sexual selection
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Sexual selection is a mode of in which members of one of the other to with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (intrasexual selection). These two forms of selection mean that some individuals have greater than others within a , for example because they are more or prefer more attractive partners to produce . Successful males benefit from frequent mating and monopolizing access to one or more fertile females. Females can have a limited number of offspring and maximize the return on the energy they invest in reproduction. The concept was first articulated by who wrote of a "second agency" of selection, in which competition between mate candidates could lead to speciation. The theory was given a mathematical basis by in the early 20th century. Sexual selection can lead males to extreme efforts to demonstrate their to be chosen by females, producing in s, such as the ornate of such as and , or the antlers of , or the manes of s, caused by a mechanism known as a , where the passing-on of the desire for a trait in one sex is as important as having the trait in the other sex in producing the runaway effect. Although the indicates that females would prefer male offspring, explains why the is most often 1:1. Sexual selection is also found in and .


History


Darwin

Sexual selection was first proposed by Charles Darwin in ' (1859) and developed in ' (1871), as he felt that natural selection alone was unable to account for certain types of non-survival adaptations. He once wrote to a colleague that "The sight of a feather in a 's tail, whenever I gaze at it, makes me sick!" His work divided sexual selection into male-male competition and female choice. These views were to some extent opposed by , mostly after Darwin's death. He accepted that sexual selection could occur, but argued that it was a relatively weak form of selection. He argued that male-male competitions were forms of natural selection, but that the "drab" peahen's coloration is itself adaptive as . In his opinion, ascribing mate choice to females was attributing the ability to judge standards of beauty to animals (such as s) far too cognitively undeveloped to be capable of feeling.


Ronald Fisher

, the and developed a number of ideas about sexual selection in his 1930 book ' including the and . The describes how sexual selection accelerates the preference for a specific ornament, causing the preferred trait and female preference for it to increase together in a runaway cycle. In a remark that was not widely understood for another 50 years he said: This causes a dramatic increase in both the male's conspicuous feature and in female preference for it, resulting in marked , until practical physical constraints halt further exaggeration. A loop is created, producing extravagant physical structures in the non-limiting sex. A classic example of female choice and potential runaway selection is the . While males have long tails that are selected for by female choice, female tastes in tail length are still more extreme with females being attracted to tails longer than those that naturally occur. Fisher understood that female preference for long tails may be passed on genetically, in conjunction with genes for the long tail itself. Long-tailed widowbird offspring of both sexes inherit both sets of genes, with females their genetic preference for long tails, and males showing off the coveted long tail itself. presents a non-mathematical explanation of the runaway sexual selection process in his book '. Females that prefer long tailed males tend to have mothers that chose long-tailed fathers. As a result, they carry both sets of genes in their bodies. That is, genes for long tails and for preferring long tails become . The taste for long tails and tail length itself may therefore become correlated, tending to increase together. The more tails lengthen, the more long tails are desired. Any slight initial imbalance between taste and tails may set off an explosion in tail lengths. Fisher wrote that: The female widowbird chooses to mate with the most attractive long-tailed male so that her progeny, if male, will themselves be attractive to females of the next generation—thereby fathering many offspring that carry the female's genes. Since the rate of change in preference is proportional to the average taste amongst females, and as females desire to secure the services of the most sexually attractive males, an additive effect is created that, if unchecked, can yield exponential increases in a given taste and in the corresponding desired sexual attribute.


After Fisher

Since Fisher's initial conceptual model of the 'runaway' process, and Peter O'Donald have provided detailed mathematical proofs that define the circumstances under which runaway sexual selection can take place. Alongside this, biologists have extended Darwin's formulation; Malte Andersson's widely-accepted 1994 definition is that "sexual selection is the differences in reproduction that arise from variation among individuals in traits that affect success in competition over mates and fertilizations". Despite some practical challenges for biologists, the concept of sexual selection is "straightforward".


Theory


Reproductive success

The of an organism is measured by the number of left behind, and their quality or probable . Sexual preference creates a tendency towards or . The general conditions of sexual discrimination appear to be (1) the acceptance of one mate precludes the effective acceptance of alternative mates, and (2) the rejection of an offer is followed by other offers, either certainly or at such high chance that the risk of non-occurrence is smaller than the chance advantage to be gained by selecting a mate. states that the sex which invests the most in producing offspring becomes a limiting resource for which the other sex competes, illustrated by the greater of an egg in a , and the limited capacity of females to reproduce; for example, in humans, a woman can only give birth every ten months, whereas a male can become a father numerous times in the same period. More recently, researchers have doubted whether Bateman was correct.


Modern interpretation

Darwin's ideas on sexual selection were met with scepticism by his contemporaries and not considered of great importance until in the 1930s biologists decided to include sexual selection as a mode of natural selection. Only in the 21st century have they become more important in ; the theory is now seen as generally applicable and analogous to natural selection. A ten-year study, experimentally varying sexual selection on s with other factors held constant, showed that sexual selection protected even an population against extinction. The of , and , holds that the male's survival until and through the age of reproduction with seemingly maladaptive traits is taken by the female as of his overall fitness. Such handicaps might prove he is either free of or resistant to , or that he possesses more speed or a greater physical strength that is used to combat the troubles brought on by the exaggerated trait. Zahavi's work spurred a re-examination of the field and several new theories. In 1984, Hamilton and introduced the "Bright Male" hypothesis, suggesting that male elaborations might serve as a marker of health, by exaggerating the effects of disease and deficiency. In 1990, Michael Ryan and A.S. Rand, working with the , proposed the hypothesis of "Sensory Exploitation", where exaggerated male traits may provide a sensory stimulation that females find hard to resist. In the late 1970s, Janzen and Mary Willson, noting that male flowers are often larger than female flowers, expanded the field of . More recently, the field has grown to include other areas of study, not all of which fit Darwin's definition of sexual selection. A "bewildering" range of models variously attempt to relate sexual selection not only to the fundamental questions of anisogamy and parental roles, but also to mechanisms such as sex ratios, parental care, , , and the "most-debated effect", namely . Elaborated characteristics that might seem costly for their bearers (e.g., the of the swordfish ') do not always have an energetics, performance or even survival cost; this may be because "compensatory traits" have evolved in concert with the sexually selected traits. File:Susa group, mountain gorilla.jpg, Male , a tournament species File:Tribolium castaneum87-300.jpg, Sexual selection protected s from extinction in a ten-year experiment. File:Tungara frog (Physalaemus pustulosus).jpg, exemplified theory of "Sensory Exploitation", where exaggerated male traits supposedly made males irresistible to females.


Toolkit of natural selection

Sexual selection may explain how characteristics such as feathers had survival value at an early stage in their evolution. proposes that the feathers of proto-birds like ' were originally sexual ornaments. The earliest proto-birds such as ' had well-developed feathers but no sign of the top/bottom asymmetry that gives wings lift. One proposal is that the feathers served as insulation, helping females incubate their eggs. But if proto-bird courtship combined displays of forelimb feathers with energetic jumps, then the transition from display to aerodynamic functions could have been relatively smooth. Sexual selection sometimes generates features that may help cause a species' extinction, as has been suggestedMiller, Geoffrey (2000). The Mating Mind. Anchor Books. for the giant antlers of the (''Megaloceros giganteus'') that became extinct in Europe. Or it may do the opposite, driving species divergence—sometimes through elaborate changes in —such that new species emerge.


Sexual dimorphism

Sex differences directly related to reproduction and serving no direct purpose in are called . Traits amenable to sexual selection, which give an organism an advantage over its rivals (such as in courtship) without being directly involved in , are called s. In most sexual species the and have different strategies, due to a difference in relative investment in producing offspring. As formulated in Bateman's principle, females have a greater initial investment in producing offspring ( in mammals or the production of the egg in and ), and this difference in initial investment creates differences in variance in expected reproductive success and bootstraps the sexual selection processes. ( and are classic examples of sex role reversal.) Also, unlike a female, a male (except in species) has some uncertainty about whether or not he is the true parent of a child, and so is less interested in spending his energy helping to raise offspring that may or may not be related to him. As a result of these factors, males can be expected to be more willing to mate than females, while females are expected to be the ones doing the choosing (except in cases of s, which has been observed in numerous species, including mammals, birds, insects and fish). The effects of sexual selection are thus often more pronounced in males than in females. Differences in secondary sexual characteristics between males and females of a species are referred to as s. These can be as subtle as a size difference (sexual size dimorphism, often abbreviated as SSD) or as extreme as horns and colour patterns. Sexual dimorphisms abound in nature. Examples include the possession of antlers by only male , the brighter coloration of many male birds in comparison with females of the same species, or even more distinct differences in basic morphology, such as the drastically increased eye-span of the male . The , with its elaborate and colourful tail feathers, which the lacks, is often referred to as perhaps the most extraordinary example of a dimorphism. Male and female s and s also differ radically in their plumage. Early naturalists even believed the females to be a separate species. The largest sexual size dimorphism in is the ' in which males are up to 30 times the size of females. Many other fish such as are sexually dimorphic. Extreme sexual size dimorphism, with females larger than males, is quite common in and . The maintenance of sexual reproduction in a highly competitive world is one of the major puzzles in biology given that can reproduce much more quickly as 50% of offspring are not males, unable to produce offspring themselves. Many non-exclusive hypotheses have been proposed, including the positive impact of an additional form of selection, sexual selection, on the probability of persistence of a species.
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Male intrasexual competition

Male-male competition occurs when two males of the same species compete for the opportunity to mate with a female. Sexually dimorphic traits, size, sex ratio, and the social situation may all play a role in the effects male-male competition has on the reproductive success of a male and the mate choice of a female. Larger males tend to win male-male conflicts due to their sheer strength and ability to ward off other males from taking over their females. For instance, in the fly ', size shows the strongest correlation to the outcome of male-male conflicts over resources like territory and females.


Influencing factors


Sex ratio

There are multiple types of male-male competition that may occur in a population at different times depending on the conditions. Competition variation occurs based on the frequency of various mating behaviours present in the population. One factor that can influence the type of competition observed is the population density of males. When there is a high density of males present in the population, competition tends to be less aggressive and therefore sneak tactics and disruptions techniques are more often employed. These techniques often indicate a type of competition referred to as scramble competition. In Japanese medaka, ', sneaking behaviours refer to when a male interrupts a mating pair during copulation by grasping on to either the male or the female and releasing their own sperm in the hopes of being the one to fertilize the female. Disruption is a technique which involves one male bumping the male that is copulating with the female away just before his sperm is released and the eggs are fertilized. However, all techniques are not equally successful when in competition for reproductive success. Disruption results in a shorter copulation period and can therefore disrupt the fertilization of the eggs by the sperm, which frequently results in lower rates of fertilization and smaller clutch size.


Resource value and social ranking

Another factor that can influence male-male competition is the value of the resource to competitors. Male-male competition can pose many risks to a male's fitness, such as high energy expenditure, physical injury, lower sperm quality and lost paternity. The risk of competition must therefore be worth the value of the resource. A male is more likely to engage in competition for a resource that improves their reproductive success if the resource value is higher. While male-male competition can occur in the presence or absence of a female, competition occurs more frequently in the presence of a female. The presence of a female directly increases the resource value of a territory or shelter and so the males are more likely to accept the risk of competition when a female is present. The smaller males of a species are also more likely to engage in competition with larger males in the presence of a female. Due to the higher level of risk for subordinate males, they tend to engage in competition less frequently than larger, more dominant males and therefore breed less frequently than dominant males. This is seen in many species, such as the Omei treefrog, '','' where larger males obtain more mating opportunities and mate with larger females.


Winner–loser effects

A third factor that can impact the success of a male in competition is . Burrowing crickets, ''Velarifictorus aspersus'', compete for burrows to attract females using their large mandibles for fighting. Female burrowing crickets are more likely to choose winner of a competition in the 2 hours after the fight. The presence of a winning male suppresses mating behaviours of the losing males because the winning male tends to produce more frequent and enhanced mating calls in this period of time.


Effect on female fitness

Male-male competition can both positively and negatively affect female fitness. When there is a high density of males in a population and a large number of males attempting to mate with the female, she is more likely to resist mating attempts, resulting in lower fertilization rates. High levels of male-male competition can also result in a reduction in female investment in mating. Many forms of competition can also cause significant distress for the female negatively impacting her ability to reproduce. An increase in male-male competition can affect a female's ability to select the best mates, and therefore decrease the likelihood of successful reproduction. However, group mating in Japanese has been shown to positively affect the fitness of females due to an increase in genetic variation, a higher likelihood of paternal care and a higher likelihood of successful fertilization. Exposure to environmental estrogens, such as some herbicides, can confuse female choice of males.


In different taxa

:''In detail, see Sexual selection in , , , , , , '' Sexual selection has been observed to occur in plants, animals and fungi. In certain and species of the throwing of s is a form of sexual selection. Certain male s of the order cement the vaginal pores of their females. Today, biologists say that certain evolutionary traits can be explained by —competition between members of the same species—distinguishing between competition before or after . Before , ''intrasexual selection''—usually between males—may take the form of ''male-to-male combat''. Also, ''intersexual selection'', or ', occurs when females choose between male mates. Traits selected by male combat are called secondary sexual characteristics (including horns, antlers, etc.), which Darwin described as "weapons", while traits selected by mate (usually female) choice are called "ornaments". Due to their sometimes greatly exaggerated nature, secondary sexual characteristics can prove to be a hindrance to an animal, thereby lowering its chances of survival. For example, the large antlers of a moose are bulky and heavy and slow the creature's flight from predators; they also can become entangled in low-hanging tree branches and shrubs, and undoubtedly have led to the demise of many individuals. Bright colourations and showy ornamenations, such as those seen in many male birds, in addition to capturing the eyes of females, also attract the attention of predators. Some of these traits also represent energetically costly investments for the animals that bear them. However, one must also consider that intersexual selection can occur with an emphasis on resources that one sex possesses rather than morphological and physiological differences. For example, males of ', a non-social bee species, form aggregations of territories considered to be leks, to defend fragrant-rich primary territories. The purpose of these aggregations is only facultative, since the more suitable fragrant-rich sites there are, the more habitable territories there are to inhabit, giving females of this species a large selection of males with whom to potentially mate. After copulation, male–male competition distinct from conventional may take the form of , as described by in 1970. More recently, interest has arisen in , a phenomenon of internally fertilised animals such as mammals and birds, where a female can get rid of a male's without his knowledge. Finally, is said to occur between breeding partners, sometimes leading to an between males and females. Sexual selection can also occur as a product of release, such as with the stingless bee, '. Female mating preferences are widely recognized as being responsible for the rapid and divergent evolution of male secondary sexual traits. Females of many animal species prefer to mate with males with external ornaments—exaggerated features of morphology such as elaborate sex organs. These preferences may arise when an arbitrary female preference for some aspect of male morphology—initially, perhaps, a result of —creates, in due course, selection for males with the appropriate ornament. One interpretation of this is known as the . Alternatively, genes that enable males to develop impressive ornaments or fighting ability may simply show off greater or a more efficient , features that also benefit females. This idea is known as the . Bright colors that develop in animals during mating season function to attract partners. It has been suggested that there is a causal link between strength of display of ornaments involved in sexual selection and free radical biology. To test this idea, experiments were performed on male . Male lizards are brightly conspicuous in their breeding coloration, but their color declines with aging. Experiments involving administration of s to these males led to the conclusion that breeding coloration is a reflection of innate anti-oxidation capacity that protects against , including . Thus color could act as a "health certificate" that allows females to visualize the underlying oxidative stress induced damage in potential mates. Darwin conjectured that traits such as beards and hairlessness in different human populations are results of . has hypothesized that many human behaviours not clearly tied to survival benefits, such as humour, music, visual art, verbal creativity, and some forms of , are courtship adaptations that have been favoured through sexual selection. In that view, many human artefacts could be considered subject to sexual selection as part of the , for instance clothing that enhances sexually selected traits . Some argue that the is a sexually selected trait, as it would not confer enough fitness in itself relative to its high maintenance costs.PLoS ONE: Sexual Selection and the Evolution of Brain Size in Primates


References


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* Andersson, M. (1994) ''Sexual selection''. . * Arnqvist, G. & Rowe, L. (2013)
Sexual conflict
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The ant and the peacock: altruism and sexual selection from Darwin to today
'. Cambridge University Press. * (1871) '. John Murray, London. * Eberhard, W. G. (1996
Female control: Sexual selection by cryptic female choice
Princeton, Princeton University Press. * (1930) '. Oxford University Press, , Chapter
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* (2006) ''La guerre des sexes chez les animaux''. Eds Odile Jacob. * Miller, G. F. (1998) How mate choice shaped human nature: A review of sexual selection and human evolution. In: C. Crawford & D. Krebs (Eds.) ''Handbook of evolutionary psychology: Ideas, issues, and applications''. Lawrence Erlbaum, pp. 87–129 * Miller, G. F. (2000) ''The Mating Mind: How sexual choice shaped the evolution of human nature''. Heinemann, London. * Rosenberg, J. & Tunney, R. J. (2008)
Human vocabulary use as display. Evolutionary Psychology, 6, 538–549
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