LS - An International Journal of Life Sciences

DOI : 10.5958/j.2319-1198.2.2.012

Pre-breeding and Population Improvement

Asif M Iqbal, Ajaz A Lone, Shafiq A Wani, Shabir H Wani*, FA Nehvi

1
Division of Plant Breeding and Genetics, Sher-e-Kashmir University and Agricultural Sciences and Technology of
Kashmir, Shalimar- 291123 Jammu and Kashmir, India.
Email id: [email protected]

ABSTRACT

Pre-breeding refers to all activities designed to identify desirable characteristics and/or genes from
unadapted (exotic or semi-exotic) materials, including those that, although adapted, have been subjected
to any kind of selection for improvement. It is an essential part of germplasm diversification strategies.
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Pre-breeding is the most promising alternative to link genetic resources and breeding programmes. As
pre-breeding is being carried out, the resulting materials are expected to have merit to be included in
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ordinary breeding programmes. There are at least three distinct uses of genetic enhancement. The first
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is to prevent genetic uniformity and consequent genetic vulnerability. Only recently has pre-breeding-
genetic enhancement-become a necessary, frequent and planned part of all plant breeding activities,
an essential part of germplasm diversification strategies. Genetic enhancement has a second important
purpose that of raising yield levels to new heights. This goal is more often hoped for than achieved, but
it is true that most breakthrough cultivars have highly diverse parentage. The semi-dwarf wheat, the high
yield dwarf rice, the first US hybrid sorghums and even the first US Corn Belt dent maize cultivars are
examples. In each case, extensive pre-breeding preceded development of the breakthrough, high-yield
cultivars. The pre-breeding was used to adapt diverse kinds of germplasm to new genetic backgrounds
and new geographic locales. Genetic enhancement is used to bring in new quality traits not found in local
cultivars. New levels of protein percentage in wheat or unusual starch properties in maize are examples.

1. PRE-BREEDING: A NEW REQUIREMENT IN et al., 1993). The main factors responsible for the
PLANT BREEDING low utilisation of plant genetic resources are lack of
documentation and adequate description of collections,
Activities related to genetic resources are characterised
by high cost and long-term return. Introduction and lack of the desired information by breeders, accessions
germplasm exchange, collection, characterisation, with restricted adaptability, insufficient plant
evaluation, documentation and conservation are breeders, particularly in developing countries, and
essential steps that cannot be overemphasised. An lack of collection evaluations. Low seed availability
appropriate synchronism among these activities is due to inadequate seed regeneration programmes
required for the bank to be effective in maintaining is another barrier to their use (Dowswell et al.,
genetic variability and to assure germplasm utilisation. 1996). Furthermore, breeder-to-breeder exchange of
The importance of genetic resources is widely materials is very common and constitutes a reasonable
recognised. Activities in germplasm banks demand alternative to extend genetic variability in breeding
qualified researchers in several areas of knowledge. programmes. In general, it seems that breeders are
Besides the conservation of genetic variability for the satisfied with the available genetic variability among
future, the actual utilisation of available accessions is agronomically advanced materials (Duvick, 1984;
another important goal. However, the low utilisation Paterniani, 1987; Peeters and Galwey, 1988; Nass et
of germplasm banks is a rule worldwide and is not al., 1993). According to Troyer (1990), actually elite
restricted to India or to developing countries (Nass inbred lines are considered the best genetic resources

Volume
188 2, Number 3, September-December, 2013, pp. 188-197 LS - An International Journal of Life Sciences
 Pre-breeding and Population Improvement

simply because each line contains a combination of collections of adapted varieties and genetic stocks
genetic traits that satisfies the marketplace. carrying defined traits through to land-races and wild-
The search for superior genotypes regarding yielding crop relatives. However, this richness of plant genetic
ability, disease and pest resistance, stress tolerance or diversity is greatly under-utilised, with important traits
better nutritional quality is very hard, competitive and not being exploited and the potential value not being
expensive. This is why breeders tend to concentrate channeled back into societal benefit. The conservation
to adapted and improved materials, avoiding wild of genetic resources must be linked to their increased
parents, landraces and exotics, available in germplasm and sustainable use if they are to play a key role in
banks, which would require long-time, high financial climate change adaptation. Bottlenecks that need to
support besides the difficulty to identify potentially be addressed include lack of information on genotypic
useful genes. Marshall (1989) emphasised that the and phenotypic level, e.g., need for evaluation for
difficulty to identify useful genes is the main factor resistance to biotic and abiotic stresses of increasing
responsible for the low utilisation of these accessions. importance. Data need to be easily accessible in
Evidently, there is a gap between available genetic standardised and searchable electronic formats
enabling strong networks linking conservation,
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resources and breeding programme activities. While

germplasm banks try to preserve as much as possible evaluation and plant breeding to be established to
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the genetic variability to be used by breeders, breeding secure the necessary level of pre-breeding activities.
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programmes do not explore efficiently the available Finally, long-term commitments and funding from
diversity, relying almost exclusively on their working both public and private stakeholders are needed if this
collection. In this way, to develop new populations is to have a major impact on European food security.
and inbred lines, commercial F1 hybrids are being
widely used, since this is a legitimate source of 3. ABIOTIC STRESSES: ADAPTING TO
parental germplasm (Parks, 1993). However, the ENVIRONMENTAL CHANGES
recurrent use of these genotypes should increase
the narrowness of the genetic base. In a short time, Tolerance to particular abiotic stresses such as
available commercial hybrids probably will be drought, cold, salinity, heat, water logging, nutrient
very similar since they partake the same genetic use efficiency and mineral toxicity has traditionally
background. Genetic vulnerability must be a constant enabled cereal crops to cope with the prevalent local
concern in plant breeding for all species. Theoretical stresses with a balance of traits that were fine tuned
expectations and actual experience show that the use to optimise economic yields in their environments.
of uniform materials can be dangerous, as occurred However, global warming is producing shifts in the
with cytoplasm T, susceptible to Helminthosporium prevalence of some abiotic stresses with drought being
maydis race T in the United States in 1970 and next increasingly important, given the widely acknowledged
year in other regions of the world effect of climate change on the amount of precipitation
and its distribution over the growing seasons in Europe.
2. SUSTAINABLE USE OF GENETIC DIVERSITY In addition, ozone levels in the troposphere will rise
notably in the near future, affecting the amount of
Gene banks have often, through necessity, focused ultraviolet radiation that reaches the Earth surface in
mainly on the immediate conservation aspects of plant turn causing large increases of plant oxidative stress.
genetic resource activities. However, there is an urgent Therefore, it is expected that the agricultural areas
need for active engagement with all stakeholders to of Europe will experience enhanced or novel abiotic
enhance the utilisation of plant genetic resources stresses making it increasingly urgent to develop
in order to assure the functionality of the entire cereals that can withstand such environmental
“Genetic resource-chain”. There are considerable changes, in order to increase or even just to maintain
genetic resources for small grain cereals including current yield levels.

Volume 2, Number 3, September-December, 2013 189

 Asif M Iqbal, Ajaz A Lone, Shafiq A Wani, Shabir H Wani, FA Nehvi

4. BIOTIC STRESS: COPING WITH EMERGING 4. Backcrossing the hybrids with agronomically
DISEASES acceptable base as recurrent parent.
Plant pathogens cause considerable yield losses 5. Chromosome doubling when genome non-
in cereal production, reducing crop quality and homology is present (4n or 6n pathway) and
threatening food safety. Disease prevention and backcrossing.
control is thus a prerequisite for competitive cereal 6. Chromosome techniques such as translocations,
production with the breeding of genetically disease- alien additions lines (MAAL and DAAL),
resistant crops being one of the most environmentally substitutions, homoeologous pairing and also
and economically desirable ways to manage plant irradiation to induce interchanges and gene
diseases. transfer.
The prevalence of different plant diseases is changing
due to changing environmental conditions, including 4.2. ALLOPLASM DEVELOPMENT
global climate change, but also changes in agricultural
production with trends towards larger areas Transfer of cytoplasm (mt and cp genomes) from
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planted to fewer and/or genetically more uniform interspecific and intergeneric origin greatly enables
varieties, reduced crop rotation and soil tillage, the exploitation of cytoplasmic genetic male sterility
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loss of biodiversity, changing use of pesticides and system (CMS)- male fertility system. Contributions
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global trade. In such a changing environmental and of cytoplasms of Triticum timopheevi, Gossypium
economical context, plant diseases will inevitably harknessii, Helianthus petiolaris, Oryza nivara and
appear and compromise crop production in regions Cajanus scarabaeoides in the development of CMS
where they did not represent a problem before, as is for harnessing heterosis are significant. Breeders
already being seen with the emergence of a new strain also needto search for many useful traits from exotic
of heat-tolerant wheat yellow rust and increasing variation that are not often considered in germplasm
problems with Fusarium head blight and Ramularia enhancement. They are physiological traits such
leaf spot. Understanding the host-pathogen biology is as enhanced growth rate, improved source sink,
the first step towards minimising the risks represented stabilising high-yield potential, insulating the preferred
by plant diseases. Durable, both race non-specific lines from environmental stresses, conferring high
and race-specific, resistance incorporated into high- adaptation and enhancing the quality.
yielding genotypes is the main method to manage
diseases of cereals. New durable and efficient sources
4.3. GERMPLASM ENHANCEMENT: GENE
of resistance will have to be sought, in the case of
TRANSFER
cereals, from landraces and from wild relatives of the
crops. 4.3.1. Wheat
Interspecific and intergeneric gene transfers were
4.1.1 TECHNIQUES FOR ALIEN GENE TRANSFER successfully done into wheat. Homologous and
Different techniques are followed to overcome barriers homoeologous pairing of chromosome induced
to wide hybridisation and to obtain viable hybrids to polyploidy and chromosome substitutions have been
affect gene transfer: frequently used mechanisms for introgression within
and between the gene pools. Different genes for
1. Embryo rescue and in vitro techniques. leaf and stem rust, stripe rust and powdery mildew
2. Involving bridge species, which are crossable with resistance (Table 1) were transferred to hexaploid
both the parental species. wheat from different Triticum sp and Aegilops, e.g.,
3. Application of exogenous plant growth regulators stem rust resistance was transferred from three or
and immunosuppressors at post pollination stage. more species:

190 LS - An International Journal of Life Sciences

 Pre-breeding and Population Improvement

Table 1: Disease resistance genes and their source homoeologous pairing as shown below (Table 4).
Genes Source Table 4: Genes transferred for disease and insect resistance
Sr 21, Sr 22 Triticum monococcum Species Genes transferred
Sr 2, Sr 9e, Sr 13 T. dicoccum O. australiensis BLB, BPH
Sr 14, Sr 17, Sr 33 Ae. squarrosa
O. brachyantha BLB
Germplasm enrichment of wheat by transfer of genes
O. latifolia BLB, BPH, WBPH
from Secale through translocations has enabled gene
O. minuta Blast, BLB, BPH
transfer for disease/insect resistance (Table 2). A few
examples are: O. officinalis BPH, BLB, WBPH, GLH
O. nivara Grassy stunt
Table 2: Transfer of genes through translocationsfor disease and
insect resistance O. perennis CMS
Translocations Resistance for Gene transfer conferring resistance to YSB and leaf
1B. R - Tr PM; stem, leaf and stripe folder is in progress.
2BS. 2RL - Tr Hessian fly
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4BL. 5 RL - Tr High copper tolerance 4.4. YIELD ENHANCEMENT

Irradiation-induced translocations Hessian fly
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Introgression from wild taxa leading to the yield

Homoeologous pairing-mediated gene transfer
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enhancement in the backcross derivatives of Avena,

was also successful for transfer of genes from A. Sorghum, pearl millet and tomato has been indicated
longissimum, Ae. umbellate and Ae. elongatum to (Zamir et al., 2001). In chickpea and chilli also such
wheat. yield enhancement through introgression from wild
Likewise, chromosome substitution method (Table related species has been reported. In tomato, systematic,
3) has helped for gene introgression in the following concentrated and mission-oriented programmes (Table
crops. 5) have achieved successful germplasm enhancement
Table 3: Gene introgression through chromosome substitution for many traits.
method Table 5: Genes transferred for biotic and abiotic stress tolerance
Sl. No. Resistance in tomato
1 Triticum aestivum × Haynaldia Powdery mildew Species Resistance genes transferred to tomato
2 Trifolium repens × T. nigrescens Cyst nematode L. hirsutum Fus. Wilt, Leaf Curl Virus, Whitefly
3 Beta vulgaris × Brassica. species. Nematode L. pimpinellifolium* Fus. Wilt, early blight
4 Cicer arietinum × C. reticulatum Cyst nematode L. chilense CMV
5 Glycine max × G. tomentella Cyst nematode L. cheesmanii High temperature, Salt
(homoeologous) L. peruvianum Nematode
High grain number per spike, high protein content, L. pennellii* Salt, potato aphid, high brix
temperature tolerance, Karnal bunt resistance and *Fruit and juice quality traits were also introgressed from these
drought tolerance are to mention some of the other two species to tomato. Homoeologous chromosome pairing was
economic traits that have been introduced from alien instrumental in gene transfer.
sources into T. aestivum wheat. The introgressed lines in tomato for various traits were
again crossed with each other lines leading to staking of
4.3.2. Rice different genes. The pyramiding has resulted in multiple
Genes conferring resistance to insects and disease and resistance breeding lines with elevated yield potential
other traits are introgressed (Table 3) as germplasm- and highly acceptable fruit and juice quality traits. Such
enhancement activity into cultivated rice through lines can be either prebred or utilised as parent lines for
hybrid development or for cultivar identification.

Volume 2, Number 3, September-December, 2013 191

 Asif M Iqbal, Ajaz A Lone, Shafiq A Wani, Shabir H Wani, FA Nehvi

In a study in cotton, the introgressed lines from various specific traits and for base broadening in pre-breeding.
wild taxa - sources were assembled and the useful Molecular techniques and bioinformatics allow more
variation was assessed for agronomic traits and lint precise and faster selection methodologies as well
qualities. Alloplasmic (cytoplasmic) lines interaction as providing a much more detailed understanding
with G. harknessi introgressed restorer parents for the of the underlying genetics. Such efficient tools are a
agronomic and yield in the hybrids have also been crucial element in the modern plant breeding process
assessed. The results (Tuteja et al., 2006) indicated to handle complicated traits efficiently. While costs
the promising nature of the introgressed lines as of molecular genotyping are constantly decreasing
new sources for hybridisation to further diversify and methods becoming more efficient, reliable and
the adapted varieties. Some of the introgressed lines precise phenotyping are costly, time consuming and
showed promise as parents in new hybrid combinations an increasing challenge. The strategy for the future
not brought out so far. should contain a broad knowledge of and access to
Domestication and continued selection result in narrow modern technology, and combining the application of
genetic base. Returning back to wild ancestors and new tools and techniques with traditional and efficient
plant breeding methods to achieve final goal rapidly.
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utilising them as donors will be helpful and rewarding.

Germplasm enhancement is an effective strategy
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for accomplishing gene transfer to the cultivated 6. POPULATION IMPROVEMENT

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background. Germplasm, those from related taxa and

progenitors wild relatives, have many useful genes and In self-pollinated crops, selection is employed to
exotic variation for almost all the traits as compared isolate plants with superior genotypes; these plants are
to the cultivar variation. A critical understanding, then used to establish separate pure lines or their seeds
evaluation and documentation of exotic variation are a are bulked to produce a mixture of pure lines. This
prerequisite to plan for transferring useful and desired is possible because self-pollinated crops are naturally
gene complexes to cultivated agronomic background. homozygous and generally do not show appreciable
inbreeding depression. In contrast, cross-pollinated
Advanced backcrosses and/or introgressed in breds crops generally show moderate-to-severe inbreeding
developed by germplasm enhancement are useful depression. Consequently, (1) inbreeding must be
improved derived prebred lines in cultivar and hybrid avoided or kept to a minimum in cross-pollinated
development. DNA markers and Quantitative Trai species. Further, (2) individual plants from such crops
Loci identification accelerate the genetic enhancement are highly heterozygous; the progeny from such plants
progress in a precise manner. Many QTLs that have would be heterogeneous and usually different from
the potential for improving yield have been transferred the parent plant due to segregation and recombination.
from exotic sources and staked on to the cultivar. Yield Therefore, desirable genes can be seldom fixed through
variation due to additivity of cultivated and exotic selection in cross-pollinated populations, except for
genes could be inferred and even dramatic increase in qualitative traits and perhaps, for easily observable
yield was evident. This is a revitalising effort in crop quantitative characters with high heritability. (3) The
improvement. The exotic QTLs conferring resistance breeder, therefore, aims at increasing the frequency
to biotic and abiotic stresses and other kinds of traits of desirable alleles in the populations. (4) This would
can be alternative to Genetically Modified Organisms result in an increase in the frequency of desirable gene
(Gur and Zamir 2004). combinations or genotypes. As a result, the phenotype
of the population would be favourably changed.
5. NEW TOOLS TO ADDRESS THE CHALLENGES
It should be clearly understood that in cross-
The recent progress in biotechnology has opened pollinated species the genotype of the individual
up enormous possibilities, both for introgression of plants is generally of little importance, particularly

192 LS - An International Journal of Life Sciences

 Pre-breeding and Population Improvement

in population improvement programmes. It is Mass selection, as applied to cross-pollinated crops, is

the frequency of desirable genes or alleles in the essentially the same as that applied to self-pollinated
population as a whole that determines the value of crops. A generalised scheme for the method is outlined.
population. Accumulation of desirable alleles in a The selection cycle may be repeated one or more times
population through various breeding techniques is to increase the frequency of favourable alleles; in such
known as population improvement and those breeding a case, the selection scheme is generally known as
procedures that are used for such work are referred to phenotypic recurrent selection. Care should be taken
as population improvement approaches. to select a sufficiency of mass selection primarily
depends upon the number of genes controlling the
7. METHODS OF POPULATION IMPROVEMENT character, gene frequencies and, more importantly,
heritability of the concerned trait.
The various breeding methods used for population
improvement may be grouped into the following two
general classes: (1) methods without progeny testing 11. MODIFICATIONS OF MASS SELECTION
and (2) methods with progeny testing. The two basic defects of mass selection, viz., (1) lack
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of control of the pollen source and (2) the confusing

effect of environment on the phenotypes of individual
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8. BREEDING METHODS WITHOUT PROGENY

plants, can be corrected by suitable modifications of
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TESTING
the selection scheme; these modifications are briefly
In breeding methods belonging to this group, plants
described below.
are selected on the basis of their phenotype, and no
progeny test is carried out, e.g., mass selection. 1. Inferior plants in the field are detasselled, and the
remaining plants are allowed to open-pollinate.
This modification exercises some control on the
9. BREEDING METHODS WITH PROGENY
pollen source, but the identification of inferior
TESTING
plants, of necessity, is based on only those
In most breeding methods, however, plants are initially characters, which are expressed before flowering.
selected on the basis of their phenotype, but the final 2. Pollen from all the selected plants is collected
selection of plants that contribute to the next generation and bulked; this pollen is used to pollinate the
is based on a progeny test. This class is population selected plants. This scheme ensures full control
improvement methods include progeny selection or on the pollen source, but it can be applied to only
ear-to-row method, and recurrent selection.
those characters that can be selected before pollen
shedding.
10. MASS SELECTION
3. Stratified Mass Selection: This modification,
Mass selection is the oldest breeding scheme available suggested by Gardner in 1961, is also known as
for cross-pollinated crops. In mass selection, a number the grid method of mass selection. The field from
of plants are selected on the basis of their phenotype, which selection is to be done, is divided into several
and the open-pollinated seed from them is bulked small plots, e.g., having 40-50 plants each. Equal
together to raise the next generation. The selected number of superior plants is selected from each of
plants are allowed to open-pollinate, i.e., to mate at the small plots, i.e., selection is done within the
random including some degree of selfing (usually plots and not among the plots. The seed from all
about 10% in maize, Z. mays). Thus, mass selection the selected plants is composited to raise the next
is based on the maternal parent only, and there is no generation. The basis for this modification is the
control on the pollen parent. Selection of plants is based consideration that variation due to environment,
on their phenotype and no progeny test is conducted. including heterogeneity in soil fertility, will be

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 Asif M Iqbal, Ajaz A Lone, Shafiq A Wani, Shabir H Wani, FA Nehvi

much smaller within the small plots than that in 12.2. Modification of Ear-To-Row Method
the whole field. Thus, selection within the plots In order to overcome the defect described above, the
is expected to be more effective than that without following modification may be used.
any stratification. Stratified mass selection has
been able to increase the yielding ability of an 1. First year: Several plants are selected on the
open-pollinated variety of maize, Hays Golden, basis of phenotype (as on item 1 above). Open-
by about 3% per cycle (or generation) for 15 pollinated seed of the selected plants is harvested
generations. separately.

4. Plants of a constant genotype, e.g., a single cross 2. Second year: Small progeny rows are grown (as in
hybrid, are planted as checks after every one, two item 2 above) and evaluated. The remaining seed
or four plants of the variety under selection. The from each of the selected plants is kept separately.
yields of plants under selection are expressed as Superior progenies are identified.
per cent of the yield of nearest check plant. This 3. Third Year: The remaining seed from those plants
scheme is designed to minimise the environmental that produced the superior progenies (identified in
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influence on the yields of plants being selected. the second year) is bulked and planted in the third
It employs the principle of contiguous control year; this constitutes the selected version of the
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discussed earlier in some detail. population. Plants are allowed to open-pollinate.

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A number of plants are selected on the basis of

12. SELECTION SCHEMES WITH PROGENY TEST phenotype and the selection cycle may be repeated
one or more times.
12.1. Progeny Selection This modification of ear-to-row method was widely
The simplest form of progeny selection is the ear- used in breeding of maize in the United States of
to-row method, which has been extensively used in America and was responsible for the development of
maize. This method was developed by Hopkins in several varieties. In this method, plants from superior
1902. In its simplest form, the ear-to-row method of progenies only are allowed to mate among themselves.
selection is as follows. But for each selection cycle, 2 years are required as
1. A number (50-100) of plants are selected on compared to only 1 in the case of ear-to-row method.
the basis of their superior phenotype. They are
allowed to be open-pollinate, and the seeds from 13. RECURRENT SELECTION
individual plants are harvested separately.
Recurrent selection is a modified form of progeny
2. A single row of 10-50 plants, i.e., a progeny row, selection. However, it differs from progeny selection
is grown from each selected plant. The progeny in two main aspects. First, the selected plants are
rows are evaluated for desirable characters and self-pollinated in recurrent selection, whereas they
superior progenies are identified. are open-pollinated in progeny selection. Second,
3. Several phenotypically superior plants are selected the progeny of selected plants are intermated in all
from the superior progenies. There is no control possible combinations in this method, whereas they
on pollination; the plants are permitted to open- are open-pollinated in progeny selection.
pollinate.
4. Small progeny rows, as in item 2, are grown from 14. TYPES OF RECURRENT SELECTION
the selected plants, and the process of selection is There are four types of recurrent selection, namely
repeated. (1) simple recurrent selection, (2) recurrent selection
for general combining ability (RSGCA), (3) recurrent

194 LS - An International Journal of Life Sciences

 Pre-breeding and Population Improvement

selection for specific combining ability (RSSCA) and 1. This method is used for genetic improvement of
(4) reciprocal recurrent selection (RSS). Each type is quantitative characters.
used under specific conditions as discussed below. 2. The selection is made on the basis of test cross
14.1. Simple Recurrent Selection performance.
3. A heterozygous tester with broad genetic base
A type of recurrent selection that does not include is used for testing general combining ability.
tester is referred to as simple recurrent selection. Generally, an open-pollinated variety is used as a
It is also known as phenotypic recurrent selection. tester.
This method is an extension of mass selection. Main
features of simple recurrent selection are given below: 4. This method is used for improving general
combining ability of a population for a character.
1. The tester is not used in this scheme.
5. This method is more effective with incomplete
2. It does not measure the combining ability. dominance and less effective with over dominance.
3. The selection is based on phenotype or simple test. 6. This method is used for the improvement of those
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4. This method is useful only for those characters characters that are governed by additive gene
that have high heritability. action.
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5. This method requires only two seasons for the 7. This method requires three seasons or years for
completion of one selection cycle. completion of each cycle of selection as given
In the first year, the superior plants for the character below.
under improvement are selected from the heterozygous First Year
base population. These plants are grown from the Superior plants for the character under
selfed seed and intermating is done among the improvement are selected from the base population.
progeny. The crossed seed is bulked in equal quantity. The selected plants are selfed and also crossed to
This completes original cycle of selection. In the third a heterozygous tester having broad genetic base.
year, bulked seed is grown and superior plants are The selfed seed is kept in cold storage.
selected and selfed like first year. In the fourth year,
progeny of selected plants are grown from selfed seed Second Year
and intermating is done like first year. The crossed The crossed seed is sown and the combining
seed is composited in equal quantity for use in the ability of the selected plants is evaluated and
next cycle of selection. This completes first cycle of plants with good general combining ability (GCA)
simple recurrent selection. Thus, selection cycles may are identified.
be repeated till the desired improvement is achieved. Third Year
The progeny of selected platens with good GCA
14.2. RSGCA are grown from their selfed seed kept in cold
A form of recurrent selection that is used to improve store. These progeny are intermated in all possible
the general combining ability of a population for a combinations and their crossed seed is composited
character and includes heterozygous tester is referred to form a new source population for further
to as RSGCA. It is also known as a half sib recurrent selection. This completes original selection cycle.
selection with heterozygous tester. This is an extension In the same way, another cycle can be completed
of the Jenkin’s (1940 scheme used for development in three years (4th-6th year). This is called first
of short-term synthetics. Main features of this scheme recurrent selection cycle. Many such cycles may
are given below. be made to obtain desired results.

Volume 2, Number 3, September-December, 2013 195

 Asif M Iqbal, Ajaz A Lone, Shafiq A Wani, Shabir H Wani, FA Nehvi

14.3. RSSCA 14.4. RSS

A form of recurrent selection that is used to improve A form of recurrent selection that is used to improve
the specific combining ability (SCA) of a population both GCA and SCA of a population for a character
for a character by using homozygous tester is referred using two heterozygous testers is known as RRS. It is
to as RSSCA. It is also known as a half sib recurrent also termed as recurrent reciprocal half sib selection.
selection with homozygous tester. This method was This scheme was proposed by Comstock et al. in 1949.
originally proposed by Hull in 1945. Main features of Main features of this method are given below.
this scheme are briefly presented below. 1. This scheme is also used for the improvement of
• This method is used for genetic improvement of polygenic characters.
polygenic characters. 2. The selection is made on the basis of the test cross
• The selection is made on the basis of test cross performance.
performance. 3. Two heterozygous populations each of which is the
• A heterozygous tester with narrow genetic base tester for other are used for in this method. The two
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is used for testing specific combining ability. In populations may be designated as A and B.
other words, an inbred is used as a tester. 4. This method is used for improving a population both
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• This method is used for improving specific for GCA and SCA for specific character.
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combining ability of a population for a character. 5. This method is equally effective with incomplete,
• This method is more effective with over dominance complete and over dominance.
and less effective with incomplete dominance. 6. This method is used for the improvement of those
• This scheme is used when a characters that characters that are governed by both additive and
are governed by non-additive (dominance and non-additive gene action.
epistasis) gene action. 7. This method also requires three seasons or years for
• This method requires three seasons or years for completion of each cycle of selection as given below.
completion of each cycle of selection. First Year
The selection procedure of this method is same as for Several phenotypically superior plans are selected
RSGCA except that the tester is an inbred line, in this from population A and B. The pollen of some
case which has narrow genetic base. The differences selected plants of A population is used to cross
in the performance of test crosses are due to difference large number of randomly selected plants of
in their specific combining ability. A comparison of population B. Similarly, pollen of some selected
recurrent selection for GCA and recurrent selection plants of B population is used to cross large
for SCA is presented. number of plants of population A. All the plants of

Comparison of RSGCA and RSSCA

Particulars RSGCA RSSCA
Application Used to improve polygenic traits Also used to improve polygeneic traits
Basis of selection Test cross performance Test cross performance
Tester used Heterozygous Heterozygous
Effectiveness More effective with incomplete dominance More effective with complete and over dominance
Condition of use Used when additive gene action is important Used when non-additive gene action is important
Impact It improves GCA of a character It improves SCA of a character

196 LS - An International Journal of Life Sciences

 Pre-breeding and Population Improvement

population A and B used as pollen parents in the population improvement can be employed in breeding
crosses are selfed. programmes so as to increase the frequency of desirable
Second Year alleles in the population. As a result, the phenotype of
the population would be favourably changed.
The progeny of test crosses made with pollen
parents of A and B populations are evaluated in
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