Male facial attractiveness and masculinity may provide sex‐ and culture‐independent cues to semen quality

Phenotype‐linked fertility hypothesis (PLFH) predicts that male secondary sexual traits reveal honest information about male fertilization ability. However, PLFH has rarely been studied in humans. The aim of the present study was to test PLFH in humans and to investigate whether potential ability to select fertile partners is independent of sex or cultural background. We found that on the contrary to the hypothesis, facial masculinity was negatively associated with semen quality. As increased levels of testosterone have been demonstrated to impair sperm production, this finding may indicate a trade‐off between investments in secondary sexual signalling (i.e. facial masculinity) and fertility or status‐dependent differences in investments in semen quality. In both sexes and nationalities (Spanish and Colombian), ranked male facial attractiveness predicted male semen quality. However, Spanish males and females estimated facial images generally more attractive (gave higher ranks) than Colombian raters, and in both nationalities, males gave higher ranks than females. This suggests that male facial cues may provide culture‐ and sex‐independent information about male fertility. However, our results also indicate that humans may be more sensitive to facial attractiveness cues within their own populations and also that males may generally overestimate the attractiveness of other men to females.     

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.     

Facial attractiveness signals different aspects of “quality” in women and men

We explored the relationships between facial attractiveness and several variables thought to be related to genotypic and phenotypic quality in humans (namely fluctuating asymmetry (FA), body mass index (BMI), health, age). To help resolve some controversy around previous studies, we used consistent measurement and statistical methods and relatively large samples of both female (n=94) and male (n=95) subjects (to be evaluated and measured), and female (n=226) and male (n=153) viewers (to rate attractiveness). We measured the asymmetry of 22 traits from three trait families (eight facial, nine body and five fingerprint traits) and constructed composite asymmetry indices of traits showing significant repeatability. Facial attractiveness was negatively related to an overall asymmetry index in both females and males, with almost identical slopes. Female facial attractiveness was best predicted by BMI and past health problems, whereas male facial attractiveness was best predicted by the socioeconomic status (SES) of their rearing environment. Composite FA indices accounted for a small (<4%) but usually significant percentage of the variation in facial attractiveness in both sexes, when factors related to asymmetry were controlled statistically. We conclude that, although facial attractiveness is negatively related to developmental instability (as measured by asymmetry), attractiveness also signals different aspects of “quality” in the two sexes, independent of FA.     

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.     

Does attractiveness in men provide clues to semen quality?

The psychological mechanisms underlying attractiveness judgements in humans are thought to be evolved adaptations for finding a high quality mate. The phenotype-linked fertility hypothesis proposes that females obtain reliable information on male fertility from male expression of sexual traits. A previous study of Spanish men reported that facial attractiveness was positively associated with semen quality. We aimed to determine whether this effect was widespread by examining a large sample of Australian men. We also extended our study to determine whether cues to semen quality are provided by components of attractiveness: masculinity, averageness and symmetry. Each male participant was photographed and provided a semen sample that was analyzed for sperm morphology, motility and concentration. Two independent sets of women rated the male photographs for attractiveness, and three further sets of 12 women rated the photographs for masculinity, symmetry or averageness. We found no significant correlations between semen quality parameters and attractiveness or attractive traits. Although male physical attractiveness may signal aspects of mate quality, our results suggest that phenotype-linked cues to male fertility may not be general across human populations.     

Multiple male traits interact: attractive bower decorations facilitate attractive behavioural displays in satin bowerbirds

Sexually selected male courtship displays often involve multiple behavioural and physical traits, but little is known about the function of different traits in mate choice. Here, we examine female courtship behaviours to learn how male traits interact to influence female mating decisions. In satin bowerbirds (Ptilonorhynchus violaceus), successful males give highly aggressive, intense behavioural displays without startling females. Males do this by modulating their displays in response to female crouching, which signals the display intensity that females will tolerate without being startled. Females typically visit multiple males for multiple courtships before choosing a mate, and females show differing tolerance for intense displays during their first courtship with each male. We test three hypotheses that may explain this: (i) familiarity with the courting male; (ii) the order of the courtship in mate-searching; and (iii) the attractiveness of the courting male. We found that females are more tolerant of intense displays during first courtships with attractive males; this increased female tolerance may allow attractive males to give higher intensity courtship displays that further enhance their attractiveness. We then examined why this is so, finding evidence that females are less likely to be startled by males with better physical displays (bower decorations), and this reduced startling then contributes to male courtship success. This role of physical displays in facilitating behavioural displays suggests a novel mechanism by which multiple physical and behavioural traits may influence female choice.     

Does Masculinity Matter? The Contribution of Masculine Face Shape to Male Attractiveness in Humans

Background

In many animals, exaggerated sex-typical male traits are preferred by females, and may be a signal of both past and current disease resistance. The proposal that the same is true in humans – i.e., that masculine men are immunocompetent and attractive – underpins a large literature on facial masculinity preferences. Recently, theoretical models have suggested that current condition may be a better index of mate value than past immunocompetence. This is particularly likely in populations where pathogenic fluctuation is fast relative to host life history. As life history is slow in humans, there is reason to expect that, among humans, condition-dependent traits might contribute more to attractiveness than relatively stable traits such as masculinity. To date, however, there has been little rigorous assessment of whether, in the presence of variation in other cues, masculinity predicts attractiveness or not.

Methodology/Principal Findings

The relationship between masculinity and attractiveness was assessed in two samples of male faces. Most previous research has assessed masculinity either with subjective ratings or with simple anatomical measures. Here, we used geometric morphometric techniques to assess facial masculinity, generating a morphological masculinity measure based on a discriminant function that correctly classified >96% faces as male or female. When assessed using this measure, there was no relationship between morphological masculinity and rated attractiveness. In contrast, skin colour – a fluctuating, condition-dependent cue – was a significant predictor of attractiveness.

Conclusions/Significance

These findings suggest that facial morphological masculinity may contribute less to men''s attractiveness than previously assumed. Our results are consistent with the hypothesis that current condition is more relevant to male mate value than past disease resistance, and hence that temporally fluctuating traits (such as colour) contribute more to male attractiveness than stable cues of sexual dimorphism.     

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.     

Egg quality and parental ornamentation in the blue tit Parus caeruleus

Components of avian egg quality often show remarkable variation between females and yet the causes of variation in this fitness-related trait remain poorly understood. We investigated the egg size and yolk carotenoid investment of blue tit Parus caeruleus females in relation to multiple parental traits. We show that females produce eggs with more colourful yolks when mated to ultraviolet (UV) attractive males. Since yolk carotenoids may enhance offspring fitness, but may be available to females in limiting amounts, the correlation we found suggests adaptive egg quality adjustment in response to mate attractiveness, but an experimental approach is required to rule out alternative explanations. Yolk colour was also correlated with laying date and yolk mass. Though the effect of laying date could not be explained by ambient temperature during egg formation, it suggests a proximate constraint of general carotenoid availability on yolk composition. Egg size was not affected by male attractiveness; however, females with a brighter crown produced larger eggs, suggesting that crown coloration might indicate individual quality in females too.     

The relative importance of intra- and intersexual selection on human male sexually dimorphic traits

Recent evidence suggests that in sexual selection on human males, intrasexual competition plays a larger role than female choice. In a sample of men (N?=?164), we sought to provide further evidence on the effects of men's physical dominance and sexual attractiveness on mating success and hence in sexual selection. Objective measures and subjective ratings of male sexually dimorphic traits purportedly under sexual selection (height, vocal and facial masculinity, upper body size from 3D scans, physical strength, and baseline testosterone) and observer perceptions of physical dominance and sexual attractiveness based on self-presentation video recordings were assessed and associated with mating success (sociosexual behaviour and number of potential conceptions) in a partly longitudinal design. Results from structural equation models and selection analyses revealed that physical dominance, but not sexual attractiveness, predicted mating success. Physical dominance mediated associations of upper body size, physical strength, as well as vocal and facial physical dominance and attractiveness with mating success. These findings thus suggest a greater importance of intrasexual competition than female choice in human male sexual selection.     

Quantifying the strength and form of sexual selection on men's traits

Although recent research has increasingly focused on human sexual selection, fundamental questions remain concerning the relative influence of individual traits on success in competition for mates and the mechanisms, form, and direction of these sexual selective pressures. Here, we explore sexual selection on men's traits by ascertaining men's dominance and attractiveness from male and female acquaintances. On a large American university campus, 63 men from two social fraternities provided anthropometric measurements, facial photographs, voice recordings, and reported mating success (number of sexual partners). These men also assessed each other's dominance, and 72 women from two socially affiliated sororities assessed the men's attractiveness. We measured facial masculinity from inter-landmark distances and vocal masculinity from acoustic parameters. We additionally obtained facial and vocal attractiveness and dominance ratings from unfamiliar observers. Results indicate that dominance and the traits associated with it predict men's mating success, but attractiveness and the traits associated with it do not. These findings point to the salience of contest competition on men's mating success in this population.     

Genetic diversity revealed in human faces

From an evolutionary perspective, human facial attractiveness is proposed to signal mate quality. Using a novel approach to the study of the genetic basis of human preferences for facial features, we investigated whether attractiveness signals mate quality in terms of genetic diversity. Genetic diversity in general has been linked to fitness and reproductive success, and genetic diversity within the major histocompatibility complex (MHC) has been linked to immunocompetence and mate preferences. We asked whether any preference for genetic diversity is specific to MHC diversity or reflects a more general preference for overall genetic diversity. We photographed and genotyped 160 participants using microsatellite markers situated within and outside the MHC, and calculated two measures of genetic diversity: mean heterozygosity and standardized mean d(2). Our results suggest a special role for the MHC in female preferences for male faces. MHC heterozygosity positively predicted male attractiveness, and specifically facial averageness, with averageness mediating the MHC-attractiveness relationship. For females, standardized mean d(2) at non-MHC loci predicted facial symmetry. Thus, attractive facial characteristics appear to provide visual cues to genetic quality in both males and females, supporting the view that face preferences have been shaped by selection pressures to identify high-quality mates.     

Facial attractiveness,developmental stability,and fluctuating asymmetry

Despite robust cross-cultural reliability of human facial attractiveness ratings, research on facial attractiveness has only superficially addressed the connection between facial attractiveness and the history of sexual selection in Homo sapiens. There are reasons to believe that developmental stability and phenotypic quality are related. Recent studies of nonhuman animals indicate that developmental stability, measured as fluctuating asymmetry in generally bilateral symmetrical traits, is predictive of performance in sexual selection: Relatively symmetrical males are advantaged under sexual selection. This pattern is suggested by our study of facial attractiveness and fluctuating asymmetry in seven bilateral body traits in a student population. Overall, facial attractiveness negatively correlated with fluctuating asymmetry; the relation for men, but not for women, was statistically reliable. Possible confounding factors were controlled for in the analysis.     

"You are so beautiful"*: Behind women's attractiveness towards the biology of reproduction: a narrative review

Female beauty has always attracted human beings. In particular, beauty has been interpreted in terms of reproductive potential and advantage in selection of mates. We have reviewed the recent literature on female facial and physical beauty with the objective of defining which parameters could influence female attractiveness. Symmetry, averageness, and sexual dimorphism with regards to facial beauty, as well as waist-to-hip ratio (WHR), breast size, and body mass index (BMI) for physical beauty, have been assessed. In current societies, it appears that facial attractiveness results from a mixture of symmetry and averageness of traits, high forehead and cheekbones, small nose and chin, full lips, thin eyebrows, and thick hair. A low WHR reliably characterized physical attractiveness, whereas inconsistencies have been observed in the evaluation of breast size and BMI. The importance of breast size appears to vary with time and sex of evaluators, whereas the impact of BMI is related to socio-economic conditions. The various hypotheses behind beauty and the role of attractiveness in mate choice and sexual selection are here described in terms of continuation of human species. Intriguing associations are emerging between features of attractiveness and some reproductive disorders, as both are substantially influenced by sex steroid hormones.     

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.     

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.     

Attractiveness of men's faces in relation to women's phase of menstrual cycle

In between-subjects studies on two groups of women of the same age, we show that women assess male's facial attractiveness differently in the follicular (F) and luteal (L) phases. In the high conception risk phase (F), women tended to give higher scores to male faces than when they were in the luteal phase. During the five first days of the cycle, i.e. when the estrogen level is still low, women assessed men's facial attractiveness relatively highly. We suggest that it is progesterone in the luteal phase that is responsible for lower attractiveness assigned then to male faces. We also tested which anthropometric facial traits or indices influence male attractiveness. We found that assessments of attractiveness were correlated only with mouth height (positively) and the angle between the middle of the mouth and the middle of the eyes (negatively). The results are compared with those from other studies and discussed in the light of evolutionary biology.     

Advanced paternal age is associated with lower facial attractiveness

In view of disease risk, Kong et al. (2012) demonstrated that most of the new mutations are explained by the age of the father at conception. Accordingly, paternal age effects have been found for a variety of offspring traits, from physical and mental health to intelligence. Here, we investigated whether facial attractiveness is significantly associated with paternal age. We used the Wisconsin Longitudinal Study (n = 4018 male and 4416 female high school graduates) to analyze the association between an individual's father's age at birth and that individual's facial attractiveness (estimated by rating the high school yearbook photographs from 1957), controlling for sex, age as well as mother's age. We find that subject's facial attractiveness decreased with advancing paternal but not maternal age, suggesting that facial attractiveness might be a cue of an individual's new mutation load.     

Sexual Conflict and Gender Gap Effects: Associations between Social Context and Sex on Rated Attractiveness and Economic Status

Human mate choice research often concerns sex differences in the importance of traits such as physical attractiveness and social status. A growing number of studies indicate that cues to social context, including other people who appear in stimulus photographs, can alter that individual’s attractiveness. Fewer studies, however, consider judgements of traits other than physical attractiveness, such as wealth. Here we manipulate the presence/absence of other people in photographs of target models, and test the effects on judgments of both attractiveness and earnings (a proxy for status). Participants (N = 2044) rated either male or female models for either physical attractiveness or social/economic status when presented alone, with same sex others or with opposite sex others. We collectively refer to this manipulation as ‘social context’. Male and female models received similar responses for physical attractiveness, but social context affected ratings of status differently for women and men. Males presented alongside other men received the highest status ratings while females presented alone were given the highest status ratings. Further, the status of females presented alongside a male was constrained by the rated status of that male. Our results suggests that high status may not directly lead to high attractiveness in men, but that status is more readily attributed to men than to women. This divide in status between the sexes is very clear when men and women are presented together, possibly reflecting one underlying mechanism of the modern day gender gap and sexist attitudes to women’s economic participation. This adds complexity to our understanding of the relationship between attractiveness, status, and sex in the light of parental investment theory, sexual conflict and economic theory.