David R. Feinberg

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). Womens 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 mens voices to feminized mens 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 mens 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.

Facial appearance is a cue to oestrogen levels in women

Although many accounts of facial attractiveness propose that femininity in womens faces indicates high levels of oestrogen, there is little empirical evidence in support of this assumption. Here, we used assays for urinary metabolites of oestrogen (oestrone-3-glucuronide, E1G) and progesterone (pregnanediol-3-glucuronide, P3G) to investigate the relationship between circulating gonadal hormones and ratings of the femininity, attractiveness and apparent health of womens faces. Positive correlations were observed between late follicular oestrogen and ratings of femininity, attractiveness and health. Positive correlations of luteal progesterone and health and attractiveness ratings were marginally significant. Ratings of facial attributions did not relate to hormone levels for women wearing make-up when photographed. There was no effect of sex of rater on the relationships between oestrogen and ratings of facial appearance. These findings demonstrate that female facial appearance holds detectable cues to reproductive health that are considered attractive by other people.

Correlated preferences for facial masculinity and ideal or actual partner's masculinity

Studies of womens preferences for male faces have variously reported preferences for masculine faces, preferences for feminine faces and no effect of masculinity–femininity on male facial attractiveness. It has been suggested that these apparently inconsistent findings are, at least partly, due to differences in the methods used to manipulate the masculinity of face images or individual differences in attraction to facial cues associated with youth. Here, however, we show that womens preferences for masculinity manipulated in male faces using techniques similar to the three most widely used methods are positively inter-related. We also show that womens preferences for masculine male faces are positively related to ratings of the masculinity of their actual partner and their ideal partner. Correlations with partner masculinity were independent of real and ideal partner age, which were not associated with facial masculinity preference. Collectively, these findings suggest that variability among studies in their findings for womens masculinity preferences reflects individual differences in attraction to masculinity rather than differences in the methods used to manufacture stimuli, and are important for the interpretation of previous and future studies of facial masculinity.

Commitment to relationships and preferences for femininity and apparent health in faces are strongest on days of the menstrual cycle when progesterone level is high.

Previous studies of changes in womens behavior during the menstrual cycle have offered insight into the motivations underpinning womens preferences for social cues associated with possible direct benefits (e.g., investment, low risk of infection) and indirect benefits (e.g., offspring viability). Here we sought to extend this work by testing for systematic variation in womens preferences for male and female faces and in their attitudes to their romantic relationship during the menstrual cycle. In Study 1, we found partnered womens reported commitment to their romantic relationship and preferences for femininity in male and female faces were strongest on days of the menstrual cycle when progesterone levels are increased (and fertility is low). Happiness in relationships did not change across the cycle. In Study 2, we found that the effect of cycle phase on womens preference for feminine faces was independent of increased attraction to apparent health in faces during the luteal phase. Collectively, these findings are further evidence that womens preferences for social cues associated with possible direct benefits and commitment to relationships are strongest during conditions characterized by raised progesterone level, while attraction to men displaying cues associated with possible indirect benefits is strongest when women are most fertile.

Menstrual cycle, pregnancy and oral contraceptive use alter attraction to apparent health in faces

Previous studies demonstrating changes in womens face preferences have emphasized increased attraction to cues to possible indirect benefits (e.g. heritable immunity to infection) that coincides with periods of high fertility (e.g. the late follicular phase of the menstrual cycle). By contrast, here we show that when choosing between composite faces with raised or lowered apparent health, womens preferences for faces that are perceived as healthy are (i) stronger during the luteal phase of the menstrual cycle than during the late follicular, fertile phase, (ii) stronger in pregnant women than in non–pregnant women and (iii) stronger in women using oral contraceptives than in women with natural menstrual cycles. Change in preference for male faces was greater for short– than long–term relationships. These findings indicate raised progesterone level is associated with increased attraction to facial cues associated with possible direct benefits (e.g. low risk of infection) and suggest that womens face preferences are influenced by adaptations that compensate for weakened immune system responses during pregnancy and reduce the risk of infection disrupting foetal development.

Effects of Menstrual Cycle Phase on Face Preferences

While many studies of face preferences have emphasized high agreement among individuals about the types of faces they consider attractive and unattractive, other studies have demonstrated systematic variation in face preferences. Here, we review the evidence that women’s preferences for masculinity, apparent health, and self-resemblance in faces change systematically during the menstrual cycle. Our review focuses on the proximate mechanisms that might underpin these changes (i.e., what changes in hormone levels are important for effects of menstrual cycle phase) and the possible functions of these changes (i.e., to maximize the likelihood that offspring inherit strong immune systems or to increase the likelihood of successful pregnancy by either promoting affiliation with individuals who will provide support and care during pregnancy or by promoting strategies to avoid contagion during social interactions). While evidence that differentiates between these two accounts of the function of cyclic shifts in face preferences is currently equivocal for masculinity preferences, there is compelling evidence that the function of the effects of menstrual cycle phase on preferences for apparent health and self-resemblance in faces is to increase the likelihood of successful pregnancy.

Voice pitch predicts reproductive success in male hunter-gatherers

The validity of evolutionary explanations of vocal sexual dimorphism hinges upon whether or not individuals with more sexually dimorphic voices have higher reproductive success than individuals with less dimorphic voices. However, due to modern birth control methods, these data are rarely described, and mating success is often used as a second-rate proxy. Here, we test whether voice pitch predicts reproductive success, number of children born and child mortality in an evolutionarily relevant population of hunter-gatherers. While we find that voice pitch is not related to reproductive outcomes in women, we find that men with low voice pitch have higher reproductive success and more children born to them. However, voice pitch in men does not predict child mortality. These findings suggest that the association between voice pitch and reproductive success in men is mediated by differential access to fecund women. Furthermore, they show that there is currently selection pressure for low-pitch voices in men.

Symmetry Is Related to Sexual Dimorphism in Faces: Data Across Culture and Species

Background Many animals both display and assess multiple signals. Two prominently studied traits are symmetry and sexual dimorphism, which, for many animals, are proposed cues to heritable fitness benefits. These traits are associated with other potential benefits, such as fertility. In humans, the face has been extensively studied in terms of attractiveness. Faces have the potential to be advertisements of mate quality and both symmetry and sexual dimorphism have been linked to the attractiveness of human face shape. Methodology/Principal Findings Here we show that measurements of symmetry and sexual dimorphism from faces are related in humans, both in Europeans and African hunter-gatherers, and in a non-human primate. Using human judges, symmetry measurements were also related to perceived sexual dimorphism. In all samples, symmetric males had more masculine facial proportions and symmetric females had more feminine facial proportions. Conclusions/Significance Our findings support the claim that sexual dimorphism and symmetry in faces are signals advertising quality by providing evidence that there must be a biological mechanism linking the two traits during development. Such data also suggests that the signalling properties of faces are universal across human populations and are potentially phylogenetically old in primates.

Facial cues of dominance modulate the short-term gaze-cuing effect in human observers

Responding appropriately to gaze cues is essential for fluent social interaction, playing a crucial role in social learning, collaboration, threat assessment and understanding others’ intentions. Previous research has shown that responses to gaze cues can be studied by investigating the gaze-cuing effect (i.e. the tendency for observers to respond more quickly to targets in locations that were cued by others’ gaze than to uncued targets). A recent study demonstrating that macaques demonstrate larger gaze-cuing effects when viewing dominant conspecifics than when viewing subordinate conspecifics suggests that cues of dominance modulate the gaze-cuing effect in at least one primate species. Here, we show a similar effect of facial cues associated with dominance on gaze cuing in human observers: at short viewing times, observers demonstrated a greater cuing effect for gaze cues from masculinized (i.e. dominant) faces than from feminized (i.e. subordinate) faces. Moreover, this effect of facial masculinity on gaze cuing decreased as viewing time was increased, suggesting that the effect is driven by involuntary responses. Our findings suggest that the mechanisms that underpin reflexive gaze cuing evolved to be sensitive to facial cues of others’ dominance, potentially because such differential gaze cuing promoted desirable outcomes from encounters with dominant individuals.

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

Many studies have shown that womens judgments of mens 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 mens judgments of womens attractiveness. To investigate this issue, we compared mens attractiveness judgments of feminized and masculinized womens and mens 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 womens 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 mens reported attraction to femininity in mens faces did not differ between high and low testosterone test sessions, suggesting that the effect of testosterone that we observed for judgments of womens faces was not due to a general response bias. Collectively, these findings suggest that changes in testosterone levels contribute to the strength of mens reported attraction to femininity in womens faces and complement previous findings showing that testosterone modulates mens interest in sexual stimuli.