Sexual selection

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The sexual selection ( Latin selectio , read ') is a intraspecific selection acting on physical characteristics and is caused by variance in reproductive success thereof between members sex. Charles Darwin recognized this "sexual selection" as one of the three types of selection in the theory of evolution . This leads to the emergence of sexually dimorphic characteristics, i. H. the secondary sexual characteristics in the appearance of the sexes of a species , explained evolutionarily .

Delimitations

Sexual selection evokes a colorful sexual dimorphism in the lavender bird of paradise . Male above; Females below. Painting by John Gerrard Keulemans (1912)

In his work The Origin of Species from 1859, Charles Darwin describes artificial and natural selection .

  • The artificial selection ( breeding ) is a focused selection of individuals with specific, desired properties by man. Individuals who do not have these characteristics are strictly excluded from reproduction . As a result, forms can develop that are less adaptable than their ancestors in the field (domestic animals, cultivated plants).
  • The natural selection will take place without the action of man. Individuals with greater biological fitness have structural or performance characteristics which, in comparison to other individuals, cause a higher number of surviving offspring in their environment. This selection pressure is subject to properties such as adaptability to environmental changes, opportunities to nod in and resistance to enemy pressure. In evolutionary biology and sociobiology , the extended term of relative selection explains altruistic behavior patterns. As an extension of natural selection, group selection was proposed, which has recently been discussed as multilevel selection .

The concept of natural selection, however, contradicted observable characteristics that are actually disadvantageous for their carriers in the respective environment . In his book The Descent of Man and Sexual Selection of 1871, Darwin describes the sexual selection with which he could explain these characteristics.

  • The sexual selection is a selection of individuals by advantages in reproductive success compared to same sex the same way . Intrasexual selection affects characteristics that play a role in same-sex competition for access to mating partners. Intersex selection acts on traits used by members of one sex to bring about an explicit choice for mating in members of the opposite sex.

Intrasexual selection: competitive struggles between members of the same sex

Intrasexual selection affects characteristics (e.g. body size, coloration, vocalizations, canines) that are important for same-sex competition in mating access. Such features are useful as a weapon (e.g. antlers) or as protection against injuries (e.g. lion's mane) in damage or commentary fights, or they serve as social signals in the case of showing off . A polygamous mating behavior is a prerequisite for the distinctive manifestation of these sexual dimorphisms . In monogamous species, such features develop only weakly or not at all. The intrasexual selection has a stronger effect on the gender, which has the lower parenting effort . In many species and most vertebrates, these are the males, in some species also the females. When males go to great lengths to recruit females, there is an incentive for males to be choosy. For example, in the monogamous marmosets and tamarin monkeys , the females compete to form pairs with attractive males.

If intrasexual selection has a symmetrical effect on both sexes, even strong intrasexual selection does not lead to pronounced sexual dimorphism. This occurs z. B. in monogamous couple relationships if, with a permanent excess of males or females, numerous extra-pair copulations undermine monogamy and the genetic quality of the possible mating partners is very different. Then there is a selective incentive for both sexes to develop signs of quality or health. Mating games , after- mating games or pair bonding rituals with the participation of both sexes have the same effect . Charles Darwin was already aware of this case. Strong dimorphism tends to be a sign of unequal systems in which the variability in reproductive success of one, mostly male, sex is higher than that of the other.

In species where the males fight for access to females, the male is usually larger than the female. In species in which the competition takes place through pronounced mating games or demonstrations, however, the males tend to be smaller. According to Rensch's rule, which was confirmed in a study in shorebirds, in species with sexual size dimorphism, the males tend to be larger in large species and smaller in small species than the females. In amphibians, the males are usually smaller than the females. In the few species with larger males, there is a significant connection with mating battles between the males.

Sperm competition

In many species, the promiscuity of the females can lead to sperm competition between the males. As a result of the selection pressure, the males had to adapt, e.g. B. Production of particularly fast and efficient sperm , control of the females, large testes that produce voluminous and sperm-rich ejaculate, special "kamikaze" sperm with a spiral-shaped tail that can wrap around competing sperm and destroy them, or masturbation for fitness increase in sperm for the next copulation. As a result of this competition, the females have developed techniques and strategies with which, after copulation with several males, they can choose which sperm is to be fertilized (“cryptic” partner choice), or according to the theory of “retained sperm” by Robin Baker and Mark Bellis what sperm she will fertilize. Because of the multiple fertilization, females have z. B. the possibility of getting good genes for your offspring and avoiding sperm that are not very viable or genetically incompatible.

Intersex selection: partner choice by members of the opposite sex

Other forms of sexual dimorphism, such as the magnificent plumage of the peacock or the bird of paradise , cannot be explained by natural or intrasexual selection, but by the preference of their bearer in the choice of mate . The sex with the higher effort chooses the partner. In many species this is the females (“female choice”) due to the higher parental effort. In some species, the males choose (e.g., Odin's and Thor's chickens ). For some species, the gender chosen can also be determined by other influences, e.g. B. through the food supply, which influences the quantity and quality of spermatophores that females receive from the males, through the effort involved in choosing a partner or through the operational gender ratio.

Examples of selection criteria:

Darwin accepted, but not explained, the evolution of intersex selection. If mating with carriers of certain characteristics results in a higher number of surviving offspring in the opposite sex, preference for those characteristics may evolve. However, some properties such as the magnificent plumage seem to have a fitness disadvantage for the female, since such plumage in natural selection causes disadvantages for their male offspring. The same applies to other features. For example, there are monogamous and polygamous males in many bird species. In general, females of polygamous males have less reproductive success due to their reduced help in raising young . However, some females mate with already mated instead of a free male.

Explanations of such cases by the theory of evolution must be based on the fact that the carriers of a selected trait will have more offspring in the longer term than those without this trait. Otherwise the characteristic is at best evolutionarily neutral. Various model assumptions are conceivable for the corresponding pairing systems, in which this applies despite the disadvantages caused by natural selection.

  1. Directly selected mechanisms: Carriers of the characteristic have an advantage through the choice of partner that leads directly to a higher number of offspring.
  2. Mechanisms selected indirectly: Carriers of the trait initially have fewer offspring, but they are more fit, which is why they prevail in the population in the longer term. In doing so, it is not the characteristic itself that is selected, but a characteristic correlated with it (e.g. louder mating call, correlated with genetic quality).
  3. Sexual conflict: The trait only benefits members of one gender. Since the parents are genetically different, z. B. a trait passed on by the father can be promoted, which gives the males a mating advantage, even if the trait can be directly disadvantageous for females.

In natural pairing systems these possibilities do not have to be realized exclusively. A certain characteristic can also be partially conditioned or promoted through selection in several ways, which makes research demanding. The same characteristic can be equally important for intra- as well as for intersexual selection, as it is e.g. As for the mop of the crested auklet Aethia cristatella is detected.

The differences in body structure and behavior of the sexes, which are the starting point for sexual selection, result from the differences in the gametes according to the classical view . The sex with the larger gametes is (by definition) the female. The emergence of this difference itself is usually interpreted through “disruptive selection”: An individual can produce very many, but then inevitably very small, or a few, but then better-nourished, gametes with a higher probability of survival. Intermediate individuals fall between the two optima. From the different gamete sizes, it is usually concluded that the male sex, due to the much higher potential reproductive rate, has a greater advantage of investing as little as possible in individual offspring and instead investing better in a higher number of offspring ( Bateman principle ). As a result, small differences in the strategy of the sexes can initially increase. However, in diploid species, the number of offspring of one sex cannot exceed that of the other (the “ Fisher condition”). Differences can therefore be based on the fact that few males fertilize a large number of females and increase the relative proportion of their genes in the gene pool. A comparable strategy for the females is not possible.

If an individual does not accept certain possible partners, i.e. is choosy, costs, e.g. B. causes search costs or risks and spent lifetime. Such a strategy therefore requires a selection mechanism. This connection has been proven experimentally z. B. in the pronghorn : if females can freely choose their mating partner, they have more offspring than if they mate randomly.

Several theories have been developed to interpret gender dimorphism and mating systems in different species. The best known are the runaway selection , i. H. Self-running processes from RA Fisher and the handicap principle .

Direct benefits

A female can gain direct advantages for the offspring by choosing a partner if the male z. B. defends a high-quality territory and participates in the rearing of young or the defense against predators . This case was considered trivial for a long time and was therefore hardly considered. A systematic review showed only slightly greater effect direct benefits through the female mate choice than by indirect (eg. As a result of the genetic makeup of the offspring) for some fitness components. To recognize such benefits, the female interprets the males' signals and must avoid cheaters who imitate the fitness signals. As with the genetic makeup, there is a strong incentive to develop forgery-proof signal systems.

Sensory preference

According to the sensory bias theory, sexual characteristics can develop through female preferences for male characteristics such as color, size or acoustic signals. According to this, females prefer males with such characteristics when choosing a partner. For example, the males of the genus Anolis perform quick upward movements in front of the female in a specific mating ritual. In the genus Xiphophorus there are males with a long extension of the caudal fin ( swordtail ) and without extension ( platys ). In selective experiments, females without an extension prefer the males with an artificially glued extension over the wild type. In some species, individuals with completely unnatural human markings are also preferred as partners. This effect occurred in studies. a. in birds in which certain males were ringed in color by the experimenter in order to investigate completely different questions. Females significantly preferred males with certain colored rings over others.

Runaway selection

The Runaway selection was developed from 1915 by the geneticist and evolutionary biologist RA Fisher and published in his book in 1930. After 1958, biologists and mathematicians took up the theory and developed it further. A self-run process is created by sensory preferences when choosing a partner, e.g. B. when females prefer male carriers of a hereditary trait to mate. If the genes for this preference and for the trait are linked, there is a positive feedback that causes extreme trait manifestations in an evolutionarily short time. The process can then only end due to external influences, e.g. B. by natural selection. Then z. B. the tail length of the peacock cock has become so expensive that it has a survival disadvantage. When the survival disadvantage and the advantage of mating success are balanced, a balance can be established. For example, by comparing the characteristic expression within the species group with the predictions of the various hypotheses about intersexual selection, the mechanism was identified as the most likely reason for the coloring and the courtship games of the males in purrbirds .

Sexy Son Hypothesis

As a variant of the runaway selection , the sexy-son hypothesis was proposed in 1979 by PJ Weatherhead and RJ Robertson. Like runaway selection , this hypothesis is difficult to test. According to this hypothesis, females of some species mate with polygynous males. B. invest a lot in the search for a partner due to particularly pronounced secondary sexual characteristics, although such a male will help less in raising young. Your advantage may be inherited polygyny and thus a possible high reproductive success of your "sexy sons" in the future. This allows the trait to spread throughout the population. Investments by males to raise the young, e.g. B. mating (re) games, couple bonding rituals or a territory are no guarantee for a fatherhood of the offspring. This hypothesis explains the behavior of the females of some songbird species such as B. the star . The females mate with polygynous males, even if this results in fewer offspring than with a monogamous partner who helps with rearing. In birds, females can in principle influence the sex of their offspring and, according to the hypothesis, they should increase the proportion of their male offspring in order to be able to spread their genes to their then also polygynous sons with this only advantage.

Handicap hypothesis

The handicap principle developed by Amotz Zahavi and Avishag Zahavi explains the development of traits through partner choice that bring a survival disadvantage for the wearer, but as a signal prove the quality of his genes. According to the hypothesis, the handicap is a forgery-proof signal from a particularly viable individual who can pass on its beneficial properties to the offspring. That is why we speak of “good genes” or “luxury” traits. Through exposure or handicap and thus endangerment from predators or food competitors through the handicap, a mating partner signals his particular fitness. A mating partner with such abnormalities is then assessed as particularly strong and healthy and thus a relatively safe guarantee for healthy and viable offspring. The intersexual and intrasexual selection are equivalent to each other. An expensive trait used to attract a partner is equivalent to an expensive trait used to fight with peers such as sex. B. the deer antlers. An extension of the handicap hypothesis on the influence of the immune defense comes from Folstad and Karter. Their hypothesis is based on the observation that a higher level of the sex hormone testosterone increases the expression of male sex-dimorphic characteristics and at the same time reduces the body's immune defense. Only particularly healthy males can therefore show distinctive characteristics and accept the associated immunodeficiency as a handicap.

Evolutionary impasse

The consequences of particularly extreme handicap characteristics are interpreted as an "evolutionary dead end" if their advantages for reproductive fitness are due to drastic changes, e.g. B. the environment, illnesses, new competitors or changed food are lost and therefore only the fitness disadvantages remain for their wearers. Due to these disadvantages, the population is then reduced if the specializations associated with extreme formations in the species concerned cannot adapt to new circumstances. As such dead ends that have led to extinction, z. B. the antlers of the ice age giant deer , the tusks of the mammoths or the canines of the saber-toothed cats interpreted. The giant deer lived in the open tundra , which at the end of the Ice Age initially turned into marshland and then into forest . According to this hypothesis, the giant deer with its weight and antlers of over 3.6 m wingspan could neither live on soft ground nor in the forest, which is why the species became extinct due to the ecological changes . However, this hypothesis is difficult to investigate and is controversial in relation to other explanatory models, such as the overkill hypothesis .

Evolutionary Suicide

If the individuals of a species benefit greatly at the expense of the population, evolutionary adaptations can lead to extinction of the species according to the hypothesis of "evolutionary suicide". Some studies have shown a correlation between traits, selection pressure and an increased risk of extinction.

Physical performance characteristics

External characteristics such as body height, weapon size or the size of the primary sex organs can be directly related to the fitness of the males. In some species, the females also check the genetic fitness of the males via their physical performance in courtship games , e.g. B. in paired courtship dance, courtship flight or courtship fight, or through their preliminary work for brood care. A female only copulates with males whose fitness she judges to be sufficient.

Examples:

  • In some weaver bird species , such as B. Textorweber , the male builds the nest and the female checks the firmness. In some species this behavior has become ritualized and only nesting material is presented.
  • With some bird species, the male brings food to the female as a "bride present" and thus demonstrates the quality of his feeding area.

The fire beetle ( Neopyrochroa flagellata ) has another form of physiological performance trait . The male ingests the poison cantharidin through food, which protects eggs and larvae from predators. This poison is mostly stored in a gland in the abdomen and transmitted with the sperm, a small part is collected in a head gland. The females only mate with males when they can taste the venom on the male's head gland.

Social signals

In some species, a signaling system has evolved which correlates with the genetic fitness of the males, but which has no direct connection with their ability to survive, reproduce or rearing. Females choose males on the basis of their most pronounced key stimuli , such as B. conspicuous colors, calls or behaviors that are presented by males during courtship . The conspicuousness of the signals reduces the general fitness of the males through natural selection, which counteracts sexual selection. This creates a balance in the expression of the characteristics and prevents hypertrophication of secondary sexual characteristics. This connection was z. B. detected in Poecilia reticulata . Brightly colored males are more attractive to females, but also more conspicuous to predators . In habitats without predators, the males are more colorful.

Examples:

  • Peacock : Decorative feathers with many and large eyes increase reproductive success.
  • Bankiva chicken : Hens prefer roosters with light, "shining" eyes and large, red combs and wattles. These traits correlate with good health and high disease resistance.
  • Barn swallow ( Hirundo rustica ): Barn swallows show no conspicuous sexual dimorphism. The males can only be recognized by the elongated edge feathers of the fork tail. They are more than an inch longer than those of the females, who prefer males with longer tail feathers.
The tail feathers in the males vary between 84 and 132 millimeters. Older males have longer tail feathers than younger ones, as these are somewhat elongated with each moult in winter quarters. Older males arrive in the breeding area earlier than younger ones, mate earlier and thus have the possibility of a second brood. The length of the tail feathers does not play a role in the competition of the males for nesting sites, but in the choice of the females, as has been established in experiments. One group of males had their tail feathers shortened by two centimeters and another group lengthened by this amount. Compared to an untreated control group, 85% of the males with the elongated tail feathers brooded a second time, but only 10% of the males with short tail feathers. The males with long tail feathers copulated twice as often with the female of a male with shortened tail feathers as the control group. With long tail feathers, flight performance is poorer when eating. Males with long tail feathers no longer capture large, fast-flying insects, but only small, slow-flying insects. Since their brood needs as much food as the males with shorter tail feathers, they have to prey more. As a result of this exertion, males develop shorter tail feathers again during the next moult. This limits the length of the feathers and the females choose the most experienced and most successful males in finding food.
  • In some bird species such as B. Manakin , Lyrebird or Bowerbirds , prepare the males before their courtship dance courses. The female selects the male based on the quality of the space or performance. Particularly attractive males can mate numerous females, while less attractive males cannot reproduce. Male bowerbirds equip their dance places with objects of striking color, the number of which attracts the female. Strong males destroy their competitors' dance floors and rob the jewelry material for their own space. Copulation takes place on the dance floor, but the simple brood nest built by the females is usually far away from the dance floor.

Heterozygosity hypothesis

A potentially important factor when choosing a partner is the genetic compatibility of a partner. Then the quality of a mating partner is determined by their own genetic makeup and therefore varies for different partners. According to the effect of heterozygosity , the “quality” of a certain gene (actually: allele ) is not determined absolutely, but only depending on the situation in connection with the genome of the respective partner. This z. B. the mating of females with several males, which is widespread in the animal kingdom, is explained as risk minimization in order to avoid partners with genetically unsuitable elements. The heterozygous offspring of genetically different parents should in particular have a particularly efficient immune system . Research on human reproductive biology can also be interpreted in this way and one hypothesis provides e.g. B. a connection between the goodness of the immune system and pheromones . The better the immune systems complement each other, i.e. the more different they are, the more attractive the partner's smell will be. Empirical tests of the hypothesis have shown an advantage in pairings with genetically compatible or different partners in some cases, and in some cases a partner choice based on appropriate markers has also been demonstrated.

Forced mating

In addition to intra- and intersexual selection, Pradhan and van Schaik point out the role of forced mating of females by males. If the females cannot evade the males, their choices may be limited by the males. The characteristics developed through intrasexual selection (e.g. body size, antlers, horns) are then not only used in the same-sex rivalry between males, but also as an alternative strategy to force mating with females. This should give the females a selective incentive to avoid such mating partners. This hypothesis can also explain why in most mammals the males have "weapons", while in birds ornaments predominate.

Another important factor is the harassment of females through unwanted male attempts to mate, even if the mating does not take place. In a study on the forest lizard it could be shown that if there is an experimentally generated excess of males in the population, the males can become a major cause of mortality for the females through constant harassment and attempts at mating. As a result, not only does the population increase fall in the case of excess males, as expected, but the population size even decreases. This creates a significant risk of extinction for the population. Something similar was found in a number of other species. In the case of fruit flies , attempts to mate by males that are specifically aimed at particularly fertile females can put them at a severe disadvantage, which weakens their advantage (in natural selection).

Gender ratio

With normal sexual reproduction, the gender distribution is in principle 1: 1. As early as 1930 RA Fisher showed that, in the absence of particular factors, being overweight in one sex leads to selection pressure on the other sex. The gender ratio is subject to sexual selection and an unequal gender ratio then has a strong effect on sexual selection. According to the theory, the gender ratio should tend in the direction of the sex with a higher potential, i.e. H. taking into account the investments of the respective parent in the offspring, certain reproductive rates may be postponed. The decisive factor is the gender ratio of the individuals involved in reproduction of reproductive age, which z. B. can be postponed by a higher youth mortality of a sex. The gender ratio that actually works biologically is called the “operational sex ratio” ( OSR). Hidden factors can have a decisive influence on this. Is z. For example, if the female is only ready to conceive a few days a year, if the male is more or less permanently ready to conceive, the number of females actually willing to mate at a given time may be much lower than that of the males, even if both are equally frequent. This shifted the operational gender ratio in favor of the males. It has the same effect if males or females reach sexual maturity earlier than the opposite sex.

Without parental care for the offspring, the superiority of the male sex in the reproductive rate caused by the size ratio of the sex cells (gametes) can often prevail and the operational gender ratio can be shifted in favor of the males. An exclusive care of the females for the offspring then reinforces this tendency and the excess of males leads to stronger competition between the males. At z. B. Most bird species, however, both sexes supply the offspring. In many species, the male is the sole provider of the offspring and the female does not participate significantly beyond the delivery of the eggs. In addition to a number of insect and fish species such as pipefish, this also includes some salamanders and bird species such as B. ratites . The sex ratio can then be shifted in favor of the females, as a result of which they then compete more strongly for mating partners and are more subject to sexual selection.

The operational sex ratio can be variable for species, e.g. B. when the mortality of one sex is more dependent on environmental factors than that of the other (e.g. larger males, lack of food). In such species, researchers have experimentally changed the ratio and observed the consequences. In the case of the mouth -brooding Galilean parsley Sarotherodon galilaeus , it was shown that the reproductive strategy is influenced by the gender ratio. In this species, sometimes both sexes, sometimes one alone, care for the offspring. If there is an excess of one sex, the other will leave its offspring more often. This can be explained by the higher relative costs an individual has when there are more potential mates available to them.

Sexually antagonistic selection

Characteristics that lead to reproductive success through sexual selection are mostly unevenly distributed between the sexes. The selection pressure on the respective sex can work in different directions if there is no common optimum for both sexes. This phenomenon is called “sexually antagonistic selection”, tends to lead to an increase in genetic variability and is possibly one of the most important factors for this.

Empirical evidence of the operation of sexually antagonistic selection has been found in a number of species, such as: B. in fruit flies or red deer . The red deer z. It has been shown, for example, that daughters of fathers who were particularly successful in reproductive terms had less reproductive success than the average. This finding is also a serious problem for hypotheses such as B. the handicap hypothesis, which predicts greater success for the offspring of both sexes. According to the model, mutated alleles should advantageously only accumulate in the male sex on the X chromosome, because here they can have an effect on the male, while their effect on the female in the heterozygous case can be reduced by the allele on the second DNA strand. This prediction was confirmed with the fruit fly.

Sexually antagonistic selection can lead to an “arms race” between the sexes. This "sexually antagonistic coevolution" was z. B. shown in the seed beetles . In the males of many species, the Aedeagus has thorns that can injure the female during copulation. The females respond by strengthening the genital tract.

Mating systems and ecological constraints

The structure and formation of social and mating systems are not subject to sexual selection alone. The interactions between pairing systems and ecological boundary conditions, i. H. The compulsion given by natural selection is the subject of a separate research program. The framework conditions for the effect of sexual selection are therefore determined by environmental factors, in particular the distribution of resources in the environment ( ecological constraints model , ECM).

The almost unmanageable variety of mating systems in the animal kingdom (for mammals, cf.) can be classified according to the way of life of the respective species. Whether an individual can monopolize one or more potential mating partners for himself, i. H. Excluding other conspecifics from mating depends in a predictable way on the type of diet and lifestyle. If the contribution of both sexes is absolutely necessary for the successful rearing of boys, this results in (obligatory) monogamy. If the environmental factors suggest a territorial way of life (ie spatial restriction to a “home territory”) for females, males can monopolize females for themselves by denying access to other males. In the case of widely dispersed females, (facultative) monogamy results - in contrast to the obligatory, the male here may or may not contribute to rearing young. Those living together in groups or herds result in polygamy. If females live together in stable groups with no territory, single males (or a coalition of such) can restrict other males from entering these groups. If none of these requirements are met, it is usually more advantageous for males to individually look for as many mating partners as possible (promiscuity). Other environmental factors, e.g. B. Predators can have similar effects. For example, in langurs (a group of Asian monkeys), two males live together with a group of females if there are monkey-hunting eagles in the habitat. If these are missing, there is only one male.

The formation of family associations also depends in a predictable way on the environment. The formation of a family from parents and (adolescent) offspring is associated with a renunciation of reproductive possibilities for the offspring. This can be beneficial when the risks and costs of dispersion and hunting are high, e.g. B. because good areas are scarce.

Speciation

By z. B. Self-running processes can split species into daughter species as a result of diverging sexual preferences. Populations of a species evolve into separate species when isolation mechanisms prevent gene flow between organisms in those populations. Prezygotic isolation caused by sexual selection, morphological peculiarities or special mating signals can prevent mating if the signals cannot be interpreted or the partners are unattractive. Species formation through prezygotic mechanisms seems to proceed faster than through postzygotic mechanisms (e.g. sterility or inability to survive of the offspring) when the respective populations are in contact with one another. Since, according to theoretical models, sexual selection can proceed faster than natural selection, isolating mechanisms can develop quickly without the need for stronger, ecologically effective adaptations . As a starting point, small differences in the preference of the females between different populations, e.g. B. in the color patterns of guppies . According to the runaway selection model, such differences are sufficient, regardless of an adaptive value, to initiate a feature shift that takes place very quickly. Through adaptive radiation then can quickly types into groups of species split. This model is used to explain the biodiversity of the extremely rapidly evolving cichlid species in the East African lakes. In speciation, assortative mating is an important mechanism by which males and females with similar specializations or adaptations prefer each other when mating.

In the event of a split, behavioral characteristics should change to morphological characteristics beforehand. As the basis of an evolutionary change, these features must be at least partially hereditary. Little is known about the genetic basis of such behavioral traits. Classical breeding experiments show that it is usually quantitative traits that are influenced by many genes. The investigations are carried out on model organisms, especially on fruit flies with quantitative trait locus (QTL). In addition to their involvement in behavioral traits, many of the genes that act are often of fundamental importance for other biological processes ( pleiotropy ).

Sexual selection in humans

Some researchers close to sociobiology apply the theory of sexual selection to the species Homo sapiens and call the research approach evolutionary psychology . Your numerous opponents from the social sciences , especially social constructivism or feminist theory , speak of biologism or essentialism . For ethical reasons, experiments can only be carried out to a limited extent on humans and research into the causes of human social behavior has an impact on our self-image or the legitimacy of political and social systems. In addition, an application of psychological knowledge that z. B. were collected on academically educated adults in industrialized countries, problematic in other cultures . Relationships in Paleolithic hunter-gatherer communities, to which there is no direct access, were probably decisive for the evolution of human social behavior. The following methods are therefore used:

For example, David Buss examined possible evolutionary reasons for human jealousy, female sexual behavior or the causes of murderous intentions. Geoffrey Miller shed light on the question of how the human brain could develop and, like Gad Saad, what role consumption plays in Homo sapiens.

Mating strategies and systems

As a special feature of humans, both sexes take part in the care and rearing of the offspring. No other ape species, the males involved in the care of the offspring, either by living in family groups chimpanzee species , nor in the Gorilla - harems , the monogamous living Gibbons or solitary orangutans . For single mothers, empirical social research has shown a significant increase in child mortality in hunter-gatherer and agricultural cultures and to a certain extent in modern societies, but not in the most highly developed industrial and welfare states.

According to the theoretical predictions of sexual selection theory, both sexes have a common evolutionary interest in investing in the welfare of the offspring. A “mixed” strategy is easier for fathers than for mothers, who are less likely to reduce the supply, at least for their babies. For men, investing in a woman's offspring is evolutionarily advantageous if they can be sure of their fatherhood and thus their reproductive success. For women, however, benefit considerations play a role when choosing a partner. B. apply to monogamous songbird species. You can bind a presumably reliably helping “provider” and his resources or a genetically attractive, but perhaps unreliable man who will pass on his “good genes” to your offspring and whose “sexy” sons then have better chances of mating. Reputation for monogamy gained through “ morality ” can also apply to men, e.g. Pay off, for example, through a reputation as a “good provider”, especially when raising children from outside your home.

There is little data on the social life of the immediate ancestors of man, and societies living “originally” are examined instead. A connection between monogamy and raising the offspring together has not been empirically proven. Monogamy occurs in less than 5% of mammalian species. When comparing human cultures, there are monogamous , polygynous and polyandric relationships. People live together in social associations, the size of which in hunter-gatherer cultures is primarily limited by natural resources, so that pairing cannot be secured by territoriality, for example. The size of these cohabiting groups is usually around 30 individuals. For example, in order to avoid inbreeding, some of the offspring of social animal species leave the territory, while the other part remains philopatric . In most animal species, the young males leave the social association, but in the case of the great apes (chimpanzees and gorillas), which are closest related to Homo sapiens, the young females. For decades, there has been a scientific controversy about philosophy in human hunter-gatherer cultures. In this regard, there appear to be great differences between human cultures and it has been argued that there was a great evolutionary advantage in this for the species of humans.

An average man is taller, a lot heavier and stronger than an average woman, which often correlates with polygynous behavior, while in purely monogamous relationships equally large genders are to be expected. In comparison to the genus Australopithecus , however, the sexual size dimorphism is considerably smaller in humans. Compared to monogamous primates, a man has larger testicles and produces "kamikaze" sperm. The human penis is unusually long for primates. B. interpreted as an ornament towards competitors or interpreted as the result of a sperm competition . From this it is concluded that mating behavior is moderately polygamous, but this was also denied for modern humans. Meanwhile, further arguments in favor of polygyny over long periods of human evolution have been advanced through genetic studies on X chromosomes.

In addition to menopause , the size of the female breasts , hidden ovulation , the strong variance in the length of the menstrual cycle with only a 28% chance of conception per cycle and the hidden coitus are unique to the great apes and are used as strategies for controlling conception and as protection against male aggressiveness interpreted. Women place different demands on long-term partners than on men for short sexual contacts. Studies have found that women can schedule meetings with lovers at the time of their ovulation , have more orgasms than with regular partners, and hold back more sperm that remains fertile in crypts for up to 7 days. Some researchers doubt such connections and see no evolutionary adaptation in the female orgasm , for example . From the 1940s onwards, depending on the study and interpretation, investigations revealed 5% to 30% cuckoo children who are sociobiologically seen as the result of a “combined reproductive strategy” (KFS). In the upper class 2%, 12% in the middle class and 20% in the lower class were found to be cuckoo children. Different studies showed different values ​​for the KFS. According to this, 20–40% of women and 30–50% of men in the US have at least one affair during their marriage. The likelihood of affairs was higher with attractive partners or personality traits such as the dark triad , narcissism or sociopathy, and women are more emotionally motivated and men are more visually sexually motivated.

Studies on the possible influence of sexual selection on the development of the human body show different tendencies of women and men to the physical appearance of the opposite sex.

The cross-cultural interest of women in older men, especially for long-term relationships such as B. Marriage or cohabitation is explained by the economic security that older men can often offer. Studies show the preference of women over men with social status and potential as well as financial viability and of men over sexually faithful and younger women of the most fertile age. To attract the attention of potential partners, men demonstrate possession and position, and women emphasize their physical appearance . For long-term relationships, both genders prefer partners who are interesting, intelligent, humorous, altruistic , reliable and familiar. When dealing with competitors , women use a number of verbal and z. T. subtle techniques that change the perception of men such as B. change associatively or ambivalently .

violence

Jealousy , stalking or violence in couples is e.g. B. by David Buss interpreted as evolutionary adaptation and such behavior as an attempt to deter, control and increase costs in the event of infidelity or change of partner. According to controversially discussed studies, this cross-cultural violence is used equally frequently by both sexes. Women often only use physical violence as part of self-defense, with 79% of women who kill their partner being younger than 40 years. According to homicide investigations, women are three times more likely to be killed by a partner than by a stranger. Young women, women with a significantly older or less attractive partner are at higher risk. In the case of a separation, the risk of being killed is 5 to 7 times higher, 78–91% of the murders are committed in the first year after the separation, with 81% of the perpetrators being under 50 years of age. Women with children from previous partnerships are 12 times more likely to be killed in a separation. Young stepchildren are 40 to 100 times more likely to be killed, and stepchildren leave home an average of 2 years earlier than biological children.

Stalking is interpreted as a behavior to e.g. B. To keep partners, to win (back) or to eliminate rivals. In a mild form, stalking is used by both sexes, in a harassing or violent form it is mainly used by men. After a breakup, stalking lasts an average of 2 years but can last from a few days to a decade, and 88% of the men who killed their wives were previously stalkers.

Forced mating is interpreted as control over the reproductive worth of partners or sperm competition. Sexually unfaithful women or women with a jealous partner are 2 to 5 times more likely to be raped . According to studies, regardless of social class, 13-25% of women are violently sexually assaulted or raped before the age of 40. Contrary to the fear of more than 90% of women of sexual violence and rape murder by strangers, according to statistics 80–90% of women are raped by acquaintances and 10–26% of married women are raped by their sexual partners. The killing rate for rape is 0.01–2% and is higher in wartime .

Gender distribution

In Homo sapiens, the gender distribution at birth is usually around 1.05 male to 1.00 female. However, the mortality of boys is somewhat higher than that of girls, so that the ratio is almost balanced when entering puberty. Through the targeted abortion of female fetuses , artificial gender selection has been practiced for several decades in several Asian countries such as China , India , Azerbaijan , Georgia or Armenia , but also in Albania, thus introducing further selection pressure. As a result, there are an estimated 160 million fewer women. For example, it is forecast for China that one in ten men in 2013 and one in five between 2020 and 2045 will not be able to find a wife.

Sexual selection as an explanatory approach for the evolution of intelligence and culture

According to researchers Robin Dunbar or Geoffrey Miller , sexual selection played an important role in the creation of the human mind. The hypothesis postulates the performance of the brain as an adaptation to the requirements of a complex social life and not in order to e.g. B. better to get food or to be able to avoid predators. One of the special requirements would be e.g. B. the specific social couple bond in humans, in which the intelligence and related skills of the partner are decisive when choosing a partner. Large parts of human social life and a large number of human cultural achievements are then interpreted as an ornament or pairing prelude in order to showcase one's own special capabilities in the possible mating partner, such as B. Music or morals. With his book The Art Instinct, the philosopher Denis Dutton also presents the theory of an evolutionary aesthetic that interprets human aesthetics as an evolutionary product.

In contrast to the pair bond, an alternative theory explains the evolution of the brain from larger social associations for humans, which are otherwise rare in the animal kingdom. The evolution of larger brains, which is advantageous for each species, and thus increased intelligence, soon reaches its limits for most species due to energetic constraints. The expenditure associated with brain growth and maturation increases the individual development time so much that even better survival rates through higher intelligence can no longer compensate for this. This limit can only be overcome if other conspecifics - in addition to the mother - take part in raising the young. This could e.g. B. explain the relatively larger average brain size in birds compared to mammals. Humans are one of the few species in which relatives do not directly participate in raising children, which is possibly a decisive starting advantage.

Evolutionary psychology has recently turned increasingly to the question of what role individual differences have played in evolution. The emotional reactions in long-term relationships, whose partners have a common life story, are constitutive for human individuality and personality. On this basis, the philosophical anthropologist Ferdinand Fellmann introduced the term “emotional selection” as an extension of sexual selection in the context of the couple bond model of the incarnation.

Remarks

  • There are some non-monogamous animal species that have no apparent sex dimorphism. In such rare cases, signals imperceptible to humans such as B. Pheromones play a role. In blue tits z. B. signals detected in the ultraviolet spectral range.
  • In some ungulates , both sexes have horns or antlers. In this case, they serve not only for sexual selection, but also have other functions, for example defense against predators . They can then be of different sizes between the sexes (example: reindeer ) or differently shaped (example: bighorn sheep ).

literature

Web links

Sexual selection on YouTube , September 19, 2014 (A brief description of the evolutionary mechanism of sexual selection)

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