Outcrossing

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Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity.

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Outcrossing in animals

Outcrossing can be a useful technique in animal breeding. The outcrossing breeder intends to remove the traits by using "new blood." With dominant traits, one can still see the expression of the traits and can remove those traits whether one outcrosses, line breeds or inbreeds. With recessive traits, outcrossing allows for the recessive traits to migrate across a population. Many traits are Mendelian and therefore exhibit a more complicated intermediate phenotype. The outcrossing breeder then may have individuals that have many deleterious genes that may be expressed by subsequent inbreeding. There is now a gamut of deleterious genes within each individual in many dog breeds. [1]

Increasing the variation of genes or alleles within the gene pool may protect against extinction by stressors from the environment among inbred animal populations. For example, in this context, a recent veterinary medicine study tried to determine the genetic diversity within cat breeds. [2]

A degree of outcrossing to avoid mating between very close relatives is believed to happen in the wild. [1]

Outcrossing in plants and fungi

Outcrossing in plants is usually enforced by self-incompatibility. The primary adaptive function of flowers is the facilitation of outcrossing, a process that allows the masking of deleterious mutations in the genome of progeny. The masking effect of outcrossing is known as “genetic complementation”, [3] an effect also recognized as hybrid vigor or heterosis. Once outcrossing is established in a lineage of flowering plants due to the benefit of genetic complementation, subsequent switching to inbreeding becomes disadvantageous because it allows expression of the previously masked deleterious recessive mutations, i.e. inbreeding depression.

Outcrossing in fungi involves syngamy between haploid cells produced by separate diploid individuals. [4]

Life-history traits are said to increase the probability of outcrossing in fungi such as long-distance dispersal and persistence of the haploid stage. Some studies even show that fungi favor outcrossing in comparison to other mating types. In a study performed with the commercial button mushroom, Agaricus bisporus, outcrossed populations of the fungi showed higher fitness than inbred ones in several fitness components. [5]

General practice

Breeders inbreed within their genetic pool, attempting to maintain desirable traits and to cull those traits that are undesirable. When undesirable traits begin to appear, mates are selected to determine if a trait is recessive or dominant. Removal of the trait is accomplished by breeding two individuals known not to carry it. [6]

Gregor Mendel used outcrossing in his experiments with flowers. He then used the resulting offspring to chart inheritance patterns, using the crossing of siblings, and backcrossing to parents to determine how inheritance functioned. [7]

Darwin's perspective

Charles Darwin, in his book The Effects of Cross and Self-Fertilization in the Vegetable Kingdom,. [8] :462 stated regarding outcrossing that "the offspring from the union of two distinct individuals, especially if their progenitors have been subjected to very different conditions, have an immense advantage in height, weight, constitutional vigor and fertility over the self-fertilizing offspring from either one of the same parents". He thought that this observation was amply sufficient to account for outcrossing sexual reproduction. The disadvantages of self-fertilized offspring (inbreeding depression) are now thought to be largely due to the homozygous expression of deleterious recessive mutations; [9] and the fitness advantages of some outcrossed offspring are thought to be largely due to the heterozygous masking of such deleterious mutations except when such mutations lead to outbreeding depression.

See also

Related Research Articles

<span class="mw-page-title-main">Inbreeding</span> Reproduction by closely related organisms

Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. 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 recessive traits resulting from incestuous sexual relationships and consanguinity. Animals avoid incest only rarely.

<span class="mw-page-title-main">Self-pollination</span> Form of

Self-pollination is a form of pollination in which pollen from the same plant arrives at the stigma of a flower or at the ovule. There are two types of self-pollination: in autogamy, pollen is transferred to the stigma of the same flower; in geitonogamy, pollen is transferred from the anther of one flower to the stigma of another flower on the same flowering plant, or from microsporangium to ovule within a single (monoecious) gymnosperm. Some plants have mechanisms that ensure autogamy, such as flowers that do not open (cleistogamy), or stamens that move to come into contact with the stigma. The term selfing that is often used as a synonym, is not limited to self-pollination, but also applies to other type of self-fertilization.

<span class="mw-page-title-main">Dog breeding</span> Mating selected dogs for specific qualities

Dog breeding is the practice of mating selected dogs with the intention of maintaining or producing specific qualities and characteristics. When dogs reproduce without such human intervention, their offspring's characteristics are determined by natural selection, while "dog breeding" refers specifically to the artificial selection of dogs, in which dogs are intentionally bred by their owners. Breeding relies on the science of genetics, hence a breeder who is knowledgeable on canine genetics, health, and the intended purpose of the dogs attempts to breed suitable dogs.

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. The heterotic offspring often has traits that are more than the simple addition of the parents' traits, and can be explained by Mendelian or non-Mendelian inheritance. Typical heterotic/hybrid traits of interest in agriculture are higher yield, quicker maturity, stability, drought tolerance etc.

<span class="mw-page-title-main">Evolution of sexual reproduction</span> How sexually reproducing multicellular organisms could have evolved from a common ancestor species

Sexual reproduction is an adaptive feature which is common to almost all multicellular organisms and various unicellular organisms. Currently, the adaptive advantage of sexual reproduction is widely regarded as a major unsolved problem in biology. As discussed below, one prominent theory is that sex evolved as an efficient mechanism for producing variation, and this had the advantage of enabling organisms to adapt to changing environments. Another prominent theory, also discussed below, is that a primary advantage of outcrossing sex is the masking of the expression of deleterious mutations. Additional theories concerning the adaptive advantage of sex are also discussed below. Sex does, however, come with a cost. In reproducing asexually, no time nor energy needs to be expended in choosing a mate and, if the environment has not changed, then there may be little reason for variation, as the organism may already be well-adapted. However, very few environments have not changed over the millions of years that reproduction has existed. Hence it is easy to imagine that being able to adapt to changing environment imparts a benefit. Sex also halves the amount of offspring a given population is able to produce. Sex, however, has evolved as the most prolific means of species branching into the tree of life. Diversification into the phylogenetic tree happens much more rapidly via sexual reproduction than it does by way of asexual reproduction.

A crossbreed is an organism with purebred parents of two different breeds, varieties, or populations. Crossbreeding, sometimes called "designer crossbreeding", is the process of breeding such an organism. While crossbreeding is used to maintain health and viability of organisms, irresponsible crossbreeding can also produce organisms of inferior quality or dilute a purebred gene pool to the point of extinction of a given breed of organism.

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 the term F1 crossbreed may be used. The term is sometimes written with a subscript, as F1 hybrid. Subsequent generations are called F2, F3, etc.

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.

Complementation refers to a genetic process when two strains of an organism with different homozygous recessive mutations that produce the same mutant phenotype have offspring that express the wild-type phenotype when mated or crossed. Complementation will ordinarily occur if the mutations are in different genes. Complementation may also occur if the two mutations are at different sites within the same gene, but this effect is usually weaker than that of intergenic complementation. When the mutations are in different genes, each strain's genome supplies the wild-type allele to "complement" the mutated allele of the other strain's genome. Since the mutations are recessive, the offspring will display the wild-type phenotype. A complementation test can test whether the mutations in two strains are in different genes. Complementation is usually weaker or absent if the mutations are in the same gene. The convenience and essence of this test is that the mutations that produce a phenotype can be assigned to different genes without the exact knowledge of what the gene product is doing on a molecular level. American geneticist Edward B. Lewis developed the complementation test.

Inbreeding depression is the reduced biological fitness that has the potential to result from inbreeding. The loss of genetic diversity that is seen due to inbreeding, results from small population size. 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, though inbreeding and outbreeding depression can simultaneously occur.

In biology, outbreeding depression happens when crosses between two genetically distant groups or populations result in a reduction of fitness. The concept is in contrast to inbreeding depression, although the two effects can occur simultaneously. Outbreeding depression is a risk that sometimes limits the potential for genetic rescue or augmentations. It is considered postzygotic response because outbreeding depression is noted usually in the performance of the progeny.

Allogamy or cross-fertilization is the fertilization of an ovum from one individual with the spermatozoa of another. By contrast, autogamy is the term used for self-fertilization. In humans, the fertilization event is an instance of allogamy. Self-fertilization occurs in hermaphroditic organisms where the two gametes fused in fertilization come from the same individual. This is common in plants and certain protozoans.

<i>Ciona</i> Genus of tunicate

Ciona is a genus of sea squirts in the family Cionidae.

<span class="mw-page-title-main">Mating in fungi</span> Combination of genetic material between compatible mating types

Fungi are a diverse group of organisms that employ a huge variety of reproductive strategies, ranging from fully asexual to almost exclusively sexual species. Most species can reproduce both sexually and asexually, alternating between haploid and diploid forms. This contrasts with most multicellular eukaryotes such as mammals, where the adults are usually diploid and produce haploid gametes which combine to form the next generation. In fungi, both haploid and diploid forms can reproduce – haploid individuals can undergo asexual reproduction while diploid forms can produce gametes that combine to give rise to the next generation.

<span class="mw-page-title-main">Plant evolution</span> Subset of evolutionary phenomena that concern plants

Plant evolution is the subset of evolutionary phenomena that concern plants. Evolutionary phenomena are characteristics of populations that are described by averages, medians, distributions, and other statistical methods. This distinguishes plant evolution from plant development, a branch of developmental biology which concerns the changes that individuals go through in their lives. The study of plant evolution attempts to explain how the present diversity of plants arose over geologic time. It includes the study of genetic change and the consequent variation that often results in speciation, one of the most important types of radiation into taxonomic groups called clades. A description of radiation is called a phylogeny and is often represented by type of diagram called a phylogenetic tree.

<span class="mw-page-title-main">Popular sire effect</span>

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. However, the popular sire effect is not just down to wanting to produce a champion. For example, in Staffordshire Bull Terriers there are several popular sires who are used by breeders to produce specific colours that are not favoured in the show ring. 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.

Genetic purging is the increased pressure of natural selection against deleterious alleles prompted by inbreeding.

Inbreeding avoidance, or the inbreeding avoidance hypothesis, is a concept in evolutionary biology that refers to the prevention of the deleterious effects of inbreeding. Animals only rarely exhibit inbreeding avoidance. 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 and natural and sexual selection, in order to prevent breeding among related individuals. 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 such mechanisms.

Inbreeding in fish is the mating of closely related individuals, leading to an increase in homozygosity. Repeated inbreeding generally leads to morphological abnormalities and a reduction in fitness in the offspring. In the wild, fish have a number of ways to avoid inbreeding, both before and after copulation.

Autogamy or self-fertilization refers to the fusion of two gametes that come from one individual. Autogamy is predominantly observed in the form of self-pollination, a reproductive mechanism employed by many flowering plants. However, species of protists have also been observed using autogamy as a means of reproduction. Flowering plants engage in autogamy regularly, while the protists that engage in autogamy only do so in stressful environments.

References

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