Women's estradiol predicts preference for facial cues of men's testosterone

A growing body of research has shown that women express stronger attraction to more masculine traits when they are tested near ovulation than when tested during other times in the menstrual cycle. Although these effects have been interpreted as increased preferences for markers of elevated testosterone during times in the cycle when conception is most likely, no previous studies have directly demonstrated that women express stronger attraction to higher testosterone men at different times in the cycle. In addition, little research has addressed which hormonal or other physiological mechanisms may regulate temporal shifts in women's attractiveness judgments. In this research, we demonstrate that women with higher estradiol concentrations exhibit stronger preferences for the faces of men with higher testosterone concentrations, and that women's testosterone preference and estradiol curves track one another across days of the cycle. The findings are the first direct demonstration in humans that hormone concentrations in one sex are associated with attraction to cues of hormonal status in the opposite sex. The results support a functional role for estradiol in calibrating women's mating psychology to indices of their current fertility, analogous to similar processes that have been documented in nonhuman species. A strong correlation between estradiol and testosterone preference specifically during the luteal phase further suggests that women's mate preferences may track their fertility between different cycles in addition to being calibrated to the timing of ovulation within individual cycles.     

Partner preferences in the context of concurrency: What Himba want in formal and informal partners

Research on human mate preferences that distinguishes between short- and long-term partners has been conducted mainly in industrialized societies, where multiple mating and concurrent partnerships are stigmatized. However, cross-culturally, there is significant variation in the frequency and the level of acceptance of such relationships. Furthermore, the dichotomy between short- and long-term partnerships does not fully describe the diversity in actual extra-pair behavior, which ranges from single sexual encounters to multi-year love affairs. Here we present another comparison, between formal (marital) and informal (non-marital) partners, which we feel better captures this diversity. We assess the traits that men and women prefer in each type of partner among Himba pastoralists, where concurrent partnerships are common and accepted for both sexes. We situate our findings in relation to three main explanations for concurrent partnerships: dual-mating, trading-up and multiple investors. We find some similarities with the existing literature in the traits that are listed as important, including physical attractiveness, wealth and intelligence. Our evidence suggest that Himba men follow a dual strategy, preferring hard-working wives and attractive girlfriends. Women's preferences align most strongly with a multiple investors explanation, most clearly articulated through their preferences for wealthy husbands and generous boyfriends. Limited support is also found for a dual-mating strategy in women. These findings suggest that local cultural norms and ecologies modulate mate preferences in important ways.     

What makes a voice masculine: Physiological and acoustical correlates of women's ratings of men's vocal masculinity

Men's voices contain acoustic cues to body size and hormonal status, which have been found to affect women's ratings of speaker size, masculinity and attractiveness. However, the extent to which these voice parameters mediate the relationship between speakers' fitness-related features and listener's judgments of their masculinity has not yet been investigated.     

Men report stronger attraction to femininity in women's faces when their testosterone levels are high

Many studies have shown that women's judgments of men's attractiveness are affected by changes in levels of sex hormones. However, no studies have tested for associations between changes in levels of sex hormones and men's judgments of women's attractiveness. To investigate this issue, we compared men's attractiveness judgments of feminized and masculinized women's and men's faces in test sessions where salivary testosterone was high and test sessions where salivary testosterone was relatively low. Men reported stronger attraction to femininity in women's faces in test sessions where salivary testosterone was high than in test sessions where salivary testosterone was low. This effect was found to be specific to judgments of opposite-sex faces. The strength of men's reported attraction to femininity in men's faces did not differ between high and low testosterone test sessions, suggesting that the effect of testosterone that we observed for judgments of women's faces was not due to a general response bias. Collectively, these findings suggest that changes in testosterone levels contribute to the strength of men's reported attraction to femininity in women's faces and complement previous findings showing that testosterone modulates men's interest in sexual stimuli.     

Effects of learning on song preferences and Zenk expression in female songbirds

Male songbirds learn to produce their songs, and females attend to these songs during mate choice. The evidence that female song preferences are learned early in life, however, is mixed. Here we review studies that have found effects of early song learning on adult song preferences, and those that have not. In at least some species, early experience with song can modify adult song preferences. Whether this learning needs to occur during an early sensitive phase, akin to male imitative vocal learning, or not remains an open question. Studies of the neural bases for female song preferences highlight activity (as measured by immediate-early gene induction) in regions of the auditory forebrain as often, but not always, being associated with song preferences. Immediate-early gene induction in these regions, however, is not specific to songs experienced early in life. On the whole, inherited factors, early experience, and adult experience all appear to play a role in shaping female songbirds preferences for male songs.     

Menstrual cycle, trait estrogen level, and masculinity preferences in the human voice

Men with low testosterone (feminine men) invest in relationships and offspring more than men with high testosterone (masculine men). Women's attraction to testosterone dependent traits (e.g. masculine face shape) is enhanced during the late-follicular, fertile phase of the menstrual cycle. Attractive, feminine women have stronger preferences for masculine men as possible long-term partners than less attractive, masculine women. We manipulated 2 testosterone related vocal traits (voice pitch and apparent vocal-tract length) in voices to test if women prefer masculinized men's voices to feminized men's voices; masculinity preferences are enhanced at the fertile (late-follicular) menstrual cycle phase; the amount that masculinity preferences shift cyclically relates to average estrone-3-glucuronide concentration (the primary urinary metabolite of estrone, E3G). We found women displayed general masculinity preferences for men's voices; masculinity preferences were greater in the fertile (late-follicular) phase of the cycle than the non-fertile (early-follicular and luteal) phase; and this effect was most pronounced for women with low average E3G concentration. As feminine women (i.e. those with high average E3G levels) are most able to obtain investment even from masculine men, these women may not need to change their mating preference or strategy during the menstrual cycle as much as masculine women.     

Mate preferences and choices for facial and body hair in heterosexual women and homosexual men: influence of sex,population, homogamy,and imprinting-like effect

Recent research has reported that male body and facial hair influence women's mate preferences. However, it is not clear whether such preferences are typical for women or for individuals who prefer males as sexual partners. Here we explored body and facial hair in preferred and actual partners among men and women who prefer men as sexual partners. Including homosexual individuals provides a unique opportunity to investigate whether evolved mating psychologies are specific to the sex of the individual or sex of the partner. Based on an online survey of 1577 participants from Brazil and the Czech Republic, we found that, on average, homosexual men preferred hairier stimuli than heterosexual women, supporting past findings that homosexual men have strong preferences for masculine traits. Preferences for facial and body hair appear to be influenced less by sex of the preferred partner than sex of the individual, pointing to a possible sex-specific mating psychology. Further, Brazilians preferred bigger beards than Czechs, which was positively associated with the self-reported amount of beardedness in Brazil, suggesting that familiarity effects underpin cross-cultural differences in preferences for facial hair. Moreover, homosexual men preferred a self-similar degree of beardedness, and Czech women preferred a similar degree of beardedness as their fathers had during their childhood. However, these effects were not associated with the level of facial hair in their actual partners; in general, mate preferences and actual mate choices for facial and body hair differed. Thus, individual differences in some self-reported characteristics, cultural factors, and aspects of personal experience may modulate differences in preferences for masculine traits.     

Hypothetical rankings of prospective husbands for female kin in lowland Nicaragua: consensus analysis indicates high agreement and associations with wealth and hunting skill

In preindustrial societies, kin may exert influence on the mating choices of women, but there have been few systematic studies of the preferred characteristics of husbands for female kin. In an indigenous Mayangna and Miskito community, photographs of 29 male household heads were presented to informants, who ranked the men on three characteristics: desirability as a spouse, hunting ability and wealth. For the desirability rankings, informants were asked to consider the advice that they would give to young female relatives and rank the men based on the qualities that such women should seek in a husband. Consensus analysis indicates that there is high agreement among informants on all three sets of rankings. There is no evidence that the age and sex of informants are associated with variation in evaluations of the desirability of men, which suggests that the evaluations by reproductively active women do not significantly differ from rankings by other informants. Multivariate analysis indicates that perceptions of both a man's wealth and his hunting ability are positively associated with his desirability as a prospective husband for female kin. By contrast, a strong kin-based social network, as measured by the presence of consanguineal kin in the community, seems unimportant to a man's desirability as a husband. Although it remains unclear to what extent hunting ability is a signal of phenotypic quality, these results support predictions that individuals will encourage female kin to marry men who are good resource providers. Finally, compared to a conventional reliability analysis, consensus analysis is demonstrated as a superior method for assessing both unidimensionality and subgroup variation in informant rankings.     

Parental preferences for the facial traits of their offspring's partners can enhance parental inclusive fitness

Physical appearance provides a wealth of information concerning an individual's biological fitness and reproductive quality, but we do not know whether parents make use of this information when evaluating potential partners for their offspring. This is critical to our understanding of human mate choice, because parents frequently influence their offspring's mating decisions, either directly, for instance through arranged marriages, or indirectly, through manipulating their offspring's partner choice. Here, we used facial images that varied in attractiveness, masculinity, health, and symmetry to assess both reproductively-aged daughters' and their parents' preferences in potential mates for the daughters. In line with our predictions, both daughters and their parents had clear preferences for markers of genetic quality, although the daughters showed significantly stronger preferences for these markers than their parents. Contrary to previous research, parents and daughters did not have stronger preferences for markers of genetic quality if they perceived the daughter to be more attractive. Parents' preferences for the facial markers of genetic quality in their offspring's partner may help maximize inclusive fitness.     

Sexual preferences during artificial menstrual cycles in social groups of rhesus monkeys (Macaca mulatta)

Eight groups of rhesus monkeys each consisting of one male and four ovariectomized females were observed while two of the females were treated with hormones to produce artificial menstrual cycles. These were either synchronized or offset by 7-day increments. Sexually preferred females, defined by the numbers of ejaculations per test, received almost twice as many ejaculations as did non-preferred females during all synchronized and offset cycles and during all cycle phases. However, short-term changes in partner preference occurred when the midcycle phase of non-preferred females coincided with the middle or late progesterone phase of preferred females, suggesting a negative effect of progesterone on behavior during the menstrual cycle. There were highly significant differences between preferred and non-preferred partners for almost all of their sexual and social interactions, and preferred partners showed longer proximity and grooming times as well as higher levels of sexual activity. Partner preferences accounted for more of the behavioral variance between pairs than did female dominance, although males sought the proximity of dominant females independently of their partner preferences. Thus, in a setting uncomplicated by male mate competition, sexual preference by male rhesus monkeys is a robust phenomenon depending on complex interactions between dominance, hormonal status, and the individual behavior of female partners.     

Men''s preferences for women''s profile waist-to-hip ratio in two societies

Women's waist-to-hip ratio (WHR) varies with age, and a lower WHR is associated with a higher estrogen-to-androgen ratio and possibly higher fecundity, at least in some populations. Consequently, it has been argued that selection has favored a universal male preference for a low female WHR. In previous studies using frontal pictures, men in the United States preferred a low WHR of 0.7, but men among Hadza hunter–gatherers and a few other small-scale societies preferred higher ratios. Unlike the actual WHR of women, measured with a tape around the waist and the hips and buttocks, the WHR in frontal pictures excludes the buttocks. Because frontal WHR gives only a partial picture, we used profile views of women to measure men's preferences for the profile WHR. Hadza men preferred a lower profile WHR (more protruding buttocks) than American men. Since Hadza men preferred higher frontal WHR but lower profile WHR, and since both contribute to the actual WHR, these results imply there is less disparity between American and Hadza preferences for the actual WHR of real women. We suggest men's preferences vary with the geographic variation in the shape of women who have wider hips in some populations and more protruding buttocks in others.     

Hormonal modulation of phonotaxis and advertisement-call preferences in the gray treefrog (Hyla versicolor)

Hormonal levels fluctuate during the breeding season in many anurans, but the identity of the hormones that modulate breeding behavior and their effects remain unclear. We tested the influence of a combined treatment of progesterone and prostaglandin on phonotaxis, the key proceptive reproductive behavior of female anurans. First, we found that female gray treefrogs (Hyla versicolor) treated with progesterone and prostaglandin exhibited phonotaxis to synthetic male advertisement signals significantly more often than animals treated with ringers vehicle or uninjected controls. Responsive females had greater levels of plasma progesterone and estradiol compared to both control groups, suggesting that these steroids may be promoting phonotaxis. Second, we found that the selectivity of hormonally-induced phonotaxis in H.versicolor was similar to that observed in freshly captured breeding animals. Females made the same choices between acoustic signals after hormone treatments in tests of frequency, call rate and pulse rate, compared to their responses without treatment immediately after collection from the breeding chorus. The preference for a longer call was, however, significantly weaker after hormonal induction of phonotaxis. Hormonally primed females were also less likely to respond in any test and took longer to respond than did freshly collected females. Consequently, our study shows how progesterone-prostaglandin induced phonotaxis in female treefrogs influences both the quality and quantity of phonotaxis, relative to that exhibited by naturally breeding females.     

Gonadal hormones provide the biological basis for sex differences in behavioral responses to cocaine

Both clinical and rodent studies show sexually dimorphic patterns in the behavioral response to cocaine in all phases of the addiction process (induction, maintenance, and relapse). Clinical and rodent studies also indicate that hormonal fluctuations during the menstrual/estrous cycle modulate cocaine-induced subjective effects in women and locomotor activity in female rats. Evidence suggests that gonadal hormones underlie these observed differences and could be the biological basis of sex-specific differences in cocaine addiction. To study the effects of gonadal hormones on cocaine-induced activity, two approaches have been used. First, studies have examined the role of endogenous hormones through gonadectomy (GDX) and side-by-side comparisons with intact rats. Second, the individual contributions of testosterone, progesterone, and estrogen have been determined by hormone replacement in GDX rats. In this review, we discuss gonadal hormones as the biological basis for the behavioral responses to cocaine, and the clinical implications of these findings.     

Men's preferences for women's breast size and shape in four cultures

The morphology of human female breasts appears to be unique among primates due to their permanent fat deposits. It has been previously suggested that female breast morphology arose as a result of sexual selection. This is supported by evidence showing that women with larger breasts tend to have higher estrogen levels; breast size may therefore serve as an indicator of potential fertility. However, breasts become less firm with age and parity, and breast shape could thus also serve as a marker of residual fertility. Therefore, cross-culturally, males are hypothesized to prefer breast morphology that indicates both high potential and residual fertility. To test this, we performed a survey on men's preferences for breast morphology in four different cultures (Brazil, Cameroon, the Czech Republic, Namibia). As stimuli, we used two sets of images varying in breast size (marker of potential fertility) and level of breast firmness (marker of residual fertility). Individual preferences for breast size were variable, but the majority of raters preferred medium sized, followed by large sized breasts. In contrast, we found systematic directional preferences for firm breasts across all four samples. This pattern supports the idea that breast morphology may serve as a residual fertility indicator, but offers more limited support for the potential fertility indicator hypothesis. Future studies should focus on a potential interaction between the two parameters, breast size and firmness, which, taken together, may help to explain the relatively large variation in women's breast sizes.     

Hormonally mediated epigenetic changes to steroid receptors in the developing brain: Implications for sexual differentiation

The establishment of sex-specific neural morphology, which underlies sex-specific behaviors, occurs during a perinatal sensitive window in which brief exposure to gonadal steroid hormones produces permanent masculinization of the brain. In the rodent, estradiol derived from testicular androgens is a principal organizational hormone. The mechanism by which transient estradiol exposure induces permanent differences in neuronal anatomy has been widely investigated, but remains elusive. Epigenetic changes, such as DNA methylation, allow environmental influences to alter long-term gene expression patterns and therefore may be a potential mediator of estradiol-induced organization of the neonatal brain. Here we review data that demonstrate sex and estradiol-induced differences in DNA methylation on the estrogen receptor α (ERα), estrogen receptor β (ERβ), and progesterone receptor (PR) promoters in sexually dimorphic brain regions across development. Contrary to the overarching view of DNA methylation as a permanent modification directly tied to gene expression, these data demonstrate that methylation patterns on steroid hormone receptors change across the life span and do not necessarily predict expression. Although further exploration into the mechanism and significance of estradiol-induced alterations in DNA methylation patterns in the neonatal brain is necessary, these results provide preliminary evidence that epigenetic alterations can occur in response to early hormone exposure and may mediate estradiol-induced organization of sex differences in the neonatal brain.     

Exposure to visual cues of pathogen contagion changes preferences for masculinity and symmetry in opposite-sex faces

Evolutionary approaches to human attractiveness have documented several traits that are proposed to be attractive across individuals and cultures, although both cross-individual and cross-cultural variations are also often found. Previous studies show that parasite prevalence and mortality/health are related to cultural variation in preferences for attractive traits. Visual experience of pathogen cues may mediate such variable preferences. Here we showed individuals slideshows of images with cues to low and high pathogen prevalence and measured their visual preferences for face traits. We found that both men and women moderated their preferences for facial masculinity and symmetry according to recent experience of visual cues to environmental pathogens. Change in preferences was seen mainly for opposite-sex faces, with women preferring more masculine and more symmetric male faces and men preferring more feminine and more symmetric female faces after exposure to pathogen cues than when not exposed to such cues. Cues to environmental pathogens had no significant effects on preferences for same-sex faces. These data complement studies of cross-cultural differences in preferences by suggesting a mechanism for variation in mate preferences. Similar visual experience could lead to within-cultural agreement and differing visual experience could lead to cross-cultural variation. Overall, our data demonstrate that preferences can be strategically flexible according to recent visual experience with pathogen cues. Given that cues to pathogens may signal an increase in contagion/mortality risk, it may be adaptive to shift visual preferences in favour of proposed good-gene markers in environments where such cues are more evident.     

Female choice for heterozygous mates changes along successive matings in a lizard

Female mate choice and female multiple mating are major focuses of studies on sexual selection. In a multiple mating context, the benefits of mate choice can change along successive matings, and female choice would be expected to change accordingly. We investigated sequential female mate choice in the moderately polyandrous common lizard (Zootoca vivipara, synonym Lacerta vivipara). Along successive mating opportunities, we found that females were relatively unselective for the first mate, but accepted males of higher heterozygosity for subsequent mating, consistent with the trade-up choice hypothesis. We discuss the evidence of trade-up mate choice in squamates and generally trade-up for mate heterozygosity in order to motivate new studies to fill gaps on these questions.     

What people prefer and what they think they prefer in short- and long-term partners. The effects of the phase of the menstrual cycle,hormonal contraception,pregnancy, and the marital and the parenthood status on partner preferences

The issue with most studies concerned with mate selection preferences in humans is that they rely on declarations and rational actions of experimental subjects, which are affected by their pre-conceived opinions and prejudices. Moreover, current research suggests that subcortical structures and processes, rather than the neocortex, play the principal role in actual partner choice behaviour. Consequently, we have only limited information on how relevant our current knowledge is in relation to real-life human ethology. To address these issues, we surveyed 2718 women and 4073 men between the ages of 16–50 and compared their declared and observed preferences for various properties in short-term and long-term partners. We found differences between what the subjects declared to prefer and what they preferred in reality: for example, men declared that wealth was the second least desirable property out of eleven in short-term partners, but we observed that in reality, they considered wealth the third most important factor after charisma and sense of humour. Similarly, while women declared that dominance and masculinity were desirable properties in short-term partners, in the observational part of the study, they showed little preference for these traits. Furthermore, we investigated the effects of the phase of the menstrual cycle, hormonal contraception, pregnancy, and partnership and parenthood status on these preferences. We found some support for the good parents hypothesis and no support for the good genes and the immunocompetence handicap hypotheses when observed, rather than declared preferences were considered. We also detected that hormonal contraception, and parenthood and partnership status influenced declared preferences in considerable ways, but had only a small, if any, impact on observed preferences. We suggest interpreting the results of studies reliant on declarations and rational actions of experimental subjects with great caution.     

Mechanisms of sexual selection: sexual swellings and estrogen concentrations as fertility indicators and cues for male consort decisions in wild baboons

Male mate-guarding episodes ('consortships'), are taxonomically widespread, yet costly to individual males. Consequently, males should bias consortships toward females with whom the probability of conception is high. We combined data on consortships with visual scoring of sexual swellings and assays of fecal estrogen concentrations (fE) in a wild population of baboons (Papio cynocephalus) to test the hypotheses that sexual swellings are reliable indicators of (1) within-cycle timing of ovulation, (2) differences in conception probability among females that differ in maturational stage, and (3) conceptive versus non-conceptive cycles of parous females. We also evaluated whether adult males might rely on swellings or other estrogen-dependent signals (e.g., fE) for mate-guarding decisions. We found that sexual swellings reflected conception probability within and among cycles. Adult males limited their consortships to the turgescent phase of cycles, and consorted more with adult females than with newly cycling adolescents. The highest ranking (alpha) males discriminated more than did males of other ranks; they (1) limited their consortships to the 5-day peri-ovulatory period, (2) consorted more with adult than with adolescent females, and (3) consorted more with adult females on conceptive cycles than on non-conceptive cycles, all to a greater extent than did males of other ranks. Male mate choice based on sexual swellings and other estrogenic cues of fertility may result in sexual selection on these female traits and enhance dominance-based reproductive skew in males. Alpha males are the least constrained in their mating behavior and can best take advantage of these cues to mate selectively.     

The ‘home advantage’ is necessary for a full winner effect and changes in post-encounter testosterone

Winning aggressive contests can both enhance future winning ability and change post-encounter hormones; however, it remains unclear if the context of a fight also influences such winner effects and hormone changes. We investigated this issue by using California mice (Peromyscus californicus) to test if the effect of residency status is necessary to improve future winning ability and alter post-encounter hormones. Male mice were subjected to an aggressive contest and their blood was collected 45 min after the fight. Upon contest initiation, focal mice had a ‘home advantage’ and three prior winning experiences, only one of these factors, or neither factor. Only individuals with a ‘home advantage’ and prior winning experience showed a full winner effect. Post-encounter changes in testosterone and progesterone resulted from an interaction between residency status and winning experience. These data indicate that in male California mice a ‘home advantage’ is necessary to produce the full winner effect and that residency status helps regulate inter-individual variation in the expression of post-encounter testosterone pulses and progesterone changes. Furthermore, these respective behavioral and physiological phenomena might be modulated in a context-specific manner, in part by the surrounding physical environment.