Module 1

Introduction to Animal Breeding

At the end of this module you are expected to:
1. Define basic concepts and terminologies used animal breeding;
2. Understand the terms, principles and aspects that govern animal breeding;
3. Know the history of animal breeding including animal domestication;
4. Learn and be familiar with the different breeds of common domesticated animals;
5. Appreciate the importance of animal breeding in our daily lives.

What is animal breeding?

 Animal breeding is about selective breeding: only use males and females for
breeding that have passed a certain quality criterion.
 With a predefined goal in mind: to genetically improve the population in a certain
direction.
 People make a plan with the intention to select the best animals according to a
predefined list of requisites (traits), and use those selected animals for breeding
the next generation so that the offspring on average will be better than the parents.
o In other words: selective breeding causes a shift in population average from
one generation to the next.
 Although at first instance you may think that animal breeding involves keeping
animals and making sure they reproduce, and it thus would involve optimizing
reproduction techniques or something along those lines, this is not the case.

Five Important Aspects of Animal Breeding

1. Most importantly, obviously for selective breeding to be successful it is essential

that the trait (e.g. running speed or milk production or coat color) under selection
is heritable.
2. That animals have different genetic backgrounds so that selection is possible.
3. The direction of selection is defined by humans and they decide which animals are
allowed to mate and produce members of the next generation.
4. Success of animal breeding can be judged by looking at a shift in population
average phenotype from one generation to the next. So, animal breeding works at
population level, not automatically at individual level.
5. Success of animal breeding can be measured as the cumulative result of multiple
generations of selection. Breeding decisions are made with the future in mind.

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Heritable Traits and Phenotype

A trait is "a distinguishing phenotypic characteristic, typically belonging to an individual".

In practice this means anything you can record or measure on an individual.

A phenotype is that what you observe or measure on the animal for a certain trait. It can
depend both on the genetic background of the animal (provided it is heritable) and
external circumstances such as level of nutrition.

 Selective breeding will only be successful in case the trait under selection is
heritable.
o Because only a fraction of the animals is selected for breeding, so is allowed
to produce offspring, and because the trait is heritable, the performance in
the offspring will resemble that of the parents.
 Therefore, only the best parents are used for breeding and the average of the next
generation will be better than that of the current.

 A trait is heritable if the performance for that trait, at least in part, depends on the
genetic make-up (DNA) of an animal.
 Differences in performance between animals can (partly) be explained by genetic
differences between animals.

Selection by Nature

Natural selection is the process whereby animals that are better adapted to their
environment have a higher change to survive and produce more offspring than less
adapted animals. The next generation thus, on average, will be more adapted than the
current generation.

 It sounds as if animal breeding is all in the hands of the humans.

o Compared to natural populations this indeed is the case, as we decide
which animals are allowed offspring and which are not: selective breeding
or in other words artificial selection.
 However, as in natural populations there is another force that plays an important
role and that is the force of natural selection.
 In natural selection it is not us but the environment that determines survival and
reproductive success of animals.
 So, after we have decided which animals we intend as parents, they still need to
be able to survive until reproductive age and to be able to reproduce successfully.
o Natural selection also results in directional change in the population
average.
o Animals adapt to their environment and the ones who can do that best will
be the most successful in survival and reproduction.
o In other words: in natural selection the direction of selection is on
adaptation to the environment.

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Natural Selection vs Artificial Selection (Selective Breeding)

 In some cases, natural selection work in the opposite direction of selective

breeding.
 In those cases, without human intervention the animals with the desired qualities
will be less successful in surviving and/or producing offspring.
 For example, the fact that in many cows there is a negative relationship between
high milk production and being able to get pregnant, shows that animals with the
desired quality: high milk production, are less likely to produce offspring unless
there is extra effort put into it by the farmer.
 Also, the fact that very high producing cows often have health problems indicates
that their chance of producing offspring is reduced compared to their more average
producing 'sisters'.
 Selective breeding often competes with natural selection.

Domestication and animal breeding

Domestication is the process of conversion of wild animals to domestic use.

 Humans have become so familiar to the fact that some of the best animals in a
breed require assistance with some aspects of their survivability and/or
reproduction that we think it is normal.
 Domestic animals are 'created' by humans and to maintain that we accept certain
disadvantages.

 Domestic animals need to live in (close) association with humans, therefore they
have to become tame.
 They also have to meet the expectations of their owners for the purpose of keeping
them.
 This can be achieved by selective breeding. Expectations of owners will change in
time, followed by a change in selective breeding plan.
 Domestication often has resulted in a type of animals that has become quite
different from their wild counterparts.
 As a result, domestication often also involves the development of a dependency
on humans so that the animals lose their ability to live in the wild.

Domestication of the dog

 The first animal species to be domesticated was the dog. Estimates of when this
happened vary a lot, but it was approximately 12,000 years ago.
 An appealing theory of how this happened is that when people started to settle
down and become farmers, they also started to accumulate waste.
 The tamer than average wolves were brave enough to eat from that waste and
thus had a secure source of food.

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  This was an advantage, so natural selection pressure was on being not very afraid
of humans.
 Eventually, a kind of symbiotic relationship developed, where these ancestors of
the dog started to perform 'tasks' like warning the humans for approaching danger,
helping in hunting, provide warmth, etc. and in return these animals would receive
food security.
 This type of symbiotic relationship is still present in village dog populations in Africa
and Asia, and also in some Southern European countries.
 It is believed that our current domestic dog breeds originate from these village
dogs. There is evidence that genetically the village dogs are in between the wolves
and the dogs.

Domestication of other species

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Prerequisites for domestication

 Domestication is not always successful.

o Despite many attempts, the zebra, for example, has not been domesticated.
o Even though it is closely related to the horse and the donkey and you can
keep it in an enclosed area where it will survive and reproduce, apart from
the exceptional case, it has not been successfully tamed.
o Several generations in captivity and some selective breeding did not make
the zebra genetically tame so that it can be ridden. Why is that? People are
not sure, but there is a list of prerequisites for successful domestication that
seem to hold. The zebra may not meet one or more of them.

The apparent prerequisites are:

1. The animals should be able to adapt to the type of feed they are offered by humans.
This may be different (in diversity) from what they were used to in the wild.
2. Animal must be able to survive and reproduce in the relatively closed quarters of
captivity. Animals that need a very large territory are not suitable to be
domesticated.

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 3. Animals need to be naturally calm. Very skittish or flighty animals will be hard to
prevent escaping.
4. Animals need to be willing to recognize humans as their superior, which means
they must have a flexible social hierarchy.

Origin of animal breeding: a history of science

The start in the 18-th century

 Until roughly the 1700's animal breeding, as in selective breeding, did not really
exist.
 People mated their animals with animals in the neighborhood that they liked.
 There was no systematic way of selecting animals for reproduction, based on
predefined characteristics that did not change from mating to mating, but remained
similar in time.
 Sir Robert Bakewell (1725 – 1795) introduced keeping accurate records of
performance of animals so that objective selection became possible.
o He used inbreeding (mating of related animals with similar traits) to fix
certain characteristics in animals and he also introduced progeny testing:
the method of evaluating performance of the first (small) group of progenies
and use that information to select the best father of future progeny.
o He promoted the idea to 'breed the best to the best'.
o Bakewell developed the New Leicester sheep from the old Lincolnshire
breed.
 The New Leicester had good quality fleece and a good fatty shoulder
that was popular at the time.
o Bakewell also noticed that Longhorn cattle were growing well and used less
feed compared to other cattle.
 So, he developed that further in order to grow more meat efficiently.
It is amazing he did this without knowing anything about
genetics.

Establishment of herdbooks

 With time the number of people increased who were using the selective breeding
approach introduced by Bakewell.
 With the growing number of generations of selective breeding, it became
increasingly difficult to remember the relationships between the animals, especially
further back in the pedigree.
 This was the reason to start recording pedigree on paper, so that correct
information could be reproduced and it could be proven that an animal was of a
certain breed.
 The first herdbook was for the thoroughbred horse and was established in England
in 1791.

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  This book did not contain all pedigree, but only those of horses that were winning
important races. Following the race horses, the Shorthorn cattle (1822) were next
to start a herdbook.
 In the rest of Europe, herdbooks only started to be established in from 1826
onwards for horses (in France), and from 1855 onwards for cattle (also in France).
 The first international herdbook was established for the American Berkshire pigs
in 1876.
 The first dog in the Netherlands was registered by the Koningklijke Nederlandsche
Jachtvereeninging Nimrod (predecessor of the Raad van Beheer op Kynologisch
Gebied in Nederland) in 1874.
 After the turn of the century animal breeding within herdbook settings became
standard.

Creation of breeds

 With the establishments of herdbooks, breeds were formed.

 There is still debate on what is a true definition of the term “breed”.
o This is nicely illustrated in dog breeding by the fact that the Fédération
Cynologique Internationale (FCI), the international federation of kennel
clubs, which are national organizations across herdbooks, recognizes 339
separate breeds, while the English Kennel club recognizes 210 breeds, and
the American Kennel club even only 162.
 It is interesting to realize that these herdbooks were established without any
knowledge about genetics. Breeders had a feeling about inheritance and that was
sufficient to invent this selective breeding.

A breed is a group of animals of a certain species that through generations of selective

breeding has become uniform in performance, appearance, and selection history.

A species is the largest group of animals that are capable of interbreeding and producing
fertile offspring.

Breeding in the 19-th century

 In 1859, Charles Darwin (1809 – 1882) published his book “On the origin of
species”, based on the findings that he collected during his voyage on “the Beagle”.
o He discovered the forces of natural selection.
o He also concluded that the individuals that fit best in their environment have
the highest chance to survive and reproduce: they are the fittest.
Consequently, different environments result in different directions of
selection pressure. He based this on his findings on the Galapagos islands,
where finches on one island were different from finches on the next island.
His conclusion was that the difference in food source, predators present,
etc. between the islands had made the finches develop differently over very
many generations. They adapted to their specific environments.

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 o Darwin translated his ideas to domesticated species as well: "We cannot
suppose that all the breeds were suddenly produced as perfect and as
useful as we see now them; indeed, in several cases, we know that this has
not been their history. The key is man's power of accumulative selection:
nature gives successive variations; man adds them up in certain directions
useful to him. In this sense he may be said to make for himself useful
breeds" C. Darwin. On the Origin of species (1859, p.30) Still, Darwin did
not know about the basic laws of inheritance.

 It was the monk Gregor Mendel, who in 1865 published the results of his studies
of genetic inheritance in garden peas.
o He showed that genetic material is inherited from both parents,
independently of each other. And that each (diploid) individual thus carries
2 copies of the same gene, of which only 1 is passed on to their offspring
and which one is a result of chance (independent assortment).
o He also showed that these gene copies (alleles) can be dominant (only 1
copy determines the expression of the gene), recessive (2 copies are
required for expression), or additive (a copy of both alleles result in an
expression that is intermediate to that of having 2 copies of either of the
alleles).
o These findings had no immediate impact on animal breeding and were
not recognized as important until 1900.

Animal breeding in the 20-th century

 Most of the animal breeding theory we are still using today, was invented in the
first half of the 20th century.

 Statistician R. A. Fisher (1890 – 1962) showed that the diversity of expression of

a trait could depend on the involvement of a large number of so-called Mendelian
factors (genes). He published a lot related to statistics and animal breeding, but
his keynote paper came out in 1918.

 Fisher, together with Sewall Wright (1889 – 1988) and J.B.S. Haldane, were the
founders of theoretical population genetics.

 Thomas Hunt Morgan (1866-1945) and coworkers connected the chromosome

theory of inheritance to the work by Mendel and created a theory where
chromosomes of cells were believed to carry the actual hereditary material.
o Morgan won the Nobel prize for it in 1933.

 Jay L. Lush (1896 – 1982), who is known as the “modern father of animal
breeding”. He advocated that instead of subjective appearance, animal breeding
should be based on a combination of quantitative statistics and genetic information.
o His book “Animal Breeding Plans” that was published in 1937 greatly
influenced animal breeding around the world.

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 Lanoy Nelson Hazel (1911-1992) was inspired by Lush's book and started
working for him.
o He received his PhD degree in 1941 and in that PhD thesis he developed
the “selection index theory”, a method used for decades to determine
what weights should be put on the different traits under selection.
o In the process of developing this method he also came up with a concept
on how to estimate genetic correlations. This is essential for assigning the
proper weight to selection traits.
o Hazel also developed a method using least squares, a statistical technique,
for more complicated data with unequal numbers of subclasses as often
occur in animal data. Until then, statistical techniques by Hazel were used
to optimize weighing the performances for various traits in animals to select
those with the most optimal combination.

 The estimated breeding value (ebv) was only developed later by the statistician
C. R. Henderson (1911 – 1989), who was a student of Hazel in Ames.
o The estimated breeding value made it possible to rank the animals
according to their estimated genetic potential (the ebv), which resulted in
more accurate selection results and thus a faster genetic improvement
across generations.
o Henderson further improved the accuracy of the estimated breeding value
by deriving the best linear unbiased prediction (BLUP) of the ebv in 1950,
but the term was only used since 1960. He also suggested to integrate the
full pedigree of the population to include genetic relationships between
individuals.
o This way performance of relatives could be included in estimating the
breeding value of an individual. The so-called animal model was born.
o Unfortunately, in those days the computer power was too limited to be able
to also calculate the breeding values using the animal model. The practical
implementation thus had to wait until the later 1980's. Current great minds
that have developed a way to incorporate large scale DNA information that
has become available in animal model (BLUP) theory to estimate the so-
called genomic breeding values are Theo Meissen (professor in Ås,
Norway) and Mike Goddard (professor in Melbourne, Australia).

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Learning Activity 1

Make a portfolio of the different breeds of the following species of animals (minimum of
10 breeds each) include brief description and use of the said breed;

1. Swine
2. Cattle
3. Carabao
4. Sheep
5. Goats
6. Horses
7. Dogs
8. Cats
9. Ducks
10. Chicken

Example:

Brahman
Description:
• Originated in India
• Able to survive on very little, poor
feed
• Insect & heat resistant
• Excess skin and large hump on
back
• White to gray, red to black
• Posses sweat glands

Guide Questions 1

1. What is animal breeding?

2. Why do we need to study animal breeding?

Answer to Guide Questions 1

Question 1: Animal breeding involves the selective breeding of domestic animals with the
intention to improve desirable (and heritable) qualities in the next generation.
Question 2: It is important to study animal breeding in order to address the challenges of
increasing human population that also increases the demands for a sustainable source
of food and other resources derived from animals.

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Key points

1. Animal breeding is about selective breeding: only use males and females for
breeding that have passed a certain quality criterion.

2. A trait is "a distinguishing phenotypic characteristic, typically belonging to an

individual" while a phenotype is that what you observe or measure on the animal
for a certain trait.

3. Natural selection is the process whereby animals that are better adapted to their
environment have a higher change to survive and produce more offspring than
less adapted animals.

4. Domestication is the process of conversion of wild animals to domestic use.

5. A breed is a group of animals of a certain species that through generations of

selective breeding has become uniform in performance, appearance, and
selection history while a species is the largest group of animals that are capable
of interbreeding and producing fertile offspring.

6. History of animal breeding started when Sir Robert Bakewell (1725 – 1795)
introduced keeping accurate records of performance of animals so that objective
selection became possible which was followed by the significant contributions of
well-known scientist like Charles Darwin and Gregor Mendel with their works on
natural selection and genetics, respectively.

7. Modern day animal breeding was attributed to Jay L. Lush (1896 – 1982), who is
known as the “modern father of animal breeding”. He advocated that instead of
subjective appearance, animal breeding should be based on a combination of
quantitative statistics and genetic information. His works was published in his book
“Animal Breeding Plans” in 1937 that greatly influenced animal breeding around
the world.

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References and Supplementary Materials

Books

1. van der Waaij, K. (2014). Textbook Animal Breeding: Animal Breeding and
Genetics for BSc Students: Centre for Genetic Resources and Animal Breeding
and Genomics Group, Wageningen University and Research Centre, the
Netherlands

Online Supplementary Reading Material

1. Introduction to Breeding and Genetics; https://jvanderw.une.edu.au/gene251-351-

lec1.pdf; Retrieved July 9, 2020

2. Dihybrid Cross; https://www.pinterest.ph/pin; Retrieved July 9, 2020

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