Mate choice, also known as intersexual selection, is an evolutionary process in which selection is dependent on the attractiveness of an individual's phenotypic traits. Evolutionary change is possible because the qualities that are desired in a mate are more frequently passed on to each generation over time. For example, if female peacocks desire mates who have a colourful plumage, then this trait will increase in frequency over time as male peacocks with a colourful plumage will have more reproductive success.
Mate choice is one of two components of sexual selection, the other being intrasexual selection. Ideas on sexual selection were first introduced in 1871, by Charles Darwin, then expanded on by Ronald Fisher in 1915. At present, there are five mechanisms that explain how mate choice has evolved over time. These are direct phenotypic benefits, sensory bias, the Fisherian runaway hypothesis, indicator traits and genetic compatibility.
In systems where mate choice exists, one sex is competitive with same-sex members and the other sex is choosy (which means they are selective when it comes to picking individuals to mate with). There are direct and indirect benefits of being the selective individual. In most species, females are the choosy sex which discriminates among competitive males, but there are several examples of reversed roles (see below). It is preferable for an individual to choose a compatible mate of the same species, in order to maintain reproductive success. Other factors that can have an impact on mate choice include pathogen stress and the Major Histocompatibility Complex (MHC).
Origins and historyEdit
Charles Darwin first expressed his ideas on sexual selection and mate choice in his book The Descent of Man, and Selection in Relation to Sex in 1871. He was perplexed by the elaborate ornamentation that males of some species have, because such features appeared to be detrimental to survival and to have negative consequences for reproductive success. Darwin proposed two explanations for the existence of such traits: these traits are useful in male-male combat or they are preferred by females. This article focuses on the latter. Darwin treated natural selection and sexual selection as two different topics, although in the 1930s biologists defined sexual selection as being a part of natural selection.
In 1915, Ronald Fisher wrote a paper on the evolution of female preference and secondary sexual characteristics. Fifteen years later, he expanded this theory in a book called The Genetical Theory of Natural Selection. There he described a scenario where feedback between mate preference and a trait results in elaborate characters such as the long tail of the male peacock (see Fisherian runaway).
In 1948, using Drosophila as a model, Angus John Bateman presented experimental evidence that male reproductive success is limited by the number of mates obtained, while female reproductive success is limited by the number of pregnancies that she can have in her lifetime. Thus a female must be selective when choosing a mate because the quality of her offspring depends on it. Males must fight, in the form of intra-sexual competition, for the opportunity to mate because not all males will be chosen by females. This became known as Bateman's principle, and although this was a major finding that added to the work of Darwin and Fisher, it was overlooked until George C. Williams emphasised its importance in the 1960s and 1970s.
In 1972, soon after Williams' revival of the subject, Robert L. Trivers presented his parental investment theory. Trivers defined parental investment as any investment made by the parent that benefits his or her current offspring at the cost of investment in future offspring. These investments include the costs of producing gametes as well as any other care or efforts that parents provide after birth or hatching. Reformulating Bateman's ideas, Trivers argued that the sex which exhibits less parental investment (not necessarily the male) will have to compete for mating opportunities with the sex that invests more. The differences in levels of parental investment create the condition that favours mating biases.
Direct and indirect benefitsEdit
Being choosy (having a bias in the context of mating) must incur a fitness advantage in order for this behaviour to evolve. Two types of fitness benefits (direct and indirect) are thought[by whom?] to drive the evolutionary mechanisms of mate choice:
- Direct benefits increase the fitness of the choosy sex through direct material advantages or resources. These benefits include but are not limited to increased territory quality, increased parental care, and protection from predators. There is much support for maintenance of mate choice by direct benefits and this approach offers the least controversial model to explain discriminate mating.
- Indirect benefits increase genetic fitness for the offspring, and thereby increase the parents' inclusive fitness. When it appears that the choosy sex does not receive direct benefits from his or her mate, indirect benefits may be the payoff for being selective. These indirect benefits may include high-quality genes for their offspring (known as adaptive indirect benefits) or genes that make their offspring more attractive (known as arbitrary indirect benefits).
Currently, there are five mechanisms that explain the evolution of mate choice:
- Direct phenotypic benefits
- Sensory bias
- Fisherian runaway
- Indicator traits
- Genetic compatibility
Direct and/or indirect benefits are driving the mating biases described in each mechanism. It is possible that these mechanisms co-occur although the relative roles of each have not been evaluated adequately.
Direct phenotypic benefitsEdit
Choosy individuals receive direct benefits from their mates and this results in immediate increased fecundity, or the number of offspring produced. If the competitive sex displays an ornamental trait that reliably indicates some direct benefit then strong selection will favor mating bias. Having a mating preference is advantageous in this situation because it directly affects reproductive fitness. Direct benefits are widespread and the evidence for this mechanism of evolution is well supported in empirical studies.
One example of a sexually selected trait with direct benefits is the bright plumage of the northern cardinal, a common backyard bird in the eastern United States. Male northern cardinals have conspicuous red feathers while the females are more cryptic in coloration. In this example, the females are the choosy sex and will use male plumage brightness as a signal when picking a mate because males with brighter plumage have been shown to feed their young more frequently than males with duller plumage. This increased help caring for the young lifts some of the burden from the mother so that she can raise more offspring than she could without help.
In the great reed warbler, females tend to be attracted to males with longer song repertoires since they tend to sire offspring with improved viability. In doing so, they gain indirect benefits for their own young. In the Utetheisa ornatrix, females select males based on body size, systemic content of pyrrolizidine alkaloid, and glandular content of hydroxydanaidal. As a result, these females demonstrate direct and indirect phenotypic benefits: they have offspring that are less vulnerable to predation because of their increased size and higher alkaloid content, increasing viability and fitness.
The sensory bias hypothesis states that the preference for a trait evolves in a non-mating context and is then exploited by one sex in order to obtain more mating opportunities. The competitive sex evolves traits that exploit a pre-existing bias that the choosy sex already possesses. This mechanism is thought to explain remarkable trait differences in closely related species because it produces a divergence in signaling systems which leads to reproductive isolation.
Sensory bias has been demonstrated in guppies, freshwater fish from Trinidad and Tobago. In this mating system, female guppies prefer to mate with males with more orange body coloration. However, outside of a mating context, both sexes prefer animate orange objects which suggests that preference originally evolved in another context, like foraging. Orange fruits are a rare treat that fall into streams where the guppies live. The ability to find these fruits quickly is an adaptive quality that has evolved outside of a mating context. Sometime after the affinity for orange objects arose, male guppies exploited this preference by incorporating large orange spots to attract females.
Another example of sensory exploitation is in the water mite Neumania papillator, an ambush predator which hunts copepods (small crustaceans) passing by in the water column. When hunting, N. papillator adopts a characteristic stance termed the 'net stance' - their first four legs are held out into the water column, with their four hind legs resting on aquatic vegetation; this allows them to detect vibrational stimuli produced by swimming prey and use this to orient towards and clutch at prey. During courtship, males actively search for females - if a male finds a female, he slowly circles around the female whilst trembling his first and second leg near her. Male leg trembling causes females (who were in the 'net stance') to orient towards often clutch the male. This did not damage the male or deter further courtship; the male then deposited spermatophores and began to vigorously fan and jerk his fourth pair of legs over the spermatophore, generating a current of water that passed over the spermatophores and towards the female. Sperm packet uptake by the female would sometimes follow. Heather Proctor hypothesised that the vibrations trembling male legs made were done to mimic the vibrations that females detect from swimming prey - this would trigger the female prey-detection responses causing females to orient and then clutch at males, mediating courtship. If this was true and males were exploiting female predation responses, then hungry females should be more receptive to male trembling - Proctor found that unfed captive females did orient and clutch at males significantly more than fed captive females did, consistent with the sensory exploitation hypothesis.
Other examples for the sensory bias mechanism include traits in auklets, wolf spiders, and manakins. Further experimental work is required to reach a fuller understanding of the prevalence and mechanisms of sensory bias.
Fisherian runaway and sexy sons hypothesisEdit
These hypotheses refer to a coupled selection for females to be attracted and, likewise, a selection for males to be attractive. This can lead to self-reinforcing coevolution. If runaway selection is strong enough, it may incur significant costs such as increased visibility to predators and energetic costs to maintain the trait's full expression. Hence peacocks' extravagant feathers, or any number of lek mating displays. This model does not predict a genetic benefit; rather, the reward is more mates.
In a study done on great reed warblers, models based on the polygyny threshold and sexy son hypotheses predict that females should gain evolutionary advantage in either short-term or long-term in this mating system. Although the importance of female choice was demonstrated, the study did not support the hypotheses. Other studies, such as those conducted on long-tailed widowbirds, have demonstrated the existence of female choice. Here, females chose males with long tails, and even preferred those males with experimentally lengthened tails over shortened tails and those of naturally occurring length. Such a process shows how female choice could give rise to exaggerated sexual traits through Fisherian runaway selection.
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Indicator traits are those that signal good overall quality of the individual. Traits that are perceived as attractive must reliably indicate broad genetic quality in order for selection to favor them and for preference to evolve. This is an example of indirect genetic benefits received by the choosy sex because mating with such individuals will result in high quality offspring. The indicator traits hypothesis is split into three highly related subtopics: the handicap theory of sexual selection, the good genes hypothesis, and the Hamilton-Zuk hypothesis.
People rate the importance of certain traits differently when referring to their own or to others ideal long term partner's. Research suggests that women consider traits indicating genetic fitness as more important for their own partner, and traits that provide benefits to others are proritised for their sister's ideal partner.
Indicator traits are condition-dependent and have associated costs. Therefore, individuals that can handle these costs well (cf. "I can do X [here, survive] with one hand tied behind my back") should be desired by the choosy sex for their superior genetic quality. This is known as the handicap theory of sexual selection.
The good genes hypothesis states that the choosy sex will mate with individuals who possess traits that signify overall genetic quality. In doing so, they gain an evolutionary advantage for their offspring through indirect benefit.
The Hamilton-Zuk hypothesis posits that sexual ornaments are indicators of parasite and disease resistance. To test this hypothesis, red jungle fowl males were infected with a parasitic roundworm and monitored for growth and developmental changes. Female preference was also evaluated. What the researchers found was that parasites affected the development and final appearance of ornamental traits and that females preferred males who were not infected. This supports the idea that parasites are an important factor in sexual selection and mate choice.
One of many examples of indicator traits is the condition-dependent patch of red feathers around the face and shoulders of the male house finch. This patch varies in brightness among individuals because the pigments that produce the red color (carotenoids) are limited in the environment. Thus, males who have a high quality diet will have brighter red plumage. In a manipulation experiment, female house finches were shown to prefer males with brighter red patches. Also, males with naturally brighter patches were better fathers and exhibited higher offspring feeding rates than duller males. This study is heavily cited in the literature and it provides solid support for the indicator traits hypothesis that is associated with direct benefits.
Genetic compatibility refers to how well the genes of two parents function together in their offspring. Choosing genetically compatible mates could result in optimally fit offspring and notably affect reproductive fitness. However, the genetic compatibility model is limited to specific traits due to complex genetic interactions (e.g. major histocompatibility complex in humans and mice). The choosy sex must know their own genotype as well as the genotypes of potential mates in order to select the appropriate partner. This makes testing components of genetic compatibility difficult and controversial.
A controversial but well known experiment suggests that human females use body odor as an indicator of genetic compatibility. In this study, males were given a plain T-shirt to sleep in for two nights in order to provide a scent sample. College women were then asked to rate odors from several men, some with similar MHC (major histocompatibility complex) genes to their own and others with dissimilar genes. MHC genes code for receptors that identify foreign pathogens in the body so that the immune system may respond and destroy them. Since each different gene in the MHC codes for a different type of receptor it is expected that females will benefit from mating with males who have more dissimilar MHC genes. This will ensure better resistance to parasites and disease in the offspring. Researchers found that women tended to rate the odors higher if the male's genes were more dissimilar to their own. They conclude that the odors are influenced by the MHC and that they have consequences for mate choice in human populations today.
Similar to the humans of the odor rating experiment, animals also choose mates based upon genetic compatibility as determined by evaluating their potential mate(s) body odor. Some animals, such as mice even assess a mate's genetic compatibility based on their urine odor.
In an experiment studying three-spined sticklebacks, researchers found that females prefer to mate with males that share a greater diversity of major histocompatibility complex (MHC) and in addition possess a MHC halotype specific to fighting the common parasite Gyrodactylus salaris. Mates that have MHC genes different from one another will be superior when reproducing with regard to parasite resistance, body condition and reproductive success and survival.
The genetic diversity of animals and life reproductive success (LRS) at the MHC level is optimal at intermediate levels rather than at its maximum, despite MHC being one of the most polymorphic genes. In a study, researchers discovered that mice heterozygous at all MHC loci were less resistant than mice homozygous at all loci to salmonella, so it appears disadvantageous to display many different MHC alleles due to the increased loss of T-cells, which aid an organism's immune system and trigger its appropriate response.
MHC diversity may also be correlated to MHC gene expression. As long as a heritable component exists in expression patterns, natural selection is able to act upon the trait. Therefore, gene expression for MHC genes might contribute to the natural selection processes of certain species and be in fact evolutionarily relevant. For example, in another study of three-spined sticklebacks, exposure to parasite species increased MHC class IIB expression by over 25%, proving that parasitic infection increases gene expression.
MHC diversity in vertebrates may also be generated by the recombination of alleles on the MHC gene.
Sex role reversal in animalsEdit
In species where mating biases exist, females are typically the choosy sex because they provide a greater parental investment than males. However, there are some examples of sex role reversals where females must compete with each other for mating opportunities with males. Species that exhibit parental care after the birth of their offspring have the potential to overcome the sex differences in parental investment (the amount of energy that each parent contributes per offspring) and lead to a reversal in sex roles. The following are examples of male mate choice (sex role reversal) across several taxa.
- Fish: Male fish typically display high levels of parental care (see pipefish, scissortail sergeant, and seahorses). This is because females will deposit their eggs in a special brooding pouch that the male possesses. She doesn't participate in parental care after this event. The male then has the burden of raising the offspring on his own which requires energy and time. Thus, males in these species must choose among competitive females for mating opportunities. Surveys across multiple species of pipefish suggest that the sex differences in the level of parental care may not be the only reason for the reversal. Mating systems (e. i. monogamy and polygamy) might also heavily influence the appearance of male mate choice.
- Amphibian: Male poison-arrow frogs (Dendrobates auratus) take on a very active parenting role. Females are lured by the males to rearing sites where they deposit their eggs. The male fertilises these eggs and accepts the burden of defending and caring for the young until they are independent. Because the male contributes a higher level of parental investment, females must compete for opportunities to leave their eggs with the limited available males.
- Bird: Bird species are typically biparental in care, and may also be maternal like the Guianan cock-of-the-rocks. However the reverse may also hold true. Male wattled jacanas provide all parental care after the eggs have been laid by the females. This means that the males must incubate the eggs and defend the nest for an extended period of time. Since males invest much more time and energy into the offspring, females are very competitive for the right to lay their eggs in an established nest.
- Mammal: There are no confirmed cases of sex role reversed mammals but female spotted hyenas have peculiar anatomy and behaviour that has warranted much attention. Female spotted hyenas are much more aggressive than males due to their high levels of androgens during development. The increased male hormones during development contribute to an enlarged pseudopenis that is involved in mating and birth. Although the anatomical and behavioural roles differ from accepted norms, spotted hyenas are not sex role reversed because the females do not compete with each other for mates.
For many years it has been suggested that sexual isolation caused by differences in mating behaviours is a precursor for reproductive isolation (lack of gene flow), and consequently speciation, in nature. Mate choice behaviours are thought to be important forces that can result in speciation events because the strength of selection for attractive traits is often very strong. Speciation by this method occurs when a preference for some sexual trait shifts and produces a pre-zygotic barrier (preventing fertilisation). These processes have been difficult to test until recently with advances in genetic modelling. Speciation by sexual selection is gaining popularity in the literature with increasing theoretical and empirical studies.
There is evidence of early speciation through mate preference in guppies. Guppies are located across several isolated streams in Trinidad and male colour patterns differ geographically. Female guppies have no coloration but their preference for these colour patterns also vary across locations. In a mate choice study, female guppies were shown to prefer males with colour patterns that are typical of their home stream. This preference could result in reproductive isolation if two populations came into contact again.
The black-throated blue warbler, a North American bird, is another example. Asymmetric recognition of local and non-local songs has been found between two populations of black-throated blue warblers in the United States, one in the northern United States (New Hampshire) and the other in the southern United States (North Carolina). Males in the northern population respond strongly to the local male songs but relatively weakly to the non-local songs of southern males. In contrast, southern males respond equally to both local and non-local songs. The fact that northern males exhibit differential recognition indicates that northern females tend not to mate with "heterospecific" males from the south; thus it is not necessary for the northern males to respond strongly to the song from a southern challenger. A barrier to gene flow exists from South to North as a result of the female choice, which can eventually lead to speciation.
Mate choice in humansEdit
In humans, males and females differ in their strategies to acquire mates and focus on certain qualities. The strategies that each gender uses differs in regards to whether they are long-term or short-term. Human mate choice depends on a variety of factors, such as genes, negative traits, and parasite stress.
Female mate choiceEdit
Although, in humans, both males and females are choosy in terms of whom they decide to mate with, as is seen in nature, females exhibit even more mate choice selection than males. According to Bateman's principle of Lifespan Reproductive Success (LRS), human females display the least variance of the two sexes in their LRS due to their high obligatory parental investment, that is a nine-month gestational period, as well as lactation following birth in order to feed offspring so that their brain can grow to the required size.
Historically, human female sexual selection can be examined by looking at ways in which males and females are sexually dimorphic, especially in traits that serve little other evolutionary purpose. For example, male traits such as the presence of beards, overall lower voice pitch, and average greater height are thought to be sexually selected traits as they confer benefits to either the women selecting for them, or to their offspring. Experimentally, women have reported a preference for men with beards and lower voices.
Female mate choice hinges on many different coinciding male traits, and the trade off between many of these traits must be assessed. The ultimate traits most salient to female human mate choice, however, are parental investment, resource provision and the provision of good genes to offspring. Many phenotypic traits are thought to be selected for as they act as an indication of one of these three major traits. The relative importance of these traits when considering mate selection differ depending on the type of mating arrangement females engage in. Human women typically employ long term mating strategies when choosing a mate, however they also engage in short term mating arrangements, so their mate choice preferences change depending on the function of the type of arrangement.
Short term mating strategiesEdit
Women do not always seek out and engage in long term mating arrangements. This is evidenced by factors such as the evolved male tendency to seek out multiple sexual partners - a trait that could not have evolved if women were not also historically engaging in short term arrangements - and by the tendency of some women to pursue affairs outside of their long-term couple pairings.
David Buss outlines several hypotheses as to the function of women's short term mate choices:
- Resource hypothesis: Women may engage in short term mating in order to gain resources that they may not be able to gain from a long-term partner, or that a long-term partner may not be able to provide consistently. These resources may be food, protection for the woman and her children from aggressive men who may capture or sexually coerce them, or status, by providing the woman with a higher social standing. Women may also benefit from having several short term mating arrangements through paternity confusion - if the paternity of her offspring is not certain, she may be able to accrue resources from several men as a result of this uncertainty.
- Genetic benefit hypothesis: Women may choose to engage in short term mating arrangements in order to aid conception if her long term partner is infertile, to gain superior genes to those of her long term partner, or to acquire different genes to those of her partner and increase the genetic diversity of her offspring. This relates to what is known as the sexy son hypothesis; if a woman acquires genes from a high quality male, her offspring will likely have higher mate value, resulting in their increased reproductive success.
- Mate expulsion and mate switching: Women may engage in a short term mating arrangement in order to cause her long term partner to end their relationship; in other words, to facilitate a break-up. Women may also use short term mating if their current partner has depreciated in value, and they wish to 'trade-up' and find a partner that they believe has higher value.
- Short term for long term goals: Women may use short term sexual relationships in order to assess a mate's value as a long term partner, or in the hopes that the short term arrangement will result in one that is long term.
Long term mating strategiesEdit
In long term mating arrangements, women typically look for males who will provide a high level of parental investment, and who can provide resources to the woman or to her offspring. The provision of economic resources, or the potential to acquire many economic resources is the most obvious cue towards the ability of a man to provide resources, and women have been shown experimentally to rate the importance of their partner's financial status twice as highly as men. However, many other traits exist that may act as cues towards a man's ability to provide resources that have been sexually selected for in women's evolutionary history. These include older age – older males have had more time to accrue resources - industriousness, dependability and stability – if a woman's long term partner is not emotionally stable or is not dependable then their provision of resources to her and her offspring are likely to be inconsistent. Additionally, the costs associated with an emotionally unstable partner such as jealousy and manipulativeness may outweigh the benefits associated with the resources they are able to provide.
Women's mate choice is not as straightforward as selecting a mate that displays all of her desired qualities. Often, potential mates will possess some qualities that are desirable and some that are not, so women must assess the relative costs and benefits of their potential partners' traits and 'trade off'. Women's mate choices will also be constrained by the context in which they are making them, resulting in conditional mate choices. Some of the conditions that may influence female mate choice include the woman's own perceived attractiveness, the woman's personal resources, mate copying and parasite stress.
Male mate choiceEdit
Generally, it is unusual for males within a species to be the choosy sex. There are many reasons for this. In humans, following sexual reproduction, the female is obliged to endure a nine-month pregnancy and childbirth. This means that females naturally provide a greater parental investment to offspring, than males. Human males have a larger quantity of gametes than females, which are replenished at a rate of approximately 12 million per hour. Conversely, female humans are born with a fixed amount of egg cells which are not restocked over the lifespan. This provides males with a greater window of opportunity to mate and reproduce than females, hence why females are usually more choosy.
Despite not being the typically choosy gender, human males can be influenced by certain traits of females when making decisions about a potential mate:
Short term mating strategiesEdit
When finding a short term mate, males highly value women with sexual experience and physical attractiveness. Men seeking short term sexual relationships are likely to avoid women who are interested in commitment or require investment.
Examples of short term mating strategies in males:
- Multiple sexual partners: When looking for short term sexual relationships, men may wish for there to be as little time as possible between each partner. When engaging in sexual intercourse with multiple partners, it's important to be aware that the risk of contracting a sexually-transmitted disease may increase if contraception isn't used.
- Physical attractiveness: Men who are interested in a short term sexual relationship are more likely to prioritise information about the body of potential partners, rather than their faces. When finding a female for a short term relationship, compared with a long term relationship, males are less likely to prioritise factors such as commitment.
- Relaxation of standards: It has been reported that men are more likely to engage in a sexual relationship with women who have a lower levels of intelligence, independence, honesty, generosity, athleticism, responsibility and cooperativeness, when this relationship is short term. Men may be more accepting of lower standards, than what they usually prefer, because they are not entering a long term relationship with this person.
- Sexual experience: Many men assume that women who have engaged in sexual experiences beforehand are likely to have a higher sex drive than women who haven't. These women may also be more accessible and require less courtship.
Long term mating strategiesEdit
Although from an evolutionary perspective women are usually the choosy sex, if a human male has the desire to reproduce he may seek certain qualities in a potential mate who could be the mother of his offspring. Humans have the ability to rely on biological signals of reproductive success and non-biological signals, such as the female's willingness to marry. Unlike many animals, humans are not able to consciously display physical changes to their body when they are ready to mate, so they have to rely on other forms of communication before engaging in a consensual relationship.
Examples of long term mating strategies in males:
- Commitment and marriage: A human male may be interested in mating with a female who seeks marriage. This is because he has exclusive sexual access to the female, so any offspring produced in the relationship will be genetically related to him (unless the female has sexual intercourse with another male outside of the marriage). This increases the likelihood of paternity certainty. With two married parents investing in the offspring, their chance of survival may increase; therefore the male's DNA will be passed on to the children of his offspring. Also, a male who is interested in committing to a female may be more attractive to potential mates. A male who can promise resources and future parental investment is likely to be more appealing to women than a male who is unwilling to commit to her.
- Facial symmetry: Symmetrical faces have been judged to signal good general health and the ability for a woman to withstand adverse environmental factors, such as illness.
- Femininity: A feminine face can be a signal of youth, which in turn signals strong reproductive value. As a woman gets older, her facial features become less feminine due to ageing. Femininity can also be linked to disease-resistance and high estrogen levels, which are factors that suggest reproductive value to a potential mate.
- Physical beauty: Observable characteristics of a woman can indicate good health and the ability to reproduce, qualities which are likely to be desired by a male. This may include smooth skin, absence of lesions, muscle tone, long hair and high energy levels.
- Resources: Men who are looking for a long term partner may strive to achieve a high status or resources, such as their own home or a job promotion. This may increase their chance of attracting a desirable mate.
- Waist-to-hip ratio: A waist-to-hip ratio of 0.7 is an indicator of fertility, lower long term health risks and suggests that the woman isn't already pregnant. A male is likely to desire these qualities in a mate, as it will increase the chance of survival of any offspring the couple have together.
- Youth: Faces that appear younger are usually rated as more attractive by males. This could include faces which do not have many wrinkles or where there is very clear skin. A female who appears younger is likely to be appealing to mates, as it suggests that she has a higher reproductive value than alternative, older, females. As a woman passes her twentieth birthday, her reproductive value declines steadily until around the age of fifty.
Parasite-stress on human mate choiceEdit
The parasite-stress theory, otherwise known as pathogen stress, is where a parasite or disease stresses the development of organisms, leading to a change in their mate preference and choice. In societies with a high prevalence of parasites or pathogens greater emphasis is placed by the members of that society, on the physical attractiveness/good looks of their mates or potential mates, compared to members of societies with a lower prevalence of parasites or diseases who put less emphasis on physical attractiveness. It indicates that physical attractiveness is a way in which humans can determine resistance to parasites, as it's believed that parasites and diseases would worsen the look of those who are suffering or have suffered from a disease, and would also limit the number of high-quality pathogen-resistant mates.
The Hamilton-Zuk hypothesis (see Indicator traits) has greatly influenced research into human mate choice. It has been found that males of all cultures have been found to rate female attractiveness very highly, but when the risk of parasitic infection was high, males rated female attractiveness as much higher. In cultures where parasitic infection is especially high, members of that society use every cue available to them to determine the physical health status of the potential mate. Regardless of the wealth or ideology, the areas of a society i.e. that are more at risk or have higher rates of parasites and diseases, the females will rate masculinity as higher.
- Scarification: In pre-industrial societies body markings, such as tattoos or scarifications, have been suggested as a way in which can attract potential mates to the reproductive quality of a person. However, Singh and Bronstad (1997) found that when there was an increase in pathogen prevalence only female stomach scarification was predicted and no other anatomical area, with no evidence for male scarification.
- Masculinity: In societies where there are high levels of parasites or diseases, the females of that society, as the overall health of its members decreases, increasingly start to place more emphasis on masculinity in their mate preference. In particular, women look for increasing signs of masculinity in areas such as the voice, face and body shape of males. The face, in particular, may hold several cues for parasitic resistance and has been the subject of most attractiveness research.
- Polygamy: Tropical areas were originally associated with polygynous societies and this was a result of the surrounding environment being both ecologically richer and homogenous. However, whilst tropical areas were associated with Polygamy, pathogen stress, is regarded as a better indicator of polygamy and has been positively correlated with it. Further to that, over the course of human evolution, areas which had high levels of parasite-stress may have shifted the polygamy threshold and increased the amount certain types of polygamy present in a society.
Gangested and Buss (2009) say that whilst research indicates that parasite stress may have only influenced mate choice, females searching for "good genes" which show parasite resistance,in areas which have high prevalence of parasites. John Cartwright also points out that females may be simply avoiding the transmission of parasites to themselves rather than it being them choosing males with good genes and that females look for more than just parasite-resistant genes.
Major Histocompatibility Complex (MHC) or in humans, Human Leukocyte Antigen (HLA), produce protein products that are essential for the functioning of the immune system. The genes of MHC have extremely high variability, assumed to be a result of frequency-dependent parasite-driven selection and mate choice. This is believed to be so it promotes heterozygosity improving the chances of survival for the offspring and avoid inbreeding.
In experiments using rats, MHC-associated mate choice indicated that odor cues played a role. In humans, it has generally been shown that a correlation men and women will rate the opposite genders odour as more pleasant, if the human has MHC-dissimilar antigens to them. However, women on contraceptive pills rate the odour of MHC-similar men as being more pleasant, it is unknown why women on contraceptive pills rate smell in this way. It was found that when processing MHC-similar smells were processed faster.
Human facial preferences have been shown to correlate with both MHC-similarity and MHC-heterozygosity. Research into MHC-similarity with regards to facial attractiveness is limited but research so far suggests that women when thinking of long-term relationships will choose males who are MHC-similar. While facial asymmetry hasn't been correlated with MHC-heterozygosity, the perceived healthiness of skin appears to be. It appears to be that only MHC-heterozygosity and no other genetic markers are correlated with facial attractiveness in males and it has been shown that so far that there is no correlation that has been found in females.
- Edward, D. A (2014). "The description of mate choice". Behavioral Ecology. 26: 301–310. doi:10.1093/beheco/aru142.
- Petrie, Marion; Halliday, Tim; Sanders, Carolyn (1991). "Peahens prefer peacocks with elaborate trains". Animal Behaviour. 42: 323–331. doi:10.1016/s0003-3472(05)80484-1.
- Andersson, Malte (1994). Sexual Selection. Princeton University Press.
- Moller, A.; Jennions, M. (2001). "How important are direct benefits of sexual selection?". Naturwissenschaften. 88 (10): 401–415. PMID 11729807. doi:10.1007/s001140100255.
- Kokko, H.; Brooks, R.; Jennions, M.; Morley, J. (2003). "The evolution of mate choice and mating biases". Proceedings of the Royal Society B. 270 (1515): 653–664. PMC . PMID 12769467. doi:10.1098/rspb.2002.2235.
- Dawkins, Marian; Guilford, Tim (Nov 1996). "Sensory Bias and the Adaptiveness of Female Choice". The American Naturalist. 148 (5): 937–942. JSTOR 2463414. doi:10.1086/285964.
- Halliday, T. R (1983). "The study of mate choice". Mate Choice. 1.
- Darwin, C. 1871. The Descent of Man, and Selection in Relation to Sex. John Murray, London.
- Miller, Geoffrey (2000). The mating mind: how sexual choice shaped the evolution of human nature, London, Heineman, ISBN 0-434-00741-2 (also Doubleday, ISBN 0-385-49516-1) p.8
- Fisher, R. A. (1915). "The evolution of sexual preference". Eugenic Review. 7 (3): 184–192. PMC . PMID 21259607.
- Bateman, A. (1948). "Intra-sexual selection in Drosophila". Heredity. 2 (Pt. 3): 349–368. PMID 18103134. doi:10.1038/hdy.1948.21.
- Williams, G.C. 1966. Adaptation and Natural Selection. Princeton University Press, Princeton, N.J.
- Williams, G. C. 1975. Sex and evolution. Princeton University Press, Princeton, N.J.
- Trivers, R.L. (1972). Parental investment and sexual selection. In B. Campbell (Ed.), Sexual selection and the descent of man, 1871-1971 (pp. 136–179). Chicago, IL: Aldine
- Andersson, M. 1994. Sexual Selection, Princeton University Press
- Price, T.; Schluter, D.; Heckman, N. (1993). "Sexual selection when the female directly benefits". Biological Journal of the Linnean Society. 48 (3): 187–211. doi:10.1111/j.1095-8312.1993.tb00887.x.
- Moller, A. P. 1994. Sexual selection and the barn swallow. Oxford University Press, Oxford
- Hill, G (28 March 1991). "Plumage coloration is a sexually selected indicator of male quality". Nature. 350 (6316): 337–339. doi:10.1038/350337a0. Retrieved 16 November 2014.
- Linville, S. U.; Breitwisch, R.; Schilling, A. J. (1998). "Plumage brightness as an indicator of parental care in northern cardinals". Animal Behaviour. 55: 119–127. doi:10.1006/anbe.1997.0595.
- Iyengar, Vikram K.; Rossini, Carmen; Eisner, Thomas (2001). "Precopulatory Assessment of Male Quality in an Arctiid Moth ( Utetheisa Ornatrix ): Hydroxydanaidal Is the Only Criterion of Choice". Behavioral Ecology and Sociobiology. 49 (4): 283–88. doi:10.1007/s002650000292.
- Boughman, J. W. (2002). "How sensory drive can promote speciation". Trends in Ecology and Evolution. 17 (12): 571–577. doi:10.1016/S0169-5347(02)02595-8.
- Rodd, F. H.; Hughes, K. A.; Grether, G. F.; Baril, C. T. (2002). "A possible non-sexual origin of mate preference: are male guppies mimicking fruit?". Proceedings of the Royal Society B. 269 (1490): 475–481. PMC . PMID 11886639. doi:10.1098/rspb.2001.1891.
- Proctor, Heather C. (1991-10-01). "Courtship in the water mite Neumania papillator: males capitalize on female adaptations for predation". Animal Behaviour. 42 (4): 589–598. doi:10.1016/S0003-3472(05)80242-8.
- Proctor, Heather C. (1992-10-01). "Sensory exploitation and the evolution of male mating behaviour: a cladistic test using water mites (Acari: Parasitengona)". Animal Behaviour. 44 (4): 745–752. doi:10.1016/S0003-3472(05)80300-8.
- Proctor, H. C. (1992-01-01). "Effect of Food Deprivation on Mate Searching and Spermatophore Production in Male Water Mites (Acari: Unionicolidae)". Functional Ecology. 6 (6): 661–665. JSTOR 2389961. doi:10.2307/2389961.
- Alcock, John. Animal Behaviour: A Evolutionary Approach (10th ed.). Sinauer. pp. 70–72. ISBN 9780878939664.
- Jones, I. L.; Hunter, F. M. (1998). "Heterospecific mating preferences for a feather ornament in least auklets". Behavioral Ecology. 9 (2): 187–192. doi:10.1093/beheco/9.2.187.
- McClinktock, W. J.; Uetz, G. W. (1996). "Female choice and pre-existing bias: Visual cues during courtship in two Schizocosawolff spiders". Animal Behaviour. 52: 167–181. doi:10.1006/anbe.1996.0162.
- Prum, R. O. (1996). "Phylogenetic tests of alternative intersexual selection mechanisms: Trait macroevolution in a polygynous clade". The American Naturalist. 149 (4): 688–692. JSTOR 2463543. doi:10.1086/286014.
- Fuller, R. C.; Houle, D.; Travis, J. (2005). "Sensory bias as an explanation for the evolution of mate preferences". American Naturalist. 166 (4): 437–446. PMID 16224700. doi:10.1086/444443.
- Biegler, R.; Kennair, L. E. O. (2016). "Sisterly love: Within-generation differences in ideal partner for sister and self". Evolutionary Behavioural Sciences. 10 (1): 29–42. doi:10.1037/ebs0000060.
- Zahavi, A. (1975). "Mate selection – a selection for the handicap". Journal of Theoretical Biology. 53 (1): 205–214. PMID 1195756. doi:10.1016/0022-5193(75)90111-3.
- Hamilton W. D., Zuk M. (1982). "Heritable true fitness and bright birds: A role for parasites?". Science. 218 (4570): 384–387. PMID 7123238. doi:10.1126/science.7123238.
- Zuk, M.; Thornhill, R.; Lignon, J. D.; Johnson, Kristine (1990). "Parasites and mate choice in red jungle fowl". American Zoology. 30 (2): 235–244. doi:10.1093/icb/30.2.235.
- Hill, G. E. (1991). "Plumage coloration is a sexually selected indicator of male quality". Nature. 350 (6316): 337–339. doi:10.1038/350337a0.
- Puurtinen, M.; Ketola, T.; Kotiaho, J. S. (2005). "Genetic compatibility and sexual selection". Trends in Ecology and Evolution. 20 (4): 157–158. PMID 16701361. doi:10.1016/j.tree.2005.02.005.
- Wedekind, C.; Seebeck, T.; Bettens, F.; Paeke, A. J. (1995). "MHC-dependent mate preferences in humans". Proceedings: Biological Sciences. 260 (1359): 245–249. PMID 7630893. doi:10.1098/rspb.1995.0087.
- Brennan, Peter A.; Zufall, Frank (2006). "Pheromonal Communication in Vertebrates". Nature. 444 (7117): 308–15. PMID 17108955. doi:10.1038/nature05404.
- Eizaguirre, C., et al. "MHC-Based Mate Choice Combines Good Genes and Maintenance of MHC Polymorphism." Molecular ecology 18.15 (2009): 3316-29.
- Reusch, Thorsten B. H.; et al. (2001). "Female Sticklebacks Count Alleles in a Strategy of Sexual Selection Explaining MHC Polymorphism". Nature. 414 (6861): 300–2.
- Kalbe, M; Eizaguirre, C; Dankert, I; et al. (2009). "Lifetime reproductive success is maximized with optimal MHC diversity". Proceedings of the Royal Society B, Biological Sciences. 276: 925–934. doi:10.1098/rspb.2008.1466.
- Nowak, MA; Tarczyhornoch, K; Austyn, J.M. (1992). "The optimal number of major histocompatibility complex-molecules in an individual". Proceedings of the National Academy of Sciences USA. 89: 10896–10899. doi:10.1073/pnas.89.22.10896.
- Woelfing, B; Traulsen, A; Milinski, M; Boehm, T (2009). "Does intra-individual MHC diversity keep a golden mean?". Philosophical Transactions of the Royal Society B. 364: 117–128. doi:10.1098/rstb.2008.0174.
- Ilmonen, P.; Penn, D.J.; Damjanovich, K.; Morrison, L.; Ghotbi, L.; Potts, W.K. (2007). "Major histocompatibility complex heterozygosity reduces fitness in experimentally infected mice". Genetics. 176: 2501–2508. doi:10.1534/genetics.107.074815.
- Schad, J.; et al. (2011). "MHC Class II DRB Diversity, Selection Pattern and Population Structure in a Neotropical Bat Species, Noctilio Albiventris". Heredity. 107 (2): 115–26. doi:10.1038/hdy.2010.173.
- Wegner, M.; Kalbe, M.; Rauch, G.; Kurtz, J.; Schaschl, H.; Reusch, T. B. H. (2006). "Genetic variation in MHC Class II expression and interactions with MHC sequence polymorphism in three-spined sticklebacks". Molecular Ecology. 15: 1153–1164. doi:10.1111/j.1365-294x.2006.02855.x.
- Schaschl, H.; et al. (2006). "Selection and Recombination Drive the Evolution of MHC Class II DRB Diversity in Ungulates". Heredity. 97 (6): 427–37. doi:10.1038/sj.hdy.6800892.
- Andersson, M. 1994. Sexual selection. Princeton University Press, Princeton, NJ
- Vincent, A.; Anhesjo, I.; Berglund, A.; Rosenqvist, G. (1992). "Pipefish and seahorses: Are they all sex role reversed?". Trends in Ecology and Evolution. 7 (7): 237–241. PMID 21236017. doi:10.1016/0169-5347(92)90052-D.
- Wells, K. (1978). "Courtship and parental behavior in a Panamanian poison-arrow frog (Dendrobates auratus)". Herpetologica. 34 (2): 148–155. JSTOR 3891667.
- Emlen, S. T.; Wrege, P. H. (2005). "Sex dimorphism, intrasexual competition and sexual selection in wattled jacana, a sex role reversed shore bird in Panama". The Auk. 121 (2): 391–403. JSTOR 4090403. doi:10.1642/0004-8038(2004)121[0391:sdicas]2.0.co;2.
- Eens, M.; Pinxten, R. (2000). "Sex-role reversal in vertebrates: behavioral and endocrinological accounts". Behavioral processes. 51 (1–3): 135–147. PMID 11074317. doi:10.1016/S0376-6357(00)00124-8.
- Glickman, S. E.; Frank, L. G.; Davidson, J. M.; Smith, E. R.; Siiteri, P. K. (1987). "Androstenedione may organize or activate sex-reversed traits in female spotted hyenas". PNAS. 84 (10): 3444–3447. PMC . PMID 3472215. doi:10.1073/pnas.84.10.3444.
- Frank, L. G. (1986). "Social organization of the spotted hyena II: Dominance and reproduction". Animal Behaviour. 35 (5): 1510–1527. doi:10.1016/S0003-3472(86)80221-4.
- Mayr, E. 1942. Systematics and Origin of Species, Belknap Press
- Ritchie, Michael G. (2007). "Sexual Selection and Speciation". Annual Review of Ecology, Evolution, and Systematics. 38: 79–102. doi:10.1146/annurev.ecolsys.38.091206.095733.
- Endler, J. A.; Houde, A. E. (1995). "Geographic variation in female preferences for male traits in Poecelia reticulata". Evolution. 49 (3): 456–468. doi:10.2307/2410270.
- Colbeck, G.J.; Sillett, T.S.; Webster, M.S. (2010). "Asymmetric discrimination of geographical variation in song in a migratory passerine". Animal Behaviour. 80: 311–318. doi:10.1016/j.anbehav.2010.05.013.
- Barrett, Louise; Dunbar, Robin; Lycett, John (2002). Human Evolutionary Psychology. Hampshire: Palgrave. ISBN 0-333-72558-1.
- Collins, Sarah A (2000). "Men's voices and women's choices" (PDF). Animal Behaviour. 60: 773–780. doi:10.1006/anbe.2000.1523.
- Barber, Nigel (1995). "The evolutionary psychology of physical attractiveness: sexual selection and human morphology". Ethology and Sociobiology. 16: 395–424. doi:10.1016/0162-3095(95)00068-2 – via Elsevier Science Direct.
- Buss, David (2016). Evolutionary Psychology, The New Science of Mind. New York: Routledge. pp. 103–104. ISBN 978-0-205-99212-6.
- Edward, Dominic; Chapman, Tracey (2011). "The evolution and significance of male mate choice". Trends in Ecology and Evolution. 26: 647–654. doi:10.1016/j.tree.2011.07.012.
- Buss, David (2016). Evolutionary Psychology, The New Science of Mind. New York: Routledge. pp. 163–176. ISBN 978-0-205-99212-6.
- Buss, David (2016). Evolutionary Psychology, The New Science of Mind. New York: Routledge. pp. 133–162. ISBN 978-0-205-99212-6.
- Singh, Devendra; Bronstad, P Matthew (1997). "Sex differences in the anatomical locations of human body scarification and tattooing as a function of pathogen prevalence". Evolution and Human Behavior. 18: 403–416. doi:10.1016/s1090-5138(97)00089-5.
- Cartwright, John (c2000). Evolution and human behavior: Darwinian perspectives on human nature. Basingstoke: Macmillan. pp. 146–147. ISBN 9780333714577. Check date values in:
- Ludvico, L.R.; Kurland, J.A. (1995). "Symbolic or not-so symbolic wounds: The behavioral ecology of human scarification". Ethology and Sociobiology. 16: 155–172. doi:10.1016/0162-3095(94)00075-i.
- DeBruine, Lisa M.; Jones, Benedict C.; Crawford, John R.; Welling, Lisa L. M.; Little, Anthony C. (2010). "The health of a nation predicts their mate preferences: cross-cultural variation in women's preferences for masculinized male faces". Proceedings of the Royal Society B. 277: 2405–2410. PMC . PMID 20236978. doi:10.1098/rspb.2009.2184.
- Jones, Benedict C.; Feinberg, David R.; Watkins, Christopher D.; Fincher, Corey L.; Little, Anthony C.; DeBruine, Lisa M. (2012). "Pathogen disgust predicts women's preferences for masculinity in men's voices, faces, and bodies". Behavioral Ecology. 24: 373–379. doi:10.1093/beheco/ars173.
- Thornhill, R; Gangestad, S. W.; Scheib, J. E. (1999). "Facial attractiveness, symmetry and cues of good genes.". Proceedings of the Royal Society B. 266: 1913–1917.
- DeBruine, Lisa M.; Little, Anthony C.; Jones, Benedict C. (2012). "Extending parasite-stress theory to variation in human mate preferences". Behavioural and Brain Sciences. 35: 86–87. doi:10.1017/s0140525x11000987.
- White, D. R.; Burton, M. L. (1988). "Causes of polygyny: Ecology, economy, kinship, and warfare". American Anthropologist. 90: 871–887. doi:10.1525/aa.1988.90.4.02a00060.
- Low, Bobbi S. (1990). "Marriage Systems and Pathogen Stress in Human Societies". American Zoologist. 30: 325–339. doi:10.1093/icb/30.2.325.
- Gangestad, Steven W.; Buss, David M. (1993). "Pathogen prevalence and human mate preferences". Ethology and Sociobiology. 14: 89–96. doi:10.1016/0162-3095(93)90009-7.
- Yamazaki, K.; Yamaguchi, M.; Baranoski, L.; Bard, J.; Boyse, E. A.; Thomas, L. (1979). "Recognition among mice. Evidence from the use of a Y-maze differentially scented by congenic mice of different major histocompatibility types". Journal of Experimental Medicine. 150: 755–760. doi:10.1084/jem.150.4.755.
- Wedekind, C.; Fu¨ri, S. (1997). "Body odour preferences in men and women: do they aim for specific MHC combinations or simply heterozygosity?". Proceedings of the Royal Society B. 264: 1471–1479. PMC . PMID 9364787. doi:10.1098/rspb.1997.0204.
- Pause, B. M.; Krauel, K.; Schraders, C.; Sojka, B.; Westphal, E.; Muller-Ruchholtz, W.; Ferstl, R. (2005). "The human brain is a detector of chemosensorily transmitted HLA-class I-similarity in same- and opposite-sex relations". Proceedings of the Royal Society B. 273: 471–478. doi:10.1098/rspb.2005.3342.
- Havlicek, Jan; Roberts, S. Craig (2009). "MHC-correlated mate choice in humans: A review". Psychoneuroendocrinology. 34: 497–512. PMID 19054623. doi:10.1016/j.psyneuen.2008.10.007.
- Roberts, S. C.; Little, A. C.; Gosling, L. M.; Jones, B. C.; Perrett, D. I.; Carter, V.; Petrie, M (2005). "MHC-assortative facial preferences in humans". Biology Letters. 1: 400–403. doi:10.1098/rsbl.2005.0343.
- Roberts, S. C.; Little, A. C.; Gosling, L. M.; Perrett, D. I.; Carter, V.; Jones, B. C.; Penton-Voak, I. S.; Petrie, M. (2005). "MHC-heterozygosity and human facial attractiveness". Evolutionary Human Behaviour. 26: 213–226. doi:10.1016/j.evolhumbehav.2004.09.002.
- Lie, H.; Simmons, L.; Rhodes, G. (2008). "Genetic diversity revealed in human faces". Evolution. 62: 2473–2486. doi:10.1111/j.1558-5646.2008.00478.x.
- Thornhill, R.; Gangestad, S. W.; Miller, R.; Scheyd, G.; McCollough, J. K.; Franklin, M. (2003). "Major histocompatibility complex genes, symmetry, and body scent attractiveness in men and women". Behavioural Ecology. 14: 668–678. doi:10.1093/beheco/arg043.
- Coetzee, V.; Barrett, L.; Greeff, J. M.; Henzi, S. P.; Perrett, D. I.; Wadee, A. A. (2007). "Common HLA alleles associated with health, but not with facial attractiveness". PLOS One. 2: e640. doi:10.1371/journal.pone.0000640.