Genetic improvement (Systems approach to genetic improvement)

There are many ways of improving the productivity of livestock. In livestock improvement these many
ways include improved feeding and management (physical environment), improved health care,
physiological or pharmacological interventions and animal breeding (genetic improvement). The above
methods of improving the productivity of livestock are under the control of the farmer except for genetic
improvement which is under the control of the animal breeders. The methods of genetic improvement
are limited and genetic improvement may be effected or achieved through selection, crossbreeding,
inbreeding, breed substitution and any combination of all these.

Selection of breeding stock:

Selection is the process of choosing certain individuals (plants and livestock) in the population as future
parents of the next generation in preference to others. Selection is carried out in order to improve the
average performance of plants and livestock for the specific trait or a combination of the required traits.
In practice, selection involves the culling of plants and livestock showing bad traits from the crop or herd.
Selection changes the gene frequency in the population, i.e. the gene frequency between the parents and
the off-spring generations; and therefore changes the performance in the population.

Various selection methods are used to meet the requirements of the breeder after setting out the selection
objectives and the measurements, i.e. the selection criterion.

1. Performance Testing or Mass / Individual Selection:

This is the selection method based on the phenotypic performance of animals as their genetic
constitution is not known. It is used for the selection of traits with high heritabilities that can easily
be measured in potential parents and hence not used for selection of traits with low heritabilities.
The advantage is that it is very easy to carry out and speeds up genetic progress. However, many
traits such as milk yield and reproductive efficiency can only be measured in cows, i.e. sex limited
traits. Therefore, for sex limited traits bulls cannot be selected according to their phenotypic
performance.

Performance testing involves measuring the performance of the potential parent individuals and
selecting as parents those with the records that meet the selection criterion. In the process of
selection, a single record of each animal's performance test is used and an estimate of its’ breeding
value (BV) or genetic merit is calculated.

BV = h2 x (Individual performance - Average of contemporaries)

or
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BV = h x (Individual deviation)

The breeding value (genetic merit) is the potential for a given animal to improve the performance of
a given population of animals. For example, if a given cow has a potential to produce 6,000 litres of
milk per lactation and another 5,000 litres per lactation, and if this difference is due to genetic
makeup then the breeding value/genetic value of the cow is 1,000 litres of milk per lactation, i.e. the
cow can improve the lactation performance of the herd by 1,000 litres of milk per lactation and
depending on the h2 of the milk yield.

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 Performance testing is mainly used to test potential sires under similar environmental conditions at
Performance Testing Stations. The method is use to reduce a large number of potential sires to a
reasonable number for further Progeny Testing.

2. Progeny Testing

Progeny testing is the selection method in which the selection decision of the potential parents is
based on the performance of an animal's progeny, i.e. the potential parents are selected on the basis
of the performance of their off-springs. Progeny testing is used to assess quantitative traits which can
be expressed in one sex only, such as estimating the genes for milk production possessed by a bull;
traits which can be measured late in life, such as carcase characteristics which can be measured after
slaughter; and traits which have low heritability (and individual selection will be highly inaccurate).

Progeny Testing is usually carried out on male animals because one male can be mated to many
females, i.e. males are able to express their potential widely by serving a number of females. Progeny
Testing is usually carried out by comparing the performance of the progeny of a particular sire with
the performance of the progeny of other sires reared under similar environmental conditions, i.e.
contemporary comparison. For example, to select bulls of high milk potential the performance of
their daughter progeny is evaluated because the bulls cannot be milked. To achieve this, the bulls are
introduced to different herds at random in order to reduce the dam effect, i.e. by assigning bulls to
the females.

The disadvantage of progeny testing bulls/sires is that it takes long to assess their breeding value.
Progeny testing takes 7 years in cattle and by then the bull would be 8 years old. However, the
technique of deep freezing semen at artificial insemination centres allow for accurate progeny testing
of the bulls to be carried out. The technique also increases the working life of valuable sires as semen
can be deep frozen and used long after the bulls’ death.

Contemporary Comparisons

In the process of Progeny Testing, semen from bulls is used to inseminate a large number of cows
(500) and their 200 – 250 heifers are then reared in different environments. After calving their
lactation milk yield (305 days) is recorded. The milk yields of a particular bull's daughters are then
averaged and compared with the average yields of other bulls (contemporary comparisons method).
For example, if 5 daughters of Bull A yield 3, 800 kg milk and daughters of other bulls yield 3, 250 kg
milk. Therefore, Bull A has a contemporary comparison of +550.

In order to find the true Relative Breeding Value (RBV), the average performance of each herd in the
country must be worked out. The average is the contemporary national breed average. The national
breed average is taken as 100% and a bull must have a RBV of 100+ to improve the performance or
production of the breed in order to be selected. For example: Bull A may have RBV = 123, Bull B may
have RBV = 113 and Bull C may have RBV = 85 (culled). Hence, bulls A and B may be selected as future
parents.

Improved Contemporary Comparisons (ICC):

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 It is a method of progeny testing whereby the ICC is expressed as a comparison with a fixed base
which is approximately the breed average. Corrections for age and season of calving are used on the
records before the ICC is calculated.

ICC = No. of heifers from bull x No. of contemporaries

No. of heifers from bull + No. of contemporaries

When the breed average increases through improved selection and breeding, the ICC should be
updated with the base average being adjusted.

3. Pedigree Selection or Evaluation / Breeding

A pedigree is a record of parentage so that the genetic relationship between individual animals in the
population is known. Pedigree breeding implies that individual animals of known ancestry are
selected for breeding as the potential parents of the next generation. Each pedigree animal possesses
a pedigree record which describes the name, breed, date of birth, appearance and the performance
of its ancestors, i.e. the parents, grandparents, great grandparents and the great great grandparents.
Therefore, pedigree Selection is the selection of potential parents based on the pedigree
performance of the ancestors.

Pedigree selection is more appropriate and efficient in measuring sex limited traits, such as milk yield,
traits with low heritability and traits that are measured late in life, such as litter size in ewes and sows
and carcase quality in beef.

Subject of Pedigree

Parent Grandparent Great Great Great

Grandparent Grandparent
½ ¼ 1/8 1/16

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4. Sib Testing or Family Selection:

Family selection is also known as Collateral Relatives Methods. Sibs are full brothers and sisters
(progeny that have two common parents). Half-sibs are half brothers and sisters (progeny that have

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 one common parent). Sib-testing is used for selection of traits with low heritability, sex limited traits
and traits that are expressed or can be assessed late in life.

Family Selection involves the selection of individual animals in the same generation according to the
average performance of related animals (full and half sibs). The principle is that related animals (full
and half sibs) have some proportions of their genes in common inherited from their parents, i.e. the
animals will have the same or similar genetic relationship. The use of the average of a number of
collateral relatives reduces environmental factors affecting production. The selection of best
individuals from best families can be more rewarding when both mass and family selections are used.

Selection methods for more than one trait

In livestock improvement it is rare to wish to select for improved performance of only one trait at any one
time. The livestock breeder or owner may be interested in selecting for improved performance of several
traits at a time instead of only one trait. However, the more traits selected for the less the selection pressure
that is exerted on each trait, i.e. the lower is the rate of progress toward the improved performance for each
trait. This occurs because most of the traits are correlated to each other and part of that correlation is genetic
and may be positive or negative, for example, in dairy production animal size positively correlated to
increased milk yield but increased milk yield is negatively correlated to increased butter fat content of the
milk.

The selection pressure on any of the traits depends upon the number of traits being selected for at any one
time. For simultaneous selection of two traits, the selection pressure is 1/√2 = 71% of that possible when only
one trait is selected for (58% for three traits and 50% for four traits). The formula becomes 1/√n when the
number of traits being selected for is n.

Methods have been designed for the selection to achieve improved performance of a number of traits as
preferred (for multiple trait selection). The methods are as follows:

1. Tandem Selection:

Tandem selection, as the name implies, involves the improvement of one trait at a time to the desired
level for several generations, followed by the next trait for several generations and then the next.
The selection technique takes many generations (too long) to achieve the desired performance for a
number of desired traits. The selection technique has been found to be inefficient and the poorest
because when the traits being selected for are negatively correlated because the selection for
improved performance of the second trait may reverse much of the improvement that was achieved
during selection for the first trait.

For example, in sheep, a positive genetic correlation exists between growth rate and mature size. As
a result, attempts to select for increased growth rate invariably results (also) in an increase in mature
size and therefore increased weight of the breeding stock. However, bigger ewes require more feed
for maintenance and this has serious implications when feed is a limiting factor as is the case under
the extensive system of management.

A negative correlation also exists between milk yield and butterfat content of the milk in cattle and
selection for increased butterfat content of the milk tends to reverse much of the improvements for

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 increased milk yield when selecting for the increased butterfat content of the milk. Hence, the
selection technique is rarely used because it is the poorest and simultaneous improvement of the
characters is preferred.

2. Independent Culling Levels:

The technique involves the simultaneous selection for improved performance for number of traits
after setting up independent culling levels for each character to be selected for, i.e. acceptable and
independent levels of performance must be set. Animals that perform above the accepted levels of
performance are chosen as potential parents and the animals which fail to reach the set levels are
discarded. For example, to simultaneously select for improved milk production and butterfat content
of the milk the acceptable minimum levels for milk yield and butterfat content should be defined,
such as, cows producing below 5, 000 litres of milk per lactation and producing milk containing below
3.8% butterfat content are to be discarded.

The disadvantage of the technique is that all the individual animals that perform below the set
independent culling levels are culled even if they may very good in one of the traits, i.e. animals with
high performance in one trait are discarded because there is no compensation for poor performance
in one trait by brilliance in another trait. The advantage of the technique is that the time taken to
effect change is short.

Trait Y 5, 000 kg
(Milk yield)

Trait X (3.8% BF)

3. Selection Index:

The method involves combining measurements for two or more traits into a single value for the
animal, this single value for the animal is referred to as the Selection Index, I. The Selection Index (I)
is valuable in comparing individual animals on a relative basis. The selection index (value) is used as
a criterion for selection and to predict the future genetic and financial worth of the animals. The
advantage of the use of the Selection Index is that it takes into consideration the heritability of each
trait, the relative economic value of each trait and the genetic and phenotypic correlation between
traits.

Though the selection index is widely used it has a number of problems in its use. These problems
include the complexity in its use because it requires computations and access to sophisticated
computation facilities. In addition, the economic value of each trait tends to vary with time (price of
milk per litre), accurate records should be maintained and heritability depend on the variation in the

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population (variation tends to reduce and animals become more uniform with continued selection).
Another weakness of Index Selection is that high merit in one trait can be used to make up for
deficiencies on other traits.

For example, if Cow A produces 6, 000 litres of milk per lactation containing 3.5% butter-fat and Cow
B produces 8, 000 litres of milk per lactation containing 3.0% butter-fat and given that the coefficients
(constants) b1 = 0.7 and b2 = -0.4 and that the Selection Index (I) = b1 M + b2 F; where M = Milk Yield
and F = Butterfat content of the milk.

Selection Index for Cow A = 0.7 x 6,000 + (-0.4 x 3.5) = 4,200 - 1.4 = 4,198.6

Selection Index for Cow B = 0.7 x 8,000 + (-0.4 x 3.0) = 5,600 - 1.2 = 5,598.9

Hence, in terms of selection indices Cow B would be selected yet Cow B produces milk with very low
BF content. Hence, high milk yield tends to compensate for low BF content of the milk.

Considering both the Independent Culling Levels and Selection Index it has been found that any
animal that falls below the set criterion in one trait it is not selected with the Independent Culling
Levels even if the animal has very good performance in one trait because it falls independently below
the selection criterion for that trait, i.e. there is no compensation for poor performance in one trait
by another. With the Selection Index a deficiency in one trait is made up by merit in another. The
Independent Culling Levels and the Selection Index can be looked at in combination as below:

Thus, when Independent Culling Levels and Selection Index are looked at in combination animals in
R and S would be selected when Selection Index is used but not with the Independent Culling Levels.
Animals in T will be selected as potential parents when Independent Culling Levels is used but not
when Selection Index is used.

Tandem selection is the least efficient and Index Selection is 10% more efficient than the Independent
Culling Levels method.

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 Breeding Practices / Systems:

There are various breeds of cattle, sheep, goats, pigs, and other livestock with different levels of
performance. The breed differences in performance can be regarded as having arisen through selection
that was carried out in the past, i.e. through both natural and artificial selection. As a result of different
objectives of past breeders, many breeds have become relatively specialised in function. For example,
there are specific breeds of cattle for milk production and some for beef production but in developing
countries, including Zambia; the local breeds available remain firmly ‘multi-purpose’ in function.

Inbreeding is the mating of individual animals that are more closely related than the average of the population
to which they belong. Inbreeding increases homozygosity and, hence, uniformity in the population, i.e.
inbreeding increases the number of gene pairs that are homozygous because of common parentage, some of
these genes are lethal and some may cause serious defects (defective animals).

The closest degree of inbreeding with the most rapid approach to complete homozygosity is found in self-
fertilising organisms, such as oats, peas, beans, barley and tomato. For example, if a self-pollinating plant
heterozygous at one locus (Aa) is selfed over four generations the following will be observed:

AA x aa

F1 Aa

F2 1AA : 2Aa : 1aa

F3 6AA : 4Aa : 6aa

F4 28AA : 8Aa : 28aa

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 Generation Genotypes Heterozygosity Homozygosity
0 AA Aa aa 100% 0
100%

F1 25 50 25 50 50

F2 37.5 25 37.5 25 75

F3 43.75 12.5 43.75 12.5 87.5

F4 46.875 6.25 46.875 6.25 93.75

Thus, the proportion of homozygous plants is raised in each generation and complete homozygosity is attained
in a little over 10 generations of self-fertilisation. The number of homozygous genotypes is increased by 50% in
each generation as a result of selfing or inbreeding. Heterozygosity also reduces by 50% in each generation. After
10 generations heterozygosity will be totally eliminated.

The continuous matings of brother and sister results in a slower increase in homozygosity and takes 18
generations to attain much homozygosity as compared to the one that occurs over a period of 10 generations
with self-fertilising plants. Matings of brother and sister (full sibs) reduces residual heterozygosity by about 19%
in each generation and mating of half-sibs reduces heterozygosity by only 11% in each generation and hence
they are much safer processes.

Continued breeding within one herd/flock without introducing new blood also results in an increase in
homozygosity. The increase in homozygosity is dependent largely upon the number of sires used to produce
each generation. In most domestic animals the number of females used for breeding purposes greatly exceeds
the number of males and in such situations heterozygosity is reduced in each generation by about 1/8N; where
N is the number of males used.

For example, if four males are used per season homozygosity is increased by about 3.1% in each generation. If
10 sires are used the increase in homozygosity is about 1.2% each generation. Gradual approaches to
homozygosity are not likely to hurt the herd/flock as long as defective males are excluded from reproduction
(and reduced further but special care must be taken to avoid the mating of close relatives).

The mating of brother and sister or parent and the off-spring constitute close inbreeding. Close inbreeding
reduces the efficiency of reproduction. In addition, lethal traits and other defects appear in the progeny. The bad
genes cause defects when they become homozygous and should be eliminated or discarded (rogued out).
However, they take with them a lot of good genes that they are linked with. Hence, highly inbred lines are poor
and do not reproduce, a condition referred to as inbreeding depression. In domestic animals, such mating leads
to the extinction of the line within 6 to 8 generations because of failure to reproduce. Inbreeding causes animals

9
that also lack vigour, for example, the production of small litters in dogs and pigs and low hatchability of eggs in
poultry.

Inbred animals will have similar genes at loci because they have a common ancestor and the intensity of
relationship or the fact that an animal has similar genes at loci is called the coefficient of inbreeding. The
coefficient of inbreeding, F, is the measure of the probability that an individual animal carries two genes at loci
that are alike (similar) by descent. The F or degree of inbreeding in any individual (x) may be calculated from the
formula by Wright:

Fx = Σ(½)n(1 + FA)
Where:
n = The number of individuals in each path linking the parents of the animal to a common ancestor. The value of
n is got by counting the parents, the common ancestor and every other individual in the path; and

FA = The Inbreeding Coefficient of a common ancestor. If it is unknown then it is taken as being zero.

Wright's coefficient is a measure of the degree of relationship between the sire and the dam and will depend on
how far back both the dam and the sire have a common ancestor. If they have more than one such common
ancestor, they are likely to be more closely related than if they had only one. Then separate contributions of
each common ancestor must be calculated. These are then added up (Σ) to determine Fx.

The ½ in the formula recognises the fact that the contribution of an ancestor's genes to the individual x is halved
with each generation that separates them. To correctly apply the formula the pedigree must be shown with a
path (arrow) diagram in order to calculate Fx.

For example:

1. Half-Sib Mating:

C x D C x E D C E

A B
A x B
x

PATH: A ---- C ---- B, hence n = 3 and FA = 0. The common ancestor is C.

Fx = Σ(½)n(1 + FA) = Σ(½)3(1 + 0) = 0.125 . Hence, the individual x is 12.5% inbred, i.e. 12.5% of the alleles at loci
are identical by descent and any deleterious effects in the genes have 12.5% chance of appearing in homozygous
form in the progeny.

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2. Full Sib Mating (Brother X Sister Mating):

C x D C x D C D

A x B A B

x
x

Paths: A ---- D ----- B, hence n = 3 and FA = 0

A ---- C ----- B, hence n = 3 and FA = 0. The common ancestors are C and D.

Fx = Σ(½)n(1 + FA) = (½)3(1 + 0) + (½)3(1 + 0) = 0.125 + 0.125 = 0.25. Hence the individual is x is 25% inbred, i.e.
25% of the alleles at loci are identical by descent and any deleterious effects in the genes have 25% chance of
appearing in homozygous form in the progeny.

3. Two Generations of Mating Brother X Sister:

Ax B A x B A x B A x B A B

M x N M x N M N

S x D S D

x
x

The common ancestors are M, N, A and B.

Paths: S ---- M ---- D, hence n = 3 and FA = 0
S ---- N ---- D, hence n = 3 and FA = 0
S ---- M ---- A ---- N ---- D, hence n = 5 and FA = 0
S ---- M ---- B ---- N ---- D, hence n = 5 and FA = 0
S ---- N ---- A ---- M ---- D, hence n = 5 and FA = 0
S ---- N ---- B ---- M ---- D, hence n = 5 and FA = 0

Fx = Σ(½)n(1 + FA)
= (½)3(1+0) + (½)3(1+0) + (½)5(1+0) + (½)5(1+0) + (½)5(1+0) + (½)5(1+0)
= 0.125 + 0.125 + 0.03125 + 0.03125 + 0.03125 + 0.03125 + 0.03125
= 0.375

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4. Sire Daughter Mating:

G D

Path: D --- C, hence n = 2 and FA = 0

C Fx = Σ(½)n(1 + FA) = (½)2 (1 + 0) = 0.25

5.
F A B

H E

G C D

The common ancestor E is inbred because its parents, A and C are full sibs.
FE = Σ(½)n(1 + FA) = (½)3 (1 + 0) = 0.25. Hence, FH = Σ(½)n(1 + FA) = (½)3 (1 + 0.25) = 0.125 x 1.25 = 0.1563

6.

C I

A K

X D J

B G L

The common ancestor D is inbred because its parents, K and G are half sibs. Similarly, the individual B is inbred
but the individual B is not a common ancestor in this path.

PATHS: n (½)n Inbreeding Contribution

A -- D -- G -- E -- B 5 0.0313 0 0.0313
A -- D -- B 3 0.125 0.125 0.1406
A--D--K--J--G--E--B 7 0.0078 0 0.0078
Fx = 0.1797

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Example 7:

A E

x D

B G

The common ancestor A has the same grand-sire E on both the dam's and sire's sides of the pedigree.

Path: C --- E --- D; hence n = 3 and FA = 0

FA = Σ(½)3 (1 + 0) = 0.125

Contribution of A and B to Fx:

Paths: A --- B; hence n = 2 and FA = 0.125.

A --- D --- G --- H --- B; hence n = 5 and FA = 0.
A --- C --- E --- D ---G --- H --- B; n = 7 and FA = 0

Fx = Σ(½)n (1 + FA)
= (½)2 (1 + 0.125) + (½)5 (1 + 0) + (½)7 (1 + 0)
= (0.25 x 1.125) + (0.03125 x 1) + (0.0078125 x 1)
= 0.28125 + 0.03125 + 0.0078125
= 0.3203125

Coefficient of Relationship:
The coefficient of relationship is used to describe how closely related two animals may be, i.e. the intensity of
relationship, Rxy. The coefficient of relationship is generally twice the Coefficient of Inbreeding, F. Rxy can be
calculated from the formula:

Rxy = Σ(½)n(1 + FA) / √ (1 + Fx)(1 + Fy)

Where n = The number of generations between the individuals x and y.

FA, Fx, Fy = The Coefficient of Inbreeding of the common ancestors, and the individuals x and y.

Since the Coefficient of Inbreeding, F is generally be half the Coefficient of Relationship, R, it is easier to work out
the F that would result when related animals are mated together and then double the figure to get the R. For
example:

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8. Full Sib Mating:

X S Paths: x -- s -- y, n = 2 and x -- d -- y, n = 2

Y D

Rxy = (½)2(1+FA) + (½)2(1+FA) / √ (1 + 0)(1 + 0)

= 0.25 + 0.25 = 0.5

9. S A Paths: S -- A -- B -- D, n = 3 and S -- B -- D, n = 2

D B FS = (½)2(1+FA) = 0.25

Hence, Rxy = (½)3(1+FA) + (½)2(1+FA) / √ 1.25 x 1 = 0.375 / 1.25 = 0.3354

10.
B

D A

F C

The common ancestor D is inbred because its parents, B and C are half sibs. Hence, FD = (½)3 (1+FA) = 0.125

Paths: D – C – E
D–B–A–C–E

Hence, RDE = (½)2 (1+FA) + (½)4 (1+FA) / √ 1 + 0.125 x 1 + 0

= 0.25 + 0.0625 / √ 1.125 = 0.3125 / 1.0606
= 0.2946

Thus, inbreeding results in the reduction in fitness and the reduction in fitness is referred to as Inbreeding
Depression. Inbreeding depression is seen as reduced fitness, i.e. reduced performance, vigour, fertility and
increased mortality rate. Hence, inbreeding is not desirable and is not a method livestock improvement. It is used
to breed animals that are genetically homozygous and is used to reveal or expose hereditary abnormalities and
lethal traits which are masked in the original animals/cattle, i.e. to expose the desirable and undesirable

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 traits/genes and the undesirable traits/genes are rogued out (culled). In this way, inbreeding is used to fix genes
(breed formation). Inbreeding is used to show defects hidden in the original cattle used.

2. Outcrossing and Cross-Breeding

Out-crossing is the mating of two unrelated pure bred lines and out-crossing results in stronger, healthier, larger and
vigorous F1 individuals, the condition referred to as heterosis or hybrid vigour that is seen as increased fitness or
vigour. For example, the maize hydrids with great vigour are a result of out-crossing known inbred maize lines.

Cross breeding is the mating of two distinct breeds or races or different species to get a heterozygous progeny in the
F1 in which the level of fitness is increased. The increase in fitness is referred to as Hybrid Vigour. Hybrid vigour is
seen as an improvement in fertility, performance and viability (it is the reverse of the loss of vigour resulting from
inbreeding). Cross-breeding increases heterozygosity, increased fitness of such individuals is also called heterosis.
Under normal conditions hybrids cannot be used for further breeding and a nucleus of pure breeds is always
necessary to create the cross-breeds.

Hybrid vigour can be defined as extra vigour exceeding that of both parent stocks as shown by hybrids from crossing
of species, breeds, strains and inbred lines. Hybrid vigour is expressed as more rapid growth, larger size, increased
productivity and better viability. The crossing of breeds is often adopted in commercial practice to advantage of
hybrid vigour. Crossbreeding by mating two different pure breeds is used to improve the productivity of tropical
breeds with the view of combining different qualities from each breed, i.e. the offspring inherits all the good qualities
of either parent. For example, hybrid maize (corn) has increased yield and the mule, a result of crossbreeding the
male donkey (Equus asimus, 2n = 62) with the female horse (Equus caballus, 2n = 64), is famous for its hardiness and
capacity to work.

Hybrid vigour is at maximum (100%) in the F1; hybrid vigour is at 50% in the F2 and continues to reduce by 50% in
later generations from then. It is for this reason that parent strains of animals must be maintained and fresh crosses
made anew for each generation. However, in some plant species hybrids can be kept going indefinitely by using
asexual reproduction forms such as budding, grafting and propagation of stems, tubers or rhizomes.

The theories about forces that might be responsible for hybrid vigour are known.

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 Generation Genotypes Heterozygosity Homozygosity
0 AA Aa aa 100% 0
100%

F1 25 50 25 50 50

F2 37.5 25 37.5 25 75

F3 43.75 12.5 43.75 12.5 87.5

F4 46.875 6.25 46.875 6.25 93.75

The first is the theory of dominant genes raising vigour and the tendency of dominant genes to suppress recessive
genes. Therefore, a cross between two animals will combines in the progeny all the dominant genes that were
homozygous in either parent plus about half of those that were heterozygous. Parents distantly related have a
greater chance of contributing differing dominant genes to their progeny and thus increasing hybrid vigour. For
example:

AAbbccDDee x aaBBccddEE
Loci with dominant genes = 2 2

AaBbccDdEe
Loci with dominant genes = 4

All the recessives will be masked by the dominant alleles (except one) and therefore such animals will be more
vigorous than either parent.

The second theory is that of heterosis or over-dominance in which the heterozygote is more vigorous than either
parents, i.e. the dominant alleles may stir up biochemical processes in the cell that will make the cell more active or
more vigorous. This condition in the heterozygote is referred to as heterosis, synonymous with hybrid vigour. Some
breeders refer to the same as over-dominance.

The merit (good performance) of crosses of the Bos taurus and Bos indicus breeds of cattle cannot be attributed to
hybrid vigour because hybrid vigour should not persist for more than one or two generations after a cross is made
except in plant hybrids that are maintained by asexual reproduction. Bos taurus and Bos indicus crosses owe their
special value to the good characters introduced by Zebus, particularly tolerance to heat, the repulsion of ticks that
makes them resist tick-borne diseases. Milk yield in crossbred cattle is also 20% higher.

The cross-bred pigs have higher litter size at birth and at weaning and get to market weight faster and on less feed.
Similarly, every broiler in the world is a cross between two or more breeds and the broilers reach market weight

16
faster and on less feed. This has today made the chicken as the cheapest high quality meat readily available on market
and this is due to hybrid vigour.

Domestic animal breeders can get the most hybrid vigour at the least expense by finding a strain that has general
combining ability - the capacity to cross well with most others and to yield profitable amounts of hybrid vigour. Some
stocks may cross well (nick) with certain other strains but not with all strains. Such are said to have specific combining
ability. A scheme to enhance specific combining ability is the use of reciprocal recurrent selection.

2.1 Grading Up

Grading up is the changing of one breed to another by continued crossing. This is achieved by mating the exotic breed
to the indigenous breed and the cross-bred females. The exotic bull (or artificial insemination bulls' semen) is mated
to the whole herd but his progeny are mated by different bulls of the same breed to avoid in-breeding. Grading up
is inappropriate if the environment is not suitable for the new breed. For example:

100% Angoni x 100% Friesian Heterosis

F1 50% A:50% F x Friesian 100%

F2 25% A:75% F x Friesian 50%

F3 12.5% A:87.5% F x Friesian 25%

F4 6.25% A:93.75% F Near 0

The cross-bred animals will be almost entirely of the exotic breed by the F4 generation with 93.75% exotic blood and
will be acceptable as pure-bred animals. Note that hybrid vigour reduces by 50% in each generation of grading up.

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2.2 Rotational Crossing

This is the continuous alternate crossing of two or more breeds.

2.2.1 Criss-Crossing

The alternate crossing of two breeds is called criss-crossing, for example:

100% Angoni x 100% Friesian Heterosis

F1 50% A : 50% F x Friesian 1.0

F2 Angoni x 25% A : 75% Friesian 0.5

F3 62.5% A : 37.5% F x Friesian 0.75

F4 31.25 A : 68.75 F 0.62

The proportion of genes settles down to a ratio of ⅔ rds of the additive genetic effects of the last sire used and 1/3rd
of the additive genetic effects of the other sire. The proportion of hybrid vigour is variable from generation to
generation.

2.2.2 Three Breed Rotation:

Crosses between two breeds to produce the F1 hybrid are sometimes followed by mating the hybrid to a third breed
to produce a three-way cross.

100% Breed A X 100% Breed B Heterosis

F1 100% Breed C x 50% A : 50% B 1.0

F2 25% A:25% B:50% C x Breed A 0.5

F3 Breed B x 62.5% A:12.5% B:25% C 0.75

F4 31.3% A:56.3% B:12.5% C 0.62

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The breed that was used last will predominate as far as the genotype is concerned. Rotational crossing is only
appropriate where the breeds used have the same function or their performance is not too different. Otherwise, the
proportion of the local blood will be variable from generation to generation and so will be the proportion of heterosis
attained. However, after sometime the population will reach equilibrium and the predominant genotype can be
determined from the formula:

50 x 2n / 2n – 1; where n = the number of breeds used.

For example, if three (3) breeds are used 50 x 2n = 50 x 8 = 400 = 57.1

2n - 1 8-1 7

Therefore, at equilibrium the proportion of genes contributed by the last sire is 57.1% when three breeds are used
in rotational crossing.

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