Inbreeding

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Common fruit fly females prefer to mate with their own brothers over unrelated males. Fruit flies.jpg
Common fruit fly females prefer to mate with their own brothers over unrelated males.

Inbreed is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. [2] By analogy, the term is used in human reproduction, but more commonly refers to the genetic disorders and other consequences that may arise from expression of deleterious or recessive traits resulting from incestuous sexual relationships and consanguinity.

Contents

Inbreeding results in homozygosity, which can increase the chances of offspring being affected by deleterious or recessive traits. [3] This usually leads to at least temporarily decreased biological fitness of a population [4] [5] (called inbreeding depression), which is its ability to survive and reproduce. An individual who inherits such deleterious traits is colloquially referred to as inbred. The avoidance of expression of such deleterious recessive alleles caused by inbreeding, via inbreeding avoidance mechanisms, is the main selective reason for outcrossing. [6] [7] Crossbreeding between populations also often has positive effects on fitness-related traits, [8] but also sometimes leads to negative effects known as outbreeding depression. However increased homozygosity increases probability of fixing beneficial alleles and also slightly decreases probability of fixing deleterious alleles in population. [9] Inbreeding can result in purging of deleterious alleles from a population through purifying selection. [10] [11] [12]

Inbreeding is a technique used in selective breeding. For example, in livestock breeding, breeders may use inbreeding when trying to establish a new and desirable trait in the stock and for producing distinct families within a breed, but will need to watch for undesirable characteristics in offspring, which can then be eliminated through further selective breeding or culling. Inbreeding also helps to ascertain the type of gene action affecting a trait. Inbreeding is also used to reveal deleterious recessive alleles, which can then be eliminated through assortative breeding or through culling. 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.

Inbreeding can significantly influence gene expression which can prevent inbreeding depression. [13]

Overview

Offspring of biologically related persons are subject to the possible effects of inbreeding, such as congenital birth defects. The chances of such disorders are increased when the biological parents are more closely related. This is because such pairings have a 25% probability of producing homozygous zygotes, resulting in offspring with two recessive alleles, which can produce disorders when these alleles are deleterious. [14] Because most recessive alleles are rare in populations, it is unlikely that two unrelated marriage partners will both be carriers of the same deleterious allele; however, because close relatives share a large fraction of their alleles, the probability that any such deleterious allele is inherited from the common ancestor through both parents is increased dramatically. For each homozygous recessive individual formed there is an equal chance of producing a homozygous dominant individual — one completely devoid of the harmful allele. 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 populations. In the short term, incestuous reproduction is expected to increase the number of spontaneous abortions of zygotes, perinatal deaths, and postnatal offspring with birth defects. [15] The advantages of inbreeding may be the result of a tendency to preserve the structures of alleles interacting at different loci that have been adapted together by a common selective history. [16]

Malformations or harmful traits can stay within a population due to a high homozygosity rate, and this will cause a population to become fixed for certain traits, like having too many bones in an area, like the vertebral column of wolves on Isle Royale or having cranial abnormalities, such as in Northern elephant seals, where their cranial bone length in the lower mandibular tooth row has changed. Having a high homozygosity rate is problematic for a population because it will unmask recessive deleterious alleles generated by mutations, reduce heterozygote advantage, and it is detrimental to the survival of small, endangered animal populations. [17] When deleterious recessive alleles are unmasked due to the increased homozygosity generated by inbreeding, this can cause inbreeding depression. [18]

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). [19]

Inbreeding history of the population should also be considered when discussing the variation in the severity of inbreeding depression between and within species. With persistent inbreeding, there is evidence that shows that inbreeding depression becomes less severe. This is associated with the unmasking and elimination of severely deleterious recessive alleles. However, inbreeding depression is not a temporary phenomenon because this elimination of deleterious recessive alleles will never be complete. Eliminating slightly deleterious mutations through inbreeding under moderate selection is not as effective. Fixation of alleles most likely occurs through Muller's ratchet, when an asexual population's genome accumulates deleterious mutations that are irreversible. [20]

Despite all its disadvantages, inbreeding can also have a variety of advantages, such as reducing the recombination load, [21] and allowing the expression of recessive advantageous phenotypes. It has been proposed that under circumstances when the advantages of inbreeding outweigh the disadvantages, preferential breeding within small groups could be promoted, potentially leading to speciation. [22]

Genetic disorders

Animation of uniparental isodisomy

Autosomal recessive disorders occur in individuals who have two copies of an allele for a particular recessive genetic mutation. [23] Except in certain rare circumstances, such as new mutations or uniparental disomy, both parents of an individual with such a disorder will be carriers of the gene. These carriers do not display any signs of the mutation and may be unaware that they carry the mutated gene. Since relatives share a higher proportion of their genes than do unrelated people, it is more likely that related parents will both be carriers of the same recessive allele, and therefore their children are at a higher risk of inheriting an autosomal recessive genetic disorder. The extent to which the risk increases depends on the degree of genetic relationship between the parents; the risk is greater when the parents are close relatives and lower for relationships between more distant relatives, such as second cousins, though still greater than for the general population. [24]

Children of parent-child or sibling-sibling unions are at an increased risk compared to cousin-cousin unions. [25] :3 Inbreeding may result in a greater than expected phenotypic expression of deleterious recessive alleles within a population. [26] As a result, first-generation inbred individuals are more likely to show physical and health defects, [27] [28] including:

The isolation of a small population for a period of time can lead to inbreeding within that population, resulting in increased genetic relatedness between breeding individuals. Inbreeding depression can also occur in a large population if individuals tend to mate with their relatives, instead of mating randomly.

Many individuals in the first generation of inbreeding will never live to reproduce. [31] Over time, with isolation, such as a population bottleneck caused by purposeful (assortative) breeding or natural environmental factors, the deleterious inherited traits are culled. [6] [7] [32]

Island species are often very inbred, as their isolation from the larger group on a mainland allows natural selection to work on 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 permits the 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. [33]

The reduced genetic diversity, for example due to a bottleneck will unavoidably increase inbreeding for the entire population. This may mean that 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. When 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 from the population those individuals who cannot survive because of illness.[ citation needed ]

There was an assumption[ by whom? ] 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 sexes, thought to be a mechanism by which the species instinctively avoids some of the genetic consequences of inbreeding. [34] In general, many mammal species, including humanity's closest primate relatives, avoid close inbreeding possibly due to the deleterious effects. [25] :6

Examples

Although there are several examples of inbred populations of wild animals, the negative consequences of this inbreeding are poorly documented.[ citation needed ] 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 was responsible for most of the 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. [35]

Heterozygous Heterozygous.jpg
Heterozygous

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 by 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 of alpha 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. [36] In fact, predators are known for low genetic variance, along with most of the top portion of the trophic levels of an ecosystem. [37] Additionally, the alpha males of two neighboring prides can 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. [36]

In Central California, sea otters were thought to have been driven to extinction due to over hunting, until a small colony was discovered in the Point Sur region in the 1930s. [38] Since then, the population has grown and spread along the central Californian coast to around 2,000 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. [39]

Cheetahs are another example of inbreeding. Thousands of years ago the cheetah went through a population bottleneck that reduced its population dramatically so the animals that are alive today are all related to one another. A consequence from inbreeding for this species has been high juvenile mortality, low fecundity, and poor breeding success. [40]

In a study on an island population of song sparrows, individuals that were inbred showed significantly lower survival rates than outbred individuals during a severe winter weather related population crash. These studies show that inbreeding depression and ecological factors have an influence on survival. [20]

Measures

A measure of inbreeding of an individual A is the probability F(A) that both alleles in one locus are derived from the same allele in an ancestor. These two identical alleles that are both derived from a common ancestor are said to be identical by descent. This probability F(A) is called the "coefficient of inbreeding". [41]

Another useful measure that describes the extent to which two individuals are related (say individuals A and B) is their coancestry coefficient f(A,B), which gives the probability that one randomly selected allele from A and another randomly selected allele from B are identical by descent. [42] This is also denoted as the kinship coefficient between A and B. [43]

A particular case is the self-coancestry of individual A with itself, f(A,A), which is the probability that taking one random allele from A and then, independently and with replacement, another random allele also from A, both are identical by descent. Since they can be identical by descent by sampling the same allele or by sampling both alleles that happen to be identical by descent, we have f(A,A) = 1/2 + F(A)/2. [44]

Both the inbreeding and the coancestry coefficients can be defined for specific individuals or as average population values. They can be computed from genealogies or estimated from the population size and its breeding properties, but all methods assume no selection and are limited to neutral alleles.

There are several methods to compute this percentage. The two main ways are the path method [45] [41] and the tabular method. [46] [47]

Typical coancestries between relatives are as follows:

Animals

Wild animals

Few studies have found evidence of regular incest in mammals, but banded mongooses are an exception. Mangoustes rayees - Banded Mongooses.jpg
Few studies have found evidence of regular incest in mammals, but banded mongooses are an exception.

Semi-domestic animals

White tiger in Gunma Safari Park White tiger-Gunma Safari Park.jpg
White tiger in Gunma Safari Park

Domestic animals

Hereditary polycystic kidney disease is prevalent in the Persian cat breed, affecting almost half the population in some countries. Shaded silver Persian Cat Missionhill Cosmic Rainstorm.jpg
Hereditary polycystic kidney disease is prevalent in the Persian cat breed, affecting almost half the population in some countries.
An intensive form of inbreeding where an individual S is mated to his daughter D1, granddaughter D2 and so on, in order to maximise the percentage of S's genes in the offspring. 87.5% of D3's genes would come from S, while D4 would receive 93.75% of their genes from S. Intense inbreeding - Continuous sire to daughter mating.svg
An intensive form of inbreeding where an individual S is mated to his daughter D1, granddaughter D2 and so on, in order to maximise the percentage of S's genes in the offspring. 87.5% of D3's genes would come from S, while D4 would receive 93.75% of their genes from S.

Breeding in domestic animals is primarily assortative breeding (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 exposes recessive alleles through increasing homozygosity. [55]

Breeders must avoid breeding from individuals that demonstrate either homozygosity or heterozygosity for disease causing alleles. [56] The goal of preventing the transfer of deleterious alleles may be achieved by reproductive isolation, sterilization, or, in the extreme case, culling. Culling is not strictly necessary if genetics are the only issue in hand. Small animals such as cats and dogs may be sterilized, but in the case of large agricultural animals, such as cattle, culling is usually the only economic option.

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. [57] 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. [58]

The BBC produced two documentaries on dog inbreeding titled Pedigree Dogs Exposed and Pedigree Dogs Exposed: Three Years On that document the negative health consequences of excessive inbreeding.

Linebreeding

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. [54] [59] This method can be used to increase a particular animal's contribution to the population. [54] 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 gene pool that may include an increased prevalence of genetic disorders and inbreeding depression.[ citation needed ]

Outcrossing

Outcrossing is where two unrelated individuals are 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. [60] 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.

Laboratory animals

Systematic inbreeding and maintenance of inbred strains of laboratory mice and rats is of great importance for biomedical research. The inbreeding guarantees a consistent and uniform animal model for experimental purposes and enables genetic studies in congenic and knock-out animals. In order to achieve a mouse strain that is considered inbred, a minimum of 20 sequential generations of sibling matings must occur. With each successive generation of breeding, homozygosity in the entire genome increases, eliminating heterozygous loci. With 20 generations of sibling matings, homozygosity is occurring at roughly 98.7% of all loci in the genome, allowing for these offspring to serve as animal models for genetic studies. [61] The use of inbred strains is also important for genetic studies in animal models, for example to distinguish genetic from environmental effects. The mice that are inbred typically show considerably lower survival rates.

Humans

Global prevalence of consanguinity Global prevalence of consanguinity.svg
Global prevalence of consanguinity

Effects

Inbreeding increases homozygosity, which can increase the chances of the expression of deleterious recessive alleles and therefore has the potential to decrease the fitness of the offspring. With continuous inbreeding, genetic variation is lost and homozygosity is increased, enabling the expression of recessive deleterious alleles in homozygotes. The coefficient of inbreeding, or the degree of inbreeding in an individual, is an estimate of the percent of homozygous alleles in the overall genome. [63] The more biologically related the parents are, the greater the coefficient of inbreeding, since their genomes have many similarities already. This overall homozygosity becomes an issue when there are deleterious recessive alleles in the gene pool of the family. [64] By pairing chromosomes of similar genomes, the chance for these recessive alleles to pair and become homozygous greatly increases, leading to offspring with autosomal recessive disorders. [64]

Inbreeding is especially problematic in small populations where the genetic variation is already limited. [65] By inbreeding, individuals are further decreasing genetic variation by increasing homozygosity in the genomes of their offspring. [66] Thus, the likelihood of deleterious recessive alleles to pair is significantly higher in a small inbreeding population than in a larger inbreeding population. [65]

The fitness consequences of consanguineous mating have been studied since their scientific recognition by Charles Darwin in 1839. [67] [68] Some of the most harmful effects known from such breeding includes its effects on the mortality rate as well as on the general health of the offspring. [69] Since the 1960s, there have been many studies to support such debilitating effects on the human organism. [66] [67] [69] [70] [71] Specifically, inbreeding has been found to decrease fertility as a direct result of increasing homozygosity of deleterious recessive alleles. [71] [72] Fetuses produced by inbreeding also face a greater risk of spontaneous abortions due to inherent complications in development. [73] Among mothers who experience stillbirths and early infant deaths, those that are inbreeding have a significantly higher chance of reaching repeated results with future offspring. [74] Additionally, consanguineous parents possess a high risk of premature birth and producing underweight and undersized infants. [75] Viable inbred offspring are also likely to be inflicted with physical deformities and genetically inherited diseases. [63] Studies have confirmed an increase in several genetic disorders due to inbreeding such as blindness, hearing loss, neonatal diabetes, limb malformations, disorders of sex development, schizophrenia and several others. [63] [76] Moreover, there is an increased risk for congenital heart disease depending on the inbreeding coefficient (See coefficient of inbreeding) of the offspring, with significant risk accompanied by an F =.125 or higher. [27]

Prevalence

The general negative outlook and eschewal of inbreeding that is prevalent in the Western world today has roots from over 2000 years ago. Specifically, written documents such as the Bible illustrate that there have been laws and social customs that have called for the abstention from inbreeding. Along with cultural taboos, parental education and awareness of inbreeding consequences have played large roles in minimizing inbreeding frequencies in areas like Europe. That being so, there are less urbanized and less populated regions across the world that have shown continuity in the practice of inbreeding.

The continuity of inbreeding is often either by choice or unavoidably due to the limitations of the geographical area. When by choice, the rate of consanguinity is highly dependent on religion and culture. [65] In the Western world some Anabaptist groups are highly inbred because they originate from small founder populations and until[ clarification needed ] today[ when? ] marriage outside the groups is not allowed for members.[ citation needed ] Especially the Reidenbach Old Order Mennonites [77] and the Hutterites stem from very small founder populations. The same is true for some Hasidic and Haredi Jewish groups.

Of the practicing regions, Middle Eastern and northern Africa territories show the greatest frequencies of consanguinity. [65] The link between the high frequency and the region is primarily due to the dominance of Islamic populations, who have historically engaged in familyline relations. [68] However, inbreeding culture in Middle East didn't begin with Islam, having roots in ancient Egypt and Mesopotamia.

Among these populations with high levels of inbreeding, researchers have found several disorders prevalent among inbred offspring. In Lebanon, Saudi Arabia, Egypt, and in Israel, the offspring of consanguineous relationships have an increased risk of congenital malformations, congenital heart defects, congenital hydrocephalus and neural tube defects. [65] Furthermore, among inbred children in Palestine and Lebanon, there is a positive association between consanguinity and reported cleft lip/palate cases. [65] Historically, populations of Qatar have engaged in consanguineous relationships of all kinds, leading to high risk of inheriting genetic diseases. As of 2014, around 5% of the Qatari population suffered from hereditary hearing loss; most were descendants of a consanguineous relationship. [78]

Royalty and nobility

Tutankhamun (c. 1341-1323 BC) (receiving flowers from Ankhesenamun). Tutankhamun, who was a descendant of one or possibly two generations of sibling or cousin parents, had two children with his own half-sister, none of which survived infancy. Anuk.PNG
Tutankhamun (c. 1341–1323 BC) (receiving flowers from Ankhesenamun). Tutankhamun, who was a descendant of one or possibly two generations of sibling or cousin parents, had two children with his own half-sister, none of which survived infancy.
Charles II of Spain. He is best remembered for his physical disabilities, believed to be the result of inbreeding. Juan de Miranda Carreno 002.jpg
Charles II of Spain. He is best remembered for his physical disabilities, believed to be the result of inbreeding.

Inter-nobility marriage was used as a method of forming political alliances among elites[ citation needed ]. These ties were often sealed only upon the birth of progeny within the arranged marriage. Thus marriage was seen as a union of lines of nobility, not as a contract between individuals as it is seen[ by whom? ] today.

Royal intermarriage was often practiced among European royal families, usually for interests of state. Over time, due to the relatively limited number of potential consorts, the gene pool of many ruling families grew progressively smaller, until all European royalty was related. This also resulted in many being descended from a certain person through many lines of descent, such as the numerous European royalty and nobility descended from the British Queen Victoria or King Christian IX of Denmark. [79] The House of Habsburg was infamous for its inbreeding, with the Habsburg lip cited as an ill-effect, although no genetic evidence has proved the claim. The closely related houses of Habsburg, Bourbon, Braganza and Wittelsbach also frequently engaged in first-cousin unions as well as the occasional double-cousin and uncle–niece marriages. Examples of incestuous marriages and the impact of inbreeding on royal families include:

See also

Related Research Articles

Small populations can behave differently from larger populations. They are often the result of population bottlenecks from larger populations, leading to loss of heterozygosity and reduced genetic diversity and loss or fixation of alleles and shifts in allele frequencies. A small population is then more susceptible to demographic and genetic stochastic events, which can impact the long-term survival of the population. Therefore, small populations are often considered at risk of endangerment or extinction, and are often of conservation concern.

Inbred strains are individuals of a particular species which are nearly identical to each other in genotype due to long inbreeding. A strain is inbred when it has undergone at least 20 generations of brother x sister or offspring x parent mating, at which point at least 98.6% of the loci in an individual of the strain will be homozygous, and each individual can be treated effectively as clones. Some inbred strains have been bred for over 150 generations, leaving individuals in the population to be isogenic in nature. Inbred strains of animals are frequently used in laboratories for experiments where for the reproducibility of conclusions all the test animals should be as similar as possible. However, for some experiments, genetic diversity in the test population may be desired. Thus outbred strains of most laboratory animals are also available, where an outbred strain is a strain of an organism that is effectively wildtype in nature, where there is as little inbreeding as possible.

Consanguinity Property of being from the same kinship as another person

Consanguinity is the property of being from the same kinship as another person. In that aspect, consanguinity is the quality of being descended from the same ancestor as another person.

The coefficient of relationship is a measure of the degree of consanguinity between two individuals. The term coefficient of relationship was defined by Sewall Wright in 1922, and was derived from his definition of the coefficient of inbreeding of 1921. The measure is most commonly used in genetics and genealogy. A coefficient of inbreeding can be calculated for an individual, and is typically one-half the coefficient of relationship between the parents.

Heterosis, hybrid vigor, or outbreeding enhancement is the improved or increased function of any biological quality in a hybrid offspring. An offspring is heterotic 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.

An F1 Hybrid (also known as filial 1 hybrid) is the first filial generation of offspring of distinctly different parental types. F1 hybrids are used in genetics, and in selective breeding, where it may appear as F1 crossbreed. The term is sometimes written with a subscript, as F1 hybrid. Subsequent generations are called F2, F3, etc.

Conservation genetics

Conservation genetics is an interdisciplinary subfield of population genetics that aims to understand the dynamics of genes in populations principally to avoid extinction. Therefore, it applies genetic methods to the conservation and restoration of biodiversity. Researchers involved in conservation genetics come from a variety of fields including population genetics, molecular ecology, biology, evolutionary biology, and systematics. Genetic diversity is one of the three fundamental levels of biodiversity, so it is directly important in conservation. Genetic variability influences both the health and long-term survival of populations because decreased genetic diversity has been associated with reduced fitness, such as high juvenile mortality, diminished population growth, reduced immunity, and ultimately, higher extinction risk.

Genetic load is the difference between the fitness of an average genotype in a population and the fitness of some reference genotype, which may be either the best present in a population, or may be the theoretically optimal genotype. The average individual taken from a population with a low genetic load will generally, when grown in the same conditions, have more surviving offspring than the average individual from a population with a high genetic load. Genetic load can also be seen as reduced fitness at the population level compared to what the population would have if all individuals had the reference high-fitness genotype. High genetic load may put a population in danger of extinction.

Molecular ecology A field of evolutionary biology that applies molecular population genetics, molecular phylogenetics, and genomics to traditional ecological questions

Molecular ecology is a field of evolutionary biology that is concerned with applying molecular population genetics, molecular phylogenetics, and more recently genomics to traditional ecological questions. It is virtually synonymous with the field of "Ecological Genetics" as pioneered by Theodosius Dobzhansky, E. B. Ford, Godfrey M. Hewitt, and others. These fields are united in their attempt to study genetic-based questions "out in the field" as opposed to the laboratory. Molecular ecology is related to the field of conservation genetics.

Inbreeding depression is the reduced biological fitness in a given population as a result of inbreeding, or breeding of related individuals. Population biological fitness refers to an organism's ability to survive and perpetuate its genetic material. Inbreeding depression is often the result of a population bottleneck. In general, the higher the genetic variation or gene pool within a breeding population, the less likely it is to suffer from inbreeding depression.

Thelytoky

Thelytoky is a type of parthenogenesis in which females are produced from unfertilized eggs, as for example in aphids. Thelytokous parthenogenesis is rare among animals and reported in about 1,500 species, about 1 in 1000 of described animal species, according to a 1984 study. It is more common in invertebrates, like arthropods, but it can occur in vertebrates, including salamanders, fish, and reptiles such as some whiptail lizards.

Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing unrelated genetic material into a breeding line. It increases genetic diversity, thus reducing the probability of an individual being subject to disease or genetic abnormalities.

In natural selection, negative selection or purifying selection is the selective removal of alleles that are deleterious. This can result in stabilising selection through the purging of deleterious genetic polymorphisms that arise through random mutations.

Edward Murray East was an American plant geneticist, botanist, agronomist and eugenicist. He is known for his experiments that led to the development of hybrid corn and his support of 'forced' elimination of the 'unfit' based on eugenic findings. He worked at the Bussey Institute of Harvard University where he performed a key experiment showing the outcome of crosses between lines that differ in a quantitative trait.

Popular sire effect

The popular sire effect occurs when an animal with desirable attributes is bred repeatedly. In dog breeding, a male dog that wins respected competitions becomes highly sought after, as breeders believe the sire possesses the genes necessary to produce champions. The popular sire is often bred extensively with many females. This can cause undetected, undesirable genetic traits in the stud to spread rapidly within the gene pool. It can also reduce genetic diversity by the exclusion of other males.

Consanguine marriage is marriage between individuals who are closely related. Though it may involve incest, it implies more than the sexual nature of incest. In a clinical sense, marriage between two family members who are second cousins or closer qualifies as consanguineous marriage. This is based on the gene copies their offspring may receive. Though these unions are still prevalent in some communities, as seen across the Greater Middle East region, many other populations have seen a great decline in intra-family marriages.

Genetic purging is the reduction of the frequency of a deleterious allele, caused by an increased efficiency of natural selection prompted by inbreeding.

The partial dominance hypothesis in genetics states that inbreeding depression is the result of the frequency increase of homozygous deleterious recessive or partially recessive alleles. The partial dominance hypothesis can be explained by looking at a population that is divided into a large number of separately inbred lines. Deleterious alleles will eventually be eliminated from some lines and become fixed in other lines, while some lines disappear because of fixation of deleterious alleles. This will cause an overall decline in population and trait value, but then increase to a trait value that is equal to or greater than the trait value in the original population. Crossing inbred lines restores fitness in the overdominance hypothesis and a fitness increase in the partial dominance hypothesis.

Inbreeding avoidance, or the inbreeding avoidance hypothesis, is a concept in evolutionary biology that refers to the prevention of the deleterious effects of inbreeding. The inbreeding avoidance hypothesis posits that certain mechanisms develop within a species, or within a given population of a species, as a result of assortative mating, natural and sexual selection in order to prevent breeding among related individuals in that species or population. Although inbreeding may impose certain evolutionary costs, inbreeding avoidance, which limits the number of potential mates for a given individual, can inflict opportunity costs. Therefore, a balance exists between inbreeding and inbreeding avoidance. This balance determines whether inbreeding mechanisms develop and the specific nature of said mechanisms.

Exposure of zebrafish to a chemical environmental agent, analogous to that caused by anthropogenic pollution, amplified the effects of inbreeding on key reproductive traits. Embryo viability was significantly reduced in inbred exposed fish and there was a tendency for inbred males to sire fewer offspring.

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