Selection in cross pollinated crops

In breeding SP crops, homozygous nature of plant is exploited. A pop of SP plants may be

composed of either a single, homozygous genotype, or a mixture of homozygous genotypes.
Plant selection is effective only within populations that are mixtures of genotypes. Mass
selection for a particular phenotype in a mixed population of a SP crop will reduce genetic
variability and increase the frequency of genes affecting the characters being selected. Pure line
selection involves isolation of a single, best pure line Breeder may resort to hybridization among
selected pure lines to promote recombination and select desirable recombinants
In breeding CP species, heterozygous nature of individual plant exploited. In pop of a CP sps
each plant has both homo-, and heterozygous loci, but it is heterozygous loci that give CP plants
their characteristic genetic structure. As a consequence of natural CP, the genes are reshuffled
each generation and regrouped into new genetic combinations. With an almost limitless number
of gene combinations possible within the gene pool, almost never two plants be found with
identical genotypes.Under natural environmental influences, CP populations are relatively fluid,
genes favouring adaptation and increased seed production tend to increase at the expense of
genes unfavourable for adaptation or fitness to reproduce. In a breeding population, the shift
toward more adapted genotypes may be accelerated by selection, and by environmental stresses
to which the breeding population is subjected
In CP crops, the focus of breeder is on populations instead of individual plant. A population is a
group of sexually interbreeding individuals. The capacity to interbreed implies that every gene
within the group is accessible to all members through the sexual process. All the individuals in a
population share the common gene pool. A gene pool is the total number and variety of genes
and alleles in a sexually reproducing population that are available for transmission to next
generation. Due to extensive heterozygosity in CP crops, there is an abundance of phenotypic
variation; hence cultivars of CP crops less uniform than SP crops. Genetic variability for
qualitative characters may be drastically reduced by rigid selections, but genetic variation in
quantitatively inherited characters continues to be present, due to inability of breeder to select
accurately for individual gene effects and to the influence of GxE interactions
Important difference between SP and CP crops, how breeding materials is evaluated. SP crop,
homozygous genotype is true breeding, evaluated by progeny tests. In CP crops, individual
plants are heterozygous, & largely pollinated by pollen from other plants, genotype is not
faithfully reproduced in progeny. A better test would be, if plant is pollinated with a
heterozygous/homozygous collection of pollen of known origin. Performance can then be
compared among progenies of plants pollinated with same source of pollen. A test comprising
progeny performance of plants or strains pollinated with a known tester line is known as a test
cross and evaluates the combining ability of mother plants or strains with the common tester line.
Average or overall performance of a plant or a genetic strain in a series of crosses with different
tester lines is measure of its general combining ability, whereas performance of a plant or genetic
strain in a specific combination in comparison to the performance of other cross combinations is
a measure of SCA. Combining ability tests used to identify desirable combinations of inbred
lines to develop hybrid cultivars or synthetic cultivars
HARDY WEINBERG LAW: Given by Hardy (1908) and Weinberg (1909). The law states that in a
large random mating population, gene and genotype frequencies remain constant generation after
generation in the absence of selection, mutation, migration or random drift. Consider a large population
in which random mating occurs, with no mutation or gene ow between this population and
others, no selective advantage for any genotype, and normal meiosis. Consider one locus, A,
with two alleles, A and a. The frequency of allele A in the gene pool is p, while the frequency of
allele a is q. Also, p + q = 1
Assume a population of N diploids (have two alleles at each locus) in which two alleles (A, a)
occur at one locus. Assuming dominance at the locus, three genotypes AA, Aa, and aa are
possible in an F
2
segregating population.
Assume the genotypic frequencies are D (for AA), H ( Aa), and Q (aa). In diploid population
there will be 2N alleles in it. The genotype AA has two A alleles.
Hence, the total number of A alleles in the population is calculated as 2D + H.
The proportion or frequency of A alleles (p) in the population is : (2D+H)/2N=(D+1/2H)/N=p
The same can be done for allele a, and designated q. Further, p + q = 1 and hence p = 1 q. If N
= 80, D = 4, and H=24, p=(4 +12)/80 =16/80 =0.2 and hence q=1 0.2 =0.8.
Random mating involving locus (A/a) will yield the genotypes: AA, Aa, and aa, with the
corresponding frequencies of p
2
, 2pq, and q
2
, respectively and p
2
+ 2pq + q
2
= 1.
This mathematical relationship is called the HardyWeinberg equilibrium. Equilibrium between
genes and genotypes is achieved in large populations. The frequency of genotypes in a
population depends on the frequency of genes in the preceding generation, not on the frequency
of the genotypes.
Considering the previous example, the genotypic frequencies for the next generation following
random mating can be calculated as follows:
AA=p2 =0.22 =0.04 Aa =2pq =2(0.2 0.8) =0.32 aa =q2 =0.82 =0.64 Total =1.00
HardyWeinberg equilibrium is hence summarized as: p
2
AA+2pqAa+q
2
aa=1(or 100%)
In a population of 80 plants, about three plants will be AA, 26 will be Aa, and 51 will be aa. The
frequencies of the genes in the next generation : p= (3 +13)/80 =0.2, and q=1 p=0.8 The allele
frequencies have remained unchanged, while the genotypic frequencies have changed from 4, 24,
and 52, to 3, 26, and 51, for AA, Aa, and aa, respectively.
However, in subsequent generations, both the genotype and gene frequencies will remain
unchanged, provided:
Random mating occurs in a very large diploid population.
Allele A and allele a are equally t
There is no differential migration of one allele into or out of the population
The mutation rate of allele A is equal to that of allele a.
The variability does not change from one generation to another in a random mating population.
The maximum frequency of the heterozygote (H) cannot exceed 0.5.
The HardyWeinberg law states that equilibrium is established at any locus after one generation
of random mating.
In order for HardyWeinberg equilibrium to be true, several conditions must be met. However,
some situations provide approximate conditions to satisfy the requirements.
The issue of population size
The HardyWeinberg equilibrium requires a large random mating population. However, in
practice, the law has been found to be approximately true for most of the genes in most cross-
pollinated species, except when non-random mating occur.
The issue of multiple loci
It is possible for alleles at two loci to be in random mating frequencies and yet not in equilibrium
with respect to each other. Further, equilibrium between two loci is not attained after one
generation of random mating but is attained slowly over many generations. Also, the presence of
genetic linkage will further slow down the rate of attainment of equilibrium. If there is no
linkage (c = 0.5), the differential between actual frequency and the equilibrium frequency is
reduced by 50% in each generation, it would take about seven generations to reach approximate
equilibrium. At c = 0.01 and c = 0.001, it would take about 69 and 693 generations, respectively,
to reach equilibrium.
Factors affecting change in gene frequency
Migration: Migration is important in small populations. It entails the entry of individuals
into an existing population from outside is via pollen transfer (gamete migration). The
impact this immigration will have on the recipient population will depend on the
immigration rate and the difference in gene frequency between the immigrants and
natives.
Mutation: Natural mutations are generally rare and would have little impact on gene
frequencies. Recurrent mutation may affect the gene frequency of the population.
Selection: Selection is the most important process for altering population gene
frequencies by plant breeders . Selection decides which individuals will be allowed to
contribute to the next generation. Its effect is to change the mean value of the progeny
population from that of the parental population.
Different types of selection
There are three basic forms of selection
Stabilizing Disruptive Directional
Directional selection being the one of most concern to plant breeders.
These forms of selection operate to varying degrees under both natural and articial selection. In
natural selection, the goal is to increase the tness of the species, whereas in plant breeding
articial selection is usually imposed to direct the population toward a specic goal (not
necessarily the ttest).
Stabilizing selection: Regarding characters that directly affect the tness of a plant (e.g.,
viability, fertility), selection will always be directionally toward the optimal phenotype
for a given habitat. However, for other characters, once optimal phenotype has been
attained, selection will act to perpetuate it as long as the habitat remains stable. Selection
will be for the population mean and against extreme expressions of the phenotype. It is
also called balancing or optimum selection. Eg for owering , stabilizing selection will
favour neither early owering nor late owering. Stabilizing selection promotes additive
variation.
Disruptive selection: Natural habitats generally consist of a number of ecological
niches distinguishable in time (seasonal or long-term cycles), space (microniches), or
function. These diverse ecological conditions favour diverse phenotypic optima in form
and function. Disruptive selection is a mode of selection in which extreme variants have
higher adaptive value than those around the average mean value. It promotes diversity
(polymorphism).
Directional selection: Plant breeders impose directional selection to change existing
populations or varieties (or other genotypes) in a predetermined way. Articial selection
is imposed on the targeted character(s) to achieve maximal or optimal expression. Hence,
directional selection leads to the establishment of dominance and/or genic interaction
(epistasis).
Selection in Mendelian populations
Of the various forces causing disturbance in gene frequencies, selection and non random mating
are of interest to the breeders.Consider a single pair of genes A/a, a random mating population
will consist of AA, Aa, aa genotypes. If AA or aa plants are selected, frequency of selected allele
will be 1 and of other allele will be 0. ( assuming AA or aa can be identified without error). In
the next generation only selected allele will be present (fixed). Here disadvantage in reproduction
i.e, coefficient of selection,s will be 1. s is proportionate reduction in contribution of a particular
genotype compared with a standard genotype, usually the most favoured one. Generally, s has
values less than 1 due to problems in exact identification (dominance, >100% heritability). This
is particularly true for quantitative characters. Thus, selection is expected to change gene
frequencies, rather than fix or eliminate one or other allele
When s is less than 1, rate of change in gene frequency (q ) would depend upon the intensity of
selection (s) and the gene frequency, q. If s is fixed, that is, for a given value of s, q will
depend on the gene frequency, q, and to some extent by the degree of dominance. When a allele
is rare in a population, q for this allele will be small.
As the value of q increases due to selection, q also increases, reaching a maximum at about
q=0.3 in case of dominant allele A, at q=0.5 when there is no dominance, and at q=0.7 in case of
recessive allele a. After this delta q again declines as the favoured allele becomes more frequent
in the population. Thus, selection in a RMP is highly effective in increasing or decreasing the
frequency of alleles, but is unable to fix or eliminate them. However, when combined with
inbreeding, selection is highly efficient in fixation or elimination of alleles.
When selection favours heterozygote, Aa, both alleles retained in population. q and p reach
equilibrium when q=Sa/(Sa+SA), where Sa and SA are selection intensities against aa and AA,
respectively. Equilibrium value of q is quite different from 0.5 although at this frequency,
frequency of heterozygote, favoured class, is maximum. This is because fitness of population as
a whole is greatest at equilibrium point of q, although frequency of heterozygote may not be
maximum. Fitness of a genotype may be defined as its reproduction rate in relation to those of
other genotypes. Gene frequency of q=0.5 does not necessarily produce best average phenotype
for population, although it produces maximum proportion of best phenotypic class.
For quantitative characters, selection intensity is measured as difference between the mean of the
population and that of selected individuals. Selection of extreme phenotypes will increase
frequency of desirable alleles in the population. q for each gene becomes progressively smaller
as the number of genes governing the character increases. With an increase in the frequency of
desirable alleles, the frequency of desirable genotypes will also increase. New genotypes
producing more extreme phenotypes will also appear and mean of the population will change.
q for each gene would be greatest when q=0.5 since additive gene action is assumed. With
higher/lower values of q, q for each gene will become smaller. Thus selection will be unable to
fix/eliminate alleles in case of quantitative traits. Further, variance may decrease to some extent,
but selection will show continuous gain for several generations
Progress under selection for quantitative characters is retarded by
Non additive gene action
Low heritability
Permissible selection intensity
Heritability for quantitative traits <100%.
Consider a population where frequency of A is 0.90 and it is desirable to fix A. If no problem in
identification, it needs 12 generations to change it to 0.95 and another 32 generations to change
to 0.98. If there are problems in identification, s is less than 1, assume it is 0.20. Then it takes 54
generations to reach 0.95 and another 155 to reach 0.98.
Therefore, selection in outcrossing populations is a distressingly feeble means of achieving near
fixation of even highly heritable characters.Permissible selection intensity is dependent upon
reproductive capacity of the species. It is not a serious problem in plants.
Another factor that limits the selection intensity is the consideration of inbreeding. Inbreeding
should be kept to a low level to maintain heterozygosity and to avoid inbreeding depression. The
above considerations are for a single character. Breeder generally has to concentrate on more
than one character. When two genetically uncorrelated characters have to be selected
simultaneously, selection intensity for each of the two characters has to be relaxed.