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Genic capture is a hypothesis explaining the maintenance of genetic variance in traits under sexual selection. A classic problem in sexual selection is the fixation of alleles that are beneficial under strong selection, thereby eliminating the benefits of mate choice. Genic capture resolves this paradox by suggesting that additive genetic variance of sexually selected traits reflects the genetic variance in total condition. [1] A deleterious mutation anywhere in the genome will adversely affect condition, and thereby adversely affect a condition-dependent sexually selected trait. Genic capture therefore resolves the lek paradox by proposing that recurrent deleterious mutation maintains additive genetic variance in fitness by incorporating the entire mutation load of an individual. Thus any condition-dependent trait "captures" the overall genetic variance in condition. Rowe and Houle argued that genic capture ensures that good genes will become a central feature of the evolution of any sexually selected trait.
The key quantity for genic capture is vaguely defined as "condition." The hypothesis only defines condition as a quantity that correlates tightly with overall fitness, such that directional selection will always increase average condition over time. Condition should, in general, reflect overall energy acquisition, such that life-history variation reflects differential allocation to survival and sexual signalling. Genetic variation in condition should be very broadly affected by any changes in the genome. Close to equilibrium any mutation should be deleterious, thereby leading to non-zero overall mutation rate, maintaining variance in fitness.
Rowe and Houle's simple model defines a trait as the result of three heritable components, a condition-independent component , epistatic modification and condition, suggesting the following function for a trait:
where is the condition of an individual. Loci contributing to are loosely linked and independent of loci contributing to and . Rowe and Houle then find the expected variance of and ignored higher-order terms (i.e. products of variances):
where represents the genetic variance in the signal and analogously for other traits. Under directional selection on , the loci underlying and may lose all genetic variance. However, there is no qualitative difference in directional selection on between stabilizing selection (i.e. no sexual selection) and directional selection on . Therefore, the second term will remain positive (due to biased mutation) and dominate under sexual selection.
Genic capture can also play a role in accelerating adaptation to new environments. [2]
Genic capture was proposed as a simpler alternative to another theory explaining the lek paradox [3] that proposed that sexual selection creates disruptive selection, i.e. positive selection for genetic variance. Genic capture does not require any particular fitness function.
Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Charles Darwin popularised the term "natural selection", contrasting it with artificial selection, which in his view is intentional, whereas natural selection is not.
Small populations can behave differently from larger populations. They are often the result of population bottlenecks from larger populations, leading to loss of heterozygosity and reduced genetic diversity and loss or fixation of alleles and shifts in allele frequencies. A small population is then more susceptible to demographic and genetic stochastic events, which can impact the long-term survival of the population. Therefore, small populations are often considered at risk of endangerment or extinction, and are often of conservation concern.
Heritability is a statistic used in the fields of breeding and genetics that estimates the degree of variation in a phenotypic trait in a population that is due to genetic variation between individuals in that population. It measures how much of the variation of a trait can be attributed to variation of genetic factors, as opposed to variation of environmental factors. The concept of heritability can be expressed in the form of the following question: "What is the proportion of the variation in a given trait within a population that is not explained by the environment or random chance?"
Fitness is the quantitative representation of natural and sexual selection within evolutionary biology. It can be defined either with respect to a genotype or to a phenotype in a given environment. In either case, it describes individual reproductive success and is equal to the average contribution to the gene pool of the next generation that is made by individuals of the specified genotype or phenotype. The fitness of a genotype is manifested through its phenotype, which is also affected by the developmental environment. The fitness of a given phenotype can also be different in different selective environments.
Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and population structure.
A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals, known as lekking, to entice visiting females which are surveying prospective partners to mate with. A lek can also indicate an available plot of space able to be utilized by displaying males to defend their own share of territory for the breeding season. A lekking species is characterised by male displays, strong female mate choice, and the conferring of indirect benefits to males and reduced costs to females. Although most prevalent among birds such as black grouse, lekking is also found in a wide range of vertebrates including some bony fish, amphibians, reptiles, and mammals, and arthropods including crustaceans and insects.
Genetic variation is the difference in DNA among individuals or the differences between populations. There are multiple sources of genetic variation, including mutation and genetic recombination. The mutation is the ultimate source of genetic variation, but mechanisms such as sexual reproduction and genetic drift contribute to it as well.
Pleiotropy occurs when one gene influences two or more seemingly unrelated phenotypic traits. Such a gene that exhibits multiple phenotypic expression is called a pleiotropic gene. Mutation in a pleiotropic gene may have an effect on several traits simultaneously, due to the gene coding for a product used by a myriad of cells or different targets that have the same signaling function.
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.
Inbreeding depression is the reduced biological fitness in a given population as a result of inbreeding, or breeding of related individuals. Population biological fitness refers to an organism's ability to survive and perpetuate its genetic material. Inbreeding depression is often the result of a population bottleneck. In general, the higher the genetic variation or gene pool within a breeding population, the less likely it is to suffer from inbreeding depression.
Stalk-eyed flies are insects of the fly family Diopsidae. The family is distinguished from most other flies by the possession of "eyestalks": projections from the sides of the head with the eyes at the end. Some fly species from other families such as Drosophilidae, Platystomatidae, Richardiidae, and Tephritidae have similar heads, but the unique character of the Diopsidae is that their antennae are located on the stalk, rather than in the middle of the head as in all other flies.
Genetic hitchhiking, also called genetic draft or the hitchhiking effect, is when an allele changes frequency not because it itself is under natural selection, but because it is near another gene that is undergoing a selective sweep and that is on the same DNA chain. When one gene goes through a selective sweep, any other nearby polymorphisms that are in linkage disequilibrium will tend to change their allele frequencies too. Selective sweeps happen when newly appeared mutations are advantageous and increase in frequency. Neutral or even slightly deleterious alleles that happen to be close by on the chromosome 'hitchhike' along with the sweep. In contrast, effects on a neutral locus due to linkage disequilibrium with newly appeared deleterious mutations are called background selection. Both genetic hitchhiking and background selection are stochastic (random) evolutionary forces, like genetic drift.
Enquiry into the evolution of ageing aims to explain why a detrimental process such as aging would evolve, and why there is so much variability in the lifespans of living organisms. The classical theories of evolution suggest that environmental factors such as predation, accidents, disease, starvation, etc. ensure that most organisms living in natural settings will not live until old age, and so there will be very little pressure to conserve genetic changes that increase longevity. Natural selection will instead strongly favor genes which ensure early maturation and rapid reproduction, and the selection for genetic traits which promote molecular and cellular self-maintenance will decline with age for most organisms.
Background selection describes the loss of genetic diversity at a non-deleterious locus due to negative selection against linked deleterious alleles. It is one form of linked selection, where the maintenance or removal of an allele from a population is dependent upon the alleles in its linkage group. The name emphasizes the fact that the genetic background, or genomic environment, of a neutral mutation has a significant impact on whether it will be preserved or purged from a population. In some cases, the term background selection is used broadly to refer to all forms of linked selection, but most often it is used only when neutral variation is reduced due to negative selection against deleterious mutations. Background selection and all forms of linked selection contradict the assumption of the neutral theory of molecular evolution that the fixation or loss of neutral alleles is entirely stochastic, the result of genetic drift. Instead, these models predict that neutral variation is correlated with the selective pressures acting on linked non-neutral genes, that neutral traits are not necessarily oblivious to selection. Because they segregate together, non-neutral mutations linked to neutral polymorphisms result in decreased levels of genetic variation relative to predictions of neutral evolution.
The lek paradox is the conundrum of how additive or beneficial genetic variation is maintained in lek mating species, in the face of consistent female preferences, sexual selection. While many studies have attempted to explain how the lek paradox fits into Darwinian theory, the paradox remains. Persistent female choice for particular male trait values should erode genetic diversity in male traits and thereby remove the benefits of choice, yet choice persists. This paradox can be somewhat alleviated by the occurrence of mutations introducing potential differences, as well as the possibility that traits of interest have more or less favorable recessive alleles.
The nearly neutral theory of molecular evolution is a modification of the neutral theory of molecular evolution that accounts for the fact that not all mutations are either so deleterious such that they can be ignored, or else neutral. Slightly deleterious mutations are reliably purged only when their selection coefficient are greater than one divided by the effective population size. In larger populations, a higher proportion of mutations exceed this threshold for which genetic drift cannot overpower selection, leading to fewer fixation events and so slower molecular evolution.
The antagonistic pleiotropy hypothesis was first proposed by George C. Williams in 1957 as an evolutionary explanation for senescence. Pleiotropy is the phenomenon where one gene controls for more than one phenotypic trait in an organism. Antagonistic pleiotropy is when one gene controls for more than one trait, where at least one of these traits is beneficial to the organism's fitness early on in life and at least one is detrimental to the organism's fitness later on due to a decline in the force of natural selection. The theme of G.C. William's idea about antagonistic pleiotropy was that if a gene caused both increased reproduction in early life and aging in later life, then senescence would be adaptive in evolution. For example, one study suggests that since follicular depletion in human females causes both more regular cycles in early life and loss of fertility later in life through menopause, it can be selected for by having its early benefits outweigh its late costs.
Genetic purging is the reduction of the frequency of a deleterious allele, caused by an increased efficiency of natural selection prompted by inbreeding.
Peter D. Keightley FRS is Professor of Evolutionary Genetics at the Institute of Evolutionary Biology in School of Biological Sciences at the University of Edinburgh.
Epistasis is a phenomenon in genetics in which the effect of a gene mutation is dependent on the presence or absence of mutations in one or more other genes, respectively termed modifier genes. In other words, the effect of the mutation is dependent on the genetic background in which it appears. Epistatic mutations therefore have different effects on their own than when they occur together. Originally, the term epistasis specifically meant that the effect of a gene variant is masked by that of a different gene.