Heterosis, also known as hybrid vigor, occurs when the offspring of two parents from genetically distinct breeds or varieties exhibit superior traits compared to their parents. This can be due to either dominance or overdominance of alleles. Two main hypotheses for the genetic basis of heterosis are the dominance hypothesis, where undesirable recessive alleles are masked in hybrids, and the overdominance hypothesis, where certain allele combinations provide advantages. Recent research also indicates an epigenetic component to heterosis involving microRNAs and histone modification. While hybrids may show enhanced traits, not all crosses exhibit heterosis, and superiority is difficult to define scientifically. The term "genetic superiority" is controversial and more objective measurable traits are used.
Heterosis, also known as hybrid vigor, occurs when the offspring of two parents from genetically distinct breeds or varieties exhibit superior traits compared to their parents. This can be due to either dominance or overdominance of alleles. Two main hypotheses for the genetic basis of heterosis are the dominance hypothesis, where undesirable recessive alleles are masked in hybrids, and the overdominance hypothesis, where certain allele combinations provide advantages. Recent research also indicates an epigenetic component to heterosis involving microRNAs and histone modification. While hybrids may show enhanced traits, not all crosses exhibit heterosis, and superiority is difficult to define scientifically. The term "genetic superiority" is controversial and more objective measurable traits are used.
Heterosis, also known as hybrid vigor, occurs when the offspring of two parents from genetically distinct breeds or varieties exhibit superior traits compared to their parents. This can be due to either dominance or overdominance of alleles. Two main hypotheses for the genetic basis of heterosis are the dominance hypothesis, where undesirable recessive alleles are masked in hybrids, and the overdominance hypothesis, where certain allele combinations provide advantages. Recent research also indicates an epigenetic component to heterosis involving microRNAs and histone modification. While hybrids may show enhanced traits, not all crosses exhibit heterosis, and superiority is difficult to define scientifically. The term "genetic superiority" is controversial and more objective measurable traits are used.
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.