Second to fourth digit ratio and face shape

The average human male face differs from the average female face in size and shape of the jaws, cheek-bones, lips, eyes and nose. It is possible that this dimorphism is determined by sex steroids such as testosterone (T) and oestrogen (E), and several studies on the perception of such characteristics have been based on this assumption, but those studies focussed mainly on the relationship of male faces with circulating hormone levels; the corresponding biology of the female face remains mainly speculative. This paper is concerned with the relative importance of prenatal T and E levels (assessed via the 2D : 4D finger length ratio, a proxy for the ratio of T/E) and sex in the determination of facial form as characterized by 64 landmark points on facial photographs of 106 Austrians of college age. We found that (i) prenatal sex steroid ratios (in terms of 2D : 4D) and actual chromosomal sex dimorphism operate differently on faces, (ii) 2D : 4D affects male and female face shape by similar patterns, but (iii) is three times more intense in men than in women. There was no evidence that these effects were confounded by allometry or facial asymmetry. Our results suggest that studies on the perception of facial characteristics need to consider differential effects of prenatal hormone exposure and actual chromosomal gender in order to understand how characteristics have come to be rated 'masculine' or 'feminine' and the consequences of these perceptions in terms of mate preferences.     

Visualizing facial shape regression upon 2nd to 4th digit ratio and testosterone

Sex steroids are supposed to moderate the differences between male and female facial characteristics. Studies on women's preferences for male faces reported increased preferences for facial architecture developed under the influence of testosterone as this may indicate masculinity, dominance and social status. Recent research demonstrates that facial sexual dimorphism does not only develop at puberty but may be organized much earlier in ontogeny. However, the actual cause and timing of variation in facial shape due to sex-steroids remains speculative. This study uses data from Neave and colleagues who measured digit ratio (2D:4D) as a proxy to prenatal testosterone and also salivary testosterone samples in order to study differential effects of androgens on perceived male facial shape. Male facial shape was regressed upon 2D:4D ratio and circulating levels of testosterone by means of geometric morphometric methods. We found some evidence for opposite effects of early androgen action (via 2D:4D ratio) on the upper and the lower face respectively (i.e. low 2D:4D ratio results in a relatively robust and prominent lower face), whereas circulating testosterone seems to cause a rather uniform elongation of the face. Local deformations primarily show pronounced and medially tailed eyebrows for the shapes associated with increasing salivary testosterone. These preliminary results suggest that prenatal and pubertal testosterone have differential effects on male facial shape that should be considered in future studies on women's preferences towards male facial appearance.     

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.     

Testosterone increases perceived dominance but not attractiveness in human males

Recent evidence suggests that certain features on the human face indicate hormonal levels during growth, and that women judge the attractiveness of potential partners based on the appearance of these features. One entrenched notion is male facial features that are affected by testosterone are used as direct cues in mate preference. Testosterone may be particularly revealing as it is purported to be an honest indicator of male fitness. Increased testosterone may impose an immunocompetence handicap on the bearer and only the best males can carry this handicap. To date, tests of this theory have been indirect, and have relied on digital manipulations that represent unrealistic continuums of masculine and feminine faces. We provide a much more direct test by manipulating digitally male faces to mimic known shape variation, caused by varying levels of testosterone through puberty. We produced a continuum of faces that ranged from low to high levels of testosterone in male faces and asked women to choose the points on the continuum that appeared most attractive and most physically dominant. Our data indicate that high testosterone faces reveal dominance. However, there is no evidence of directional selection for increased (or decreased) testosterone in terms of attractiveness to the opposite sex. We discuss the relevance and applicability of evolutionary interpretations of our data and, contrary to predictions, provide evidence of stabilizing selection acting on testosterone through mate preferences.     

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.     

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”.     

Reproductive strategy, sexual development and attraction to facial characteristics

Sexual reproduction strategies vary both between and within species in the level of investment in offspring. Life-history theories suggest that the rate of sexual maturation is critically linked to reproductive strategy, with high investment being associated with few offspring and delayed maturation. For humans, age of puberty and age of first sex are two developmental milestones that have been associated with reproductive strategies. Stress during early development can retard or accelerate sexual maturation and reproduction. Early age of menarche is associated with absence of younger siblings, absence of a father figure during early life and increased weight. Father absence during early life is also associated with early marriage, pregnancy and divorce. Choice of partner characteristics is critical to successful implementation of sexual strategies. It has been suggested that sexually dimorphic traits (including those evident in the face) signal high-quality immune function and reproductive status. Masculinity in males has also been associated with low investment in mate and offspring. Thus, women's reproductive strategy should be matched to the probability of male investment, hence to male masculinity. Our review leads us to predict associations between the rate of sexual maturation and adult preferences for facial characteristics (enhanced sexual dimorphism and attractiveness). We find for men, engaging in sex at an early age is related to an increased preference for feminized female faces. Similarly, for women, the earlier the age of first sex the greater the preference for masculinity in opposite-sex faces. When we controlled sexual dimorphism in male faces, the speed of sexual development in women was not associated with differences in preference for male facial attractiveness. These developmental influences on partner choice were not mediated by self-rated attractiveness or parental relationships. We conclude that individuals assort in preferences based on the rapidity of their sexual development. Fast developing individuals prefer opposite-sex partners with an increased level of sexually dimorphic facial characteristics.     

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.     

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.     

Populational differences in attractiveness judgements of male and female faces: Comparing British and Jamaican samples

In the UK and Japan, both men and women prefer somewhat feminised opposite-sex faces, especially when choosing a long-term partner. Such faces are perceived as more honest, caring, and sensitive; traits that may be associated with successful male parental investment. By contrast, women prefer less feminised faces for short-term relationships and when they are near ovulation. As genetic quality may be associated with facial masculinity, women may ‘trade-off’ cues between genetic quality and paternal investment in potential partners. No analogous trade-off has been suggested to influence men's preferences, as both attributions of prosociality and potential cues to biological quality are associated with facial femininity in female faces. Ecological and cultural factors may influence the balance of trade-offs leading to populational differences in preferences. We predicted that Jamaican women would prefer more masculine faces than British women do because parasite load is higher in Jamaica, medical care less common (historically and currently), and male parental investment less pronounced. Male preferences, however, were predicted to vary less cross-culturally, as no trade-off has been identified in female facial characteristics. We constructed masculinised and feminised digital male and female face stimuli of three populations (Jamaican, Japanese, and British) and presented them to men and women in Jamaica and in Britain. The results demonstrated that Jamaican women preferred more masculine male faces than their British counterparts did. Jamaican men tended to prefer more masculine female faces than did British men did, but this effect was complicated by an interaction suggesting that more feminised faces were preferred within culture.     

Is cooperativeness readable in static facial features? An inter-cultural approach

There is evidence in the literature that non-verbal physical features are used as cues for a propensity to cooperate. However, further studies of the human ability to visually detect cooperativeness are required. In particular, the existence of static facial cues of altruism remains questionable. Moreover, an investigation of both sex differences and cross-cultural applicability with respect to altruism detection skills is crucial in the context of the evolution of human cooperation. In this study, we used both a public good game and a charitable contribution to assess the cooperativeness of 156 men and 172 women in rural Senegal and took facial photographs of these individuals. The second portion of the study was conducted in France. In total, 194 men and 171 women were asked to distinguish the most and least selfish individual from a series of 80 pairs of Senegalese facial photographs, each pair consisting of the highest and the lowest contributor from a group in the public good game. Using mixed modeling techniques, we controlled for facial masculinity, age and socio-economic status. For male pairs, both male and female French raters were able to identify more often than by chance which individual made the smallest contribution to the public good in each group; however, detection was not successful with female faces. These results suggest that sex-specific traits are involved and that only male facial traits indicating cooperative skills are, at least inter-culturally, readable. The specific facial traits involved are investigated. However, the charitable contribution was not correlated with the contribution to the public good, and further work is necessary to identify which specific altruistic traits are detectable and to assess the generality of these results.     

Female and male responses to cuteness,age and emotion in infant faces

Neonatal features in the newborn are thought to trigger parental care, the most fundamental prosocial behaviour. The underlying mechanisms that release parental care have not yet been resolved. Here we report sex differences in the ability to discriminate cues to cuteness despite equivalence in the capability to discriminate age and facial expression. These differences become apparent in a task where adults were asked to choose the cuter of two babies. While women could reliably choose the cuter infant, men had more difficulty in doing so. When showing the exact same face pairs but asking to choose the younger or the happier baby, there was no sex difference. These results suggest that the sex difference in the ability to discriminate cues to cuteness in infants underlies female-specific emotive responses. We argue that this reactivity expressed by women evolved to ensure that a female allocates her caretaking resources to her youngest offspring while it needs mothering care.     

Testosterone responses to competition in men are related to facial masculinity

Relationships between androgens and the size of sexually dimorphic male traits have been demonstrated in several non-human species. It is often assumed that a similar relationship exists for human male faces, but clear evidence of an association between circulating testosterone levels and the size of masculine facial traits in adulthood is absent. Here we demonstrate that, after experimentally determined success in a competitive task, men with more a masculine facial structure show higher levels of circulating testosterone than men with less masculine faces. In participants randomly allocated to a 'winning' condition, testosterone was elevated relative to pre-task levels at 5 and 20 min post-task. In a control group of participants allocated to a 'losing' condition there were no significant differences between pre- and post-task testosterone. An index of facial masculinity based on the measurement of sexually dimorphic facial traits was not associated with pre-task (baseline) testosterone levels, but was associated with testosterone levels 5 and 20 min after success in the competitive task. These findings indicate that a man's facial structure may afford important information about the functioning of his endocrine system.     

Testosterone reactivity to facial display of emotions in men and women

Previous studies have examined testosterone's role in regulating the processing of facial displays of emotions (FDEs). However, the reciprocal process – the influence of FDEs, an evolutionarily ancient and potent class of social signals, on the secretion of testosterone – has not yet been studied. To address this gap, we examined the effects of emotional content and sex of facial stimuli in modulating endogenous testosterone fluctuations, as well as sex differences in the endocrine responses to faces. One hundred and sixty-four young healthy men and women were exposed, in a between-subjects design, to happy or angry same-sex or opposite-sex facial expressions. Results showed that in both men (n = 85) and women (n = 79), extended exposure to faces of the opposite sex, regardless of their apparent emotional content, was accompanied by an accumulation in salivary testosterone when compared to exposure to faces of the same sex. Furthermore, testosterone change in women exposed to angry expressions was greater than testosterone change in women exposed to happy expressions. These results add emotional facial stimuli to the collection of social signals that modulate endocrine status, and are discussed with regard to the evolutionary roles of testosterone.     

Facial masculinity and fluctuating asymmetry

Recently, women have been found to prefer the scent of symmetrical men and relatively masculine male faces more during the fertile (late follicular and ovulatory) phases of their menstrual cycles than during their infertile (e.g., luteal) phases. These findings make most theoretical sense if men's symmetry is associated with the masculinity of their faces and, therefore, men's symmetry and facial masculinity tap a shared underlying quality. This study examined associations between masculine facial features and nonfacial body symmetry as well as facial symmetry in samples of 141 men and 154 women. As predicted, a component of facial features that discriminates the sexes and reflects masculinization of the face significantly covaried with symmetry in men. No significant correlation was observed for women. These findings suggest that men's facial masculinity partly advertises underlying developmental stability.     

Is facial width-to-height ratio reliably associated with social inferences?

Theoretical considerations and early empirical findings suggested facial width-to-height ratio (fWHR) may be relevant to person perception because it is associated with behavioral dispositions. More recent evidence failing to find fWHR-behavior links suggests that mismatch or byproduct hypotheses may be necessary to explain fWHR-based trait inferences; however, these explanations may not be needed because it is not clear that fWHR is reliably associated with trait inferences. To investigate the robustness of fWHR-inference links, we conducted secondary analyses of a cross-national dataset consisting of ratings by 11,481 participants across 11 world regions who judged 60 male and 60 female faces on one of 13 social traits (ns per trait range from 760 to 975). In preregistered analyses—and exploratory analyses of a subset of traits in the larger sample of 597 faces from which the 120 faces were drawn—we found mixed evidence for fWHR-based social judgments. In multilevel models, fWHR was not reliably linked to raters' judgments of male faces for any of the 13 trait-inferences but was negatively associated with ratings of female faces' dominance, trustworthiness, sociability, emotional stability, responsibility, confidence, attractiveness, and intelligence. In exploratory analyses of a subset of traits using the larger sample of faces, fWHR was associated positively with perceptions of meanness and aggressiveness in male but not female faces, negatively with attractiveness and dominance in female but not male faces, and negatively with trustworthiness in male but not female faces. We interpret these mixed findings to suggest that (1) fWHR-inference links are likely to be smaller and less reliable than expected from prior research; (2) fWHR may play a larger role in perceptions of female faces than would be predicted from the theory underpinning fWHR hypotheses; and (3) future research should more closely examine the extent to which robust fWHR-inferences reflect mismatch in the reliability of fWHR-behavior links between ancestral and modern environments versus byproducts of other person perception mechanisms.     

The Thatcher illusion in humans and monkeys

Primates possess the remarkable ability to differentiate faces of group members and to extract relevant information about the individual directly from the face. Recognition of conspecific faces is achieved by means of holistic processing, i.e. the processing of the face as an unparsed, perceptual whole, rather than as the collection of independent features (part-based processing). The most striking example of holistic processing is the Thatcher illusion. Local changes in facial features are hardly noticeable when the whole face is inverted (rotated 180°), but strikingly grotesque when the face is upright. This effect can be explained by a lack of processing capabilities for locally rotated facial features when the face is turned upside down. Recently, a Thatcher illusion was described in the macaque monkey analogous to that known from human investigations. Using a habituation paradigm combined with eye tracking, we address the critical follow-up questions raised in the aforementioned study to show the Thatcher illusion as a function of the observer''s species (humans and macaques), the stimulus'' species (humans and macaques) and the level of perceptual expertise (novice, expert).     

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.     

Stable face representations

Photographs are often used to establish the identity of an individual or to verify that they are who they claim to be. Yet, recent research shows that it is surprisingly difficult to match a photo to a face. Neither humans nor machines can perform this task reliably. Although human perceivers are good at matching familiar faces, performance with unfamiliar faces is strikingly poor. The situation is no better for automatic face recognition systems. In practical settings, automatic systems have been consistently disappointing. In this review, we suggest that failure to distinguish between familiar and unfamiliar face processing has led to unrealistic expectations about face identification in applied settings. We also argue that a photograph is not necessarily a reliable indicator of facial appearance, and develop our proposal that summary statistics can provide more stable face representations. In particular, we show that image averaging stabilizes facial appearance by diluting aspects of the image that vary between snapshots of the same person. We review evidence that the resulting images can outperform photographs in both behavioural experiments and computer simulations, and outline promising directions for future research.     

Facial shape provides a valid cue to sociosexuality in men but not women

Existing work suggests that observers' perceptions of sociosexuality from strangers' faces are positively associated with individuals' self-reported sociosexuality. However it is not clear what cues observers use to form these judgements. Over two studies we examined whether sociosexuality is reflected in faces, which cues contain information about sociosexuality, and whether observers' perceptions of sociosexuality from faces are positively associated with individuals' self-reported sociosexuality. In Study One, Geometric Morphometric Modelling (GMM) analysis of 103 Caucasian participants revealed that self-reported sociosexuality was predicted by facial morphology in male but not female faces. In Study Two, 65 Caucasian participants judged the sociosexuality of opposite sex faces (faces from Study One) at zero acquaintance. Perceived sociosexuality predicted self-reported sociosexuality for men, but not women. Participants were also presented with composites of faces of individuals with more unrestricted sociosexuality paired with composites of faces of individuals with more restricted sociosexuality and asked to indicate which was more unrestricted. Participants selected the more unrestricted sociosexuality male, but not female, facial composites at rates significantly above chance. GMM analyses also found that facial morphology statistically significantly predicted perceived sociosexuality in women's and, to a greater extent, in men's faces. Finally, facial shape mediated the relationship between perceived sociosexuality and self-reported sociosexuality in men's but not women's faces. Our results suggest that facial shape acts as a valid cue to sociosexuality in men's but not women's faces.