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.     

Life History of Female Preferences for Male Faces

Although scientific interest in facial attractiveness has developed substantially in recent years, few studies have contributed to our understanding of the ontogeny of facial preferences. In this study, attractiveness of 30 male faces was evaluated by four female groups: girls at puberty, nonpregnant and pregnant young women, and middle-aged women. The main findings are as follows: (1) Preference for sexy-looking faces was strongest in young, nonpregnant women. (2) Biologically more mature girls displayed more adultlike preferences. (3) The intragroup consistency for postmenopausal women was relatively low. (4) In terms of the preference pattern, pregnant women were more similar to perimenopausal women than they were to their nonpregnant peers. (5) Preference for youthful appearance decreased with the age of the women. I argue that the life history of female preferences for male faces is, to a large extent, hormone-driven and underpinned by a set of evolutionary adaptations.     

Facial appearance is a cue to oestrogen levels in women

Although many accounts of facial attractiveness propose that femininity in women's 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 women's 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.     

Facial movement varies by sex and is related to attractiveness

Facial movement has received little attention in studies of human attractiveness, yet dynamic displays are an important aspect of courtship in many species. This experiment investigated whether facial movement could be used to identify sex, and whether the ease of identification was associated with attractiveness. We removed shape cues to sex by applying movement from individual faces to a standardised facial model. Participants were able to distinguish between male and female animations of this model at levels above chance. Furthermore, there was a positive association between ease of sex identification and attractiveness for female, but not male, faces. Analysis of facial movement suggested several behaviours that are more frequent in women than men (blinking, tilting, nodding, shaking, and amount of movement). Although some of these behaviours may be cues to sex identification, none alone was directly linked to attractiveness. Our findings suggest that feminine motion is attractive in female faces, but sexually recognisable movement has no clear influence on male attractiveness, in agreement with work on static faces employing composite images.     

African and European perception of African female attractiveness

Majority of research on attractiveness is restricted to faces of European origin. The perception of attractiveness may, however, vary across communities due to variations in both facial morphology and local standards of beauty. We investigated the relative contribution of four facial markers of attractiveness based on 101 female facial portraits (standardized, non-manipulated) from Cameroon and Namibia, which were assessed by local male raters and by raters from a distant European population, the Czech Republic. Images from Cameroon include only women of Bantu origin, while Namibians are represented by women of both Bantu (Owambo/Herero) and Nama origin. While controlling for age and BMI, we explored the relationship between female attractiveness and a set of facial traits: fluctuating asymmetry, averageness, shape sexual dimorphism, and skin color (rated and measured in CIELab color space).In the Cameroonian sample, local male raters favored lighter-skinned female faces with morphology closer to average. The attractiveness of Nama women as rated by Nama men positively correlated with lighter complexion, but this did not extend to rating by Cameroonian men. The attractiveness of Namibian Owambo/Herero women was positively associated with facial femininity and lighter complexion when judged by both Cameroonian and Nama male raters. In all samples, the attractiveness as rated by Czech men was predicted by age and BMI, but not by skin color. We found no significant association between attractiveness and fluctuating asymmetry in any of the tested samples. When controlling for age, the effect of skin color on attractiveness turned to be non-significant in the Owambo/Herrero and Nama sample, but remained significant in the Cameroonian sample. Variations in skin color thus represent an important factor of African female attractiveness within the African context, but they do not seem to affect judgements made by European raters. Sensitivity to some facial markers of female attractiveness thus seems to be restricted to regional populations and/or constrained by shared ethnicity.     

Fertility in the cycle predicts women's interest in sexual opportunism

Research over the past decade has documented clear, robust changes in women's sexual preferences and interests across the ovarian cycle. When fertile, women are particularly attracted to a number of masculine male features (e.g., masculine faces, voices, scents and bodies) and other traits, and especially when they evaluate men's “sexiness” rather than their attractiveness as long-term partners. The current research extended this line of research by examining changes in women's self-reported sexual interests across the cycle. We asked 68 normally ovulating women in committed romantic relationships to fill out questionnaires about their sexual preferences and interests (at that time, not in general) twice across their cycles: once when fertile and once during the luteal phase. Relative to during the luteal phase, fertile women expressed (a) greater emphasis on the physical attractiveness of a partner; (b) greater arousal at the sight or thought of attractive male bodily features; (c) greater willingness to engage in and interest in sex with attractive men, even ones who they do not know well (interest in sexual opportunism). These findings importantly extend our understanding of women's fertile-phase sexuality.     

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

Previous studies demonstrating changes in women's 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, women's 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 women's face preferences are influenced by adaptations that compensate for weakened immune system responses during pregnancy and reduce the risk of infection disrupting foetal development.     

Second to fourth digit ratio, testosterone and perceived male dominance

Previous studies have shown that male faces with extreme features associated with testosterone are perceived as dominant and masculine. Women have been reported to prefer more masculinized male faces as they may consider testosterone markers to be an 'honest' indication of good health, and such considerations may underlie their aesthetic preferences. However, pronounced testosterone facial markers are also associated with dominance, and several negative personality traits. This suggests that female aesthetic preferences may be an adaptive compromise between positive attributes associated with higher than average testosterone, and negative attributes associated with more extreme masculinization. This current study attempts to clarify the role of hormone markers in female perceptions of dominance, masculinity and attractiveness, in male facial images. Recent evidence suggests that the relative length of the 2nd to 4th finger (2D : 4D ratio) is a pointer to prenatal testosterone levels and may thus serve as a window to the prenatal hormonal environment. We measured 2D : 4D in a sample of male college students and took salivary samples to analyse circulating levels of testosterone. Women rated facial images of these males for dominance, masculinity and attractiveness. Our results show that male 2D : 4D was significantly negatively related to perceived dominance and masculinity but not attractiveness. Circulating testosterone levels were not related to dominance, masculinity or attractiveness. These findings suggest that: (i) high prenatal levels of testosterone serve to 'organize' male facial features to subsequently reflect dominance and masculine characteristics presumably activated during puberty; and (ii) attractiveness is not directly related to testosterone levels. We conclude that facial dominance and masculinity reflect a male's perceived status rather than his physical attraction to women.     

Women's attractiveness judgments of self-resembling faces change across the menstrual cycle

Two lines of reasoning predict that women's preferences for people exhibiting cues to kinship will be lower in the follicular phase than in the luteal phase of the menstrual cycle. Women may avoid kinship cues during the follicular phase when they are most fertile due to the costs of inbreeding. Alternatively, women may seek kinship cues during the luteal phase as a byproduct of the benefits of associating with kin during pregnancy, which is also characterized by high progesterone. We find that preferences for facial resemblance, a putative kinship cue, follow this predicted pattern and are positively correlated with estimated progesterone levels based on cycle day. Neither estimated estrogen levels nor conception risk predicted preferences for self-resemblance, and the cyclic shift was stronger for preferences for female faces than male faces. These findings lead to the possibility that this cyclic change in preference for self-resemblance may be a byproduct of a hormonal mechanism for increasing affiliative behavior toward kin during pregnancy rather than a mechanism for preventing inbreeding during fertile periods.     

Facial attractiveness, symmetry and cues of good genes.

Cues of phenotypic condition should be among those used by women in their choice of mates. One marker of better phenotypic condition is thought to be symmetrical bilateral body and facial features. However, it is not clear whether women use symmetry as the primary cue in assessing the phenotypic quality of potential mates or whether symmetry is correlated with other facial markers affecting physical attractiveness. Using photographs of men's faces, for which facial symmetry had been measured, we found a relationship between women's attractiveness ratings of these faces and symmetry, but the subjects could not rate facial symmetry accurately. Moreover, the relationship between facial attractiveness and symmetry was still observed, even when symmetry cues were removed by presenting only the left or right half of faces. These results suggest that attractive features other than symmetry can be used to assess phenotypic condition. We identified one such cue, facial masculinity (cheek-bone prominence and a relatively longer lower face), which was related to both symmetry and full- and half-face attractiveness.     

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.     

Men's facial masculinity predicts changes in their female partners' sexual interests across the ovulatory cycle,whereas men's intelligence does not

A substantial body of work demonstrates that women's mate preferences change across the ovulatory cycle. When fertile in their cycles, women are especially attracted to masculine features (e.g., faces, voices, bodies), socially dominant behavior, and male scents associated with body symmetry and social dominance. Women may also find intelligent men particularly attractive when fertile, though findings are mixed. Related research shows that, on average, romantically-involved women report stronger sexual attraction to men other than their pair-bond partners, but not partners, when fertile, and especially when their partners lack features fertile women prefer (e.g., symmetry). In the current study, we examined whether women's patterns of sexual interests across the cycle are similarly moderated by partners' facial masculinity, facial attractiveness, and intelligence. Results revealed predicted effects of male partners’ facial masculinity but none for partners’ intelligence. Facial attractiveness may have effects, but we find no evidence that it does so independently of facial masculinity.     

Symmetry, sexual dimorphism in facial proportions and male facial attractiveness

Facial symmetry has been proposed as a marker of developmental stability that may be important in human mate choice. Several studies have demonstrated positive relationships between facial symmetry and attractiveness. It was recently proposed that symmetry is not a primary cue to facial attractiveness, as symmetrical faces remain attractive even when presented as half faces (with no cues to symmetry). Facial sexual dimorphisms ('masculinity') have been suggested as a possible cue that may covary with symmetry in men following data on trait size/symmetry relationships in other species. Here, we use real and computer graphic male faces in order to demonstrate that (i) symmetric faces are more attractive, but not reliably more masculine than less symmetric faces and (ii) that symmetric faces possess characteristics that are attractive independent of symmetry, but that these characteristics remain at present undefined.     

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.     

Female Preferences for Male Vocal and Facial Masculinity in Videos

Vocal and facial masculinity are cues to underlying testosterone in men and influence women’s mate preferences. Consistent with the proposal that facial and vocal masculinity signal common information about men, prior work has revealed correlated female preferences for male facial and vocal masculinity. Previous studies have assessed women’s preferences for male facial and vocal masculinity by presenting faces and voices independently and using static face stimuli. By contrast, here we presented women with short video clips in which male faces and voices were simultaneously manipulated in masculinity. We found that women who preferred masculine faces also preferred masculine voices. Furthermore, women whose faces were rated as relatively more attractive preferred both facial and vocal masculinity more than did women whose faces were rated as less attractive. These findings complement other evidence for cross‐modal masculinity preferences among women and demonstrate that preferences observed in studies using still images and/or independently presented vocal stimuli are also observed when dynamic faces and voices are displayed simultaneously in video format.     

Face perception is modulated by sexual preference

Face perception is mediated by a distributed neural system in the human brain . The response to faces is modulated by cognitive factors such as attention, visual imagery, and emotion ; however, the effects of gender and sexual orientation are currently unknown. We used fMRI to test whether subjects would respond more to their sexually preferred faces and predicted such modulation in the reward circuitry. Forty heterosexual and homosexual men and women viewed photographs of male and female faces and assessed facial attractiveness. Regardless of their gender and sexual orientation, all subjects similarly rated the attractiveness of both male and female faces. Within multiple, bilateral face-selective regions in the visual cortex, limbic system, and prefrontal cortex, similar patterns of activation were found in all subjects in response to both male and female faces. Consistent with our hypothesis, we found a significant interaction between stimulus gender and the sexual preference of the subject in the thalamus and medial orbitofrontal cortex, where heterosexual men and homosexual women responded more to female faces and heterosexual women and homosexual men responded more to male faces. Our findings suggest that sexual preference modulates face-evoked activation in the reward circuitry.     

Facial correlates of sociosexuality

Previous studies have documented variation in sexual behaviour between individuals leading to the notion of ‘restricted’ individuals (i.e., people who prefer long-term relationships) and ‘unrestricted’ individuals (i.e., people who are open to short-term relationships). This distinction is often referred to as sociosexual orientation. Observers have been previously found to distinguish sociosexuality from video footage of individuals, although the specific cues used have not been identified. Here we assessed the ability of observers to judge sexual strategy based specifically on cues in both facial composites and real faces. We also assessed how observers' perceptions of the masculinity/femininity and attractiveness of faces relate to the sociosexual orientation of the pictured individuals. Observers were generally able to identify restricted vs. unrestricted individuals from cues in both composites and real faces. Unrestricted sociosexuality was generally associated with greater attractiveness in female composites and real female faces and greater masculinity in male composites. Although male observers did not generally associate sociosexuality with male attractiveness, female observers generally preferred more restricted males' faces (i.e., those with relatively strong preferences for long-term relationships). Collectively, our results support previous findings that androgenisation in men is related to less restricted sexual behaviour and suggest that women are averse to unrestricted men.     

Are human female preferences for symmetrical male faces enhanced when conception is likely?

Human females show a preference for the scent of symmetrical male bodies and appearance of masculine male faces only when conception is likely. These traits are thought to be signs of male quality. We examined whether females show an enhanced visual preference for another putative sign of mate quality, facial symmetry, when conception is likely. Twenty-nine females not taking oral contraceptives (nonpill users) rated the attractiveness of male faces varying in symmetry level (low, normal, high and perfect) for a short-term sexual partner at two phases of the menstrual cycle (low and high conception risk). They also rated the attractiveness of male faces for a long-term sexual partner and female faces for general attractiveness. We also tested a control group of 27 pill users. An overall preference for symmetry was found in all participants. However, the hypothesized enhanced cyclic preference for symmetrical male faces in nonpill users was not found. Nor was there an effect of relationship context. These results may be more consistent with a direct benefit or sensory bias model for preference evolution than with an indirect genetic benefit model. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

The Relative Contributions of Facial Shape and Surface Information to Perceptions of Attractiveness and Dominance

Although many studies have investigated the facial characteristics that influence perceptions of others’ attractiveness and dominance, the majority of these studies have focused on either the effects of shape information or surface information alone. Consequently, the relative contributions of facial shape and surface characteristics to attractiveness and dominance perceptions are unclear. To address this issue, we investigated the relationships between ratings of original versions of faces and ratings of versions in which either surface information had been standardized (i.e., shape-only versions) or shape information had been standardized (i.e., surface-only versions). For attractiveness and dominance judgments of both male and female faces, ratings of shape-only and surface-only versions independently predicted ratings of the original versions of faces. The correlations between ratings of original and shape-only versions and between ratings of original and surface-only versions differed only in two instances. For male attractiveness, ratings of original versions were more strongly related to ratings of surface-only than shape-only versions, suggesting that surface information is particularly important for men’s facial attractiveness. The opposite was true for female physical dominance, suggesting that shape information is particularly important for women’s facial physical dominance. In summary, our results indicate that both facial shape and surface information contribute to judgments of others’ attractiveness and dominance, suggesting that it may be important to consider both sources of information in research on these topics.     

The masculinity paradox: facial masculinity and beardedness interact to determine women's ratings of men's facial attractiveness

In many species, male secondary sexual traits have evolved via female choice as they confer indirect (i.e. genetic) benefits or direct benefits such as enhanced fertility or survival. In humans, the role of men's characteristically masculine androgen‐dependent facial traits in determining men's attractiveness has presented an enduring paradox in studies of human mate preferences. Male‐typical facial features such as a pronounced brow ridge and a more robust jawline may signal underlying health, whereas beards may signal men's age and masculine social dominance. However, masculine faces are judged as more attractive for short‐term relationships over less masculine faces, whereas beards are judged as more attractive than clean‐shaven faces for long‐term relationships. Why such divergent effects occur between preferences for two sexually dimorphic traits remains unresolved. In this study, we used computer graphic manipulation to morph male faces varying in facial hair from clean‐shaven, light stubble, heavy stubble and full beards to appear more (+25% and +50%) or less (?25% and ?50%) masculine. Women (N = 8520) were assigned to treatments wherein they rated these stimuli for physical attractiveness in general, for a short‐term liaison or a long‐term relationship. Results showed a significant interaction between beardedness and masculinity on attractiveness ratings. Masculinized and, to an even greater extent, feminized faces were less attractive than unmanipulated faces when all were clean‐shaven, and stubble and beards dampened the polarizing effects of extreme masculinity and femininity. Relationship context also had effects on ratings, with facial hair enhancing long‐term, and not short‐term, attractiveness. Effects of facial masculinization appear to have been due to small differences in the relative attractiveness of each masculinity level under the three treatment conditions and not to any change in the order of their attractiveness. Our findings suggest that beardedness may be attractive when judging long‐term relationships as a signal of intrasexual formidability and the potential to provide direct benefits to females. More generally, our results hint at a divergence of signalling function, which may result in a subtle trade‐off in women's preferences, for two highly sexually dimorphic androgen‐dependent facial traits.