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Reinforcement (speciation)

Reinforcement is a process of speciation where natural selection increases the reproductive isolation between two populations of species. This occurs as a result of selection acting against the production of hybrid individuals of low fitness. The idea was originally developed by Alfred Russel Wallace and is sometimes referred to as the Wallace effect. The modern concept of reinforcement originates from Theodosius Dobzhansky. He envisioned a species separated allopatrically, where during secondary contact the two populations mate, producing hybrids with lower fitness. Natural selection results from the hybrid's inability to produce viable offspring; thus members of one species who do not mate with members of the other have greater reproductive success. This favors the evolution of greater prezygotic isolation. Reinforcement is one of the few cases in which selection can favor an increase in prezygotic isolation, influencing the process of speciation directly. This aspect has been particularly appealing among evolutionary biologists.

History
Reinforcement has had a complex history in that its popularity among scholars has changed over time. Jerry Coyne and H. Allen Orr contend that the theory of reinforcement went through three phases of historical development: His hypothesis differed markedly from the modern conception in that it focused on post-zygotic isolation, strengthened by group selection. In 1930, Ronald Fisher laid out the first genetic description of the process of reinforcement in The Genetical Theory of Natural Selection, and in 1965 and 1970 the first computer simulations were run to test for its plausibility. and quantitative genetic studies were conducted showing that completely unfit hybrids lead unequivocally to an increase in prezygotic isolation. It was first used to describe the observed mating call differences in Gastrophryne frogs within a secondary contact hybrid zone. The Wallace effect is similar to reinforcement, but is rarely used. Daniel J. Howard considered reproductive character displacement to represent either assortive mating or the divergence of traits for mate recognition (specifically between sympatric populations). Servedio and Noor include any detected increase in prezygotic isolation as reinforcement, as long as it is a response to selection against mating between two different species. Coyne and Orr contend that, "true reinforcement is restricted to cases in which isolation is enhanced between taxa that can still exchange genes". == Models ==
Models
s form in the zone of contact 4. Genome recombination results in speciation of the two populations, with an additional hybrid species. All three species are separated by intrinsic reproductive barriers One of the strongest forms of reproductive isolation in nature is sexual isolation: traits in organisms involving mating. This pattern has led to the idea that, because selection acts so strongly on mating traits, it may be involved in the process of speciation.). It necessitates two forces of evolution that act on mate choice: natural selection and gene flow. Selection acts as the main driver of reinforcement as it selects against hybrid genotypes that are of low-fitness, regardless if individual preferences have no effect on survival and reproduction. A common signature of reinforcement's occurrence in nature is that of reproductive character displacement; characteristics of a population diverge in sympatry but not allopatry. Further, gene flow can diminish the isolation found in sympatric populations. In instances of peripatric speciation, reinforcement is unlikely to complete speciation in the case that the peripherally isolated population comes into secondary contact with the main population. Genetics The underlying genetics of reinforcement can be understood by an ideal model of two haploid populations experiencing an increase in linkage disequilibrium. Here, selection rejects low fitness Bc or bC allele combinations while favoring combinations of BC alleles (in the first subpopulation) and bc alleles (in the second subpopulation). The third locus A or a (the assortive mating alleles) have an effect on mating pattern but is not under direct selection. If selection at B and C cause changes in the frequency of allele A, assortive mating is promoted, resulting in reinforcement. Both selection and assortive mating are necessary, that is, that matings of A \times A and a \times a are more common than matings of a \times A and A \times a. A restriction of migration between populations can further increase the chance of reinforcement, as it decreases the probability of the differing genotypes to exchange. with some relying on one locus per trait and others on polygenic traits. Population structures The structure and migration patterns of a population can affect the process of speciation by reinforcement. It has been shown to occur under an island model, harboring conditions with infrequent migrations occurring in one direction, . In direct selection, the frequency of the selected allele is favored to the extreme. In cases where an allele is indirectly selected, its frequency increases due to a different linked allele experiencing selection (linkage disequilibrium). Any traits that promote isolation may be subjected to reinforcement such as mating signals (e.g. courtship display), signal responses, the location of breeding grounds, the timing of mating (e.g. seasonal breeding such as in allochronic speciation), or even egg receptivity. Many of these examples are described below. == Evidence ==
Evidence
, divergence is exhibited by changes in mating traits. These patterns of reproductive character displacement detected in species populations that exist in zones of overlap indicate that the process of speciation by reinforcement has occurred. The evidence for reinforcement comes from observations in nature, comparative studies, and laboratory experiments. Assortive mating is expected to increase among sympatric populations experiencing reinforcement. Laboratory experiments Laboratory studies that explicitly test for reinforcement are limited, == Alternative hypotheses ==
Alternative hypotheses
Various alternative explanations for the patterns observed in nature have been proposed. and that of allelic dominance: any of the alleles experiencing selection for isolation should be dominate. Species interactions can also result in reproductive character displacement (in both mate preference or mating signal). Those that have weaker prezygotic isolation will eventually fuse, losing their distinctiveness. Furthermore, patterns detected in Drosophila find high levels of prezygotic isolation in sympatry but not in allopatry. The fusion hypothesis predicts that strong isolation should be found in both allopatry and sympatry. Two other ways to determine if reinforcement occurs (as opposed to sympatric speciation) are: • if two recently speciated taxa do not show signs of post-zygotic isolation of both sympatric and allopatric populations (in sympatric speciation, post-zygotic isolation is not a prerequisite); • if a cline exists between two species over a range of traits (sympatric speciation does not require a cline to exist at all). Sexual selection In a runaway process (not unlike Fisherian runaway selection), selection against the low-fitness hybrids favors assortive mating, increasing mate discrimination rapidly. Post-zygotic isolation is not needed, initiated simply by the fact that unfit hybrids cannot get mates. == Arguments against reinforcement ==
Arguments against reinforcement
A number of objections were put forth, mainly during the 1980s, arguing that reinforcement is implausible. However, if they are selectively advantageous, gene flow should allow the alleles to spread throughout both populations. For example, the two species Drosophila santomea and D. yakuba on the African island São Tomé occasionally hybridize with one another, resulting in fertile female offspring and sterile male offspring. This effect would result in the extinction of one of the populations. This objection is overcome by when both populations are not subject to the same ecological conditions. For reinforcement to occur, prezygotic isolation must happen quickly. == References ==
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