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Sexual selection in humans concerns the concept of sexual selection, introduced by Charles Darwin as an element of his theory of natural selection,^[1] as it affects humans. The role of sexual selection in human evolution has not been firmly established although neoteny has been cited as being caused by human sexual selection.^[2] It has been suggested that the human brain is itself a product of sexual selection, i.e. it has developed as a sexual ornamentation to be used in courtship rather than for survival itself,^[3] and that it has developed in ways outlined by Ronald Fisher in the Fisherian runaway model.^{[4][5][6][7][8]} Fisher also stated that the development of sexual selection was "more favourable" in humans.^[9]

General hypotheses

Some hypotheses about the evolution of the human brain argue that it is a sexually selected trait, as it would not confer enough fitness in itself relative to its high maintenance costs (a quarter to a fifth of the energy and oxygen consumed by a human).^[10]

Sexual selection's role in human evolution cannot be definitively established, as features may result from an equilibrium among competing selective pressures, some involving sexual selection, others natural selection, and others pleiotropy. In the words of Richard Dawkins:

"When you notice a characteristic of an animal and ask what its Darwinian survival value is, you may be asking the wrong question. It could be that the characteristic you have picked out is not the one that matters. It may have "come along for the ride", dragged along in evolution by some other characteristic to which it is pleiotropically linked."^[11]

Darwin's sexual selection hypothesis

Charles Darwin described sexual selection as depending on "the advantage which certain individuals have over others of the same sex and species, solely in respect of reproduction".^[12] Darwin noted that sexual selection is of two kinds and concluded that both kinds had operated on humans:^[13] "The sexual struggle is of two kinds; in the one it is between the individuals of the same sex, generally the male sex, in order to drive away or kill their rivals, the females remaining passive; whilst in the other, the struggle is likewise between the individuals of the same sex, in order to excite or charm those of the opposite sex, generally the females, which no longer remain passive, but select the more agreeable partners."^[14]

Charles Darwin conjectured that the male beard, as well as the hairlessness of humans compared to nearly all other mammals, were results of sexual selection. He reasoned that since the bodies of females are more nearly hairless, the loss of fur was due to sexual selection of females at a remote prehistoric time when males had overwhelming selective power, and that it nonetheless affected males due to genetic correlation between the sexes. He also hypothesized that contrasts in sexual selection acting along with natural selection were significant factors in the geographical differentiation in human appearance of some isolated groups, as he did not believe that natural selection alone provided a satisfactory answer. Although not explicit, his observation that in Khoisan women "the posterior part of the body projects in a most wonderful manner" (known as steatopygia)^[15] implies sexual selection for this characteristic. In The Descent of Man, and Selection in Relation to Sex, Darwin viewed many physical traits which vary around the world as being so trivial to survival^[16] that he concluded some input from sexual selection was required to account for their presence. He noted that variation in these features among the various peoples of the world meant human mate-choice criteria would also have to be quite different if the focus was similar, and he himself doubted that, citing^[17] reports indicating that ideals of beauty did not, in fact, vary in this way around the world.

Sexual dimorphism

Men are generally hairier than women, and Darwin was of the opinion that hairlessness was related to sexual selection; however, several other explanations have been advanced to explain human hairlessness, a leading one is loss of body hair to facilitate sweating.^[18] This idea closely relates to that of the suggested need for increased photoprotection and is part of the most-commonly-accepted scientific explanation for the evolution of pigmentary traits.^[19]

Indicating that a trait is under sexual selection can be difficult to prove through correlational methods, as characters may result from different selective pressures, some involving sexual selection, others natural selection, and some may be accidental and due to pleiotropy. For example, monogamous primates are known to typically exhibit little sexual dimorphism such as particularly large males armed with huge canines; however, powerful big-toothed males can provide protection against predators and may be bigger for that reason, rather than in order to win confrontations over females. Males and females differing in size can specialize in, and more fully exploit, different food resources while avoiding competing with each other; furthermore, body size can be useful in avoiding predators and may also be of assistance in securing a mate. This is further complicated by the consideration that with larger body size, the skeleton of mammals becomes much more robust and massive (relatively speaking).^[20] Bearing these caveats in mind, levels of sexual dimorphism are generally seen as a marker of sexual selection. Studies have shown the earliest homininae were highly dimorphic and that this tendency lessened over the course of human evolution, suggesting humans have become more monogamous. In contrast, gorillas living in harems exhibit a much stronger sexual dimorphism (see: homininae).^[21]

Sexual anatomy

The theory of sexual selection has been used to explain a number of human anatomical features. These include rounded breasts, facial hair, pubic hair and penis size. The breasts of primates are flat, yet are able to produce sufficient milk for feeding their young. The breasts of non-lactating human females are filled with fatty tissue and not milk. Thus it has been suggested the rounded female breasts are signals of fertility.^[22] Richard Dawkins has speculated that the loss of the penis bone in humans, when it is present in other primates, may be due to sexual selection by females looking for a clear sign of good health in prospective mates. Since a human erection relies on a hydraulic pumping system, erection failure is a sensitive early warning of certain kinds of physical and mental ill health.^[23]

Homo has a thicker penis than the other great apes, though it is no longer than the chimpanzee's.^[24] It has been suggested the evolution of the human penis towards larger size was the result of female choice rather than sperm competition, which generally favors large testicles.^[25] However, penis size may have been subject to natural selection, rather than sexual selection, due to a larger penis' efficiency in displacing the sperm of rival males during sexual intercourse. A model study showed displacement of semen was directly proportional to the depth of pelvic thrusting, as an efficient semen displacement device.^[26]

Selection preferences and biological drivers

There are a variety of factors that drive sexual selection in humans. Current available research indicates that selection preferences are biologically driven,^[27] that is, by the display of phenotypic traits that can be both consciously and unconsciously evaluated by the opposite sex to determine the health and fertility of a potential mate.^[28] This process can be affected, however, by social factors, including in cultures where arranged marriage is practiced, or psychosocial factors, such as valuing certain cultural traits of a mate, including a persons social status, or what is perceived to be an ideal partner in various cultures.^[29]

Selection preferences in females

Some of the factors that affect how females select their potential mates for reproduction include voice pitch, facial shape, muscular appearance, and height.^[30] Several studies suggest that there is a link between hormone levels and partner selection among humans. In a study measuring female attraction to males with varying levels of masculinity, it was established that women had a general masculinity preferences for men's voices, and that the preference for masculinity was greater in the fertile phase of the menstrual cycle than in the non-fertile phase.^[30] There is further evidence from the same study that in fertile stages of the menstrual cycle, women also had a preference for other masculine traits such as body size, facial shape, and dominant behavior, which are indicators of both fertility and health.^[30] This study did not exclude males with feminine traits from being selected, however, as feminine traits in men indicate a higher probability of long-term relationship commitment,^[30] and may be one of several survival strategies.^[31] Further research also backs up the idea of using phenotypic traits as a means of assessing a potential mates fitness for reproduction as well as assessing whether a partner has high genetic quality.^[32]

Another factor affecting the selection process is the environment which the person inhabits. In biological terms, certain environmental conditions may bring about demands for or the disregarding of certain traits. One such example is a preference for males whose facial structure indicates certain hormonal ratios, such as testosterone-cortisol levels (sex and stress hormones). Research shows that, for example, in countries with varying Human Development Index (HDI) levels, females have different preferences for sex-stress hormone ratios, as expressed in the male's face. A Royal Society research showed a significant correlation between a measure of societal development and preferences for indication of higher testosterone levels, as manifested in facial features, and the interaction between preferences for testosterone and cortisol."^[33] It was concluded that societal-level ecological factors impact the valuation of traits by combinations of sex- and stress-hormones.^[33]

Selection preferences in males

Like their female counterparts, males also use visual information about a potential mate, as well as voice, body shape, and an assortment of other factors in selecting a partner. Research shows that males tend to prefer feminine women's faces and voices as opposed to women with masculine features in these categories.^[34] Furthermore, males also evaluate skin coloration, symmetry, and apparent health, as a means by which the select a partner for reproductive purposes.^[34] Males are particularly attracted to femininity in women's faces when their testosterone levels are at their highest, and the level of attraction to femininity may fluctuate as hormone levels fluctuate.^[35] Studies on men have also been done to show the effects of exogenous testosterone and its effects on attraction to femininity, and the results concluded that throughout several studies, men have shown decreased preference for feminine female faces in the long-term context, when given exogenous testosterone, but this difference did not occur with placebo.^[36]

Common preferences in either sex

Sexual selection preferences are general terms by which the mating and reproductive process are understood. As one article states, sexual selection is in essence a process which favors sexual displays for attraction, aggressiveness, dominance, size, and strength, and the ability to exclude competitors by force if necessary, or by using resources to win.^[37] Both male and female use voice, face, and other physical characteristics^[38] to assess a potential mate's ability to reproduce, as well as their health^[39]. Together with visual and chemical signals, these crucial characteristics which are likely to enhance the ability to produce offspring, as well as long term survival prospects, can be assessed and selections made.^[40][41]

Phenotype

Sexual selection has continued to be suggested as a possible explanation for geographical variation in appearance within the human species; in modern hypotheses, marriage practices are proposed as the main determinant of sexual selection. John Manning^[42] suggests that where polygyny is common, men face intense competition for wives and are more likely to be completely unsuccessful in reproducing, and the result is strong selection of males for traits which are adaptive for successful reproduction. He proposes a link to skin color through selection of males for testosterone-mediated traits which confer an ability to successfully compete for females. He suggests testosterone makes the human immune system less competent to resist pathogens. In this view the antimicrobial properties of melanin help mitigate the susceptibility to disease that polygyny induces by increasing testosteronization. According to this argument, the anti-infective qualities of melanin were more important than protection from ultraviolet light in the evolution of the darkest skin types. Manning asserts that skin color is more correlated with the occurrence of polygyny – explicable by it having an antimicrobial function – than the latitudinal gradient in intensity of ultraviolet radiation, and he points to the lack of very dark skin at equatorial latitudes of the New World and the relatively light skin of Khoisan people in Africa.^[42][43]

Research seems to contradict Manning's explanation about skin color. In 1978, NASA launched the Total Ozone Mapping Spectrometer, which was able to measure the ultraviolet radiation reaching Earth's surface. Jablonski and Chaplin took the spectrometer's global ultraviolet measurements and compared them with published data on skin color in indigenous populations from more than 50 countries. There was an unmistakable correlation: The weaker the ultraviolet light, the fairer the skin.^[44] Rogers et al. (2004) performed an examination of the variation in MC1R nucleotide sequences for people of different ancestry and compared the sequences of chimpanzees and humans from various regions of the Earth. Rogers concluded that, at the time of the evolutionary separation of chimpanzees and humans, the common ancestors of all humans had light skin that was covered by dark hair. Additionally, our closest extant relative, the chimpanzee, has light skin covered by thick body hair.^[45] Over time human hair disappeared to allow better heat dissipation through sweating^[46] and the skin tone grew darker to increase the epidermal permeability barrier^[47] and protect from folate depletion due to the increased exposure to sunlight.^[48] When humans started to migrate away from the tropics, there was less-intense sunlight, partly due to clothing to protect against cold weather. Under these conditions there was less photodestruction of folate, and so the evolutionary pressure stopping lighter-skinned gene variants from surviving was reduced. In addition, lighter skin is able to generate more vitamin D (cholecalciferol) than darker skin, so it would have represented a health benefit in reduced sunlight if there were limited sources of vitamin D.^[46] The genetic mutations leading to light skin experienced selective pressure due to settlement in northern latitudes.^[49]

Anthropologist Peter Frost has proposed that sexual selection was responsible for the evolution of pigmentary traits of women in Northern and Eastern European populations. He contends that the diversity of hair and eye color in Northeast European populations originated as a consequence of intense female-female competition, and is an adaptation for reproductive success in women.^[50][51]

Geoffrey Miller hypothesis

Homo habilis - Forensic facial reconstruction

Geoffrey Miller, drawing on some of Darwin's largely neglected ideas about human behavior, has hypothesized that many human behaviors not clearly tied to survival benefits, such as humor, music, visual art, some forms of altruism, verbal creativity or the fact that most humans have a far greater vocabulary than that which is required for survival,^[52] Miller (2000) has proposed that this apparent redundancy is due to individuals using vocabulary to demonstrate their intelligence, and consequently their "fitness", to potential mates. This has been tested experimentally, and it appears that males do make greater use of lower-frequency (more unusual) words when in a romantic mindset compared to a non-romantic mindset, suggesting that vocabulary is likely to be used as a sexual display (Rosenberg & Tunney, 2008). All these qualities are considered courtship adaptations that have been favored through sexual selection.^[53]

Miller is critical of theories that imply that human culture arose as accidents or by-products of human evolution. He believes that human culture arose through sexual selection for creative traits. In that view, many human artifacts could be considered subject to sexual selection as part of the extended phenotype, for instance clothing that enhances sexually selected traits.^[54] During human evolution, on at least two occasions, hominid brain size increased rapidly over a short period of time followed by a period of stasis. The first period of brain expansion occurred 2.5 million years ago, when Homo habilis first began using stone tools. The second period occurred 500,000 years ago, with the emergence of archaic Homo sapiens. Miller argues that the rapid increases in brain size would have occurred by a positive feedback loop resulting in a Fisherian runaway selection for larger brains. Tor Nørretranders, in The Generous Man conjectures how intelligence, musicality, artistic and social skills, and language might have evolved as an example of the handicap principle, analogously with the peacock's tail, the standard example of that principle. Another hypothesis^[55] proposes that human intelligence is a courtship indicator of health and resistance against parasites and pathogens which are deleterious to human cognitive capabilities.^[56]

Opposing arguments

The role of sexual selection in human evolution has been considered controversial from the moment of publication of Darwin's book on sexual selection (1871). Among his vocal critics were some of Darwin's supporters, such as Alfred Wallace, who argued that animals and birds do not choose mates based on their beauty or fine plumages, and that the artistic faculties in humans belong to their spiritual nature and therefore cannot be connected to natural selection, which only affects the animal nature.^[57] Darwin was accused of looking to the evolution of early human ancestors through the moral codes of the 19th century Victorian society. Joan Roughgarden, citing elements of sexual behavior in animals and humans that cannot be explained by the sexual-selection model, suggested that the function of sex in human evolution was primarily social.^[58]

Joseph Jordania suggested in 2011 that in explaining such human morphological and behavioral characteristics as singing, dancing, body painting, wearing of clothes, Darwin and proponents of sexual selection neglect another important evolutionary force, intimidation of predators and competitors with the ritualized forms of warning display, which uses the same arsenal of visual, audio, olfactory and behavioral features as sexual selection. According to Jordania, most of these warning displays were incorrectly attributed to the forces of sexual selection. Jordania proposed an aposematic model of human evolution, where most of the human morphological and behavioral features that had been considered by Darwin as the result of sexual selection, via female choice, are explained by the aposematic (intimidating) display.^[59]

See also

• Physical attractiveness


• Marriage squeeze


• Hypergamy


• Human mating strategies


• Strategic pluralism

References

Further reading

• 
  The Mating Mind, by Geoffrey Miller, published by Anchor Books, 2001.
  ISBN 978-0-385-49517-2.