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

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

Preferences for symmetry in human faces in two cultures: data from the UK and the Hadza, an isolated group of hunter-gatherers

Many studies show agreement within and between cultures for general judgements of facial attractiveness. Few studies, however, have examined the attractiveness of specific traits and few have examined preferences in hunter-gatherers. The current study examined preferences for symmetry in both the UK and the Hadza, a hunter-gatherer society of Tanzania. We found that symmetry was more attractive than asymmetry across both the cultures and was more strongly preferred by the Hadza than in the UK. The different ecological conditions may play a role in generating this difference. Such variation in preference may be adaptive if it reflects adaptation to local conditions. Symmetry is thought to indicate genetic quality, which may be more important among the Hadza with much higher mortality rates from birth onwards. Hadza men who were more often named as good hunters placed a greater value on symmetry in female faces. These results suggest that high quality Hadza men are more discriminating in their choice of faces. Hadza women had increased preferences for symmetry in men's faces when they were pregnant or nursing, perhaps due to their increased discrimination and sensitivity to foods and disease harmful to a foetus or nursing infant. These results imply that symmetry is an evolutionarily relevant trait and that variation in symmetry preference appears strategic both between cultures and within individuals of a single culture.     

Symmetry and Human Facial Attractiveness

Symmetry may act as a marker of phenotypic and genetic quality and is preferred during mate selection in a variety of species. Measures of human body symmetry correlate with attractiveness, but studies manipulating human face images report a preference for asymmetry. These results may reflect unnatural feature shapes and changes in skin textures introduced by image processing. When the shape of facial features is varied (with skin textures held constant), increasing symmetry of face shape increases ratings of attractiveness for both male and female faces. These findings imply facial symmetry may have a positive impact on mate selection in humans.     

The Scent of Symmetry: A Human Sex Pheromone that Signals Fitness?

A previous study by the authors showed that the body scent of men who have greater body bilateral symmetry is rated as more attractive by normally ovulating (non-pill-using) women during the period of highest fertility based on day within the menstrual cycle. Women in low-fertility phases of the cycle and women using hormone-based contraceptives do not show this pattern. The current study replicated these findings with a larger sample and statistically controlled for men's hygiene and other factors that were not controlled in the first study. The current study also examined women's scent attractiveness to men and found no evidence that men prefer the scent of symmetric women. We propose that the scent of symmetry is an honest signal of phenotypic and genetic quality in the human male, and chemical candidates are discussed. In both sexes, facial attractiveness (as judged from photos) appears to predict body scent attractiveness to the opposite sex. Women's preference for the scent associated with men's facial attractiveness is greatest when their fertility is highest across the menstrual cycle. The results overall suggest that women have an evolved preference for sires with good genes.     

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.     

Symmetry, sexual dimorphism in facial proportions and male facial attractiveness

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

Face perception is modulated by sexual preference

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

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.     

Life History of Female Preferences for Male Faces

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

Facial attractiveness, symmetry and cues of good genes.

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

Does attractiveness in men provide clues to semen quality?

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

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.     

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.     

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.     

Do women's preferences for symmetry change across the menstrual cycle?

In many species, symmetry enhances physical attractiveness of the face and body. In humans, facial attractiveness is also enhanced by symmetrical decoration in the form of facial paint [Cárdenas, R. A., & Harris, L. J. (2006)]. According to the good-genes hypothesis [e.g., Thornhill, R., & Gangestad, S. W. (1999)], symmetry is preferred because it is associated with mate quality. According to the receiver bias hypothesis [e.g., Enquist, M., & Johnstone, R. (1997)], it is a by-product of how the visual system is designed. Proponents of the good-genes hypothesis have suggested that a preference for symmetry may vary with fertility, namely, that it will be enhanced in women in the high-fertility phase of the menstrual cycle. Previous research does demonstrate that, during this phase, women prefer the scent of more symmetrical men [e.g., Gangestad, S. W., & Thornhill, R. (1998)]. However, research employing assessment of faces fails to find a similar effect [Koehler, N., Rhodes, G., & Simmons, L. W. (2002)]. Previous research asked subjects to judge faces one at a time during high fertility (around ovulation) and low fertility (menstruation). We used a different face-presentation method, tested women during the other low-fertility (midluteal) phase, and used decorated as well as undecorated faces. As in our prior study [Cárdenas, R. A., & Harris, L. J. (2006)], symmetry of facial features and symmetry of decoration enhanced attractiveness, but, contrary to the possible prediction of the good-genes hypothesis, the effects did not vary across the cycle. The results as they are, therefore, can be equally accommodated by both hypotheses.     

Social transmission of face preferences among humans

Previous studies demonstrating mate choice copying effects among females in non-human species have led many researchers to propose that social transmission of mate preferences may influence sexual selection for male traits. Although it has been suggested that social transmission may also influence mate preferences in humans, there is little empirical support for such effects. Here, we show that observing other women with smiling (i.e. positive) expressions looking at male faces increased women's preferences for those men to a greater extent than did observing women with neutral (i.e. relatively negative) expressions looking at male faces. By contrast, the reverse was true for male participants (i.e. observing women with neutral expressions looking at male faces increased male participant's preferences for those men to a greater extent than did observing women smiling at male faces). This latter finding suggests that within-sex competition promotes negative attitudes among men towards other men who are the target of positive social interest from women. Our findings demonstrate that social transmission of face preferences influences judgments of men's attractiveness, potentially demonstrating a mechanism for social transmission of mate preferences.     

The Body and Face of Woman: One Ornament that Signals Quality?

Evidence has accumulated in recent years supporting the hypothesis that both facial and bodily physical attractiveness in humans are certifications of developmental and hormonal health. Such evidence indicates that physical attractiveness is an honest or Zahavian signal of phenotypic and genetic quality. The hypothesis that physical beauty connotes health was first proposed by Westermarck and was discussed later by Ellis and Symons. It has been suggested that facial attractiveness in women is a deceptive signal of youth, unrelated to phenotypic and genetic quality. This sensory-bias or super-stimulus hypothesis is not supported by this study of men’s ratings of the attractiveness of photographs of 92 nude women. Independent ratings of photographs of faces, fronts with faces covered, and backs of the same women are significantly, positively correlated. The correlation between the ratings of different photos implies that women’s faces and external bodies comprise a single ornament of honest mate value, apparently constructed during puberty by estrogen and also probably by developmental adaptations for symmetry. Thus, women’s physical attractiveness in face and body honestly signal hormonal and perhaps developmental health.     

Symmetrical decorations enhance the attractiveness of faces and abstract designs

In humans and several other species, face and body symmetry have been found to enhance physical attractiveness. A proposed explanation is that symmetry is a phenotypic indicator of biological fitness. Throughout the world, symmetrical designs also are a common feature in face and body painting and the decorative arts. The implication is that symmetrical designs might provide an additional way to enhance physical attractiveness. To find out, we conducted three experiments, two with human faces and one with abstract or nonrepresentational designs. In Experiments 1 and 2, we showed undergraduate students photographs of pairs of faces and instructed them to choose the more attractive face in each pair. The photographs were of physically symmetrical and asymmetrical faces (as indexed by facial features) that had been decorated with either symmetrical or asymmetrical designs of the kind used in many preindustrial societies. As indexed by the number of times they were chosen, symmetrical faces were judged to be more attractive than asymmetrical faces; adding asymmetrical designs to symmetrical faces decreased their attractiveness; and adding symmetrical designs to asymmetrical faces increased their attractiveness. In Experiment 3, undergraduates made similar choices from pairs of abstract designs taken from several cultures and modified in shape, coloration, and orientation of design features. Symmetrical designs again were judged to be more attractive, with shape and coloration playing the more important roles. We interpret the results as suggesting that the same mechanisms underlying the judgment of physical attractiveness also underlie cultural practices of face painting and abstract art.     

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.     

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

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