Re-evaluating the relationship between pathogen avoidance and preferences for facial symmetry and sexual dimorphism: A registered report

Over the past decade, a small literature has tested how trait-level pathogen-avoidance motives (e.g., disgust sensitivity) and exposure to pathogen cues relate to preferences for facial symmetry and sexual dimorphism. Results have largely been interpreted as suggesting that the behavioral immune system influences preferences for these features in potential mates. However, findings are limited by small sample sizes among studies reporting supportive evidence, the use of small stimulus sets to assess preferences for symmetry and dimorphism, and design features that render implications for theory ambiguous (namely, largely only investigating women's preferences for male faces). Using a sample of 954 White young adult UK participants and a pool of 100 White young adult stimuli, the current registered report applied a standard two-alternative forced-choice approach to evaluate both men's and women's preferences for both facial symmetry and dimorphism in both same- and opposite-sex targets. Participants were randomly assigned to either a pathogen prime or a control prime, and they completed instruments assessing individual differences in pathogen avoidance (disgust sensitivity and contamination sensitivity). Results revealed overall preferences for both facial symmetry and dimorphism. However, they did not reveal a relation between these preferences and disgust sensitivity or contamination sensitivity, nor did they reveal differences in these preferences across control and pathogen prime conditions. Null results of pathogen-avoidance variables were consistent across participant sex, target sex, and interactions between participant sex and target sex. Overall, findings cast doubt on the hypothesis that pathogen-avoidance motives influence preferences for facial symmetry or dimorphism.     

Cross-cultural variation in men's preference for sexual dimorphism in women's faces

Both attractiveness judgements and mate preferences vary considerably cross-culturally. We investigated whether men''s preference for femininity in women''s faces varies between 28 countries with diverse health conditions by analysing responses of 1972 heterosexual participants. Although men in all countries preferred feminized over masculinized female faces, we found substantial differences between countries in the magnitude of men''s preferences. Using an average femininity preference for each country, we found men''s facial femininity preferences correlated positively with the health of the nation, which explained 50.4% of the variation among countries. The weakest preferences for femininity were found in Nepal and strongest in Japan. As high femininity in women is associated with lower success in competition for resources and lower dominance, it is possible that in harsher environments, men prefer cues to resource holding potential over high fecundity.     

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.     

Environment contingent preferences: Exposure to visual cues of direct male–male competition and wealth increase women's preferences for masculinity in male faces

Previous studies show that parasite prevalence and mortality/health are related to cultural variation in women's preferences for attractive and masculine traits in men. Other studies have suggested that both male–male competition and wealth may also be important correlates of cross-cultural variation in women's masculinity preferences. Here we examined whether exposure to cues of direct male–male competition, violence, or wealth influenced women's face preferences. We showed women slideshows of images with cues of low and high direct male–male competition/violence or wealth and measured their visual preferences for masculine face traits. Recent visual experience changed women's preferences for facial masculinity, with women preferring more masculine male faces after exposure to images of men engaged in direct physical competition, images of weapons, or images depicting items of high monetary value. Recent visual experience had no significant effects on preferences for masculinity in same-sex faces. Given that high levels of direct physical competition and violence among males may increase the importance of direct intra-sexual competition, it may be adaptive for women to shift visual preferences in favor of males with face cues indicating physical strength and dominance over investment in such environments. Similarly, in wealthy environments investment may be less important than other aspects of quality and so it may be adaptive for women to shift visual preferences in favor of males with face cues indicating other aspects of quality over investment. Overall, our data demonstrate that preferences can be strategically flexible according to recent visual experience and support the notion of environment contingent preferences.     

Men's Preferences for Women's Femininity in Dynamic Cross-Modal Stimuli

Men generally prefer feminine women''s faces and voices over masculine women''s faces and voices, and these cross-modal preferences are positively correlated. Men''s preferences for female facial and vocal femininity have typically been investigated independently by presenting soundless still images separately from audio-only vocal recordings. For the first time ever, we presented men with short video clips in which dynamic faces and voices were simultaneously manipulated in femininity/masculinity. Men preferred feminine men''s faces over masculine men''s faces, and preferred masculine men''s voices over feminine men''s voices. We found that men preferred feminine women''s faces and voices over masculine women''s faces and voices. Men''s attractiveness ratings of both feminine and masculine faces were increased by the addition of vocal femininity. Also, men''s attractiveness ratings of feminine and masculine voices were increased by the addition of facial femininity present in the video. Men''s preferences for vocal and facial femininity were significantly and positively correlated when stimuli were female, but not when they were male. Our findings complement other evidence for cross-modal femininity preferences among male raters, and show 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.     

Neural activation in women in response to masculinized male faces: mediation by hormones and psychosexual factors

Women's preference for masculine faces varies with hormonal state, sociosexuality, and relationship status, but the underlying mechanisms are poorly understood. We hypothesized that hormones and psychosexual factors (sociosexuality, sexual inhibition/excitation) mediate the perception and evaluation of male faces thereby influencing women's preferences. We used functional magnetic resonance imaging to measure brain activity in 12 women as they evaluated pictures of male faces (half 30% masculinized, half 30% feminized). Participants were heterosexual women, age 23–28 years, who were not in a committed relationship and not using hormonal contraception. Women were tested during both the follicular and luteal phase of their menstrual cycle. We found five brain regions related to face and risk processing that responded more to the masculinized than to the feminized faces, including the superior temporal gyrus, precentral gyrus, posterior cingulate cortex, inferior parietal lobule, and anterior cingulate cortex. Increased activation in the anterior cingulate cortex, specifically, may indicate that women perceive masculinized faces to be both more risky and more attractive. We did not see any areas that were more strongly activated by feminized faces. Levels of activation were influenced by hormonal and psychosexual factors. The patterns of hormonally and psychosexually mediated neural activation observed may offer insight into the cognitive processes underlying women's partner preferences.     

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.     

Effects of exogenous testosterone and mating context on men's preferences for female facial femininity

Correlational research suggests that men show greater attraction to feminine female faces when their testosterone (T) levels are high. Men's preferences for feminine faces also seem to vary as a function of relationship context (short versus long-term). However, the relationship between T and preferences for female facial femininity has yet to be tested experimentally. In the current paper, we report the results of two experiments examining the causal role of T in modulating preferences for facial femininity across both short and long-term mating contexts. Results of Experiment 1 (within-subject design, n = 24) showed that participants significantly preferred feminized versus masculinized versions of women's faces. Further, participants showed a stronger preference for feminine faces in the short versus the long-term context after they received T, but not after they received placebo. Post-hoc analyses suggested that this effect was driven by a lower preference for feminine faces in the long-term context when on T relative to placebo, and this effect was found exclusively for men who received placebo on the first day of testing, and T on the second day of testing (i.e., Order x Drug x Mating context interaction). In Experiment 2 (between-subject design, n = 93), men demonstrated a significant preference for feminized female faces in the short versus the long-term context after T, but not after placebo administration. Collectively, these findings provide the first causal evidence that T modulates men's preferences for facial femininity as a function of mating context.     

Environmental threat influences preferences for sexual dimorphism in male and female faces but not voices or dances

During evolution, humans faced the trade-off between preferences for feminine and masculine traits which are presumably connected to parental care, and genetic quality or provisioning abilities, respectively. Recent research has shown that environmental factors influence preferences for femininity/masculinity in potential mates. However, studies mainly focus on women's preferences for isolated cues in men. We examined the influence of pathogen and resource threat on women's and men's preference for femininity/masculinity in opposite sex unmanipulated faces, voices, and dances. Three hundred seventy (206 women) students, aged 18–35 years, from universities across the state of Sao Paulo, Brazil, were primed with newspaper-like articles on either pathogen threat (Aedes aegypti, and its mosquito-borne diseases) or resource threat (=economic crisis), and compared to a control-priming condition (lions poisoned in a Kenyan Reserve). Participants were randomly assigned to rate attractiveness of the stimuli in one of the mating strategies (i.e., short-term or long-term relationship context). After each priming article, participants rated attractiveness of pre-rated masculine and feminine stimuli of the opposite sex in a standalone-rating design. The environmental threats and the control-priming conditions were shown in a random sequence. We found that the environmental threat and the mating strategies had no systematic effect on preferences for faces, voices and behavior, suggesting that different modalities can cue to different individual qualities and can be used differently according to the environmental context and relationship strategy. The environmental threat only had a weak effect on preferences for facial femininity-masculinity: women found feminine male faces more attractive under pathogen threat, while men found feminine female faces more attractive under resource threat. This is in contrast to some previous studies, and shows that effect of ecological factors on human psychology should be submitted to a more rigorous scrutiny. The mating strategies only interacted with voice attractiveness: women rated male voices as more attractive after being primed with any type of environmental threat when considering long-term relationship context, while in men this was true when considering short-term relationship. Finally, feminine dance videos were perceived as attractive under environmental threat, irrespective of sex, type of environmental threat or mating strategy.     

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.     

Cross-Cultural Agreement in Facial Attractiveness Preferences: The Role of Ethnicity and Gender

Previous work showed high agreement in facial attractiveness preferences within and across cultures. The aims of the current study were twofold. First, we tested cross-cultural agreement in the attractiveness judgements of White Scottish and Black South African students for own- and other-ethnicity faces. Results showed significant agreement between White Scottish and Black South African observers'' attractiveness judgements, providing further evidence of strong cross-cultural agreement in facial attractiveness preferences. Second, we tested whether cross-cultural agreement is influenced by the ethnicity and/or the gender of the target group. White Scottish and Black South African observers showed significantly higher agreement for Scottish than for African faces, presumably because both groups are familiar with White European facial features, but the Scottish group are less familiar with Black African facial features. Further work investigating this discordance in cross-cultural attractiveness preferences for African faces show that Black South African observers rely more heavily on colour cues when judging African female faces for attractiveness, while White Scottish observers rely more heavily on shape cues. Results also show higher cross-cultural agreement for female, compared to male faces, albeit not significantly higher. The findings shed new light on the factors that influence cross-cultural agreement in attractiveness preferences.     

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.     

Lack of support for relation between woman's masculinity preference,estradiol level and mating context

It has been proposed that women's preferences for male facial sexual dimorphism are positively correlated with conception probability and differ between short- and long-term mating contexts. In this study, we tested this assumption by analyzing relationships between estradiol levels to the women's preferences of male faces that were manipulated to vary in masculinity. Estradiol was measured in daily saliva samples throughout the entire menstrual cycle collected by Polish women with regular menstrual cycles. In our analyses, we included the three most commonly used definitions of the fertile window in the literature. After computing the overall masculinity preference of each participant and measuring hormone levels, we found that i) the timing of ovulation varied greatly among women (between − 11 and − 17 days from the onset of the next menses, counting backwards), ii) there was no relationship between daily, measured during the day of the test (N = 83) or average for the cycle (N = 115) estradiol levels and masculinity preferences, iii) there were no differences in masculinity preferences between women in low- and high-conception probability phases of the cycle, and iv) there were no differences in masculinity preferences between short- and long-term mating contexts. Our results do not support the idea that women's preferences for a potential sexual partner's facial masculinity fluctuate throughout the cycle.     

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.     

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.     

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

Many studies have shown that women's judgments of men's attractiveness are affected by changes in levels of sex hormones. However, no studies have tested for associations between changes in levels of sex hormones and men's judgments of women's attractiveness. To investigate this issue, we compared men's attractiveness judgments of feminized and masculinized women's and men's faces in test sessions where salivary testosterone was high and test sessions where salivary testosterone was relatively low. Men reported stronger attraction to femininity in women's faces in test sessions where salivary testosterone was high than in test sessions where salivary testosterone was low. This effect was found to be specific to judgments of opposite-sex faces. The strength of men's reported attraction to femininity in men's faces did not differ between high and low testosterone test sessions, suggesting that the effect of testosterone that we observed for judgments of women's faces was not due to a general response bias. Collectively, these findings suggest that changes in testosterone levels contribute to the strength of men's reported attraction to femininity in women's faces and complement previous findings showing that testosterone modulates men's interest in sexual stimuli.     

Facial attractiveness judgements reflect learning of parental age characteristics

Mate preferences are shaped by infant experience of parental characteristics in a wide variety of species. Similar processes in humans may lead to physical similarity between parents and mates, yet this possibility has received little attention. The age of parents is one salient physical characteristic that offspring may attend to. The current study used computer-graphic faces to examine how preferences for age in faces were influenced by parental age. We found that women born to 'old' parents (over 30) were less impressed by youth, and more attracted to age cues in male faces than women with 'young' parents (under 30). For men, preferences for female faces were influenced by their mother's age and not their father's age, but only for long-term relationships. These data indicate that judgements of facial attractiveness in humans reflect the learning of parental characteristics.     

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.     

Concordant preferences for opposite-sex signals? Human pheromones and facial characteristics.

We have investigated whether preferences for masculine and feminine characteristics are correlated across two modalities, olfaction and vision. In study 1, subjects rated the pleasantness of putative male (4,16-androstadien-3-one; 5alpha-androst-16-en-3-one) and female (1,3,5 (10),16-estratetraen-3-ol) pheromones, and chose the most attractive face shape from a masculine-feminine continuum for a long- and a short-term relationship. Study 2 replicated study 1 and further explored the effects of relationship context on pheromone ratings. For long-term relationships, women's preferences for masculine face shapes correlated with ratings of 4,16-androstadien-3-one and men's preferences for feminine face shapes correlated with ratings of 1,3,5(10),16-estratetraen-3-ol. These studies link sex-specific preferences for putative human sex pheromones and sexually dimorphic facial characteristics. Our findings suggest that putative sex pheromones and sexually dimorphic facial characteristics convey common information about the quality of potential mates.     

Holistic gaze strategy to categorize facial expression of varying intensities

Using faces representing exaggerated emotional expressions, recent behaviour and eye-tracking studies have suggested a dominant role of individual facial features in transmitting diagnostic cues for decoding facial expressions. Considering that in everyday life we frequently view low-intensity expressive faces in which local facial cues are more ambiguous, we probably need to combine expressive cues from more than one facial feature to reliably decode naturalistic facial affects. In this study we applied a morphing technique to systematically vary intensities of six basic facial expressions of emotion, and employed a self-paced expression categorization task to measure participants' categorization performance and associated gaze patterns. The analysis of pooled data from all expressions showed that increasing expression intensity would improve categorization accuracy, shorten reaction time and reduce number of fixations directed at faces. The proportion of fixations and viewing time directed at internal facial features (eyes, nose and mouth region), however, was not affected by varying levels of intensity. Further comparison between individual facial expressions revealed that although proportional gaze allocation at individual facial features was quantitatively modulated by the viewed expressions, the overall gaze distribution in face viewing was qualitatively similar across different facial expressions and different intensities. It seems that we adopt a holistic viewing strategy to extract expressive cues from all internal facial features in processing of naturalistic facial expressions.