Inbreeding - Knowledge

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found to decrease fertility as a direct result of increasing homozygosity of deleterious recessive alleles. Fetuses produced by inbreeding also face a greater risk of spontaneous abortions due to inherent complications in development. 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. Additionally, consanguineous parents possess a high risk of premature birth and producing underweight and undersized infants. Viable inbred offspring are also likely to be inflicted with physical deformities and genetically inherited diseases. Studies have confirmed an increase in several genetic disorders due to inbreeding such as blindness, hearing loss, neonatal diabetes, limb malformations,

199: 316:, 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. 980:, or the degree of inbreeding in an individual, is an estimate of the percent of homozygous alleles in the overall genome. 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. 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. However, these deleterious effects are common for very close relatives but not for those related on the 3rd cousin or greater level, who exhibit increased fitness. 799: 915:. 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. 783: 710: 963: 227:

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

767: 47: 243:, 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. When deleterious recessive alleles are unmasked due to the increased homozygosity generated by inbreeding, this can cause inbreeding depression. 4976: 231:, 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. 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. 288: 497: 493:, 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. 936:

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. 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.

525:. 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 258:

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

758:: The single male offspring mite mates with all the daughters when they are still in the mother. The females, now impregnated, cut holes in their mother's body so that they can emerge. The male emerges as well, but does not look for food or new mates, and dies after a few hours. The females die at the age of 4 days, when their own offspring 2912: 991:

in 1839. 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. Since the 1960s, there have been many studies to support such debilitating effects on the human organism. Specifically, inbreeding has been

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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.

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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

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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

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Inbreeding is especially problematic in small populations where the genetic variation is already limited. By inbreeding, individuals are further decreasing genetic variation by increasing homozygosity in the genomes of their offspring. Thus, the likelihood of deleterious recessive alleles to pair is

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Inbreeding increases homozygosity, which can increase the chances of the expression of deleterious or beneficial recessive alleles and therefore has the potential to either decrease or increase the fitness of the offspring. Depending on the rate of inbreeding, natural selection may still be able to

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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

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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

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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

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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

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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. This is

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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. 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

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mating system depend on the ability to produce sons to mate with as a means of ensuring a mate can be found if no other male is available. It has been proposed that under circumstances when the advantages of inbreeding outweigh the disadvantages, preferential breeding within small groups could be

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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

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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

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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

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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

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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

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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

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when these alleles are deleterious. Because most recessive alleles are rare in populations, it is unlikely that two unrelated 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

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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.

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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

292: 136:. However, increased homozygosity increases the probability of fixing beneficial alleles and also slightly decreases the probability of fixing deleterious alleles in a population. Inbreeding can result in purging of deleterious alleles from a population through 1063:

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.

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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

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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

862:. 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. 1016:

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. In the Western world, some

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Breeders must avoid breeding from individuals that demonstrate either homozygosity or heterozygosity for disease causing alleles. The goal of preventing the transfer of deleterious alleles may be achieved by reproductive isolation,

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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. In general, many mammal species, including humanity's closest

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Despite all its disadvantages, inbreeding can also have a variety of advantages, such as ensuring a child produced from the mating contains, and will pass on, a higher percentage of its mother/father's genetics, reducing the

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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.

746:: 'It turns out that females in these hermaphrodite insects are not really fertilizing their eggs themselves, but instead are having this done by a parasitic tissue that infects them at birth,' says Laura Ross of 131:

mechanisms, is the main selective reason for outcrossing. Crossbreeding between populations sometimes has positive effects on fitness-related traits, but also sometimes leads to negative effects known as

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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

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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

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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.

2502:"Impact of consanguineous marriages and degrees of inbreeding on fertility, child mortality, secondary sex ratio, selection intensity, and genetic load: a cross-sectional study from Northern India" 319:

Children of parent-child or sibling-sibling unions are at an increased risk compared to cousin-cousin unions. Inbreeding may result in a greater than expected phenotypic expression of deleterious

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eliminate deleterious alleles. With continuous inbreeding, genetic variation is lost and homozygosity is increased, enabling the expression of recessive deleterious alleles in homozygotes. The

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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.

290: 2231:"The existence of species rests on a metastable equilibrium between inbreeding and outbreeding. An essay on the close relationship between speciation, inbreeding and recessive mutations" 159:. 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 438:. 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. 3028:

Casas AM, Igartua E, Valles MP, Molina-Cano JL (November 1998). "Genetic diversity of barley cultivars grown in Spain, estimated by RFLP, similarity and coancestry coefficients".

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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

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Johnson, Warren E., David P. Onorato, Melody E. Roelke, E. Darrell Land, Mark Cunningham, Robert C. Belden, Roy McBride et al. "Genetic restoration of the Florida panther."

564:(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 3983:

Speicher MR, Motulsky AG, Antonarakis SE, Bittles AH, eds. (2010). "Consanguinity, Genetic Drift, and Genetic Diseases in Populations with Reduced Numbers of Founders".

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married his sister but had no issue) were married to their brothers and sisters, so as to keep the Ptolemaic blood "pure" and to strengthen the line of succession. King

750:. 'It seems that this infectious tissue derives from left-over sperm from their father, who has found a sneaky way of having more children by mating with his daughters.' 2843:

Bodkin JL, Ballachey BE, Cronin MA, Scribner KT (December 1999). "Population Demographics and Genetic Diversity in Remnant and Translocated Populations of Sea Otters".

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population was reduced to about 30 animals, so inbreeding became a problem. Several females were imported from Texas and now the population is better off genetically.

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Nabulsi MM, Tamim H, Sabbagh M, Obeid MY, Yunis KA, Bitar FF (February 2003). "Parental consanguinity and congenital heart malformations in a developing country".

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García-Cortés LA, Martínez-Ávila JC, Toro MA (2010-05-16). "Fine decomposition of the inbreeding and the coancestry coefficients by using the tabular method".

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was often practiced among European royal families, usually for interests of state. Over time, due to the relatively limited number of potential consorts, the

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Gilbert DA, Packer C, Pusey AE, Stephens JC, O'Brien SJ (1991-10-01). "Analytical DNA fingerprinting in lions: parentage, genetic diversity, and kinship".

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Peer K, Taborsky M (February 2005). "Outbreeding depression, but no inbreeding depression in haplodiploid Ambrosia beetles with regular sibling mating".

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Although there are several examples of inbred populations of wild animals, the negative consequences of this inbreeding are poorly documented. In the

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Fareed M, Afzal M (April 2016). "Increased cardiovascular risks associated with familial inbreeding: a population-based study of adolescent cohort".

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is found. Normally, the old ruler's eldest son and daughter (who could be either siblings or half-siblings) became the new rulers. All rulers of the

4504: 996:, schizophrenia and several others. Moreover, there is an increased risk for congenital heart disease depending on the inbreeding coefficient (See 734:

found that bedbugs, in contrast to most other insects, tolerate incest and are able to genetically withstand the effects of inbreeding quite well.

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decade. Population growth is limited by the fact that all Californian sea otters are descended from the isolated colony, resulting in inbreeding.

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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

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Fareed M, Afzal M (2014). "Evidence of inbreeding depression on height, weight, and body mass index: a population-based child cohort study".

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mortality rates, some individuals in the first generation of inbreeding will not live on to reproduce. Over time, with isolation, such as a

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Hartl, D.L., Jones, E.W. (2000) Genetics: Analysis of Genes and Genomes. Fifth Edition. Jones and Bartlett Publishers Inc., pp. 105–106.

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Intensive selection for higher yield has increased relationships among animals within breed and increased the rate of casual inbreeding.

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Among these populations with high levels of inbreeding, researchers have found several disorders prevalent among inbred offspring. In

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within a population. As a result, first-generation inbred individuals are more likely to show physical and health defects, including:

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are increased when the biological parents are more closely related. This is because such pairings have a 25% probability of producing

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than expected. In fact, predators are known for low genetic variance, along with most of the top portion of the trophic levels of an

434:, as the inbreeding first removes many deleterious genes, and permits the expression of genes that allow a population to adapt to an 119:), which is its ability to survive and reproduce. An individual who inherits such deleterious traits is colloquially referred to as 5323: 5134: 2899: 1884: 3593: 5129: 4961: 2886:

Wielebnowski N (1996). "Reassessing the relationship between juvenile mortality and genetic monomorphism in captive cheetahs".

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Swindell WR, et al. (2006). "Selection and Inbreeding Depression: Effects of Inbreeding Rate and Inbreeding Environment".

1084: 4545: 3371: 846:). Without the sorting of individuals by trait, a breed could not be established, nor could poor genetic material be removed. 4487: 3992: 3481: 3401: 2383: 2358: 2213: 1990: 889: 198: 580:

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.

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Of the practicing regions, Middle Eastern and northern Africa territories show the greatest frequencies of consanguinity.

4663: 4592: 453:, the population may fall below a minimum whereby the forced interbreeding between the remaining animals will result in 5065: 4834: 4628: 2119:"Does morality have a biological basis? An empirical test of the factors governing moral sentiments relating to incest" 1234: 445:, for example due to a bottleneck will unavoidably increase inbreeding for the entire population. This may mean that a 4581: 1021:

groups are highly inbred because they originate from small founder populations that have bred as a closed population.

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Kirkpatrick M, Jarne P (February 2000). "The Effects of a Bottleneck on Inbreeding Depression and the Genetic Load".

2284: 1646: 3573: 1513:, Alaks G, Graham L, Lacy RC (October 1994). "An experimental study of inbreeding depression in a natural habitat". 587:

There are several methods to compute this percentage. The two main ways are the path method and the tabular method.

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Taft, Robert et al. "Know thy mouse." Science Direct. Vol. 22, No. 12, Dec. 2006, pp. 649-653. Trends in Genetics.

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Bernstein H, Byerly HC, Hopf FA, Michod RE (September 1985). "Genetic damage, mutation, and the evolution of sex".

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Girotto G, Mezzavilla M, Abdulhadi K, Vuckovic D, Vozzi D, Khalifa Alkowari M, Gasparini P, Badii R (2014-01-01).

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Bittles AH, Grant JC, Sullivan SG, Hussain R (2002-01-01). "Does inbreeding lead to decreased human fertility?".

731: 3441: 2659: 1362: 1126: 5198: 81:, but more commonly refers to the genetic disorders and other consequences that may arise from expression of 3302:"The evolution of hermaphroditism by an infectious male-derived cell lineage: an inclusive-fitness analysis" 2706: 927:

and rats is of great importance for biomedical research. The inbreeding guarantees a consistent and uniform

5438: 5423: 5162: 4920: 2716:. Department of Anthropology and School of Biological & Biomedical Sciences, University of Durham, U.K. 2404:"Evolution in health and medicine Sackler colloquium: Consanguinity, human evolution, and complex diseases" 1352: 1130: 993: 987:

The fitness consequences of consanguineous mating have been studied since their scientific recognition by

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The issue of casual breeders who inbreed irresponsibly is discussed in the following quotation on cattle:

798: 267: 5252: 4512: 4343:"Does consanguinity increase the risk of schizophrenia? Study based on primary health care centre visits" 1982: 1219: 782: 5082: 4668: 3591:

S1008: Genetic Selection and Crossbreeding to Enhance Reproduction and Survival of Dairy Cattle (S-284)

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There may also be other deleterious effects besides those caused by recessive diseases. Thus, similar

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Offspring of biologically related persons are subject to the possible effects of inbreeding, such as

3077: 2779: 1114:. Thus marriage was seen as a union of lines of nobility and not as a contract between individuals. 517:

lions, it is known that a population bottleneck has occurred. Researchers found far greater genetic

5099: 4829: 4648: 3644: 3243:"Incestuous sisters: mate preference for brothers over unrelated males in Drosophila melanogaster" 1893:"Gene-Expression Changes Caused by Inbreeding Protect Against Inbreeding Depression in Drosophila" 513:

are usually related. Two lines are then being "line bred". Also, in some populations, such as the

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subject to higher health costs than cows of lower genetic merit for production (Cassell, 2001).

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significantly higher in a small inbreeding population than in a larger inbreeding population.

5373: 5338: 5257: 4956: 4790: 4755: 3471: 1292: 883: 759: 743: 412: 329: 236: 223: 116: 3675:"Consanguineous marriages : Preconception consultation in primary health care settings" 2577: 709: 5203: 5077: 5050: 5005: 4855: 4673: 4215: 4073: 3590: 3510: 3254: 3129: 2852: 2411: 2028: 1605: 1522: 1416: 1150: 966:

Consanguineous marriages (second-degree cousins or closer) in the world, in percentage (%).

565: 320: 128: 85: 40: 3099: 2743: 8: 5183: 5040: 4935: 4925: 4885: 4785: 4765: 4658: 4653: 1978:

The Natural History of Inbreeding and Outbreeding: Theoretical and Empirical Perspectives

1117: 1073: 404: 301: 259: 207: 137: 4219: 4077: 3514: 3258: 3133: 2856: 2415: 2032: 1609: 1526: 1420: 270:, and allowing the expression of recessive advantageous phenotypes. Some species with a 5267: 5210: 5111: 4895: 4780: 4775: 4750: 4403: 4359: 4342: 4283: 4258: 4239: 4183: 4166: 4147: 4037: 4012: 3965: 3922: 3897: 3870: 3843: 3699: 3674: 3531: 3498: 3417: 3342: 3277: 3242: 3192: 3167: 3145: 3041: 3005: 2980: 2961: 2868: 2792: 2674: 2640: 2482: 2434: 2403: 2323: 2298: 2257: 2230: 2143: 2118: 2099: 1957: 1917: 1892: 1873: 1830: 1782: 1757: 1733: 1708: 1689: 1676: 1659: 1491: 1448: 1357: 1347: 1322: 1297: 1107: 1052: 1048: 843: 839: 450: 416: 313: 160: 148: 144: 78: 5368: 4096: 4061: 5428: 5262: 5035: 4979: 4910: 4900: 4890: 4724: 4614: 4483: 4460: 4395: 4364: 4323: 4288: 4231: 4188: 4139: 4101: 4042: 3988: 3969: 3927: 3875: 3813: 3743: 3704: 3536: 3477: 3397: 3334: 3282: 3197: 3010: 2864: 2747: 2632: 2597: 2558: 2523: 2474: 2439: 2379: 2354: 2328: 2280: 2262: 2209: 2186: 2182: 2148: 2103: 2091: 2056: 2051: 2016: 1986: 1922: 1865: 1861: 1834: 1822: 1787: 1738: 1681: 1642: 1621: 1538: 1483: 1452: 1440: 1432: 1404: 1267: 1177: 1164:, royal women were believed to carry the bloodlines and so it was advantageous for a 1146: 1111: 1110:

among elites. These ties were often sealed only upon the birth of progeny within the

962: 953: 514: 442: 228: 108: 4151: 3346: 2965: 2872: 2796: 2707:"Genetic diversity and population genetic structure in the South American sea lion ( 2578:"Consanguinity as a determinant of reproductive behaviour and mortality in Pakistan" 2554: 2005:

Shields, W. M. 1982. Philopatry, Inbreeding, and the Evolution of Sex. Print. 50–69.

1961: 1877: 1495: 1382: 5167: 4940: 4915: 4587: 4450: 4440: 4384:"Towards a Complete North American Anabaptist Genealogy II: Analysis of Inbreeding" 4354: 4315: 4278: 4270: 4243: 4223: 4178: 4131: 4091: 4081: 4032: 4024: 4013:"Inbreeding effects on fertility in humans: evidence for reproductive compensation" 3957: 3917: 3909: 3865: 3855: 3805: 3775: 3735: 3694: 3686: 3657: 3526: 3518: 3360: 3324: 3316: 3301: 3272: 3262: 3187: 3179: 3149: 3137: 3045: 3037: 3000: 2992: 2981:"Estimation of the coancestry coefficient: basis for a short-term genetic distance" 2953: 2895: 2860: 2823: 2784: 2739: 2644: 2624: 2589: 2550: 2513: 2486: 2466: 2429: 2419: 2318: 2310: 2252: 2242: 2178: 2138: 2130: 2083: 2046: 2036: 1949: 1940:

Livingstone FB (1969). "Genetics, Ecology, and the Origins of Incest and Exogamy".

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Great-grandfather/great-granddaughter or great-grandmother/great-grandson → 6.25% (

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Khlat M (December 1989). "Inbreeding effects on fetal growth in Beirut, Lebanon".

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Michod RE. Eros and Evolution: A Natural Philosophy of Sex. (1994) Perseus Books,

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The general negative outlook and eschewal of inbreeding that is prevalent in the

957: 924: 721: 713: 549: 408: 372: 187: 3577: 3499:"Deleterious alleles in the context of domestication, inbreeding, and selection" 1908: 1660:"The Genetic Interpretation of Inbreeding Depression and Outbreeding Depression" 1168:

to marry his sister or half-sister; in such cases a special combination between

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and Lebanon, there is a positive association between consanguinity and reported

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Proceedings of the National Academy of Sciences of the United States of America

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Tadmouri GO, Nair P, Obeid T, Al Ali MT, Al Khaja N, Hamamy HA (October 2009).

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Proceedings of the National Academy of Sciences of the United States of America

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Proceedings of the National Academy of Sciences of the United States of America

1428: 1249: 1000:) of the offspring, with significant risk accompanied by an F =.125 or higher. 988: 912: 518: 427: 164: 4135: 3961: 3766:

Woodley MA (2009). "Inbreeding depression and IQ in a study of 72 countries".

3739: 3690: 3661: 3141: 505: 5417: 5363: 5156: 5124: 5088: 5072: 4399: 3747: 3361:"The Evolution of Hermaphroditism by an Infectious Male-Derived Cell Lineage" 2314: 1758:"Inbreeding depression and purging in a haplodiploid: gender-related effects" 1436: 1224: 1201: 1193: 1161: 1044: 1009: 385: 271: 247: 93: 4319: 2788: 2660:"Establishment of New Population Centers with Changes in Migration Patterns" 2424: 2351:

Inbreeding, incest, and the incest taboo: the state of knowledge at the turn

1617: 1534: 5402: 5215: 4880: 4760: 4464: 4368: 4292: 4143: 4086: 3948:

Van Den Berghe PL (2010). "Human inbreeding avoidance: Culture in nature".

3913: 3879: 3860: 3708: 3540: 3338: 3286: 3201: 3183: 3014: 2636: 2593: 2562: 2527: 2478: 2443: 2266: 2247: 2190: 2152: 2134: 2095: 2060: 2041: 1926: 1869: 1826: 1791: 1742: 1685: 1487: 1444: 1307: 1254: 1197: 1096: 1056: 1032: 949: 928: 847: 754: 481:

relatives, avoid close inbreeding possibly due to the deleterious effects.

343: 151:, breeders may use inbreeding when trying to establish a new and desirable 4327: 4274: 4235: 4192: 4046: 3931: 3817: 3723: 2751: 2601: 2332: 2017:"Male-male competition magnifies inbreeding depression in wild house mice" 1625: 1542: 103:, which can increase the chances of offspring being affected by recessive 5343: 5094: 4860: 4455: 4105: 2828: 2811: 1773: 1578: 1557: 1479: 1342: 1262: 1189: 1153:

also frequently engaged in first-cousin unions as well as the occasional

1138: 791: 770: 168: 107:. In extreme cases, this usually leads to at least temporarily decreased 4407: 3388:

Freeman S, Herran JC (2007). "Aging and other life history characters".

3049: 2518: 2501: 1848:

Gulisija D, Crow JF (May 2007). "Inferring purging from pedigree data".

893:

that document the negative health consequences of excessive inbreeding.

5348: 4865: 4734: 4714: 4689: 4383: 3476:. Food and Agriculture Organization of the United Nations. p. 50. 2678: 2470: 1185: 1181: 1141:

often cited as an ill-effect. The closely related houses of Habsburg,

1018: 533: 510: 469: 454: 431: 362: 305: 276: 215: 112: 100: 51: 4445: 4428: 3522: 3329: 2900:

10.1002/(SICI)1098-2361(1996)15:4<353::AID-ZOO1>3.0.CO;2-A

39:

For reproduction by which offspring arise from a single organism, see

4227: 2087: 1402: 1121: 1092: 1088: 855: 522: 435: 339: 176: 62: 3092:"4.5 Calculation of inbreeding and relationship, the tabular method" 2941: 2765:

Ramel C (1998). "Biodiversity and intraspecific genetic variation".

2374:

Griffiths AJ, Miller JH, Suzuki DT, Lewontin RC, Gelbart WM (1999).

740:

females prefer to mate with their own brothers over unrelated males.

5225: 4905: 4875: 4719: 4709: 4637: 4028: 3554:

G2036 Culling the Commercial Cow Herd: BIF Fact Sheet, MU Extension

3320: 2957: 2628: 1953: 1818: 1403:

de Boer RA, Vega-Trejo R, Kotrschal A, Fitzpatrick JL (July 2021).

1204:

following their father's death, are the most widely known example.

1173: 1169: 1134: 1103: 932: 309: 70: 4426: 4259:"Consanguinity and recurrence risk of stillbirth and infant death" 3982: 2499: 1192:'s mother is reported to have been the half-sister to his father, 3168:"Evidence for frequent incest in a cooperatively breeding mammal" 2500:

Fareed M, Kaisar Ahmad M, Azeem Anwar M, Afzal M (January 2017).

1244: 1165: 1028: 859: 727: 478: 446: 312:. Except in certain rare circumstances, such as new mutations or 202:

Inbreeding coefficients of various populations in Europe and Asia

156: 5149: 5021: 4729: 4704: 3166:

Nichols HJ, Cant MA, Hoffman JI, Sanderson JL (December 2014).

3119: 2169:

Pusey A, Wolf M (May 1996). "Inbreeding avoidance in animals".

1332: 1302: 1040: 945: 496: 124: 89: 66: 794:

breed, affecting almost half the population in some countries.

560:

A measure of inbreeding of an individual A is the probability

4533:

King Tut Mysteries Solved: Was Disabled, Malarial, and Inbred

4167:"Decreased fecundability in Hutterite couples sharing HLA-DR" 3027: 2842: 2373: 1405:"Meta-analytic evidence that animals rarely avoid inbreeding" 1060: 1036: 526: 473: 465: 378: 4256: 1558:"Comparison of inbreeding and outbreeding in hermaphroditic 881:

The BBC produced two documentaries on dog inbreeding titled

179:. Inbreeding in plants also occurs naturally in the form of 123:. The avoidance of expression of such deleterious recessive 4606: 4121: 3987:(4th ed.). Berlin: Springer-Verlag. pp. 507–528. 3985:

Vogel and Motulsky's human genetics problems and approaches

3612:"Inbreeding: Its Meaning, Uses and Effects on Farm Animals" 3165: 3078:

How to compute and inbreeding coefficient (the path method)

1890: 923:

Systematic inbreeding and maintenance of inbred strains of

620:

Grandfather/granddaughter or grandmother/grandson → 12.5% (

4257:

Stoltenberg C, Magnus P, Skrondal A, Lie RT (April 1999).

3724:"Inbreeding and extinction: Effects of rate of inbreeding" 3497:

Bosse M, Megens HJ, Derks MF, Cara ÁM, Groenen MA (2019).

3240: 2729: 1670:(3). Oregon: Society for the Study of Evolution: 622–629. 332:

levels and higher incidence rates of being affected by an

4990: 4699: 4694: 4568:

Inbreeding: Its Meaning, Uses and Effects on Farm Animals

1709:"Fixation probability and time in subdivided populations" 1595: 931:

for experimental purposes and enables genetic studies in

576:

also denoted as the kinship coefficient between A and B.

2299:"ABC of clinical genetics. Genetics of common disorders" 1508: 1465: 4482:. Richmond, US: Kensington House Books. pp. 1–13. 3841: 3722:

Reed DH, Lowe EH, Briscoe DA, Frankham R (2003-05-01).

3721: 2913:

UCF report on complex genetic health of Florida panther

724:

females regularly mate with their fathers and brothers.

716:

females regularly mate with their fathers and brothers.

590:

Typical coancestries between relatives are as follows:

4381: 2116: 423:

factors, the deleterious inherited traits are culled.

4165:

Ober C, Elias S, Kostyu DD, Hauck WW (January 1992).

2978: 1850:

Evolution; International Journal of Organic Evolution

1807:

Evolution; International Journal of Organic Evolution

1664:

Evolution; International Journal of Organic Evolution

541:

mortality, low fecundity, and poor breeding success.

252:

Major histocompatibility complex and sexual selection

4429:"Consanguinity and hereditary hearing loss in Qatar" 4164: 3496: 3418:"Polycystic kidney disease | International Cat Care" 2704: 802:

An intensive form of inbreeding where an individual

4480:

The Gotha: Still a Continental Royal Family, Vol. 1

4206:Diamond JM (1987). "Causes of death before birth". 4062:"Effect of inbreeding on IQ and mental retardation" 3844:"Consanguinity and reproductive health among Arabs" 2979:Reynolds J, Weir BS, Cockerham CC (November 1983). 509:there is more than one dominant male, the group of 250:may be more vulnerable to infectious diseases (see 4566:Dale Vogt, Helen A. Swartz and John Massey, 1993. 4340: 3563:. Extension.missouri.edu. Retrieved on 2013-03-05. 3241:Loyau A, Cornuau JH, Clobert J, Danchin E (2012). 1891:García C, Avila V, Quesada H, Caballero A (2012). 1591: 1589: 814:and so on, in order to maximise the percentage of 633:Half-brother/half-sister, Double cousins → 12.5% ( 4593:"Cross-Fertilization in Animals and in Man" 3796:Kamin LJ (1980). "Inbreeding depression and IQ". 3465: 3463: 3461: 3234: 2725: 2723: 2695:", College of Agriculture, Utah State University. 2575: 2344: 2342: 2164: 2162: 5415: 3837: 3835: 3833: 3831: 3829: 3827: 3609: 3353: 2614: 2206:Philopatry, inbreeding, and the evolution of sex 2014: 1637: 1635: 4341:Bener A, Dafeeah EE, Samson N (December 2012). 4117: 4115: 4010: 3891: 3889: 3761: 3759: 3757: 2397: 2395: 2117:Lieberman D, Tooby J, Cosmides L (April 2003). 1755: 1586: 3947: 3458: 3096:Genetic calculation applets and other programs 2720: 2339: 2208:. Albany: State University of New York Press. 2159: 672:Half-uncle/niece or half-aunt/nephew → 6.25% ( 5006: 4622: 3824: 3791: 3789: 3387: 2942:"Coefficients of inbreeding and relationship" 2378:. New York: W. H. Freeman. pp. 726–727. 1632: 32:"Inbred" redirects here. For other uses, see 4471: 4394:(4). Wayne State University Press: 533–545. 4112: 4006: 4004: 3976: 3943: 3941: 3886: 3754: 3610:Vogt D, Swartz HA, Massey J (October 1993). 2885: 2812:"The sea otter in the eastern Pacific Ocean" 2576:Bittles AH, Grant JC, Shami SA (June 1993). 2534: 2493: 2401: 2392: 1847: 1804: 1468:American Journal of Medical Genetics. Part A 1398: 1396: 239:or having cranial abnormalities, such as in 4382:Agarwala R, Schäffer AA, Tomlin JF (2001). 4053: 4011:Ober C, Hyslop T, Hauck WW (January 1999). 3299: 2540: 2456: 2450: 2290: 2015:Meagher S, Penn DJ, Potts WK (March 2000). 1939: 1700: 5013: 4999: 4629: 4615: 3786: 3216:"Insect Incest Produces Healthy Offspring" 3161: 3159: 2348: 1756:Tien NS, Sabelis MW, Egas M (March 2015). 838:Breeding in domestic animals is primarily 4586: 4454: 4444: 4358: 4308:American Journal of Physical Anthropology 4282: 4182: 4095: 4085: 4036: 4001: 3938: 3921: 3898:"Incest, inbreeding and mental abilities" 3869: 3859: 3698: 3603: 3600:. Nimss.umd.edu. Retrieved on 2013-03-05. 3530: 3328: 3276: 3266: 3191: 3004: 2827: 2778: 2517: 2433: 2423: 2322: 2256: 2246: 2168: 2142: 2050: 2040: 1974: 1916: 1781: 1732: 1675: 1577: 1393: 1051:. Furthermore, among inbred children in 830:would receive 93.75% of their genes from 748:Oxford University's Department of Zoology 218:zygotes, resulting in offspring with two 190:which can prevent inbreeding depression. 27:Reproduction by closely related organisms 5324:2019 South Wales paternal sex abuse case 4582:Consanguineous marriages with global map 3647:. Petplace.com. Retrieved on 2013-03-05. 2651: 2296: 2073: 1706: 1083: 961: 797: 781: 765: 708: 495: 304:disorders occur in individuals who have 286: 197: 96:. Animals avoid inbreeding only rarely. 45: 4962:List of genetics research organizations 4505:"Women in Ancient Egypt, Women and Law" 4205: 3895: 3765: 3156: 3089: 2203: 1137:was known for its intermarriages; the 1067: 568:. This probability F(A) is called the " 461:who cannot survive because of illness. 186:Inbreeding can significantly influence 171:are used as stocks for the creation of 14: 5416: 4477: 4059: 3672: 2939: 2809: 2705:Freilich S, Hoelzel AR, Choudhury SR. 1200:, who married and became co-rulers of 4994: 4610: 4502: 4305: 3795: 3473:Inbreeding and brood stock management 3067:. Paris: Masson et Cie. p. 1048. 2935: 2933: 2764: 2744:10.1093/oxfordjournals.jhered.a111107 2582:International Journal of Epidemiology 2402:Bittles AH, Black ML (January 2010). 2110: 1657: 1059:cases. Historically, populations of 918: 890:Pedigree Dogs Exposed: Three Years On 594:Father/daughter or mother/son → 25% ( 5120:Article 809 of the Korean Civil Code 4570:. University of Missouri, Extension. 4375: 3469: 3377:from the original on April 22, 2023. 2657: 2228: 1555: 646:Uncle/niece or aunt/nephew → 12.5% ( 282: 175:lines to make use of the effects of 3062: 2376:An introduction to genetic analysis 1749: 818:s genes in the offspring. 87.5% of 777: 308:for a particular recessive genetic 24: 4171:American Journal of Human Genetics 4017:American Journal of Human Genetics 3042:10.1111/j.1439-0523.1998.tb01968.x 2930: 2410:. 107 Suppl 1 (suppl 1): 1779–86. 1677:10.1111/j.1558-5646.1991.tb04333.x 1365:(how some plants avoid inbreeding) 1235:Cousin marriage in the Middle East 1091:, a member of the famously inbred 297:Animation of uniparental isodisomy 143:Inbreeding is a technique used in 77:. By analogy, the term is used in 54:alleles through an inbred pedigree 25: 5450: 4559: 4543: 4263:American Journal of Public Health 3300:Gardner A, Ross L (August 2011). 3218:. 8 December 2011. Archived from 2459:American Journal of Human Biology 2349:Wolf AP, Durham WH, eds. (2005). 2171:Trends in Ecology & Evolution 275:promoted, potentially leading to 4975: 4974: 4347:Mental Health in Family Medicine 3098:. Genetics pages. Archived from 2927:329, no. 5999 (2010): 1641-1645. 2865:10.1046/j.1523-1739.1999.98124.x 2123:Proceedings. Biological Sciences 1862:10.1111/j.1558-5646.2007.00088.x 1240:Evolution of sexual reproduction 1106:was used as a method of forming 4537: 4526: 4496: 4420: 4334: 4299: 4250: 4199: 4158: 3715: 3666: 3650: 3638: 3584: 3566: 3547: 3490: 3434: 3410: 3381: 3293: 3208: 3113: 3083: 3071: 3065:Les Mathématiques de l'hérédité 3056: 3021: 2972: 2917: 2906: 2879: 2836: 2803: 2758: 2698: 2685: 2608: 2569: 2555:10.1016/j.annepidem.2016.03.001 2367: 2273: 2222: 2197: 2067: 2008: 1999: 1968: 1933: 1841: 1798: 1196:(also called Cleopatra VI) and 896: 732:North Carolina State University 704: 2693:ADVS 3910 Wild Horses Behavior 1651: 1549: 1502: 1459: 1409:Nature Ecology & Evolution 1376: 1363:Self-incompatibility in plants 1097:extremely pronounced lower jaw 905: 760:eat them alive from the inside 69:or breeding of individuals or 13: 1: 3950:Behavioral and Brain Sciences 3679:Journal of Community Genetics 2353:. Stanford University Press. 1370: 1003: 5163:Prohibited degree of kinship 4921:Missing heritability problem 4636: 3896:Roberts DF (November 1967). 3780:10.1016/j.intell.2008.10.007 3574:"Genetic Evaluation Results" 3268:10.1371/journal.pone.0051293 2667:Journal of Field Ornithology 2183:10.1016/0169-5347(96)10028-8 1353:Prohibited degree of kinship 1131:King Christian IX of Denmark 994:disorders of sex development 7: 3442:"Polycystic Kidney Disease" 1983:University of Chicago Press 1909:10.1534/genetics.112.142687 1220:Coefficient of relationship 1207: 1157:and uncle–niece marriages. 858:, or, in the extreme case, 555: 484: 193: 10: 5455: 5083:Parallel and cross cousins 5020: 3810:10.1037/0033-2909.87.3.469 2997:10.1093/genetics/105.3.767 2767:Pure and Applied Chemistry 2297:Kingston HM (April 1989). 1725:10.1093/genetics/164.2.767 1429:10.1038/s41559-021-01453-9 1077: 1071: 970: 943: 911:trait. This is called the 699: 127:caused by inbreeding, via 38: 31: 5393:Genetic sexual attraction 5385: 5316: 5281: 5249:Coefficient of inbreeding 5241: 5234: 5176: 5110: 5028: 4970: 4949: 4848: 4799: 4743: 4682: 4644: 4136:10.1080/03014460110075657 4060:Morton NE (August 1978). 3962:10.1017/S0140525X00014850 3691:10.1007/s12687-011-0072-y 3662:10.1016/j.tig.2006.09.010 3503:Evolutionary Applications 3142:10.1007/s10592-010-0084-x 2810:Kenyon KW (August 1969). 1707:Whitlock MC (June 2003). 1288:Identical ancestors point 1278:Genetic sexual attraction 998:coefficient of inbreeding 978:coefficient of inbreeding 939: 806:is mated to his daughter 788:polycystic kidney disease 570:coefficient of inbreeding 212:chances of such disorders 92:sexual relationships and 3080:, Braque du Bourbonnais. 2584:(Submitted manuscript). 2315:10.1136/bmj.298.6678.949 2229:Joly E (December 2011). 822:s genes would come from 342:both in litter size and 208:congenital birth defects 4546:"The House of Ptolomey" 4320:10.1002/ajpa.1330800407 4124:Annals of Human Biology 3902:British Medical Journal 3740:10.1023/A:1024081416729 3645:Top Cat Breeds for 2004 3309:The American Naturalist 2789:10.1351/pac199870112079 2732:The Journal of Heredity 2617:The American Naturalist 2425:10.1073/pnas.0906079106 1618:10.1126/science.3898363 1535:10.1126/science.7939661 1388:Encyclopædia Britannica 1328:List of coupled cousins 1318:Intellectual inbreeding 1080:List of coupled cousins 685:First cousins → 6.25% ( 532:In Central California, 491:South American sea lion 334:intellectual disability 306:two copies of an allele 241:Northern elephant seals 163:or through culling. In 34:Inbred (disambiguation) 5221:Jewish views on incest 5046:Child incestuous abuse 4599:Encyclopedia Americana 4087:10.1073/pnas.75.8.3906 3914:10.1136/bmj.4.5575.336 3861:10.1186/1742-4755-6-17 3798:Psychological Bulletin 3673:Hamamy H (July 2012). 3620:University of Missouri 3394:Pearson Education, Inc 3184:10.1098/rsbl.2014.0898 2543:Annals of Epidemiology 2248:10.1186/1745-6150-6-62 2135:10.1098/rspb.2002.2290 2042:10.1073/pnas.97.7.3324 1338:Outbreeding depression 1283:Heterozygote advantage 1099: 967: 879: 835: 795: 774: 744:Cottony cushion scales 717: 607:Brother/sister → 25% ( 501: 419:) breeding or natural 415:caused by purposeful ( 298: 203: 134:outbreeding depression 99:Inbreeding results in 88:traits resulting from 55: 5374:Sheffield incest case 5339:Colt clan incest case 5258:Inbreeding depression 4957:List of genetic codes 4275:10.2105/ajph.89.4.517 3728:Conservation Genetics 3390:Evolutionary Analysis 3122:Conservation Genetics 1975:Thornhill NW (1993). 1293:Inbreeding depression 1180:uninterruptedly from 1087: 965: 884:Pedigree Dogs Exposed 867: 801: 785: 769: 712: 499: 413:population bottleneck 330:intelligence quotient 296: 201: 117:inbreeding depression 61:is the production of 49: 5135:cousin marriage laws 5130:In the United States 5078:Incest between twins 5051:Consanguine marriage 4856:Behavioural genetics 2845:Conservation Biology 2829:10.3996/nafa.68.0001 2816:North American Fauna 2594:10.1093/ije/22.3.463 1942:Current Anthropology 1774:10.1038/hdy.2014.106 1579:10.1038/hdy.1993.163 1480:10.1002/ajmg.a.10020 1068:Royalty and nobility 790:is prevalent in the 773:in Gunma Safari Park 566:identical by descent 393:cardiovascular risks 161:assortative breeding 129:inbreeding avoidance 41:Asexual reproduction 5439:Kinship and descent 5424:Population genetics 5329:Álvarez incest case 5199:film and television 5041:Avunculate marriage 4936:Population genomics 4926:Molecular evolution 4886:Genetic engineering 4220:1987Natur.329..487D 4078:1978PNAS...75.3906M 3848:Reproductive Health 3580:on August 27, 2001. 3515:2019EvApp..12....6B 3446:www.vet.cornell.edu 3259:2012PLoSO...751293L 3134:2010ConG...11.1945G 2946:American Naturalist 2857:1999ConBi..13.1378B 2519:10.1038/pr.2016.177 2416:2010PNAS..107.1779B 2204:Shields WM (1982). 2033:2000PNAS...97.3324M 1610:1985Sci...229.1277B 1527:1994Sci...266..271J 1421:2021NatEE...5..949D 1215:Álvarez incest case 1118:Royal intermarriage 1108:political alliances 1074:Royal intermarriage 1049:neural tube defects 302:Autosomal recessive 138:purifying selection 75:related genetically 5268:Westermarck effect 5066:in the Middle East 4896:Genetic monitoring 4575:2012-03-08 at the 3596:2006-09-10 at the 3559:2016-04-16 at the 3370:. April 26, 2019. 2506:Pediatric Research 2471:10.1002/ajhb.22599 1562:(L.) (land snail)" 1560:Arianta arbustorum 1358:Selective breeding 1348:Proximity of blood 1323:Legality of incest 1298:Inbreeding in fish 1100: 968: 919:Laboratory animals 844:selective breeding 836: 796: 775: 718: 502: 314:uniparental disomy 299: 268:recombination load 204: 149:livestock breeding 147:. For example, in 145:selective breeding 109:biological fitness 79:human reproduction 56: 5411: 5410: 5312: 5311: 5263:Pedigree collapse 5036:Accidental incest 4988: 4987: 4911:He Jiankui affair 4901:Genetic genealogy 4891:Genetic diversity 4820:the British Isles 4725:Genetic variation 4489:978-0-9771961-7-3 4478:Beeche A (2009). 4446:10.1159/000360475 3994:978-3-540-37654-5 3523:10.1111/eva.12691 3483:978-92-5-104340-0 3403:978-0-13-227584-2 2940:Wright S (1922). 2773:(11): 2079–2084. 2709:Otaria flavescens 2385:978-0-7167-3771-1 2360:978-0-8047-5141-4 2215:978-0-87395-618-5 1992:978-0-226-79854-7 1604:(4719): 1277–81. 1268:Genetic diversity 1178:Ptolemaic dynasty 1112:arranged marriage 954:Pedigree collapse 443:genetic diversity 400:mating randomly. 351:genetic disorders 321:recessive alleles 294: 283:Genetic disorders 229:natural selection 224:produce disorders 220:recessive alleles 73:that are closely 16:(Redirected from 5446: 5378:Whittaker family 5239: 5238: 5168:Ten Abominations 5015: 5008: 5001: 4992: 4991: 4978: 4977: 4941:Reverse genetics 4916:Medical genetics 4631: 4624: 4617: 4608: 4607: 4603: 4595: 4554: 4553: 4541: 4535: 4530: 4524: 4523: 4521: 4520: 4511:. Archived from 4500: 4494: 4493: 4475: 4469: 4468: 4458: 4448: 4424: 4418: 4417: 4415: 4414: 4379: 4373: 4372: 4362: 4338: 4332: 4331: 4303: 4297: 4296: 4286: 4254: 4248: 4247: 4228:10.1038/329487a0 4203: 4197: 4196: 4186: 4162: 4156: 4155: 4119: 4110: 4109: 4099: 4089: 4057: 4051: 4050: 4040: 4008: 3999: 3998: 3980: 3974: 3973: 3945: 3936: 3935: 3925: 3893: 3884: 3883: 3873: 3863: 3839: 3822: 3821: 3793: 3784: 3783: 3763: 3752: 3751: 3719: 3713: 3712: 3702: 3670: 3664: 3654: 3648: 3642: 3636: 3635: 3633: 3631: 3626:on March 8, 2012 3622:. Archived from 3607: 3601: 3588: 3582: 3581: 3576:. Archived from 3570: 3564: 3551: 3545: 3544: 3534: 3494: 3488: 3487: 3467: 3456: 3455: 3453: 3452: 3438: 3432: 3431: 3429: 3428: 3414: 3408: 3407: 3392:(4th ed.). 3385: 3379: 3378: 3376: 3365: 3357: 3351: 3350: 3332: 3306: 3297: 3291: 3290: 3280: 3270: 3238: 3232: 3231: 3229: 3227: 3212: 3206: 3205: 3195: 3178:(12): 20140898. 3163: 3154: 3153: 3117: 3111: 3110: 3108: 3107: 3087: 3081: 3075: 3069: 3068: 3060: 3054: 3053: 3025: 3019: 3018: 3008: 2976: 2970: 2969: 2952:(645): 330–338. 2937: 2928: 2921: 2915: 2910: 2904: 2903: 2883: 2877: 2876: 2840: 2834: 2833: 2831: 2807: 2801: 2800: 2782: 2762: 2756: 2755: 2727: 2718: 2717: 2715: 2702: 2696: 2689: 2683: 2682: 2664: 2658:Leck CF (1980). 2655: 2649: 2648: 2612: 2606: 2605: 2573: 2567: 2566: 2538: 2532: 2531: 2521: 2497: 2491: 2490: 2454: 2448: 2447: 2437: 2427: 2399: 2390: 2389: 2371: 2365: 2364: 2346: 2337: 2336: 2326: 2309:(6678): 949–52. 2294: 2288: 2277: 2271: 2270: 2260: 2250: 2226: 2220: 2219: 2201: 2195: 2194: 2166: 2157: 2156: 2146: 2129:(1517): 819–26. 2114: 2108: 2107: 2088:10.1554/05-493.1 2082:(5): 1014–1022. 2071: 2065: 2064: 2054: 2044: 2012: 2006: 2003: 1997: 1996: 1972: 1966: 1965: 1937: 1931: 1930: 1920: 1888: 1882: 1881: 1845: 1839: 1838: 1802: 1796: 1795: 1785: 1753: 1747: 1746: 1736: 1704: 1698: 1697: 1679: 1658:Lynch M (1991). 1655: 1649: 1639: 1630: 1629: 1593: 1584: 1583: 1581: 1553: 1547: 1546: 1506: 1500: 1499: 1463: 1457: 1456: 1400: 1391: 1380: 1313:Insular dwarfism 1057:cleft lip/palate 810:, granddaughter 778:Domestic animals 738:Common fruit fly 694: 693: 689: 681: 680: 676: 668: 667: 663: 655: 654: 650: 642: 641: 637: 629: 628: 624: 616: 615: 611: 603: 602: 598: 369:infant mortality 357:facial asymmetry 295: 260:Muller's ratchet 181:self-pollination 21: 5454: 5453: 5449: 5448: 5447: 5445: 5444: 5443: 5414: 5413: 5412: 5407: 5381: 5369:Patrick Stübing 5359:Moe incest case 5354:Maria Ersdotter 5308: 5304:Phaedra complex 5299:Oedipus complex 5294:Jocasta complex 5289:Electra complex 5277: 5273:Kin recognition 5230: 5194:Popular culture 5172: 5106: 5056:Cousin marriage 5024: 5019: 4989: 4984: 4966: 4945: 4844: 4835:the Middle East 4801:Archaeogenetics 4795: 4739: 4678: 4640: 4635: 4577:Wayback Machine 4562: 4557: 4542: 4538: 4531: 4527: 4518: 4516: 4501: 4497: 4490: 4476: 4472: 4439:(1–4): 175–82. 4425: 4421: 4412: 4410: 4380: 4376: 4339: 4335: 4304: 4300: 4255: 4251: 4214:(6139): 487–8. 4204: 4200: 4163: 4159: 4120: 4113: 4058: 4054: 4009: 4002: 3995: 3981: 3977: 3946: 3939: 3908:(5575): 336–7. 3894: 3887: 3840: 3825: 3794: 3787: 3764: 3755: 3720: 3716: 3671: 3667: 3655: 3651: 3643: 3639: 3629: 3627: 3608: 3604: 3598:Wayback Machine 3589: 3585: 3572: 3571: 3567: 3561:Wayback Machine 3552: 3548: 3495: 3491: 3484: 3470:Tave D (1999). 3468: 3459: 3450: 3448: 3440: 3439: 3435: 3426: 3424: 3416: 3415: 3411: 3404: 3396:. p. 484. 3386: 3382: 3374: 3363: 3359: 3358: 3354: 3304: 3298: 3294: 3239: 3235: 3225: 3223: 3214: 3213: 3209: 3172:Biology Letters 3164: 3157: 3118: 3114: 3105: 3103: 3090:Christensen K. 3088: 3084: 3076: 3072: 3061: 3057: 3026: 3022: 2977: 2973: 2938: 2931: 2922: 2918: 2911: 2907: 2884: 2880: 2841: 2837: 2808: 2804: 2780:10.1.1.484.8521 2763: 2759: 2728: 2721: 2713: 2703: 2699: 2690: 2686: 2662: 2656: 2652: 2613: 2609: 2574: 2570: 2539: 2535: 2498: 2494: 2455: 2451: 2400: 2393: 2386: 2372: 2368: 2361: 2347: 2340: 2295: 2291: 2278: 2274: 2227: 2223: 2216: 2202: 2198: 2167: 2160: 2115: 2111: 2072: 2068: 2013: 2009: 2004: 2000: 1993: 1973: 1969: 1938: 1934: 1889: 1885: 1846: 1842: 1803: 1799: 1754: 1750: 1705: 1701: 1656: 1652: 1640: 1633: 1594: 1587: 1556:Chen X (1993). 1554: 1550: 1521:(5183): 271–3. 1507: 1503: 1464: 1460: 1401: 1394: 1381: 1377: 1373: 1368: 1273:Genetic purging 1230:Cousin marriage 1210: 1133:. The House of 1102:Inter-nobility 1082: 1076: 1070: 1006: 973: 960: 958:Cousin marriage 942: 925:laboratory mice 921: 908: 899: 780: 722:Banded mongoose 714:Banded mongoose 707: 702: 691: 687: 686: 678: 674: 673: 665: 661: 660: 652: 648: 647: 639: 635: 634: 626: 622: 621: 613: 609: 608: 600: 596: 595: 558: 550:Florida panther 487: 397: 373:child mortality 287: 285: 196: 188:gene expression 50:The passage of 44: 37: 28: 23: 22: 15: 12: 11: 5: 5452: 5442: 5441: 5436: 5431: 5426: 5409: 5408: 5406: 5405: 5400: 5395: 5389: 5387: 5383: 5382: 5380: 5379: 5376: 5371: 5366: 5361: 5356: 5351: 5346: 5341: 5336: 5334:Armando Lucero 5331: 5326: 5320: 5318: 5314: 5313: 5310: 5309: 5307: 5306: 5301: 5296: 5291: 5285: 5283: 5279: 5278: 5276: 5275: 5270: 5265: 5260: 5255: 5245: 5243: 5236: 5232: 5231: 5229: 5228: 5223: 5218: 5213: 5208: 5207: 5206: 5201: 5191: 5186: 5180: 5178: 5174: 5173: 5171: 5170: 5165: 5160: 5153: 5146: 5145: 5144: 5143: 5142: 5127: 5122: 5116: 5114: 5108: 5107: 5105: 5104: 5103: 5102: 5092: 5085: 5080: 5075: 5070: 5069: 5068: 5063: 5053: 5048: 5043: 5038: 5032: 5030: 5026: 5025: 5018: 5017: 5010: 5003: 4995: 4986: 4985: 4983: 4982: 4971: 4968: 4967: 4965: 4964: 4959: 4953: 4951: 4947: 4946: 4944: 4943: 4938: 4933: 4931:Plant genetics 4928: 4923: 4918: 4913: 4908: 4903: 4898: 4893: 4888: 4883: 4878: 4873: 4871:Genome editing 4868: 4863: 4858: 4852: 4850: 4849:Related topics 4846: 4845: 4843: 4842: 4837: 4832: 4827: 4822: 4817: 4812: 4806: 4804: 4797: 4796: 4794: 4793: 4788: 4783: 4778: 4773: 4771:Immunogenetics 4768: 4763: 4758: 4753: 4747: 4745: 4741: 4740: 4738: 4737: 4732: 4727: 4722: 4717: 4712: 4707: 4702: 4697: 4692: 4686: 4684: 4683:Key components 4680: 4679: 4677: 4676: 4671: 4666: 4661: 4656: 4651: 4645: 4642: 4641: 4634: 4633: 4626: 4619: 4611: 4605: 4604: 4584: 4579: 4561: 4560:External links 4558: 4556: 4555: 4536: 4525: 4495: 4488: 4470: 4433:Human Heredity 4419: 4374: 4333: 4298: 4249: 4198: 4157: 4111: 4052: 4029:10.1086/302198 4000: 3993: 3975: 3937: 3885: 3823: 3804:(3): 469–478. 3785: 3774:(3): 268–276. 3753: 3734:(3): 405–410. 3714: 3665: 3649: 3637: 3602: 3583: 3565: 3546: 3489: 3482: 3457: 3433: 3409: 3402: 3380: 3352: 3321:10.1086/660823 3315:(2): 191–201. 3292: 3253:(12): e51293. 3233: 3222:on 16 May 2017 3207: 3155: 3128:(5): 1945–52. 3112: 3082: 3070: 3055: 3030:Plant Breeding 3020: 2971: 2958:10.1086/279872 2929: 2916: 2905: 2894:(4): 353–369. 2878: 2851:(6): 1378–85. 2835: 2802: 2757: 2719: 2697: 2684: 2673:(2): 168–173. 2650: 2629:10.1086/303312 2623:(2): 154–167. 2607: 2568: 2533: 2492: 2449: 2391: 2384: 2366: 2359: 2338: 2289: 2272: 2235:Biology Direct 2221: 2214: 2196: 2158: 2109: 2066: 2007: 1998: 1991: 1967: 1954:10.1086/201009 1932: 1883: 1856:(5): 1043–51. 1840: 1819:10.1554/04-128 1797: 1748: 1699: 1650: 1631: 1585: 1572:(5): 456–461. 1548: 1501: 1458: 1415:(7): 949–964. 1392: 1374: 1372: 1369: 1367: 1366: 1360: 1355: 1350: 1345: 1340: 1335: 1330: 1325: 1320: 1315: 1310: 1305: 1300: 1295: 1290: 1285: 1280: 1275: 1270: 1265: 1260: 1252: 1250:Founder effect 1247: 1242: 1237: 1232: 1227: 1222: 1217: 1211: 1209: 1206: 1127:Queen Victoria 1072:Main article: 1069: 1066: 1005: 1002: 989:Charles Darwin 972: 969: 941: 938: 920: 917: 913:founder effect 907: 904: 898: 895: 842:breeding (see 779: 776: 764: 763: 751: 741: 735: 725: 706: 703: 701: 698: 697: 696: 683: 670: 657: 644: 631: 618: 605: 557: 554: 519:heterozygosity 486: 483: 403:Due to higher 396: 395: 389: 382: 375: 365: 359: 353: 347: 336: 325: 284: 281: 248:immune systems 195: 192: 165:plant breeding 26: 9: 6: 4: 3: 2: 5451: 5440: 5437: 5435: 5432: 5430: 5427: 5425: 5422: 5421: 5419: 5404: 5401: 5399: 5396: 5394: 5391: 5390: 5388: 5384: 5377: 5375: 5372: 5370: 5367: 5365: 5364:Mongelli case 5362: 5360: 5357: 5355: 5352: 5350: 5347: 5345: 5342: 5340: 5337: 5335: 5332: 5330: 5327: 5325: 5322: 5321: 5319: 5315: 5305: 5302: 5300: 5297: 5295: 5292: 5290: 5287: 5286: 5284: 5280: 5274: 5271: 5269: 5266: 5264: 5261: 5259: 5256: 5254: 5250: 5247: 5246: 5244: 5240: 5237: 5233: 5227: 5224: 5222: 5219: 5217: 5214: 5212: 5209: 5205: 5202: 5200: 5197: 5196: 5195: 5192: 5190: 5187: 5185: 5182: 5181: 5179: 5175: 5169: 5166: 5164: 5161: 5159: 5158: 5157:Muth v. Frank 5154: 5152: 5151: 5147: 5141: 5138: 5137: 5136: 5133: 5132: 5131: 5128: 5126: 5125:Consanguinity 5123: 5121: 5118: 5117: 5115: 5113: 5109: 5101: 5098: 5097: 5096: 5093: 5091: 5090: 5089:Snokhachestvo 5086: 5084: 5081: 5079: 5076: 5074: 5073:Covert incest 5071: 5067: 5064: 5062: 5061:notable cases 5059: 5058: 5057: 5054: 5052: 5049: 5047: 5044: 5042: 5039: 5037: 5034: 5033: 5031: 5027: 5023: 5016: 5011: 5009: 5004: 5002: 4997: 4996: 4993: 4981: 4973: 4972: 4969: 4963: 4960: 4958: 4955: 4954: 4952: 4948: 4942: 4939: 4937: 4934: 4932: 4929: 4927: 4924: 4922: 4919: 4917: 4914: 4912: 4909: 4907: 4904: 4902: 4899: 4897: 4894: 4892: 4889: 4887: 4884: 4882: 4879: 4877: 4874: 4872: 4869: 4867: 4864: 4862: 4859: 4857: 4854: 4853: 4851: 4847: 4841: 4838: 4836: 4833: 4831: 4828: 4826: 4823: 4821: 4818: 4816: 4813: 4811: 4808: 4807: 4805: 4802: 4798: 4792: 4789: 4787: 4784: 4782: 4779: 4777: 4774: 4772: 4769: 4767: 4764: 4762: 4759: 4757: 4754: 4752: 4749: 4748: 4746: 4742: 4736: 4733: 4731: 4728: 4726: 4723: 4721: 4718: 4716: 4713: 4711: 4708: 4706: 4703: 4701: 4698: 4696: 4693: 4691: 4688: 4687: 4685: 4681: 4675: 4672: 4670: 4667: 4665: 4662: 4660: 4657: 4655: 4652: 4650: 4647: 4646: 4643: 4639: 4632: 4627: 4625: 4620: 4618: 4613: 4612: 4609: 4601: 4600: 4594: 4589: 4585: 4583: 4580: 4578: 4574: 4571: 4569: 4564: 4563: 4551: 4547: 4540: 4534: 4529: 4515:on 2010-12-27 4514: 4510: 4506: 4503:Seawright C. 4499: 4491: 4485: 4481: 4474: 4466: 4462: 4457: 4456:11577/3455561 4452: 4447: 4442: 4438: 4434: 4430: 4423: 4409: 4405: 4401: 4397: 4393: 4389: 4388:Human Biology 4385: 4378: 4370: 4366: 4361: 4356: 4352: 4348: 4344: 4337: 4329: 4325: 4321: 4317: 4313: 4309: 4302: 4294: 4290: 4285: 4280: 4276: 4272: 4269:(4): 517–23. 4268: 4264: 4260: 4253: 4245: 4241: 4237: 4233: 4229: 4225: 4221: 4217: 4213: 4209: 4202: 4194: 4190: 4185: 4180: 4176: 4172: 4168: 4161: 4153: 4149: 4145: 4141: 4137: 4133: 4130:(2): 111–30. 4129: 4125: 4118: 4116: 4107: 4103: 4098: 4093: 4088: 4083: 4079: 4075: 4072:(8): 3906–8. 4071: 4067: 4063: 4056: 4048: 4044: 4039: 4034: 4030: 4026: 4023:(1): 225–31. 4022: 4018: 4014: 4007: 4005: 3996: 3990: 3986: 3979: 3971: 3967: 3963: 3959: 3955: 3951: 3944: 3942: 3933: 3929: 3924: 3919: 3915: 3911: 3907: 3903: 3899: 3892: 3890: 3881: 3877: 3872: 3867: 3862: 3857: 3853: 3849: 3845: 3838: 3836: 3834: 3832: 3830: 3828: 3819: 3815: 3811: 3807: 3803: 3799: 3792: 3790: 3781: 3777: 3773: 3769: 3762: 3760: 3758: 3749: 3745: 3741: 3737: 3733: 3729: 3725: 3718: 3710: 3706: 3701: 3696: 3692: 3688: 3685:(3): 185–92. 3684: 3680: 3676: 3669: 3663: 3659: 3653: 3646: 3641: 3625: 3621: 3617: 3613: 3606: 3599: 3595: 3592: 3587: 3579: 3575: 3569: 3562: 3558: 3555: 3550: 3542: 3538: 3533: 3528: 3524: 3520: 3516: 3512: 3508: 3504: 3500: 3493: 3485: 3479: 3475: 3474: 3466: 3464: 3462: 3447: 3443: 3437: 3423: 3419: 3413: 3405: 3399: 3395: 3391: 3384: 3373: 3369: 3362: 3356: 3348: 3344: 3340: 3336: 3331: 3326: 3322: 3318: 3314: 3310: 3303: 3296: 3288: 3284: 3279: 3274: 3269: 3264: 3260: 3256: 3252: 3248: 3244: 3237: 3221: 3217: 3211: 3203: 3199: 3194: 3189: 3185: 3181: 3177: 3173: 3169: 3162: 3160: 3151: 3147: 3143: 3139: 3135: 3131: 3127: 3123: 3116: 3102:on 2010-03-27 3101: 3097: 3093: 3086: 3079: 3074: 3066: 3059: 3051: 3047: 3043: 3039: 3036:(5): 429–35. 3035: 3031: 3024: 3016: 3012: 3007: 3002: 2998: 2994: 2991:(3): 767–79. 2990: 2986: 2982: 2975: 2967: 2963: 2959: 2955: 2951: 2947: 2943: 2936: 2934: 2926: 2920: 2914: 2909: 2901: 2897: 2893: 2889: 2882: 2874: 2870: 2866: 2862: 2858: 2854: 2850: 2846: 2839: 2830: 2825: 2821: 2817: 2813: 2806: 2798: 2794: 2790: 2786: 2781: 2776: 2772: 2768: 2761: 2753: 2749: 2745: 2741: 2738:(5): 378–86. 2737: 2733: 2726: 2724: 2712: 2710: 2701: 2694: 2688: 2680: 2676: 2672: 2668: 2661: 2654: 2646: 2642: 2638: 2634: 2630: 2626: 2622: 2618: 2611: 2603: 2599: 2595: 2591: 2587: 2583: 2579: 2572: 2564: 2560: 2556: 2552: 2549:(4): 283–92. 2548: 2544: 2537: 2529: 2525: 2520: 2515: 2511: 2507: 2503: 2496: 2488: 2484: 2480: 2476: 2472: 2468: 2465:(6): 784–95. 2464: 2460: 2453: 2445: 2441: 2436: 2431: 2426: 2421: 2417: 2413: 2409: 2405: 2398: 2396: 2387: 2381: 2377: 2370: 2362: 2356: 2352: 2345: 2343: 2334: 2330: 2325: 2320: 2316: 2312: 2308: 2304: 2300: 2293: 2286: 2285:0-7637-1511-5 2282: 2276: 2268: 2264: 2259: 2254: 2249: 2244: 2240: 2236: 2232: 2225: 2217: 2211: 2207: 2200: 2192: 2188: 2184: 2180: 2176: 2172: 2165: 2163: 2154: 2150: 2145: 2140: 2136: 2132: 2128: 2124: 2120: 2113: 2105: 2101: 2097: 2093: 2089: 2085: 2081: 2077: 2070: 2062: 2058: 2053: 2048: 2043: 2038: 2034: 2030: 2027:(7): 3324–9. 2026: 2022: 2018: 2011: 2002: 1994: 1988: 1984: 1980: 1979: 1971: 1963: 1959: 1955: 1951: 1947: 1943: 1936: 1928: 1924: 1919: 1914: 1910: 1906: 1903:(1): 161–72. 1902: 1898: 1894: 1887: 1879: 1875: 1871: 1867: 1863: 1859: 1855: 1851: 1844: 1836: 1832: 1828: 1824: 1820: 1816: 1813:(2): 317–23. 1812: 1808: 1801: 1793: 1789: 1784: 1779: 1775: 1771: 1768:(3): 327–32. 1767: 1763: 1759: 1752: 1744: 1740: 1735: 1730: 1726: 1722: 1719:(2): 767–79. 1718: 1714: 1710: 1703: 1695: 1691: 1687: 1683: 1678: 1673: 1669: 1665: 1661: 1654: 1648: 1647:0-201-40754-X 1644: 1638: 1636: 1627: 1623: 1619: 1615: 1611: 1607: 1603: 1599: 1592: 1590: 1580: 1575: 1571: 1567: 1563: 1561: 1552: 1544: 1540: 1536: 1532: 1528: 1524: 1520: 1516: 1512: 1505: 1497: 1493: 1489: 1485: 1481: 1477: 1473: 1469: 1462: 1454: 1450: 1446: 1442: 1438: 1434: 1430: 1426: 1422: 1418: 1414: 1410: 1406: 1399: 1397: 1390: 1389: 1384: 1379: 1375: 1364: 1361: 1359: 1356: 1354: 1351: 1349: 1346: 1344: 1341: 1339: 1336: 1334: 1331: 1329: 1326: 1324: 1321: 1319: 1316: 1314: 1311: 1309: 1306: 1304: 1301: 1299: 1296: 1294: 1291: 1289: 1286: 1284: 1281: 1279: 1276: 1274: 1271: 1269: 1266: 1264: 1261: 1259: 1257: 1253: 1251: 1248: 1246: 1243: 1241: 1238: 1236: 1233: 1231: 1228: 1226: 1225:Consanguinity 1223: 1221: 1218: 1216: 1213: 1212: 1205: 1203: 1202:ancient Egypt 1199: 1195: 1194:Cleopatra VII 1191: 1187: 1183: 1179: 1175: 1171: 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