Heterosis

Heterosis, hybrid vigor, or outbreeding enhancement, is the improved or increased function of

any biological quality in a hybrid offspring. The adjective derived from heterosis is heterotic.
An offspring exhibits heterosis if its traits are enhanced as a result of mixing the genetic contributions
of its parents. These effects can be due to Mendelian or non-Mendelian inheritance
Controversy
The term heterosis often causes confusion and even controversy, particularly in selective breeding
of domestic animals, because it is sometimes claimed that all crossbred plants and animals are
"genetically superior" to their parents, due to heterosis. However, there are two problems with this
claim:
First, "genetic superiority" is an ill-defined term and not generally accepted terminology within the
scientific field of genetics. A related term fitness is well defined, but it can rarely be directly
measured. Instead, scientists use objective, measurable quantities, such as the number of seeds a
plant produces, the germination rate of a seed, or the percentage of organisms that survive to
reproductive age.
[3]
Within this perspective, crossbred plants and animals exhibiting heterosis may
have "superior" production on these scales, but this does not necessarily equate to evidence of
"genetic superiority". Use of the term "genetic superiority" is a value judgement, generally in the
realm of politics, and is not science.
Second, not all hybrids exhibit heterosis (see outbreeding depression).
A clearly ambiguous counter-example to any value judgement on hybrids and hybrid vigor is the mule.
While mules are almost always infertile, they are valued for a combination of hardiness and
temperament that is different from either of their horse or donkey parents. While these qualities may
make them "superior" for particular uses by humans, the infertility issue implies that these animals
would most likely become extinct without the intervention of humans through animal husbandry,
making them "inferior" in terms of natural selection.
Some modern geneticists refrain from even using the terms inferior and superior due to the
association of these words with political movements that espouse genocide.
Genetic and epigenetic bases of heterosis
Since the early 1900s (as discussed in the article Dominance versus overdominance) two competing
genetic hypotheses, not necessarily mutually exclusive, have been developed to explain hybrid vigor.
More recently, an epigenetic component of hybrid vigor has also been established.

The genetic dominance hypothesis attributes the superiority of hybrids to the masking of expression
of undesirable (deleterious) recessive alleles from one parent by dominant (usually wild-type) alleles
from the other (see Complementation (genetics)). It attributes the poor performance of inbred strains
to the expression of homozygous deleterious recessive alleles. The genetic overdominance
hypothesis states that some combinations of alleles (which can be obtained by crossing two inbred
strains) are especially advantageous when paired in a heterozygous individual. This hypothesis is
commonly invoked to explain the persistence of some alleles (most famously the Sickle cell trait allele)
that are harmful in homozygotes. In normal circumstances, such harmful alleles would be removed
from a population through the process of natural selection. Like the dominance hypothesis, it
attributes the poor performance of inbred strains to expression of such harmful recessive alleles. In
any case, outcross matings provide the benefit of masking deleterious recessive alleles in progeny.
This benefit has been proposed to be a major factor in the maintenance of sexual reproduction among
eukaryotes, as summarized in the article Evolution of sexual reproduction.
An epigenetic contribution to heterosis has been established in plants and it has also been reported in
animals.MicroRNAs (miRNAs), discovered in 1993, are a class of non-coding small RNAs which repress
the translation of messenger RNAs (mRNAs) or cause degradation of mRNAs. In hybrid plants, most
miRNAs have non-additive expression (it might be higher or lower than the levels in the parents)

This
suggests that the small RNAs are involved in the growth, vigor and adaptation of hybrids.

It was also shown that hybrid vigor in an allopolyploid hybrid of two Arabidopsis species was due to
epigenetic control in the upstream regions of two genes, which caused major downstream alteration in
chlorophyll and starch accumulation. The mechanism involves acetylation and/or methylation of
specific amino acids in histone H3, a protein closely associated with DNA, which can either activate or
repress associated genes.
One example of where particular genes may be important in vertebrate animals for heterosis is
the major histocompatibility complex. Vertebrates inherit several copies of both MHC class I and MHC
class II from each parent, which are used inantigen presentation as part of the adaptive immune
system. Each different copy of the genes is able to bind and present a different set of potential
peptides to T-lymphocytes. These genes are highly polymorphic throughout populations, but will be
more similar in smaller, more closely related populations. Breeding between more genetically distant
individuals will decrease the chance of inheriting two alleles which are the same or similar, allowing a
more diverse range of peptides to be presented. This therefore gives a decreased chance that any
particular pathogen will not be recognised, and means that more antigenic proteins on any pathogen
are likely to be recognised, giving a greater range of T-cell activation and therefore a greater
response. This will also mean that the immunity acquired to the pathogen will be against a greater
range of antigens, meaning that the pathogen must mutate more before immunity is lost. Thus
hybrids will be less likely to be succumb to pathogenic disease and will be more capable of fighting off
infection.


INTRODUCTION to ANIMAL BREEDING

From the very early days human beings depend on animals and animal products for food
and other requirements. In dairy and poultry farms high yielding animals are reared. These
high yielding animals are produced by hybridisation experiments. Previously the animals
were developed basing on unscientific methods. Before the discovery of principles of
heredity human beings have selected the animals with required characters and learned to
develop the plants having the selected characters. This phenomenon is called Artificial
selection. However, an increased knowledge of biology, especially genetics, has helped in
improving the quality of animals and animal products as per the human requirements.
ANIMAL BREEDING-PRINCIPLES
The animal breeder faces many complex problems during hybridisation experiments
because many traits of animals are dependent on the interaction of multiple genes. When
the attempts are made only to increase the size of eggs in fowls,it was observed that the
progeny produced yielded few number of eggs or even they die sometimes. That is if only
one character is taken for improvement of the animals, the other characters will degenerate
or result in harmful effects. Hence at the time of selection all the desirable characters are to
be taken into consideration. The techniques for the improvement of animals involve
principles of selection based on quantitative variations. It is not possible for all of the
desirable traits to be obtained in one individual. The successful product must contain
maximum number of desirable traits and a minimum number of undesirable traits.
1. The body form
It is an important factor in selecting racially improved variety of animals. A certain body
form in cattle and broilers will be having high market value. They yield delicious mutton if
they are having well built body form.
2. Productivity
This is of great significance to the breeder. Some times it has first priority over other traits.
For example the number of eggs, quantity of milk, or wool per animal is an important
criterion in any programme of improvement of animals.
3. Quality of the product
In addition to the quantity, the quality of the productivity is also to be taken into
consideration during breeding experiments. The cattle which yield low quantity of milk but
having high percentage of fat content are more prominent than those which yield high
quantity of milk but with low percentage of fat content. Similarly he quality of wool in a
sheep is more important than the quantity of wool.
4. Resistance to diseases
The ability of the animal to resist diseases, to withstand adverse environmental conditions
are also important in the animals produced by hybridisation experiments.
5. Early maturity
It is another trait that the animal breeders look into for improvement of animals. The
earlier, the animals mature to the productive age, lesser is the cost of maintaining them. If
a hen matures early and begins egg production, it is more valuable than that which matures
later.
6. Economy in the use of food
If the amount of food required to produce a certain quantity and quality of animal product is
comparatively higher, the commercial value of such an animal is said to be very low.
In milk yielding cattle and egg yielding fowls if most of their food material is converted into
productivity, such cattle and fowls are considered as more valuable. The above-mentioned
are only few of the desirable qualities that the animal breeders select for improvement of
animals.


Inbreeding

Inbreeding is reproduction from the mating of pairs who are closely related genetically. Inbreeding
results inhomozygosity, which can increase the chances of offspring being affected by recessive or
deleterious traits. This generally leads to a decreased fitness of a population, which is
called inbreeding depression. An individual who results from inbreeding is referred to as inbred. The
avoidance of expression of deleterious recessive alleles caused by inbreeding is thought to be the main
selective force maintaining the outcrossing aspect of sexual reproduction. (See also Inbreeding
depression.)
Livestock breeders often practice controlled breeding to eliminate undesirable characteristics within a
population, which is also coupled with culling of what is considered unfit offspring, especially when
trying to establish a new and desirable trait in the stock.
In plant breeding, inbred lines are used as stocks for the creation of hybrid lines to make use of the
effects of heterosis. Inbreeding in plants also occurs naturally in the form of self-pollination.
Offspring of biologically related persons are subject to the possible impact of inbreeding, such
as congenital birth defects, which is significantly increased due to the closeness of the relationship of
the biological parents (see coefficient of relationship). This is because such pairing increases the
proportion of zygotes that are homozygous, in particular for deleterious recessive alleles that produce
such disorders

(see also inbreeding depression). Because most of such alleles are rare in populations,
it is unlikely that two unrelated marriage partners will both be heterozygous carriers. However,
because close relatives share a large fraction of their alleles, the probability that any such rare
deleterious allele present in the common ancestor will be inherited from both related parents is
increased dramatically with respect to non-inbred couples. Contrary to common belief, inbreeding
does not in itself alter allele frequencies, but rather increases the relative proportion of homozygotes
to heterozygotes. However, because the increased proportion of deleterious homozygotes exposes the
allele to natural selection, in the long run its frequency decreases more rapidly in inbred population. In
the short term, incestuous reproduction is expected to produce increases in spontaneous abortions of
zygotes, perinatal deaths, and postnatal offspring with birth defects.

There may also be other
deleterious effects besides those caused by recessive diseases. Thus, similar immune systems may be
more vulnerable to infectious diseases (see Major histocompatibility complex and sexual selection).

Results
Inbreeding may result in a far higher phenotypic expression of deleterious recessive genes within a
population than would normally be expected. As a result, first-generation inbred individuals are more
likely to show physical and health defects, including:
Reduced fertility both in litter size and sperm viability
Increased genetic disorders
Fluctuating facial asymmetry
Lower birth rate
Higher infant mortality
Slower growth rate
Smaller adult size
Loss of immune system function
Inbreeding can occur just because a small population has been isolated during some time, so that all
breeding individuals became genetically related. It can also occur in a large population if individuals
tend to mate with their relatives, instead of mating at random.
Many individuals in the first generation of inbreeding will never live to reproduce. Over time, with
isolation such as apopulation bottleneck caused by purposeful (assortative) breeding or
natural environmental factors, the deleterious inherited traits are culled.
Island species are often very inbred, as their isolation from the larger group on a mainland allows
natural selection to work upon their population. This type of isolation may result in the formation
of race or even speciation, as the inbreeding first removes many deleterious genes, and allows
expression of genes that allow a population to adapt to an ecosystem. As the adaptation becomes
more pronounced the new species or race radiates from its entrance into the new space, or dies out if
it cannot adapt and, most importantly, reproduce.
The reduced genetic diversity that results from inbreeding may mean a species may not be able to
adapt to changes in environmental conditions. Each individual will have similar immune systems, as
immune systems are genetically based. Where a species becomes endangered, the population may fall
below a minimum whereby the forced interbreeding between the remaining animals will result
in extinction.
Natural breedings include inbreeding by necessity, and most animals only migrate when necessary. In
many cases, the closest available mate is a mother, sister, grandmother, father, brother, or
grandfather. In all cases, the environment presents stresses to remove those individuals who cannot
survive because of illness from the population.
There was an assumption that wild populations do not inbreed; this is not what is observed in some
cases in the wild. However, in species such as horses, animals in wild or feral conditions often drive off
the young of both genders, thought to be a mechanism by which the species instinctively avoids some
of the genetic consequences of inbreeding. In general, many mammal species including humanity's
closest primate relatives avoid close inbreeding possibly due to the deleterious effects.
Examples
Although there are several examples of inbred populations of wild animals, the negative consequences
of this inbreeding are poorly documented.
The cheetah has very low levels of genetic variation, suggesting a population bottleneck (of unknown
cause) and subsequent inbreeding sometime in the past several thousand years. All cheetahs now
come from this small gene pool. Theoretically, their lack of genetic variance could put cheetahs at
greater risk from infectious diseases. One outbreak offeline infectious peritonitis in a captive cheetah
population which was studied over a 5-year period had a morbidity rate of over 90%, and a mortality
rate of 60%. Conversely, inbreeding can purge a population of deleterious alleles, and the cheetah is
known for few genetic illnesses.
In the South American sea lion, there was concern that recent population crashes would reduce
genetic diversity. Historical analysis indicated that a population expansion from just two matrilineal
lines were responsible for most individuals within the population. Even so, the diversity within the lines
allowed great variation in the gene pool that may help to protect the South American sea lion from
extinction.
In lions, prides are often followed by related males in bachelor groups. When the dominant male is
killed or driven off by one of these bachelors, a father may be replaced with his son. There is no
mechanism for preventing inbreeding or to ensure outcrossing. In the prides, most lionesses are
related to one another. If there is more than one dominant male, the group ofalpha males are usually
related. Two lines are then being "line bred". Also, in some populations such as the Crater lions, it is
known that a population bottleneck has occurred. Researchers found far greater
genetic heterozygosity than expected. In fact, predators are known for low genetic variance, along
with most of the top portion of the tropic levels of anecosystem. Additionally, the alpha males of two
neighboring prides can potentially be from the same litter; one brother may come to acquire
leadership over another's pride, and subsequently mate with his 'nieces' or cousins. However, killing
another male's cubs, upon the takeover, allows the new selected gene complement of the incoming
alpha male to prevail over the previous male. There are genetic assays being scheduled for lions to
determine their genetic diversity. The preliminary studies show results inconsistent with the
outcrossing paradigm based on individual environments of the studied groups.
In Central California, the Sea Otters were thought to have been driven to extinction due to over
hunting, until a colony of about 30 breeding pairs was discovered in the Big Sur region in the 1930s.
Since then the population has grown and spread along the central Californian coast to around 2000
individuals, a level that has remained stable for over a decade. Population growth is limited by the fact
that all Californian Sea Otters are descended from the isolated colony resulting in inbreeding.

Domestic animals
Breeding in domestic animals is assortative breeding primarily (see selective breeding). Without the
sorting of individuals by trait, a breed could not be established, nor could poor genetic material be
removed. Homozygosity is the case where similar or identical alleles combine to express a trait that
is not otherwise expressed (recessiveness). Inbreeding, through homozygosity, exposes recessive
alleles. Inbreeding is used to reveal deleterious recessive alleles, which can then be eliminated
through assortative breeding or through culling.
Inbreeding is used by breeders of domestic animals to fix desirable genetic traits within a population
or to attempt to remove deleterious traits by allowing them to manifest phenotypically from the
genotypes. Inbreeding is defined as the use of close relations for breeding such as mother to son,
father to daughter, brother to sister.
Breeders must cull unfit breeding suppressed individuals and/or individuals who demonstrate either
homozygosity or heterozygosity for genetic based diseases. The issue of casual breeders who
inbreed irresponsibly is discussed in the following quotation on cattle:
Meanwhile, milk production per cow per lactation increased from 17,444 lbs to 25,013 lbs from 1978
to 1998 for the Holstein breed. Mean breeding values for milk of Holstein cows increased by 4,829
lbs during this period. High producing cows are increasingly difficult to breed and are subject to
higher health costs than cows of lower genetic merit for production (Cassell, 2001).
Intensive selection for higher yield has increased relationships among animals within breed and
increased the rate of casual inbreeding.
Many of the traits that affect profitability in crosses of modern dairy breeds have not been studied in
designed experiments. Indeed, all crossbreeding research involving North American breeds and
strains is very dated (McAllister, 2001) if it exists at all.
Linebreeding is a form of inbreeding. There is no clear distinction between the two terms, but
linebreeding may encompass crosses between individuals and their descendants or two
cousins. This method can be used to increase a particular animal's contribution to the
population. While linebreeding is less likely to cause problems in the first generation than does
inbreeding, over time, linebreeding can reduce the genetic diversity of a population and cause
problems related to a too-small genepool that may include an increased prevalence of genetic
disorders and inbreeding depression.
Outcrossing is where two unrelated individuals have been crossed to produce progeny. In
outcrossing, unless there is verifiable genetic information, one may find that all individuals are
distantly related to an ancient progenitor. If the trait carries throughout a population, all individuals
can have this trait. This is called the founder effect. In the well-established breeds, that are
commonly bred, a large gene pool is present. For example, in 2004, over 18,000 Persian cats were
registered. A possibility exists for a complete outcross, if no barriers exist between the individuals to
breed. However it is not always the case, and a form of distant linebreeding occurs. Again it is up to
the assortative breeder to know what sort of traits both positive and negative exist within the diversity
of one breeding. This diversity of genetic expression, within even close relatives, increases the
variability and diversity of viable stock.
In the registered dog population, the onset of large numbers of casual breeders has corresponded
with an increase in the number of genetic illnesses of dogs by not understanding how, why and
which traits are inherited. The dog sites indicate that the largest percentage of dog breeders in the
US are casual breeders. Therefore the investment in a papered animal, with an expected short term
profit, motivates some to ignore the practice of culling. Casual breeders in companion animals often
ignore breeding restrictions within their contracts with source companion animal breeders. The
casual breeders breed the very culls that a genetics based breeder has released as a pet. The
casual breeder was also cited in the quotes above on cattle raising.


Crossbreed
A crossbreed or crossbred usually refers to an animal with purebred parents of two different breeds,
varieties, or populations. Crossbreeding, sometimes called "designer crossbreeding", refers to the
process of breeding such an animal, often with the intention to create offspring that share the traits of
both parent lineages, or producing an animal with hybrid vigor. While crossbreeding is used to
maintain health and viability of animals, irresponsible crossbreeding can also produce animals of
inferior quality or dilute a purebred gene pool to the point of extinction of a given breed of animal.
[

The term is also used at times to refer to a domestic animal of unknown ancestry where the breed
status of only one parent or grandparent is known, though the term "mixed breed" is technically more
accurate. The term outcross is used to describe a type of crossbreeding used within a purebred breed
to increase the genetic diversity within the breed, particularly when there is a need to
avoid inbreeding.
In animal breeding, crossbreed describes crosses within a single species, while hybrid refers to crosses
between different species. In plant breeding terminology, the term crossbreed is uncommon. No
universal term is used to distinguish hybridization or crossing within a population from those between
populations, or even those between species.


Crossbreeds in specific animals
Cattle
In cattle, there are systems of crossbreeding. One is used when the purebred females are particularly
adapted to a specific environment, and are crossed with purebred bulls from another environment to
produce a generation having traits of both parents.
[1]

Sheep
The large number of breeds of sheep, which vary greatly, creates an opportunity for crossbreeding to
be used to tailor production of lambs to the goal of the individual stockman.
[2]

Llamas
Results of crossbreeding classic and woolly breeds of llama are unpredictable. The resulting offspring
displays physical characteristics of either parent, or a mix of characteristics from both, periodically
producing a fleeced llama. The results are increasingly unpredictable when both parents are
crossbreeds, with possibility of the offspring displaying characteristics of a grandparent, not obvious in
either parent.
[3]

Dogs
A crossbred or hybrid dog is a cross between two (sometimes more) known breeds, and is usually
distinguished from amixed-breed dog, which has ancestry from many sources, some of which may not
be known. Crossbreeds are popular, due to the belief that they have increased hybrid vigor without
loss of attractiveness of the dog. Certain planned crossbreeding between purebred dogs of different
breeds can produce puppies worth more than their purebred parents, due to a high demand.
[citation
needed]

Horses
Crossbreeding in horses is often done with the intent of ultimately creating a new breed of horse. One
type of modern crossbreeding in horses is used to create many of the warmblood breeds. Warmbloods
are a type of horse used in the sport horsedisciplines, usually registered in an open stud book by
a studbook selectionprocedure that evaluates conformation, pedigree and, in some animals, a training
or performance standard. Most warmblood breeds began as a cross of draft horsebreeds
on Thoroughbreds, but have, in some cases, developed over the past century to the point where they
are considered to be a true-breeding population and have a closed stud book. Other types of
recognized crossbreeding include that within the American Quarter Horse, which will register horses
with one Thoroughbred parent and one registered Quarter Horse parent in the "Appendix" registry,
and allow such animals full breed registration status as Quarter Horses if they meet a certain
performance standard. Another well-known crossbred horse is the Anglo-Arabian, which may be
produced by a purebred Arabian horse crossed on a Thoroughbred, or by various crosses of Anglo-
Arabians with other Anglo-Arabians, as long as the ensuing animal never has more than 75% or less
than 25% of each breed represented in its pedigree.

Hybrid animals
A hybrid animal is one with parentage of two separate species, differentiating it from crossbred
animals, which have parentage of the same species. Hybrids are usually, but not always, sterile.
One of the most ancient types of hybrid animal is the mule, a cross between a female horse and a
male donkey or ass. Theliger is a hybrid cross between a male lion and female tiger. The yattle is a
cross between a cow and a yak. Other crosses include the tigon (between a female lion and male
tiger) and yakalo (between a yak and buffalo). The Incas recognized that hybrids of Lama glama
(llama) and Lama pacos (alpaca) resulted in a hybrid with none of the advantages of either parent.
[5]

At one time it was thought that dogs and wolves were separate species, and the crosses between
dogs and wolves were called wolf hybrids. Today wolves and dogs are both recognized as Canis
lupus, but the old term "wolf hybrid" is still used.
Mixed breeds
A mixed-breed animal is defined as having undocumented or unknown parentage, while a
crossbreed generally has known, usually purebred parents of two distinct breeds or varieties. A dog
of unknown parentage is often called a mixed-breed dog, "mutt" or "mongrel." A cat of unknown
parentage is often referred to as domestic short-haired or domestic long-haired cat generically, and
in some dialects is often called a "moggy". A horse of unknown bloodlines is a grade horse.

Grading-up
It is a type of breeding system used to bring about radical improvement in unproductive breeds.
Simple grading-up involves mating animals of two breeds (one to be improved and the other to do the
improving). The resulting hybrids are mated for several generations with males of the improving breed
until the desired type of animal is obtained. Highly productive hybrids of the fourth, fifth, and sixth
generations that attain the desired traits of the improving breed are inbred, sometimes resulting in the
creation of a new breed. Grading-up involving several improving breeds is the quickest and most
efficient method of radically improving unproductive livestock and transforming breeds (for example,
changing coarse-wooled breeds of sheep into fine-wooled and semifine-wooled breeds).
The rate at which a breed can be transformed and improved depends on the extent of hereditary
differences between the animals of the breeds being crossed, the degree of hereditary stability of the
breeds, the care taken in selecting and matching the hybrids, and the feeding and maintenance
conditions of the young hybrids. Grading-up is used to breed almost all species of agricultural animals.
In the USSR it was widely used from 1925 to 1950 to increase the breed purity and productivity of
commercial and breeding herds.