Mate choice trade-offs and women’s preference for physically attractive men

Researchers studying human sexuality have repeatedly concluded that men place more emphasis on the physical attractiveness of potential mates than women do, particularly in long-term sexual relationships. Evolutionary theorists have suggested that this is the case because male mate value (the total value of the characteristics that an individual possesses in terms of the potential contribution to his or her mate’s reproductive success) is better predicted by social status and economic resources, whereas women’s mate value hinges on signals conveyed by their physical appearance. This pattern may imply that women trade off attractiveness for resources in mate choice. Here I test whether a trade-off between resources and attractiveness seems to be occurring in the mate choice decisions of women in the United States. In addition, the possibility that the risk of mate desertion drives women to choose less attractive men as long-term mates is tested. The results were that women rated physically attractive men as more likely to cheat or desert a long-term relationship, whereas men did not consider attractive women to be more likely to cheat. However, women showed no aversion to the idea of forming long-term relationships with attractive men. Evidence for a trade-off between resources and attractiveness was found for women, who traded off attractiveness, but not other traits, for resources. The potential meaning of these findings, as well as how they relate to broader issues in the study of sex differences in the evolution of human mate choice for physical traits, is discussed.     

Gender Differences in Cooperation: Experimental Evidence on High School Students

The emergence of cooperation among unrelated human subjects is a long-standing conundrum that has been amply studied both theoretically and experimentally. Within the question, a less explored issue relates to the gender dependence of cooperation, which can be traced back to Darwin, who stated that "women are less selfish but men are more competitive". Indeed, gender has been shown to be relevant in several game theoretical paradigms of social cooperativeness, including prisoner''s dilemma, snowdrift and ultimatum/dictator games, but there is no consensus as to which gender is more cooperative. We here contribute to this literature by analyzing the role of gender in a repeated Prisoners'' Dilemma played by Spanish high-school students in both a square lattice and a heterogeneous network. While the experiment was conducted to shed light on the influence of networks on the emergence of cooperation, we benefit from the availability of a large dataset of more 1200 participants. We applied different standard econometric techniques to this dataset, including Ordinary Least Squares and Linear Probability models including random effects. All our analyses indicate that being male is negatively associated with the level of cooperation, this association being statistically significant at standard levels. We also obtain a gender difference in the level of cooperation when we control for the unobserved heterogeneity of individuals, which indicates that the gender gap in cooperation favoring female students is present after netting out this effect from other socio-demographics factors not controlled for in the experiment, and from gender differences in risk, social and competitive preferences.     

Physical attractiveness and cooperation in a prisoner's dilemma game

The modulating role of age on the relationship between physical attractiveness and cooperativeness in a prisoner's dilemma game (PDG) was investigated. Previous studies have shown that physical attractiveness is negatively related to cooperative choices among young men but not young women. Following the argument that the negative relationship between physical attractiveness and cooperation is a product of short-term mating strategies among attractive men, we predicted that this relationship is unique to young men and absent among women and older men. We tested this hypothesis with 175 participants (aged 22–69 years). The results showed that physical attractiveness was negatively related to cooperative behavior among young men but not among women or older men. We further observed that the negative relationship between physical attractiveness and cooperation among young men was particularly strong when attractiveness was judged by women.     

Cues to fertility: perceived attractiveness and facial shape predict reproductive success

Attractive facial features in women are assumed to signal fertility, but whether facial attractiveness predicts reproductive success in women is still a matter of debate. We investigated the association between facial attractiveness at young adulthood and reproductive life history—number of children and pregnancies—in women of a rural community. For the analysis of reproductive success, we divided the sample into women who used contraceptives and women who did not. Introducing two-dimensional geometric morphometric methodology, we analysed which specific characteristics in facial shape drive the assessment of attractiveness and covary with lifetime reproductive success. A set of 93 (semi)landmarks was digitized as two-dimensional coordinates in postmenopausal faces. We calculated the degree of fluctuating asymmetry and regressed facial shape on facial attractiveness at youth and reproductive success. Among women who never used hormonal contraceptives, we found attractive women to have more biological offspring than less attractive women. These findings are not affected by sociodemographic variables. Postmenopausal faces corresponding to high reproductive success show more feminine features—facial characteristics previously assumed to be honest cues to fertility. Our findings support the notion that facial attractiveness at the age of mate choice predicts reproductive success and that facial attractiveness is based on facial characteristics, which seem to remain stable until postmenopausal age.     

Facial attractiveness and fertility in populations with low levels of modern birth control

Evolutionary models of human mate choice generally assume that physical attractiveness has evolved through sexual selection, i.e., it has been associated with higher mating opportunities and subsequent reproductive success across our evolutionary history. Here we investigate whether facial attractiveness is related to fertility in order to understand the extent to which selection can operate on attractive traits in modern populations. We used data from two populations where the prevalence of modern birth control methods is low and thus unlikely to disconnect mating opportunities from reproductive success: men and women from contemporary rural Senegal and men from the West Point Military Academy in the USA who graduated in 1950. We found that facial attractiveness negatively predicts age-specific reproduction in both sexes in Senegal and is independent from lifetime reproductive success in men from the USA. Overall, the results suggest that facial attractiveness is not under positive selection and raise questions about methodological approaches currently used to assess attractiveness.     

Sex ratio influences the motivational salience of facial attractiveness

The sex ratio of the local population influences mating-related behaviours in many species. Recent experiments show that male-biased sex ratios increase the amount of financial resources men will invest in potential mates, suggesting that sex ratios influence allocation of mating effort in humans. To investigate this issue further, we tested for effects of cues to the sex ratio of the local population on the motivational salience of attractiveness in own-sex and opposite-sex faces. We did this using an effort-based key-press task, in which the motivational salience of facial attractiveness was assessed in samples of faces in which the ratio of male to female images was manipulated. The motivational salience of attractive opposite-sex, but not own-sex, faces was greater in the own-sex-biased (high competition for mates) than in the opposite-sex-biased (low competition for mates) condition. Moreover, this effect was not modulated by participant sex. These results present new evidence that sex ratio influences human mating-related behaviours. They also present the first evidence that the perceived sex ratio of the local population may modulate allocation of mating effort in women, as well as men.     

Physical attractiveness and reproductive success in humans: evidence from the late 20th century United States

Physical attractiveness has been associated with mating behavior, but its role in reproductive success of contemporary humans has received surprisingly little attention. In the Wisconsin Longitudinal Study (1244 women, 997 men born between 1937 and 1940), we examined whether attractiveness assessed from photographs taken at age 18 years predicted the number of biological children at age 53–56 years. In women, attractiveness predicted higher reproductive success in a nonlinear fashion, so that attractive (second highest quartile) women had 16% and very attractive (highest quartile) women 6% more children than their less attractive counterparts. In men, there was a threshold effect so that men in the lowest attractiveness quartile had 13% fewer children than others who did not differ from each other in the average number of children. These associations were partly but not completely accounted for by attractive participants' increased marriage probability. A linear regression analysis indicated relatively weak directional selection gradient for attractiveness (β=0.06 in women, β=0.07 in men). These findings indicate that physical attractiveness may be associated with reproductive success in humans living in industrialized settings.     

Cooperation and conflict between women in the family

Here I review recent research on reproductive conflict between females in families and how it influences their reproductive behaviour. Kin selection can favor cooperation between parent and offspring, siblings, or unrelated co‐residents who share interests in other family members such as grand‐offspring. However, these are also the individuals most likely to be sharing resources, and so conflict can also emerge. While substantial interest has arisen in evolutionary anthropology, especially over the last two decades, in the possibility of cooperative breeding in humans, less attention has been paid to reproductive conflict among female kin. Communal breeding in animals is generally understood as emerging from competition over the resources needed to breed. Competition for household resources is a problem that also faces human families. Models suggest that in some circumstances, inclusive fitness can be maximized by sharing reproduction rather than harming relatives by fighting with them, even if the shares that emerge are not equal. Thus, competition and cooperation turn out to be strongly related to each other. Reproductive competition within and between families may have underpinned the biological evolution of fertility patterns (such as menopause) and the cultural evolution of marriage, residence, and inheritance norms (such as late male marriage or primogeniture), which can enhance cooperation and minimize the observed incidence of such conflicts.     

Female competition and its evolutionary consequences in mammals

Following Darwin's original insights regarding sexual selection, studies of intrasexual competition have mainly focused on male competition for mates; by contrast, female reproductive competition has received less attention. Here, we review evidence that female mammals compete for both resources and mates in order to secure reproductive benefits. We describe how females compete for resources such as food, nest sites, and protection by means of dominance relationships, territoriality and inter‐group aggression, and by inhibiting the reproduction of other females. We also describe evidence that female mammals compete for mates and consider the ultimate causes of such behaviour, including competition for access to resources provided by mates, sperm limitation and prevention of future resource competition. Our review reveals female competition to be a potentially widespread and significant evolutionary selection pressure among mammals, particularly competition for resources among social species for which most evidence is currently available. We report that female competition is associated with many diverse adaptations, from overtly aggressive behaviour, weaponry, and conspicuous sexual signals to subtle and often complex social behaviour involving olfactory signalling, alliance formation, altruism and spite, and even cases where individuals appear to inhibit their own reproduction. Overall, despite some obvious parallels with male phenotypic traits favoured under sexual selection, it appears that fundamental differences in the reproductive strategies of the sexes (ultimately related to parental investment) commonly lead to contrasting competitive goals and adaptations. Because female adaptations for intrasexual competition are often less conspicuous than those of males, they are generally more challenging to study. In particular, since females often employ competitive strategies that directly influence not only the number but also the quality (survival and reproductive success) of their own offspring, as well as the relative reproductive success of others, a multigenerational view ideally is required to quantify the full extent of variation in female fitness resulting from intrasexual competition. Nonetheless, current evidence indicates that the reproductive success of female mammals can also be highly variable over shorter time scales, with significant reproductive skew related to competitive ability. Whether we choose to describe the outcome of female reproductive competition (competition for mates, for mates controlling resources, or for resources per se) as sexual selection depends on how sexual selection is defined. Considering sexual selection strictly as resulting from differential mating or fertilisation success, the role of female competition for the sperm of preferred (or competitively successful) males appears particularly worthy of more detailed investigation. Broader definitions of sexual selection have recently been proposed to encompass the impact on reproduction of competition for resources other than mates. Although the merits of such definitions are a matter of ongoing debate, our review highlights that understanding the evolutionary causes and consequences of female reproductive competition indeed requires a broader perspective than has traditionally been assumed. We conclude that future research in this field offers much exciting potential to address new and fundamentally important questions relating to social and mating‐system evolution.     

Beautiful parents have more daughters: a further implication of the generalized Trivers-Willard hypothesis (gTWH)

The generalized Trivers-Willard hypothesis (gTWH) [Kanazawa, S., 2005. Big and tall parents have more sons: further generalizations of the Trivers-Willard hypothesis. J. Theor. Biol. 235, 583-590) proposes that parents who possess any heritable trait which increases the male reproductive success at a greater rate than female reproductive success in a given environment will have a higher-than-expected offspring sex ratio, and parents who possess any heritable trait which increases the female reproductive success at a greater rate than male reproductive success in a given environment will have a lower-than-expected offspring sex ratio. One heritable trait which increases the reproductive success of daughters much more than that of sons is physical attractiveness. I therefore predict that physically attractive parents have a lower-than-expected offspring sex ratio (more daughters). Further, if beautiful parents have more daughters and physical attractiveness is heritable, then, over evolutionary history, women should gradually become more attractive than men. The analysis of the National Longitudinal Study of Adolescent Health (Add Health) confirm both of these hypotheses. Very attractive individuals are 26% less likely to have a son, and women are significantly more physically attractive than men in the representative American sample.     

Women's attractiveness is linked to expected age at menopause

A great number of studies have shown that features linked to immediate fertility explain a large part of the variance in female attractiveness. This is consistent with an evolutionary perspective, as men are expected to prefer females at the age at which fertility peaks (at least for short‐term relationships) in order to increase their reproductive success. However, for long‐term relationships, a high residual reproductive value (the expected future reproductive output, linked to age at menopause) becomes relevant as well. In that case, young age and late menopause are expected to be preferred by men. However, the extent to which facial features provide cues to the likely age at menopause has never been investigated so far. Here, we show that expected age at menopause is linked to facial attractiveness of young women. As age at menopause is heritable, we used the mother's age at menopause as a proxy for her daughter's expected age of menopause. We found that men judged faces of women with a later expected age at menopause as more attractive than those of women with an earlier expected age at menopause. This result holds when age, cues of immediate fertility and facial ageing were controlled for. Additionally, we found that the expected age at menopause was not correlated with any of the other variables considered (including immediate fertility cues and facial ageing). Our results show the existence of a new correlate of women's facial attractiveness, expected age at menopause, which is independent of immediate fertility cues and facial ageing.     

Differences in Time Use for Mating and Nepotistic Effort as a Function of Male Attractiveness in Rural Belize

This paper explores whether physical attractiveness was a determinant of reproductive strategy in a sample of men living in rural Belize. A theoretical argument is presented to explain why differences in male physical attractiveness should lead to differences in strategy as evidenced by time-use, and why these differences should be especially apparent in nonindustrialized societies. Retrospective data were collected on men’s time use during their last day off from work. The results were that more facially attractive men spent more time in mating effort and less time in nepotistic effort than less facially attractive men. Another component of physical attractiveness, fluctuating asymmetry, was not successful in predicting differences in time use. The results suggest that facially attractive men spend their leisure time seeking sexual access rather than spending it with kin, because their potential fitness returns are higher for this activity, whereas less attractive men receive higher returns to time spent with kin. This could be due directly to fitness returns to nepotism received by less attractive men, or because family involvement displays potential parental investment skills that are attractive to women. This may help build a reputation for reliability; in other words, time spent in nepotistic effort could be an alternative mating tactic that appeals to women’s desire for a responsible paternally investing mate.     

The threat premium in economic bargaining

Costly punishment is thought to have evolved because it promotes cooperation and the equitable sharing of resources, but the costs associated with punishment – for both the punisher and the punished – limit the efficiency of this enforcement system in economic interactions. Reputation may also guide decision-making, but this information is not always available (e.g., in interactions involving strangers). Across several bargaining studies, we provide evidence of an efficient and flexible “threat-based” bargaining system that can influence the division of resources without the need for costly punishment and reputational information. We found that participants, without prompting, dynamically adjusted bargaining based on the perceived threat-potential (resource holding power and aggressiveness) of the bargaining partner, giving larger offers to individuals who appeared more threatening. These effects of perceived threat-potential were strongest among participants who were most vulnerable to harm in physical contests (women vs men and weaker men vs stronger men), despite that offers were made on-line and anonymously to photographs of the individuals rather than in face-to-face interactions. These results may reflect an overgeneralization of a real-world threat heuristic that allows low threat individuals to extract resources when possible, while avoiding physical retaliation and harm, and high threat individuals to appropriate larger shares of a resource through static facial cues of threat rather than by physically expressing their propensity to punish. Previously, researchers have highlighted the monetary advantages of attractiveness (the “beauty premium”), but the effects of threat either trumped, devalued, or were equivalent to those of attractiveness.     

Exploring the effects of working for endowments on behaviour in standard economic games

In recent years, significant advances have been made in understanding the adaptive (ultimate) and mechanistic (proximate) explanations for the evolution and maintenance of cooperation. Studies of cooperative behaviour in humans invariably use economic games. These games have provided important insights into the mechanisms that maintain economic and social cooperation in our species. However, they usually rely on the division of monetary tokens which are given to participants by the investigator. The extent to which behaviour in such games may reflect behaviour in the real world of biological markets--where money must be earned and behavioural strategies incur real costs and benefits--is unclear. To provide new data on the potential scale of this problem, we investigated whether people behaved differently in two standard economic games (public goods game and dictator game) when they had to earn their monetary endowments through the completion of dull or physically demanding tasks, as compared with simply being given the endowment. The requirement for endowments to be 'earned' through labour did not affect behaviour in the dictator game. However, the requirement to complete a dull task reduced cooperation in the public goods game among the subset of participants who were not familiar with game theory. There has been some effort to test whether the conclusions drawn from standard, token-based cooperation games adequately reflect cooperative behaviour 'in the wild.' However, given the almost total reliance on such games to study cooperation, more exploration of this issue would be welcome. Our data are not unduly worrying, but they do suggest that further exploration is needed if we are to make general inferences about human behaviour from the results of structured economic games.     

Facial attractiveness is related to women's cortisol and body fat,but not with immune responsiveness

Recent studies suggest that facial attractiveness indicates immune responsiveness in men and that this relationship is moderated by stress hormones which interact with testosterone levels. However, studies testing whether facial attractiveness in women signals their immune responsiveness are lacking. Here, we photographed young Latvian women, vaccinated them against hepatitis B and measured the amount of specific antibodies produced, cortisol levels and percentage body fat. Latvian men rated the attractiveness of the women''s faces. Interestingly, in women, immune responsiveness (amount of antibodies produced) did not predict facial attractiveness. Instead, plasma cortisol level was negatively associated with attractiveness, indicating that stressed women look less attractive. Fat percentage was curvilinearly associated with facial attractiveness, indicating that being too thin or too fat reduces attractiveness. Our study suggests that in contrast to men, facial attractiveness in women does not indicate immune responsiveness against hepatitis B, but is associated with two other aspects of long-term health and fertility: circulating levels of the stress hormone cortisol and percentage body fat.     

MHC-assortative facial preferences in humans

Individuals tend to choose mates who are sufficiently genetically dissimilar to avoid inbreeding. As facial attractiveness is a key factor in human mate preference, we investigated whether facial preferences were related to genetic dissimilarity. We asked female volunteers to rate the attractiveness of men from photographs and compared these results with individual genotypes at the major histocompatibility complex (MHC). In contrast to previously reported preferences based on odour, we found a non-significant tendency for women to rate MHC-similar faces as more attractive, suggesting a preference for cues to a self-similar MHC in faces. Further analysis revealed that male faces received higher attractiveness scores when rated by women who were MHC-similar than by MHC-dissimilar women. Although unexpected, this MHC-similar facial preference is consistent with other studies documenting assortative preferences in humans, including for facial phenotype.     

Physical attractiveness influences reproductive success of modern men

Theory suggests that reproductive success is positively associated with an individual’s genetic quality. However, the association between physical attractiveness and reproductive success (i.e., number of offspring) in modern humans remains less clear. Here we examined associations between men’s reproductive success and physical attractiveness from retrospective data obtained from married, divorced, and single samples of Slovakian men. As predicted, facially more attractive and taller men were more likely to engage in marriage. In turn, married men had higher reproductive success than single men. Even when men’s marital status was considered, facially more attractive men had higher reproductive success than their less attractive counterparts. This supports the importance of physical attractiveness in sexual selection in modern humans.     

Intrasexual competition and eating restriction in heterosexual and homosexual individuals

Restrictive eating attitudes and behaviors have been hypothesized to be related to processes of intrasexual competition. According to this perspective, within-sex competition for status serves the adaptive purpose of attracting mates. As such, status competition salience may lead to concerns of mating desirability. For heterosexual women and gay men, such concerns revolve around appearing youthful and, thus, thinner. Following this logic, we examined how exposure to high-status and competitive (but not thin or highly attractive) same-sex individuals would influence body image and eating attitudes in heterosexual and in gay/lesbian individuals. Results indicated that for heterosexuals, intrasexual competition cues led to greater body image dissatisfaction and more restrictive eating attitudes for women, but not for men. In contrast, for homosexual individuals, intrasexual competition cues led to worse body image and eating attitudes for gay men, but not for lesbian women. These findings support the idea that the ultimate explanation for eating disorders is related to intrasexual competition.     

Major histocompatibility complex genes, symmetry, and body scent attractiveness in men and women

Previous research indicates that the scent of developmentalstability (low fluctuating asymmetry, FA) is attractive to womenwho are fertile (at high-conception risk points in their menstrualcycles), but not to other women or men. Prior research alsoindicates that the scent of dissimilarity in major histocompatibilitycomplex (MHC) genes may play a role in human mate choice. Westudied the scent attractiveness to the opposite sex of t-shirtsworn for 2 nights' sleep. Our results indicate that the twoolfactory systems are independent. We repeated previous resultsfrom studies of the scent of symmetry. We repeated previousresults from MHC research in part; men, but not women, showeda preference for t-shirts with the scent of MHC dissimilarity.Women's scent ratings of t-shirts were uncorrelated with thewearer's MHC dissimilarity and allele frequency, but positivelycorrelated with the wearer's MHC heterozygosity. Fertile womendid not exhibit any MHC trait preferences. Women's preferencefor the scent of men who were heterozygous for MHC alleles maybe stronger in women who are at infertile cycle points. Menpreferred the scent of common MHC alleles, which may functionto avoid mates with rare alleles that exhibit gestational drive.Men also preferred the scent of women at fertile cycle points.The scent of facially attractive women, but not men, was preferred.Neither FA nor facial attractiveness in either sex correlatedwith MHC dissimilarity to others, MHC heterozygosity, or MHCallelic rarity.     

The role of facial hair in women's perceptions of men's attractiveness,health, masculinity and parenting abilities

Facial hair strongly influences people's judgments of men's socio-sexual attributes. However, the nature of these judgments is often contradictory. The levels of intermediate facial hair growth presented to raters and the stage of female raters' menstrual cycles might have influenced past findings. We quantified men's and women's judgments of attractiveness, health, masculinity and parenting abilities for photographs of men who were clean-shaven, lightly or heavily stubbled and fully bearded. We also tested the effect of the menstrual cycle and hormonal contraceptive use on women's ratings. Women judged faces with heavy stubble as most attractive and heavy beards, light stubble and clean-shaven faces as similarly less attractive. In contrast, men rated full beards and heavy stubble as most attractive, followed closely by clean-shaven and light stubble as least attractive. Men and women rated full beards highest for parenting ability and healthiness. Masculinity ratings increased linearly as facial hair increased, and this effect was more pronounced in women in the fertile phase of the menstrual cycle, although attractiveness ratings did not differ according to fertility. Our findings confirm that beardedness affects judgments of male socio-sexual attributes and suggest that an intermediate level of beardedness is most attractive while full-bearded men may be perceived as better fathers who could protect and invest in offspring.