drosophila.
 In plants, Stadler in 1928 first used X rays for induction of
mutations in Barley.

8
CHEMICAL MUTAGENS

9
10
CLASSIFICATION OF MUTATION
(1) Based on source (5) Based on site
Spontaneous mutation Nuclear mutation
Induced mutation Cytoplasmic mutation
(2) Based on Direction
Forward mutation (6) Based on character
Reverse mutation Morphological
(3) Based on survival Biochemical
Lethal (7) Based on visibility
Sub lethal Macro – mutation
Sub vital
Micro – mutation
Vital
(4) Based on tissue (8) Based on dominance relationship
Somatic mutation Dominant mutation
Germinal mutation Recessive mutation
Codominant mutation
Incompletely dominant mutation
11
 CLASSIFICATION OF MUTATION
(9) Based on Intensity of character expression
Amorphic mutations
Hypo-morphic mutation
Hyper-morphic mutation
(10) Based on Expression of neighbouring gene
Polar mutations
Non-polar mutations
(11) Based on amino acid replacement in the polypeptide
Mis sense
Nonsense
Frame-shift

12
❖ Mutations produced by changes in the base sequence of genes are known as
gene or point mutations some mutations may be produced by changes in
chromosome structure or even in chromosome number they are termed as
chromosomal mutation.

➢ There are three types of mutations based on genetic basis of heritable change :

➢ 1. Gene mutations : These are produced by change in the base sequence of

genes.
The change may be due to base substitutions, deletion or addition.

➢ 2. Chromosomal mutation : These arise due to change in chromosome number

that may leads to polyploidy or aneuploidy or change in chromosome structure
that result in deletions duplication, inversion and translocation.

➢ 3. Cytoplasmic or plasmagene mutation : These are due to change in the base

sequence of plasma genes. The plasma genes are present in mitochondria or
chloroplast. Here the mutant character occurrs in buds or somatic tissues which
are used for propagation in clonal crops.
13
MOLECULAR BASIS OF GENE MUTATION

 Mutation in a character would generally be produced by a

change in the amino acid sequence of the protein
determining the expression of that trait.

 This change would be due to a change in the base

sequence of the DNA molecule coding for the protein, which
may result from
1. Base substitution
2. Base addition or deletion
3. Transposition

14
 MOLECULAR BASIS OF GENE
MUTATION
➢ Base substitution :-
When one base in a DNA molecule is replaced by another, it
is called base substitution. It is of two types
i. Transition
ii. Transversion
Transversion

A T

Transition
Transition

G Transversion C
15
 Classification of mutations :
❑ Based on origin, the mutations are classified as spontaneous and induced mutations.

1. Spontaneous mutations : Mutations occur in natural populations at a low rate

(10-6) but different genes may show different mutation rates. Here the different genes
show different mutation rate. For example : in maize R-locus mutates at the frequency
of 4.92 x 10-4 i.e. (1 in 20000 population), when as Su locus at 2.4 x 10-6 ( 1 in 25
lakhs). The Wx locus considered to be highly stable. The difference in mutation rate
may be due to a) Genetic back ground i.e. presence of mutator genes b) Genes them
selves c) Environment.

2. Induced mutation : Mutations may be artificially induced by treatment with

certain physical or chemical agents. Available evidence indicates that induced mutation
rarely produce new alleles they produce alleles which are already known to occur
spontaneously. Induced mutations are comparable to spontaneous mutations in their
effects and in the variability they produce. Induced mutation occur at a relatively
higher frequency so that it is practical to work with them.

16
3. Based on magnitude of phenotypic effects mutation
as classified as
Macro mutations : Oligogenic Mutation – Large
phenotypic effect and recognizable on individual plant
basis and can be seen easily in M2 generations – Eg. Ancon
breed in sheep, pod maize to cob maize.

Micro mutations : Polygenic mutations – Small

phenotypic effect which can not be recognized on
individual plant basis but can be recognize only in a group
of plants. Selection should be done in M3 or later
generations.

17
Procedure for irradiation : The plant material may be treated in any of the
following source. 1. Seeds, 2. Seedlings, 3. Flowers, 4. Cuttings

1. Seeds : Seeds are used after soaking to get greater frequency of induced
mutations than air dried.

2. Seedlings : At any stage of life cycle can be subjected to radiation but usually
seedlings neither too young nor too old are irradiated due to their convenience in
handling in pots transportation from nursery easily.

3. Flowers : Meiotic cells have been found more sensitive than the mitotic cells
and therefore plants are irradiated in the flowering stage in order to affect the
developing gametes.

4. Cuttings : In case of fruit tree when they are propagated by clones – the
desirable cuttings are exposed to irradiation.

18
 Selection of the variety for mutagen treatment
❖ The variety selected for mutagenesis should be the best available in the crop.
❖ Dose of the Mutagen
An optimum dose of the mutagen should be used. An optimum dose is the one
which produces the maximum frequency of mutations and causes the minimum
killing. Many workers feel that a dose close to LD50 should be optimum. LD50
is that dose of a mutagen, which would kill 50% of the treated individuals.

Mutation Breeding for oligogenic traits

The handling procedure described here is based on the selection for a recessive mutant
allele of an oligogene.
1. M1. Several hundred seeds are treated with a mutagen and are speceplanted. In
general, the number of treated seeds is so adjusted as to give rise to -500 fertile M1
plants at the harvest. Care should be taken to avoid out crossing; this can be achieved
either by planting the M1 population in isolation or by bagging the inflorescenes of M1
plants or even the whole M1 plants. M1 plants will be chimeras for the mutations
present in heterozygous state. About 20 to 25 seeds from each M1 spike are harvested
separately to raise the M2 progeny rows.
19
2. M2. About 2,000 progeny rows are grown. Careful and regular
observations are made on the M2 rows. But only distinct mutations are detected
in M2 because the observations are based on single plants. All the plants in M2
rows suspected of containing new mutations are harvested separately to raise
individual plant progenies in M3. if the mutant is distinct, it is selected for
multiplication and testing. However, most of the mutations will be useless for
crop improvement. Only 1-3 per cent of M2 rows may be expected to have
beneficial mutations.
3. M3. Progeny rows from individual selected plants are grown in M3. Poor
and inferior mutant rows are eliminated. If the mutant progenies are
homogeneous, two or more M3 progenies containing the same mutation may be
bulked. Mutant M3 rows are harvested in bulk for a preliminary yield trial in M4.
4. M4. A preliminary yield trial is conducted with a suitable check, and
promising mutant lines are selected for replicated multilocation trials.
5. M5-M7. Replicated multilocation yield trials are conducted. The out-
standing line may be released as a new variety. The low yielding mutant lines,
however, should be retained for use in hybridization programmes.

20
Mutation breeding for polygenic traits :Mutagenesis does produce genetic
variation in polygenic traits; this variation is usually as much as 50% of that generated
in F2 generation, but sometimes it may be as much as or even greater than the latter.
1. M1 and M2. M1 and M2 are grown in the same way as in the case of oligogenic
traits. In M2, vigorous, fertile and normal looking plants that do not exhibit a mutant
120 phenotype are selected and their seeds are harvested separately to raise individual
plant progeny rows in M3.

2. M3. Progeny rows from individual selected plants are grown. Careful
observations are made on M3 rows for small deviations in phenotype from the parent
variety. Inferior rows are discarded. Few rows may be homogeneous and would be
harvested in bulk. Selection in done in M3 rows showing segregation; a majority of M3
rows would show segregation. Intensive and careful evaluation of a large number of M3
progeny rows allows identification of mutants with altered quantitative traits, e. g.,
partial or horizontal disease resistance. Such mutants occur in high frequencies that
approach 1% or even high, so that their isolation becomes quite cost effective.

21
3. M4. Bulked seed from homogeneous M3 rows may be planted in a
preliminary yield trial with a suitable check; superior progenies are selected for
replicated multilocation yield trials. Individual plant progenies from M3 are critically
observed. Progenies showing segregation may be subjected to selection only if they
are promising. Super ior homogeneous progenies are harvested in bulk for
preliminary yield tests in M5.

4. M5-M8. Preliminary yield trials and / or multi-location trials are conducted

depending upon the stage when the progenies become homogeneous. Outstanding
progenies may be released as new varieties.

22
 Applications of Mutation Breeding
Mutation breeding has been used for improving both oligogenic as well as polygenic
characters. Mutagenesis has been used to improve morphological and physiological
characters including yielding ability. Various applications of mutation breeding are :
1. Induction of desirable mutant alleles which may not be available in the
germplasm.
2. It is useful in improving specific characteristics of a well adapted high yielding
variety.
3. Mutagenesis has been successfully used to improve various quantitative character
including yield.
4. F1 hybrids from intervarietal crosses may be treated with mutagens in order to
increase genetic variability by inducing mutation and to facilitate recombination
of linked genes.
5. Irradiation of interspecific (distant) hybrids has been done to produce
translocations.

23
Advantages :
1. Mutation create inexhaustible variation.

2. When no improvement is possible this method has to be adopted.

Limitations :
1. Frequency of desirable mutations is very low about 0.1 percent. To detect the
desirable one in M2 considerable time, labour & other resources are to be employed.

2. To screen large population, efficient quick and unexpensive selection techniques are
needed.

3. Desirable mutations may be associated with undesirable side effects due to other
mutations thus extending the mutation breeding programme.

4. Detection of recessive mutations in polyploids and clones is difficult and larger doses
of mutagen have to be applied and larger populations are to be grown.

24
 Achievements :
a) Natural mutants :
Rice : GFB 24 – arose as a mutant from Konamani variety Dee – Gee – Woo –
Gen – Arose as a mutant from rice in China
MTU 20 – arose as a mutant from MTU-3
Sorghum Co. 18 – arose as a mutant from Co. 2
Cotton : DB 3-12 from G. heroaccum variety Western 1

b) Induced mutants :
Rice : Jagannath-gamma ray induced mutant from T.141
Wheat : Sarbati Sonora Gamma radiation from Sonora 64
NP 836 mutants, through irradiation from NP 709
Cotton : Indore 2 Induced from Malwa upland 4
MLU 7 gamma ray induced mutant from culture 1143 EE
MLU 10 gamma ray induced mutant from MLU 4
Mustard : Primax whicte (1950)
Summer Pope seed Regina I (1953)
Sugarcane : Co.8152 gamma ray induced mutant from Co. 527
Groundnut : NC 4
Cas tor : Aruna (NPH1) – Fast neutrons induced mutant from HC 6 25
Table:- Some important varieties developed in India through various mutagens in
different crops

Crop species Varieties developed and

mutagens used
X – rays Gamma rays rothers

Wheat NP 836, JRC 7447 Pusa lerma, Sharbati

sonora -
Rice Biraj, CNM 20, LNM 25, AU I, AU 4, K 84 hybrid PL 56 (EMS) IIT
Jagananth mutant, mohan 60(EMS) indira
padmini, DL 253 (EMS)
Cotton Indore 2, MA 9, MCU 7 MCU 10, Rasmi Pusa
Ageti -
chickpea Pusa 408, Pusa 413 Kiran (N)
- Pusa 417
Groundnut TG 3, TG 4, TG 17, Vikram BP1, BP2 Co 2 (EMS)

Papaya Pusa Nanha

- -
26
27
27
Unit 5
Breeding for disease and insects
resistance

Dr. Chetariya Chana P.

Assistant Professor,
Dept. of GPB,
School of Agriculture,
L.P.U., Punjab.
 Contents

1. Mechanism of insects and disease resistance

2. Methods of breeding used for disease and insects resistance

3. Genetics of disease and insects resistance

 STRESS

Stress refers to adverse condition for the crop growth and production
imposed by either environmental factors or biological factors.
 Types of stress

1. Biotic
2. Abiotic

Biotic: The stress that is caused by biological agents, such as disease,

insects and parasitic weed.

Abiotic: When the stress is caused by environmental factors or non

biological factors, such as moisture, temp., light etc.
Mechanisms of Insect Resistance
 Mechanisms of Insect Resistance

There are four mechanisms of insect resistance

1. Non-preference
2. Antibiosis
3. Tolerance
4. Avoidance or escape

The first three mechanisms were given by Painter (1951).

A resistant variety may have one, two or more of these
mechanisms.
1. Non – preference

Non preference refers to various features of host plant that make

the host undesirable or unattractive to insects for food, shelter, or
reproduction. This type of insect resistance is also known as non
acceptance and antixenosis.

Various plant character which are associated with non preference include
colour, light penetration, hairiness, leaf angle, odour and taste.

For example, in cotton red plant body, smooth leaves, leaf, open canopy,
thickness and long pedicel are examples of non preference to bollworms,
and hairiness of leaf and stem is non preference for jassids.
 Non Preference Mechanism of Insect Resistance in Some Crop
Plants

Host Crop Insect Pest Non-preference Preference

Wheat Strem Sawfly Solid stem Hollow stem
Rice i)Rice stem borer Lignified stem Nonlignified stem
ii) Brown plant hopper Low asparagines High asparagines
Red Pericarp White Pericarp
Purple stigma Yellow stigma
Maize Corn earworm Toughness of husk Soft husk
Corn leaf aphid High DIMBOA Low DIMBOA
Cotton Bollworms Smooth leaves Hairy leaves
Nectariless Nectaried
Long Pedicel Short Pedicel
Okra leaf Normal leaf
Thick boll rind Soft boll rind
Boll weevil Frego bract Normal Bract
Red Plant body Green plant body
Jassids Hairiness of leaves Smoothness
 Host Crop Insect Pest Non-preference Preference
Potato leaf
Soybean hopper Hairiness Smoothness
Pea Pea aphid Yellow green leaves Blue grren leaves
Leaves with high ligth Leaves with low light
Cabbage Cabbage aphid reflection refelction
Sugarbeet Aphid Low free sugar High free sugar
Brassica Cabbage aphid Low sinigrin High sinigrin
Sweet clover Flying weevil Low coumairn High coumairn
2. Antibiosis

Antibiosis refers to the adverse effect of host plant on the

development and reproduction of insect pests which feed on
resistant plant.

In some cases, antibiosis may lead even to death of an insect.

In cotton, antibiosis is related with high level of gossypol, tannins and

silica contents, antibiosis may involve morphological, physiological and
biochemical features of the host plant.
Antibiosis Mechanism of Insect Resistance in Some Crop Plants

Host Crop Insect Pest Cause of Antibiotics

Wheat Hessian fly H1 to H3 genes
Stem sawfly Solid stem
Green bugs High benzyl alcohol
Barley Cereal leaf beetle Waxiness of leaves
Green bugs High benzyl alcohol
Rice Rice stem borer High silica content
European corn borer High DIMBOA in leaves
Maize stem borer High Aspartic acid
Low nitrogen and sugar
Cotton Bollworms High gossypol
High gossypol
High tanins
High silica content
Host Crop Insect Pest Cause of Antibiotics
Sugarbeet Aphid Low free sugar
Spotted aphid and pea
Alfalfa aphid High saponin
Brassica Cabbage aphid Low sinigrin
Potato Aphid Gummy Trichome exudates
Exudates of glandular leaf
Tobacco Mites hairs
Exudates of secondary
Medicago Alfalfa weevil trichomes on leaves.
3. Tolerance

Tolerance refers to the ability of a variety to produce greater

yield than susceptible variety at the same level of insect
attack.

In other words, a tolerant variety will give higher yield than

susceptible one despite the insect attack.
4. Avoidance or Escape

Avoidance refers to escape of a variety from insect attack

either due to earliness or its cultivation in the season where
insect population is very low.

For example, early maturing cotton varieties escape pink

bollworm infestation which occurs late in the season.
 Mechanism of Disease Resistance

The various mechanisms of disease resistance are as follows:

1) Mechanical
2) Hypersensitivity
3) Nutritional
4) Systemic Acquired Resistance (SAR)
1) Mechanical

Certain mechanical and or anatomical features of the host may prevent

infection.
For example, closed flowering habit of wheat and barley prevents
infection by the spores of ovary infecting fungi.

2) Hypersensitivity

In a large number of cases, immune reaction is due to the hypersensitive

reaction of the host.
This mechanism is found in case of biotrophic organism or obligate
parasites.
The HR is one kind of programmed cell death (PCD)
associated with the death of a small number of cells at and
around the site of infection.
1. The hypersensitive response (HR) is a complex, early
defense response that causes necrosis and cell death
to restrict the growth of a pathogen.

2. The HR is characterized by the rapid death of cells in

the local region surrounding an infection.

3. The HR serves to restrict the growth and spread of

pathogens to other parts of the plant.
 3) Nutritional

The reduction is growth and in spore production is generally

supposed to be due to an unfavourable physiological conditions
within the host.

Most likely, a resistant host does not fulfil the nutritional

requirements of the pathogen and thereby limits its growth and
reproduction.
4. SYSTEMIC ACQUIRED RESISTANCE (SAR)

The signals that lead to systemic expression of the

antimicrobial PR (pathogenesis-related) genes in the un inoculated
distal tissue to protect the rest of the plant from secondary
infection. This phenomenon is called systemic acquired resistance
(SAR).
SYSTEMIC ACQUIRED RESISTANCE
 IMPORTANCE

❖Systemic acquired resistance (SAR) is an important component of plant

defense against pathogen infection.
❖Systemic acquired resistance (SAR) is a mechanism of induced defense
that confers long-lasting protection against a broad spectrum of
microorganisms.
❖SAR requires the signal molecule salicylic acid (SA) and is associated
with accumulation of pathogenesis-related proteins, which are thought to
contribute to resistance.
❖SA is apparently not the translocated signal but is involved in
transducing the signal in target tissues.
 SAR
Avr R gene
SA

NPR1

PR-1 PR-2 PR-5

SAR
Methods of Breeding for Disease and Insect Resistance

The following breeding methods have commonly used,

1) Introduction
2) Selection
3) Mutation
4) Hybridization
5) Somaclonal Variation
6) Genetic engineering
1. Introduction

plant introduction is the process of taking / introducing plants/

genotype or group of genotype into new environment where they
were not being grown before.

This is easy and rapid method of developing disease resistant variety.

The resistant variety may be introduced and after testing, if found
suitable, can be released in the disease prone area.
Introduction may be classified into two categories:
a) Primary
b) Secondary

a) Primary Introduction:
When the introduced variety is well suited to the new environment and
is directly released for commercial cultivation without any change the
original genotype, known as primary introduction.

Ex. Introduction of semi dwarf wheat varieties Sonora-64, Lerma Rojo

and semi dwarf Rice Var. TN-1, IR-8, IR-28, and IR-36.
2) Secondary Introduction:
The introduced variety is subjected to selection or may be hybridized
with local variety to transfer one or few desirable characters to the
local variety, known as secondary introduction.

Ex. Kalyan sona and sonalika varieties selected from the material
introduced from CIMMYT.
2. Selection

When the source of resistance is a cultivated variety, mass selection

and pure lines selection in self pollinated crops, mass and recurrent
selection in cross pollinated species, and clonal selection in the
vegetatively propagated crops will be ideal for isolating disease
resistant plants.

The resistant plants may be multiplied, screened for disease

resistance and released a variety.
3. Hybridization

Hybridization is used when resistant genes are available either in

the germplasm or in wild species of crop plants.

After hybridization, the hybrid material is handled either by

pedigree method or by backcross method.

The pedigree method is used when the resistance is governed by

polygene.

The backcross method is used when resistance to governed by

oligogenes.
4. Mutation

Induced mutations are also use for disease resistance. Many disease
resistant varieties have been developed in various crops through
induced mutations.
5. Somaclonal Variation

Somaclonal variation is the variation seen in plants that have

been produced by plant tissue culture.

Disease resistant somaclonal variants can be obtained in the

following two ways,

Firstly, plants regenerated from cultured cells or their progeny are

subjected to disease test and resistant plants are isolated.

Secondly, cultured cells are selected for resistance to the toxin or

culture filterate produced by the pathogen and plants are
regenerated from the selected cell.
6. Genetic Engineering

Genetic engineering, also called genetic modification, is the direct

manipulation of an organism's genome using biotechnology.

It is a set of technologies used to change the genetic makeup of cells,

including the transfer of genes within and across species boundaries to
produce improved or novel organisms.

New DNA may be inserted in the host genome by first isolating and
copying the genetic material of interest using molecular
cloning methods to generate a DNA sequence, or by synthesizing the
DNA, and then inserting this construct into the host organism.

Ex. Bt-Cotton
 Genetics of Insect Resistance in Plant

The inheritance of insect resistance may be governed in three ways

1) Oligogenes
2) Polygenes
3) Cytoplasmic gene
In some cases, insect resistance is governed by one or few genes.
Examples of oligogenic insect resistance include in wheat to Hessian fly
and stem sawfly, in barley to green bugs, in alfalfa to pea aphid, in cotton
to Jassids, in apple to woolly aphid etc. The oligogenic resistance from
one host to another is simple.

In some cases, insect resistance is governed by several genes each gene

having small and additive effect.
Examples of polygenic insect resistance are: in wheat to cereal leaf
beetle, in rice to stem borer, in maize to ear worm and leaf aphid, in
alfalfa to spotted aphid, etc.

Sometimes, insect resistance is governed by cytoplasmic genes.

Examples, resistance to European corn borer in maize.
 Genetics of Insect Resistance in Some Crop Plants

Genetic Control of
Resistance Crop Species Insect Pest
Oligogenic Wheat Grenbugs
Hessian fly
Barley Greenbugs
Rice Plant hopper
Maize Corn borer
Cotton Jassids
Alfalfa Pea Aphid
Raspberry Rubus Aphid
Apple Woolly Aphid
Genetic Control of Crop Insect
Resistance Species Pest
Polygenic Wheat Leaf beetle
Rice Stem borer
Maize Earworm
Leaf Aphid
Brassica Aphid
Alfalfa Spotted Aphid
Cytoplasmic Maize European
Corn borer
Lettuce Root Aphid
 Genetics of Disease Resistance in Plant

1. Oligogenic Inheritance

2. Polygenic Inheritance

3. Cytoplasmic Inheritance
Genetics of Disease Resistance in Some Crop Plants
 Types of resistance
Resistance
• Types of resistance
Resistance to diseases is also a genetically controlled
character.
• Plants possess two different types of resistance:

• Monogenic- It is controlled by specific single gene. This

type possesses high resistance to a given strain or race of
the pathogen but it susceptible to other races.
• Polygenic- It is controlled by many genes and is not so
high but at the same time does not easily breakdown due
to the evolution of new races. This is also referred to as
durable resistance.
 Vertical vs. horizontal resistance
• The concept of vertical and horizontal resistance was
suggested by Vander Plank in 1968.
• Resistance is vertical (differential) when it is
completely effective against some races of a pathogen
but not against the others.
• Vertical resistance is complete but is not permanent.
• Horizontal resistance is effective against all races of a
pathogen.
• Horizontal resistance, though incomplete, is of
permanent nature.
 Vertifolia Effect
• Van der Plank introduced the term Vertifolia effect derived from
the German potato variety ‘Vertifolia’ having the late blight
resistance genes R3 and R4.
• The variety became susceptible when the appropriate Pathotypes
P(3,4) evolved in the pathogen leading to a nearly complete failure
of the crop; such a total failure of vertical resistance leading to a
disease epidemic is known as “Vertifolia effect”.
• Thus Vertifolia effect refers to an epidemic development in a variety
carrying vertical resistance genes and a low level of HR, leading to
heavy economic losses.

The failure occurs because of the following two reasons:

• 1) The level of horizontal resistance in the varieties carrying
oligogenes for resistance is usually low, and
• 2) The pathogen is able to evolve the virulent Pathotypes specific to
the oligogenes for vertical resistance.
 Boom and Bust Cycle
• It may be expected that when a new variety carrying VR is developed, it is resistant
to the prevalent Pathotypes of the pathogen in question. Therefore, it becomes
popular rapidly and occupies a large proportion of the growing area under the
concerned crop.
• This period of boom (high popularity and cultivation in a large proportion of area)
continues till the Pathotypes specific to the VR gene (s) employed in the variety
evolves and /or increases in frequency to cause an epidemic in the field occupied by
the variety.
• The now susceptible variety loses popularity and area under it declines sharply thus
the variety is effectively busted by the virulent Pathotypes. This has been called
‘boom and bust’ cycle by Priestey (1970).
• When a variety goes bust, breeders develop new variety with a new VR gene,
which makes it resistant to the predominant Pathotypes. This variety also undergoes
the ‘boom and bust’ cycle, and the breeders put forth a new variety and so on.
• The period for which such a variety may last is , on an average, -5 years in the case
of air-borne fungal pathogens. But in many cases, this period is much shorter, in
extreme cases, the variety go bust even before it reaches farmers fields.
 Gene for gene hypothesis
• It was first proposed by Flor (1942, 1945) as the simplest explanation of
the result of studies of the inheritance of resistance and pathogenicity flax-
rust host-pathogen system, and now applies to most of the combinations.
• According to this hypothesis, “for a resistance gene in the host there is a
complementary avirulence gene in the pathogen.”
• The coexistence of host plant and pathogen side by side in nature indicate
that they have co-evolved; and changes in virulence of pathogen have been
continuously governed by the changes in host or vice versa.
• This concept has been shown to operate in many other diseases like rusts,
smuts, apple scab, late blight of potato and many other fungal , bacterial ,
virus and higher parasitic plant diseases.
• Generally, but not always in the host, gene for resistance is dominant (R)
and gene for susceptibility is recessive (r).
• In pathogen, however, gene for avirulence that have inability to infect is
dominant (A) and for virulence it is recessive (a).
 Gene for gene hypothesis
Thus plant variety when carrying gene for resistance (R) for certain pathogen and
other lacking gene (R) i.e. carrying gene for susceptibility (r) are inoculated with
two races of pathogen, one of which carries a gene for avirulence (A) and other
carrying the gene for virulence (a) against (R); it gives a 4 gene combinations as
below:
1. Out of the four combinations as above; only AR interaction is resistant or
incompatible i.e. host has certain gene for resistance(R) that recognizes the
corresponding genes for avirulence (A), so there is incompatibility.
2. In Ar combination, infection results because host lacks the gene
for resistance so pathogen can attack with other gene for virulence.
3. In aR combination, infection occurs, although the host has gene
for resistance but the pathogen lacks the gene for avirulence (A) which is
recognized by specific gene for resistance; hence no defence mechanism is
activated.
4. Finally in ar gene combination, as plant lacks the gene for resistance and have
gene for susceptibility (r) and pathogen have the gene for virulence (a), it
results in infection.
 Breakdown of resistance
• A new gene for virulence that attacks existing gene
for resistance appears by mutation of an existing avr gene
which then avoids gene for gene recognition and
the resistance breaks down.
• Plant breeders introduce another gene for resistance (R) in
plant which recognizes the protein of new gene for
virulence of pathogen and extend the resistance of host
beyond the range of new gene for virulence in pathogen.
• This produces a variety that is resistant to all races that have
an avirulence gene corresponding to specific gene
for resistance until another gene for virulence appears in
pathogen.
Breeding vegetatively (clonally)
propagated species
 Breeding Method in Asexually Propagated Crop –
Hybridization
• Crops which are propagated asexually or by vegetative means are known as asexually propagated
or vegetatively propagated or clonal crops. There are some agricultural (Sugarcane, Potato, Sweet
Potato, etc) and horticultural (Banana, mango, citrus, pears, peaches, litchi, etc) crops that
propagates by asexual means. The main reasons of asexual reproduction are:

• 1) Reduced flowering and seed set.

2) Non flowering in many cases.
3) To avoid inbreeding depression in certain crops and
4) Apomixis in some species.

• Asexual reproduction produces the progeny, which are exactly identical to their parent in genotype,
because the progeny is derived from the vegetative cells through mitosis. Therefore, the main
advantage of asexually reproduction is that it preserves the genotype of an individual indefinitely.
 Characteristics of Asexually propagated crops
1. A great majority of them are perennials e.g. sugarcane, fruit trees etc.,
2. Many of them show reduced flowering and seed set.
3. They are invariably cross pollinated
4. These crops are highly heterozygous ands show severe inbreeding
depression
5. A vast majority of asexually propagated crops are either polyploids
eg., sugarcane, potato.
6. Many species are interspecific hybrids eg. Banana and sugarcane.
7. This crop consists of a large number of clones that is progeny derived
from a single plant through asexual reproduction.
 CLONE
• A clone is a group of plants produced from a single plant through
asexual reproduction. Thus asexually propagated crops consists of a
large number of clones These crops are also known as clonal crops.
• All the members of a clone have the same genotype as the parent
plant. As a result, they are identical with each other in genotype.
• Consequently the phenotypic differences within a clone do not have a
genetic basis and are purely due to the environmental effects.
 Characteristics of a clone
1. All the individuals belonging to a single clone are identical in
genotype.
2. The phenotype variation with in a clone is due to the environment only
3. The phenotype of a clone is due to the effects of genotype (G) the
environment (E) and the G x E interaction over the population mean (r)
Phenotype (P) = M+G+E+ GE
4. Theoretically clones are immortal i.e. a clone can be maintained
indefinitely through asexual reproduction. But clones usually degenerate
due to viral or bacterial infections.
5. Clones are generally, highly heterozygous and show severe loss in
vigour due to inbreeding.
 CLONAL SELECTION
• Some agricultural crops and a large number of horticultural crops are
asexually propagated. Some common asexually propagated crops are
sugarcane, potato, sweet potato colocasia, Discorea (gams), Mentha,
Ginger, turmeric, banana etc., almost all the fruit trees.
• Segregation and recombination produce new gene combinations due to
which the progeny differ from their parents in genotype and
phenotype.
• Asexual reproduction, on the other hand, produces progeny exactly
identical to their parents in genotype because the progeny are derived
from vegetative cells through mitosis.
• It preserves the genotype of an individual indefinitely. Any genotype is
preserved and maintained through asexual reproduction.
Genetic variation within clones
• It may arise due to somatic mutation, mechanical mixture and
occasional sexual reproduction.

Clonal degeneration:-
• Theoretically, clones are immortal.
• The loss in vigour and productivity of clones with time is known as
clonal degeneration.
• The clonal degeneration may result from (1) mutation (2) viral
diseases and (3) bacterial diseases.
Mutation: It is a recurrent process, it may become a problem over a
long period of time.

Viral diseases: They are easily transmitted through vegetative

propagules. Viruses are perhaps responsible for more cases of clonal
degeneration than any other single cause.

Bacterial diseases:- By the result of bacterial infections making clonal

degeneration.
Natural Vegetative Propogation

New plants grow from parts of the parent

plant. They include:

Stems:-
Runners are stems that grow horizontally
above the ground. They have nodes where
buds are formed. These buds grow into a
new plant.
Roots
• New plants will grow out of swollen, modified roots called tubers. Buds develop
at the base of the stem and then grow into new plants.
Leaves
• Leaves of some plants will grow into a new plant if they become detached from the parent plant.
Other plants grow small plants called plantlets on the edge of their leaves. i.e. cactus, thorns etc.
Bulbs
• A bulb contains an underground stem. Leaves are attached to the stem. These leaves contain much
stored food. At the center of the bulb is an apical bud. Also attached are lateral buds. The apical
bud will produce leaves and a flower while the lateral buds will produce new shoots. As the plant
grows and develops it will form a new bulb underground.
Artificial Vegetative Reproduction
• Horticulturists and farmers use artificial means to produce plants that are identical to the parent
plant. Some of the methods used are:
Cuttings
• Cuttings are part of the plant that is cut off of the parent plant. Shoots with leaves attached are
usually used. New roots and leaves will grow from the cutting. The shoot is cut at an angle. A
growth promoter may be used to help with the growth of the roots.
Grafting
• In grafting 2 plants are used to develop a new plant with combined traits from the 2 parent plants.
In grafting the scion is the above ground part of one plant. The scion is attached to the stock which
is the rooted part of the second plant.
Brimto : Most recent achievement of ICAR
Layering
• In layering a shoot of a parent plant is bent until it can be covered by soil. The tip of the shoot
remains above ground. New roots and eventually a new plant will grow. These plants can then be
separated.
 Methods of improvement of Asexually propagated crops

• A single outstanding plant selected form a population forms the basis

of a new variety. The breeding behaviour or genotypes of the plant is
not important, Since there would be no further sexual reproduction.
The outstanding plant may be selected from an old unimproved
variety, an improved variety that has become variable or from a
population produced by crossing two or more clones. The procedure of
selection used for such crops is known as "clonal selection". Since the
selected plants are used to produce new clones.
 Breeding approaches used in asexual crops
Several breeding approaches are used in the breeding of asexually propagated species.

Planned introduction

• Just like seed, vegetative material (whole plants or parts) may be introduced into a new
production area for evaluation and adaptation to the new area.

• Seedlings or cut- tings may be introduced.

• However, the technology of tissue culture allows a large variety of small samples to be
introduced in sterile condition.

• These disease-free samples are easier to process through plant quarantine.

Hybridization with clonal selection

• This procedure is applicable to species that are capable of producing seed in appreciable quantities.

• Because heterosis can be fixed in clonal populations, the breeder may conduct a combining ability analysis to
determine the best combiners to be used in hybridization
 Interspecific hybridization in the improvement of clonal crops
• Interspecific hybridization has been successfully used in the improvement of
clonal crops like, potato, sugarcane, strawberries etc., potato variety "Kufri Kuber'
was developed from a complex cross (Solanum curtilobum x S. tubersoum) x
Solanum andigena. This vareity shows much less clonal degeneration in the plains
than the vareity up to date.
• Generally interspecific crosses are made to transfer specific characters such as
disease resistance from the wild species to the cultivated potato. For example S.
demissum has been extensively used as a source of late blight resistance.
• All sugarcane varieties now in cultivation have developed from complex crosses
between sugarcane varieties have been developed from complex crosses between
Saccharum officinarum (Noble cane), S. barberi (Indian cane) and Saccharum
sp. like S. spontaneous has been used to combine its and high yielding ability of S.
officinarum.
• The principle reason for such a great success is then asexual reproduction this
completely avoids segregation and recombination. Another reason in that most of
them are not seed crops, hence flowering and fertility are not essential for their
success as varieties.
Problems in the breeding asexually propagated
crops
• There are several problems peculiar to clonal crops, which are difficult
to resolve. There are three major problems in their breeding

1. Reduced flowering and fertility

2. Difficulties in genetic analysis and
3. Perennial life cycle.
• Other techniques that are applicable include backcrossing to transfer specific traits and wide crossing.

• The challenges with backcrossing are several.

• As previously indicated, clonal species are very heterozygous and prone to inbreeding depression.

• Backcrossing to one parent (the recurrent parent) provides an opportunity for homozygosity and
consequently inbreeding depression.

• To prevent this, breeders may cross the backcross to another clone instead of the recurrent parent,
followed by selection to identify superior plants.

• The process is repeated as needed.

Breeding implications, advantages, and limitations of clonal
propagation

Advantages

1.1 Sterility is not a factor in clonal propagation because seed is not involved.

1.2 Because clonal plants are homogeneous, the commercial product is uniform.

1.3 Micropropagation can be used to rapidly multiply planting material.

2.4 Heterozygosity and heterosis are fixed in clonal populations.

Disadvantages

1 Clonal propagules are often bulky to handle (e.g., stems, bulbs).

2 Clones are susceptible to devastation by an epidemic. Because all plants in the clonal population are
identical, they are susceptible to the same strain of pathogen.

3 Clonal propagules are difficult to store for a long time because they are generally fresh and
succulent materials.
Genetic issues in asexual breeding

Genetic makeup

• All the progeny from an individual propagated asexually are

genetically identical (clones) and uniform.

• Clones are products of mitosis. Any variation occurring among

them is environmental in origin.
Heterozygosity and heterosis

• Many species that are asexually propagated are highly heterozygous; they are
highly heterotic.

• Consequently, they are susceptible to inbreeding depression.

• For those species that can be hybridized without problems, an advantage of

asexual propagation is that heterosis, where it occurs, is fixed for as long as the
cultivar is propagated asexually.

Ploidy

• Many known species that are asexually propagated are interspecific hybrids or
have high ploidy.
 THANK YOU
“Happy Learnings”
Unit 6
 Unit VI

Plant biotechnology in crop improvement:

biotechnological tools-dna markers and
marker assisted selection

Intellectual property rights: participatory

plant breeding, intellectual property rights,
patenting, plant breeders and farmer’s rights
Plant biotechnology in crop improvement:

Biotechnological tools:-
➢DNA markers and
➢marker assisted selection (MAS)
 What is a marker?

• Any genetic element (locus, allele, DNA sequence or

chromosome feature) which can be readily detected by
phenotype, biochemical, cytological or molecular
techniques.

• A marker is a signpost linked to the trait of interest and

is co-inherited along with the trait.
TYPES OF MARKERS:

Morphological markers

Cytological markers

Biochemical markers

DNA markers
Molecular markers and their type that are used in plant
breeding
 Hybridization
based • Restricted Fragment Length
markers Polymorphism (RFLP)

• Simple Sequence Repeats (SSRs)

Types of • Inter Simple Sequence Repeats
(ISSRs)
Molecular/ PCR based • Random Amplified Polymorphic
markers DNA (RAPD)
• DNA Amplification
DNA markers Fingerprinting (DAF)
• Sequence Tagged Sites (STS)

DNA
sequence
based • Single Nucleotide
Polymorphisms (SNPs)
markers
7
 MORPHOLOGICAL MARKERS
➢ These are qualitative traits that can be scored visually. They are
generally dominant or recessive.
e.g. flower colour, seed shape,
growth habits or pigmentation etc.
Merits:
➢ Easy to score,
➢ not costly, etc.
Demerits:
➢ They are highly influenced by the environment, etc.
➢ Their effect may be undesirable in breeding programs.
➢ They may mask the effects of linked genes making it nearly
impossible to identify desirable linkages for selection.
➢ Limited in number and are generally dominant. Besides, they
often interfere with other traits.
➢Growth stage specific
 Biochemical markers
➢ These are the proteins produced by gene expression which can

be isolated and identified by electrophoresis and staining.

➢ Most of the biochemical makers are polymorphic and insensitive

to environment just like molecular markers.

➢ Isozyme – These are the different molecular form of same

enzymes and these isozymes used as biochemical markers

 DNA Based Markers or molecular markers
➢These DNA based makers differentiate the organisms at DNA
level and are inherited in simple Mendelian fashion.

➢A DNA segment of known or unknown structure and function,

which can be used to detect the presence of specific sequence of
nucleotides in another DNA or RNA molecule.
BASIS OF DNA POLYMORPHISM
• Base substitution
• Insertion/Deletion
• Duplication
• Inversion
• Translocation
• Transposition – Transposons
• Recombination – unequal crossing over
• Variation in the number of tandem repeats
 Properties desirable for ideal DNA
markers
• Highly polymorphic nature
• Co-dominant inheritance (determination of homozygous and
heterozygous states of diploid organisms)
• Frequent occurrence in genome
• Selective neutral behavior (the DNA sequences of any organism are
neutral to environmental conditions or management practices)
• Easy access (availability)
• Easy and fast assay
• High reproducibility
Some commonly used types of genetic markers are
Non-PCR Based or Hybridization based marker,
➢ RFLP- Restriction fragment length polymorphism.

PCR Based
➢ RAPD- Random amplification of polymorphic DNA.
➢ AFLP-Amplified fragment length polymorphism.
➢ SCAR-Sequence characterize amplified region.
➢ STS- Sequence tagged sites.
➢ EST-Express sequence tags.
➢ SNP-Single nucleotide polymorphism.
➢ SSR-Simple sequence repeats
➢ CAPS-Cleaved amplified polymorphic sequences
 Marker assisted selection (MAS)

The use of DNA markers that are tightly-linked to target loci

as a substitute for or to assist phenotypic screening.
Why Marker Assisted Selection?
➢Selection at seedling stage possible
➢Selection of traits with low heritability
➢Distinguishing homozygotes from heterozygotes
➢Pyramiding of Resistance Genes
➢Selection for recessive gene, etc.
Main feature of MAS
Main feature of MAS
Five main considerations for the use of DNA
markers in MAS
❑Reliability: Molecular markers should co-segregate or tightly linked
to traits of interest, preferably less than 5 cM genetic distance. The
use of flanking markers or intragenic markers will greatly increase the
reliability of the markers to predict phenotype.
❑DNA quantity and quality: Some marker techniques require large
amounts and high quality DNA, which may sometimes be difficult to
obtain in practice, and this adds to the cost of the procedures.
❑Technical procedure: Molecular markers should have high
reproducibility across laboratories and transferability between
researchers. The level of simplicity and time required for the technique
are critical considerations. High-throughput simple and quick methods
are highly desirable.
❑Level of polymorphism: Ideally, the marker should be highly
polymorphic in breeding material and it should be co-dominant for
differentiation of homozygous and heterozygous individuals in
segregating progenies.
❑Cost: Molecular markers should be user-friendly, cheap and easy to use
for efficient screening of large populations. The marker assay must be
cost-effective in order for MAS to be feasible.
 Markers must be tightly-linked to target loci

• Ideally markers should be <5 cM from a gene or QTL

Marker A

QTL
5 cM

Marker A Marker B

QTL
5 cM 5 cM
 Markers must be polymorphic
RM84 RM296

1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8

P1 P2
P1 P2

Not polymorphic Polymorphic!

 WHY USE MARKER-ASSISTED
SELECTION
(i) traits that are difficult to manage through conventional
phenotypic selection-because they are expensive or time-
consuming to measure, or have low penetrance or complex
inheritance;
(ii) traits whose selection depends on specific environments or
developmental stages that influence the expression of the
target phenotype;
(iii) maintenance of recessive alleles during backcrossing or
for speeding up backcross breeding in general; and
(iv) pyramiding multiple monogenic traits (such as pest and
disease resistances or quality traits) or several QTL for a
single target trait with complex inheritance (such as
drought tolerance or other adaptive traits).
Overview of (1) LEAF TISSUE
MAS SAMPLING
MAPPING
POPULATION
(2) DNA EXTRACTION
• F2 progenies
• F2 derived F3
• DH lines (3) PCR
• BC progenies
• RILs
• NILs (4) GEL ELECTROPHORESIS

(5) MARKER ANALYSIS

Advantages of MAS over conventional
phenotypic selection
1. It may be simpler than phenotypic screening, which can save
time, resources and effort. Classical examples of traits that are difficult
and laborious to measure are cereal cyst nematode and root lesion
nematode resistance in wheat
2. Selection can be carried out at the seedling stage. This may be
useful for many traits, but especially for traits that are expressed at later
developmental stages. Therefore, undesirable plant genotypes can be
quickly eliminated.
3. Single plants can be selected. With MAS, individual plants can
be selected based on their genotype. For most traits, homozygous and
heterozygous plants cannot be distinguished by conventional
phenotypic screening.
Intellectual Property Rights (IPR):
Participatory Plant Breeding,
Intellectual property rights,
Patenting,
Plant Breeders and Farmer’s rights
 PARTICIPATORY PLANT BREEDING

• Participatory plant breeding is the process by which farmers

are routinely involved in a plant breeding programme with
opportunities to make decisions throughout.
What are the goals of PPB?

• Increase production and profitability of crop production through the

development and enhanced adoption of suitable, usually improved,
varieties.
• Provide benefits to a specific type of user, or to deliberately address
the needs of a broader range of users.
• Build farmer skills to enhance farmer selection and seed production
efforts.
What activities can PPB include?
• Identifying breeding objectives
• Generating genetic variability (including the provision of plants to be
included in breeding program)
• Selecting within variable populations to develop experimental
varieties
• Evaluating experimental varieties (PVS - participatory variety
selection)
• Variety release
• Popularization (diffusion of information about new variety and how it
is managed)
• Seed production
INTELLECTUAL PROPERTY RIGHTS

• Intellectual property refers to creations of the mind: inventions;

literary and artistic works; and symbols, names and images used in
commerce.

• The right of an inventor to derive economic benefits from his

intellectual property is known as “Intellectual Property Rights”
IPR PROTECTION

• Main forms of IPR protection are:

I. Trade Secret
II. Patent
III. Copyright
IV. Geographical Indications (GI)
V. Trademarks:
VI. Trade names
VII. Plant Breeder’s Right
Trade Secret

• A trade secret is anything [a formula, process, method,

mechanism, tool, pattern or device] which the disclosing
party desires to keep secret
PATENT

• The right granted by a government to an inventor to exclude others

from imitating, manufacturing, using or selling the invention in
question for commercial purpose during the specified period.
• Patents are granted for:
✓ an invention
✓ innovation in an invention
✓ a concept
 Patent requirement

• Novelty
• Inventiveness
• Industrial applicability
• Patentability
Limits of a patent

1. Limitation of time:
A patent is valid for specified period of time from the date of its
award; in most countries this period is 15-20 years

2. Limitation of space:
A patent is valid only in the country of its award; it is not valid in other
countries.
 copyright

• The legal right given to an author, composer, playwright,

publisher, or distributor to exclusive publication, production,
sale, or distribution of a literary, musical, dramatic, or artistic
work is called copyright.
• In India the copyright is protected by [The Indian Copyright
Act (1957), amended in 1999] are per as international
standards.
 Geographical Indications (GI):
• GI are signs used on goods that have a specific
geographical origin and possess qualities or a reputation that
are due to that place of origin
Trademarks:

• A trademark is a distinctive sign that identifies certain goods

or services as those produced or provided by a specific
person or enterprise. It may be one or a combination of
words, letters, and numerals.

Trade name: A trade name is the name which a

business trades under for commercial
purpooses, although its registered, legal name,
used for contracts and other situation, may be
another.
PLANT BREEDER’S RIGHT

• Rights granted by a government to a plant breeder, originator or

owner of a variety to exclude others from producing or
commercializing the propagating material of that variety for a
minimum period of 15-20 yrs.
• The plant breeder right are also known as plant
variety rights.
• The protection of the PBR is valid only in the
country where it has been registered.
• The four basic requirements for protection of a
plant variety under PBR, viz., novelty,
distinctiveness, uniformity and stability.

• The PBR provides right to breeder or owner to

prevent others from commercial production and
marketing of his variety without permission.

• The PBR provides exclusive rights to the owner for

commercial production and marketing of his variety.
REQUIREMENTS OF PBR
• Novelty:
A variety should not have been commercially exploited for more than
one year before the grant of PBR protection

Distinctiveness:
A new variety must be distinguishable from other varieties by one or
more identifiable morphological, physiological or other characteristics

• Uniformity
A variety must be uniform in appearance under specified environment of
its adaptation

• Stability
A variety must be stable in appearance and its clonal characteristics over
successive generations under specified environment
Benefits from PBR

1) It allows breeders to benefit from the varieties developed by them

2) Encourage the plant breeding activities
3) Private sector is encouraged to invest in plant breeding and seed
industry
4) Increased competition among various organizations engaged in the
plant breeding is likely to be beneficial to both the farmers and the
nation
5) It will enable access to varieties developed in other countries and
protected by IPR laws.
• Farmer’s Rights
• Farmer’s variety
• Extant variety
 Farmers rights
• The farmers rights refer to the rights arising from the past, present
and future contribution of farmers in conserving, improving and
making available plant genetics resources, particularly in the centre of
origin.
• The farmer rights have been recognised by FAO conference held in
Rome from 11-29 November 1989.
• In India the protection of farmers and plant breeders right, new plant
variety is provided by the Protection of Plant Varieties and farmer’s
Right Act (PPVFR, 2001).
• The PPVFR provides rights to farmers to save, exchange, share and
sell his farm produce including seed of a protected variety.
 Farmers’ variety

➢A variety which (i) has been traditionally cultivated and

evolved by the farmers in their fields, or

➢(ii) is a wild relative or land race of a variety about which

the farmers possess common knowledge.
 Extant variety

➢A variety available in India which is

(i) notified under section 5 of Seeds Act, 1966, or
(ii) farmers’ variety, or
(iii) a variety about which there is common
knowledge, or
(iv) any other variety which is in public domain.
•UPOV and PPVFR
 UPOV
International Union for the
Protection of New Varieties of
Plants
➢ The International Union for the Protection of
New Varieties of Plants (UPOV) is an
intergovernmental organization based in Geneva,
Switzerland.
➢ UPOV was established in 1961 by the
International Convention for the Protection of
New Varieties of Plants (the “UPOV
Convention”).
➢ The mission of UPOV is to provide and promote
an effective system of plant variety protection,
with the aim of encouraging the development of
new varieties of plants, for the benefit of society.
➢Most countries and intergovernmental
organizations which have introduced a plant variety
protection (PVP) system have chosen to base their
system on the UPOV Convention in order to
provide an effective, internationally recognized
system
➢ The UPOV Convention provides the basis for
UPOV members to encourage plant breeding by
granting breeders of new plant varieties an
intellectual property right: the breeder’s right.
➢ In order to obtain protection, the breeder needs to
file individual applications with the authorities of
UPOV members entrusted with the task of granting
breeders' rights.
➢ The UPOV Convention specifies the acts that
require the breeder’s authorization in respect of the
propagating material of a protected variety and,
under certain conditions, in respect of the
harvested material.
 Aim of UPOV
The introduction of the UPOV system of plant variety
protection and UPOV membership were found to be
associated with:
(a) increased breeding activities,
(b) greater availability of improved varieties,
(c) increased number of new varieties,
(d) diversification of types of breeders (e.g. private breeders,
researchers),
(e) increased number of foreign new varieties,
(f) encouraging the development of a new industry
competitiveness on foreign markets, and
(g) improved access to foreign plant varieties and enhanced
domestic breeding programs.
 PPVFR

➢ The protection of Plant Varieties and Farmers

Rights Act 2001 was approved by the
government of India in 2004 but came in to force
with effect from January 2005.
➢ The main objective of this act is to encourage
development of new plant varieties and promote
development of seed industries.
➢ Headquarter of PPV and FRA located in Pusa
campus (National Agricultural Complex), New
Delhi.
 Objectives of PPV and FRA Act 2001

➢ To establish an effective system for protection of

plant varieties, farmer’s rights and plant breeders
rights.
➢ To recognize and protect farmer’s rights for their
contribution in conserving, improving and making
available plant genetic resources for development of
new plant varieties.
➢ To stimulate investment for research and
development both public and private sector for
development of new plant varieties.
➢ To promote the growth of seed industries in country
to ensure availability of high quality seeds to farmers.
 Plant material Protected under PPV and FRA

➢ All new varieties of self pollinated species

such pure line and multi line varieties.
➢ New varieties of cross pollinated crop
species such as open pollinated synthetic and
composite varieties
➢ Varieties of asexually propagated crop.
➢ Single, three way and double cross hybrids.
➢ Inbred parental lines of hybrid varieties.
➢ Trees, Fruits, ornamental and vines etc.
 Requirements for registration under PPV and
FRA

DUS testing-
Novelty
Distinctness
Uniformity
Stability
 Duration of Protection Under PPV and
FRA

➢According to PPV and FRA Act, 2001

Protection period of annual and
biannual crop is 15 years.
➢ For perennial plants such as trees and
vines is 18 years.
Merits of PPV and FRA, 2001