Immune function during early adolescence positively predicts adult facial sexual dimorphism in both men and women

Evolutionary theories suggest that humans prefer sexual dimorphism in faces because masculinity in men and femininity in women may be an indicator of immune function during development. In particular, the immunocompetence handicap hypothesis proposes that sexual dimorphism indicates good immune function during development because the sex hormones, particularly testosterone in men, required for the development of sexually dimorphic facial features also taxes the immune system. Therefore, only healthy males can afford the high level of testosterone for the development of sexually dimorphic traits without compromising their survival. Researchers have suggested that a similar mechanism via the effects of oestrogen might also explain male preferences for female femininity. Despite the prominence of the immunocompetence handicap hypothesis, no studies have tested whether immune function during development predicts adult facial sexual dimorphism. Here, using data from a longitudinal public health dataset, the Western Australian Pregnancy Cohort (Raine) Study (Generation 2), we show that some aspects of immune function during early adolescence (14 years) positively predict sexually dimorphic 3D face shape in both men and women. Our results support a fundamental assumption that facial sexual dimorphism is an indicator of immune function during the development of facial sexual dimorphism.     

Self-perceived attractiveness influences human female preferences for sexual dimorphism and symmetry in male faces

Exaggerated sexual dimorphism and symmetry in human faces have both been linked to potential 'good-gene' benefits and have also been found to influence the attractiveness of male faces. The current study explores how female self-rated attractiveness influences male face preference in females using faces manipulated with computer graphics. The study demonstrates that there is a relatively increased preference for masculinity and an increased preference for symmetry for women who regard themselves as attractive. This finding may reflect a condition-dependent mating strategy analogous to behaviours found in other species. The absence of a preference for proposed markers of good genes may be adaptive in women of low mate value to avoid the costs of decreased parental investment from the owners of such characteristics.     

Manipulating Shape Cues in Dynamic Human Faces: Sexual Dimorphism is Preferred in Female but not Male Faces

Sexual dimorphism in the human face has been linked to attractiveness, and computer‐graphic techniques have been useful in this field by allowing experimental manipulation of dimorphism. However, a limitation of much research is its reliance on static pictorial stimuli, whereas real faces are dynamic, as is much courtship behaviour throughout nature. Furthermore, little is known about possible interactions between static and dynamic facial cues and attractiveness. We adapted well‐established face‐morphing technology to manipulate sexual dimorphism in male and female dynamic facial displays depicting prosocial and antisocial behaviour. Masculinised and feminised versions of these videos were presented as a two‐alternative forced choice task to measure preferences. Feminised female videos were preferred in both movement contexts, as expected. More surprisingly, no directional preference for masculinity or femininity was evident for the male videos in either context. Further analysis showed that the movement of the faces (prosocial vs. antisocial) had no effect on attractiveness ratings. Results were the same with static stimuli and did not conflict with findings from the wider literature using static faces. These findings suggest that the new technique we describe is a valid way to manipulate facial shape in videos and can now be widely applied to future studies of facial morphing.     

Preferences for masculinity in male bodies change across the menstrual cycle

In human females cyclic shifts in preference have been documented for odour and physical and behavioral male traits. Women prefer the smell of dominant males, more masculine male faces and men behaving more dominantly when at peak fertility than at other times in their menstrual cycle. Here we examine variation in preferences for body sexual dimorphism. Across two studies, both between- and within-participant, we show that women prefer greater masculinity in male bodies at times when their fertility is likely highest, in the follicular phase of their cycle. Shifts were seen when rating for a short-term but not when rating for a long-term relationship. In line with studies showing similar effects for facial sexual dimorphism, we also show that women prefer greater masculinity when they think themselves attractive than when they think themselves less attractive. These results indicate that women's preferences for sexual dimorphism in male bodies follow a similar pattern as found for sexual dimorphism and dominance in other domains and such differences in preference may serve a similar function. Cyclic preferences could influence women to select partners when most likely to become pregnant that possess traits that may be most likely to maximize their offspring's quality via attraction to masculinity or serve to help acquire investment via attraction to femininity.     

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

Studies of women's 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 women's 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 women's 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 women's 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.     

The health of a nation predicts their mate preferences: cross-cultural variation in women's preferences for masculinized male faces

Recent formulations of sexual selection theory emphasize how mate choice can be affected by environmental factors, such as predation risk and resource quality. Women vary greatly in the extent to which they prefer male masculinity and this variation is hypothesized to reflect differences in how women resolve the trade-off between the costs (e.g. low investment) and benefits (e.g. healthy offspring) associated with choosing a masculine partner. A strong prediction of this trade-off theory is that women''s masculinity preferences will be stronger in cultures where poor health is particularly harmful to survival. We investigated the relationship between women''s preferences for male facial masculinity and a health index derived from World Health Organization statistics for mortality rates, life expectancies and the impact of communicable disease. Across 30 countries, masculinity preference increased as health decreased. This relationship was independent of cross-cultural differences in wealth or women''s mating strategies. These findings show non-arbitrary cross-cultural differences in facial attractiveness judgements and demonstrate the use of trade-off theory for investigating cross-cultural variation in women''s mate preferences.     

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.     

The relationship between sexual dimorphism in human faces and fluctuating asymmetry

Previous studies have found both support and lack of support for a positive relationship between masculinity and symmetry, two putative signs of mate quality, in male faces. We re-examined this relationship using an explicit measure of facial fluctuating asymmetry, as well as other measures of asymmetry, and measures of facial masculinity/femininity. We also used ratings of these traits for faces. Further, we examined the relationship between facial sexual dimorphism and body asymmetry. We found no significant correlations between facial masculinity and any of our measures of asymmetry or ratings of symmetry in males. Facial femininity was not consistently associated with facial symmetry in females, but was associated with body symmetry. Therefore, for females, but not males, facial femininity and body symmetry may reflect similar aspects of mate quality. We also examined the relationships between trait ratings and measurements. Our results provide validation of our ability to measure aspects of asymmetry that are perceived to be symmetrical, and aspects of sexual dimorphism that are perceived as feminine in females and masculine in males.     

Male-Biased Sex Ratios in Offspring of Attractive Males in the Guppy

The attractiveness hypothesis predicts that females produce offspring with male-biased sex ratios when they mate with attractive males because their male offspring will inherit the paternal sexual attractiveness and may have high reproductive success. In this study, we examined the effect of the attractiveness of the male guppy Poecilia reticulata in terms of the conspicuousness of its orange spot patterns, important criteria affecting female choice in this species, on the offspring sex ratios. We found that food-manipulation treatment altered the conspicuousness of the orange spot patterns in a full-sibling male pair. When females were presented to these males, they showed a greater mate preference for males having brighter orange spots than for those having duller orange spots. Subsequently, half of the females were mated with the preferred males and the remaining females were mated with the less preferred males. When the females exhibited a greater preference for their mates, their offspring sex ratios were more male biased. These results appear to be consistent with the prediction of the attractiveness hypothesis. In the guppy, as male sexual attractiveness is heritable, the male-biased sex ratios of the broods of attractive males may be adaptive.     

Partnership status and the temporal context of relationships influence human female preferences for sexual dimorphism in male face shape

Secondary sexual characteristics may indicate quality of the immune system and therefore a preference for masculinity may confer genetic benefits to offspring; however, high masculinity may be associated with costs of decreased paternal investment. The current study examined women's preferences for masculinity in male faces by using computer graphics to allow transformation between feminine and masculine versions of individual male faces. We found that preferences for masculinity are increased when women either have a partner or are considering a short-term relationship. Such preferences are potentially adaptive, serving to: (i) maximize parental investment and cooperation in long-term relationships by biasing choices towards feminine faced males, and (ii) maximize possible good-gene benefits of short-term or extra-pair partners by biasing choices towards masculine faced males. We also found that individuals using oral contraception do not show the above effects, indicating that such hormonal intervention potentially disrupts women's choices for evolutionarily relevant benefits from males.     

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.     

A Lack of Sexual Dimorphism in Width-to-Height Ratio in White European Faces Using 2D Photographs, 3D Scans, and Anthropometry

Facial width-to-height ratio has received a great deal of attention in recent research. Evidence from human skulls suggests that males have a larger relative facial width than females, and that this sexual dimorphism is an honest signal of masculinity, aggression, and related traits. However, evidence that this measure is sexually dimorphic in faces, rather than skulls, is surprisingly weak. We therefore investigated facial width-to-height ratio in three White European samples using three different methods of measurement: 2D photographs, 3D scans, and anthropometry. By measuring the same individuals with multiple methods, we demonstrated high agreement across all measures. However, we found no evidence of sexual dimorphism in the face. In our third study, we also found a link between facial width-to-height ratio and body mass index for both males and females, although this relationship did not account for the lack of dimorphism in our sample. While we showed sufficient power to detect differences between male and female width-to-height ratio, our results failed to support the general hypothesis of sexual dimorphism in the face.     

Body Shape Preferences: Associations with Rater Body Shape and Sociosexuality

There is accumulating evidence of condition-dependent mate choice in many species, that is, individual preferences varying in strength according to the condition of the chooser. In humans, for example, people with more attractive faces/bodies, and who are higher in sociosexuality, exhibit stronger preferences for attractive traits in opposite-sex faces/bodies. However, previous studies have tended to use only relatively simple, isolated measures of rater attractiveness. Here we use 3D body scanning technology to examine associations between strength of rater preferences for attractive traits in opposite-sex bodies, and raters’ body shape, self-perceived attractiveness, and sociosexuality. For 118 raters and 80 stimuli models, we used a 3D scanner to extract body measurements associated with attractiveness (male waist-chest ratio [WCR], female waist-hip ratio [WHR], and volume-height index [VHI] in both sexes) and also measured rater self-perceived attractiveness and sociosexuality. As expected, WHR and VHI were important predictors of female body attractiveness, while WCR and VHI were important predictors of male body attractiveness. Results indicated that male rater sociosexuality scores were positively associated with strength of preference for attractive (low) VHI and attractive (low) WHR in female bodies. Moreover, male rater self-perceived attractiveness was positively associated with strength of preference for low VHI in female bodies. The only evidence of condition-dependent preferences in females was a positive association between attractive VHI in female raters and preferences for attractive (low) WCR in male bodies. No other significant associations were observed in either sex between aspects of rater body shape and strength of preferences for attractive opposite-sex body traits. These results suggest that among male raters, rater self-perceived attractiveness and sociosexuality are important predictors of preference strength for attractive opposite-sex body shapes, and that rater body traits –with the exception of VHI in female raters– may not be good predictors of these preferences in either sex.     

Vocal and Facial Attractiveness Judgments of Children, Adolescents and Adults: the Ontogeny of Mate Choice

Physical traits are thought to be used as indicators of mate quality, allowing individuals to select mates most likely to help them bear the fittest offspring. As the capacity for human sexual behaviour emerges at puberty, we investigated whether adult‐like judgments of the relative attractiveness of opposite‐sex individuals also arise at puberty. Following previous research, we focussed on facial and vocal attractiveness, which are known to be used in human mate choice and to carry concordant information regarding mate quality. Here we show that males with more attractive faces have more attractive voices as judged by female adults and adolescents, but not by female children. This suggests that cues of facial and vocal attractiveness provide similar information, but that awareness of these cues does not develop fully until reproductive capability, when mate choice judgments become relevant. Adolescents’ judgments also mirrored those of adults in that, like adults, they preferred lower‐pitched male voices, and as a group made concordant judgments about facial attractiveness. However, they did not make similarly concordant judgments about vocal attractiveness, suggesting that a further period of maturation and learning is required to fully develop optimal judgments for mate choice.     

The Morphometrics of “Masculinity” in Human Faces

In studies of social inference and human mate preference, a wide but inconsistent array of tools for computing facial masculinity has been devised. Several of these approaches implicitly assumed that the individual expression of sexually dimorphic shape features, which we refer to as maleness, resembles facial shape features perceived as masculine. We outline a morphometric strategy for estimating separately the face shape patterns that underlie perceived masculinity and maleness, and for computing individual scores for these shape patterns. We further show how faces with different degrees of masculinity or maleness can be constructed in a geometric morphometric framework. In an application of these methods to a set of human facial photographs, we found that shape features typically perceived as masculine are wide faces with a wide inter-orbital distance, a wide nose, thin lips, and a large and massive lower face. The individual expressions of this combination of shape features—the masculinity shape scores—were the best predictor of rated masculinity among the compared methods (r = 0.5). The shape features perceived as masculine only partly resembled the average face shape difference between males and females (sexual dimorphism). Discriminant functions and Procrustes distances to the female mean shape were poor predictors of perceived masculinity.     

Phenotypic correlates of male reproductive success in western gorillas

Sexual selection is thought to drive the evolution of sexually dimorphic traits that increase male reproductive success. Despite a large degree of sexual dimorphism among haplorhine primates, phenotypic traits that may influence the reproductive success of males are largely unstudied due to long life spans and the difficulties in quantifying such traits non-invasively. Here we employ digital photogrammetry of body length and crest size, as well as ranking of the gluteal muscle size, to test whether these sexually dimorphic traits are associated with long-term measures of male reproductive success in western gorillas. Among 19 adult male gorillas monitored for up to 12.5 years, we found that all three phenotypic traits were positively correlated with the average number of mates per male, but only crest size and gluteal muscle size were significantly correlated with offspring survival and the annual rate of siring offspring that survive to weaning age. We discuss why such sexually dimorphic traits might be under ongoing selection in gorillas and other species.     

Evidence of adaptation for mate choice within women's memory

Sexually dimorphic characteristics in men may act as cues, advertising long-term health, dominance, and reproductive potential to prospective mates. Evolution has accordingly adapted human cognition so that women perceive sexually dimorphic facial features as important when judging the attractiveness and suitability of potential mates. Here we provide evidence showing, for the first time, that women's memory for details encountered in recently experienced episodes is also systematically biased by the presence of men's facial cues signaling enhanced or reduced sexual dimorphism. Importantly, the direction and strength of this bias are predicted by individual differences in women's preferences for masculine versus feminine facial features in men and are triggered specifically while viewing images of male but not female faces. No analogous effects were observed in male participants viewing images of feminized and masculinized women's faces despite the fact that male participants showed strong preferences for feminized facial features. These findings reveal a preference-dependent memory enhancement in women that would promote retention of information from encounters with preferred potential mates. We propose that women's memory for recently experienced episodes may therefore be functionally specialized for mate choice and in particular for the comparative evaluation of alternative potential mates. This also raises the possibility that similar specialization may be present in other species where it has been established that precursor, ‘episodic-like’ forms of memory exist.     

Facing aggression: cues differ for female versus male faces

The facial width-to-height ratio (face ratio), is a sexually dimorphic metric associated with actual aggression in men and with observers'' judgements of aggression in male faces. Here, we sought to determine if observers'' judgements of aggression were associated with the face ratio in female faces. In three studies, participants rated photographs of female and male faces on aggression, femininity, masculinity, attractiveness, and nurturing. In Studies 1 and 2, for female and male faces, judgements of aggression were associated with the face ratio even when other cues in the face related to masculinity were controlled statistically. Nevertheless, correlations between the face ratio and judgements of aggression were smaller for female than for male faces (F_1,36 = 7.43, p = 0.01). In Study 1, there was no significant relationship between judgements of femininity and of aggression in female faces. In Study 2, the association between judgements of masculinity and aggression was weaker in female faces than for male faces in Study 1. The weaker association in female faces may be because aggression and masculinity are stereotypically male traits. Thus, in Study 3, observers rated faces on nurturing (a stereotypically female trait) and on femininity. Judgements of nurturing were associated with femininity (positively) and masculinity (negatively) ratings in both female and male faces. In summary, the perception of aggression differs in female versus male faces. The sex difference was not simply because aggression is a gendered construct; the relationships between masculinity/femininity and nurturing were similar for male and female faces even though nurturing is also a gendered construct. Masculinity and femininity ratings are not associated with aggression ratings nor with the face ratio for female faces. In contrast, all four variables are highly inter-correlated in male faces, likely because these cues in male faces serve as “honest signals”.     

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.     

A multivariate approach to human mate preferences

Human mate choice is complicated, with various individual differences and contextual factors influencing preferences for numerous traits. However, focused studies on human mate choice often do not capture this multivariate complexity. Here, we consider multiple factors simultaneously to demonstrate the advantages of a multivariate approach to human mate preferences. Participants (N = 689) rated the attractiveness of opposite-sex online dating profiles that were independently manipulated on facial attractiveness, perceived facial masculinity/femininity, and intelligence. Participants were also randomly instructed to either consider short- or long-term relationships. Using fitness surfaces analyses, we assess the linear and nonlinear effects and interactions of the profiles' facial attractiveness, perceived facial masculinity/femininity, and perceived intelligence on participants' attractiveness ratings. Using hierarchical linear modeling, we were also able to consider the independent contribution of participants' individual differences on their revealed preferences for the manipulated traits. These individual differences included participants' age, socioeconomic status, education, disgust (moral, sexual, and pathogen), sociosexual orientation, personality variables, masculinity, and mate value. Together, our results illuminate various previously undetectable phenomena, including nonlinear preference functions and interactions with individual differences. More broadly, the study illustrates the value of considering both individual variation and population-level measures when addressing questions of sexual selection, and demonstrates the utility of multivariate approaches to complement focused studies.