Unit 3

Continuous Variation

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 3.1 Introduction

• Continuous variation refers to variation without natural discontinuities

Example: Height, weight, length etc.

• Characters that exhibit such variation are called quantitative characters or metric
characters

• Thus, their study depends on measurement instead of on counting.

• The branch of genetics concerned with metric characters is called quantitative genetics
or biometrical genetics.

• It embraces most of the characters of economic value to plant and animal breeders and
most of the changes concerned in micro-evolution.

• Thus, this branch of genetics has its most important application to practical problems
and also its most direct bearing on evolutionary theory.
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 Question

How does the intrinsically discontinuous variation caused by genetic

segregation is translated into the continuous variation of metric
characters?

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• Quantitative variation is the result of:

1. the simultaneous segregation of many genes affecting the character,

2. the superimposition of truly continuous variation arising from non-

genetic causes.

• Thus, Mendelian genes and metric genes differ in the magnitude of their effects
relative to other sources of variation.

• A gene with an effect large enough to cause a recognizable discontinuity even in

the presence of segregation at other loci and of non-genetic variation can be
Mendelian gene

• Whereas a gene whose effect is not large enough to cause a discontinuity cannot
be studied individually.
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 This distinction is reflected in the terms major gene and minor gene.

 There are, however, all intermediate grades, genes that cannot properly be classed as
major or as minor.

 Furthermore, as a result of pleiotropy, the same gene may be grouped as major with
respect to one character and minor with respect to another character.

• Variation caused by the simultaneous segregation of many genes may be called

polygenic variation, and the minor genes concerned are sometimes referred to as
Q) Brief Major Genes, Polygenes, and QTLs; Structural vs Regulatory genes
polygenes.

3.2 Metric characters

• Refers to any attribute that varies continuously and can be measured

• Example:- anatomical dimensions and proportions, physiological functions of all

sorts, and mental or psychological qualities.
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• The frequency distributions of most metric characters approximate more or
less closely to normal curves.

• Sometimes, however, the scale of measurement must be modified

(transformed) if a distribution approximating to the normal is to be obtained.

• Q) What are the grounds for believing that metric characters really are
controlled by genes at many loci?

• There are two kinds of evidence:

• The first is indirect observations that agree well with what is expected from
the theory of polygenic inheritance.

• The second is direct experimental evidence suggesting that, loci affecting

quantitative traits can be mapped on chromosomes
• Loci identified in this way are called Quantitative Trait Loci, or QTLs
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 3.3 Properties of metric characters

• There are two basic genetic phenomena concerned with metric characters

A. Resemblance between relatives

• Relatives tend to resemble each other, and the degree increases with closeness.

• However, the degree of resemblance varies with the character, some showing
more, some less.

• The degree of resemblance between relatives is one of the aims of quantitative

genetics in predicting the outcome of selective breeding.

B. Inbreeding depression (conversely, hybrid vigor, or heterosis)

• Since most characters of economic value in domestic animals and plants are
aspects of vigor or fertility, inbreeding is generally deleterious.

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 The reduced vigor and fertility of inbred lines is restored on crossing, and in
certain circumstances, hybrid vigor could be used for improvement.

Values and Means in Metric Characters

• A quantitative differences exhibited in a metric character is called value

• The value observed when the character is measured on an individual is the

phenotypic value of that individual.

• The genetic properties of the populations in the phenotypic value could be

partitioned into component parts:

♣ Genotype (assemblage of genes ) and

♣ Environment (all the non-genetic circumstances that influence the

phenotypic value).
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 Thus, genotype and environment are by definition the only determinants of
phenotypic value.

• Symbolically, it is given by:

• If the mean environmental deviation in the population as a whole is taken to be

zero, then the mean phenotypic value is equal to the mean genotypic value.

• The term population mean then refers equally to phenotypic or to genotypic

values.

• When dealing with successive generations we shall assume for simplicity that the

environment remains constant from generation to generation, so that the

population mean is constant in the absence of genetic change.

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 Population mean

• How does the gene frequencies influence the mean of the character in the population

as a whole?

• Suppose the gene frequencies of A1 and A2 be p and q respectively, the three

genotypes would be A1A1, A1A2, and A2A2.

• The mean value in the whole population is obtained by multiplying the value of each

genotype by its frequency and summing over the three genotypes.

• i.e

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 Assumptions
• d and a are arbitrarily assigned values

• A1 is assumed to be the allele that increases the value.

• The origin, or point of zero value, is assumed to be the mid-way between the values
of the two homozygotes (A1A1 and A2A2).

• The value d of the heterozygote depends on the degree of dominance. Thus,

• If there is no dominance, d = 0;

• If A1 is dominant over A2, d is positive, and

• If A2 is dominant over A1, d is negative.

• If dominance is complete, d is equal to +a or -a,

• If there is overdominance, d is greater than +a or less than -a.

• The degree of dominance may be expressed as d/a.

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• On summary, the population mean would be given by;

• This is both the mean genotypic value and the mean phenotypic value of the population
with respect to the character.

• The contribution of any locus to the population mean thus has two terms:

Ω a(p — q) attributable to homozygotes, and

Ω 2dpq attributable to heterozygotes.

 If there is no dominance (d = 0), the second term is zero, and the mean is proportional to
the gene frequency: M = a(1— 2q).

 If there is complete dominance (d = a) and the mean is proportional to the square of the
gene frequency: M = a(1 — 2q2).
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How genes at different loci combine to produce a joint effect on the
character?

• To find the joint effect of genes at several loci on the mean, one needs to
consider:

Combination by addition (additive gene effect):

• It implies that the value of a genotype with respect to several loci is the sum of
the values attributable to the separate loci.
For example;

If the genotypic value of A1A1 is aA and that of B1B1 is aB, then the genotypic
value of A1A1B1B1 is aA + aB

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•Thus, the population mean resulting from the joint effects of several loci is the sum of the
contributions of each of the separate loci, and given by:

Average effect
•The transmission of value from parent to offspring could be done not by means of genotypic

values alone

•This is because parents pass on their genes and not their genotypes

•That is genotypes are created afresh in each generation and thus, a new measure of value

involving genes and not genotypes is needed

•Such new measure is called a ‘breeding value’ (a value associated with the genes carried by

the individual and transmitted to its offspring).

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• Thus, average effects refer to the new value associated with genes as distinct

from genotypes.

• They depend on the genotypic values, a and d as previously defined, and also on
the gene frequencies.

• They are therefore properties of populations as well as of the genes concerned.

• The concept is tough but fundamental to understanding the inheritance of

quantitative characters.

• In random breeding populations, the average effect of a particular gene (allele)

could be defined as the mean deviation from the population mean of individuals
which received that gene from one parent

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• Example: If a number of gametes all carrying A1 unite at random with gametes from
the population; then the mean of the genotypes so produced deviates from the
population mean by an amount which is the average effect of the A1 gene.

• See table below;

• To elaborate further, consider a locus with two alleles, A1 and A2, at frequencies p
and q respectively, and take first the average effect of the gene A1 for which we shall
use the symbol ά1.
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• If gametes carrying A1 unite at random with gametes from the population, the
frequencies of the genotypes produced will be p of A1A1 and q of A1A2.

• The genotypic value of A1A1 is +a and that of A1A2 is d, and the mean of these,
taking account of the proportions in which they occur, is pa + qd.

• The difference between this mean value and the population mean is the average
effect of the gene A1. Taking the value of the population mean from the table; we get

• Similarly, the average effect of the gene A2 is

• The reason why average effects depend on gene frequencies can be seen in the words
‘taken at random’ in the definition, because the content of a random sample
depends on the gene frequency in the population.
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 Breeding value

• Refers to the value of an individual judged by the mean value of its progeny.

• Breeding value, unlike average effect, can therefore be measured.

• If an individual is mated to a number of individuals taken at random from the

population, then its breeding value is twice the mean deviation of the progeny
from the population mean.

• The deviation has to be doubled because the parent in question provides only half
the genes in the progeny, the other half coming at random from the population.

• Breeding values can be expressed in absolute units, but are usually more
conveniently expressed in the form of deviations from the population mean.

• In terms of average effects, the breeding value of an individual is equal to the sum
of the average effects of the genes it carries
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 The summation is being made over the pair of alleles at each locus and over all
loci. Thus, for a single locus with two alleles, the breeding values of the genotypes
are as follows:

• In a population with Hardy-Weinberg equilibrium, the mean breeding value must

be zero; or

• If breeding values are expressed in absolute units, the mean breeding value must
be equal to the mean genotypic value and to the mean phenotypic value.

• Multiplying the breeding value by the frequency of each genotype and summing
gives the mean breeding value (expressed as a deviation from the population
mean) as Selale University 19
• The breeding value is sometimes referred to as the ‘additive genotype’, and
variation in breeding value ascribed to the ‘additive effects’ of genes.

Dominance deviation

• The breeding value is separated as a component part of the genotypic value of an

individual.

• Thus, when a single locus is under consideration, the difference between the
genotypic value G and the breeding value A of a particular genotype is known as
the dominance deviation D, so that G=A+D

• The dominance deviation arises from the property of dominance among the alleles
at a locus, since in the absence of dominance, breeding values and genotypic
values coincide. Selale University 20
 Interaction deviation

• When only a single locus is under consideration, the genotypic value is made up
of the breeding value and the dominance deviation only.

• But when the genotype refers to more than one locus, the genotypic value may
contain an additional deviation due to non-additive combination.

• Let GA be the genotypic value of an individual attributable to one locus, GB that

attributable to a second locus, and G the aggregate genotypic value attributable to
both loci together. Then

where IAB is the deviation from additive combination of these genotypic values.

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• If I is not zero for any combination of genes at different loci, those genes are said
to ‘interact’ or to exhibit ‘epistasis’.

• The deviation is called the interaction deviation or epistatic deviation.

• If the interaction deviation is zero the genes concerned are said to ‘act additively’
between loci.

• Thus, ‘additive action’ may mean either genes at one locus (absence of
dominance), or genes at different loci (absence of epistasis).

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• For interactions involving more than two loci, the interaction effect is given by:

G=A+D+I

where A is the sum of the breeding values attributable to the separate loci, and D

is the sum of the dominance deviations

Variance

• The genetics of a metric character centers around the study of its variation, since most
primary genetic questions are formulated in terms of variation.

• The basic idea in the study of variation is its partitioning into components attributable
to different causes.

• Thus, the relative magnitude of the components determines the genetic properties of
the population, in particular the degree of resemblance between relatives.
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 Components of variance

• The amount of variation is measured and expressed as the variance (deviations

from the population mean).

• The components include:

Genotypic variance

The variance of genotypic values

Environmental variance

The variance of environmental deviations.

Total variance

The phenotypic variance, or the variance of phenotypic values, and is the sum of the
separate components.
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•There are also other components. See table below

•The total variance is given by:

•The partitioning of the variance into its components allows us to estimate the

relative importance of the various determinants of the phenotype,

• It particularly help us to detect the role of heredity versus environment, or

nature and nurture.

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 Components as proportions of the total

• The partitioning of the variance into its components allows us to estimate the
relative importance of the various determinants of the phenotype,

• The relative importance of heredity in determining phenotypic values is called the

heritability of the character.

• There are, however, two distinctly different meanings of ‘heredity’ and

heritability, depending on whether they refer to genotypic values or to breeding
values.

• A character can be ‘hereditary’ if:

♣ It is determined by the genotype or

♣ It is being transmitted from parents to offspring

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• The extent to which it is hereditary in the two senses may not be the same.

• The ratio VG/VP expresses the extent to which individuals’ phenotypes are
determined by the genotypes.

• This determination is caned the heritability in the broad sense or the degree of
genetic determination.

• The ratio VA/Vp expresses the extent to which phenotypes are determined by the
genes transmitted from the parents.

• Such determination is called the heritability in the narrow sense, or simply the
heritability.

• VA/Vp determines the degree of resemblance between relatives and is therefore of

the greatest importance in breeding programs.

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 Estimation of the degree of genetic determination, VG/VP

• Neither the genotypic nor the environmental components of variance can be

estimated directly from observations on a single population

• But in certain circumstances, they can be estimated in experimental populations.

• Environmental variance cannot be removed because it includes by definition all

non-genetic variance, and much of this is beyond experimental control.

• Elimination of genotypic variance can, however, be achieved experimentally.

Genetic components of variance

• The partition into genotypic and environmental variance does led to the
understanding of the genetic properties of a population (cause of resemblance
between relatives).

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Thus, the genotypic variance must be further divided into breeding value,
dominance deviation, and interaction deviation.

• The additive variance (variance of breeding values) is the important component

since it is the chief cause of resemblance between relatives (already covered)

• Therefore, it is the chief determinant of the observable genetic properties of the

population and of the response of the population to selection.

• Moreover, it is the only component that can be readily estimated from

observations made on the population.

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• In practice, therefore, the important partition is into additive genetic variance
versus all the rest, or non-additive genetic and environmental variance.

• This partitioning yields the ratio VA/Vp, which is the heritability of the character.

Correlation and interaction between genotype and environment

Correlation

• Correlation between genotype and environment is seldom important and can

usually be neglected in experimental populations where randomization of
environment is one of the chief objects of experimental design.

• There are some situations, however, in which the correlation exists.

• Since genotypic and phenotypic values could be correlated, there is also a

correlation between genotypic value and environmental deviation.
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• When a correlation is present the phenotypic variance is increased by twice the
covariance of genotypic values and environmental deviations

• Thus, the equation becomes:

Interaction between genotypes and environments

• When interaction between genotypes and environments is present, the phenotypic

value of an individual is amended into: P = G + E + IGE .

• Thus, the source variation becomes Vp = VG + VE +2covGE +VGE

• Genotype-environment interaction becomes very important if individuals of a

particular population are to be reared under different conditions.

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 Environmental variance

• Environmental variance is a source of error that reduces precision in genetic

studies

• Good examples are: Nutritional and climatic factors, maternal effects, error of
measurements,

• Thus, it should be reduced by careful management or proper design of the

experiments.

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 Unit 4

Heritability

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 Important components of heritability includes:

• Phenotypic variance, σ2P

• Genotypic variance, σ2G

• Additive

• Non-additive

• Environmental variance, σ2E

• Heritable fraction of the total variance

• Broad-sense, σ2G/ σ2P

• Narrow-sense, σ2A/ σ2P

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 4.1 Heritability in the broad sense (H2):

 Is the proportion of the phenotypic variance that is due to all genetic effects
( additive, dominance and epistasis):

 It measures the strength of the relationship between the phenotypic values of the
individuals and their genotypic values.

4.2 Heritability in the narrow sense (h2):

 Is the proportion of the phenotypic variance that is due to additive genetic

effects only

 Thus, it is give as:

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It measures two things:

1.The degree to which the offspring resemble their parents in the phenotype for a
trait.

2.The strength of the relationship between the phenotypic values and the additive
genetic effects (the relationship between P and A).

Important Reminder:

1. Heritability is a measure on a population of individuals in a given environment

for a given character. It is NOT measured on one individual.

2. Heritability can be estimated for each quantitative trait.

3. It varies from population to another and from environment to another for the
same traits.
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 Example:

 If you are given a body weight data taken on Selale Horse population recorded at
their 5 years age : VA= 42, VD=9, VI=3, VE=55; where V represents variance

Calculate

a) heritability in the broad sense and

b) heritability in the narrow sense for this trait.

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• Heritability is always positive ranging from 0 to 1.0.

• It could be expressed as:

Low when h2< 0.20 or less than 20%:

• Example: reproductive traits like litter size, and conception rate, longevity or
productive live ( about 0.10), weaning weight in swine ( about 0.10)

Moderate if h2 is between 0.2 to 0.4 or between 20% and 40 %:

 Example: Milk yield, fat yield and protein yield (0.25-0.35), Birth weight in
sheep, weight in sheep

High if h2> 0.4 or greater than 40%

Example: Carcass traits and traits related to skeletal dimensions like mature body
weight, Fat and protein% in milk
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•Heritability expresses the proportion of the total variance that is attributable to differences of
breeding values
• It determines the degree of resemblance between relatives.
•In estimating heritability, only the phenotypic values of individuals can be directly measured
•But, in practical sense, it is the breeding value that determines the fate of subsequent
generations
•Thus, heritability could be defined as the ratio of additive genetic variance to phenotypic
variance:

•Regarding heritability as the regression of breeding value on phenotypic value, an individual’s

estimated breeding value is the product of its phenotypic value and the heritability

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Thus,

• Heritability expresses the reliability of phenotypic value as a guide to the breeding

value

• Or, it expresses the degree of correspondence between phenotypic value and breeding
value.

• The value of the heritability depends on the magnitude of all the components of
variance, a change in any one of these will affect it.

• With regards to population size, small populations maintained long enough are
expected to show lower heritability than large populations.

• The environmental variance is dependent on the conditions of culture or management:

• Thus, more variable conditions reduce the heritability; more uniform conditions
increase it.
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 4.3 Estimating Heritability from ANOVA

• In practical sense, heritability is estimated either by using variance components or

ANOVA.

• In using variance components, the genetic components of variance in families or lines

containing multiple individuals related in some systematic way can be developed.

• The families can be evaluated in replicated trials either in a single or multiple

locations.

• This permits partitioning of the phenotypic variance into a component due to

genotype, genotype-by-environment interaction and error effects.

• In estimating heritability from ANOVA, observed mean squares are expected to be

equated to their expectations and the variance components are estimated algebraically
as a function of mean squares.
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 h2B = Genotypic variance/phenotypic variance
=

= +
=
Over-locations (from ANOVA):

=
=

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from Parent-offspring:
h2B =

From realized h2 h2r =

d e r m a ting
l b e cov e r e d un
This w il
h2 from NC-designs:
design
h2 N =

σ2A = 4σ2m
= 4( )

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 4.4. How to maximize heritability?

Heritability = proportion of the phenotype which is due to the genotype

It is then given by G2/(G2+E2/r)

Thus, to maximize its estimate, one needs to:

• Increase genetic variance or (G2 )

• Reduce error variance (E2)

• Increase number of replications (r)

• Improved statistical design and analysis techniques

• Choice of field and layout of trials

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 4.5 Applications of Heritability Estimates

• It enables to estimate the expected gains from selection based on alternate strategies.

• To predict gain from selection experiments under different experimental designs.

• To design cost effective breeding strategies.

Basis of Response to Selection

• Heritability could be defined as a fraction of the selection deferential expected to be

gained when selection is practiced on a defined selection units

• In order to apply the definition, an understanding of the response to selection is

important.

• Thus, response of selection could be expressed as a change in the mean progeny

phenotypic values due to change in the mean value of selection units.

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• The selection deferential is the difference between the mean of the selection units
and the overall mean of the initial population.

• Thus, the expected response in any variable, y, due to change in a related variable, x,
can be expressed as

Δy = b(Δx)
where b is the coefficient of regression of y on x,

Δy = the change in y,

Δx = the change in x.

Look at the next slide for more clarification

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 Thus, realized heritability could be defined as the relation between the observed
response to selection and the selection differential.

 It is given by

Reference Populations

• Heritability estimates must refer to a defined population of genotypes.

• Reference populations are generally assumed to be random mating populations

in Hardy-Weinberg equilibrium

• Defining the reference populations of genotypes and environments, could help

in defining the statistical model that will be used to estimate heritability.

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 Unit 5

Selection

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• This chapter considers the changes brought about by the action of a breeder or
experimenter.

• There are two ways in which the action of the breeder can change the genetic
properties of the population:

♣ The choice of individuals to be used as parents, which constitutes selection,

♣ Controlling the way in which the parents are mated, which embraces
inbreeding and crossbreeding.

• Selection is among the ways of improvements in animals and plants.

• Selection means breeding from the ‘best’ individuals, whatever ‘best’ may be.

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• The simplest form of selection is to choose individuals on the basis of their own
phenotypic values.

• Selection could be:

• Artificial selection: is resulted from the action of the breeder in the choice of parents

• Natural selection: operates through differences of fertility among the parent

individuals, or of viability among their progeny

• It acts on the natural differences of viability among the individuals of the offspring
generation

• Both could bring change in genes frequencies

• Though natural differences of fertility and viability are always present, they are not
necessarily always connected with the genes concerned with the metric character.

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 Response to selection (R)

• An important change produced by selection is the change in population mean.

• R is thus the difference in mean of phenotypic values between the offspring of

the selected parents and the whole of the parental generation before selection.

• The measure of the selection applied is the average superiority of the selected

parents, which is called the selection differential, and symbolized by S.

• It is the mean phenotypic value of the individuals selected as parents

• It is expressed as a deviation from the population mean (from the mean

phenotypic value of all the individuals in the parental generation before selection).

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• The connection between the response and selection differential is given by:
R = h2S

• Deviation of the progeny from the population mean is, by definition, the breeding value
of the parents

• Thus, the response is equivalent to the breeding value of the parents.

Selection differential and intensity of selection

• The selection differential can be predicted in advance provided that two conditions hold:-

♣ The phenotypic values of the character being selected are normally distributed

♣ Selection is by truncation.

• In truncation selection, individuals are chosen strictly in order of merit as judged by their
phenotypic values

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• The selection differential, S and the intensity of selection, i refer to the mean
superiority of all the parents used.

• Males and females may differ in the amount of selection that can be applied to
them.

• Some characters can be measured only on one sex, so that no selection can be
applied to the other sex.

• Example, if the selection applied to males and females differs, thus, the values of
S or i to be used are the unweighted means for the two sexes.
• Hence, S = ½(Sm + Sf)
i = ½(im + if)

• Thus, if only females, for example, can be selected, S = 1/2 Sf and the expected
response is R = 1/2 h2Sf

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• Response could be improved (heritability can be increased) through;

A. reducing the environmental variation

B. Careful experimental design and measurements

C. Conducting experiments at multiple sites

D. Slightly inviting assortative mating

Realized heritability

• The response to selection can be used as a means of estimating the heritability in the base
population

• Thus, the expected value of the ratio R/S is the heritability and on rearrangement gives

h2 = R/S

• The heritability estimated in this way is called the realized heritability

• It is primarily a description of the response and may not provide a valid estimate of the
heritability in the base population Selale University 55
 Change of gene frequency under artificial selection

• Change in population mean resulting from selection is the result of changes in

gene frequencies at the loci that influence the character selected.

• But since the effects of the loci cannot be individually identified, the changes of
gene frequency cannot be followed in practice.

• It is possible, however, to deduce an approximate expression connecting the

intensity of selection, i, with the coefficient of selection, s, acting on individual
loci.

• The coefficient of selection is deduced by finding the relative fitness of the

genotypes, i.e., the relative proportions that survive, through being selected.

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 The effects of experiment result on Selection

• There are many consequences of selection that can be discovered only by

experiment.

• The experiment results could cast either short term or long-term effects

A. Short-term results

Repeatability of response

• The variability of the responses from one generation to the next is caused by:

♣ Random drift because of restricted number of parents

♣ Sampling error in estimating the generation mean

♣ Variation of selection differentials

♣ Environmental factors
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• The changes due to drift are cumulative, any change in one generation being
carried on as the starting point for the change in the next generation.
Sampling variance

• The important conclusion is that the standard error of the regression seriously
underestimates the standard error of the realized heritability

• There are then two sources of error in the estimation of the response: variance
due to. random drift. Ϭ2d, and variance due to measurement error Ϭ2e .

• The first depends on the effective number of parents, Ne and the second on the
number of individuals measured, M, from which the mean is estimated.
• The drift variance of a line bred without selection is approximately equal to 2FVA

• The inbreeding coefficient F is approximately equal to t∆F, where t is the

number of generations and ∆F is the rate of inbreeding, which is equal to 1/2 Ne
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• So Ϭ2d = t VA/ Ne.

Asymmetry of response

• Asymmetry of response has important practical consequences because:

1. The prediction of a response is made from the heritability estimated in the base
population.

2. Prediction of the mean of the responses is carried out in two directions, and if there
is asymmetry the response in one direction will fall short of expectation.

• Thus if a character of economic importance is selected in one direction only, the

response may disappoint the breeder by being less than was expected.

• The main causes of asymmetrical responses are as follows:

1. Random drift: can bring asymmetry of response if there is only one selection line
in each direction

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 2. Selection differential: The selection differential may differ between the upward
and downward selected lines, because of

(i) Natural selection that may aid artificial selection in one direction or hinder it in
the other,

(ii) The fertility may change so that a higher intensity of selection is achieved in one
direction than in the other,

(iii) The variance may change as a result of the change of mean: the selection
differential will increase as the variance increases and vice versa.

3. Inbreeding depression: most experiments on selection are made with

populations not very large in size

• Thus, there is usually an appreciable amount of inbreeding during the progress of

the selection.
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• If the character selected is one subject to inbreeding depression, there will be a tendency
for the mean to decline through inbreeding.

• This will reduce the rate of response in the upward direction and increase it in the
downward direction, thus giving rise to asymmetry.

4. Maternal effects: an asymmetry of response associated with the maternal component

of the character.

5. Genetic asymmetry: Both additive genetic variance and the heritability depend on
gene frequencies.

• Additive genes contribute maximally to the heritability when the gene frequency is 0.5,
and recessive genes when the recessive allele has a frequency of 0.75.

• These will be called the ‘symmetrical’ gene frequencies.

Asymmetry will result if the initial population is not at the point, in respect of gene
frequencies, where the additive variance is maximal
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 6. Genes with large effects: Asymmetry of response that appears immediately in
the first generation.

7. Scalar asymmetry: The genetic and environmental variation may be skewed to

different degrees or in opposite directions.

• The genetic variation will then make up a larger proportion of the total at one end
of the distribution than at the other.

8. Indirect selection: Sometimes the criterion of selection is not quite the same
as the character measured for assessing the response.

• Then, if the measured character is not linearly related to the selection criterion,
asymmetry of response may result.

B. Long-term results

• The outcome of selection over a long period is unpredictable because of;

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1. The outcome depend on the properties of the individual gene contributing to the
response which cannot be determined by observation at the outset

2. Mutation produces new variation whose nature could not be predicted.

Selection using information from relatives

• An individual's own phenotypic value is not the only source of information about its
breeding value

• Thus, additional information is provided by the phenotypic values of relatives,

particularly by those of full or half sibs.

• The use of information from relatives is of great importance in the application of

selection to animal breeding because;

1. the characters to be selected may bear low heritability, and thus the mean value of a
number of relatives often provides a more reliable guide to breeding value than the
individual's own phenotypic value. Selale University 63
2. when the outcome of selection is a matter of economic gain, even quite a small
improvement of the response will repay the extra effort of applying the best technique.

• If the family structure of the population is taken into account one could compute the
mean phenotypic value of each family called the family mean

Criteria for selection

• The phenotypic value of an individual, P, measured as a deviation from the

population mean, is the sum of two parts:

P= Pf + Pw

Where Pf is the deviation of its family mean from the population mean

Pw is the deviation of the individual from the family mean (the within-

family deviation)

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• The procedure of selection, then, varies according to the attention paid, or the weight
given, to these two parts.

• There are three simple procedures that can be followed:

• First, we may select on the basis of individual values only, giving equal weights to
the two components Pf and Pw.

• This is called individual selection.

• Second, we may select on the basis of the family mean Pf only, giving zero weight to
the within-family deviation Pw.

• This is known as family selection.

• Third, we may select on the basis of the within-family deviation Pw alone, giving
zero weight to the family mean Pf

• This is known as within-family selection.

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• Instead of one or other of these three simple procedures, one may take account of
both components, Pf and Pw, but give them different weights

• This is known as selection by optimum combination or combined selection more

generally, index selection.

• This method represents the general solution for obtaining the maximum rate of
response
Simple methods

• The features of the above mentioned simpler methods is presented below

Individual selection: Individuals are selected solely in accordance with their

own phenotypic values.

• This method is usually the simplest to operate and in many circumstances it yields
the most rapid response.
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• Mass selection is a term often used for individual selection, especially when the selected
individuals are put together in mass for mating.

Family selection: Whole families are selected or rejected as units, according to

the mean phenotypic value of the family.

• Individual values are thus not acted on except in so far as they determine the family mean.

• In other words, the within-family deviations are given zero weight.

• The families may be of full sibs or half sibs, families of more remote relationship being of
little practical significance.

Within-family selection:

• The criterion of selection is the deviation of each individual from the mean value of the
family to which it belongs

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• Those that exceed their family mean by the greatest amount are regarded as the most
desirable.

• It is the reverse of family selection the family means being given zero weight.

Prediction of response

• Deducing the response expected from each selection could help in evaluating the
relative importance of the methods.

• Thus, the expected response is given by R = iϬph2 where i is the intensity of selection,

Ϭp is the standard deviation and h2 the heritability of the phenotypic values of individuals

• In family selection , Rf = iϬfhf2 where i is the intensity of selection, Ϭf is the observed

standard deviation of family means, and hf2 is the heritability of family means

• For within-family selection, Rw= iϬwhw2 where Ϭw is the standard deviation, and hw2 the
heritability of within-family deviations.
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How to detect heritability?

• Heritability is simply the proportion of the phenotypic variance that is made up

of additive genetic variance.

• Thus, to compute heritability, partitioning of the variance between and within

families of large size is essential.

• Accordingly,

• This partitioning of both the additive and the phenotypic variance leads to the
heritability's of family means and of within-family deviations, since these
heritability's are simply the ratios of the additive variance to the phenotypic
variance. Selale University 69
• Thus, heritability of family means is rVA/tVp , or (r/t)h2 since VA/Vp is the
heritability of individual values, h2.

• The values of r for different relationships could be ½ for full sibs, ¾ for half sibs.

• The intraclass correlation t between members of families is the between-group

component divided by the total variance: t = Ϭ2B/Ϭ2T

• The total variance, written here as Ϭ2T is the phenotypic variance and written as
Vp

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