PLANT BREEDING

1) History of plant breeding

2) Modes of reproduction, selfing, and crossing techniques.

3) Origin, evolution, and domestication of crop plants, centre of origin, law of

homologous series, crop genetic resources conservation and utilisation.

4) Application of principles of plant breeding, improvement of crop plants

5) Molecular markers and their application in plant improvement

6) Pure-line selection, pedigree, mass and recurrent selections, combining ability, and its
significance in plant breeding.

7) Heterosis and its exploitation.

8) Somatic hybridisation.

9) Breeding for disease and pest resistance.

10) Role of interspecific and intergeneric hybridisation.

11) Genetically modified crop plants.

History of plant breeding, origin, evolution, and domestication of crop plants, centre of
origin, law of homologous series, crop genetic resources, conservation and utilization

1. Describe the history of plant breeding in India. Write the objectives of plant breeding and
methods for creation of variability for crop improvement. (20) 2014

2. Explain the law of homologous series. How is it helpful in genetic resources, conservation
and utilization? (12.5) 2017

3. What do you understand by centre of origin? Discuss the law of Homologous series of
variation. (15) 2013

4. Discuss the concept of centre of origin of crops and its role in plant breeding. Describe the
role of N.I. Vavilov in identifying centre of origin of various crops with appropriate examples.
(20) 2016

5. Write down about the origin and domestication of rice. (10) 2019

6. Gene Banks (12.5) 2011

7. Discuss the importance of conservation of plant genetic resources for crop improvements.
(12.5) 2018

8. Describe the role of plant genetic resources conservation in crop improvement. (10) 2020
9. What do you mean by gene pool concept and genetic erosion? Classify the gene pool of a
crop. (10) 2015

10. Give a brief account of conservation of crop genetic resources. (10) 2012

11. Discuss the importance of crop genetic resource conservation and utilization. (20) 2014

12. Discuss orthodox and recalcitrant seeds with examples. (10) 2017

13. What do you mean by conservation of plant genetic resources? Explain their
contribution in crop improvement in India. (15) 2016

14. What do you understand by passport data of genetic resources? Write its importance in
germplasm collection and conservation. (8) 2018

15. Explain in situ and ex situ germplasm conservations. (10) 2013

16. Explain the right approaches for breeding of crop varieties for wide adaptability. (8M IFS-
2024) 2024

Plant breeding can be defined as an art, a science, and a technology of improving the genetic
makeup of plants in relation to their economic use for mankind. Plant breeding deals with
the genetic improvement of crop plants also known as science of crop improvement.

History and development of plant breeding:

- 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 continues today and is likely to continue for some time in future

- Ex: In case of timber trees, medicinal plants, microbes

- During the long period of historic cultivation natural selection has definitely acted on the
domesticated species.

- Movement of man from one place to another brought about the movement of his
cultivated plant species

Early historical developments:

- 700 BC - Babylonians and Assyrians pollinated date palm artificially

- 1717 - Thomas Fairchild - produced the first artificial hybrid, popularly known as Fairchild’s
mule, by using carnation with sweet William

- 1727 - The first plant breeding company was established in France by the Vilmorins.
- 1760-1766 - Joseph Koelreuter, a German, made extensive crosses in tobacco.

- 1759-1835 – Knight was perhaps the first man to use artificial hybridisation to develop
several new fruit varieties.

1840 – Le Couteur & Shireff:

Applied individual plant selection and progeny testing to develop better cereal varieties.

1856 – Vilmorin (France):

Improved the progeny test method and used it successfully in sugar beet improvement.

1900 – Nilsson & Associates (Sweden):

Developed and popularized individual plant selection at Svalöf Experiment Station.

1903 – Johannsen’s Pure Line Theory:

Provided the genetic basis for selection by defining uniform, genetically pure lines.

Genetic Foundations of Modern Breeding

1900 – Rediscovery of Mendel’s Work:

Mendel’s laws of inheritance became the foundation of scientific plant breeding.

Cytogenetic Advances:
Discovery that chromosomes carry genes led to chromosome manipulation and cytogenetic
breeding.

Development of Hybrid Breeding

G.H. Shull’s Maize Studies (1908):

Found that inbreeding reduces vigour, but crossing inbred lines restores it (heterosis).

Hybrid Variety Revolution:

Hybrid breeding became standard in crops like maize, sorghum, and pearl millet for higher
yields.

Modern Phase of Plant Breeding

Integration with Genetics and Biotechnology:

Modern plant breeding uses molecular tools, cytogenetics, and biotechnology for precision
improvement.

Focus of Modern Breeding:

Emphasis on yield, pest and disease resistance, stress tolerance, and nutritional
enhancement.

History of plant breeding in India

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

- 1905 – The Imperial Agricultural Research Institute was established in Pusa, Bihar
- 1934 – The buildings of the institute were damaged in an earthquake

- 1936 – Shifted to New Delhi

- 1946 – Name was changed Indian Agricultural Research Institute

- 1901-05 – Agricultural Colleges were established at Kanpur, Pune, Sabour, Llyalpur,

Coimbatore

- 1929 – Imperial Council of Agricultural Research was established

- 1946 – Name was changed to the Indian Council of Agricultural Research

- 1921 – Indian Central Cotton Committee was established – Notable research on breeding
and cultivation of cotton. Eg, 70 improved varieties of cotton

- 1956 – Project for intensification of regional research on cotton, oilseeds and millets
(PIRRCOM) was initiated to intensify research on these crops – located at 17 different
centres throughout the country

- 1957 – All India Coordinated maize improvement project was started with the objective of
exploiting heterosis

- 1961 - The first hybrid maize varieties released by the project

- ICAR initiated coordinated projects for improvement of the other crops

- 1960 – First Agricultural University established at Pantnagar, Nainital, U.P.

Objectives of Plant Breeding:

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

2. Improved quality: Quality of produce varies from crop to crop. E.g., grain size, colour,
milling and backing quality in wheat. Cooking quality in rice, malting quality in barley, colour
and size of fruits, nutritive and keeping quality in vegetables, protein content in pulses, oil
content in oilseeds, fibre length in cotton.

3. Abiotic resistance: Crop plants also suffer from abiotic factors such as drought, soil
salinity, extreme temperatures, heat, wind, cold and frost. The 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. 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, It

requires less crop management period, less insecticidal sprays, permits new crop rotations
and often extends the crop area. Thus, breeding for early maturing crop varieties is
necessary. Maturity has been reduced from 270 days to 170 days in cotton, from 270 days
to 120 days in pigeonpea, and from 360 days to 270 days in sugarcane.

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

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

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

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. Tallness, high tillering and profuse branching
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, 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.

10. Non-shattering characteristics: The shattering of pods is a 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. It have permitted
their cultivation in new areas. Rice is now cultivated in Punjab,vb wheat in West Bengal.

13. Wider adaptability: Adaptability refers to the suitability of a variety for general
cultivation over a wide range of environmental conditions. 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. Mung is grown as a summer crop in addition to
the main kharif crop.

Activities in plant breeding:

The process involves a series of connected steps — variation creation, selection, evaluation,
multiplication, and distribution.

1. Creation of Variation
Importance of Variation: No genetic improvement is possible without variability in the
population.

Sources of Variation: Domestication, germplasm collection, introduction, hybridisation,

mutation, polyploidy, and somaclonal variation generate diversity.

2. Selection

Purpose of Selection: To identify and isolate the most desirable genotypes from a mixed
population.

Basis of Selection: Selection is based on observable phenotype, though phenotype–

genotype correlation can be variable.

Selection Schemes: Different schemes are designed for specific situations — e.g., self-
pollinated vs. cross-pollinated crops.

3. Evaluation

Meaning of Evaluation: Evaluation involves comparing new genotypes with existing varieties
for yield and adaptability.

Duration and Scope: Conducted over multiple years and locations to assess performance
stability.

Role of ICAR Projects: The All India Coordinated Crop Improvement Projects (AICRPs)
efficiently handle evaluation across locations.

4. Release and Multiplication

Variety Release: If a strain outperforms existing ones, it is officially released for commercial
cultivation.

Seed Multiplication: Large-scale multiplication and distribution of new varieties are done by
national and state seed agencies.

5. Coordination and Efficiency

Need for Coordination: All breeding steps must be well-coordinated for an efficient crop
improvement programme.

Consequences of Inefficiency: Neglect or weakness in any step reduces programme

efficiency and wastes valuable resources.

Continuous Review: Regular monitoring helps identify problems, apply corrective measures,
and ensure sustained breeding progress.
Undesirable effects

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

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

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

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

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

combinations. The examples of 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, in some cases, produced severe epidemics. 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 plant species under human management for
cultivation, marking the beginning of agriculture and plant breeding.
 • Present-day cultivated crops evolved from wild and weedy ancestors.

• Evolution occurred through continuous natural and artificial (human) selection.

• Domestication resulted in crops that are genetically, morphologically, and

physiologically distinct from their wild progenitors.

Early Domestication

• Most crop plants were domesticated by prehistoric humans.

• Selection was conscious or unconscious, based on:

o Larger grains or fruits

o Better taste

o Non-shattering habit

o Ease of harvest

• Over generations, domesticated crops became so different that:

o Many are now considered distinct species

o In some cases, wild progenitors are unknown

Modern Domestication

• Domestication is a continuous and ongoing process.

• Modern domestication targets include:

o Crop plants

o Microorganisms (antibiotics, nitrogen fixation)

o Forest trees (timber)

o Medicinal and industrial plants

Selection in Domestication

Variability and Selection

• Early domesticated crops showed high variability

• Variability arose due to:

o Hybridization

o Recombination

o Occasional mutations
 • This variability acted as raw material for selection

What is Selection?

Selection is the process by which some genotypes leave more offspring than others,
leading to preferential survival of desirable traits.

• Desirable plants are retained

• Undesirable plants are gradually eliminated

• Determines which traits persist in future generations

Natural Selection

• Occurs without human involvement

• Driven by environmental factors:

o Temperature

o Soil conditions

o Climate

o Pests and diseases

• Maintains adaptability and variability

• Does not eliminate all variation

Artificial Selection

• Deliberate human selection

• Based on visible traits (phenotype), such as:

o Bigger grains

o Sweeter fruits

o Higher yield

• Selected plants alone are allowed to reproduce

• Results in:

o Faster improvement

o Reduced genetic variability

Result of Selection

• Early selection was primitive but highly effective

 • Combined natural + artificial selection led to:

o Major changes in crop plants

o Loss or obscurity of wild ancestors in many crops

Changes in Crops Due to Domestication

Almost all plant characters have been altered under domestication due to continuous
artificial selection, often supported by natural selection. The major changes and well-
documented crop examples are given below.

1. Reduction or Elimination of Shattering

Meaning: Loss of natural seed dispersal mechanism to facilitate harvesting.

Examples

• Rice: Oryza sativa

o Non-shattering controlled by sh4 gene

o Wild ancestor O. rufipogon shows complete shattering

• Wheat: Triticum aestivum

o Cultivated wheat non-shattering vs wild einkorn (T. boeoticum)

• Sorghum: Sorghum bicolor – domesticated types non-shattering

2. Elimination of Dormancy

Meaning: Seeds germinate immediately after harvest.

Examples

• Wheat: Modern cultivars of Triticum aestivum

• Barley: Hordeum vulgare (malting varieties prone to pre-harvest sprouting)

• Mung bean: Vigna radiata cultivars show reduced dormancy compared to wild types

3. Reduction of Toxic or Undesirable Substances

Meaning: Removal of bitter, poisonous, or anti-nutritional compounds.

Examples

• Cucumber & Pumpkin: Cucumis sativus, Cucurbita spp.

o Reduced cucurbitacin (bitterness)

• Cassava: Manihot esculenta

o Low cyanogenic glycosides in sweet cassava

 • Cotton: Gossypium hirsutum

o Reduced gossypol in cultivated seed types

4. Modification of Plant Type

Meaning: Altered architecture for better yield and management.

Examples

• Rice: Semi-dwarf, erect leaf varieties (e.g., IR-series)

• Maize: Zea mays

o Reduced branching compared to teosinte

• Wheat: Compact plant type vs tall, spreading wild forms

5. Decrease in Plant Height

Meaning: Reduced lodging and higher harvest index.

Examples

• Rice: Semi-dwarf varieties carrying sd-1 gene

o IR8, IR36

• Wheat: Rht-B1, Rht-D1 genes

o Kalyan Sona, Sonalika

6. Increase in Plant Height

Meaning: Selected for fibre or biomass yield.

Examples

• Jute: Corchorus capsularis

• Sugarcane: Saccharum officinarum

• Napier grass: Pennisetum purpureum (fodder)

7. Shortening of Life Cycle

Meaning: Early maturity enables multiple cropping.

Examples

• Cotton: Early maturing Gossypium hirsutum varieties

• Pigeonpea (Arhar): Short-duration varieties (e.g., UPAS-120)

• Rice: Short-duration varieties like ADT-36

8. Increase in Grain or Fruit Size

Meaning: Direct selection for consumer preference and yield.

Examples

• Rice: Long-grain basmati vs small-grain wild rice

• Wheat: Bold-grained durum wheat

• Tomato: Large-fruited cultivars vs wild Solanum pimpinellifolium

9. Increase in Economic Yield

Meaning: Greater proportion of biomass allocated to harvestable parts.

Examples

• Rice & Wheat: Modern HYVs with high harvest index

• Maize: Hybrid maize with increased cob size

• Oilseeds: High oil-yielding groundnut varieties

10. Promotion of Asexual Reproduction

Meaning: Vegetative propagation ensures uniformity and yield stability.

Examples

• Sugarcane: Stem cuttings

• Potato: Solanum tuberosum (tubers)

• Sweet potato: Ipomoea batatas (vine cuttings)

• Banana: Musa spp. (suckers)

11. Preference for Polyploidy

Meaning: Polyploids show larger organs, vigour, and adaptability.

Examples

• Wheat: Hexaploid (Triticum aestivum, 2n=6x=42)

• Potato: Tetraploid (2n=4x=48)

• Tobacco: Nicotiana tabacum

• Sweet potato: Hexaploid

12. Shift in Sex Form

Meaning: Transition toward bisexuality and self-fertility.

Examples

• Papaya: Carica papaya

o Wild dioecious → cultivated hermaphrodite types

• Grape: Vitis vinifera

o Dioecious wild → bisexual cultivated forms

• Brinjal: Elimination of self-incompatibility in cultivated lines

13. Reduction in Variability

Meaning: Increased genetic uniformity for stability and uniform yield.

Examples

• Pureline wheat varieties (e.g., HD-2967)

• Rice varieties released as purelines

• Vegetatively propagated crops (banana, sugarcane)

Domestication of Rice

Origin and Ancestors

• Cultivated rice: Oryza sativa

• Wild progenitors:

o Oryza rufipogon

o Oryza nivara

CENTRES OF ORIGIN:

The concept was proposed by Nikolai Ivanovich Vavilov (Leningrad geneticist) based on
extensive studies of global crop diversity. He observed that cultivated crop plants evolved
from their wild relatives in regions showing maximum variability, which he termed as
Centres of Origin.

Primary Centres of Origin

Primary centres of origin are geographical regions where crops first evolved and show
maximum dominant gene diversity, crops were first domesticated from wild species.

They exhibit High genetic diversity Presence of wild and weedy relatives. Dominant genes
occur frequently. They are usually geographically small but genetically rich.

Secondary Centres of Origin

secondary centres are regions where crops were later introduced (human migration,
cultivation and selection) and show high variability mainly due to recessive genes.

Wild relatives may be absent.

Gene micro centre: In some cases, small areas within the centers of diversity exhibit
tremendous genetic diversity of crop plants. These areas are called as micro centres.

Centre of Diversity

A centre of diversity is a geographical area where a crop shows maximum genetic variability,
irrespective of whether it originated there.

All primary centres of origin are centres of diversity, but not all centres of diversity are
centres of origin.

These regions are valuable for Plant breeding, Conservation of germplasm, Genetic
improvement programs.

Maize originated in Southern Mexico and Guatemala but shows very high diversity in South
America → therefore South America = centre of diversity, not origin.

Types of Centres of Diversity: Vavilov and later researchers recognized three main types:

Type Description Example Crops

Primary Centre of Original site of domestication with wild Rice in India, Potato in
Diversity relatives and maximum variation Andes

Secondary Centre of Region where crop was introduced later Maize in China, Wheat in
Diversity and developed variability Ethiopia

Kashmir Valley (apple,

Micro-centre / Local A small localised area within a larger
walnut), Ethiopian
Diversity Pocket region containing exceptional variability
Highlands

Importance of Centres of Origin and Diversity

• Provide raw material (germplasm) for crop improvement and breeding

• Aid in the identification of wild relatives and resistant genes

• Support biodiversity conservation strategies and help trace the evolution and
domestication history of crops.

Main centres of origin proposed by Vavilov.

1. Chinese centre
 2. Hindustan Centre

3. Central Asiatic Centre

4. Near Eastern centre

5. Mediterranean centre

6. Abyssinian centre

7. South Mexican and Central American centre.

8. South American centre

Geographical Primary Crops Secondary

Centre Significance
Area Covered Originated Centre Crops

Soybean, Radish, Foxtail

& Proso millet,
Buckwheat, Opium,
Central & Western Largest and oldest
Several Brassica spp.,
1. Chinese Centre China, adjacent Maize, Cowpea, centre; exceptional
Several Allium spp.,
(East Asia) lowlands, Korea, Turnip, Sesame diversity in temperate
Brinjal, Some Cucurbita
Taiwan fruits and vegetables.
spp., Pear, Peach,
Apricot, Plum, Apple,
Orange, Chinese tea

Rice, Pigeon pea (Arhar),

Chickpea, Green gram,
Brinjal, Turnip,
Cucumber, Lettuce, Important for spices,
India, Assam, Wheat forms,
Sugarcane (S. fruits, tropical cereals
2. Hindustan Myanmar, Barley, Minor
officinarum), Several and fibre crops;
(Indian) Centre Malaysia, Java, pulses and
cotton species (G. monsoon-adapted
Sumatra cucurbits
arboreum), Hemp, Black species evolved here.
pepper, Mango, Orange,
Citrus spp., Coconut,
Banana, Turmeric

NW India (Punjab, Wheat (T. aestivum),

3. Central Asiatic Kashmir), Known as
Club wheat, Pea, Rye
Centre Afghanistan Centre;
Afghanistan, Linseed, Safflower,
rich in temperate
Tajikistan, Cotton (G. herbaceum),
 Geographical Primary Crops Secondary
Centre Significance
Area Covered Originated Centre Crops

Uzbekistan, Tian- Muskmelon, Carrot, fruits, vegetables and

Shan (China) Spinach, Onion, Almond, fibre crops.
Apricot, Pear, Grapes,
Apple (cultivated types)

Nine wheat species, Rye,

Barley, Lentil, Pea,
Known as Persian or
Alfalfa, Persian clover,
Turkey, Iraq, Syria, Asia Minor centre;
4. Near Eastern / Cabbage, Carrot, Oats, Rape, Black
Lebanon, Israel, birthplace of major
Southwest Asian Lettuce, Fig, mustard, Turnip,
Iran, cereals and
Centre Pomegranate, Apple Apricot
Transcaucasia legumes—cradle of
(wild & cultivated
early agriculture.
forms), Grapes,
Almonds, Chestnut

Emmer wheat (Triticum

dicoccum) and related
Triticum spp., Several Rich source of leafy
Spain, Italy,
Avena spp., Barley, Citrus spread, vegetables, pulses,
5. Mediterranean Greece, North
Lentil, Pea, Broad bean, Wheat oilseeds and forages;
Centre African coastal
Lathyrus spp., Chickpea, diversification important secondary
region
Multiple Brassica spp., wheat pool.
Onion, Garlic, Lettuce,
Beet

Barley (Hordeum
vulgare), Wheat (T.
durum, T. turgidum, T.
Unique for ancient
6. Abyssinian dicoccum), Sorghum (S.
Ethiopia, Eritrea, Broad bean cereal domestication,
(Ethiopian) bicolor), Bajra, Chickpea,
parts of Somalia (Vicia faba) oil crops and origin of
Centre Lentil, Linseed,
Arabica coffee.
Safflower, Sesame,
Castor, Coffee, Onion,
Okra, Finger millet

Maize, Common bean,

7. South Mexican Centre of
Southern Mexico, Pumpkin, Sweet potato, Tomato forms
& Central domestication for
Guatemala, Chillies (Capsicum spp.),
American globally important
Cotton (G. hirsutum),
 Geographical Primary Crops Secondary
Centre Significance
Area Covered Originated Centre Crops

(Mesoamerican) Honduras, Costa Arrowroot, Papaya, crops like maize, chilli

Centre Rica Guava, Avocado and beans.

Potato, Cassava, Quinoa,

Lima bean, Peanut,
Extremely high
Pineapple, Pumpkin (C.
Peru, Bolivia, Sweet potato diversity in tubers
8. South maxima), Tomato,
Ecuador, diversification, and root crops, and
American Tobacco, Rubber,
Colombia, Chile, Tomato major neo-tropical
(Andean) Centre Quinine tree, Sweet
Brazil, Paraguay evolution fruit and industrial
potato (wild types),
crops.
Egyptian cotton (G.
barbadense)

Law of homologous series in variation:

The Law of Homologous Series in Variation, proposed by Leningrad Nikolai I. Vavilov (1920),
states that genetically related species exhibit parallel or repetitive patterns of hereditary
variation.

Vavilov postulated this law of homologues series and this law states that characters found in
one species also occur in other related species.

In wheat diploid (2x), tetraploid (4x) and hexaploid (6x) wheats show a series of identical
contrasting characters.

If a trait is missing in one species, but present in related species, it can often be found by
exploring its centre of origin or diversity, where maximum variability occurs.

Triticeae (Wheat, Barley, Oats) Awn length, grain colour, spike morphology

Solanaceae (Potato, Tomato, Brinjal, Pepper) Fruit size, pigmentation, disease resistance

Significance

• Serves as a predictive tool for locating unknown or missing traits in related taxa.

• Helps identify centres of origin and diversity through comparative trait mapping.

• Strengthens theoretical understanding of evolutionary relationships and genetic

parallelism.
 • Facilitates resistance breeding by identifying wild relatives as gene donors (e.g., rust
resistance in wheat, late blight resistance in potato).

Applications in Modern Plant Breeding

• Used in creation of core and mini-core germplasm collections (e.g., ICAR-NBPGR).

• Guides resistance breeding by identifying related species as gene donors (e.g., for
rust in wheat or late blight in potato).

• Supports predictive gene mining for abiotic stress, nutritional traits and climate-
resilient breeding.

Role of N. I. Vavilov in Identifying Centres of Origin:

• Proposed the concept of Centres of Origin/Centres of Diversity, stating that crops

originate in specific geographic regions where maximum genetic variability and wild
relatives are found.
• Established the principle that greatest diversity indicates the place of origin,
providing a scientific criterion for locating crop origins.
• Identified eight primary centres of origin (later expanded) and associated them with
specific crop groups, laying the foundation for crop evolutionary studies.
• Formulated the Law of Homologous Series in Variation, showing that related species
exhibit parallel patterns of variation, enabling prediction of useful traits.
• Emphasized the importance of wild relatives as reservoirs of genes for disease
resistance, abiotic stress tolerance, and quality traits.
• Laid the foundation of modern germplasm conservation, leading to the
establishment of organized gene banks and ex situ conservation strategies.
• Provided a theoretical and practical framework for plant breeding, guiding breeders
to target centres of origin for genetic resource collection.

Origin and Domestication of Rice:

• Rice (Oryza sativa) is believed to have originated in the Indo-Burma, South China,
and Southeast Asian region, as recognised by Vavilov and confirmed by genomic
evidence.

• Archaeological remains (8000–10,000 years old) from the Yangtze River Valley
(China) and Ganga Basin (India) support early domestication.

Pathway of Domestication

1. Wild progenitors: Oryza rufipogon, O. nivara

2. Early selection for domestication traits: Non-shattering panicles, Reduced seed

dormancy, larger grains and improved yield
 3. Modern improvement: Incorporation of dwarfing gene sd-1, Improved lodging
resistance and nitrogen use efficiency

Current Diversity in Cultivated Rice

• Indica: Long-grained, tropical adaptation (India, Southeast Asia)

• Japonica: Short-grained, temperate adaptation (Japan, China)

• Aromatic group: Includes Basmati and Jasmine rice

Breeding Value of Wild Relatives

Species Useful Traits

Bacterial blight resistance (Xa genes), salinity tolerance, pest

O. officinalis, O. minuta
resistance

O. barthii, O.
Drought tolerance and weedy rice competitiveness
longistaminata

Plant Introduction

Plant introduction is the intentional transfer of plant genotypes (varieties, wild relatives, or
new crop species) from their native or existing areas to new agro-ecological environments
where they were not previously cultivated, for cultivation, evaluation, or use in crop
improvement.
Introduction may occur between countries or within a country.

Procedure of Plant Introduction

1. Procurement – Collection of seeds or propagules from indigenous or foreign sources

with complete passport data (origin, taxonomy, ecology).

2. Quarantine – Mandatory inspection and treatment under the Destructive Insects and
Pests Act (1914) to prevent entry of exotic pests and diseases.

3. Cataloguing – Assignment of Plant Introduction (P.I.) numbers and systematic

documentation.

4. Evaluation – Multilocational testing for adaptability, yield, quality, and resistance to

stresses.

5. Multiplication – Controlled increase of selected material maintaining genetic purity.

6. Distribution – Supply to breeders, research institutions, or farmers for direct

cultivation or hybridization.
Advantages of Plant Introduction

• Introduces entirely new crops or superior varieties

• Provides novel genes for disease resistance, stress tolerance, and quality
improvement

• Rapid and economical method of crop improvement

• Essential for germplasm conservation and prevention of genetic erosion

• Enables cultivation in disease-free regions

Disadvantages of Plant Introduction

• Introduction of exotic weeds (e.g., Eichhornia crassipes)

• Introduction of diseases (e.g., late blight of potato)

• Introduction of insect pests (e.g., potato tuber moth)

• Ornamentals becoming invasive weeds (e.g., Lantana camara)

• Potential ecological imbalance (e.g., excessive water use by Eucalyptus)

Examples of Plant Introduction

• Primary introduction: Sonora-64 and Lerma Roja wheat; IR-8, IR-36 rice

• Secondary introduction: Kalyan Sona and Sonalika wheat (from CIMMYT material)

• Within India: Rice in Punjab, wheat in West Bengal, grape in Haryana

Institutions Involved in Plant Introduction in India

• National Bureau of Plant Genetic Resources (NBPGR) – National authority for

germplasm introduction, quarantine, and conservation

• Botanical Survey of India (BSI) – Exploration, identification, and documentation of

plant resources

• Forest Research Institute (FRI) – Introduction and acclimatization of exotic forest tree
species

Gene Pool:

A gene pool refers to the total genetic variability present in a population or species. For
plant breeding and crop improvement, understanding the gene pool helps determine the
accessibility of desirable genes such as disease resistance, yield, stress tolerance, and quality
traits.
Harlan & de Wet, 1971 classified the gene pool into three major categories based on the
ease of gene transfer and reproductive compatibility.

1. Primary Gene Pool (GP-1)

This includes individuals belonging to the same species or forms that are freely
interbreeding. Crossing within this group results in normal meiosis and fully fertile hybrids.
The gene transfer is easy and does not require special techniques.
Examples: Cultivated rice (Oryza sativa) cultivars, bread wheat varieties (Triticum
aestivum).

Significance: Most commercial crop improvement programs operate within GP-1 due to high
compatibility and stable inheritance.

2. Secondary Gene Pool (GP-2)

This consists of species that are closely related to the cultivated forms. Although crossing is
possible, hybrids often show reduced fertility, chromosomal irregularities, and may require
repeated backcrossing or techniques like embryo rescue.

Examples: Triticum dicoccum with wheat, Oryza rufipogon with cultivated rice.

Significance: Valuable source of traits like disease and pest resistance, but requires semi-
advanced breeding approaches.

3. Tertiary Gene Pool (GP-3)

This group includes distantly related wild species, where hybridization with cultivated
species is difficult or may result in complete sterility.

Gene transfer from this pool demands advanced biotechnological tools such as protoplast
fusion, genetic engineering, chromosome manipulation, and bridge crosses.

Examples: Aegilops spp. for wheat rust resistance, Oryza minuta for rice stress tolerance.

Significance: Critical for long-term crop resilience, especially against emerging biotic and
abiotic stresses.

Passport data of genetic resources

Passport data refers to the initial and essential documentation collected at the time of
acquiring, exploring, or conserving a biological sample, particularly plant germplasm. It acts
as the unique identity record (similar to an Aadhaar card) for an accession and remains
associated with it throughout its existence in genebanks, research programs, and breeding
pipelines.

Institutions like the ICAR-National Bureau of Plant Genetic Resources (NBPGR) use
standardized global formats such as Multi-Crop Passport Descriptor (MCPD) developed by
FAO and Bioversity International for data interoperability.
Genetic Erosion

• Modern agriculture favours pureline varieties (self-pollinated crops) and hybrids

(cross-pollinated crops).

• This leads to Loss of local or land races (desi varieties) and open-pollinated types —
both important reservoirs of variability. Destruction of wild species and breeding
grounds due to cultivation and grazing. Elimination of wild relatives by weedy or
cultivated forms.

• The gradual loss of variability in cultivated and wild relatives is termed genetic
erosion. This variability evolved naturally over long periods and, if lost, cannot be
restored quickly.

Germplasm conservation:

The sum total of hereditary material (genes) present in a species is called its germplasm. A
germplasm collection is a collection of a large number of genotypes of a crop species and its
wild relatives. Germplasm collections are also known as gene banks or world collections
(when large enough to include global genotypes).

Importance

• Regular crossing with wild relatives has played a key role in the evolution of crop
plants. Germplasm collections furnish the richest source of variability.

• Crop improvement depends on utilizing this variability in breeding programmes.

Types of germplasm conservation

Germplasm conservation refers to the preservation of genetic material (seeds, tissues, or

whole plants) to ensure the availability of genetic variability for crop improvement and
biodiversity protection.
It can be broadly classified into in situ (on-site) and Ex situ (off-site) conservation.

In situ conservation

• Conservation of plant species in their natural habitats or ecosystems, where they

continue to evolve under natural selection.

• Maintains natural evolutionary processes, gene flow, and ecological balance.

1. Gene Sanctuaries: Specific protected areas where wild relatives of cultivated plants are
preserved in their natural habitats. Citrus gene sanctuary in Meghalaya and Musa
(banana) gene sanctuary in Meghalaya.

2. Biosphere Reserves: Large, multi-purpose conservation areas that combine ecosystem

protection with sustainable use of biodiversity. Include core, buffer, and transition zones.
Example: Nilgiri Biosphere Reserve, Nanda Devi Biosphere Reserve.
3. National Parks and Wildlife Sanctuaries: Protect wild flora and fauna in their natural
environments. Help conserve wild crop relatives and their natural gene pools. Example:
Kaziranga National Park, Silent Valley, Gir Sanctuary.

4. On-farm Conservation: Local or traditional farmers cultivate and maintain landraces, folk
varieties, and local cultivars in their fields. Promotes dynamic conservation, as these
varieties continue evolving under farmer selection. Example: Traditional rice varieties in
Odisha and Kerala and Millets in Karnataka and Tamil Nadu.

5. Home Gardens: Common in tropical and subtropical regions (e.g., Kerala, Northeast
India). Farmers grow a diverse mix of crops (fruits, vegetables, spices, medicinal plants)
near homes, ensuring species and varietal diversity.

Advantages of In Situ Conservation

• Maintains natural evolutionary processes and gene flow.

• Conserves co-adapted gene complexes (naturally balanced traits).

• Preserves ecosystem stability and biodiversity.

• Enables continuous adaptation to changing environments.

Limitations of In Situ Conservation

• Requires large land areas and long-term protection.

• High management cost and need for continuous monitoring.

• Vulnerable to natural disasters, climate change, and human interference

(deforestation, encroachment).

• Difficult to access conserved material for breeding or research.

Ex situ conservation

Conservation of genetic resources outside their natural habitats under controlled conditions.
Ideal for immediate use in breeding programs and long-term storage.

I. Gene bank: A Gene Bank is a repository for storing germplasm (seeds, tissues, DNA,
pollen, etc.) of different species for long-term conservation and future use. It maintains
viable samples under controlled temperature and humidity to minimize deterioration. The
goal is to preserve genetic variability of crop species and their wild relatives for use in
breeding and research.
 Examples / Suitable
Type Material Stored
Crops

Seed Gene Orthodox seeds stored under low temperature (-18°C)

Wheat, rice, pulses
Bank and low humidity

Field Gene Coconut, banana,

Living plants maintained in fields or plantations
Bank sugarcane

In vitro Gene Tissue culture or cell suspension under controlled lab

Potato, yam, banana
Bank conditions

Cryo Gene Plant cells, embryos, pollen, or shoot tips stored in Cassava, coffee,
Bank liquid nitrogen (-196°C) banana

DNA / Pollen DNA, pollen grains, or genomic extracts stored for Mango, cocoa, fruit
Bank research and molecular breeding crops

1. Seed Gene Banks

• Most common and economical method. Viable seeds are dried to low moisture (3–
7%) and stored at low temperature (-18°C to -20°C).
• Suitable for orthodox seeds that can withstand drying and freezing (e.g., wheat, rice,
maize). Seeds are the most efficient propagating unit of plants.
• By controlling temperature and moisture, the metabolic activity of seeds can be
slowed down, extending their viability and life span.
• Hence, seed gene banks aim to store seeds in dormant condition to prevent
germination and deterioration.
• Seeds differ in their tolerance to desiccation (drying) and low temperature, and are
categorized into Orthodox and Recalcitrant types (Roberts, 1973).

2. Field Gene Banks

• Used for species with recalcitrant seeds or vegetative propagation.

• Plants are grown and maintained as living collections in fields.

• Examples: Banana, coconut, sugarcane, cassava, coffee.

Advantages: Easy observation of plant traits and breeding access. Maintains entire plant
structure and genetic diversity.

Limitations: High maintenance cost (irrigation, pest control). Susceptible to pests, diseases,
and environmental changes.
3. In Vitro Conservation (Tissue Culture)

Uses tissue or cell cultures maintained under slow-growth, sterile conditions in laboratories.
Suitable for vegetatively propagated and recalcitrant-seeded crops (e.g., potato, yam,
banana, garlic). Can be combined with cryopreservation for long-term storage.

Advantages: Space-efficient and protected from field hazards. Rapid multiplication and
recovery of plants when needed.

Limitations: Requires specialised infrastructure and technical expertise. Risk of somaclonal

variation (genetic change in culture).

4. Cryopreservation

Ultra-low temperature (-196°C) preservation using liquid nitrogen. Plant material (cells,
pollen, embryos, shoot tips) remains metabolically inert. Used for very long-term storage
(decades to centuries).

Advantages: Very long-term and stable conservation. Minimal maintenance after freezing.

Limitations: High setup cost and technical complexity. Limited to specific tissues or species.

5. Pollen Banks

• Stores genetic material at molecular level (pollen grains).

• Useful for recalcitrant species and biotechnological research.

• Example: Pollen of coconut and cocoa stored for breeding programs.

Advantages: Requires very little space. Ideal for gene-level studies and hybridization
programs.

Limitations: Cannot regenerate entire plants directly from DNA/pollen.

6. DNA Bank

A DNA Bank is a specialized repository that stores purified DNA samples, cloned genes, or
genomic libraries from various plant species. It allows molecular-level conservation of
genetic information, independent of viability of seeds or tissues. It serves as a
complementary system to traditional gene banks.

Institute Location Major Facilities

National gene bank for crop germplasm; seed,

National Bureau of Plant
New Delhi tissue, DNA, cryo-banks; coordinates
Genetic Resources (NBPGR)
exploration and documentation.
Institute Location Major Facilities

Conserves dryland crops (sorghum, pearl

ICRISAT Gene Bank Hyderabad
millet, chickpea, pigeon pea, groundnut).

Central Institute for

Research on Cotton Mumbai Cotton germplasm conservation.
Technology (CIRCOT)

Central Plantation Crops Kasaragod,

Field gene banks for coconut, arecanut, cocoa.
Research Institute (CPCRI) Kerala

National Centre for Plant Molecular-level conservation; plant genome

New Delhi
Genome Research (NIPGR) sequencing and DNA banking.

National Cryo-preservation Cryo-preservation of vegetatively propagated

New Delhi
Facility (NBPGR) crops.

National Facility for Plant

NBPGR, New In vitro conservation of elite and endangered
Tissue Culture Repository
Delhi species.
(NFPTCR)

II. Botanical Gardens and Arboreta

Maintain living collections of diverse plant species for education, research, and conservation.
Examples: Indian Botanic Garden, Howrah and National Orchidarium, Yercaud (Tamil Nadu).

Advantages: Promote public awareness and scientific study and serve as ex situ repositories
for rare and threatened species.

Limitations: Limited species number and high maintenance cost, and not suitable for long-
term genetic preservation.

Germplasm Utilization

Germplasm utilization is the strategic use of genetic resources—landraces, wild relatives,

improved varieties and engineered lines—to develop superior crop varieties through
systematic evaluation, selection and breeding.

Importance of Germplasm Utilization

• Provides raw material for breeding new varieties.

• Ensures genetic diversity, reducing genetic vulnerability.

 • Helps in the development of crops with Higher yield, Stress tolerance (drought,
salinity, heat), Pest and disease resistance, Improved nutritional quality.

• Essential for achieving climate resilience and sustainable agriculture.

Categories of Germplasm Used in Breeding

Type of Germplasm Examples Use in Breeding

Landraces: Locally evolved and

Indigenous farmer-saved Base material for resistance
genetically diverse; useful for stress
varieties and adaptation
tolerance and adaptive traits.

Wild Relatives: Provide lost or rare

Wild rice, teosinte (maize Source of genes for
genes for disease, pest and abiotic
ancestor) resistance/tolerance
stress resistance.

Obsolete Varieties: Once widely

Unique alleles and stability
grown; contain useful but lost alleles Old high-yielding varieties
traits
valuable for modern breeding.

Modern Cultivars: High-performing

cultivars with desirable agronomic HYVs, hybrids Direct use or donor parents
traits.

Mutants: Variation created artificially; Novel traits (e.g., dwarfism in

Induced/spontaneous
helps introduce new useful traits. wheat & rice)

Genetically Engineered Lines:

Developed via biotechnology for
Bt cotton, Golden rice Targeted trait improvement
specific traits like pest resistance or
nutrition.

Methods of Utilizing Germplasm

Direct Utilization: Landraces or old varieties used directly as cultivars. Some basmati rice
landraces still grown.

Indirect Utilization: Germplasm used as parents in breeding programs. Wild species transfer
disease resistance genes.

Pre-breeding: Introgression of beneficial genes from wild or unadapted materials into usable
breeder lines.

Challenges in Germplasm Utilization

 • Poor characterisation and documentation.

• Crossability barriers with wild species.

• Undesirable linkages (linkage drag).

• Long breeding cycles and Genetic erosion due to monocropping and modern
agriculture.

Modes of reproduction, selfing, and crossing techniques

1. Discuss the relevance of mode of reproduction in plant breeding. (12.5M CS-2014)

2. Describe self-pollination, cross-pollination and often cross-pollination. Discuss different

mechanisms for facilitating self-pollination and cross-pollination. (20M CS-2021)

3. Synchrony in flowering and planting ratio of parents. (15M IFS-2017)

4. Gametogenesis and sporogenesis. (10M IFS-2016)

5. Explain the mechanisms which promote cross-pollination. (8M IFS-2021)

MODES OF REPRODUCTION

The mode of reproduction determines whether a crop is naturally homozygous or

heterozygous, which directly influences the breeding objective and method.

• In self-pollinated crops (e.g., wheat), plants are largely homozygous, so breeding

aims to develop and maintain pure, uniform homozygous lines.

• In cross-pollinated crops (e.g., maize), plants are naturally heterozygous, so breeding

focuses on maintaining or creating genetically diverse heterozygous populations or
hybrids.

The modes of reproduction in crop plants may be broadly grouped into two categories:
asexual and sexual.

I. Asexual Reproduction

Asexual reproduction does not involve the fusion of male and female gametes. New plants
may develop from vegetative parts of the plant (vegetative reproduction) or may arise from
embryos that develop without fertilisation (apomixis).

Vegetative Reproduction
In nature, a new plant develops from a portion of the plant body. This may occur through
modified underground and sub-aerial stems, and through bulbils.

Underground Stems

The underground modifications of the stem generally serve as storage organs and contain
many buds. These buds develop into shoots and produce plants after rooting. Examples of
such modifications are given below.

Tuber: Potato, Bulb: Onion, Garlic, Rhizome: Ginger, turmeric

Sub-aerial Stems

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

Bulbils

Bulbils are modified flowers that develop into plants directly without the formation of seeds.
These are vegetative bodies; their development does not involve fertilisation and seed
formation. The lower flowers in the inflorescence of garlic naturally develop into bulbils.

Artificial Vegetative Reproduction

Vegetative reproduction is widely used for multiplying many crop species, even when it does
not occur naturally in them. Several commercial methods are Stem cuttings: for crops like
sugarcane, grapes, and rose.

• Layering, budding and grafting: commonly used in fruit crops and ornamental
shrubs.

• Tissue culture (micropropagation): enables rapid multiplication in many species and

is being further developed for others.

Although these crops can reproduce sexually, vegetative propagation is preferred because it
preserves the genetic uniformity and desirable traits of the parent plant.

Significance of Vegetative Reproduction:

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

Apomixis

In apomixis, seeds are formed, but the embryos develop without fertilisation. Consequently,
the plants resulting from them are identical in genotype to the parent plant.
In apomictic species, sexual reproduction is either suppressed or absent (generally
facultative).

When sexual reproduction does occur, the apomixis is termed as facultative.

But when sexual reproduction is absent, it is referred to as obligate. The details of apomictic
reproduction vary so widely that a confusing terminology has resulted.

classification of apomixis.

Adventive Embryony

In this case, embryos develop directly from vegetative cells of the ovule, such as nucellus,
integument. Development of embryo does not involve production of embryo sac. Adventive
embryony occurs in mango, citrus, etc.

Apospory

Some vegetative cells of the ovule develop into unreduced embryo sacs after meiosis. The
embryo may develop from egg cell or some other cell of the embryo sac. Apospory occurs in
some species of hawkweed and ferns.

Diplospory

Embryo sac is produced from the megaspore, which may be haploid or, more generally,
diploid. Generally, the meiosis is so modified that the megaspore remains diploid. Diplospory
leads to parthenogenesis or apogamy.

Parthenogenesis:

Asexual reproduction in which a new individual develops from an unfertilized egg (ovule)
without the involvement of male gametes or fertilization. Offspring are usually genetically
identical (haploid or diploid) depending on the species. It is of two types

Gonial parthenogenesis – embryos develop from an egg cell,

Somatic parthenogenesis – embryos develop from any cell of the embryo sac other than the

egg cell.

Apogamy

In apogamy, synergids or antipodal cells develop into an embryo. Like parthenogenesis,

apogamy may be haploid or diploid depending upon the haploid or diploid state of the
embryo sac. Diploid apogamy occurs in Allium and many other plant species.

Significance of Apomixis

• Fixation of Hybrid Vigour (Heterosis): Apomixis allows hybrids to reproduce true-to-

type, enabling the perpetual maintenance of hybrid vigour across generations.
 • Genetic Uniformity: Produces genetically identical progeny, maintaining the
desirable genotype without segregation or variation.

• Stability of Traits: Traits such as yield, quality, and resistance remain stable over
successive generations.

• Cost-effective Seed Production: Farmers can save and reuse seeds without loss of
performance, reducing dependency on repeat hybrid seed purchase.

• Avoids Inbreeding Depression: Maintains heterozygosity and prevents reduction in

vigour seen in sexually reproducing heterozygous crops.

• Useful in Breeding Programs: Once a superior genotype is developed, apomixis helps

preserve and multiply it efficiently.

• Propagation of Sterile Hybrids: Useful in crops where sexual reproduction is difficult

or seeds are sterile (e.g., some triploids).

• Environment-independent Reproduction: Reproduction is not dependent on

pollination, making seed formation possible even under adverse conditions.

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
specialized structures known as flowers.

Flower

A typical flower consists of sepals, petals, stamens, and pistil.

• A flower with both stamens and pistil is called a perfect or hermaphrodite flower.

• A flower with only stamens is staminate (male), while one with only a pistil is
pistillate (female).

• In monoecious plants (e.g., maize, castor, coconut, Colocasia), staminate and

pistillate flowers occur on the same plant.

• In dioecious plants (e.g., papaya, date palm, pistachio), male and female flowers
occur on separate plants.

In crop plants, meiosis in the stamen produces microspores, and meiosis in the pistil
produces megaspores. Subsequent mitotic divisions form the male and female gametes,
respectively.

1. 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 degenerate leaving one functional megaspore per ovule. This completes
megasporogenesis.

2. Gametogenesis

The production of male and female gametes in the microspores and the megaspores,
respectively, is known as gametogenesis.
Microgametogenesis.

It refers to the formation of male gametes (sperms) in flowering plants. During pollen
maturation, the microspore nucleus divides mitotically to form two nuclei — a generative
nucleus and a vegetative (tube) nucleus. Pollen is usually released at this binucleate stage.

After pollination, the pollen germinates on the stigma and forms a pollen tube. As the tube
grows through the style, the generative nucleus divides mitotically to produce two male
gametes (sperms). The pollen grain along with its tube constitutes the microgametophyte.

The pollen tube finally enters the ovule through the micropyle and releases the two sperms
into the embryo sac for fertilization.

Megagametogenesis.

Megagametogenesis is the process by which a functional megaspore develops into the

female gametophyte (embryo sac). The nucleus of the megaspore undergoes three mitotic
divisions, forming eight nuclei.

These nuclei become organized as follows

• Three nuclei move to the micropylar end forming One egg cell and Two synergids

• Three nuclei migrate to the chalazal end and form three antipodal cells.

• The remaining two nuclei stay in the centre as polar nuclei, which later fuse to form
the diploid secondary nucleus.

Thus, the mature embryo sac typically contains 1 egg cell, 2 synergids, 3 antipodal cells, 1
diploid secondary nucleus

This entire development of the embryo sac from a megaspore is known as

megagametogenesis.
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.

MODES OF POLLINATION

Pollination refers to the transfer of pollen grains from anthers to stigmas. Pollen from an
anther may fall on the same flowers, leading to "self-pollination" or " autogamy". When
pollen from flowers of one plant are transmitted to the stigmas of flowers of another plant,
it is known as " cross pollination" or allogamy.

A third situation " Geitonogamy" results when pollen from a flower of one plant falls on the
stigmas of other flower of the same plant e.g. maize.

SELF POLLINATION

Autogamy is the transfer of pollen from the anther to the stigma of the same flower. It is the
closest form of inbreeding and results in high homozygosity.

Species that predominantly self-pollinate become genetically stable and homozygous and
therefore show little or no inbreeding depression.

Autogamy occurs only in crops with hermaphrodite flowers, although a small degree of
cross-pollination (up to ~5%) may still occur depending on the crop variety, temperature,
humidity, and environmental conditions.
Several natural mechanisms exist to promote self-pollination, ensuring genetic uniformity in
such crops.

1. CLEISTOGAMY - In this case, flowers do not open at all. This ensures complete self-
pollination. Since, foreign pollen cannot reach the stigma of closed flowers. It occurs in some
varieties of wheat (Triticum sp.) Oats (Avena sp.), Barley (Hordeum vulgare) and in a
number of other grasses.

2. CHASMOGAMY - In some sp. the flower open only after pollination has taken place. This
occurs in many cereals, such as wheat, barley, rice and oats.

3. Position of Anthers.
In some species, stigmas are surrounded by anthers in such a way that self-pollination is
ensured. Such a 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. Pollination generally occurs after the
flowers open. But the position of anthers in relation to the stigma ensures self-pollination.

4. In a few sp. stigmas become receptive and elongate through stamina columns. This
ensures pre-dominant self-pollination.

5. 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.

6. 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.

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 in breeding
depression, but may exhibit considerable heterosis. Therefore, the aim of breeding methods
generally is to develop homozygous varieties.

CROSS-POLLINATION:

In cross-pollination species, the transfer of pollen from a flower the stigmas of the other
may be brought about the wind (anemophilic), water (hydrophilic) or insects (entomophilic).
This is common form of outbreeding and leads to heterozygosity. Many of the crop plants
are naturally cross-pollinated. In many sp. A small amount (up to 5-10 per cent) of selfing
may occur. There are several mechanisms that facilitate cross-pollination. Various
mechanisms are,

1. Dicliny. It refers to unisexual flowers. This is of two types: viz.

i) monoecy. When male and female flowers are separate but present in the same plants. 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, strawberries, cassava and rubber.

ii) dioecy 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. 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 in linseed.

4. Herkogamy. Hindrance to self-pollination due to some physical barriers such as presence

of a 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 non-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.

Genetic consequences of cross-pollination

Cross-pollination preserves and promotes heterozygosity in a population. Cross- pollinated

sp. 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.

OFTEN CROSS-POLLINATED SPECIES:

In many crop plants, cross pollination often exceeds 5 per cent and may reach 30 percent.
Such sp. are generally known as " often cross-pollinated crops" e.g. Sorghum, cotton,
redgram and safflower etc.,

The genetic architecture of such crops is intermediate between self-pollinated and cross-
pollinated sp. Consequently, in such species breeding methods suitable for both of them
may be profitably applied. But often hybrid varieties are superior to others.
 [Link] Self pollination Cross pollination

1. Self-pollination leads to a very rapid Cross-pollination preserves and

increase in homozygosity. Therefore,
promotes heterozygosity in a
populations of self–pollinated species
population. Cross-pollinated species are
are highly homozygous. highly heterozygous and show mild to
severe inbreeding depression and a
considerable amount of heterosis.

2. Self-pollinated species do not show The breeding methods in such species

inbreeding depression, but may exhibit aim at improving the crop species

considerable heterosis. without reducing heterozygosity to an

appreciable degree.

3. The aim of breeding methods Usually, hybrid or synthetic varieties are

Generally is to develop homozygous the aim of a breeder wherever the seed

varieties. The inbreeding mechanisms production of such varieties is

are generally under precise genetic economically feasible.

control, but can be influenced by both

the genetic background as well as the

environment.

Pure-line selection, pedigree, mass and recurrent selections, Combining ability, its
significance in plant breeding.

1. Define pureline. Discuss the relevance of pureline selection in self- and cross-pollinated
crops. (12.5) CSE-2014

2. Define 'purelines' along with their characteristics. How are they used in crop
improvement? (10) CSE-2016

3. Define mass selection. Explain its applications in self-pollinated crops. (10m)

4. Define the pedigree method and give its importance in crop improvement, with suitable
examples. (15) CSE-2019

5. Write the various methods of plant breeding and describe the backcross method with its
importance in the development of crop varieties. (20) CSE-2019
6. What is mass selection? Discuss its applications in varietal improvement. (10) CSE-2020

7. Describe the advantages and disadvantages of mass selection, simple recurrent selection
and clonal selection. (10) CSE-2021

8. What are the applications of mass selection in self-pollinated crops? Describe its merits
and demerits (10) CSE-2022

9. Describe different approaches for improving the characteristics of inbred lines. (10) CSE-
2024

10. Give an account of the bulk method of breeding. Discuss its merits and demerits. (10)
CSE-2024

11. Describe in brief various schemes suggested to improve characteristics of inbred lines.
(10) IFS-2015

12. Explain the breeding methods of rice crop. (8) IFS-2019

13. Pedigree and Bulk method of plant breeding. (10) IFS-2019

14. Write the various methods of plant breeding. Describe backcross method with its
importance in development of crop varieties. (15) IFS-2020

15. Describe backcross method of plant breeding and its application in development of biotic
stress tolerant or resistant varieties with suitable examples. (15) IFS-2022

16. Explain the method of recurrent selection. Write the importance of combining ability in
crop improvement. (15) IFS-2022

17. Explain the methods for creating the variability. How are the types of selection
responsible for population improvement in crop plants? (15) IFS-2022

18. Write the breeding objectives for improvement of wheat and gram. Describe the
pedigree method of breeding for crop improvement. (15) IFS-2023

19. Give a concise account on merits and demerits of mass selection. (8) IFS-2024

20. Define general combining ability. Describe the procedure for recurrent selection for
general combining ability. (12.5) CSE-2014

21. Define 'general combining ability'. Explain the genetic basis and methods for estimating
SCA in a diallel mating system. (12.5) CSE-2016

SELECTION IN SELF-POLLINATED CROPS

Selection involves permitting reproduction only in those plants that possess desirable
characteristics. The next generation is raised from the seeds of selected plants only; seeds of
other plants are rejected.

Basis of selection: Plant phenotype (visible traits).

Effectiveness of Selection

• The success of selection depends on how accurately the phenotype reflects the
genotype.

• Two main characteristics/limitations:

1. Effective only for heritable differences — depends on the heritability of the

trait.

2. Does not create new variation — it only utilizes existing variation within the
population.

Requirements for Effective Selection

1. Presence of variation in the population.

2. Heritability of that variation (traits must be genetically controlled).

Purpose

• To isolate and multiply desirable plant types from the population.

• Selection is fundamental to crop improvement, forming one of the two basic steps
in any breeding programme:

1. Creation of variation

2. Selection of desirable types

THE PROGENY TEST

Progeny test = Evaluation of the worth or breeding value of a plant based on the
performance of its progenies.

Developed by: Louis de Vilmorin, hence also known as the Vilmorin Isolation Principle or
Vilmorin Principle.

Vilmorin’s Experiment (Sugarbeet)

• Selected sugarbeet plants with high sugar content.

• Classified them based on progeny performance:

1. Progenies high in sugar content

 2. Progenies mixed (some high, some low)

3. Progenies low in sugar content

Plants similar in phenotype may produce different progenies. Therefore, the true value of a
plant can be known only through its progeny.

Functions of the Progeny Test

1. Determines Breeding Behaviour: Reveals whether a plant is homozygous or

heterozygous. e.g. In barley, both VV and Vv have two-rowed spikes, but their
progeny segregation reveals genetic constitution.

2. Determines Heritability of the Character: Shows whether a selected trait is

genetically controlled (heritable) or environmental. If differences persist in progeny
→ genetic, If differences disappear in progeny → environmental

Significance

• Basis of all modern breeding methods.

• Ensures selection for true genetic differences rather than environmental effects.

THE PURELINE THEORY

A pureline refers to the progeny of a single homozygous plant in a self-pollinated species,

where all individuals in the line are genetically identical. A pureline is a genetically uniform
population that exhibits no genetic variation within the line, with any observed variation
being purely environmental.

Characteristics of a Pureline:

1. Genetic Uniformity: All individuals within a pureline share the same genotype.

2. Environmental Variation: The variation seen in purelines is environmental rather

than genetic, and it is non-heritable.

3. Homozygosity: Purelines are characterized by a high degree of homozygosity (AA,

aa), with an absence of heterozygosity (Aa) as the result of prolonged self-pollination.

4. No New Genetic Variation: Over time, genetic diversity is lost, and the population
becomes uniform.

5. Phenotypic Variation: Despite being genetically identical, phenotypic variation may

still be observed due to environmental factors.

Johannsen’s Experiment (1903)

Wilhelm Johannsen's groundbreaking work on the pureline theory was based on his
experiments with Phaseolus vulgaris (bean). He introduced the concept through the
following key observations:

1. Variation in Seed Size: Johannsen observed variation in seed size within a

commercial seed lot.

2. Selection and Progeny Growth: He selected large, medium, and small seeds and
grew them separately. Progenies from large seeds produced large-seeded plants,
while progenies from small seeds produced small-seeded plants.

3. Genetic Basis of Variation: The variation in the original lot was genetic, indicating
that different purelines were present within the mixture.

4. Pureline Formation: Johannsen raised 19 progeny lines, each derived from a single
seed. Each line exhibited distinct mean seed weights, and within each pureline,
variation was minimal, attributed only to environmental factors.

5. Conclusion: He concluded that variation within a pureline is environmental and non-

heritable, while variation between purelines is genetic and heritable. Thus, pureline
selection is effective only between purelines, not within them.

Relevance in Crop Breeding of Self-Pollinated Crops

Pureline theory plays a fundamental role in the breeding of self-pollinated crops. By

repeatedly self-pollinating a plant, homozygosity increases, leading to the establishment of a
genetically uniform population. The following points illustrate its significance:

1. Genetic Uniformity: Purelines, once established, are genetically identical, ensuring

uniformity in traits, making them ideal for true-breeding crops.

2. Fixation of Desirable Traits: Traits of interest (e.g., disease resistance, high yield, or
specific quality traits) can be stabilized through pureline selection, as the population
becomes genetically fixed.

3. Homozygosity and True-Breeding: Continuous selfing leads to a population that

breeds true, where offspring exhibit the same traits as the parent, providing
consistency in crop production.

Example: In wheat breeding, pureline selection is used to develop stable, high-yielding

varieties. Over multiple generations of self-pollination, a uniform population is achieved,
making it possible to select for desirable traits such as disease resistance or specific grain
quality.

Relevance in Crop Breeding of Cross-Pollinated Crops

In cross-pollinated crops, the application of pureline theory is more challenging due to the
inherent genetic variability introduced by cross-pollination. However, pureline theory still
has relevance in the following ways:

1. Pureline Hybridization: The concept of purelines is used in hybrid breeding

programs. In these programs, purelines of different genetic backgrounds are crossed
to create hybrids with superior traits (hybrid vigor).

2. Selection of Parental Lines: While cross-pollination introduces genetic diversity, the

selection of purelines for hybrid production ensures that the desired traits are fixed
in the parental lines. These lines serve as stable parents that, when crossed, produce
high-yielding hybrids.

Example: In maize breeding, purelines are developed and then crossed to create hybrids.
The purelines serve as genetically uniform sources of traits that, when combined, result in
hybrids with improved performance, such as increased yield and resistance to pests and
diseases.

ORIGIN OF VARIATION IN PURELINES

Although purelines are genetically uniform, they may show variation over time. This
variation arises due to mechanical mixture, natural hybridisation, chromosomal
aberrations, and mutations.

Mechanical Mixture

• Mechanical mixture occurs when seeds of other genotypes accidentally get mixed
with the pureline during cultivation, harvesting, threshing, or storage. Such
contamination leads to the presence of off-types within the pureline.

• It can be prevented by careful handling of seeds and by practising mass selection

periodically to remove contaminants and maintain purity.

Natural Hybridization

• In most self-pollinated crops, a small amount of cross-pollination may occur under

certain environmental conditions.

• When purelines are grown near other genotypes, cross-pollination may take place,
resulting in natural hybrids.

• Subsequent selfing of these hybrids produces new genotypes, thereby introducing

variation in the pureline.

• This can be avoided by isolating the pureline with two to three border rows of the
same variety on all sides of the plot.
Chromosomal Aberrations

• In some crops like wheat, oats, and barley, meiotic abnormalities such as chromatin
bridges and chromosome fragments occur occasionally.

• These may cause duplications or deficiencies of small chromosome segments,

leading to genetic variation.

• The effects of such minor chromosomal changes are often similar to gene mutations,
and therefore, their exact contribution to variation in purelines is uncertain.

Mutation

• Mutation is a sudden, heritable change in a gene, often caused by chemical changes

within the DNA molecule.

• It is spontaneous, random, recurrent, and specific in its phenotypic effect.

Mutations are considered the primary source of new genetic variability in plant
species.

• For example, Stadler (1942) reported different mutation rates in maize genes — gene
R (colour) showed a rate of 4.92 × 10⁻⁴, gene Su (non-sugary endosperm) had 2.4 ×
10⁻⁸, while gene Wx (non-waxy endosperm) showed no mutant alleles among
1,503,744 gametes tested.

Pure-Line Selection

Pure-line selection is a method of crop improvement used in self-pollinated crops, where a

new variety is developed from the progeny of a single homozygous plant selected from a
genetically variable population.

A large number of superior-looking plants are selected from the base population and
harvested individually. Their progenies are grown in separate rows, evaluated over
successive generations, and the best performing progeny is yield tested and finally released
as a variety.

Characteristics of Pure Lines

1. All plants within a pure line possess the same genotype.

2. Variation within a pure line is environmental and non-heritable.

3. Over time, pure lines may show genetic variation due to mutation, natural
hybridization, or mechanical mixtures.

General Steps in Pure-Line Selection

 • First Season:
Select superior individual plants from the base population and harvest them
separately.

• Second Season:
Grow selected plants in progeny rows and evaluate performance; reject inferior
progenies.

• Third Season:
Repeat progeny row evaluation and selection.

• Fourth Season:
Test selected progenies in replicated preliminary yield trials along with suitable
check varieties.

• Fifth Season:
Conduct comparative yield trials and select the best culture.

• Sixth Season:
Conduct multilocation trials across different research stations with local checks.

• Seventh Season:
Conduct adaptive research trials in farmers’ fields and release the best yielder
through the Variety Release Committee.

Wheat: NP 4, Kalyansona

Rice: Adt-1, 3, 5

Groundnut: TMV 3, 4, 7, 8

Green gram (Moong) – Pusa Baisakhi

Advantages of Pure-line Selection

1. Maximum improvement over the original variety

Pure-line selection isolates the best homozygous genotype from a population,
resulting in the highest possible improvement.
Example: Wheat varieties NP-4 and NP-52 selected from heterogeneous local
populations.

2. High genetic and phenotypic uniformity

All plants in a pure-line variety have the same genotype, leading to uniform growth,
maturity, and yield.
Example: Rice variety IR-8 shows uniform plant type and maturity.

3. Ease of identification and seed certification

Extreme uniformity simplifies varietal identification and maintenance of genetic
 purity during seed production.
Example: Pure-line wheat varieties are easily distinguished during certification
compared to mass-selected varieties.

Disadvantages of Pure-line Selection

1. Limited adaptability and yield stability

Pure-line varieties are often less stable across environments than heterogeneous
local varieties.
Example: Pure-line wheat varieties may perform poorly under stress compared to
traditional landraces.

2. Time-consuming and costly process

Development requires several generations of selection and extensive yield trials.
Example: Development of Kalyan Sona wheat involved multiple years of testing
before release.

3. Improvement limited by existing genetic variability

No improvement is possible if the base population lacks sufficient genetic variation.
Example: Uniform old varieties cannot be improved further unless mutations occur.

4. Greater demand on breeder’s time and effort

Continuous selection, testing, and evaluation reduce the breeder’s ability to manage
multiple programmes.
Example: Long-term pure-line breeding programmes in self-pollinated crops like
wheat and rice.

Scientific Basis and Significance

Pure-line selection involves three major phases:

1. Selection of superior individual plants from a variable population.

2. Evaluation of progenies over several generations by observation.

3. Extensive yield trials with precise measurements to confirm superiority.

Any progeny that consistently outperforms existing varieties is released as a pure-line

variety. The success of this method in the early 20th century was largely due to the
availability of genetically diverse landraces, which served as rich reservoirs of variation.
Although its importance has declined in major crops, pure-line selection remains valuable in
less-improved and minor crop species.
Applications of Pure-Line Selection

Pure-line selection is extensively used in the improvement of self-pollinated crops,

particularly for refining local, introduced, and existing varieties and for isolating superior
genotypes.

1. Improvement of Local (Desi) Varieties

Pure-line selection is highly effective in improving local varieties that possess substantial
genetic variability. Superior homozygous lines are isolated from heterogeneous populations.

Examples:

• NP-4 and NP-52 – wheat (Triticum aestivum)

• Pusa Sawani – okra (selection from a Bihar variety with field resistance to yellow
mosaic virus)

2. Improvement of Introduced Varieties

Introduced germplasm is subjected to pure-line selection to identify genotypes better

adapted to local agro-climatic conditions and resistant to local pests and diseases.

Example:
 • Kalyan Sona – leaf rust-resistant selection from CIMMYT material (Mexico)

3. Improvement of Old Pure-Line Varieties

Old pure-line varieties may develop variability over time due to mutation or mechanical
mixtures. Pure-line selection helps in isolating new, superior lines from such populations.

4. Selection for New or Previously Unimportant Characters

Pure-line selection is used to isolate plants possessing novel or rare traits, such as disease
resistance, improved plant type, or grain quality, from an existing variety.

Example:

• A root-rot-resistant sorghum (Sorghum bicolor) variety was developed by selecting

resistant plants from the susceptible Dwarf Yellow Milo following an outbreak of
Periconia circinata in Kansas (1926).

5. Selection from Segregating Generations of Crosses

Pure lines are frequently isolated from segregating populations obtained after hybridization.
This is achieved through breeding methods such as:

• Pedigree method

• Bulk method

• Backcross method

Mass Selection

Mass selection is the earliest and simplest method of crop improvement in which a large
number of phenotypically superior plants are selected from a heterogeneous population,
and their seeds are bulked to raise the next generation. Selection is based solely on
phenotypic performance, without progeny testing.

Selection criteria include plant height, maturity, ear or panicle type, grain colour and size,
disease resistance, tillering ability, and productivity. When variation exists for seed traits
such as seed colour or size, selection is carried out before seed bulking.

The resulting population is more uniform than the original, especially for qualitative traits
controlled by few major genes (e.g., plant height, awns, seed colour), but considerable
variability remains for quantitative traits such as yield, quality, and adaptability. In self-
pollinated crops, mass-selected varieties represent a mixture of several similar pure lines,
while in cross-pollinated crops they consist of multiple genotypes. Hence, mass-selected
varieties retain genetic variability, permitting further improvement by selection.

Genetic Constitution

• Homozygous but heterogeneous (self-pollinated crops)

 • Highly heterogeneous (cross-pollinated crops)

Types of Mass Selection

1. Positive Mass Selection

Desirable plants are selected from old varieties or landraces.
Used for improvement of yield, plant type, and disease resistance.

2. Negative Mass Selection

Undesirable or off-type plants are removed from the population.
Used mainly for varietal purification in seed production and certification.

Applications and Scope

1. Improvement of Local or Desi Varieties

Local varieties are genetically diverse but well adapted and stable. Mass selection removes
inferior plant types and improves average yield, uniformity, and general appearance
without affecting adaptability.

Example: Improvement of traditional rice and wheat landraces.

2. Purification of Existing Pure-line Varieties

Pure lines become impure over time due to mechanical mixtures, natural hybridization, and
mutations. Regular mass selection helps restore genetic purity.
This is the most important present-day use of mass selection in self-pollinated crops.

3. Importance in Cross-pollinated Crops

Mass selection is especially important in cross-pollinated species, where maintaining

heterogeneity ensures buffering capacity and prevents inbreeding depression.

General Procedure of Mass Selection

• First Year:
Grow a heterogeneous base population (old variety or landrace). Select 200–2000
phenotypically superior plants and bulk their seeds.

• Second Year:
Grow bulked seed in a preliminary yield trial along with standard and/or original
variety as check. Reject inferior phenotypes.

• Third to Sixth Year:

Evaluate the bulk in coordinated or multilocation yield trials (IET → main yield trials)
for yield and adaptability.
 • Seventh Year:
If superior, multiply seed, name, and release the variety. Seed becomes available for
distribution by the eighth year.

Merits of Mass Selection

• Applicable to both self- and cross-pollinated crops

• Useful for improvement of landraces and purification of varieties

• Retains genetic variability, allowing future selection

• Varieties show wide adaptability and yield stability due to buffering effect

• Effective against disease epidemics

• Less time-consuming and less expensive than pure-line selection

• Improvement possible for traits with high heritability and few genes

• Important in forage crops and minor crops

Demerits of Mass Selection

• Varieties are less uniform than pure-line varieties

• Selection based only on phenotype; true breeding value not known

• No progeny test, so superiority may be environmental

• In cross-pollinated crops, no control over pollination, causing rapid deterioration

• Improvement is not permanent; selection must be repeated

• Traits with low heritability and polygenic control cannot be effectively improved

• Does not create new genotypes, only utilizes existing variability

• Seed certification is difficult due to heterogeneity

• In self-pollinated crops, pure-line selection is more effective

PURE VS MASS SELECTION:

[Link] Pureline selection Mass selection

1. The new variety is a pureline The new variety is a mixture of pure-

lines.
2. The new variety is highly uniform. In The variety has genetic variation for
fact the variation within pureline quantitative characters, although it is
variety is purely environmental relatively uniform in general
appearance

3. The selected plants are subjected to a The progeny test is generally not
progeny test carried out

4. The variety is generally best pureline The variety is inferior to the best
present in the original population. The pureline because of pureline included
pureline selection brings about the in it will be inferior to the best
greatest improvement over original pureline
variety

5. Generally, a pureline variety is expected Usually the variety has a wider

to have a narrower adaptation and adaptation and greater stability than
lower stability in performance than a a pureline variety
mixture of purelines.

6. The plants are selected for their The selected plants have to be similar
desirability. It is not necessary that they in phenotype since their seeds are
should have a similar phenotype. mixed to make up the new variety,

7. It is more demanding; requires detailed Less demanding; yield trials are not
progeny tests and yield trials. essential even when large
populations are used.

8. Generally, Takes about 9–10 years to Takes about 5–7 years to develop a
develop a new variety. new variety.

9. Selection within a pureline is Selection within a mass-selected

ineffective unless it has become population remains effective due to
genetically variable retained genetic variability.

10. Produce is uniform in quality. Produce is not uniform, as different

purelines may vary in yield and
quality.

11. Variety is easily identified in seed Variety is relatively difficult to

certification programmes. identify in seed certification.

12. Used mainly in self-pollinated crops, Used in both self- and cross-
occasionally in cross-pollinated ones. pollinated crops.
Bulk Method of Breeding

The Bulk Method, first used by Nilsson-Ehle (1908) at Svalöf, is a selection method
employed mainly in self-pollinated crops. It is also known as the Mass Method or Hill
Population Method. In this approach, the segregating generations (F₂ onward) are grown in
bulk, and seed from the entire population is harvested and advanced without individual
plant selection during the early generations.

Bulking Period

• Bulking is continued for 6–7 generations or even 30+ generations, depending on

crop biology and breeding objectives.

• During this period:

o Natural selection operates strongly (competition, disease pressure, climatic

adaptation).

o Artificial selection is minimal or absent.

• Genotypic frequencies change primarily due to natural selective forces.

Procedure

1. Hybridization
Selected parents are crossed to produce simple or complex crosses.

2. F₁ Generation
F₁ plants are space-planted and harvested in bulk.

3. F₂–F₆ Generations (Bulking Phase)

Populations are grown at commercial seed rates and spacings and harvested in bulk.
Natural selection modifies population structure; deliberate selection is generally
avoided.

4. F₇ Generation
Plants are space-planted.
Seeds from phenotypically superior individual plants are harvested separately.

5. F₈ Generation
Individual plant progenies are grown in single or multiple rows.
Superior progenies are selected and harvested in bulk.

6. F₉ Generation
Preliminary Yield Trials (PYT) are conducted with standard commercial checks.
Evaluation includes yield, plant height, lodging, disease resistance, flowering and
maturity.
 7. F₁₀–F₁₂ Generations
Superior lines are tested in replicated multilocation yield trials for yield stability,
adaptability, resistance, and quality traits.
Lines superior to checks are identified for release.

8. F₁₃ Generation
Seed multiplication of the selected line is undertaken.
Breeder supplies breeder seed for foundation seed production.

Key Features of the Bulk Method

• Relies primarily on natural selection in early generations

• Minimal record keeping compared to pedigree method

• Effective for selecting widely adapted genotypes

• Particularly suited to traits with high heritability and environmental stability

• Less labour-intensive but slower in eliminating undesirable types

Applications of the Bulk Method

The bulk method of handling segregating generations has three major applications in crop
improvement:

1. Isolation of Homozygous Lines

 • In the bulk method, populations are advanced without selection until a high level of
homozygosity is attained.

• Individual plant selection is initiated after the bulking period, when segregation is
largely complete.

• A Preliminary Yield Trial (PYT) can be conducted as early as the second year after
bulking.

• This approach allows isolation of homozygous lines with minimum labour, cost, and
record keeping.

Example:
Isolation of pure lines in wheat and barley after prolonged bulking.

2. Waiting for the Opportunity for Selection

• Favourable environments for effective selection do not occur every year (e.g.,
disease epidemics, drought stress).

• The bulk method allows segregating generations to be safely maintained without

loss of genetic variability, as artificial selection is usually avoided.

• When favourable conditions arise, the population can be shifted to pedigree

selection.

This combined approach is known as the Mass–Pedigree Method (Harlan).

Example:
Bulking wheat populations until a natural rust outbreak enables effective disease-resistance
selection.

3. Allowing Natural Selection to Alter Genetic Composition

• Prolonged bulking permits natural selection to act on the population under field
conditions.

• Genotypes with:

o Better adaptation

o Higher competitive ability

o Greater stress tolerance

tend to increase in frequency.

• This results in a higher probability of isolating superior lines compared to direct

selection from F₂.
 • Due to this evolutionary shift, Suneson referred to the bulk method (especially long-
term bulks) as the Evolutionary Method of Breeding.

Example:
Selection for wide adaptability in cereals grown under rainfed or marginal environments.

Merits of the Bulk Method

1. Natural selection increases the frequency of superior genotypes over time.

2. Large population size increases the probability of isolating transgressive segregants.

3. Selection is practiced after near-homozygosity, making it more effective for

quantitative traits like yield.

4. Particularly suited for small-grain cereals (wheat, barley, oats) grown at high
densities.

5. Useful for studying gene and genotype survival under natural field conditions.

6. Simple, economical, and less labour-intensive, allowing breeders to manage

multiple breeding programmes simultaneously.

Demerits of the Bulk Method

1. Effective natural selection becomes prominent only after F₁₀ or later, often requiring
bulking up to F₂₀ or beyond, making the method time-consuming.

2. Short-term bulks are useful only for isolating homozygous lines; natural selection has
minimal impact.

3. Provides limited scope for breeder’s skill in early generations.

4. Inheritance patterns cannot be studied due to lack of pedigree records.

5. Natural selection may favour competitive but agronomically inferior types (e.g., tall
plants over high-yielding dwarfs).

6. Off-season nurseries and greenhouses are unsuitable, as natural selection under

those conditions may not reflect the target environment.

Modern Perspective & Scientific Relevance

• Bulk method is increasingly relevant in:

o Evolutionary breeding

o Climate-resilient crop development

o Low-input and organic agriculture

• Modern adaptations integrate:

 o Periodic phenotypic screening

o Molecular tools at later stages (bulk → selection)

Modified Bulk Method

In the modified bulk method, selection is practiced in F₂, F₃, and subsequent generations.
Pedigree records are not maintained; instead, superior plants are selected, bulked, and
advanced generation after generation.

In F₄, superior plants are selected and harvested on a single-plant basis. In F₅, these selected
plants are evaluated in progeny rows, and the best progenies are selected and harvested. In
F₆, preliminary yield trials (PYT) are conducted to identify the most promising families. In the
subsequent generations, regular yield trials are undertaken for further evaluation and
selection.

This modification of the bulk method provides the breeder with ample opportunity to apply
skill and judgment in selection, while the absence of pedigree record maintenance remains a
major advantage of the method.

Merits

• Allows selection in early generations (F₂–F₃), improving efficiency for highly heritable
traits.

• Combines natural selection with breeder’s conscious selection, enhancing

adaptability.

• No maintenance of pedigree records, reducing labour, time, and cost.

• Superior plants contribute more seed, increasing the frequency of desirable

genotypes.

• Suitable for handling large segregating populations.

Demerits

• Early selection may lead to loss of superior genotypes due to environmental effects.

• Not suitable for traits with low heritability (e.g. yield).

• Genetic variability gradually reduces because of repeated selection and bulking.

• Performance of selected plants in early generations may not reflect true genetic
potential.

• Less precise than pedigree method for detailed genetic studies.

Single Seed Descent (SSD) Method

(Modification of Bulk Method)

Single Seed Descent (SSD) is a modification of the bulk method in which one seed from each
plant in the segregating generations (F₂ onward) is advanced to the next generation
without selection, until near-homozygosity is attained. The method was formalized by C.A.
Brim (1966) during soybean breeding. SSD is mainly used in self-pollinated crops to rapidly
obtain homozygous and homogeneous lines.

Procedure

• A large F₂ population is raised from the selected cross.

• One random seed from each F₂ plant is bulked to raise the F₃ generation.

• The same procedure is followed in F₃, F₄, and subsequent generations.

• Generations are advanced rapidly at high plant density, often using off-season
nurseries or greenhouses.

• By F₅ or F₆, plants become nearly homozygous.

• In F₅/F₆, individual plants are selected and grown as plant progeny rows.

• Selection is practiced among progenies, and superior rows are bulked for PYT in
F₇/F₈.

• Coordinated yield trials are conducted in F₈/F₉.

Merits

• Enables maximum speed of generation advancement in conventional breeding.

• Requires minimum space, labour, and record-keeping.

• Makes efficient use of greenhouse and off-season facilities.

• Maintains a random sample of genotypes from the original F₂ population.

• Ideal for achieving rapid homozygosity.

Demerits

• No selection is possible during segregating generations.

• Progressive reduction in population size due to germination failure and plant loss.

• Not suitable where early phenotypic selection is essential.

• Loss of weak but genetically superior plants may occur, especially in pulses.

Role in Crop Improvement

• Rapid development of homozygous lines

Example: Production of recombinant inbred lines (RILs) in rice and wheat.
 • Time-efficient cultivar development
Example: Accelerated breeding of soybean varieties using off-season nurseries.

• Preservation of genetic variability until fixation

Example: Maintenance of wide variability in wheat segregating populations.

• Pre-breeding and genetic studies

Example: Use of SSD-derived RILs for QTL mapping and trait dissection.

Pedigree Method of Breeding

The pedigree method is a selection procedure used in self-pollinated crops, first described
by H.H. Lowe (1927). In this method, selection starts from the F₂ generation and continues
through successive segregating generations, with individual plant selection and progeny
testing in each generation.

Pedigree refers to the record of ancestry of a selected plant, tracing its descent generation
by generation. Throughout the breeding programme, a complete parent–offspring record is
maintained, known as the pedigree record.

Pedigree Record

A pedigree record documents the ancestry and selection history of each progeny line.

Example: Pedigree number 7911

• 79 → year of cross (1979)

• 11 → cross number made in that year

Purpose of Pedigree Records

• To trace each progeny back to the original F₂ plant

• To determine genetic relatedness among selections

• To infer possible shared genes and inheritance patterns

• To assist in accurate selection and comparison of lines

Applications of the Pedigree Method

The pedigree method is the most widely used approach for handling segregating
generations in self-pollinated crops.

Major Applications
 1. Improvement of established varieties (combination breeding)
– Correction of specific defects such as disease susceptibility, tall plant height, or late
maturity

2. Recovery of superior recombinants

– Identification of desirable gene combinations following hybridization

3. Transgressive breeding
– Selection of segregants exceeding both parents for yield or other traits

4. Improvement of specific traits

– Disease resistance, maturity duration, plant height, lodging resistance, quality traits

5. Simultaneous improvement of yield and agronomic traits

6. Applicable to simple and complex crosses

Procedure of Pedigree Method

1. Hybridization
Selected parents are crossed to produce F₁ seed (simple or complex cross).

2. F₁ Generation
F₁ plants are space-planted to produce sufficient F₂ seed.
Usually 15–30 F₁ plants are adequate.

3. F₂ Generation
200–10,000 plants are space-planted.
100–500 superior individual plants are selected based on phenotype, and seeds are
harvested separately.

4. F₃ Generation
Progeny rows of selected F₂ plants are grown.
Superior plants are selected from high-performing families.

5. F₄ Generation
Selection is practiced within and between progenies.
Inferior families are discarded; superior progenies are advanced.

6. F₅ Generation
Many lines approach homozygosity.
25–100 progenies are bulk-harvested and evaluated in preliminary yield trials.

7. F₆ Generation
Lines are grown in multi-row plots, evaluated visually, and harvested in bulk.
Progenies are largely homozygous.
 8. F₇ Generation
Replicated preliminary yield trials are conducted.
2–5 superior lines outperforming checks are selected for multilocation testing.

9. F₈–F₁₀ Generations
Replicated multilocation trials for yield, adaptability, and resistance.

10. F₁₁ Generation

Seed multiplication and variety release.

Merits of Pedigree Method:

1. Breeder’s maximum control: Allows careful visual selection in every generation.

e.g. Selecting ideal plant type in wheat F₂.
 2. Helps understand inheritance of qualitative traits: Pedigree records show how traits
pass from parent to progeny.
e.g. Tracing awned/awnless or grain-colour inheritance.

3. High chance of obtaining transgressive segregants: Superior recombinants better

than both parents often appear.
e.g. Rice lines with more tillers than either parent.

4. Early removal of defective progenies: Weak, diseased, or lodging-prone lines are

discarded early.
e.g. Rejecting rust-susceptible wheat lines in F₂–F₃.

5. Best for simply inherited and easily visible traits: Traits like height, color, or major-
gene resistances.
e.g. Selecting dwarf vs tall plants in barley.

Demerits of Pedigree Method:

1. Pedigree record maintenance is tedious: Requires detailed tracking of each progeny.

e.g. Long numbers like 7911-7-4-14-3 (A plant selected in F₆, from row 3, whose
parents trace back to→ F₅ row 14 → F₄ row 4 → F₃ row 7 → cross no. 7911).

2. Labour-intensive and time-consuming: Difficult to manage many crosses at once.

e.g. Handling large wheat populations.

3. No natural selection: Only breeder-driven selection; weaker genotypes may persist.

e.g. Retaining visually good but stress-susceptible lines.

4. Useful genotypes may be lost early: Early selection when heterozygosity is high may
eliminate good types.
e.g. Discarding an F₂ rice plant that looks poor due to environment.

5. Success depends heavily on breeder’s skill: Requires experience to judge early-

generation plants.
e.g. F₂ yield cannot be judged directly—needs expert intuition.
Backcross method

The backcross method is a breeding technique in which an F₁ hybrid is repeatedly

crossed with one of its parents (recurrent parent) to transfer a specific desirable gene from
a donor parent into an otherwise well-adapted variety, without altering its genetic
background.

Procedure

• A recurrent parent (elite but deficient in one trait) is crossed with a donor parent
carrying the desired gene.

• The F₁ hybrid is repeatedly backcrossed to the recurrent parent.

• Selection is practiced in each generation for the target trait and resemblance to the
recurrent parent.

• After 5–6 backcrosses, plants contain ≈98% recurrent parent genome.

• Selected plants are selfed, progenies are evaluated, and the best line is tested and
released.

(Procedure differs for dominant and recessive genes; recessive traits require selfing and
testing in alternate generations.)

Merits

• New variety remains nearly identical to the recurrent parent, except for the added
gene.

• Predictable and reproducible results.

• Requires less yield testing, saving time.

• Smaller populations are sufficient due to single-trait selection.

• Efficient use of off-season nurseries and greenhouses.

• Only practical method for interspecific gene transfer.

Demerits

• Yield and overall performance cannot exceed the recurrent parent.

• Linkage drag may introduce undesirable traits.

• Repeated hybridization is labour-intensive and costly.

• Recurrent parent may become obsolete during programme duration.

Role in Crop Improvement

 • Transfer of disease and insect resistance
Example: Rust resistance in wheat; downy mildew resistance in pearl millet (Tift 23A).

• Correction of specific defects in elite varieties

Example: Adding blast resistance to popular rice varieties without changing grain
quality.

• Interspecific gene transfer

Example: Disease resistance introgressed from Saccharum spontaneum into
sugarcane.

• Development of cytoplasmic male sterile (CMS) lines

Example: CMS transfer in hybrid rice and pearl millet.

• Production of isogenic lines

Example: Near-isogenic wheat lines for disease resistance studies.

A. Transfer of a Dominant Gene

Example: Rust resistance in wheat (R)

1. Selection of parents

o Recurrent parent (A): high-yielding but susceptible

o Donor parent (B): resistant (dominant gene)

2. Hybridization (Year 1)

o A (♀) × B (♂)

3. F₁ generation (Year 2)

o All plants express resistance (Rr)

o F₁ plants are backcrossed to recurrent parent A

4. BC₁ generation (Year 3)

o Segregation occurs (Rr : rr = 1:1)

o Resistant plants selected and backcrossed to A

5. BC₂–BC₅ generations (Years 4–7)

o In each generation:

▪ Select resistant plants

▪ Backcross to recurrent parent A

6. BC₆ generation (Year 8)

 o Plants possess ~98% recurrent parent genome

o Resistant plants are selfed

7. BC₆F₂–BC₆F₃ (Years 9–10)

o Homozygous resistant plants selected

o Progenies similar to A are bulked

8. Yield testing & release (Years 11–12)

o Compared with recurrent parent A

o Seed multiplication and distribution

B. Transfer of a Recessive Gene

Example: Recessive disease resistance (r)

1. Selection of parents

o Recurrent parent (A): susceptible

o Donor parent (B): resistant (rr)

2. Hybridization (Year 1)

o A (♀) × B (♂)

3. F₁ generation (Year 2)

o All plants susceptible (Rr)

o Backcrossed to recurrent parent A

4. BC₁ generation (Year 3)

o No resistant plants expressed

o All plants are selfed

5. BC₁F₂ generation (Year 4)

o Segregation occurs (rr appears)

o Resistant plants selected and backcrossed to A

6. BC₂ generation (Year 5)

o No resistance expression

o Plants selected for resemblance to A and backcrossed

7. BC₂F₂ / BC₃F₂ generations (Years 6–8)

o Alternate selfing and testing for resistance

o Resistant plants backcrossed to A

8. Final backcross (Year 9–10)

o Resistant plants selfed

o Homozygous resistant progenies selected

9. Yield testing & release (Years 11–13)

o Tested against recurrent parent

o Seed multiplication and release

Merits of Backcross Method

1. New variety remains almost identical to the recurrent parent, except for the
transferred gene; results are predictable and reproducible.
e.g. Adding blast resistance to a popular rice variety while keeping yield and grain
quality unchanged.

2. Extensive yield testing is not required, as the recurrent parent’s performance is

already known, saving 4–5 years.
e.g. Introducing rust resistance into an established wheat variety without repeating
multi-location trials.
 3. Method is not environment-dependent, except for trait expression; off-season
nurseries and greenhouses can produce 2–3 generations per year.
e.g. Conducting rapid backcrossing for bacterial blight resistance in rice using off-
season facilities.

4. Smaller populations are sufficient because selection targets a single character.

e.g. Screening only a few plants for stem-borer resistance in maize backcross lines.

5. Specific defects can be corrected without altering desirable traits of well-adapted

varieties. e.g. Improving a farmer-preferred cotton variety for jassid resistance
without changing its fibre quality.

6. Only suitable method for interspecific gene transfer from related or wild species.
e.g. Transferring late blight resistance from wild potato (Solanum demissum) to
cultivated potato.

Demerits of Backcross Method

1. New variety cannot exceed the recurrent parent except for the added trait.
Example: Yield remains the same even after adding disease resistance to a rice
variety.

2. Linkage drag may occur, transferring undesirable linked genes along with the desired
gene.
e.g. Introducing insect resistance in tomato may also bring small fruit size due to
linked genes.

3. Hybridization must be done in every backcross, which is laborious, expensive, and

time-consuming. e.g. Repeated hand-emasculation and pollination required in cotton
backcrossing

4. Recurrent parent may become outdated by the time the programme finishes, as
superior varieties may emerge. e.g. a new higher-yielding wheat variety may already
be released.

RECURRENT SELECTION AND COMBINING ABILITY:

Combining Ability (CA): refers to the capacity or ability of a genotype to transmit superior
performance to its crosses.

The concept of CA as a measure of gene action in maize was proposed by Sprague & Tatum
(1942).

General Combining Ability (GCA): Average performance of a strain/genotype/inbred line in

a series of hybrids/cross combinations.

Estimated from the performance of F1’s from the crosses.

GCA estimated from half-sib families.
Specific Combining Ability (SCA): the performance of a parent in a specific cross.

Refers to the deviation of a particular cross from general combining ability.

Full sib: Progeny of the mating of two individuals/parents.

Half sib: Progeny having one parent in common.

Tester: A common parent mated to a number lines/strain/plants.

Open pollinated variety: variety developed in cross pollinated species, pollination occurring
naturally without restriction.

GENETIC BASE:

Genetic Base: The amount of genetic diversity available in a breeding population is referred
to as genetic base of that population.

1. Broad genetic base:

Plant populations that are composed of several pure lines, inbred lines or heterozygotes and
homozygotes are said to have broad genetic base.

Examples: Mass selected varieties, multiline varieties, synthetics, composites and land races.

2. Narrow genetic base:

Those plant populations that have been developed from single homozygote or heterozygote
are said to have narrow genetic base.

Examples: pure line variety, varieties developed by backcross, pedigree method bulk
method, and single seed decent method, clonal variety.

RECURRENT SELECTION:

• Idea of recurrent selection was first independently suggested by Hayes and Garber
(1919) and East and Jones (1920).

• The procedure of recurrent selection described by Jenkins (1940).

• The term recurrent selection given by Hull (1945).

• It is a cyclic selection that used to improve the frequency of desirable alleles for a
character in breeding population.

Hull defined recurrent selection as reselection generation after generation with inbreeding
of selects to provide for genetic recombination.

• Recurrent selection is defined as reselection generation after generation, with inter-

mating of selected plant to produce the population for the next cycle of selection.
 • The idea of this method was to ensure the isolation of superior inbreds from the
population subjected to recurrent selection.

The isolation of an outstanding inbred line depends on two factors:

1. The proportion of superior genotypes present in the base population from which
lines are isolated.

2. The effectiveness of selection during the inbreeding of desirable genes.

TYPES RECURRENT SELECTION:

1. Simple Recurrent selection (SRS)

2. Recurrent selection General Combining Ability (RSGCA)

3. Recurrent selection Specific Combining Ability (RSSCA)

4. Reciprocal Recurrent selection (RRS)

1. Simple recurrent selection:

• It was given by Sprague and Brown in 1930.

• A type of recurrent selection that does not include tester is referred as simple
recurrent selection.

• It is also known as phenotypic recurrent selection.

• This method is an extension of mass selection.

• The selection based on phenotype.

• This method is useful only for those characters that have high heritability.

• A number of plants with desirable phenotypes are selected and self-pollinated.

• In the second year, separate progeny rows are grown for selfed seeds.

• Progenies are intercrossed by hand in all possible combinations, and an equal

amount of seed from each is composited to raise the next generation.

• For the first recurrent cycle, the same process is repeated, but this time with a
composited seed.

• Recurrent selection is effective in increasing the frequency of desirable genes in the

selected population. Suited for highly heritable characters.

• In some cases, we can see large variability than the population.

• Recurrent selection is considered to be more effective than selection with self-

pollination.
Inbreeding can be kept low by following ways:

1. Population from inter crosses is may be allowed to mate randomly for one
generation, and the seed is subjected to reselection

2. Each intercrops maybe grown separately, and it should be ensured that selected
plants are not related by any parentage

1. Simple Recurrent Selection (SRS)

Concept:

Simple Recurrent Selection (SRS) is the most basic form of recurrent selection. It focuses on
selecting the best individuals based on their phenotype (observable traits) and using those
to form the next generation. The process is repeated over multiple generations to gradually
improve the population by increasing the frequency of desirable traits.

Procedure:

1. Year 1:

o Select phenotypically superior plants from the population.

 o Self-pollinate each of these plants.

o Harvest the seeds separately for evaluation in the next year.

2. Year 2:

o Grow progeny rows from the selfed seeds of the selected plants.

o Cross all the plants within each progeny row in all possible combinations
(intercrossing).

o Collect equal amounts of seed from each intercross and mix them together to
form the next generation’s seed population.

3. Year 3:

o Grow the new generation, select superior plants, and self-pollinate them.

o Discard inferior plants.

o Harvest and store seeds for evaluation.

4. Subsequent Years:

o Repeat the selection and crossing process for several cycles, depending on
the desired improvement.

Advantages:

• Straightforward and easy to implement.

• Suitable for traits with high heritability, where the phenotype is a good indicator of
genetic potential.

Limitations:

• Limited to traits that are easy to measure on individual plants.

• Ineffective for traits with low heritability or those that are influenced by
environmental factors.

2. Recurrent Selection for General Combining Ability (RSGCA)

Concept:

Recurrent Selection for General Combining Ability (RSGCA) aims to improve the general
combining ability (GCA) of a population. GCA is a measure of how well an individual’s genes
combine with the genes of other individuals in the population. RSGCA focuses on improving
traits that are influenced by additive genetic effects (where the combined effect of genes is
simply the sum of their individual effects).

Procedure:
 1. Year 1:

o Select several phenotypically superior plants from the source population.

o Self-pollinate the selected plants and also cross each of them with a broad-
based tester (e.g., open-pollinated variety or synthetic variety).

o Harvest seeds separately from both the self-pollinations and the crosses.

2. Year 2:

o Conduct a replicated yield trial using the seeds from the test crosses.

o Identify superior progenies based on performance.

3. Year 3:

o Plant the selfed seeds from the superior progenies in separate rows.

o Intercross the selected progeny in all possible combinations.

o Mix seeds from all intercrosses to create the source population for the next
selection cycle.

4. Year 4 and Beyond:

o Repeat the process, selecting plants that show superior combining ability,
both in terms of selfed progeny and test cross performance.

Advantages:

• Useful for improving population yield or other important traits.

• Can be used to increase the frequency of desirable genes in the population.

Limitations:

• Requires the use of a broad genetic base tester, which may be difficult to find or
maintain.

• Limited to traits with additive genetic effects, not effective for traits influenced by
non-additive genetic factors.

3. Recurrent Selection for Specific Combining Ability (RSSCA)

Concept:

Recurrent Selection for Specific Combining Ability (RSSCA) is used to improve the ability of a
population to combine well with a specific inbred line. Unlike GCA, which considers the
general combining ability with many lines, SCA focuses on how well an individual combines
with a particular inbred tester. This method is especially useful when dominance and
epistasis (gene interactions) play a significant role in heterosis (hybrid vigor).
Procedure:

1. Year 1:

o Select phenotypically superior plants from the source population.

o Self-pollinate them and cross each with a specific inbred tester (narrow
genetic base).

o Harvest seeds separately from both self-pollinations and test crosses.

2. Year 2:

o Conduct yield trials on progenies from the test crosses.

o Identify superior progenies based on performance when crossed with the

tester inbred.

3. Year 3:

o Plant the selfed seeds from superior progenies and intercross the selected
progeny in all possible combinations.

o Mix the seeds from all intercrosses to form the source population for the next
cycle.

4. Year 4 and Beyond:

o Continue the cycle, selecting for specific combining ability based on the
performance of progeny derived from test crosses with the specific inbred.

Advantages:

• Effective for improving traits controlled by non-additive gene action, such as

dominance and epistasis.

• Useful for breeding lines that will combine well with a specific inbred parent.

Limitations:

• Requires an inbred tester, which may be more difficult to maintain than a broad-
based tester.

• Less effective for traits controlled by additive gene effects.

4. Reciprocal Recurrent Selection (RRS)

Concept:

Reciprocal Recurrent Selection (RRS) is a breeding method designed to improve two

populations simultaneously. It is a more complex form of recurrent selection that combines
both General Combining Ability (GCA) and Specific Combining Ability (SCA). RRS is
particularly useful when two populations are being improved and both need to combine well
with each other.

Procedure:

1. Year 1:

o Two source populations (A and B) are selected. Phenotypically superior plants

from both populations are selected.

o Each selected plant is self-pollinated.

o Plants from population A are crossed with random plants from population B,
and vice versa.

o Harvest seeds separately from self-pollinations and test crosses.

2. Year 2:

o Conduct replicated yield trials for the test crosses from both populations.

o Identify superior progenies based on performance in the yield trials.

3. Year 3:

o Plant selfed seeds from superior progenies and intercross plants within each
population.

o Mix equal amounts of seeds from intercrosses within each population (A and
B) to create new source populations for the next cycle.

4. Year 4 and Beyond:

o Continue to select superior plants from both populations.

o Repeat the process of self-pollination, test crossing, progeny evaluation, and

intercrossing in subsequent years.

Advantages:

• Simultaneously improves two populations.

• Allows for the selection of both GCA and SCA, making it a more powerful method for
genetic improvement.

• Especially effective in breeding hybrid varieties by improving both parental

populations.

Limitations:

• Requires more resources and effort than other recurrent selection methods due to
the complexity of working with two populations simultaneously.
 • Can be difficult to implement, especially when both populations require significant
improvement.

Composite Varieties

A composite variety is a commercial variety produced by mixing seeds of several

phenotypically outstanding lines and allowing open-pollination to produce crosses in all
possible combinations. Unlike synthetic varieties, the parental lines in a composite are not
tested for general combining ability (GCA), and the yield cannot be predicted with certainty.

Production:

• Mixing Lines: Seeds from multiple superior lines (typically up to 20) are mixed and
planted in isolation.

• Open Pollination: Crosses occur among all the mixed lines through open pollination,
creating a genetically diverse population.

• No Testing for GCA: The parental lines are not tested for GCA, meaning the
performance of the composite cannot be precisely predicted.

• Maintenance: Composites are maintained through open-pollination and may be

improved through further population improvement.

Merits:

• Broad Genetic Base: Composites are more genetically diverse, which can provide
stability across varied environmental conditions.

• Simple Seed Production: Less expertise is needed in seed production compared to

hybrid varieties.

• Lower Seed Cost: Seed production costs are lower compared to hybrids.

• No Need for Reconstitution: Unlike hybrids, composites do not require

reconstitution.

Demerits:

• Unpredictable Performance: Since the parental lines are not tested for GCA, the
yield and other traits cannot be predicted.

• Lower Uniformity: The genetic diversity might lead to inconsistent performance

across generations.

Example: Maize composites such as Amber, Jawahar, and Kisan are widely used in India.

Synthetic Varieties
A synthetic variety is created by crossing several inbred lines (usually 4-6) that combine well
with each other. The selected inbreds are tested for general combining ability (GCA), and
once the variety is synthesized, it is maintained by open-pollination.

Production:

• Inbred Line Selection: Inbred lines are tested for GCA, and those with good
combining ability are chosen as parents.

• Crossing: The selected inbreds are crossed in all possible combinations to create the
synthetic variety.

• Open Pollination: Once synthesized, the population is maintained through open-

pollination, with progeny such as Syn1, Syn2, etc.

• Maintenance: Synthetics can be multiplied for one or more generations to produce

commercial seed.

Merits:

• Feasible for Cross-Pollinated Crops: Especially useful in species where pollination

control is difficult, such as maize, forage crops, and sugar beets.

• Seed Reuse: Farmers can use harvested grain as seed for subsequent crops.

• Wider Genetic Base: They are more stable across variable environments due to
broader genetic diversity.

• Lower Seed Cost: Cheaper than hybrids, and seed production requires less
specialized expertise.

Demerits:

• Lower Performance: Generally, synthetics perform lower than hybrids because they
only exploit GCA, while hybrids exploit both GCA and SCA.

• Less Consistent Performance: Performance can drop if lines with poor GCA are
included.

Example: Maize synthetic varieties developed by CIMMYT, such as Syn1 and Syn2, are
widely used for population improvement.

Hybrid Varieties

A hybrid variety is the first-generation cross (F1) between two genetically distinct parent
plants (often inbred lines). Hybrids exploit both general combining ability (GCA) and specific
combining ability (SCA), leading to enhanced traits like yield, disease resistance, and vigor
(known as heterosis or hybrid vigor).

Production:
 • Parent Selection: Two genetically distinct inbred lines or populations with
complementary traits are selected as parents.

• Controlled Cross-pollination: The selected parents are crossed in controlled

conditions (often using male-sterility techniques to ensure cross-pollination).

• Hybrid Seed Production: The resulting hybrid seeds are harvested and distributed for
commercial use.

Merits:

• Superior Performance: Hybrids exhibit superior growth, yield, and disease resistance
due to heterosis.

• High Predictability: The performance of hybrids can be predicted because they are
carefully selected for desirable traits.

• Uniformity: Hybrids are generally more uniform, with consistent performance across
fields.

Demerits:

• High Seed Cost: Hybrid seed production is costly due to the controlled breeding
process.

• Seed Cannot Be Reused: Farmers cannot save and replant hybrid seeds, as the
offspring typically do not perform as well as the parent plants.

Example: Maize hybrids such as Pioneer, Dekalb, and Syngenta hybrids are commonly used
in commercial farming for high yield and disease resistance.

Clonal Selection
Clonal selection is the process of selecting superior plants from a population of asexually
propagated crops (e.g., sugarcane, potato, banana) and using them to produce new clones
with desirable traits.

Production:

• Selection: Superior plants are selected from a population.

• Cloning: These plants are propagated asexually (e.g., through cuttings or grafting) to
maintain genetic uniformity.

• Testing: Selected clones are tested for traits like yield, disease resistance, and quality.
Merits:

• Preserves Genotype: Clonal selection avoids inbreeding depression and preserves

desirable gene combinations.

• Hybridization: Can be combined with hybridization for variability.

• Purity: Maintains the purity of clones over generations.

Demerits:

• Limited Variability: Uses only the existing genetic variability in the population.

• Clonal Degeneration: Clones can degenerate over time due to viral or bacterial
diseases, and mutations can accumulate.

Example: Sugarcane, Banana, Potato, and other clonally propagated crops.

Different approaches for improving the characteristics of inbred lines

Improving the characteristics of inbred lines is a central focus in plant breeding, especially
when developing new varieties or hybrids with superior agronomic traits such as higher
yield, disease resistance, and adaptability. Inbred lines, which are genetically uniform due to
repeated self-pollination, serve as the foundation for both hybrid and open-pollinated
variety development.

However, inbreeding can lead to inbreeding depression, reducing the vigor and overall
performance of these lines. Therefore, various breeding strategies are employed to enhance
the genetic quality of inbred lines, minimize inbreeding depression, and improve desirable
traits.

These strategies include:

1. Pedigree Selection Method

2. Bulk Method
3. Recurrent Selection
4. Heterosis (Hybrid Vigor)
5. Mutation Breeding

Heterosis and its exploitation.

1. "Heterosis is observed in both self and cross-pollinated crops, but hybrid varieties are
more common in cross-pollinated crops". Comment on this statement on the basis of
relevant information. (10) 2014

2. Define heterosis. How is heterosis being exploited in the development of hybrids? (20)
2017

3. What do you mean by heterosis? Describe the physiological basis of heterosis. (10) 2018

4. Describe the physiological and molecular basis of heterosis. (15) 2021

5. Explain inbreeding depression and its effect in crops. Also, explain the degrees of
inbreeding depression. (10M CS-2023)

6. Heterosis breeding and Mutation breeding. (10) 2018

7. Discuss the role of heterosis in crop improvement with suitable examples. (8) 2019

8. What do you mean by heterosis? Describe the genetic basis of heterosis. (8) 2020

9. Discuss the genetic basis of heterosis and inbreeding depression. (8) 2021

Cross-pollinated and many asexually reproducing species maintain high heterozygosity.

Inbreeding/selfing increases homozygosity and often causes loss of vigour and fertility
(inbreeding depression). Hybridization between unrelated lines restores heterozygosity and
may produce heterosis (hybrid superiority). These are often treated as opposite outcomes of
changes in heterozygosity.

Heterosis: Superiority of an F₁ hybrid over its parents for yield or other traits (term first used
by Shull, 1914).

Types of heterosis (common estimates)

• Mid-parent / Average (Relative) heterosis: F₁ > mean of parents

𝐹1 − 𝑀𝑃 𝑃1 + 𝑃2
%𝐻 = × 100, 𝑀𝑃 =
𝑀𝑃 2

• Heterobeltiosis (Better-parent heterosis): F₁ > better parent

𝐹1 − 𝐵𝑃
%𝐻 = × 100
𝐵𝑃

• Economic / Standard heterosis: F₁ > best commercial variety (most useful)

𝐹1 − 𝐶𝑉
%𝐻 = × 100
𝐶𝑉

• Negative heterosis: F₁ inferior to mid-parent/better parent (sometimes desirable for

traits like short duration/earliness).

Hybrid vigour is often used synonymously; practically it refers to superiority over the better
parent.

Manifestations of heterosis

• Higher yield

• Higher reproductive ability (seed set, fertility, propagule yield)

• Greater size and vigour (biomass, leaf area, faster early growth)

• Better quality (keeping quality, processing traits)

• Earlier flowering/maturity in some crops

• Greater resistance to diseases/pests

• Greater adaptability/stability across environments

Exploitation of heterosis in hybrid development

Hybrid varieties
A hybrid variety is the F₁ produced from genetically dissimilar parents (inbreds/pure
lines/clones/populations) and grown commercially.

Steps in heterosis breeding (core pipeline)

1. Germplasm collection & divergence assessment (choose diverse parents; e.g., D²

grouping conceptually).

2. Development of inbred lines/pure lines

o Cross-pollinated crops: repeated selfing + selection → inbreds after ~5–7

generations (vigour reduces initially then stabilizes).

3. Combining ability testing

o GCA (general combining ability): average performance across crosses (often

via topcross/tester).

o SCA (specific combining ability): performance of specific pairwise crosses

(diallel/line×tester).

4. Hybrid synthesis and evaluation

o Select best combinations for standard/economic heterosis, stability,

adaptation.

5. Commercial seed production system

o Ensure controlled pollination (detasseling/CMS/hand emasculation, isolation,

rouging) depending on crop.

Common hybrid types (especially in maize and other cross-pollinated crops)

• Single cross: A × B (two inbreds) — high performance, but seed production cost may
be higher.

• Double cross: (A × B) × (C × D) — easier/cheaper seed production historically due to

vigorous single-cross parents.

• Three-way cross: (A × B) × C — intermediate.

• Topcross: inbred × tester/variety — mainly for testing GCA.

Economic heterosis is the key criterion for release and adoption.

Heterosis in self-pollinated vs cross-pollinated crops

Improving Heterosis in Cross-Pollinated Crops

Cross-pollinated crops already maintain heterozygosity and show high inbreeding
depression. Therefore, heterosis improvement focuses on population improvement and
heterotic pattern development.

A) Development of Heterotic Groups

• Divide germplasm into distinct heterotic pools.

• Improve each group separately.

• Cross between groups → maximum heterosis.

• Widely practiced in maize breeding.

B) Reciprocal Recurrent Selection (RRS)

Most powerful method.

• Two populations are improved simultaneously.

• Selection is based on cross performance between populations.

• Enhances both GCA and SCA.

• Increases frequency of favorable dominant alleles in complementary populations.

C) Recurrent Selection

• Improves population mean.

• Accumulates favorable alleles.

• Reduces genetic load.

• Maintains variability for continued improvement.

D) Exploiting Genetic Divergence

• Use genetically diverse parents.

• Optimum divergence gives maximum heterosis.

• Too much divergence may reduce combining ability.

E) Purging Deleterious Alleles

• Limited selfing + selection reduces genetic load.

• Leads to stronger inbreds and better hybrids.

Improving Heterosis in Self-Pollinated Crops

Self-pollinated crops are naturally homozygous and show little inbreeding depression.
Heterosis is usually lower but can be enhanced by the following:
A) Creation of Artificial Divergence

• Select parents from genetically distant sources.

• Use exotic germplasm.

• Develop heterotic pools.

B) Population Improvement Before Inbreeding

• Intermate diverse lines.

• Improve population through recurrent selection.

• Then extract inbreds.

• Produces superior combining ability.

C) Combining Ability Based Selection

• Select parents based on SCA.

• Identify crosses showing high economic heterosis.

D) Cytoplasmic Male Sterility (CMS) System

• Enables large scale hybrid production.

• Makes heterosis exploitation practical (e.g., rice).

E) Mutation and Introgression

• Introduce new alleles from wild species.

• Increases allelic diversity → may enhance heterosis.

Physiological and molecular basis of heterosis

Physiological basis

• Often associated with:

o faster early growth rate

o higher leaf area index and biomass

o more efficient resource mobilization and metabolism

• A classic interpretation: hybrids may have more efficient enzyme systems and
altered growth regulators (earlier proposal: Whaley, 1952). Evidence is crop/trait
dependent; no single physiological mechanism explains all cases.

Molecular/genetic “working models”

• Improved metabolic balance in heterozygotes (intermediate/optimal product levels).

 • Complementation: both alleles contribute functionally distinct products → broader
functional capacity.

• Hybrid/novel enzyme forms (heteromultimers/“hybrid substance” concept in older

literature).

• Tissue-specific expression balance: intermediate expression across tissues can

improve whole-plant performance.

• Modern molecular interpretations often map onto:

o dominance/complementation (masking deleterious alleles),

o overdominance (true heterozygote advantage at some loci),

o epistasis (favourable interactions among loci).

Genetic basis of heterosis (classical hypotheses)

(A) Dominance hypothesis (Davenport, 1908; expanded later)

• Dominant alleles favourable; recessives deleterious.

• In F₁, harmful recessives are masked → heterosis.

• In inbreeding, recessives become homozygous → depression.

• Key issue historically: difficulty isolating inbreds “as vigorous as hybrids”; explained
partly by many genes + linkage (“dominance of linked genes” idea) and polygenic
nature/environment effects.

(B) Overdominance hypothesis (East & Shull, 1908)

• At some loci, heterozygote > both homozygotes (Aa > AA and aa).

• Predicts it is impossible to fix hybrid performance fully in homozygous lines if

heterosis is mainly overdominant.

• Clear examples exist for some major genes/oligogenes; for complex traits,
distinguishing overdominance from linkage/epistasis is difficult.

(C) Epistasis (gene interaction)

• Heterosis may result from favourable interactions among non-allelic genes

assembled in the hybrid.

Role of heterosis in crop improvement

• Yield enhancement and productivity jumps (hybrid technology).

• Stability and adaptability across environments (often lower variance than inbreds).
 • Improved quality, earliness, and biotic stress resistance in specific hybrids.

• Enables structured breeding systems: heterotic groups, combining ability

improvement, hybrid-based seed industry.

INBREEDING DEPRESSION

Mating between individuals related by descent/ancestry (sib mating). Selfing produces the
highest inbreeding.

Reduction/loss of vigour and fertility due to inbreeding/selfing; especially pronounced in

cross-pollinated species.

Genetic change with selfing: heterozygosity is reduced by ½ each generation.

Effects of inbreeding (on crops/populations)

• Expression of lethal/sub-lethal alleles (chlorophyll deficiency, rootless seedlings,

malformations).

• Reduced vigour (dwarfing, weaker plants, smaller organs).

• Reduced reproductive ability (low seed set, sterility).

• Population splits into distinct inbred lines (more uniform within lines).

• Increased homozygosity, reduced within-line variation.

• Yield reduction (in maize, inbreds may yield ~half of original open-pollinated source).

Degrees of inbreeding depression

1. High: severe depression/lethality; lines hard to maintain after few selfing generations
e.g., alfalfa (lucerne), carrot

2. Moderate: depression noticeable but inbreds can be developed and maintained

e.g., maize, sorghum (jowar), pearl millet (bajra)

3. Low: small depression

e.g., cucurbits, sunflower, onion/rye often cited as low in some notes

4. Nil/none: typical of self-pollinated crops (though heterosis may still occur)

Genetic basis of inbreeding depression

• Mainly due to increased homozygosity, leading to:

1. unmasking of deleterious recessive alleles (genetic load concept), and/or

2. loss of heterozygote advantage at loci where heterozygotes are superior

(overdominance), plus
 3. disruption of favourable epistatic interactions.

• Heterozygous vs homozygous balance (Mather concept):

o Cross-pollinated species tend to maintain heterozygous balance and carry

more genetic load, hence strong depression on inbreeding.

o Self-pollinated species evolve homozygous balance, with deleterious

recessives purged over time.

Exploitation of inbreeding in crop improvement

Even though inbreeding can reduce vigour, it is deliberately used to:

• Develop inbred lines/pure lines (uniform, stable, true-breeding).

• Expose deleterious alleles → purging through selection.

• Create parents for hybrid breeding, where:

o inbreds provide genetic uniformity and predictable combining ability,

o hybridization restores vigour → heterosis exploitation.

Practical strategy in cross-pollinated crops: controlled selfing + rigorous selection to obtain

stable inbreds → test GCA/SCA → produce commercial hybrids.

Somatic hybridization, Role of interspecific and intergeneric hybridization.

1. Explain the importance of somatic hybridization in agriculture. (10) CSE-2021

2. Explain the techniques involved in somatic hybridization in crop plants. (10) CSE-2022

3. Highlight the various contributions of interspecific hybridisation in crop improvement.

(10) CSE-2014

4. What is distant hybridisation? Discuss its applications, achievements and limitations in

plant breeding. (20) CSE-2024

5. Interspecific and intergeneric hybrids. (10) IFS-2016

6. What is distant hybridization? Explain how it is used for developing new varieties. (15)
IFS-2021

Hybridization

Hybridization is the mating or crossing of two plants or lines with dissimilar genotypes. It is
performed by transferring pollen from the male parent to the stigma of the female parent.
The resulting seeds and progeny are called hybrids or F₁ generation.

The progeny obtained by selfing or inter-mating F₁ plants form the segregating generations
(F₂, F₃, etc.).
The first natural hybridization was recorded by Cotton Mather (1716) in corn. The first
artificial interspecific plant hybrid was produced by Thomas Fairchild in 1717. It is commonly
known as ‘Fairchild Mule’.

Objectives of Hybridization

The main objective of hybridization is to create genetic variation.

When two genotypically different plants are crossed, their genes combine in F₁, and
segregation and recombination in later generations produce new gene combinations.

The extent of variation depends on the number of genes for which the parents differ —

• Closely related parents → few differing genes → less variation.

• Distantly related parents → many differing genes → high variation.

Major Aims of Hybridization

(a) Combination Breeding

1. The aim is to combine desirable characters from two or more varieties into a single
improved variety.

2. These traits may be controlled by oligogenes (few major genes) or polygenes (many
minor genes).

(b) Transgressive Breeding

1. The aim is to produce plants in F₂ and later generations that are superior to both
parents — a phenomenon known as transgressive segregation.

2. This occurs due to the accumulation of favourable (plus) genes from both parents
through recombination.

(c) Hybrid Varieties

1. In most self-pollinated crops, the F₁ generation often shows hybrid vigour (heterosis)
— higher yield, vigour, and adaptability than the parents.

2. Wherever it is commercially feasible, the F₁ hybrid may be released directly as a

variety.

Need for Hybridisation:

1. Natural variability in self-pollinated crops is quickly lost due to repeated selection.

2. As landraces are replaced by purelines, genetic variation becomes exhausted.

3. Therefore, plant breeders must create new variability artificially.

 4. This is most effectively achieved through crossing two genetically different
purelines.

Procedure of Hybridization:

It involves the following steps:

(i) Selection of parents.

(ii) Selfing of parents or artificial self-pollination.

(iii) Emasculation.

(iv) Bagging

(v) Tagging

(vi) Crossing

(vii) Harvesting and storing the F, seeds

(viii) Raising the F1 generation.

(i) Selection of parents:

The selection of parents depends upon the aims and objectives of breeding. Parental plants
must be selected from the local areas and are supposed to be the best suited to the existing
conditions.

(ii) Selfing of parents or artificial self-pollination:

It is essential for inducing homozygosity for eliminating the undesirable characters and
obtaining inbreeds.

(iii) Emasculation:

It is the third step in hybridization. Inbreeds are grown under normal conditions and are
emasculated. Emasculation is the removal of stamens from female parent before they burst
and shed their pollens.

It can be defined as the removal of stamens or anthers or the killing of the pollen grains of a
flower without affecting in any way the female reproductive organs. Emasculation is not
required in unisexual plants but it is essential in bisexual or self-pollinated plants.

Various methods used for emasculation are:

(a) Hand Emasculation or Forceps or Scissor Method:

This method is generally used in those plants which have large flowers. In this method the
corolla of the selected flowers is opened and the anthers carefully removed with the help of
fine-tip forceps.
(b) Hot Water Treatment:

Removal of stamens with the help of forceps is very difficult in minute flowers. In such small
hermaphrodite flowers (e.g., Bajra, Jowar) emasculation is done by dipping the flowers in
hot water for a certain duration (1-10 minutes) of time.

The time varies from species to species. This method Is based on the fact that gynoecia can
withstand the hot temperature at which the anthers are killed

(c) Cold Water Treatment:

Like hot water cold water also kills pollen grains without damaging the gynoecium. In rice 0-
6°C temperature is maintained to kill the pollen grains. This method is less effective than hot
water treatment.

(d) Alcohol Treatment Method:

This method is not commonly used for emasculation because duration of treatment is an
important factor since a very short duration is required failing which even the gynoecium
may be damaged. Flowers or inflorescences are immersed in alcohol of a suitable
concentration for a brief period. In alfa-alfa, a treatment of even 10 seconds with 57 %
alcohol is sufficient to kill the pollen grains.

(e) Suction Method:

It is a mechanical method and is suitable for the crops having minute flowers. In this method
the amount of pressure is applied in such a way that only anthers are sucked out and other
parts of the flower like gynoecium remain intact. However, in this method 10-15% self
pollination takes place. It is one of the major drawback of this method.

(f) Male Sterility or Self-incompatibility Method:

Emasculation option can be eliminated by the use of male-sterile plants, In some self-
pollinated plants for example, Sorghum, Onion, Barley etc. anthers are sterile and do not
produce any viable pollens! Similarly self-incompatibility may also be used to avoid
emasculation.

g) Chemical Gametocides:

Certain chemicals are capable of causing male sterility, when sprayed before flowering e.g.,
2, 4-D, naphthalene acetic acid (NAA), maleic-hydrazide (MA), tribenzoic acid etc. FW450 in
cotton may be used for bringing about emasculation.

(iv) Bagging:

It is the fourth step and completed with emasculation. The emasculated flower or
inflorescence is immediately bagged to avoid pollination by any foreign pollen. The bags may
be made of paper, butter paper, glassine or fine cloth. Butter paper or vegetable parchment
bags are most commonly used.

The bags are tied to the base of the inflorescence or to the stalk of the flower with the help
of thread, wire or pins. The bagging is done with the emasculation in bisexual plants and
before the stigma receptivity and dehiscence of the anthers in unisexual plants. Both male
and female flowers are bagged separately to prevent contamination in male flowers and
cross-pollination in female flowers

(v) Tagging:

The emasculated flowers are tagged just after bagging. Generally circular tags of about 3 cm
or rectangular tags of about 3 x 2 cm are used. The tags are attached to the base of flower or
inflorescence with the help of thread.

The information on tag must be as brief as possible but complete bearing the following
information:

(i) Number referring to the field record

(ii) Date of emasculation

(iii) Date of crossing

(iv) Name of the female parent is written first followed by a cross sign (x) and then the male
parent, e.g., C x D denotes that C is the female parent and D is the male parent.

(vi) Crossing:

It is the sixth step. It can be defined as the artificial cross-pollination between the genetically
unlike plants. In this method mature, fertile and viable pollens from the male parent are
placed on the receptive stigma of emasculated flowers to bring about fertilization.

Pollen grains are collected in petridishs (e.g., Wheat, cotton etc.) or in paper bags (e.g.,
maize) and applied to the receptive stigmas with the help of a camel hair brush, a piece of
paper, tooth pick or forceps. In some crops (e.g., Jowar, Bajra) the inflorescences of both
parents are enclosed in the same bag.

(vii) Harvesting and Storing the F1 Seeds:

Crossed heads or pods of desirable plants are harvested, and after complete drying, they are
threshed. Seeds are stored properly with original tags.

(viii) Raising the F1 generation:

In the coming season, the stored seeds are sown separately to raise the F1 generation. The
plants of F1 generation are progenies of cross seeds and therefore are hybrids.
Types of hybridisation:

Close / Narrow Hybridisation

Parents are closely related and belong to the same species.

• Intervarietal hybridisation: Cross between two varieties of the same species.

Example: Wheat (HD2967 × PBW343)

• Intravarietal/Intrapopulation hybridisation: Within a variety or within a line.

Outcome: Hybrids are generally fertile and used for commercial crop improvement.

Distant / Wide Hybridisation (Parents are genetically distant)

Distant hybridization is the crossing of genetically distant individuals belonging to:

• Different species (interspecific) or

• Different genera (intergeneric)

within the same family, to introduce desirable alien genes into cultivated crops.

Objectives of Distant Hybridization

• Transfer resistance genes (diseases, pests, abiotic stress)

• Broaden genetic base

• Improve yield, quality, adaptability

• Create novel crop types

Interspecific hybridisation: Interspecific hybridisation involves the crossing of two different
species within the same genus. This method facilitates the transfer of specific traits between
closely related plants, leading to the creation of hybrids with enhanced qualities.

o Apples (Malus spp.) - Many modern apple varieties are hybrids between different
Malus species, such as M. domestica (common apple), M. sieversii (Central Asian
apple), and M. fusca. Popular hybrids include 'Fuji', 'Gala', and 'Pink Lady'.

o Citrus fruits - Oranges, grapefruits, lemons, and other citrus crops are primarily
interspecific hybrids involving Citrus maxima, C. paradisi, C. reticulata, C. sinensis and
other species. Varieties like oranges originated from crosses between pummelos and
mandarins.

o Wheat (Triticum spp.) - Modern bread wheat is a hybrid between two grass species,
T. turgidum and Aegilops tauschii. This hybridisation event occurred only 10,000
years ago.

Intergeneric hybridisation: Different genera.

Intergeneric hybridization takes hybridization a step further by crossing plants from different
genera. This process broadens the genetic pool, creating hybrids with diverse characteristics.

Intergeneric Hybridization in Plants: Examples

o Wheat rye cross

Radish cabbage cross (Karpechenko, 1928): Radish (Raphanus sativus) x Cabbage (Brassica
oleracea) = F1 sterile→ Colchicine treatment = Raphanobrassica
Limitations of Distant Hybridization in Crop Improvement:

Distant hybridization, while a valuable tool in the field of crop breeding, is subject to several
significant limitations that can hinder its effectiveness and applicability. Understanding these
limitations is crucial for optimizing the benefits of distant hybridization in developing
improved crop varieties.

• Incompatible crosses: Many distant crosses, despite various techniques, remain

unachievable. This often leads to a trial-and-error approach, especially in intergeneric
and interfamily crosses, where the reasons for failure are not always clear.

• F1 hybrid sterility: F1 hybrids from distant crosses frequently exhibit sterility, ranging
from partial to complete. Overcoming this requires specialized techniques, such as
chromosome doubling, which may not always be feasible.

• Difficulties in new species creation: The development of new crop species through
allopolyploidy is often challenged by issues like reduced yield, poor agronomic traits,
and sterility, necessitating extensive breeding efforts.

• Reduced homoeology: The lack of chromosome pairing between parental species

limits the transfer of oligogenes, requiring complex chromosome manipulations,
mainly viable in polyploid species.

• Linkage drags: Often, desirable genes are linked with undesirable ones, complicating
their use in crop improvement. Although continued breeding can sometimes resolve
this, it demands significant time and resources.

• Transfer of recessive and quantitative traits: Transferring recessive traits and

quantitative characters through distant hybridization is often impractical, with
dominant monogenic traits being the most successfully transferred.

• Flowering issues in F1 hybrids: Some F1 hybrids, like those in the genus Glycine, may
not flower at all, posing a significant barrier to further breeding.

• Crossing challenges with improved varieties: Often, the best varieties of a crop
species are not easily crossable with wild species, necessitating the use of older, less
improved varieties for distant hybridization.

• Seed Dormancy: In some interspecific hybrids, such as in Arachis, F1 seeds exhibit

prolonged dormancy, complicating their use in crop improvement programs

Enhancing Crop Improvement through Distant Hybridization: Distant hybridization has been
instrumental in advancing crop improvement, leveraging the genetic diversity found across
different species. This approach not only enriches the genetic pool but also introduces
beneficial traits into cultivated crops.
 • Creation of new crop species: By crossing species and inducing chromosome doubling
in sterile hybrids, new allopolyploid crop species are developed. A prime example is
Triticale, a successful synthetic crop derived from wheat and rye.

• Development of alien addition and substitution lines: This involves incorporating

chromosomes or chromosome segments from one species into another. In wheat,
alien addition lines have introduced disease resistance traits from wild relatives.

• Transfer of small chromosome segments for specific traits: This method is crucial for
introducing specific desirable genes, like disease resistance, from one species to
another. An example is the transfer of black arm resistance from Gossypium
barbadense to Gossypium hirsutum.

• Utilization as new varieties: Some distant hybrids are directly used as new crop
varieties due to their advantageous traits. The hybrid cotton variety 'Varalakshmi' is
an example, offering high yield and quality.

• Transfer of cytoplasm: This technique involves transferring the cytoplasm of one

species to another to produce desirable traits, such as cytoplasmic male sterility.

• Development of disease resistance: Many crops have benefitted from genes for
disease resistance transferred from wild relatives, as seen in wheat with rust
resistance genes from related species.

• Improving adaptation and quality: Distant hybridization has contributed to broader

adaptability and improved quality in crops. For instance, cold tolerance in wheat and
increased protein content in rice are outcomes of this approach.

• Yield improvement: Wild relatives of crops like oat and chickpea have been used to
increase yield significantly through distant hybridization.

• Enhancing other characteristics: This method has also improved various other traits
in crops, such as plant stature in wheat and oil quality in oil palm.

Attributes Interspecific Hybridization Intergeneric Hybridization

Definition Crossing between two Crossing between two different

different species of the same genera of the same family.
genus

Desirable character Used when the desirable Used when the desirable genes
source character is not found within are not found in different
the crop's species. species of the same genus.
Gene transfer method Effective for transferring genes Utilised for the transfer of
from related cultivated or wild specific characters from allied
species into cultivated plants genera into cultivated species

Propagation success More successful in vegetatively Generally used in asexually

propagated species like propagated species.
sugarcane and potato.

Types of crosses Can produce fully sterile, F1 hybrids are typically sterile;
produced partially fertile, or fully fertile fertility may be restored
crosses through chromosome doubling.

introgression Leads to introgression, which is While challenging, introgression

the transfer of some genes is achievable with advanced
into the genome of another breeding techniques and is
species used to incorporate specific
genes from one genus to
another.

Use in crop Widely used and successful in Rarely used in crop

improvement crop improvement programs. improvement due to various
associated problems.

Seed setting More than one seed setting is Seed setting is not possible.
possible.

Fertility Hybrids can vary from Hybrids are always sterile;

completely sterile to fully sterility can be overcome by
fertile. chromosome doubling.

Parents involved Involves two species of the Involves two genera of the
same genus. same family

Evolution of few crops Possible but sometimes Not possible with interspecific
difficult. crosses; less possible than
intergeneric crosses.

Example G. hirsutum x G. barbadense, T. Triticale, a cross between wheat

aestivum x T. compactum, T. and rye, Raphanobrassic, a
aestivum x T. durum cross between radish and
cabbage
SOMATIC HYBRIDISATION:

Somatic hybridization is a biotechnological technique in which two genetically

distinct plant cells are fused to form a hybrid cell, which can subsequently regenerate into a
whole plant. The fusion is achieved at the protoplast level, bypassing normal sexual
reproduction.

This technique is particularly useful when:

• Sexual hybridization is impossible (interspecific/intergeneric barriers)

• Hybrids are sterile

• Cytoplasmic traits need to be transferred

• Vegetatively propagated crops require genetic improvement

The core process involved is protoplast fusion.

Protoplast Isolation

A protoplast is a plant cell from which the cell wall has been enzymatically removed, leaving
the plasma membrane intact.

Source Material

• Young, healthy mesophyll tissue (preferred)

• Cell suspension cultures

• Hypocotyls, cotyledons, callus tissue

Enzymatic Isolation

Cell walls are degraded using:

• Cellulase (degrades cellulose)

• Pectinase / Macerozyme (degrades pectin)

The process is carried out in hypertonic solution (0.5–0.8 M mannitol or sorbitol) to prevent
osmotic bursting.

After 16–18 hours incubation:

• Naked spherical protoplasts are released

• They are purified by filtration and centrifugation

Protoplast Fusion

Protoplast fusion refers to the physical and physiological union of two protoplasts resulting
in cytoplasmic continuity and nuclear coexistence within a single hybrid cell.
Fusion may occur:

• Naturally (spontaneous)

• Artificially (induced: chemical or electrical)

Spontaneous Fusion

Spontaneous fusion occurs unintentionally during enzymatic cell wall degradation.

During protoplast isolation:

• Adjacent cells are connected via plasmodesmata.

• Enzymatic digestion weakens cell walls.

• Closely associated protoplasts fuse due to membrane instability.

Mostly intraspecific

Produces homokaryons (multiple nuclei of same genotype)

May contain 2–40 nuclei

Low frequency but unavoidable

To minimize spontaneous fusion:

• Sequential isolation methods

• Strong plasmolyticum pre-treatment

• Gentle agitation during enzymatic digestion

Significance

Spontaneous fusion has little value in somatic hybridization because:

• It does not create genetic novelty

• It mainly produces same-species fusions

Induced Fusion

Induced fusion is deliberately carried out to fuse protoplasts of different genotypes.

It can be achieved through:

1. Chemical methods

2. Electrofusion

Chemical Fusion

Chemical fusogens alter membrane surface properties, promoting adhesion and fusion.
Polyethylene Glycol (PEG) Mediated Fusion

Most widely used method.

Procedure:

• Mix protoplasts of both parents

• Treat with 15–45% PEG solution (MW 1500–6000)

• Incubate for 15–30 minutes

• Gradual washing with culture medium

Mechanism:

Fusion occurs in three phases:

1. Agglutination – membranes brought into close proximity

2. Localized membrane fusion – cytoplasmic bridges form

3. Rounding off – formation of spherical heterokaryons

PEG acts by:

• Dehydrating membrane surfaces

• Redistributing surface charges

• Forming molecular bridges (often mediated by Ca²⁺ ions)

Limitations:

• Toxicity in some species

• Random multi-cell aggregates

• Less precise control

Other Chemical Methods

• NaNO₃ treatment

• High pH (10.5) + High Ca²⁺ treatment

These are less efficient compared to PEG.

Electrofusion

Electrofusion is a more controlled and efficient technique.

Principle:

1. Low-voltage alternating current aligns protoplasts (pearl-chain formation)

 2. A short high-voltage pulse creates transient membrane pores

3. Adjacent membranes fuse

Advantages:

• Rapid and synchronous

• Reduced chemical toxicity

• Higher fusion efficiency

• Controlled fusion events

Selection of Hybrid Cells

Because heterokaryons occur at low frequency, selection is critical.

Visual Markers

• Chlorophyll pigmentation

• Anthocyanin pigmentation

• Fluorescent dyes

Complementation

Hybrid restores a trait missing in one parent.

Example:

• One parent sensitive to antibiotic

• Other resistant

• Hybrid survives selective medium

Selectable Genetic Markers

• Antibiotic resistance genes

• Metabolic resistance (e.g., 6-azauracil)

Culture of Entire Population

Hybrids identified later by:

• Chromosome count

• Isozyme patterns

• Organelle DNA analysis

• Molecular markers
Regeneration of Hybrid Plants

Hybrid cells must:

1. Regenerate cell wall

2. Undergo mitotic division

3. Form callus or somatic embryos

4. Regenerate shoots and roots

Regeneration is often the limiting step in somatic hybridization.

Types of Somatic Hybrids

Symmetric Hybrids

• Retain full chromosome complements of both parents

• Allopolyploid in nature

• Example: Potato × Tomato (“Pomato”)

Applications:

• Widening genetic base

• Novel species formation

• Direct gene transfer

Asymmetric Hybrids

• Chromosome elimination of one parent occurs

• Partial introgression of genetic material

Useful for:

• Targeted gene transfer

• Disease resistance incorporation

Cybrids (Cytoplasmic Hybrids)

Contain:

• Nuclear genome of one parent

• Cytoplasmic organelles from both parents

Produced by:

• Fusion with enucleated protoplasts (cytoplasts)

 • Nuclear inactivation by irradiation

• Chromosome elimination during mitosis

Hybrid Cell Selection and Identification in Somatic Hybridization

(Concise, scientific presentation)

After protoplast fusion, the resulting population is heterogeneous and consists of:

• Unfused parental protoplasts

• Homokaryons (fusion between same genotype)

• Heterokaryons (true hybrid cells)

Since heterokaryons occur at low frequency (generally 0.5–10%), efficient selection systems
are essential.

Hybrid Cell Selection

Biochemical (Selective) Methods

These methods exploit genetic or metabolic differences between parental cells.

(a) Drug/Antibiotic Resistance

One parent is resistant to a selective agent (e.g., actinomycin D, kanamycin), while the other
is sensitive.

• Sensitive parent → cannot grow on selective medium

• Resistant parent → may survive but often lacks regeneration capacity

• Hybrid → combines resistance and regeneration ability

Thus, only hybrid cells proliferate on selective medium.

(b) Auxotrophic Complementation

Auxotrophic mutants cannot synthesize essential metabolites and fail to grow on minimal
medium.

If:

• Parent A lacks pathway X

• Parent B lacks pathway Y

Neither parent grows on minimal medium, but the hybrid restores both pathways through
genetic complementation and survives.

This method provides strong and specific selection.

Visual Selection

Used when parents differ in visible traits.

(a) Pigmentation Markers

• Green (chlorophyll-containing) vs albino

• Anthocyanin-pigmented vs non-pigmented

Hybrids show combined or intermediate pigmentation.

(b) Fluorescent Labeling

Parents are labeled with different fluorescent dyes.

Hybrid cells exhibit dual fluorescence and can be detected microscopically or sorted.

Limitation: Labor-intensive and less efficient for large populations.

Cytometric Techniques

Modern methods improve precision and efficiency.

Flow Cytometry / FACS

• Fluorescently labeled protoplasts are analyzed by laser detection.

• Dual-labeled cells are sorted automatically as hybrids.

Advantages:

• Rapid

• Quantitative

• High throughput

Identification of Hybrid Plants

After regeneration, hybridity must be confirmed using multiple criteria.

1. Morphological Analysis

Hybrids often show intermediate traits between both parents:

• Leaf morphology

• Growth habit

• Flower characteristics

Useful as preliminary evidence but not definitive.

Chromosomal Constitution
Cytological examination (chromosome counting):

• Symmetric hybrids → sum of parental chromosomes (amphidiploid)

• Asymmetric hybrids → partial chromosome elimination

• Cybrids → nuclear genome of one parent only

Provides direct cytogenetic confirmation.

Isoenzyme Analysis

Isoenzymes are different molecular forms of the same enzyme.

Electrophoretic banding patterns show:

• Parent A bands

• Parent B bands

• Hybrid exhibits bands of both parents simultaneously

Confirms biparental nuclear contribution.

Molecular Techniques

Highly reliable confirmation methods:

• PCR with species-specific primers

• RFLP analysis

• RAPD, AFLP, SSR markers

• rDNA analysis

• Organelle DNA analysis (mitochondrial and chloroplast)

Hybrids display nuclear DNA markers from both parents and may show recombinant
organelle genomes.

Regeneration of Hybrid Plants

Hybrid cells must:

1. Regenerate cell wall

2. Undergo mitotic division

3. Form callus or somatic embryos

4. Regenerate shoots and roots

Regeneration is often the limiting step in somatic hybrids

Applications of Somatic Hybridisation:

• Crop Improvement: Somatic hybridisation is used to develop crop varieties with

desirable traits such as disease resistance, stress tolerance, and improved yield.

• Genetic Diversity Enhancement: By combining genetic material from different plant

species, somatic hybridisation contributes to the enrichment of genetic diversity in
breeding programs.

• Creation of Novel Varieties: This technique enables the creation of novel plant
varieties with unique combinations of traits not found in traditional breeding
programs.

• Biotechnological Research: Somatic hybridisation is instrumental in studying gene

expression, functional genomics, and understanding the mechanisms of plant
development and stress responses.

• Disease Resistance: Disease-resistance genes have been able to spread from one
plant to many others due to somatic hybridisation. The spotted wilt virus, TMV,
insect pests, and cold tolerance can no longer harm tomatoes.

Somatic Hybridization Examples

• Pomato: Created by merging tomato and potato protoplasts, resulting in a single

plant that produces both tomatoes and potatoes.

• Wheat-Agropyron hybrids: Used to transfer genes for disease resistance and

environmental adaptation from wild grasses to cultivated wheat.

• Citrus intergeneric hybrids: Generated by fusing protoplasts from different citrus

species, producing hybrids with improved fruit quality and disease resistance.

• Brassica napus (rapeseed) hybrids: Utilized to introduce novel traits like herbicide
resistance and increased oil content from related Brassica species.

• Banana breeding: Employed to combine traits for disease resistance and improved
fruit quality from different banana varieties.

Advantages of Somatic Hybridization

• Somatic hybridization can be performed on young, immature plants.

• It is now simple to research cytoplasmic genes and how they work.

• Somatic hybridization was used to transfer genes into plants to provide

environmental tolerance against cold and frost.

• Selective somatic hybrids with quality traits, such as the generation of high nicotine
concentration, have been created.
 • In the hybrid cell, the protoplast fusion results in a distinctive nuclear-cytoplasmic
combination.

• It produces novel plants with desirable traits.

• An alternate technique for creating remote hybrids with desirable qualities

considerably across species or genera, which cannot be achieved by the traditional
approach of sexual hybridization, is somatic cell hybridization, parasexual
hybridization, or protoplast fusion.

Limitations of Somatic Hybridization

• The created plants are not always healthy and fruitful.

• The regenerated plants produced through somatic hybridization occasionally display

diversity for a variety of causes, including soma clonal variations, chromosomal
elimination, organelle segregation, etc.

• However, it does not necessarily result in the production of viable seeds, making the
fusion of distant plant genera feasible.

• Genetic instability can occasionally result from protoplast fusion.

• The effective manifestation of a certain characteristic is not necessarily ensured by

somatic hybridization.

• There are only a few selection criteria.

Development of Hybrids

Hybrids are the first generation (F1) from a cross between two pure lines, open-pollinated
varieties or clones that are genetically dissimilar. Most of the commercial hybrids are F1s
from two or more pure lines (tomato, rice, Jowar) or inbred lines (maize, sunflower, caster
etc.)

Pure line: It is the progeny of single self-fertilised homozygous plant.

Inbred line:

It is a near-homozygous line obtained by continuous inbreeding in a cross-pollinated crop

followed by selection.

Single cross: When two inbred lines or pure lines are crossed to produce the Fl hybrid, it is
known as a single cross.

Double cross: when two single crosses are crossed, the resulting hybrid population is known
as a double cross.

Three-way cross: It is a cross between a single cross and an inbred to give a hybrid
population. Top cross: When an inbred is crossed with an open-pollinated variety it is known
as an inbred variety cross or a top cross. The purpose of top cross is to estimate the GCA of
the inbred line crossed with OPV. When the cross is made to assess the combining ability, it
is known as a test cross.

Test cross: A test cross may be made with an inbred (for SCA), hybrid, synthetic or OPV (for
GCA). The common parent used in the test cross is known as the tester, and the progeny
derived from these crosses are known as test cross progeny.

Polycross: It is the progeny of a line produced through random pollination by a number of

selected lines.

Varietal cross: when two open pollinated varieties are mated it is known as varietal cross or
population cross.

Synthetic Variety: Definition, Production (Operations) and Merits

The idea of developing and commercially using synthetic varieties in maize was first
proposed by Hayes and Garber in the year 1919. Over time, synthetic varieties became
important in the improvement of many cross-pollinated crops, especially where controlled
pollination is difficult, expensive, or impractical on a large scale such as alfalfa, red clover,
different forage grasses, maize, ryegrass, and other forage crop species.

A synthetic variety is a cross-pollinated variety developed by intercrossing a number of

selected inbred lines or clones having good general combining ability (GCA) and
subsequently maintained by open pollination.

They are popular because:

• They maintain a high degree of heterozygosity.

• They preserve genetic diversity.

• They are stable under variable agro-climatic conditions.

• Farmers can reuse seed for several generations.

• They show moderate hybrid vigour, which improves yield without requiring yearly
purchase of hybrid seed.

Essential Characteristics of Synthetic Varieties

 • Genetically heterogeneous and heterozygous

• Developed from multiple parents (usually 5–10)

• Express moderate but stable heterosis

• Performance remains fairly constant over generations

• Maintained by random mating

• Mainly used in cross-pollinated crops

Production of Synthetic Varieties (Operations Involved)

The production of a synthetic variety involves a series of well-defined operations,

combining selection, testing, intercrossing, bulking and maintenance.

A. Selection of Parental Material:

• 5–10 inbred lines or clones are selected

• Selection criteria: High yield potential, Resistance to diseases and pests, to abiotic
stresses, Wide adaptability, Good agronomic performance

• Parents must show good general combining ability, not just individual superiority.
Good GCA ensures that each parent contributes favourably when crossed with
others.

Parent Material Used for Producing Synthetic Varieties

Synthetic varieties can be produced from many different types of genetic material such as:

1. Inbred lines

• These are genetically uniform lines obtained after several generations of self-
pollination. Inbred lines are highly homozygous and genetically stable. Only those
inbred lines that exhibit high levels of general combining ability are selected.

2. Clones

• Used mainly in crops that are vegetatively propagated, especially cross-pollinated

forage grasses or perennial crops.

3. Open-pollinated varieties

• These are genetically heterogeneous populations that have been maintained by

natural cross-pollination for many generations.

4. Genotypes obtained through recurrent selection

 • These genotypes have been improved through repeated cycles of selecting superior
individuals and intercrossing them, resulting in increased frequency of favourable
alleles.

• Normally, five to eight genotypes with proven high general combining ability are
selected to produce a synthetic variety, though in some cases the number may range
from two to ten genotypes.

B. Testing for General Combining Ability (GCA),

The purpose of this stage is to determine which parental genotypes possess the ability to
combine favourably with many other genotypes.

Genotypes used in this evaluation include: Inbred lines, Clones, Plants selected from open-
pollinated varieties, Short-term inbred lines derived from recurrent selection programmes

Why evaluate general combining ability?

Because synthetic varieties exploit the portion of hybrid vigour that is associated with
general combining ability, not specific combining ability.
Therefore, only genotypes with high general combining ability are used.

Methods used for evaluating general combining ability:

1. Top-cross method

• Each inbred or genotype is crossed with a common tester.

• The resulting progenies are evaluated in replicated field trials for Yield, Traits
contributing to yield, Growth performance

2. Polycross method

• Selected genotypes are grown together in an isolated block.

• These genotypes are allowed to intermate naturally using open pollination.

• This method is widely used in perennial forage species.

3. Single-cross method

• All possible single crosses among the selected genotypes are produced.

• Each cross is evaluated for general combining ability.

All genotypes with high general combining ability are selected for use in forming the
synthetic variety.

C. Intercrossing of Selected Parents

• Parents with proven GCA are intermated

 • Methods: All possible controlled crosses among parents, or Growing parents
together in a polycross nursery

• Self-pollination is prevented

• Ensures random mating among selected parents

Significance: Creates maximum recombination and heterozygosity.

D. Seed Collection and Bulking

• Seeds harvested from all intercrossed parents

• Equal quantity of seed from each parent or cross is mixed

• Bulked seed constitutes Synthetic-1 (Syn-1) generation

Syn-1 represents: Maximum genetic diversity, Highest heterozygosity, Initial expression of

heterosis

E. Maintenance and Seed Multiplication

• Syn-1 is grown in isolation

• Maintained by open pollination

• Subsequent generations: Syn-2, Syn-3, etc.

• Periodic roguing of off-types carried out

• Seed from later generations is supplied as commercial seed

Although heterosis slightly declines over generations, overall performance remains
stable.

Merits of Synthetic Varieties

1Yield Stability: Yield remains consistent across seasons, Less sensitive to environmental
fluctuations

2. Seed Reusability: Farmers can reuse seed for several generations, Unlike hybrids, no
compulsory annual seed replacement

3. Wide Adaptability: Performs well under diverse agro-climatic conditions

4. Genetic Diversity: Broad genetic base, Better buffering against Diseases, Pests, Abiotic
stresses

5. Cost Effectiveness: Lower seed cost. Suitable for small and marginal farmers

6. Ease of Seed Production: No complex hybrid seed production system. Easier maintenance
compared to hybrids
7. Suitability for Low-Input Agriculture: Ideal for Rainfed areas and Resource-poor farming
systems

Demerits

• Yield lower than single-cross hybrids. Lower yield compared to the best single-cross
and double-cross hybrids.

• Can only be produced in cross-pollinated crops.

• Strongly dependent on the general combining ability of parents. Less uniform than
hybrid varieties.

• Moderate, but not maximum, exploitation of hybrid vigor.

• Gradual reduction in heterosis over generations

• Not suitable where very high yield is the sole objective

Examples of Crops with Synthetic Varieties: Maize, Sorghum, Pearl millet, Fodder crops and
grasses

Synthetic varieties represent a balanced breeding approach, combining moderate heterosis,

yield stability, genetic diversity and seed economy. They are particularly valuable for
sustainable agriculture, especially under rainfed and low-input conditions, and serve as an
effective alternative to hybrids for resource-constrained farmers.

Achievements of Synthetic Varieties (Exact Notes)

• Synthetic varieties have been developed in cross-pollinated crops like maize, pearl
millet, sunflower, sugarbeet, alfalfa, lucerne, etc.

• In India synthetic varieties have been evolved in pearl millet at the International
Crops Research Institute for the Semi-Arid Tropics and in sugarbeet at Pantnagar.

• Example:
Sugarbeet – plant synthetic – 3
Cauliflower – synthetic – 3
Bajara – ICMS-7703

COMPOSITE VARIETY

• In cross-pollinated crops, the mixture of genotypes from several sources is

maintained in bulk from one generation to the next is referred as a composite
variety.

• Composite variety is a variety derived from advanced generation of random-mated

outstanding lines (germplasm inbreds, varieties, hybrids, advanced generation lines).
 • Mixing the seeds of several phenotypically outstanding lines produces a composite
variety, and encouraging open pollination to produce crosses in all combinations
among the mixed lines.

• The lines used to produce a composite variety are rarely tested for combining ability
with each other like synthetic composite are commercial varieties and are
maintained by open-pollination in isolation.

• The line used to produce a composite variety are rarely tested for combining ability
with each other.

• Mixing the seeds of various genotypes, which are similar in maturity height, seed
size, colour, etc. develops composite varieties.

• The variety is maintained by open pollination. Farmers can use their own seed for 3
to 4 years.

• The yields of composite varieties cannot be predicted in advance, which is contrast to

synthetic; this is because of yield of all first-generation crosses among the
component lines are not available. Like synthetic, composite are commercial varieties
maintained by open pollination in isolation. Another type of population, called
germplasm complexes are produced by mixing seeds from several lines or
populations of diverse genetic origin.

Procedure for Development of Composite Variety

• There is no restriction to the number of lines included in the development of

composite, but the line possessing desirable characteristics should be selected like
earliness, insect resistance, drought and frost resistance can be included in the
development.

• The seeds of desired selected lines mixed together and random mating is allowed for
4–5 generations.

• Such uniform population is tested in replicated trials, across environment along with
standards checks and high yielding stable type can be released as composite variety.

• The combining ability of lines used in composite are not tested, whereas in synthetic
each of the component is tested for its combining ability.

Features of Composite Variety

• Heterogeneous.

• Relevant to cross-pollinated species only. Can be developed from an open-pollinated

variety or any other heterozygous variety.
 • A farmer can use his own saved seed for 3 to 4 years; after that seed should be
replaced.

• There can be two or more constituent genotypes.

Achievement of Composite Varieties

• In India six composite varieties of maize like Amber, Jawahar, Kisan, Vikram, Sona
and Vijay were released in 1967.

• Subsequently number of composite varieties has been released in bajra and other
crops.

• Some of the released composite varieties of maize are Co-1, NLD, Renuka and
Kanchan, Diara-3 etc., while in bajra released composite are WCC-75, RCBIC-9, ICTP-
8203.

Synthetic Varieties Composite Varieties

Crossing in all combination of Mixing the seeds of several phenotypically outstanding

number of lines that combine well lines and encouraging open pollination to produce
with each other. crosses in all combination among the lines.

4–10 numbers of parents are

Many parents are involved
involved

It is tested for general combining

Not tested for general combining ability
ability

Cost of seed less than composite

Cost of seed is less than hybrid.
variety.

It shows less heterosis than

It shows more heterosis than synthetic
hybrid.

Reconstituted is done. Reconstituted is never done.

Maintenance of variety easy. Maintenance of variety is difficult but easy.

Prediction of performance
Prediction of performance is not possible.
possible.