Presentation

On

MATING SYSTEM
Speaker : Dr. Deepak
 MATING SYSTEM
Mating is a process that determines which males
(selected) are bred to which females (selected). In
other words, it can be defined as a set of rules for
making mating decision.

Primarily mating system can be classified as:

a) Random mating

b) Non-random mating
 Random Mating: (Panmixia or Panmixis)

It is defined as the type of mating in which any

individual of one sex has an equal chance of mating
with any other individual of the opposite sex in the
population.

In random mating all individuals contribute an equal

number of offspring to the next generation, since all
individuals are assumed to be equally viable without
any difference in fertility.

This is the simplest of all mating systems. It is a type

of mating in which any female in a population has an
equal chance to be mated with any male in a
population.
 Non-random mating:

Artificial mating in which the mating is decided or planned

or controlled by the breeder can be considered as non
random mating.
These mating strategies can be either based on animal
performance or on pedigree relationship.
 Non-random mating:
Non random mating

Based on phenotypic resemblance Based on genotypic resemblance

Assortative mating
Inbreeding Outbreeding

Positive Negative
Assortative mating Assortative mating
Based on phenotypic resemblance:
It is also known as assortative mating: Here mating is
based on phenotypic resemblance.

It is the mating of either similar individuals or dissimilar

individuals.

If the mated pairs are phenotypically similar it is called as positive

assortative mating. If it is phenotypically different it is called as
negative assortative mating.
a)Positive assortative mating or Like to like mating:
If animals who are phenotypically alike are allowed to mate among
themselves, it is called as “like to like” mating or phenotypic or positive
assortative mating.

Positive assortative mating tends to create more genetic and phenotypic

variation in the offspring generation than would be found in a comparable
randomly mated population
The consequence of assortative mating with a single locus in terms of
genotypic frequencies among the progeny is to increase the frequencies of
homozygotes and reduce that of heterozygotes.

b) Negative assortative mating or Disassortative mating or Unlike mating:

Under this mating system, animals which are phenotypically unlike are
allowed to mate. It results in loss in homozygosity, but with a gradual
increase in heterozygosity. Negative assortative mating tends to decrease
variation and increase phenotypic uniformity in the population. This is also
called as “Corrective mating”.
II. Based on genetic resemblance:
This system of mating was introduced after the discovery of a
measure of genetic relationship (coefficient of relationship) by
Wright in 1922.

Genetic assortative mating or Inbreeding: Mating of related

animals. In this individuals which are more closely related genetically
are mated. Since the mating is between individuals which are more
closely related, the homozygosity increases and genetic variability
decreases.

The effect of positive assortative mating is similar to those of

inbreeding but to a lesser extent.

Outbreeding: When the relationship between individuals which are

mated together is less close than the average relationship within the
population, the mating system is referred as outbreeding. It
increases heterozygosity and genetic variability which offers more
opportunity for effective selection.
Inbreeding
Inbreeding: When the mates are more closely related than the average members
of the breed, the mating system can be called as inbreeding. The mated
individuals should have one or more common ancestor in their pedigree up to 4-6
generations. This means that inbreeding is the mating of related animals within
4-6 generations. This is because the contribution of ancestral genes to the total
genetic constitution of the descendent of seventh generation is less than one
percent on the basis of the halving nature of inheritance. Thus, the maximum limit
of considering an ancestor/common in the pedigree of an animal is up to 6
generation.
Inbreeding may either be occasional or consistently carried out for several
generations. If it is consistently carried out for several generations (recurrent
inbreeding), it can be classified as
1) Close inbreeding
2) Line breeding
Close inbreeding:
Close inbreeding is a severe or strict form of inbreeding. The
matings are made between sibs or between parents and
progeny. This type of mating is carried out to produce inbred
lines with relatively high degree of homozygosity.

Example: Mating between sibs or between parents and

progeny / offspring i.e. sire x daughter or son x dam. The
most often used method is ‘full sib’ mating. Same effect can
also be achieved by consistently back crossing the progeny to
the younger parent. Sometimes half sib mating is also taken
as close breeding. Half-sib mating is very much slower in
reaching homozygosity but it is less risky.
The close breeding is practiced for some specific purposes, which
are:
•To develop highly inbred lines.
•To discover undesirable recessive genes with the help of parents-
offspring mating.
•To get more uniform progeny.

•However, the close breeding has some disadvantages like

inbreeding depression and intensification of undesirable
characteristics.
 INBREEDING DEPRESSION
Inbreeding is generally detrimental in domestic animals. Increased inbreeding is
accompanied by reduced fertility, slower growth rates, greater susceptibility to
disease, and higher mortality rates. As a result, producers try to avoid mating
related animals.

Inbreeding depression-
The reduction in mean phenotypic performance associated with inbred animals
from the mean phenotypic value before inbreeding in a population
Inbreeding can have dramatic effects on a herd.
These effects are the result of individuals
receiving identical genes from each parent.
If the parents are related, it is more likely that
they have genes that are identical. An individual
receiving identical genes from each parent is said
to be homozygous for that pair of genes.
This would be desirable if the gene the individual
received from each parent leads to superior
performance. However, most animals carry
undesirable genes that usually remain hidden
unless the animal is homozygous.
An inbred individual is more likely to be
homozygous for any gene, so the animal is more
likely to express undesirable genes, and hence,
undesirable traits.
Inbreeding does not create undesirable recessive
genes, but it does tend to bring to light these
unfavorable genetics.
This leads to a decline in average phenotypic
performance called inbreeding depression.
 Line breeding
Line breeding implies a system of mating in which the
relationships of individuals are kept as close as possible
to outstanding animal. In general line breeding is a
milder form of inbreeding.

Line breeding can be an effective tool for perpetuating

the genes from an outstanding ancestor. It should be
used only in herds that are superior and only those bulls
that are outstanding.

D G1 G2 G3
 Line breeding
In general, line breeding is a special form of inbreeding where
unlike inbreeding, relationship is not so close. In this method an
attempt is made to concentrate the inheritance of one ancestor or
line of ancestor in the line bred offspring.

The genetic effects of line breeding are same as those of

inbreeding. But the primary purpose is not to increase the
homozygosity but to retain or concentrate a good proportion of
genes/ traits of a particular outstanding ancestor (Sire/Dam)
among its descendants.

Usually a sire is not mated to his own daughter and matings are
made between half-sibs or grand sire and granddaughter.
Identical by Descent

Identical by state
Alike in state: Genes which are same in function but are
not derived from the same ancestrol chromosome are
called alike in state.

Identical by descent: Genes which are direct replication

of the same gene from the co-ancestor in an earlier
generation. or gene derived from the same location on a
single chromosome of an ancestor

Synonymous of inbreeding coefficient:

Malecot (1948) Called it as Coefficient de parent or

kinship and denoted it by Φxy

Kempthore (1957) Called it as Coefficient of parentage.

 Type of relationship
Direct relationship: This is the relationship between an
ancestor and its any descendent which are related to each
other on direct genes donar-receipient basis.

Collateral relationship: The relationship due to the genetic

contribution of common parent to the descendents is called
collateral relationship

Pedigree: It is document containing the origin of an

animal. It is a systematical record of the ancestors in the
past few generations of the animal.

Common ancestor: Common ancestor contributes its

inheritance to the inbred individual through both its
parents.
Full sibs: Sibs means brother and sisters. Two individuals
having both parents common are called full sibs.

Half sibs: Two individuals having one common parent are called
Half sibs.

First cousins: The progeny of full sibs from single family

are known as either full cousin or cousin or single first cousin.

Double first cousin: The progeny of full sibs from two families
are called as double first cousin. e.g. When two full brother are
married with two full sisters of another family and
produce the progeny.

Inbred:: Any individual will be inbred when its parents are related
either directly or collaterally related.
 Regular system of matings:

The mating of relatives of the same degree of relationship

in
each generation is called the regular system of inbreeding.

All individual in the same generation will have same

inbreeding coefficient because the same mating system is
practiced in all generations.

F increases in a regular and predictable fashion.

Such system includes Selfing, Halfsibs mating, fullsibs

mating, parent offspring mating mating of cousins of various
degree of relationship.
 GENETIC EFFECTS OF INBREEDING
1.The foremost effect of inbreeding is that it increases the
homozygosity at the expense of heterozygosity without
affecting the gene frequency.

2.Hidden recessive genes come to light and get chance to

express themselves. There are many hidden recessive genes in
outbred population. They are hidden by being paired with
dominant genes. The inbreeding brings them to light by
increasing homozygosity.

3.Inbreeding leads to gene fixation. As a consequence of

increased homozygosity the characters are fixed in an inbred
population due to gene fixation. These fixed characters may be
favourable or unfavourable. It requires the culling of animals
in which the unfavourable character is fixed.
4. The inbred homozygous animals particularly for dominant genes are more
prepotent than non inbreds.

5. Inbreeding depression: This is defined as the impairment of performance of

inbred organisms, which show improvement upon crossbreeding. These traits
are denoted as heterotic traits.

a) Effect of inbreeding on population mean: There is reduction in the mean of an

inbred population compared to random mating and it equals to amount
-2pqdF.

b) The amount of change in mean (inbreeding depression) depends on gene

frequencies and it will be maximum when q = p= ½ .
c) There will be no change in mean under inbreeding
when there is no dominance (d=0).

d) Epistasis, which is interaction between genes that are

not alleles may also be responsible for harmful effect
of inbreeding.

e) When there is no dominance epistasis alone does not

cause inbreeding depression.
6. Effect on variance: Due to Inbreeding within line variance
decreases and between line variance increases.

The total genetic variance with no dominance is increased

due to inbreeding.

7. Environmental variance: The undesirable consequences of

inbreeding appears to include a lowering of the
physiological homeostasis of individuals.
 Homeostasis
It denotes the power of an organism to hold the
physiological processes within normal limits inspite
of varying external conditions.

Homozygosity appears to lead to increased

sensitivity of the organism towards environmental
influences and disturbances and thus increased
environmental variance.

Since better homeostasis must be due to better

functioning of endocrine system and/or other
physiological mechanism.
 PHENOTYPIC EFFECTS OF INBREEDING

The inbreeding has adverse effect particularly on fitness traits like reproduction, vigour
(vitality) and growth traits.

1. The inbreeding tends to depress the growth rate in farm animals. This results in
small margin of profit when the live weight decides the profit in meat animals. The
growth rate also affects the production and reproduction of animals.

2. The reproductive efficiency is reduced with inbreeding in farm animals. The

reproductive efficiency is affected adversely in terms of delay in puberty and
testicular development, reduction in gametogenesis (number of ova shed by
females), and increase in embryonic losses. However, the results are variable.
3. Inbreeding results into an increase in death rate and
hence loss in vigour. The inbred animals are more
susceptible to environmental conditions. However,
the results are not consistent.

4. The inbreeding also causes the appearance of genetic

defects which are mostly recessive in inheritance and
remain hidden in out bred animals. The inbreeding
brings them in homozygous state and hence the
abnormalities are expressed and identified.
5. The phenotypic uniformity among inbred animals within an
inbred line is increased for the characters which are
monofactorial. However, it is not true for polygenic traits. The
variations within inbred lines are more due to environmental effect
than to genetic causes.

6. The productive traits show inbreeding depression which means a

decrease in performance records

DISADVANTAGES OF INBREEDING

a) The frequency of undesirable recessive genes is increased

b) It results in inbreeding depression which causes reduction in

growth, reproductive efficiency, vigour in terms of vitality.

c) The inbred animals are more prone (susceptible) to environmental

changes.
APPLICATIONS OF INBREEDING
The inbreeding is a useful tool in animal breeding. Though inbreeding results in
inbreeding depression but there are certain cases when inbreeding is advantageous.

1. Inbreeding is used to test for recessive alleles the sire may have. The inbreeding
unhidden the undesirable recessive genes by bringing them in homozygous state. This
is done by mating a sire to his 20-25 daughters. Thus it helps to eliminate the
undesirable genes.

2. Inbreeding is used to produce distinct families within a breed. The selection between
families for traits of low h2 is more effective.
3. The inbreeding is used to develop inbred lines as a seed
stock which can be crossed according to their combining
ability. The inbreeding with selection had helped in the past
to develop several breeds of livestock viz. long horn cattle,
Shire horse of Robert Bakewell and later on Merino,
Rambouillet breeds of sheep were developed.

4. Inbreeding also helps to know the type of gene action

affecting a trait. The traits governed by additive gene
action do not show inbreeding depression.

5. Inbreeding increases the prepotency which is the ability of

an individual to stamp its characteristics on the progeny.
The prepotency depends on the homozygosity of dominant
genes.
Genetic effects of inbreeding:
1) It makes more pairs of genes in the population homozygous
regardless of the kind of gene action involved (i.e. desirable/
undesirable or favourable / unfavourable). All phenotypic
effects of inbreeding result from this effect of increasing
homozygosity.
2) Inbreeding does not create new genes in the population
3) Inbreeding changes the frequency of the genes in the
population. It brings many recessives to light as it increases
the frequency of both dominant and recessive
homozygotes.
4) The inbreeding reduces the genetic variability within inbred
lines but leads to genetic differentiation between lines.
 CONSEQUENCES OF INBREEDING:
Consequences of inbreeding:
Inbreeding does not increase the number of recessive alleles in a
population but merely brings them to light through increasing the
frequency of homozygotes.
Inbreeding fixes character in an inbred population through
increased homozygosity whether or not the effects are
favourable.
One consequence of the increase in homozygosity caused by
inbreeding is greater prepotency in inbreds. Individuals are said to
be prepotent if the performance of their offspring is especially
like their own and (or) is especially uniform. An inbred individual
is more likely to be prepotent if its homozygous loci contain
chiefly dominant alleles.
Inbreeding accompanied by selection may increase the
phenotypic uniformity among the animals with in the population.
 PHENOTYPIC EFFECTS OF INBREEDING

Effect on growth

Reproduction

Vigour

Hereditary lethals and Abnormalities

Inbreeding depression is generally greatest for traits

associated with natural fitness such as viability and
reproductive ability
 The traits which are commonly affected by inbreeding include:
fertility, number of services per conception, embryonic death, litter
size in pigs, reduction in milk yield and growth rate.

Inbreeding also tends to reduce fitness. The characters like fat

percent in milk and back fat thickness in pigs do not show inbreeding
depression.

In poultry a reduction in egg production of 0.43% for each 1%

increase of inbreeding coefficient was reported. Inbreeding increase
chick mortality.

As inbreeding increases, there is increase in the average number of

AI/conception/cow and decrease in body weight of calves noticed.
Many of the adverse effects of inbreeding in animals are known to be
due to the action of several pairs of recessive genes, each of which has
detrimental effect on the trait.

Sometimes it may either result in the failure to produce required

enzymes or abnormal production of proteins or other compounds.
Inbred animals are usually less able to cope with their environment
when compared to non-inbred animals.

This is evidenced by reduction in fertility, viability and growth rate. This

may be due to the undesirable effects of inbreeding, resulting from
physiological insufficiency and due to a deficiency or lack of balance of
hormone from the endocrine system.
 EFFECT OF INBREEDING ON DIFFERENT KINDS OF GENE
ACTION

Effect of Inbreeding on Different Kinds of Gene Action

When considering the genotypes it is true that inbreeding increases the homozygosity, which affect all
pairs of segregating genes. But the effect of inbreeding on the phenotype varies depending upon the kind of
gene action involved.
Dominance and recessiveness
As we discussed earlier, the reason for the decline in vigour due to inbreeding is the upshot of the
uncovering of detrimental recessive genes through increased homozygosity, which normally remain hidden in
non-inbred populations due to dominant genes. As the degree of homozygosity increases, the average value
of the population move towards the recessive phenotype. But if complete homozygosity were attained there
would be no further decline, as there would be no further uncovering of recessive genes.
Over dominance
In overdominance, the heterozygote genotype is superior to either of the homozygotes. But as a result
of inbreeding there is a decrease in the number of heterozygous and increase in the number of homozygous
individuals in the population which result in deterioration of the trait. But if we couple selection also with
inbreeding, superior animals will get selected, which will favour heterozygotes. So the rate of increase in
homozygosity will be slower in the population.

Epistasis
It may also result in the deterioration of the trait when inbreeding is practiced. Since epistasis involve
at least two different pair of genes, it is very difficult to predict the effect of epistatic gene action under actual
conditions. But theoretically, developing inbred lines, crossing them and then developing a new synthetic
population from the cross will be an option to fix a larger number of favourable combinations of genes with
epistatic effects.
 Additive gene action
The role of additive gene action is prominent, if selection is practiced
along with inbreeding. Selection of superior animals will improve the
population, if additive is the only kind of gene action involved for a trait until
genetic variation is exhuasted.
But once the population reaches this point, no further improvement
can be made. However, economic traits are affected by both additive and
non-additive types of gene action. So it is not easy to develop a superior
inbred line. Therefore it is necessary to fixing superior genes in inbred lines
and then cross them to get combinations of genes that will give hybrid vigor.
WAYS TO MINIMIZE INBREEDING
Ne = Effective population size

ΔF = Rate of inbreeding

Ne = 4 Nm Nf
Nm + Nf

ΔF = 1/2Ne

ΔF = 1/8Nm + 1/8Nf

Nm = No of males

Nf = No of females
 WAYS TO MINIMIZE INBREEDING

1.Mating Plan: The mating should be among the unrelated

animals. The inbreeding is reduced by mating females from one sire
line to males from another sire line.

2.Population size: In India the population size of organized herds

is small. The inbreeding is inevitable in a closed herd of limited size
having about 100 females and 5-10 sires. Therefore, Ne should be
such that ⌂F is not more than 1.0% per generation. It is thus
essentially required to use more number of males.

3.Purchase or exchange of sires: In a single herd, the ⌂F is

increased by use of limited number of males. The way to
reduce ⌂F is to introduce new breeding males in the herd from
outside.
Out breeding systems can be of different types.
 Crossbreeding
 Outcrossing
 Grading up
 Top crossing
 Composite crossing
 Species hybridization or species crossing
 FORMS OF OUT BREEDING
1. Out crossing:
Mating of unrelated pure bred animals of the same
breed is called out crossing.

The progeny produced by out crossing are called out­

cross.

The out crossing system is to use the selected best

purebred sires on the females of the same herd or
different herd but of the same breed.
Advantages of out crossing:

1. This system of selection and out crossing is very

effective for characters governed by additive
effect of genes having high heritability.

2. Out crossing does not allow the fixation of

undesirable genes and hence brings improvement.

3. This system is most widely used for the

production of pure breds
3 It is advantageous to the breeders who feel they
can purchase from other breeder than to produce
in their own herd.

4 Out crossing is practiced when selection limit has

been reached.

5 Out crossing combine with selection is effective to

offset the adverse effect of inbreeding.
 Selection plateau:
Due to continued selection the response to selection
after 10-15 generations starts declining and in some
cases after 20-30 generations of continuous selection
the response is ceased.

This point or level at which there is a failure to respond

is often called as selection plateau and the population is
called plateaued population.
 Forms of out crossing
a) Selective breeding: The out crossing within a
herd by use of selected sires is called as selective
breeding.

b) Top crossing :This mating system is a form of

out crossing and is like grading up system. It is the
mating of females to the last male in the top side of
the pedigree.
 Crossbreeding:
The mating of animals from different established breeds is
called crossbreeding. The progeny produced is called
crossbred.

This system has major role in livestock improvement by

developing new breeds of livestock.

Advantages or Purpose of crossbreeding:

a. To take advantage of heterosis: The two breeds differ

genetically and hence the crossbreds are expected to show
heterosis or hybrid vigour. This make the crossbred more
productive and better than either of the parental breeds.
The crossbreeding has become quite common in pigs and
poultry.
b. To combine good qualities of two or more breeds:

The distinct breeds have different good qualities. One

breed may be superior for one or more traits ,while
other breed(s) may be superior for some other desired
traits.

For example, crossbreeding of exotic and zebu breeds

may combine good qualities of the two breeds in one.
This is known as the complementarity .
c. The crossbreeding is effective for traits that can not
be improved by selection within a breed due to less
genetic variability.

d. New breeds of livestock can be evolved/developed by

crossbreeding.

e. The crossbreeding is also used to study the

behaviour of characters in hereditary transmission.
The crossing of inbred line is particularly more useful
in this respect.
Disadvantages of crossbreeding

1. The breeding merit of crossbred animals may be

reduced because of heterozygosity.

2. Crossbreeding require maintenance of more then two

or more pure breeds in order to produce the
crossbreds which involve high investment.

3. Crossbreds are useless for further breeding purposes

because their offsprings will be of low merits than
themselves.
 Types of crossbreeding
The major forms are the regular crossing and
composite crossing.

1. Regular or systematic crossing

This is mating of the same cross on regular basis to

take the advantages of heterosis and complementarity.

This type of crossing exploits non-additive gene

effects through heterosis and the additive gene effects
through complementarity when two or more characters
complement each other.
 2. Composite crossing to produce synthetic breed

It is done by producing one or few crosses between

two or more populations to produce a single population
having genes from each of the population.

This single population is a mixture of various

crossbred populations and is called as synthetic or
composite. This synthetic population is improved by
selection within it.
 Forms of regular crossing

1. Two pure breed crosses: When two purebreds are

mated together and their crossbred progeny are not used
in breeding programme. This produces offsprings which are
100 % heterozygous showing 100% individual heterosis but
no parental heterosis because the parents are not
crossbreds.

2. Inter se mating: This is the crossing of crossbred

progeny having the same level of inheritance of two breeds
like crossing of F1 with F1. This creates a number of genetic
groups
 3.Back crossing: It is the mating of crossbred animals to
purebred animal of either breed used to produce the
crossbred progeny (F1 ) like F1 x P1 or F1 x P2 where P1 and
P2 are two purebreds used to produce F1 animal.

Advantage of back cross

This cross is made to exploit maternal or paternal heterosis.

As the maternal heterosis is generally of more importance,
the crossbred parents should be the females (F1 females). The
back crossing between crossbred female and purebred males
shows 100% maternal heterosis.
 4. Criss crossing: It is similar to back crossing except that
both the parental breeds (P1 and P2) are used alternately
in each generation.

5. Three breeds cross or triple crossing: In this system of

crossbreeding three breeds are used. The first cross
animal (F1) is mated to an animal of a third breed like
(AB) female x C male.

This results in full utilization of maternal heterosis

because the dam is 100% heterozygous being crossbred as
well as of individual heterosis because the progeny are
also 100% heterozygous.
6. Four breed crosses or double two breed crosses: This
involves the crossing of crossbred females produced by
crossing two breeds (A and B) with crossbred males
produced from crossing another two breeds (C and D).
Thus, mating is between (AB) x (CD). This is used to
produce commercial progeny.

This enables full exploitation of both maternal and

paternal heterosis as well as individual heterosis for the
reason that both parents are crossbred.

7. Rotational crossing: The males of two or three breeds are

used in regular sequence (rotation) in successive
generations on crossbred females of the previous
generation. Thus it is called as the rotational crossing
which may involve two or three breeds.
In crossing: Crossing of inbred lines of the same
breed.

In crossbreeding: Crossing of inbred lines of different

breeds.

Recombination loss: A loss in gene combination value

caused by gradual breaking up of favourable epistatic
blocks of linked loci in advanced generations of
certain hybrids
Grading up

The sires of a pure breed are mated to the females of non-

descript, scrub or native (deshi) type.

The crossbred females are back crossed to the pure bred sires
to produce the progeny with 75% genes from the pure breed.

Thus after six generations of crossing the graded females with

the purebred sires produce the progeny which have 98.4%
genes of the purebreed.

Thus upgrading system of mating changes the genetic

composition without creating new genes but by transmitting
good quality genes of an improved breed.
 Advantages of grading up:

1. Purebreds can be obtained just after few

generations (7th to 8th generation)

2. It is less expensive because it can be started with

little money in comparison to the purchase of an
entire herd of purebreds.

3. This system helps to prove the genetic potentialities

of the sire and adds to its market value.
 Limitations of grading up

1. The most important is that the genotype-environmental

interaction puts a limit to use this system. This is
because the purebred sire which gives good results in
one environment may not give similar results in other
environment.

2. Purebreds are not always better than the graded.

 3. Species hybridization
The crosses between two species are the widest (extreme) possible
type of outbreeding. This is known as species hybridization. The
following are the example of species hybridization:

Species hybrid obtained Species crossed

1. Mule Male ass (Jack) x females horse (mare)
2 Hinny Female ass (Jennet) x Male horse (Stalion)
3. Zebroid Zebra-horse hybrid
4. Asbra Ass x Zebra (in Africa)
5. Pien niu Cattle- Yak cross in Tibet
6. Cattalo American buffalo bull (B. bison) x Domestic
cow (B. taurus)
7. Jatsa (F1 male) Mithun x Cow
8. Jatsamin (F1 female) Mithun x Cow
9. Jechha (F1 males) Mithun x siri cow
Mithun x siri cow
10. Jessam (F1 females)
 Heterosis or Hybrid vigour
•The superiority of outbreds/crossbreds over the average of
their parents is called the heterosis or hybrid vigour.

•The hybrid vigour is the opposite to the phenomenon of

inbreeding depression.

•The outbreds are heterozygotes in which the effects of

undesirable recessive genes are hidden by the effect of
favourable genes (dominant). Therefore, the outbreds are
superior to the average of their parents.
•The heterosis is the measure of the effect of out breeding and is
defined in terms of the difference of outbred animals from the
average of purebred parents. Thus heterosis may be positive or
negative.

•The positive heterosis is often called as hybrid vigour. This is

the phenomenon by which crossbred progenies are better than
the average value of the two purebreds used to produce
crossbred progeny. In some cases, the crossbred progeny exceed
even the better parent.
•The heterosis is caused by heterozygosity created due to crossing
two purebred populations with different gene frequencies.

•The cause of heterosis, like inbreeding depression is the non­

additive gene action (dominance, over dominance and epistasis).

•No heterosis is observed for traits governed by additive gene action.

The traits showing heterosis are called often as heterotic traits.

HF1 = dy2

d is the degree of dominance

y2 is the square of differene in gene frequency of the two population

crossed.
 Estimation of heterosis

The amount of heterosis is estimated by comparing the mean value

of purebred and crossbred animals as follows:

Heterosis = Mean of F1 progeny - Mean of parental breeds

This can also be expressed in terms of percentage as:

% Heterosis = Mean of F1 progeny- mean of parent breeds X 100

Mean of parent breed
 Genetic basis of heterosis
Additive gene action is not responsible for heterosis. It is the
non additive type of gene action which causes heterosis.

In this case the mean of the F1 may even be higher than the
better parent or lower than the inferior parent. When F1
exceed the better parent it is called useful heterosis.

(i)Dominance hypothesis:- The degree of heterosis depends

upon the type of trait and type of mating. The early expressed
traits, like survival and growth rate to weaning are more
influenced. The traits which are more adversely affected by
inbreeding also show greatest degree of heterosis.
The mating of unrelated individuals (mating of inbred
lines, two purebreds) show heterosis in their offspring.
 There are two reasons for this.
a. The animals of one purebred will be homozygous for some loci and the
animals of other group (lines, purebreds) will be homozygous for other loci.
When lines or breeds homozygous for different genes will be crossed they will
produce heterozygous offspring. The favourable dominant genes in the
offspring will mask the unfavourable recessives. Thus the performance of the
hybrid offspring will exceed to that of the average of the parents and sometimes
exceed even to that of better parent.

For example, for polygenic traits, one line or breed may be homozygous
dominat for some pairs and homozygous recessive for another like AA, bb, CC
and DD whereas the other breed may be respectively homozygous recessive and
dominant at other loci like aa, BB, cc, dd. On crossing them the resulting
progeny (F1) will carry dominant alleles at all loci and would be superior to
both parents for that trait.
b. The crosses between two lines or breeds having different
gene frequencies (p and q) produce a lower frequency of
recessive than the average of the two parents. This is because
the frequency of the recessive in the cross will be qq' instead
of ½ (q2 + q'2).

For example if q = 0.2 and q' = 0.6, then qq' = 0.12 and ½
(q2+q'2) = ½ (0.04 + 0.36) = 0.20. This is always true except
for equal gene frequencies.
 (ii) Overdominance theory:- The overdominace is the
interaction between genes that are alleles and it results in the
heterozygous individuals being superior to the best
homozygous parent. The crossbreeding results in superior
animals if overdominance is important for the reason that the
animal produced by crossbreeding has a maximum number
of heterozygous loci.

For example Crow (1952) suggested that the dominance

hypothesis can account for only 5% of the increase in
performance in crosses of lines of corn. Thus the dominance
theory appears to be insufficient to explain heterosis in corn.
However, heterosis observed in corn can be only due to
overdominance at least at few loci affecting corn yield.
(iii) Epistatic effects:- The epistasis is a phenomenon of
interacting genes which are not alleles. The epistasis is the
effect of genes resulting from the new combination of genes
from different loci. The different genes coming together in
the hybrid interact with each other and produce greater
effect than when they are alone in different parents.

Physiological basis of heterosis:

It has been concluded on the basis of some studies that

crossbreds have a more efficient metabolic system.
 General and specific combining ability

This concept of general and specific combining ability was

given by Sprague and Tatum (1942)

Diallel Crossing: It is called as all possible mating between a

number of genotypes (inbred lines, clones or individuals)

General combining ability (g.c.a.): The term is used to

designate the superiority in average performance of a line
(strain or breed) in hybrid combination. The variation in g.c.a.
is due to additive genetic variance (VA and VAA).
 Specific combining ability (SCA)

It is used to designate those cases in which certain

combination do relatively better or worse than would be
expected on the basis of average performance of lines
involved.

The variation in s.c.a. is due to non-additive genetic variance

(dominance, over dominance and epistasis).

It is the deviation of performance of a particular cross from

the sum of the g.c.a. of the two lines.
Methods of selection for general and specific combining ability

1. Recurrent selection (RS) for general combining ability: This

method consist in testing two segregating populations
against a common broad based tester population and then
selecting individuals within the two populations on the
merit of their cross progeny performance.

The selected individuals are mated to members of their own

population to produce the next generation. The process is
repeated through successive cycles. The two selected
populations are then crossed which is expected to result in
the production of superior hybrids.
 Selection for specific combining ability:
When the SCA is caused by dominance:
•The dominant gene action can be utilized by two ways. One by
producing dominant homozygous individuals for all pair of genes which
effect the trait, and secondly by producing heterozygous individuals.

•It is very difficult to distinguish homozygous dominant from

heterozygotes and that the quantitative traits are polygenic which
reduces the probability of getting all dominant genes into one
homozygous strain.

•The second method uses the production of heterozygous individuals

which can be achieved by producing two complementary homozygous
lines. Though the production of two complementary homozygous lines is
problematic but can be achieved by directed process (directional
selection) of selection during inbreeding.

When the SCA is caused by over dominance and epistasis: This type of
gene action can only be exploited by producing heterozygous
individuals.
Recurrent selection for specific combining ability:
This method of selection differ with from RRS and RS for g.c.a.
in that instead of starting with two segregating populations and
selecting both for combining ability with others, one starts with
only one segregating population and select it for combining
ability with constant tester line.

Advantage:
This reduces the amount of effort spent on testing and is expected
to yield more rapid progress at the beginning because the gene
frequencies will be highly differentiated between tester
population and segregating population than both segregating
populations.

Disadvantage:
The ultimate gain is expected to be less because when the tester
population is kept constant the general combining ability of only
one population can be improved.
 Recurrent Selection For S.C.A
Segregating Population A Inbred tester line-B
Segregating Inbred tester line
population

Test cross
progeny

Selection of animals on the Random breeding

basis of test cross progeny
performance

Segregating population Inbred tester line

Test cross
progeny
2. Reciprocal Recurrent selection:
This system of breeding and selection is to improve both General
combining ability and specific combining ability. This method of
selection involves two segregating population, say A and B. A
number of males of A population is crossed with female of B
population and vice versa.

The males and females of each of the two populations are then
selected on the basis of their crossbred progeny performance. The
rest of the parents are discarded, together with all the crossbred
progeny which are only used to test the combining ability of the
parents.

Selected males and females are remated to members of their own

population to produce the next generation of parents, say A1, B1.
The whole cycle is repeated in each generation.
 Reciprocal Recurrent Selection For SCA & GCA
Segregating Population A Segregating Population B
B-males
A-males
B- females
A- females
Test cross progeny

AB BA
progeny Progeny

A- males selected on the basis B- males selected on the basis of

of AB progeny performance BA progeny performance

A-females selected on the basis B-females selected on the basis

of BA progeny performance of AB progeny performance

Progeny of selected A parents Progeny of selected B parents

Test cross
 COLLATERAL RELATIVES - FAMILY SELECTION

The selection criteria to estimate the B.V. of an individual

may be the information (performance records) of its collateral
relatives.

An individual's collateral relatives are those which are not

related with the individual on direct genes donor - recipient
basis but receive common genes from their common ancestor
(s) e.g. full sibs, half sibs, cousins, uncle/aunt, and
nephew/niece.

The more closely related collaterals to the individual are likely to

have more common genes possessed by the individual and can
provide more accurate information about the individual. e.g. full sib
families and half sib families.
 Family selection: The selection criteria is called as family
selection when the individual's own record is included to
estimate the family mean.

Sib selection: The selection criterion is called as the sib

selection when the individual's own record is not included in
estimating the family mean.

Two types: Full sib selection and half sib selection.

 The sib selection is practiced for the following traits for
which the measurements on the individual are not available
or recorded

1. Slaughter traits

2. Sex limited traits

3. Threshold traits like disease resistance

4. Traits with low heritability in species with high

reproductive rate so as many sibs are measured in short
time

5. The full sib (F.S.) selection is more accurate than half sib
(H.S.) selection.
The H.S. selection is favoured for the following reasons

Half sibs are easily available in more numbers than F.S. viz. The
rate of inbreeding is more for F.S. selection than H.S. selection.
The inbreeding counter balances the effect of selection.

The F.S. correlation is more likely to be increased by c-effects

(common environment shared by F.S. ). The intra class
correlation (t) is rh2 for H.S. and rh2+c2 for F.S. where C2 is the
added contribution of maternal or common environmental
effects. This reduces the accuracy of F .S. selection.

Accuracy of sib selection = r√nh2/(1+(n-1)t)

 Family selection

The selection criteria are known as family selection when based on

the performance of the sibs plus the individual's own record. The
family selection like the sib selection is of two types depending on the
type of sibs viz Full sib family selection and Half sib family selection.

Common environment (c - effects)

The environmental effects which are different for different

families but same for all members of one and the same family are
known as common environmental effects denoted as c-effects
(Lerner, 1950). The family members share common environment
during pre and post natal stage.
Factors affecting accuracy of sib and family selection:

1. The accuracy of sib selection and family selection is more

for traits of low heritability. The relative efficiency of
family and sib selection compared to individual selection
decreases with increase in heritability. When h2 is greater
than 0.5 the selection on any number of FS is never as
accurate as individual selection and same is true for HS
selection when h2 is greater than 0.25.

2. The accuracy of sib selection based on one FS is 0.5 h and

based on one HS is 0.25 h. The accuracy of HS and FS
selection as well as family selection is increased with
increase in n but never exceeds more than 0.5 for HS
(√0.25) and 0.71 for FS (√0.50) selection irrespective of n
and h2 of the trait.
3. The type of family viz. HS and FS families in terms of r
between sibs also affects the accuracy of selection. The more
close family members (FS and HS) provide more reliable
information for selection compared to remote families. In
other words, the accuracy of family selection decreases when
r among family members decreases. Thus FS selection is
more efficient than HS selection.

4. The degree of correlation between the phenotypes of sibs (t)

affects the accuracy of selection. The family selection is
better (even better than individual selection) when the
phenotypic correlation (t) is low. When t is
Advantages of family selection:

The family selection can improve the characters of low

heritability in species with high reproductive rates (pig;
poultry) so as to get many sibs in a short time.

The family selection does not allow the generation interval

to increase.

Family selection is a support to individual selection

because it is better to select an individual from a superior
family.
Disadvantages or limitation of family selection

Family selection is costly of space particularly when the breeding

space and testing facilities are limited. The limited facilities reduce
the intensity of selection.

The family selection as a unit of selection results in inbreeding

and thus limits the genetic diversity. This is because only few
families represent the next generation.

The F.S. family selection can only be applied in species with high
reproductive rates to get large family size.
 Within family selection

It is the selection criteria when individuals are selected on the

basis of their performance expressed as deviation from their
family mean. The individuals that exceed their family mean by
the greatest amount are selected

Combined selection

The selection of an individual on the basis of two or more

sources of information is called as the combined selection or
index selection. It is better to select on the basis of an index
combining information from various relatives (dam, sibs and or
progeny) with or without individual's own record.
PROGENY TESTING UNDER FIELD CONDITIONS
A MODEL

CLUSTER OF 40 Test bulls

VILLAGES 30-35
TOTAL BREEDABLE daughters Per
Duration = 1.5
POPULATION =10000 bull
year

Heritability of milk yield = 0.2

Repeatability of milk yield = 0.3
Average Generation Interval from four paths = 8 years

Annual Genetic Gain = 1 -1.5%