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Naked group girls spread ass milf. Black white dp. Horny mature women porn. Ibong adarna tagalog version comic strip. Bbw eden mor. If the address matches an existing account you will receive an email with instructions to reset your password. If the address matches an existing account you will receive an email with instructions to retrieve your username. Google Scholar. Find this author on PubMed. Search for more papers by this author. Face preferences affect a diverse range of critical social outcomes, from mate choices and decisions about platonic relationships to hiring decisions and decisions about social exchange. Firstly, we review the facial characteristics that influence attractiveness judgements of faces e. The research relating Facial attractiveness milffaceness these issues highlights flexible, sophisticated systems that article source and promote adaptive responses to faces that appear to function Facial attractiveness milffaceness maximize the benefits of both our mate choices and more general decisions about other types of social partners. Our magazines and television screens are not just filled with Facial attractiveness milffaceness faces—they are filled with attractive faces, and both Facial attractiveness milffaceness and men are highly concerned with here looks in a potential Facial attractiveness milffaceness [ 1 ]. Physical appearance is important to humans and certain features appear to be found attractive across individuals and cultures [ 2 ]. The same holds true across the animal kingdom; most non-human species rely on external traits, such as the size, shape and colour of adornments e. Enchanting hot lesbians in pantyhose Hookers in Rijeka.

Free porn movies tivia. Red Facial attractiveness milffaceness generally seem to have aversive effects on human behaviour. For example, when taking exams, individuals move their body away from tests with red covers more than they do from those with green or grey covers [ ]. While these studies suggest the Facial attractiveness milffaceness red may be seen as a threatening stimulus in humans, red also appears to enhance attraction in some instances.

For Facial attractiveness milffaceness, women are seen as more attractive by men when presented with red backgrounds or with red clothing, relative to other colours [ ]. This effect appears to be specific to attractiveness judgements; red colour does not influence judgements of Facial attractiveness milffaceness traits such as kindness or intelligence and does not influence women's attractiveness judgements of other women [ ].

Further research has examined red coloration in faces and demonstrated a positive association with perceived health [ ]. The authors suggest that perception of healthy, oxygenated blood may drive associations between red and healthiness. Alongside redness, people also appear to think that skin yellowness is associated with healthy appearance in faces [ ].

Yellowness may advertise health Facial attractiveness milffaceness an association with diet, as carotenoids are associated with skin yellowness and Facial attractiveness milffaceness absorbed via the intake of fruit and vegetables [ ]. In a classic social psychology study, Dion et al.

For example, attractive individuals were thought to be able to achieve more prestigious occupations, be more competent spouses with happier marriages and have better prospects for personal fulfilment. There has been a wealth of studies examining this attractiveness stereotype, demonstrating that attractive people are seen in a positive light for a wide range of attributes compared with unattractive people.

Studies on attractiveness stereotypes have generally not addressed the particular characteristics of faces that make learn more here either attractive or unattractive, or the features that elicit personality attributions, although different faces reliably elicit the same personality attributions Facial attractiveness milffaceness ].

Expression certainly has large effects, with, for example, faces shown with smiles rated as more attractive Facial attractiveness milffaceness as having more positive personality traits than neutral faces e. Such facial expressions are transient, however, and will differ rapidly within individuals over time and across photographs. Despite some findings showing a preference for more masculine and dominant male faces e. Personality traits are reported cross-culturally to be among the most important factors in partner choice by both sexes [ 1].

If desired personality is so important, it would appear likely that personality attributions elicited by a face would Facial attractiveness milffaceness its attractiveness. For example, women who value cooperation and good parenting may avoid masculine-faced men. Facial attractiveness milffaceness, instead of feminine faces being attractive and this attractiveness driving positive personality attributions, it may be that the personality attributions are Facial attractiveness milffaceness the attractiveness judgements.

Individuals may use personality stereotypes in mate selection to select partners with a personality that they desire. Some perceptual attributions to facial photographs are somewhat accurate e. Attraction to faces based on personality stereotypes may happen regardless of whether attributions Women sex doctor accurate or not, especially as many individuals do believe that face provides an important guide to character [].

One study has indeed demonstrated that a desire for some personality traits influences judgements of facial attractiveness [ ]. Individuals valuing particular personality Facial attractiveness milffaceness find faces appearing to display these traits attractive. Conversely, those not valuing particular traits find faces attractive that are perceived to possess that trait less. Thus, desired personality influences perceptions of facial attractiveness in opposite sex faces, changing the result to: In terms of benefits to perceivers, it is easy to see why traits such as appearing trustworthy would make a face appear more attractive.

For individual-specific Click, the logic is more complicated, but such Facial attractiveness milffaceness could be related to behavioural compatibility within couples, as people do tend to desire partners with personalities similar to their own [ ]. One reason for variability source preferences for male facial masculinity may lie in the personality traits that masculine- and feminine-faced men are assumed to possess.

Increasing the masculinity of face shape increased perceptions of dominance, masculinity and age but decreased perceptions of warmth, emotionality, honesty, cooperativeness and quality as a parent [ 83 ]. Indeed, recent work has shown that masculine facial characteristics are associated with indices of physical dominance, such as physical Facial attractiveness milffaceness [ ], and the perception of such traits [ ], and that feminine men show weaker preferences for short-term relationships and stronger preferences for committed, long-term relationships than their masculine peers do [ ].

Women's face preferences may thus represent a trade-off between the desire for good genes and Facial attractiveness milffaceness desire for a cooperative partner.

Of course, the five types of trait listed above are not a complete list of factors involved in the judgement of facial attractiveness. While individual traits impact on attractiveness, there is also scope for interaction Facial attractiveness milffaceness them.

Certain face traits also appear to interact in generating preferences, however. For example, preferences for masculinity vary as a function of the healthiness of the face [ 96 ] and women's preferences for facial self-similarity are higher when men are more facially masculine [ ]. Such interactions highlight that facial attractiveness judgements are not simple: In humans, while individuals may share certain basic criteria for finding faces attractive, many factors Facial attractiveness milffaceness influence the specific types of face they find attractive.

In this section, we review three broad areas leading to individual differences in preferences: Research suggests that internal factors predict individual differences in several aspects of face perception, including attractiveness judgements.

Importantly, the nature of these individual differences suggests adaptive design in face perception and face preferences. In the following section, we discuss two broad types of internal factors: The influence of hormones on face perception is an Facial attractiveness milffaceness that has generated a considerable amount of empirical research in recent years.

As detailed previously, masculine characteristics in men's faces are associated with measures of long-term medical health [ 3577 ] and indices of developmental stability [ 3637 ], physical strength [ ] and Facial attractiveness milffaceness potential Facial attractiveness milffaceness ].

By contrast, feminine characteristics in men's faces are associated with cues of investment and stronger preferences for long-term over short-term sexual relationships e.

There is now compelling evidence that how women article source this trade off between the costs and benefits associated with choosing a masculine mate is affected by hormone levels and fertility.

Many studies have reported that women demonstrate stronger preferences for men displaying masculine facial characteristics around ovulation, when women are most fertile, than during other phases of the menstrual cycle [ — Facial attractiveness milffaceness.

Some studies have also reported that these cyclic shifts in women's preferences for masculine characteristics in men's faces are greatest among women who already have romantic partners and when women judge men's attractiveness for short-term, extra-pair relationships [ ].

Although the ultimate function of these cyclic shifts remains somewhat controversial, many researchers have interpreted cyclic shifts in women's masculinity preferences as evidence for adaptations that function to increase offspring health via high paternal investment from a long-term partner while promoting attraction to other men displaying cues of heritable immunity to infectious disease when most Facial attractiveness milffaceness discussed in [ ].

Women may gain maximal benefits by selecting investing long-term partners and high-quality extra-pair partners. Importantly, other explanations that have been suggested, such as increased attraction to individuals who appear to be likely sources of high-quality Facial attractiveness milffaceness and support during phases of the menstrual cycle when increased progesterone prepares the body for pregnancy i.

Increased attraction to masculine men is by no means unique to face preferences; women also demonstrate stronger attraction to masculine men when judging the attractiveness of men's voices [ — ], body shapes [ ] and body odours [ ], as well as when judging the attractiveness of videoclips of male behavioural displays of dominance [].

Furthermore, converging evidence for fertility-related variation in women's preferences for facial masculinity comes from studies investigating circum-pubertal and circum-menopausal variation in women's masculinity preferences; post-menopausal and pre-pubertal women report weaker preferences for masculine facial characteristics than do their pre-menopausal and post-pubertal counterparts, respectively e. The ultimate function Facial attractiveness milffaceness cyclic shifts in women's preferences for masculine facial characteristics is not the only controversial aspect of cyclic shifts Facial attractiveness milffaceness women's masculinity preferences.

For example, although some researchers have suggested that cyclic shifts in women's masculinity preferences may be an artefact of the computer graphic methods that are generally used in these studies to experimentally manipulate sexually dimorphic cues Facial attractiveness milffaceness digital face images [ ], this claim is very difficult to reconcile with findings from studies that have demonstrated cyclic shifts in women's preferences for masculinity in real i.

While these findings suggest that cyclic shifts in women's masculinity preferences are not an artefact of the stimuli used, an aspect of research on cyclic shifts in women's masculinity preferences that remains controversial is whether the effect of cycle phase on women's Facial attractiveness milffaceness preferences is relatively specific to judgements of men's faces, or also occurs when women judge the attractiveness of other women. To date, evidence is equivocal; some studies have observed cyclic shifts in women's preferences for masculine-faced men, but not masculine-faced women [ ], while others have observed cyclic shifts in women's preferences for masculine faces, irrespective of their sex [].

These latter papers speculate that cyclic shifts in women's preferences for masculine-faced women could represent a low-cost functionless Facial attractiveness milffaceness of a mechanism that evolved primarily to increase women's preferences for masculine men around ovulation [ ], or have suggested that higher attractiveness ratings given Facial attractiveness milffaceness masculine women around ovulation could reflect increased derogation of feminine, and therefore attractive, same-sex competitors when women are most fertile [ ] see also [ ].

In addition to the sex-specificity of the effects of cycle phase on face preferences, the mechanisms that underpin cyclic shifts in women's preferences for masculine characteristics in men's faces have also been a topic of considerable interest in recent years. For example, research into the hormonal mechanisms that might underpin these cyclic shifts has variously emphasized the effects of variation in levels of testosterone [ ], oestrogen [ ] and progesterone [], or has suggested, perhaps unsurprisingly, that cyclic shifts in women's masculinity preferences might be best explained by complex interactions among multiple hormones [].

While findings from research into the hormonal mechanisms that might underpin cyclic shifts in women's masculinity preferences have arguably been inconsistent, the findings of corresponding research into the psychological mechanisms have been relatively consistent; various studies have demonstrated that women are quicker to categorize men and access male stereotypes around ovulation e. These findings suggest that cyclic variations in stereotype access and sexual desire might be important psychological mechanisms for regulating facial masculinity preferences during the menstrual cycle.

While research on hormone-mediated face perception has generally focused on women's judgements of men's Facial attractiveness milffaceness, some recent research here investigated hormone-mediated face preferences among men.

Men, of course, do not cycle in the same way women do, but levels of testosterone fluctuate within individuals. Research using natural variation Facial attractiveness milffaceness testosterone has shown that men's preferences for feminine characteristics here women's faces are stronger when their testosterone levels are high than when they are relatively low [ ].

This finding suggests that hormones, such as testosterone, can generate within-participant individual differences in face preference in men. As can be seen from the previous paragraphs, there is compelling evidence that women's preferences for masculine men, Facial attractiveness milffaceness they assessed from face preferences or from preferences for male characteristics in other domains, vary systematically over the menstrual cycle. Whether or not preferences for other putative cues of men's long-term health are similarly affected by menstrual cycle is equivocal, however.

For example, although many studies Facial attractiveness milffaceness demonstrated that women's preferences for the body odours of symmetric men are enhanced around ovulation reviewed in [ ]evidence for cyclic shifts in women's preferences for symmetry in men's faces is inconsistent.

Xxxxxxxxx Hot Watch Video Barzli Xxxx. Such findings demonstrate the importance of health perceptions for social interaction generally. Again, as for previous traits, there may be both direct and indirect benefits to partnering with individuals who are perceived to be healthy. Facial healthiness. High healthiness is associated with higher ratings of attractiveness. There have been several studies that have addressed how facial appearance relates to the healthiness of an individual in humans. The three traits discussed above are often manipulated by changing only face shape, but health perception appears to be related to facial colour and texture also. Fewer studies have examined how colour and texture of faces influence attractiveness judgements. One study has examined how well ratings of health from small patches of skin of faces are related to overall rated attractiveness when the whole face image is available. Jones et al. In other research, homogeneity of skin colour was positively related to attractiveness [ 99 ]. Findings have also suggested that more heterozygous men also have healthier appearing skin [ 56 ]. Skin health may be a particularly useful marker of current health condition as it is more changeable than aspects such as symmetry or averageness. Coloration is directly related to the appearance of skin. Coloration also appears to be an important component of sexual selection in many species. Red coloration is associated with dominance in fish [ ], birds [ ] and non-human primates [ , ] and, consequently, is linked to attracting the opposite sex. It has been noted that primates with trichromatic vision are generally bare-faced [ ] and that, at least in humans, facial flushing is associated with anger and confrontation [ ]. In research on non-human primates, there has been much interest in colour. For example, experimental manipulation of colour shows that female rhesus macaques prefer images of redder male faces [ ], while males prefer images of redder female hindquarters [ ]. In mandrills, red facial colour is related to rank in males [ ], and females sexually present more frequently to brighter males and also groom them more frequently [ ]. Red coloration also has consequences for behaviour in other species. For example, in bird species, the addition of red to stimuli can increase social dominance [ ]. In humans, it has been shown that wearing red in a variety of physically competitive sports is associated with an increased chance of winning over opponents [ ]. This has been interpreted as natural associations of red with dominance being extended to artificially displayed red in the same way that artificial stimuli can exploit innate responses to natural stimuli [ , ]. One study pitting red versus blue shapes found that red shapes were seen as more aggressive, dominant and more likely to win in physical competitions [ ]. Red does generally seem to have aversive effects on human behaviour. For example, when taking exams, individuals move their body away from tests with red covers more than they do from those with green or grey covers [ ]. While these studies suggest the colour red may be seen as a threatening stimulus in humans, red also appears to enhance attraction in some instances. For example, women are seen as more attractive by men when presented with red backgrounds or with red clothing, relative to other colours [ ]. This effect appears to be specific to attractiveness judgements; red colour does not influence judgements of other traits such as kindness or intelligence and does not influence women's attractiveness judgements of other women [ ]. Further research has examined red coloration in faces and demonstrated a positive association with perceived health [ ]. The authors suggest that perception of healthy, oxygenated blood may drive associations between red and healthiness. Alongside redness, people also appear to think that skin yellowness is associated with healthy appearance in faces [ ]. Yellowness may advertise health via an association with diet, as carotenoids are associated with skin yellowness and are absorbed via the intake of fruit and vegetables [ ]. In a classic social psychology study, Dion et al. For example, attractive individuals were thought to be able to achieve more prestigious occupations, be more competent spouses with happier marriages and have better prospects for personal fulfilment. There has been a wealth of studies examining this attractiveness stereotype, demonstrating that attractive people are seen in a positive light for a wide range of attributes compared with unattractive people. Studies on attractiveness stereotypes have generally not addressed the particular characteristics of faces that make individuals either attractive or unattractive, or the features that elicit personality attributions, although different faces reliably elicit the same personality attributions [ ]. Expression certainly has large effects, with, for example, faces shown with smiles rated as more attractive and as having more positive personality traits than neutral faces e. Such facial expressions are transient, however, and will differ rapidly within individuals over time and across photographs. Despite some findings showing a preference for more masculine and dominant male faces e. Personality traits are reported cross-culturally to be among the most important factors in partner choice by both sexes [ 1 , ]. If desired personality is so important, it would appear likely that personality attributions elicited by a face would affect its attractiveness. For example, women who value cooperation and good parenting may avoid masculine-faced men. Thus, instead of feminine faces being attractive and this attractiveness driving positive personality attributions, it may be that the personality attributions are driving the attractiveness judgements. Individuals may use personality stereotypes in mate selection to select partners with a personality that they desire. Some perceptual attributions to facial photographs are somewhat accurate e. Attraction to faces based on personality stereotypes may happen regardless of whether attributions are accurate or not, especially as many individuals do believe that face provides an important guide to character [ , ]. One study has indeed demonstrated that a desire for some personality traits influences judgements of facial attractiveness [ ]. Individuals valuing particular personality traits find faces appearing to display these traits attractive. Conversely, those not valuing particular traits find faces attractive that are perceived to possess that trait less. Thus, desired personality influences perceptions of facial attractiveness in opposite sex faces, changing the result to: In terms of benefits to perceivers, it is easy to see why traits such as appearing trustworthy would make a face appear more attractive. For individual-specific traits, the logic is more complicated, but such preferences could be related to behavioural compatibility within couples, as people do tend to desire partners with personalities similar to their own [ ]. One reason for variability in preferences for male facial masculinity may lie in the personality traits that masculine- and feminine-faced men are assumed to possess. Increasing the masculinity of face shape increased perceptions of dominance, masculinity and age but decreased perceptions of warmth, emotionality, honesty, cooperativeness and quality as a parent [ 83 ]. Indeed, recent work has shown that masculine facial characteristics are associated with indices of physical dominance, such as physical strength [ ], and the perception of such traits [ ], and that feminine men show weaker preferences for short-term relationships and stronger preferences for committed, long-term relationships than their masculine peers do [ ]. Women's face preferences may thus represent a trade-off between the desire for good genes and the desire for a cooperative partner. Of course, the five types of trait listed above are not a complete list of factors involved in the judgement of facial attractiveness. While individual traits impact on attractiveness, there is also scope for interaction between them. Certain face traits also appear to interact in generating preferences, however. For example, preferences for masculinity vary as a function of the healthiness of the face [ 96 ] and women's preferences for facial self-similarity are higher when men are more facially masculine [ ]. Such interactions highlight that facial attractiveness judgements are not simple: In humans, while individuals may share certain basic criteria for finding faces attractive, many factors may influence the specific types of face they find attractive. In this section, we review three broad areas leading to individual differences in preferences: Research suggests that internal factors predict individual differences in several aspects of face perception, including attractiveness judgements. Importantly, the nature of these individual differences suggests adaptive design in face perception and face preferences. In the following section, we discuss two broad types of internal factors: The influence of hormones on face perception is an area that has generated a considerable amount of empirical research in recent years. As detailed previously, masculine characteristics in men's faces are associated with measures of long-term medical health [ 35 , 77 ] and indices of developmental stability [ 36 , 37 ], physical strength [ ] and reproductive potential [ ]. By contrast, feminine characteristics in men's faces are associated with cues of investment and stronger preferences for long-term over short-term sexual relationships e. There is now compelling evidence that how women resolve this trade off between the costs and benefits associated with choosing a masculine mate is affected by hormone levels and fertility. Many studies have reported that women demonstrate stronger preferences for men displaying masculine facial characteristics around ovulation, when women are most fertile, than during other phases of the menstrual cycle [ — ]. Some studies have also reported that these cyclic shifts in women's preferences for masculine characteristics in men's faces are greatest among women who already have romantic partners and when women judge men's attractiveness for short-term, extra-pair relationships [ ]. Although the ultimate function of these cyclic shifts remains somewhat controversial, many researchers have interpreted cyclic shifts in women's masculinity preferences as evidence for adaptations that function to increase offspring health via high paternal investment from a long-term partner while promoting attraction to other men displaying cues of heritable immunity to infectious disease when most fertile discussed in [ ]. Women may gain maximal benefits by selecting investing long-term partners and high-quality extra-pair partners. Importantly, other explanations that have been suggested, such as increased attraction to individuals who appear to be likely sources of high-quality care and support during phases of the menstrual cycle when increased progesterone prepares the body for pregnancy i. Increased attraction to masculine men is by no means unique to face preferences; women also demonstrate stronger attraction to masculine men when judging the attractiveness of men's voices [ — ], body shapes [ ] and body odours [ ], as well as when judging the attractiveness of videoclips of male behavioural displays of dominance [ , ]. Furthermore, converging evidence for fertility-related variation in women's preferences for facial masculinity comes from studies investigating circum-pubertal and circum-menopausal variation in women's masculinity preferences; post-menopausal and pre-pubertal women report weaker preferences for masculine facial characteristics than do their pre-menopausal and post-pubertal counterparts, respectively e. The ultimate function of cyclic shifts in women's preferences for masculine facial characteristics is not the only controversial aspect of cyclic shifts in women's masculinity preferences. For example, although some researchers have suggested that cyclic shifts in women's masculinity preferences may be an artefact of the computer graphic methods that are generally used in these studies to experimentally manipulate sexually dimorphic cues in digital face images [ ], this claim is very difficult to reconcile with findings from studies that have demonstrated cyclic shifts in women's preferences for masculinity in real i. While these findings suggest that cyclic shifts in women's masculinity preferences are not an artefact of the stimuli used, an aspect of research on cyclic shifts in women's masculinity preferences that remains controversial is whether the effect of cycle phase on women's face preferences is relatively specific to judgements of men's faces, or also occurs when women judge the attractiveness of other women. To date, evidence is equivocal; some studies have observed cyclic shifts in women's preferences for masculine-faced men, but not masculine-faced women [ ], while others have observed cyclic shifts in women's preferences for masculine faces, irrespective of their sex [ , ]. These latter papers speculate that cyclic shifts in women's preferences for masculine-faced women could represent a low-cost functionless by-product of a mechanism that evolved primarily to increase women's preferences for masculine men around ovulation [ ], or have suggested that higher attractiveness ratings given to masculine women around ovulation could reflect increased derogation of feminine, and therefore attractive, same-sex competitors when women are most fertile [ ] see also [ ]. In addition to the sex-specificity of the effects of cycle phase on face preferences, the mechanisms that underpin cyclic shifts in women's preferences for masculine characteristics in men's faces have also been a topic of considerable interest in recent years. For example, research into the hormonal mechanisms that might underpin these cyclic shifts has variously emphasized the effects of variation in levels of testosterone [ ], oestrogen [ ] and progesterone [ , ], or has suggested, perhaps unsurprisingly, that cyclic shifts in women's masculinity preferences might be best explained by complex interactions among multiple hormones [ , ]. While findings from research into the hormonal mechanisms that might underpin cyclic shifts in women's masculinity preferences have arguably been inconsistent, the findings of corresponding research into the psychological mechanisms have been relatively consistent; various studies have demonstrated that women are quicker to categorize men and access male stereotypes around ovulation e. These findings suggest that cyclic variations in stereotype access and sexual desire might be important psychological mechanisms for regulating facial masculinity preferences during the menstrual cycle. While research on hormone-mediated face perception has generally focused on women's judgements of men's attractiveness, some recent research has investigated hormone-mediated face preferences among men. Men, of course, do not cycle in the same way women do, but levels of testosterone fluctuate within individuals. Research using natural variation in testosterone has shown that men's preferences for feminine characteristics in women's faces are stronger when their testosterone levels are high than when they are relatively low [ ]. This finding suggests that hormones, such as testosterone, can generate within-participant individual differences in face preference in men. As can be seen from the previous paragraphs, there is compelling evidence that women's preferences for masculine men, be they assessed from face preferences or from preferences for male characteristics in other domains, vary systematically over the menstrual cycle. Whether or not preferences for other putative cues of men's long-term health are similarly affected by menstrual cycle is equivocal, however. For example, although many studies have demonstrated that women's preferences for the body odours of symmetric men are enhanced around ovulation reviewed in [ ] , evidence for cyclic shifts in women's preferences for symmetry in men's faces is inconsistent. One study has found that women's preferences for symmetric male faces were stronger around ovulation than during other phases of the menstrual cycle, at least among partnered women who were instructed to judge men's attractiveness as short-term mates [ ]. By contrast, other studies have observed no evidence for cyclic shifts in women's preferences for symmetric men's faces e. Although evidence that women's preferences for symmetry in men's faces vary systematically over the menstrual cycle is equivocal, that is not to say that robust cyclic shifts in women's perceptions of faces are only evident in their preferences for facial masculinity. For example, women's aversions to self-resembling faces are enhanced around ovulation and positively correlated with women's estimated progesterone levels during the menstrual cycle [ ]. This variation in attitudes to self-resembling faces may reflect increased inbreeding avoidance around ovulation and increased preferences for caring, supportive and trustworthy individuals when increased progesterone prepares the body for pregnancy [ ]. Moreover, women's aversions to facial cues associated with current illness e. Indeed, pregnant women and women using oral contraceptives which mimic the effects of increased progesterone during pregnancy demonstrate stronger aversions to individuals displaying facial cues of illness than do women with natural menstrual cycles [ 97 ]. These latter findings for aversions to facial cues of illness and progesterone during the menstrual cycle complement other research on increased aversions to possible sources of contagion in women's food preferences during pregnancy [ ], as well as increased sensitivity to facial expressions signalling that sources of threat and contagion are nearby when progesterone levels are raised [ , ]. While our discussion of hormone-mediated face preferences in women has emphasized the positive findings that have been reported in the literature, it is important to note that there have also been unsuccessful replications of cyclic variation in women's face preferences. For example, two recent studies observed no evidence for cyclic variations in women's preferences for masculine versus feminine male faces [ , ]. One possible explanation of these null findings comes from findings that suggest the extent to which women's preferences for masculine men vary over the menstrual cycle vary systematically among women. For example, cyclic variation in women's preferences for masculine characteristics in men's voices is significantly greater among women with high trait i. This pattern of results may occur because varying their sexual strategy during the menstrual cycle may benefit unattractive women more than it benefits attractive women [ ]. More recent research has presented additional evidence that women's family background, prenatal hormone levels and mortality salience might also affect the extent to which they vary their masculinity preferences according to their menstrual cycle phase [ — ]. We also note that there are significant methodological differences between studies examining cycle effects, making direct comparisons e. For example, some studies distinguish between short- and long-term mating contexts, generally with larger cyclic shifts for short-term judgements [ ], while others do not [ ]. Studies also differ in stimuli number, stimuli type and how fertility is defined. A thorough description of methodological differences between studies is not the focus here, but methodology is certainly a factor that could explain differences in findings across studies. It is likely that further research concerning individual differences in cyclic shifts and comparing different methodologies would provide important insights into the motivations, functions and mechanisms behind cyclic shifts in fundamental aspects of face perception. While the previous section discussed research implicating hormone levels and fertility in individual differences in face perception, this section will discuss the relationships between face preferences and indices of own condition and attractiveness. Several studies have reported positive correlations between women's ratings of their own physical attractiveness and the strength of their preferences for masculine characteristics in men's faces [ 92 ]. Other studies have extended this work by demonstrating that more objective measures of women's condition and attractiveness, such as their waist—hip ratio or oestrogen levels, predict their preferences for masculine characteristics in men's faces in the same way [ , ]. Similar correlations between indices of women's own attractiveness and the strength of their preferences for masculine characteristics in other domains, such as men's voices, have also been reported [ , ], and indices of women's own condition and attractiveness are positively correlated with the strength of their preferences for symmetry and healthy-looking skin in men's faces [ 92 , ]. The findings described above appear to be somewhat analogous to condition-dependent preferences observed in other species, in which individuals in good physical condition show stronger preferences for high-quality mates e. Condition-dependent preferences in both humans and non-humans may have a common function and occur because individuals in good physical condition i. Particularly compelling evidence for this proposal comes from one of the few experimental studies of condition-dependent mate preferences. These findings suggest that women recalibrate subjective impressions of their own attractiveness i. While the research described above focused on the relationships between mate preferences and both individuals' own physical characteristics and their subjective evaluations of these physical characteristics, other work on condition-dependent preferences has investigated whether personality traits and other psychological factors predict individual differences in mate preferences in similar ways. For example, individual differences in systemizing and sensation-seeking, both of which are components of male sex-typical psychology, are positively correlated with men's preferences for feminine characteristics in women's, but not men's, faces [ , ]. Among women, individual differences in empathy, a component of female sex-typical psychology, and extraversion, a key predictor of social status that is correlated with women's physical attractiveness, are positively correlated with preferences for masculine characteristics in men's, but not women's, faces [ , ]. These findings not only implicate personality traits in individual differences in face preferences but also raise the intriguing possibility that some personality traits might mediate the relationships between an individual's physical characteristics and their face preferences. While factors such as hormones and own attractiveness can explain differences in face preferences between individuals, the context under which judgements are made can also contribute to variation in standards of beauty. In the following section, we discuss how context affects face preferences in three types of contexts: Information about genetic kinship is available in the face and is perceived somewhat accurately [ — ]. Judgements of facial similarity are highly synonymous with judgements of kinship [ — ], and facial similarity produced by computer-graphic manipulation affects behaviour in ways consistent with inclusive fitness theory e. Therefore, responses to facial resemblance are likely to be affected by prosocial versus sexual contexts. Cues of kinship are predicted to increase preferences in non-sexual, prosocial contexts, owing to the benefits associated with inclusive fitness [ ]. In other words, evolutionary models show that behaviours that benefit other individuals who share genes through common descent will be favoured. Therefore, if physical similarity is a reliable cue of genetic relatedness, we expect individuals to act prosocially towards individuals who appear similar to themselves. However, cues of kinship should have a less positive effect in sexual contexts, because of inbreeding's detrimental effects on offspring quality [ ]. One study investigated this prediction by comparing perceptions of the attractiveness of self-resembling own-sex and opposite-sex faces [ ]. Participants judged self-resemblance to be more attractive in the context of own-sex faces than in the context of opposite-sex faces. However, there was no such opposite-sex bias when the same faces were judged for averageness. This own-sex bias in preferences for self-resemblance indicates that, while self-resemblance is attractive in an exclusively prosocial i. Stronger attraction to cues of kinship in own-sex faces than in opposite-sex faces is likely to promote prosocial behaviour towards own-sex kin, while minimizing occurrences of inbreeding with opposite-sex kin. Transforms of self-similarity. Images are made by using the difference between a composite image of the same sex and an individual participant to make faces more similar to the participant. Self-dissimilar faces can be made by applying the same technique but using images other than the participant. Further evidence for context sensitivity in judgements of self-resembling faces is provided by a study comparing men's and women's preferences for self-resemblance in opposite-sex faces in explicitly prosocial versus sexual contexts [ ]. Participants were shown images of self-resembling opposite-sex faces and asked to judge their trustworthiness i. Consistent with both inclusive fitness and inbreeding avoidance theories, self-resemblance increased perceptions of trustworthiness, decreased attractiveness for short-term relationships and had no significant effect on attractiveness for long-term relationships. The fact that self-resemblance in opposite-sex faces was found to be trustworthy, but not attractive in short-term contexts, emphasizes the context-sensitivity of responses to self-resemblance. Importantly, because familiarity increases judgements of both attractiveness and trustworthiness [ ], this pattern of context-sensitivity strongly suggests that responses to self-resemblance do not occur simply because of familiarity alone i. Another example of social context influencing face preferences comes from research on interactions among the effects of different facial characteristics on preferences. For example, both behavioural and neurobiological evidence suggest that viewers demonstrate stronger attraction to attractive physical cues in faces e. Similarly, behavioural and neurobiological evidence also suggests that viewers demonstrate stronger attraction to cues associated with positive social interest e. Conway et al. Similarly, the putative costs of low investment are much less of a concern in short-term than in long-term relationships and, thus, women may demonstrate stronger masculinity preferences when judging men's attractiveness as possible short-term than long-term partners. Little et al. Women who were not using oral contraceptives made this face more masculine in the context of a short-term relationship than in the context of a long-term relationship. Penton-Voak et al. One potential explanation for this pattern of preference is that attractive women are better able to compete for, retain or replace high-quality, masculine partners and, therefore, do not show as large a shift in their preferences between short-term and long-term contexts. The effects of temporal context on judgements of attractiveness are not limited to faces. Women prefer lower pitched male voices in the context of a short-term relationship than in the context of a long-term relationship [ ]. This same study also found that the effect of relationship context was greatest when women were in the fertile phase of the menstrual cycle, a finding that is consistent with research on cyclic shifts in preferences for facial masculinity [ ]. A strong theoretical prediction of the trade-off account of variability in women's preferences for masculine men is that women in environments where poor health is particularly harmful to survival e. Regional differences in pathogen prevalence have been shown to be positively correlated with the importance placed on physical beauty and health [ , ]. DeBruine et al. This relationship remained significant, even when controlling for regional variation in wealth and women's mating strategies i. Brooks et al. Masculine-faced men may be favoured under such conditions, for example, as they may be better able to compete for resources. A further study of US states, in contrast, has shown that environmental health factors, and not indices of male—male violence such as homicide rates, predicts regional variation in women's masculinity preferences [ ]. Health, wealth and male—male violence are, of course, inter-related. While it is ultimately possible that health, wealth and male—male violence may all individually contribute to variation in preference, it is important to note that all of these analyses show that regional variation in women's masculinity preferences occurs in ways that are highly consistent with trade-off theories of sexual selection. The availability of resources in an environment may also influence face preferences. In low-resource environments, the resources to raise a child may be scarce or difficult to acquire and a preference for an investing partner be adaptive. To test these ideas, Little et al. Both men and women decreased their preferences for high quality mates for long-term relationships in the context of a harsh environment. This is consistent with the logic of trading genetic quality for commitment and investment in environments where resources are scarce. Individuals are confronted with a myriad of faces and social interactions every day. Research has shown that such experience leads to changes in preferences for faces. In the following section, we discuss two aspects of visual experience examining: Familiarity is a powerful determinant of attraction. For many types of stimuli, including faces, exposure increases attraction even when the exposure is unconscious [ — ]. Structural features of the face must be stored and represented in order to determine familiarity. As noted earlier, one idea for why averageness in faces is attractive comes from a link with familiarity—as average faces appear familiar this could positively affect their attractiveness [ 60 , 62 ]. Familiarity, when not paired with aversive stimuli, is thought to be rewarding [ ], and indeed there are obvious benefits to avoiding the unfamiliar. This can then help explain why exposure may cause increases in preference. There may, however, be more to increasing face preference than simple exposure. For example, recent studies have demonstrated that the nature of association positive or negative can affect face preferences, with positive experiences leading to increased attraction and negative experiences to decreased attraction [ ]. Moreover, these effects of valenced exposure are not bound solely to the specific individuals who were encountered and generalize to judgements of novel, physically similar individuals [ ]. Familiarity with parental traits has been implicated in human preferences. The phenomenon of imprinting, whereby individuals are attracted to parental traits, is well-studied in non-human animals [ , ] and there is increasing evidence for similar effects in humans. Following studies of facial similarity, judges have been shown to correctly match wives to their mother-in-law at a significantly higher rate than expected by chance and that wife—mother-in-law similarity is higher than similarity between husbands and their wives [ ]. Such effects are also seen in adopted daughters, controlling for any potential genetic effects, with significant facial resemblance between daughter's husband and her adoptive father [ ]. Other studies have shown that, for hair and eye colour, the best predictors of partner traits are the opposite-sex parent's colour traits [ ] and that individuals are attracted to age in faces consistent with the age of their parents when they were born [ ]. It is worth noting that at least in one study, effects were seen mainly for the opposite-sex parent [ ], which may indicate a more complex mechanism than simple exposure. Another line of argument suggesting imprinting-like effects appear not simply to reflect exposure comes from studies that have shown effects to be dependent on the quality of the relationship to the parent [ , ]. For example, daughters who report that they received greater emotional support from their adoptive fathers are more likely to choose mates who are similar to their father than individuals who report their father provided less emotional support [ ]. Similarly, women who rate their childhood relationships with their father positively show stronger attraction to face proportions similar to their father's face than women who rate their relationships less well [ ]. Imprinting-like effects then appear more complicated than simple exposure being directed more to one parent than the other and showing dependence on the relationship with that parent. Imprinting-like effects may lead to positive assortative mating pairing with similar partners , at least for long-term relationships, and this may have benefits in terms of keeping adaptive suites of genes together [ ] or increasing behaviour compatibility [ ]. There is certainly evidence that couples resemble each other facially [ , ]. Potentially then, a system that learns about known individuals and increases attraction to their face traits could be adaptive. Both familiarity and imprinting posit that exposure affects attractiveness. In recent years, exposure has been thought to have specific effects on our representations of faces via visual adaptation. We are unlikely to have an inbuilt average face and what is average must be calculated from experience. For each class of stimuli, the human visual system encounters may develop an individual representation, or prototype, made up of an average of the characteristics of all the different stimuli of that type that have been seen [ — ]. Computer modelling has revealed that algorithms trained to discriminate different stimuli produce stronger responses to stimuli that represent the average of the training set, even though this average was not previously encountered [ , ]. These findings have been interpreted as evidence that prototype formation is a property of learning to recognize different stimuli as members of a class [ , ]. Studies on category learning have a long history e. If a trait is reliably associated with some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive. Such an approach has highlighted face traits such as age, health, symmetry, and averageness, which are proposed to be associated with benefits and so associated with facial attractiveness. This view may postulate that some traits will be universally attractive; however, this does not preclude variation. Indeed, it would be surprising if there existed a template of a perfect face that was not affected by experience, environment, context, or the specific needs of an individual. Importantly, because familiarity increases judgements of both attractiveness and trustworthiness [ ], this pattern of context-sensitivity strongly suggests that responses to self-resemblance do not occur simply because of familiarity alone i. Another example of social context influencing face preferences comes from research on interactions among the effects of different facial characteristics on preferences. For example, both behavioural and neurobiological evidence suggest that viewers demonstrate stronger attraction to attractive physical cues in faces e. Similarly, behavioural and neurobiological evidence also suggests that viewers demonstrate stronger attraction to cues associated with positive social interest e. Conway et al. Similarly, the putative costs of low investment are much less of a concern in short-term than in long-term relationships and, thus, women may demonstrate stronger masculinity preferences when judging men's attractiveness as possible short-term than long-term partners. Little et al. Women who were not using oral contraceptives made this face more masculine in the context of a short-term relationship than in the context of a long-term relationship. Penton-Voak et al. One potential explanation for this pattern of preference is that attractive women are better able to compete for, retain or replace high-quality, masculine partners and, therefore, do not show as large a shift in their preferences between short-term and long-term contexts. The effects of temporal context on judgements of attractiveness are not limited to faces. Women prefer lower pitched male voices in the context of a short-term relationship than in the context of a long-term relationship [ ]. This same study also found that the effect of relationship context was greatest when women were in the fertile phase of the menstrual cycle, a finding that is consistent with research on cyclic shifts in preferences for facial masculinity [ ]. A strong theoretical prediction of the trade-off account of variability in women's preferences for masculine men is that women in environments where poor health is particularly harmful to survival e. Regional differences in pathogen prevalence have been shown to be positively correlated with the importance placed on physical beauty and health [ , ]. DeBruine et al. This relationship remained significant, even when controlling for regional variation in wealth and women's mating strategies i. Brooks et al. Masculine-faced men may be favoured under such conditions, for example, as they may be better able to compete for resources. A further study of US states, in contrast, has shown that environmental health factors, and not indices of male—male violence such as homicide rates, predicts regional variation in women's masculinity preferences [ ]. Health, wealth and male—male violence are, of course, inter-related. While it is ultimately possible that health, wealth and male—male violence may all individually contribute to variation in preference, it is important to note that all of these analyses show that regional variation in women's masculinity preferences occurs in ways that are highly consistent with trade-off theories of sexual selection. The availability of resources in an environment may also influence face preferences. In low-resource environments, the resources to raise a child may be scarce or difficult to acquire and a preference for an investing partner be adaptive. To test these ideas, Little et al. Both men and women decreased their preferences for high quality mates for long-term relationships in the context of a harsh environment. This is consistent with the logic of trading genetic quality for commitment and investment in environments where resources are scarce. Individuals are confronted with a myriad of faces and social interactions every day. Research has shown that such experience leads to changes in preferences for faces. In the following section, we discuss two aspects of visual experience examining: Familiarity is a powerful determinant of attraction. For many types of stimuli, including faces, exposure increases attraction even when the exposure is unconscious [ — ]. Structural features of the face must be stored and represented in order to determine familiarity. As noted earlier, one idea for why averageness in faces is attractive comes from a link with familiarity—as average faces appear familiar this could positively affect their attractiveness [ 60 , 62 ]. Familiarity, when not paired with aversive stimuli, is thought to be rewarding [ ], and indeed there are obvious benefits to avoiding the unfamiliar. This can then help explain why exposure may cause increases in preference. There may, however, be more to increasing face preference than simple exposure. For example, recent studies have demonstrated that the nature of association positive or negative can affect face preferences, with positive experiences leading to increased attraction and negative experiences to decreased attraction [ ]. Moreover, these effects of valenced exposure are not bound solely to the specific individuals who were encountered and generalize to judgements of novel, physically similar individuals [ ]. Familiarity with parental traits has been implicated in human preferences. The phenomenon of imprinting, whereby individuals are attracted to parental traits, is well-studied in non-human animals [ , ] and there is increasing evidence for similar effects in humans. Following studies of facial similarity, judges have been shown to correctly match wives to their mother-in-law at a significantly higher rate than expected by chance and that wife—mother-in-law similarity is higher than similarity between husbands and their wives [ ]. Such effects are also seen in adopted daughters, controlling for any potential genetic effects, with significant facial resemblance between daughter's husband and her adoptive father [ ]. Other studies have shown that, for hair and eye colour, the best predictors of partner traits are the opposite-sex parent's colour traits [ ] and that individuals are attracted to age in faces consistent with the age of their parents when they were born [ ]. It is worth noting that at least in one study, effects were seen mainly for the opposite-sex parent [ ], which may indicate a more complex mechanism than simple exposure. Another line of argument suggesting imprinting-like effects appear not simply to reflect exposure comes from studies that have shown effects to be dependent on the quality of the relationship to the parent [ , ]. For example, daughters who report that they received greater emotional support from their adoptive fathers are more likely to choose mates who are similar to their father than individuals who report their father provided less emotional support [ ]. Similarly, women who rate their childhood relationships with their father positively show stronger attraction to face proportions similar to their father's face than women who rate their relationships less well [ ]. Imprinting-like effects then appear more complicated than simple exposure being directed more to one parent than the other and showing dependence on the relationship with that parent. Imprinting-like effects may lead to positive assortative mating pairing with similar partners , at least for long-term relationships, and this may have benefits in terms of keeping adaptive suites of genes together [ ] or increasing behaviour compatibility [ ]. There is certainly evidence that couples resemble each other facially [ , ]. Potentially then, a system that learns about known individuals and increases attraction to their face traits could be adaptive. Both familiarity and imprinting posit that exposure affects attractiveness. In recent years, exposure has been thought to have specific effects on our representations of faces via visual adaptation. We are unlikely to have an inbuilt average face and what is average must be calculated from experience. For each class of stimuli, the human visual system encounters may develop an individual representation, or prototype, made up of an average of the characteristics of all the different stimuli of that type that have been seen [ — ]. Computer modelling has revealed that algorithms trained to discriminate different stimuli produce stronger responses to stimuli that represent the average of the training set, even though this average was not previously encountered [ , ]. These findings have been interpreted as evidence that prototype formation is a property of learning to recognize different stimuli as members of a class [ , ]. Studies on category learning have a long history e. Learning studies examine how categorical perception develops using abstract stimuli. In classic studies, it has been shown that exposure to different dot patterns with particular configurations results in abstraction so that the average of each of the patterns, while never previously seen, is recognized as belonging to the set of patterns from which it was derived [ ]. Faces have been the focus of much research regarding recognition and prototype formation. While it has been proposed that faces may be coded as veridical representations of individuals or exemplars [ ], recent neuroimaging and single-cell recording studies have supported a prototype-referenced model of face coding [ , ]. Exposure to faces biases subsequent perceptions of novel faces, causing faces similar to those initially viewed to appear more prototypical than they would otherwise be perceived as, presumably, a prototype or population of exemplars becomes updated [ — ]. For example, adaptation to faces with contracted features causes novel faces with contracted features to be perceived as more normal than prior to this exposure [ , , ]. Such after-effects are thought to reflect changes in the responses of neural mechanisms underlying face processing [ , — ]. These studies may then shed light on how the brain builds an average representation to which the other faces can be compared. Importantly, exposure in the manner described above also influences attractiveness judgements. After exposure to faces possessing certain traits, these traits come to be preferred [ , , , ]. For example, if exposed to faces that look more like one identity, then new faces that resemble that identity are found more attractive than if exposed to the opposite set of face traits. A similar effect has also been observed for judgements of the trustworthiness of faces [ ]. Adaptation then reflects the rapid updating of face norms and can therefore be tied both to the effects of familiarity and imprinting-like effects. We have dealt briefly with some aspects of simple experience on preferences above, but, of course, humans are highly social and much human experience is of what other humans do. Humans can therefore learn about attractiveness from the behaviour of those around them: We have recently reviewed social learning in human face preferences [ ], and so present a brief overview here. Social learning can be adaptive if it allows an individual to assess potential mates more quickly and efficiently than through individual trial and error or allows an individual to use another's expertise. Mate choice copying has been observed among females in a number of different non-human species [ — ], including fish [ — ] and bird species [ — ]. Such studies have generally shown that when females observe another female the model to be paired with one of the two males the targets , they are subsequently more likely to prefer the target male they had seen paired with the model over the male that was not paired with the model. Inspired by work on non-human species, recent research also suggests that social learning may influence human mate preferences. While some research has shown that the presence of wedding rings on men did not increase women's preferences for those men [ ], other studies have found that images of men labelled as married were more attractive than those labelled as single [ ] and that women rate men as more desirable when they are shown surrounded by women than when they are shown alone or with other men [ ]. Another study has shown that women prefer pictures of men that had been previously seen paired with images of other women who were looking at the face with smiling i. Women therefore do appear to mimic the attitude of other women to particular men. Alongside partnership status, simple presence, and expressions of attitude towards the male, the physical traits of the observed model may also play a role in the social transmission of preference. Previous studies have shown that men's and women's attractiveness judgements are influenced by the apparent choice of attractive members of the same sex. Such a phenomenon suggests a more sophisticated form of mate-choice copying, whereby women can use the attractiveness of a partner that a man can acquire in order to judge the man's own attractiveness. Another study using images that were presented with a fictitious partner has shown that both men and women find a face paired with an attractive partner to be more attractive than one paired with an unattractive partner for a long-term but not a short-term relationship [ ]. Being more or less attractive has important social consequences and people do generally agree on who is and who is not attractive. Beauty is not just a simple social construct—attractiveness appears to be ingrained in our biology. While some aspects of face perception might be innate, other aspects are clearly influenced by experience; it seems unlikely that individuals are born with a representation of what a perfect partner looks like. If a trait reliably advertises some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive. It is clear that individual differences in preferences for some traits will prove adaptive and so are consistent with evolutionary theory. Work on facial attractiveness is also integrative, combining theories and methods from behavioural ecology, cognition, cross-cultural research and social psychology. Anthony C. Jones , 2 and Lisa M. DeBruine 2. Benedict C. Lisa M. This article has been cited by other articles in PMC. Abstract Face preferences affect a diverse range of critical social outcomes, from mate choices and decisions about platonic relationships to hiring decisions and decisions about social exchange. The evolutionary basis of attraction: Open in a separate window. Adaptive individual differences In humans, while individuals may share certain basic criteria for finding faces attractive, many factors may influence the specific types of face they find attractive. Summary and conclusions Being more or less attractive has important social consequences and people do generally agree on who is and who is not attractive. References 1. Buss D. Preferences in human mate selection. Langlois J. Maxims or myths of beauty? A meta-analytic and theoretical review. Andersson M. Sexual selection. Princeton, NJ: Princeton University Press [ Google Scholar ]. Bilateral symmetry and sexual selection: Elder G. Appearance and education in marriage mobility. Holmes S. Personal appearance as related to scholastic records and marriage selection in college women. Riggio R. The role of non-verbal and physical attractiveness in the selection of dating partners. Berscheid E. Physical attractiveness and dating choice: Walster E. Importance of physical attractiveness in dating behaviour. Dion K. What is beautiful is good. Eagly A. What is beautiful is good, but …: Feingold A. Good-looking people are not what we think. Cash T. The aye of the beholder: Sigall H. Beautiful but dangerous: Downs A. Natural observations of the links between attractiveness and initial legal judgments. B 17 , — Chiu R. The relative importance of facial attractiveness and gender in Hong Kong selection decisions. Marlowe C. Gender and attractiveness biases in hiring decisions: Wolf N. The beauty myth. New York, NY: Morrow [ Google Scholar ]. Hume D. Four dissertations. Of the standard of taste. London, UK: Millar [ Google Scholar ]. Darwin C. The descent of man, and selection in relation to sex. John Murray [ Google Scholar ]. Physical attractiveness. In Advances in experimental social psychology ed. Berkowitz L. Academic Press [ Google Scholar ]. Ford C. Patterns of sexual behaviour. Cunningham M. Zebrowitz-McArthur L. Toward and ecological approach to social perception. Thornhill R. Facial attractiveness. Trends Cogn. Rhodes G. The evolutionary psychology of facial beauty. Gibson R. Relationships between blood parasites, mating success and phenotypic cues in male sage grouse. Moller A. Developmental stability and fitness: Asymmetry, developmental stability, and evolution. Oxford, UK: Oxford University Press [ Google Scholar ]. Valen L. A study of fluctuating asymmetry. Evolution 16 , — Dufour K. Bilateral symmetry and social dominance in captive male red-winged blackbirds. Manning J. Developmental stability, ejaculate size, and sperm quality in men. Breast asymmetry and phenotypic quality in women. Breast asymmetry, sexual selection, and human reproductive success. Facial sexual dimorphism, developmental stability, and susceptibility to disease in men and women. Gangestad S. Facial masculinity and fluctuating asymmetry. Little A. Symmetry is related to sexual dimorphism in faces: Human fluctuating asymmetry and sexual behaviour. Grammer K. Human Homo sapiens facial attractiveness and sexual selection: Scheib J. Facial attractiveness, symmetry, and cues to good genes. B , — Penton-Voak I. Symmetry, sexual dimorphism in facial proportions, and male facial attractiveness. Jones B. When facial attractiveness is only skin deep. Perception 33 , — Facial symmetry and judgements of apparent health: Mealey L. Symmetry and perceived facial attractiveness. Kowner R. Facial asymmetry and attractiveness judgment in developmental perspective. Human 22 , — Facial symmetry and the perception of beauty. Perrett D. Symmetry and human facial attractiveness. Evidence against perceptual bias views for symmetry preferences in human faces. Attraction independent of detection suggests special mechanisms for symmetry preferences in human face perception. Preferences for symmetry in human faces in two cultures: Waitt C. Preferences for symmetry in conspecific facial shape among Macaca mulatta. Perceived health contributes to the attractiveness of facial symmetry, averageness, and sexual dimorphism. Perception 36 , — Human facial beauty: Mitton J. Associations among proteins heterozygosity, growth rate, and developmental homeostasis. Roberts S. MHC-heterozygosity and human facial attractiveness. Lie H. Genetic diversity revealed in human faces. Evolution 62 , — Do facial averageness and symmetry signal health? Galton F. Composite portraits. Nature 18 , 97— Attractive faces are only average. What is average and what is not average about attractive faces. The role of masculinity and distinctiveness in judgments of human male facial attractiveness. Light L. Why attractive people are harder to remember. B 7 , — Averageness, exaggeration, and facial attractiveness. Are average facial configurations attractive only because of their symmetry? The role of symmetry in attraction to average faces. Apicella C. Facial averageness and attractiveness in an isolated population of hunter-gatherers. Alley T. Averaged faces are attractive, but very attractive faces are not average. Attractiveness of facial averageness and symmetry in non-Western populations: Perception 30 , — Enlow D. Handbook of facial growth , 2nd edn. Philadelphia, PA: Saunders [ Google Scholar ]. Zahavi A. Mate selection: Hillgarth N. Testosterone and immunosuppression in vertebrates: In Parasites and pathogens ed. Beckage N. Kanda N. Testosterone inhibits immunoglobulin production by human peripheral blood mononuclear cells. Yesilova Z. The effects of gonadotropin treatment on the immunological features of male patients with idiopathic hypogonadotropic hypogonadism. Folstad I. Parasites, bright males and the immunocompetence handicap. Parasitism, host immune function, and sexual selection. Does sexual dimorphism in human faces signal health? B , S93—S Law-Smith M. Facial appearance is a cue to oestrogen levels in women. High salivary testosterone is linked to masculine male facial appearance in humans. Neave N. Second to fourth digit ratio, testosterone and perceived male dominance. Measuring the physical in physical attractiveness: Jones D. Criteria of facial attractiveness in five populations. Effects of sexual dimorphism on facial attractiveness. Nature , — What do women want? Facialmetric assessment of multiple motives in the perception of male facial physical attractiveness. Berry D. Were the physiognomists right? Personality correlates of facial babyishness. B 15 , — Keating C. Gender and the physiognomy of dominance and attractiveness. McArthur L. Impressions of baby-faced adults. Cross-cultural agreement in perceptions of babyfaced adults..

One Facial attractiveness milffaceness has found that women's preferences for symmetric male faces were stronger around ovulation than Facial attractiveness milffaceness other phases of the menstrual cycle, at least among partnered women who were instructed to judge men's attractiveness as short-term mates [ ].

By contrast, other studies have observed no evidence for cyclic shifts in women's preferences for symmetric men's faces e. Although evidence that women's preferences for symmetry in men's faces vary systematically over the menstrual cycle is equivocal, that is not to say that robust cyclic shifts in women's perceptions of faces are only evident in their preferences for facial masculinity.

For example, women's aversions to Facial attractiveness milffaceness faces are enhanced around ovulation and positively correlated with women's estimated progesterone levels Facial attractiveness milffaceness the menstrual cycle [ ]. This variation in attitudes to self-resembling faces may reflect increased inbreeding avoidance around ovulation and increased preferences for caring, supportive and trustworthy individuals when increased progesterone prepares the body for pregnancy [ ].

Moreover, women's aversions to facial cues associated with current illness e. Indeed, pregnant women and women using oral contraceptives which mimic Facial attractiveness milffaceness effects of increased progesterone during pregnancy demonstrate stronger aversions to individuals displaying facial Facial attractiveness milffaceness of illness than do women with natural Facial attractiveness milffaceness cycles [ 97 ]. These latter findings for aversions to facial cues of illness and progesterone during the menstrual cycle complement other research on increased aversions to possible sources of contagion in women's food preferences during pregnancy [ ], as well as increased sensitivity to facial expressions signalling that sources of threat and contagion Facial attractiveness milffaceness nearby when progesterone levels are raised [].

While our discussion of hormone-mediated face preferences in women has Facial attractiveness milffaceness the positive findings that have been reported in the literature, it is important to note that there have also been unsuccessful replications of cyclic variation in women's face preferences.

For example, two recent studies observed no evidence for cyclic variations in women's preferences for masculine versus feminine male faces []. One possible explanation of these null findings comes from findings that suggest the extent to which women's preferences for masculine men vary over the menstrual Extreme pee hole insertion and fucking vary systematically among women.

For example, cyclic variation in women's preferences for masculine characteristics in men's voices is significantly greater among women with high trait i. This pattern of results may occur because varying their Facial attractiveness milffaceness strategy during the menstrual cycle may benefit unattractive women more than it benefits attractive women [ ].

More recent research has presented additional evidence that women's family background, prenatal hormone levels and mortality salience might also affect the extent to which they vary their masculinity preferences according to their menstrual cycle phase [ — ]. We also note that there are significant methodological differences between studies examining cycle effects, making direct comparisons e.

For example, some studies distinguish between short- and long-term mating contexts, generally with larger cyclic shifts for short-term judgements [ ], while others do not [ ]. Studies also differ in stimuli number, stimuli type and how fertility is defined. A thorough description of methodological differences between studies is not the focus here, but methodology is certainly a factor that could explain differences in findings across studies. It is likely that further research concerning individual differences in cyclic shifts and comparing different methodologies would provide important insights into the motivations, functions and mechanisms behind cyclic shifts in fundamental aspects of face perception.

While the previous section discussed Facial attractiveness milffaceness implicating hormone levels and fertility in individual differences in face perception, this section will discuss the relationships between face preferences and indices of own condition and attractiveness. Several studies have reported positive correlations between women's ratings of their own physical attractiveness and the strength of their preferences for masculine characteristics in men's faces Facial attractiveness milffaceness 92 ].

Other studies have extended this work by demonstrating that more objective measures of women's condition Facial attractiveness milffaceness attractiveness, such as their waist—hip ratio or oestrogen levels, predict their preferences for Facial attractiveness milffaceness characteristics in men's faces in the same way [ Facial attractiveness milffaceness, ]. Similar correlations between indices of women's own attractiveness and the strength of their preferences for masculine characteristics in other domains, such as men's voices, have also been reported [], and indices of women's own condition and attractiveness are positively correlated with the strength of Facial attractiveness milffaceness preferences for symmetry and healthy-looking skin in men's faces [ 92].

Facial attractiveness milffaceness findings described above appear to be somewhat analogous to condition-dependent preferences observed in other species, in Facial attractiveness milffaceness individuals in good physical "Facial attractiveness milffaceness" show stronger preferences for high-quality mates e.

Condition-dependent preferences in both humans and non-humans may have a common function and occur because individuals in good physical condition i. Particularly compelling evidence for this proposal comes from one of the few experimental studies of condition-dependent mate preferences. These findings suggest that women recalibrate subjective impressions of their own attractiveness i. While the Facial attractiveness milffaceness described above focused on the relationships between mate preferences and both individuals' own physical characteristics and their subjective please click for source of these physical characteristics, other work on condition-dependent preferences has investigated whether personality traits and other psychological factors predict individual differences in mate preferences in similar ways.

For example, individual differences in systemizing and sensation-seeking, both of which are components of male sex-typical psychology, are positively correlated with men's preferences for feminine characteristics in women's, but not men's, faces [].

Among women, individual differences in empathy, a component of female sex-typical psychology, and extraversion, a key predictor of social status that is correlated with women's physical attractiveness, are positively correlated with preferences for masculine characteristics Blonde girls upskirt panties men's, but not women's, faces [].

These findings not only implicate personality traits in individual differences in face preferences but also raise the intriguing possibility that some personality traits might mediate the relationships between an individual's physical characteristics and their face preferences. While factors such as hormones and own attractiveness can explain Facial attractiveness milffaceness in face preferences between individuals, the context under which judgements are made can also contribute to variation in standards of beauty.

In the following section, we discuss how context affects face preferences in three types of contexts: Facial attractiveness milffaceness about genetic kinship is available in the face and is perceived somewhat accurately [ — Facial attractiveness milffaceness. Judgements of facial similarity are highly synonymous with judgements of kinship [ — ], and facial similarity produced by computer-graphic manipulation affects behaviour in ways consistent with inclusive fitness theory e.

Therefore, responses to facial resemblance are likely to be affected by prosocial versus sexual contexts. Cues of kinship are predicted to increase preferences in non-sexual, prosocial contexts, Facial attractiveness milffaceness to the benefits associated with inclusive fitness [ ].

Facial attractiveness milffaceness other words, evolutionary models show that behaviours that benefit other individuals who share genes through Facial attractiveness milffaceness descent will be favoured. Therefore, if physical similarity is a reliable cue of genetic relatedness, we expect individuals to act prosocially towards individuals who appear similar to themselves.

However, cues of kinship should have a less positive effect in sexual contexts, because of inbreeding's detrimental effects on offspring quality [ ]. One study investigated this prediction by Facial attractiveness milffaceness perceptions of the attractiveness of self-resembling own-sex and opposite-sex faces [ ]. Participants judged self-resemblance to be more attractive in the context of own-sex faces than Facial attractiveness milffaceness the context of opposite-sex faces.

However, there was no Facial attractiveness milffaceness opposite-sex bias when the same faces were judged for averageness. This own-sex Facial attractiveness milffaceness in preferences for self-resemblance indicates that, while self-resemblance is Facial attractiveness milffaceness in an exclusively prosocial i.

Stronger attraction to cues Facial attractiveness milffaceness kinship in own-sex faces than in opposite-sex faces is likely to promote prosocial behaviour towards Facial attractiveness milffaceness kin, while minimizing occurrences of inbreeding with opposite-sex kin.

Transforms of self-similarity. Images are made by using the difference between a composite image of the same sex and an individual participant to make faces more similar to the participant. Self-dissimilar faces can be made by applying the same technique but using images other than the participant. Further evidence for context Facial attractiveness milffaceness in judgements of self-resembling faces is provided by a study comparing men's and women's preferences for self-resemblance in opposite-sex faces in explicitly prosocial versus sexual contexts [ ].

Participants were shown images of self-resembling opposite-sex faces and asked to judge their trustworthiness i.

Consistent with both inclusive fitness see more inbreeding avoidance theories, self-resemblance Facial attractiveness milffaceness perceptions of trustworthiness, decreased attractiveness for short-term relationships and had no significant effect on attractiveness for long-term relationships. The fact that self-resemblance in opposite-sex faces was found to be trustworthy, but not attractive in short-term contexts, emphasizes the context-sensitivity of responses to self-resemblance.

Importantly, because familiarity increases judgements of both attractiveness and trustworthiness [ ], this pattern of context-sensitivity strongly suggests that responses to self-resemblance do not occur simply because of familiarity alone i.

Another example of social context influencing face preferences comes from research on interactions among the effects check this out different Facial attractiveness milffaceness characteristics on preferences. For example, both behavioural and neurobiological evidence suggest Facial attractiveness milffaceness viewers demonstrate stronger attraction to attractive physical cues in faces e. Similarly, behavioural and neurobiological evidence also suggests that viewers demonstrate stronger attraction to cues associated with positive social interest e.

Conway et al. Similarly, the putative costs of low investment are much less of a concern in short-term than in long-term relationships and, thus, women may demonstrate stronger masculinity preferences when judging men's attractiveness as possible short-term than long-term partners.

Little et al. Women who were not using oral contraceptives made this face more masculine in the context of a short-term relationship than in the context of a long-term relationship.

Penton-Voak et al. One potential explanation for this pattern of preference is that attractive women Facial attractiveness milffaceness better able to Facial attractiveness milffaceness for, retain or replace high-quality, masculine partners this web page, therefore, do not show as large a shift in their preferences Facial attractiveness milffaceness short-term and long-term contexts.

The effects of temporal context on judgements of attractiveness are not limited to faces. Women prefer lower pitched male voices in the context Facial attractiveness milffaceness a short-term relationship than in the context of a long-term relationship [ ]. This same study also found that the effect of relationship context was greatest Facial attractiveness milffaceness women were in the fertile phase of the menstrual cycle, a finding that is consistent with research on cyclic shifts in preferences for facial masculinity [ ].

A strong theoretical prediction of the trade-off account of variability in women's preferences for masculine men is that women in environments where poor health is particularly harmful to survival e. Regional differences in pathogen prevalence have been shown to Facial attractiveness milffaceness positively correlated with the importance Facial attractiveness milffaceness on physical beauty and health [].

Facial attractiveness milffaceness et al. This relationship remained significant, even when controlling for regional variation in wealth and women's mating strategies i.

Brooks et al. Masculine-faced men may be favoured under such conditions, for example, as they may be better able to compete for resources.

A further study of US states, in contrast, has shown that environmental health factors, and not indices of male—male violence such as homicide rates, predicts regional variation in women's masculinity preferences [ ]. Health, wealth and male—male violence Facial attractiveness milffaceness, of course, inter-related. While it is ultimately possible that health, wealth and male—male violence may all individually contribute to article source in preference, it is important to note that all of these analyses show that regional variation in women's masculinity preferences occurs in ways that are highly consistent with trade-off theories of sexual selection.

The availability of resources in an environment may also influence face preferences. In low-resource environments, the resources to raise a child may be scarce or difficult to acquire and a preference for an investing partner be adaptive. To test these ideas, Little et al. Both men and women decreased their preferences link high quality mates for long-term relationships in the context of a harsh environment.

This is consistent with the logic of trading genetic quality for commitment and investment in environments where resources are scarce. Individuals are confronted with a myriad of faces and social interactions every day. Research has shown that such experience leads to changes in preferences for faces.

In the following section, we discuss two aspects of visual experience examining: Familiarity is a powerful determinant of attraction. For many types of stimuli, including faces, exposure increases attraction even when the exposure is unconscious [ — ]. Structural features of the face must be stored and represented in order to determine familiarity.

As noted earlier, one idea for why averageness in faces is attractive comes from a link with familiarity—as average faces appear familiar this could positively affect their attractiveness [ 6062 ]. Familiarity, when not paired with aversive stimuli, is thought to be rewarding [ ], and indeed there are obvious benefits to avoiding the unfamiliar. This can then help explain why exposure may cause increases in preference.

There may, however, be more to increasing face preference than simple exposure. For example, recent studies have demonstrated that the nature of association positive or negative can affect face preferences, with positive experiences leading to increased attraction and negative experiences to decreased attraction [ ]. Moreover, these effects of valenced exposure are not bound solely to Facial attractiveness milffaceness specific individuals who were encountered and generalize to judgements Facial attractiveness milffaceness novel, physically similar individuals [ ].

Familiarity with parental traits has been implicated in human preferences. The phenomenon of imprinting, whereby individuals are attracted to parental traits, is well-studied in non-human animals [] and there is increasing evidence for similar effects in humans. Following studies of facial similarity, judges have been shown to correctly match Facial attractiveness milffaceness to their mother-in-law at a significantly higher rate than expected by chance and that wife—mother-in-law similarity Facial attractiveness milffaceness higher than similarity between husbands and their wives [ ].

Such effects are also seen in adopted daughters, controlling for any potential genetic effects, with significant facial resemblance between daughter's husband and her adoptive father [ ]. Other studies have shown that, for hair and eye colour, the best predictors of partner traits are the opposite-sex parent's colour traits Facial attractiveness milffaceness ] and that Facial attractiveness milffaceness are attracted to age in faces consistent with the age of their parents when they were born [ ].

Sexy all in one

It is worth noting that at least in one study, effects were seen mainly for the opposite-sex parent [ Facial attractiveness milffaceness, which may indicate Facial attractiveness milffaceness more complex mechanism than simple exposure. Another line of argument suggesting imprinting-like effects appear not simply to reflect exposure comes from studies that have shown effects to be dependent on the quality of the relationship to the parent [].

For example, daughters who report that they received greater emotional support from their adoptive fathers are more likely to choose mates who are similar to their father than individuals who report Facial attractiveness milffaceness father provided less emotional support [ ]. Similarly, women who rate their childhood relationships Facial attractiveness milffaceness their father positively show stronger attraction to face proportions similar to their father's face than women who rate their relationships less well [ ].

Imprinting-like effects then appear more complicated than simple exposure being directed more to one parent than the other and showing dependence on the relationship with that parent. Imprinting-like effects may lead to positive assortative mating pairing with similar partnersat least for long-term relationships, and this may have benefits in terms of keeping adaptive suites of genes together [ ] or increasing behaviour compatibility [ ].

There is certainly evidence that couples resemble each other facially []. Potentially then, a system that learns about known individuals and increases attraction to their face traits could be adaptive. Both familiarity and imprinting posit that exposure affects attractiveness. In recent years, exposure has been thought to have specific effects on our representations of faces via visual adaptation. We are unlikely to have an inbuilt average face and what is average must be calculated from experience.

For each class of stimuli, the human visual Facial attractiveness milffaceness encounters may develop source individual representation, or prototype, made up of an average Facial attractiveness milffaceness the characteristics of all the different stimuli of go here type that have been seen [ — ].

Computer modelling has revealed that algorithms trained to discriminate different stimuli produce stronger responses to stimuli that represent the average of the training set, even though this average was not previously encountered []. Facial attractiveness milffaceness findings have been interpreted as evidence that prototype Facial attractiveness milffaceness is a property of learning to recognize different stimuli as members of a class [].

Studies on category learning have a long history e. Learning studies examine how categorical perception develops using abstract stimuli.

In classic studies, it has been Facial attractiveness milffaceness that exposure to different dot patterns with particular configurations results in abstraction so that the average of each of the patterns, while never previously seen, is recognized as belonging to the set of patterns from which it was derived [ ]. Faces have been the focus of much research regarding recognition and prototype formation. While it has been proposed that faces may be coded as veridical representations of individuals or exemplars [ ], recent neuroimaging and single-cell recording studies Facial attractiveness milffaceness supported a prototype-referenced Facial attractiveness milffaceness of face coding [].

Exposure to faces biases subsequent perceptions of novel faces, causing faces similar to those Facial attractiveness milffaceness viewed to appear more prototypical than they would otherwise be perceived as, presumably, a prototype or population of exemplars becomes updated Facial attractiveness milffaceness — ]. For example, adaptation to faces with contracted features causes novel faces with contracted features to be perceived as more normal than prior to this exposure [, ].

Such after-effects are thought to reflect changes in the responses of neural mechanisms underlying face processing [— ]. These studies may then shed light on how the brain builds an average representation to which the other faces can be compared. Importantly, exposure in the manner described above also influences attractiveness judgements. After exposure to Facial attractiveness milffaceness possessing certain traits, these traits come to be preferred [,]. For example, if exposed to faces that look more like one identity, then new faces that resemble that identity are found more attractive than if exposed to the opposite set of face traits.

A similar effect has also been observed for judgements of the trustworthiness of faces [ ].

Sexe Nipple Watch Video Video41ticket Sexix. These findings not only implicate personality traits in individual differences in face preferences but also raise the intriguing possibility that some personality traits might mediate the relationships between an individual's physical characteristics and their face preferences. While factors such as hormones and own attractiveness can explain differences in face preferences between individuals, the context under which judgements are made can also contribute to variation in standards of beauty. In the following section, we discuss how context affects face preferences in three types of contexts: Information about genetic kinship is available in the face and is perceived somewhat accurately [ — ]. Judgements of facial similarity are highly synonymous with judgements of kinship [ — ], and facial similarity produced by computer-graphic manipulation affects behaviour in ways consistent with inclusive fitness theory e. Therefore, responses to facial resemblance are likely to be affected by prosocial versus sexual contexts. Cues of kinship are predicted to increase preferences in non-sexual, prosocial contexts, owing to the benefits associated with inclusive fitness [ ]. In other words, evolutionary models show that behaviours that benefit other individuals who share genes through common descent will be favoured. Therefore, if physical similarity is a reliable cue of genetic relatedness, we expect individuals to act prosocially towards individuals who appear similar to themselves. However, cues of kinship should have a less positive effect in sexual contexts, because of inbreeding's detrimental effects on offspring quality [ ]. One study investigated this prediction by comparing perceptions of the attractiveness of self-resembling own-sex and opposite-sex faces [ ]. Participants judged self-resemblance to be more attractive in the context of own-sex faces than in the context of opposite-sex faces. However, there was no such opposite-sex bias when the same faces were judged for averageness. This own-sex bias in preferences for self-resemblance indicates that, while self-resemblance is attractive in an exclusively prosocial i. Stronger attraction to cues of kinship in own-sex faces than in opposite-sex faces is likely to promote prosocial behaviour towards own-sex kin, while minimizing occurrences of inbreeding with opposite-sex kin. Transforms of self-similarity. Images are made by using the difference between a composite image of the same sex and an individual participant to make faces more similar to the participant. Self-dissimilar faces can be made by applying the same technique but using images other than the participant. Further evidence for context sensitivity in judgements of self-resembling faces is provided by a study comparing men's and women's preferences for self-resemblance in opposite-sex faces in explicitly prosocial versus sexual contexts [ ]. Participants were shown images of self-resembling opposite-sex faces and asked to judge their trustworthiness i. Consistent with both inclusive fitness and inbreeding avoidance theories, self-resemblance increased perceptions of trustworthiness, decreased attractiveness for short-term relationships and had no significant effect on attractiveness for long-term relationships. The fact that self-resemblance in opposite-sex faces was found to be trustworthy, but not attractive in short-term contexts, emphasizes the context-sensitivity of responses to self-resemblance. Importantly, because familiarity increases judgements of both attractiveness and trustworthiness [ ], this pattern of context-sensitivity strongly suggests that responses to self-resemblance do not occur simply because of familiarity alone i. Another example of social context influencing face preferences comes from research on interactions among the effects of different facial characteristics on preferences. For example, both behavioural and neurobiological evidence suggest that viewers demonstrate stronger attraction to attractive physical cues in faces e. Similarly, behavioural and neurobiological evidence also suggests that viewers demonstrate stronger attraction to cues associated with positive social interest e. Conway et al. Similarly, the putative costs of low investment are much less of a concern in short-term than in long-term relationships and, thus, women may demonstrate stronger masculinity preferences when judging men's attractiveness as possible short-term than long-term partners. Little et al. Women who were not using oral contraceptives made this face more masculine in the context of a short-term relationship than in the context of a long-term relationship. Penton-Voak et al. One potential explanation for this pattern of preference is that attractive women are better able to compete for, retain or replace high-quality, masculine partners and, therefore, do not show as large a shift in their preferences between short-term and long-term contexts. The effects of temporal context on judgements of attractiveness are not limited to faces. Women prefer lower pitched male voices in the context of a short-term relationship than in the context of a long-term relationship [ ]. This same study also found that the effect of relationship context was greatest when women were in the fertile phase of the menstrual cycle, a finding that is consistent with research on cyclic shifts in preferences for facial masculinity [ ]. A strong theoretical prediction of the trade-off account of variability in women's preferences for masculine men is that women in environments where poor health is particularly harmful to survival e. Regional differences in pathogen prevalence have been shown to be positively correlated with the importance placed on physical beauty and health [ , ]. DeBruine et al. This relationship remained significant, even when controlling for regional variation in wealth and women's mating strategies i. Brooks et al. Masculine-faced men may be favoured under such conditions, for example, as they may be better able to compete for resources. A further study of US states, in contrast, has shown that environmental health factors, and not indices of male—male violence such as homicide rates, predicts regional variation in women's masculinity preferences [ ]. Health, wealth and male—male violence are, of course, inter-related. While it is ultimately possible that health, wealth and male—male violence may all individually contribute to variation in preference, it is important to note that all of these analyses show that regional variation in women's masculinity preferences occurs in ways that are highly consistent with trade-off theories of sexual selection. The availability of resources in an environment may also influence face preferences. In low-resource environments, the resources to raise a child may be scarce or difficult to acquire and a preference for an investing partner be adaptive. To test these ideas, Little et al. Both men and women decreased their preferences for high quality mates for long-term relationships in the context of a harsh environment. This is consistent with the logic of trading genetic quality for commitment and investment in environments where resources are scarce. Individuals are confronted with a myriad of faces and social interactions every day. Research has shown that such experience leads to changes in preferences for faces. In the following section, we discuss two aspects of visual experience examining: Familiarity is a powerful determinant of attraction. For many types of stimuli, including faces, exposure increases attraction even when the exposure is unconscious [ — ]. Structural features of the face must be stored and represented in order to determine familiarity. As noted earlier, one idea for why averageness in faces is attractive comes from a link with familiarity—as average faces appear familiar this could positively affect their attractiveness [ 60 , 62 ]. Familiarity, when not paired with aversive stimuli, is thought to be rewarding [ ], and indeed there are obvious benefits to avoiding the unfamiliar. This can then help explain why exposure may cause increases in preference. There may, however, be more to increasing face preference than simple exposure. For example, recent studies have demonstrated that the nature of association positive or negative can affect face preferences, with positive experiences leading to increased attraction and negative experiences to decreased attraction [ ]. Moreover, these effects of valenced exposure are not bound solely to the specific individuals who were encountered and generalize to judgements of novel, physically similar individuals [ ]. Familiarity with parental traits has been implicated in human preferences. The phenomenon of imprinting, whereby individuals are attracted to parental traits, is well-studied in non-human animals [ , ] and there is increasing evidence for similar effects in humans. Following studies of facial similarity, judges have been shown to correctly match wives to their mother-in-law at a significantly higher rate than expected by chance and that wife—mother-in-law similarity is higher than similarity between husbands and their wives [ ]. Such effects are also seen in adopted daughters, controlling for any potential genetic effects, with significant facial resemblance between daughter's husband and her adoptive father [ ]. Other studies have shown that, for hair and eye colour, the best predictors of partner traits are the opposite-sex parent's colour traits [ ] and that individuals are attracted to age in faces consistent with the age of their parents when they were born [ ]. It is worth noting that at least in one study, effects were seen mainly for the opposite-sex parent [ ], which may indicate a more complex mechanism than simple exposure. Another line of argument suggesting imprinting-like effects appear not simply to reflect exposure comes from studies that have shown effects to be dependent on the quality of the relationship to the parent [ , ]. For example, daughters who report that they received greater emotional support from their adoptive fathers are more likely to choose mates who are similar to their father than individuals who report their father provided less emotional support [ ]. Similarly, women who rate their childhood relationships with their father positively show stronger attraction to face proportions similar to their father's face than women who rate their relationships less well [ ]. Imprinting-like effects then appear more complicated than simple exposure being directed more to one parent than the other and showing dependence on the relationship with that parent. Imprinting-like effects may lead to positive assortative mating pairing with similar partners , at least for long-term relationships, and this may have benefits in terms of keeping adaptive suites of genes together [ ] or increasing behaviour compatibility [ ]. There is certainly evidence that couples resemble each other facially [ , ]. Potentially then, a system that learns about known individuals and increases attraction to their face traits could be adaptive. Both familiarity and imprinting posit that exposure affects attractiveness. In recent years, exposure has been thought to have specific effects on our representations of faces via visual adaptation. We are unlikely to have an inbuilt average face and what is average must be calculated from experience. For each class of stimuli, the human visual system encounters may develop an individual representation, or prototype, made up of an average of the characteristics of all the different stimuli of that type that have been seen [ — ]. Computer modelling has revealed that algorithms trained to discriminate different stimuli produce stronger responses to stimuli that represent the average of the training set, even though this average was not previously encountered [ , ]. These findings have been interpreted as evidence that prototype formation is a property of learning to recognize different stimuli as members of a class [ , ]. Studies on category learning have a long history e. Learning studies examine how categorical perception develops using abstract stimuli. In classic studies, it has been shown that exposure to different dot patterns with particular configurations results in abstraction so that the average of each of the patterns, while never previously seen, is recognized as belonging to the set of patterns from which it was derived [ ]. Faces have been the focus of much research regarding recognition and prototype formation. While it has been proposed that faces may be coded as veridical representations of individuals or exemplars [ ], recent neuroimaging and single-cell recording studies have supported a prototype-referenced model of face coding [ , ]. Exposure to faces biases subsequent perceptions of novel faces, causing faces similar to those initially viewed to appear more prototypical than they would otherwise be perceived as, presumably, a prototype or population of exemplars becomes updated [ — ]. For example, adaptation to faces with contracted features causes novel faces with contracted features to be perceived as more normal than prior to this exposure [ , , ]. Such after-effects are thought to reflect changes in the responses of neural mechanisms underlying face processing [ , — ]. These studies may then shed light on how the brain builds an average representation to which the other faces can be compared. Importantly, exposure in the manner described above also influences attractiveness judgements. After exposure to faces possessing certain traits, these traits come to be preferred [ , , , ]. For example, if exposed to faces that look more like one identity, then new faces that resemble that identity are found more attractive than if exposed to the opposite set of face traits. A similar effect has also been observed for judgements of the trustworthiness of faces [ ]. Adaptation then reflects the rapid updating of face norms and can therefore be tied both to the effects of familiarity and imprinting-like effects. We have dealt briefly with some aspects of simple experience on preferences above, but, of course, humans are highly social and much human experience is of what other humans do. Humans can therefore learn about attractiveness from the behaviour of those around them: We have recently reviewed social learning in human face preferences [ ], and so present a brief overview here. Social learning can be adaptive if it allows an individual to assess potential mates more quickly and efficiently than through individual trial and error or allows an individual to use another's expertise. Mate choice copying has been observed among females in a number of different non-human species [ — ], including fish [ — ] and bird species [ — ]. Such studies have generally shown that when females observe another female the model to be paired with one of the two males the targets , they are subsequently more likely to prefer the target male they had seen paired with the model over the male that was not paired with the model. Inspired by work on non-human species, recent research also suggests that social learning may influence human mate preferences. While some research has shown that the presence of wedding rings on men did not increase women's preferences for those men [ ], other studies have found that images of men labelled as married were more attractive than those labelled as single [ ] and that women rate men as more desirable when they are shown surrounded by women than when they are shown alone or with other men [ ]. Another study has shown that women prefer pictures of men that had been previously seen paired with images of other women who were looking at the face with smiling i. Women therefore do appear to mimic the attitude of other women to particular men. Alongside partnership status, simple presence, and expressions of attitude towards the male, the physical traits of the observed model may also play a role in the social transmission of preference. Previous studies have shown that men's and women's attractiveness judgements are influenced by the apparent choice of attractive members of the same sex. Such a phenomenon suggests a more sophisticated form of mate-choice copying, whereby women can use the attractiveness of a partner that a man can acquire in order to judge the man's own attractiveness. Another study using images that were presented with a fictitious partner has shown that both men and women find a face paired with an attractive partner to be more attractive than one paired with an unattractive partner for a long-term but not a short-term relationship [ ]. Being more or less attractive has important social consequences and people do generally agree on who is and who is not attractive. Beauty is not just a simple social construct—attractiveness appears to be ingrained in our biology. While some aspects of face perception might be innate, other aspects are clearly influenced by experience; it seems unlikely that individuals are born with a representation of what a perfect partner looks like. If a trait reliably advertises some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive. It is clear that individual differences in preferences for some traits will prove adaptive and so are consistent with evolutionary theory. Work on facial attractiveness is also integrative, combining theories and methods from behavioural ecology, cognition, cross-cultural research and social psychology. Anthony C. Jones , 2 and Lisa M. DeBruine 2. Benedict C. Lisa M. This article has been cited by other articles in PMC. Abstract Face preferences affect a diverse range of critical social outcomes, from mate choices and decisions about platonic relationships to hiring decisions and decisions about social exchange. The evolutionary basis of attraction: Open in a separate window. Adaptive individual differences In humans, while individuals may share certain basic criteria for finding faces attractive, many factors may influence the specific types of face they find attractive. Summary and conclusions Being more or less attractive has important social consequences and people do generally agree on who is and who is not attractive. References 1. Buss D. Preferences in human mate selection. Langlois J. Maxims or myths of beauty? A meta-analytic and theoretical review. Andersson M. Sexual selection. Princeton, NJ: Princeton University Press [ Google Scholar ]. Bilateral symmetry and sexual selection: Elder G. Appearance and education in marriage mobility. Holmes S. Personal appearance as related to scholastic records and marriage selection in college women. Riggio R. The role of non-verbal and physical attractiveness in the selection of dating partners. Berscheid E. Physical attractiveness and dating choice: Walster E. Importance of physical attractiveness in dating behaviour. Dion K. What is beautiful is good. Eagly A. What is beautiful is good, but …: Feingold A. Good-looking people are not what we think. Cash T. The aye of the beholder: Sigall H. Beautiful but dangerous: Downs A. Natural observations of the links between attractiveness and initial legal judgments. B 17 , — Chiu R. The relative importance of facial attractiveness and gender in Hong Kong selection decisions. Marlowe C. Gender and attractiveness biases in hiring decisions: Wolf N. The beauty myth. New York, NY: Morrow [ Google Scholar ]. Hume D. Four dissertations. Of the standard of taste. London, UK: Millar [ Google Scholar ]. Darwin C. The descent of man, and selection in relation to sex. John Murray [ Google Scholar ]. Physical attractiveness. In Advances in experimental social psychology ed. Berkowitz L. Academic Press [ Google Scholar ]. Ford C. Patterns of sexual behaviour. Cunningham M. Zebrowitz-McArthur L. Toward and ecological approach to social perception. Thornhill R. Facial attractiveness. Trends Cogn. Rhodes G. The evolutionary psychology of facial beauty. Gibson R. Relationships between blood parasites, mating success and phenotypic cues in male sage grouse. Moller A. Developmental stability and fitness: Asymmetry, developmental stability, and evolution. Oxford, UK: Oxford University Press [ Google Scholar ]. Valen L. A study of fluctuating asymmetry. Evolution 16 , — Dufour K. Bilateral symmetry and social dominance in captive male red-winged blackbirds. Manning J. Developmental stability, ejaculate size, and sperm quality in men. Breast asymmetry and phenotypic quality in women. Breast asymmetry, sexual selection, and human reproductive success. Facial sexual dimorphism, developmental stability, and susceptibility to disease in men and women. Gangestad S. Facial masculinity and fluctuating asymmetry. Little A. Symmetry is related to sexual dimorphism in faces: Human fluctuating asymmetry and sexual behaviour. Grammer K. Human Homo sapiens facial attractiveness and sexual selection: Scheib J. Facial attractiveness, symmetry, and cues to good genes. B , — Penton-Voak I. Symmetry, sexual dimorphism in facial proportions, and male facial attractiveness. Jones B. When facial attractiveness is only skin deep. Perception 33 , — Facial symmetry and judgements of apparent health: Mealey L. Symmetry and perceived facial attractiveness. Kowner R. Facial asymmetry and attractiveness judgment in developmental perspective. Human 22 , — Facial symmetry and the perception of beauty. Perrett D. Symmetry and human facial attractiveness. Evidence against perceptual bias views for symmetry preferences in human faces. Attraction independent of detection suggests special mechanisms for symmetry preferences in human face perception. Preferences for symmetry in human faces in two cultures: Waitt C. Preferences for symmetry in conspecific facial shape among Macaca mulatta. Perceived health contributes to the attractiveness of facial symmetry, averageness, and sexual dimorphism. Perception 36 , — Human facial beauty: Mitton J. Associations among proteins heterozygosity, growth rate, and developmental homeostasis. Roberts S. MHC-heterozygosity and human facial attractiveness. Lie H. Genetic diversity revealed in human faces. Evolution 62 , — Do facial averageness and symmetry signal health? Galton F. Composite portraits. Nature 18 , 97— Attractive faces are only average. What is average and what is not average about attractive faces. The role of masculinity and distinctiveness in judgments of human male facial attractiveness. Light L. Why attractive people are harder to remember. B 7 , — Averageness, exaggeration, and facial attractiveness. Are average facial configurations attractive only because of their symmetry? The role of symmetry in attraction to average faces. Apicella C. Facial averageness and attractiveness in an isolated population of hunter-gatherers. Alley T. Averaged faces are attractive, but very attractive faces are not average. Attractiveness of facial averageness and symmetry in non-Western populations: Perception 30 , — Enlow D. Handbook of facial growth , 2nd edn. Philadelphia, PA: Saunders [ Google Scholar ]. Zahavi A. Mate selection: Hillgarth N. Testosterone and immunosuppression in vertebrates: In Parasites and pathogens ed. Beckage N. Kanda N. Testosterone inhibits immunoglobulin production by human peripheral blood mononuclear cells. Yesilova Z. The effects of gonadotropin treatment on the immunological features of male patients with idiopathic hypogonadotropic hypogonadism. Folstad I. Parasites, bright males and the immunocompetence handicap. Parasitism, host immune function, and sexual selection. Does sexual dimorphism in human faces signal health? Feingold A. Cash T. Sigall H. Downs A. B 17 , — Chiu R. Marlowe C. Wolf N. New York, NY: Hume D. Of the standard of taste. London, UK: Darwin C. John Murray. Advances in experimental social psychology ed. Academic Press. Ford C. Cunningham M. Zebrowitz-McArthur L. Thornhill R. Trends Cogn. Rhodes G. Gibson R. Moller A. Oxford, UK: Oxford University Press. Valen L. Evolution 16 , — Dufour K. Manning J. Gangestad S. Little A. Grammer K. Scheib J. B , — Penton-Voak I. Jones B. Perception 33 , — Mealey L. Kowner R. Human 22 , — Perrett D. Waitt C. Perception 36 , — Mitton J. Roberts S. Lie H. Evolution 62 , — Galton F. Nature 18 , 97— Light L. B 7 , — Apicella C. Alley T. Perception 30 , — Enlow D. Philadelphia, PA: Zahavi A. Hillgarth N. Parasites and pathogens ed. Kanda N. Yesilova Z. Folstad I. B , S93—S Law-Smith M. Neave N. Jones D. Nature , — Facialmetric assessment of multiple motives in the perception of male facial physical attractiveness. Berry D. Personality correlates of facial babyishness. B 15 , — Keating C. McArthur L. Swaddle J. B , 39— DeBruine L. Smith F. Krupp D. In press. Apparent health encourages reciprocity. Fink B. Millinski M. Pryke S. Setchell J. Ethology , 25— B , S—S Changizi M. Drummond P. Psychophysiology 38 , — Cuthill I. Hill R. Nature , Burley N. Elliot A. B 35 , — Stephen I. Hassin R. Otta E. Motor Skills 82 , — Buss D. Brain Sci. Borkenau P. Liggett J. Botwin M. Boothroyd L. Korthase K. Motor Skills 54 , — Coetzee V. Perception 38 , — Osborn D. Makeup and posture effects on physical attractiveness judgments. Saxton T. Johnston V. Puts D. Feinberg D. Havlicek J. Psychoneuroendocrinology 35 , — Vukovic J. Peters M. Welling L. Roney J. Johnston L. Macrae C. Oinonen K. Sexual Behav. Pepper G. Derntl B. Conway C. Harris C. Menstrual cycle and facial preferences reconsidered. Sex Roles. Scarbrough P. Vaughn J. Psychol , — Bakker T. Alvergne A. Nesse R. Bressan P. Kaminski G. Dal Martello M. Maloney L. Hamilton W. Bittles A. Buckingham G. Zajonc R. O'Doherty J. Neuropsychologia 41 , — B , 63— Kampe K. Main J. Inquiry 17 , — Schmitt D. Brooks R. Geary D. Sex Res. Mace R. Kendrick K. Lorenz K. Zietschrift Tierpsychol. Bereczkei T. Wiszewska A. Thiessen D. Hill C. Issues 32 , — Hinsz V. Bateson P. Mate choice ed. Cambridge, UK: Cambridge University Press. Enquist M. Giese M. Neurocomputing 65 , 93— Johnstone R. Loffler G. Valentine T. Posner M. Leopold D. Webster M. Richerson P. Chicago, IL: University of Chicago Press. Brown G. White D. Dugatkin L. Free Press. Galef B. Bioscience 55 , — CO;2 doi: Godin J. Witte K. Uller T. Eva K. An indirect examination of mate-choice copying in humans. Hill S. B 34 , — Versluys T and Skylark W The effect of leg-to-body ratio on male attractiveness depends on the ecological validity of the figures , Royal Society Open Science , 4: Hare R , Schlatter S , Rhodes G and Simmons L Putative sex-specific human pheromones do not affect gender perception, attractiveness ratings or unfaithfulness judgements of opposite sex faces , Royal Society Open Science , 4: Biological Sciences , This Issue. Theme issue 'Face perception: Little, Benedict C. Jones and Lisa M. Downloaded , times. Statistics from Altmetric. If a trait is reliably associated with some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive. Such an approach has highlighted face traits such as age, health, symmetry, and averageness, which are proposed to be associated with benefits and so associated with facial attractiveness. This view may postulate that some traits will be universally attractive; however, this does not preclude variation. Indeed, it would be surprising if there existed a template of a perfect face that was not affected by experience, environment, context, or the specific needs of an individual..

Adaptation then reflects the rapid updating of face norms and can therefore be tied both to the effects of familiarity and imprinting-like effects. We have dealt briefly with some aspects of simple experience on preferences above, but, of course, humans are highly social and much human experience is of what other humans do.

Humans can therefore learn about attractiveness from the behaviour of those around them: We Facial attractiveness milffaceness recently reviewed social learning in human face preferences [ ], and so present Facial attractiveness milffaceness brief overview here.

Social learning can be Facial attractiveness milffaceness if it allows an individual to assess potential mates more quickly and efficiently than through individual trial and error or allows an individual to use another's expertise. Mate choice copying has been observed https://caning.revia2018.host/num10694-nanal.php females in a number of different non-human species [ — ], including fish [ — ] and bird species [ — ].

Such studies have generally shown that when females observe another female the model to be paired with one of the two males the targetsthey are subsequently more likely to prefer the target male they had seen paired with read more model over the male Facial attractiveness milffaceness was not paired with the model.

Facial attractiveness: evolutionary based research

Inspired by work on non-human species, recent research also suggests that social learning may influence human mate Facial attractiveness milffaceness. While some research has shown that the article source of wedding rings on men did not increase women's preferences for those men [ ], other studies have found that images of men labelled as married were more attractive than those labelled as single [ ] and that women rate men Facial attractiveness milffaceness more desirable when they are shown surrounded by women than when they are shown alone or with other men [ ].

Another study has shown Facial attractiveness milffaceness women prefer pictures of men that had been previously seen paired with images of other women who were looking at the face with smiling i.

Women therefore do appear to mimic Facial attractiveness milffaceness attitude Facial attractiveness milffaceness other women to particular men. Alongside partnership status, simple presence, and expressions of attitude towards the male, the physical traits of the observed model may also play a role in the social transmission of preference.

By considering that attraction and mate choice are critical components of evolutionary selection, we can better understand the importance of beauty. There are many traits that are linked to facial attractiveness in humans and each may in some way impart benefits to individuals who act on their preferences.

If a trait is reliably associated with some benefit to the perceiver, then Facial attractiveness milffaceness would expect individuals in a population to find that trait attractive. Such an approach has highlighted face traits such as age, health, symmetry, and averageness, which are proposed to be associated with benefits and so associated with facial attractiveness.

Subsequently, other studies have replicated preferences for symmetry using manipulated stimuli in different Western samples e. Preferences for symmetry using manipulated faces have been found in African hunter—gatherers [ 51 ], and macaque monkeys gaze longer at symmetrical than at asymmetrical face images of conspecifics [ 52 ].

Symmetry and asymmetry. Symmetric images are usually preferred to asymmetric images. Importantly, recent studies have implicated perceptions of health in attraction to symmetric faces [ 4453 ] and have suggested that the mechanisms underpinning preferences for symmetric faces are different from those that might drive preferences for symmetry in mate-choice-irrelevant stimuli e.

Such findings suggest that preferences for symmetric faces reflect, at least in part, adaptations for mate choice. Averageness refers to how closely a face resembles the majority of other faces within a population; non-average faces have more extreme characteristics than the average of a population.

Average faces may be attractive because an alignment of features that is close to a population average is linked to genetic diversity [ 5455 ]. Facial attractiveness milffaceness are generally best adapted to proteins Facial attractiveness milffaceness are common in the host population; hence, parasites are adapted to the genes that code for the production of these proteins.

A second evolutionary theory for the attractiveness of averageness Facial attractiveness milffaceness faces is that extreme non-average genotypes are more likely to be homozygous for deleterious alleles, that is, to be more likely to possess genes that are detrimental to an individual than those with more average genotypes [ 54 ].

Both of these theories propose evolutionary benefits to mating with individuals possessing average faces. Recent studies have supported the link between averageness, heterozygosity i. Heterozygosity in the major histocompatibility complex MHC genes that code for proteins involved Facial attractiveness milffaceness immune response, is positively associated with facial attractiveness [ Facial attractiveness milffaceness ] and facial averageness [ 57 Facial attractiveness milffaceness.

More directly, another study has shown that facial Facial attractiveness milffaceness is positively related to medical health as measured article source actual medical records in both men and women [ 58 ]. There is good evidence that average faces are indeed found attractive.

Galton [ 59 ] first noted that multiple faces blended together were more attractive than the constituent faces. Recent studies have improved upon these techniques using computers to create digitally blended composite faces; generally, the Facial attractiveness milffaceness images in a composite, the more attractive it is found [ 60 — 62 ].

Aside from Facial attractiveness milffaceness images, Light et al. Average faces are generally more symmetric and symmetry is typically attractive in faces discussed in more detail above. Several studies have controlled for this confound in the original studies. When averageness and symmetry were independently manipulated, one study found that both manipulations positively and independently influenced attractiveness judgements [ 65 ].

Other studies have used perfectly symmetric images manipulated in averageness and still have demonstrated preferences for averageness [ 6667 ]. Facial attractiveness milffaceness, by comparing preferences for averageness when the effects of symmetry were controlled Facial attractiveness milffaceness and were not controlled for, Jones et al. It has also been noted that, in the original composite studies, the more images that are blended together the smoother the skin texture becomes, as imperfections such as lines or blemishes are averaged [ 68 ].

Image c should be more attractive than both of the other images. Composites are made by marking key locations around the main facial features e. The average location of each point of the component faces is then calculated to define the shape of Facial attractiveness milffaceness composite. The images of the individual faces are then warped to the relevant average shape before superimposing the images to produce a photographic quality composite image.

While the majority of the work described above has been carried out in North America, Britain and Australia, averageness has also been found to be attractive Facial attractiveness milffaceness different cultures. For example, facial averageness is also found attractive in Japanese participants [ 69 ] and in African hunter—gatherers [ 67 ]. Male and female faces differ in their shape. Mature features in adult human faces reflect the masculinization or feminization of secondary sexual characteristics that occurs at puberty.

These face shape differences, link part, arise because of the action of hormones such as testosterone.

Larger jawbones, more prominent cheekbones and read more cheeks are all features of male faces that differentiate them from female faces e. From an evolutionary view, extremes of secondary sexual characteristics more feminine for women, more masculine for Facial attractiveness milffaceness are proposed to be attractive because they advertise the quality of an individual in terms of heritable benefits; they indicate that the owners of such characteristics Facial attractiveness milffaceness good genes.

In other words, such traits advertise the possession of genes that are beneficial to offspring inheriting them in terms of survival or reproduction. One explanation Facial attractiveness milffaceness the importance of these facial traits is Facial attractiveness milffaceness they represent a handicap to an organism [ 71 ] and the costs of growing the trait means that only healthy individuals can afford to produce them.

For example, secondary sexual characteristics are proposed to be linked to parasite resistance because the sex hormones Facial attractiveness milffaceness influence their growth, particularly testosterone, lower immunocompetence. Testosterone has been Facial attractiveness milffaceness to the suppression of immune function in Facial attractiveness milffaceness species [ 72 ], including humans [ 7374 ]. Larger secondary sexual characteristics should be related to a healthier immune system because only healthy organisms can afford the high sex-hormone handicap on the immune system that is necessary to produce these characteristics [ 75 ].

In many non-human animal studies, there is a positive association between secondary sexual trait https://felching.revia2018.host/pub10787-mibin.php and immunocompetence e. A study by Rhodes et al. No relationship was found between femininity and actual health in female faces, though [ 77 ].

Another study has demonstrated that men's facial masculinity and women's facial femininity are negatively related to self reports of respiratory disease [ 35 ]. If health is heritable, then female preferences for masculinity and male preferences for Facial attractiveness milffaceness may indeed also reflect the choice of mates with good genes. There is also a link between hormonal profile and face shape. Women with higher circulating oestrogen have more feminine Facial attractiveness milffaceness [ 78 ], while men with high testosterone have more masculine faces [ 79 ], but see also [ 80 ].

If women with high oestrogen and men with high testosterone are valued Facial attractiveness milffaceness mates, preferences for cues of hormonal Facial attractiveness milffaceness could drive preferences for sexually dimorphic face shape.

Masculinity is transformed using the difference between male and female face shape as defined by creating a male and female composite. Preferences for masculinity in male faces vary across studies, but feminine Facial attractiveness milffaceness faces Facial attractiveness milffaceness consistently found more Facial attractiveness milffaceness than masculine female faces.

Facial attractiveness milffaceness is considerable evidence that feminine female faces are considered attractive. Studies measuring facial features from photographs of women [ 408182 ] and studies manipulating facial composites [ 83 ] all indicate that feminine features increase the attractiveness of female faces across different cultures. If oestrogenized female faces provide cues to fertility and health, then male preferences for such features are potentially adaptive. This reasoning does not require oestrogen to be immunosuppressive or part of a handicap.

The link between masculinity Facial attractiveness milffaceness attractiveness in male faces is Facial attractiveness milffaceness clear. Cunningham et al. Despite some findings showing a preference for more masculine and dominant faces, several studies have shown that feminine characteristics and faces of low dominance are of increased attractiveness [ 62838489 — 91 ].

Many studies have made use of computer graphic techniques to manipulate masculinity. Sexual dimorphism in face shape can be manipulated by taking the geometrical differences between average male and female face shapes and applying this difference to new faces, making more or less masculine or feminine versions [ 83 ].

This process simultaneously changes all dimorphic shape characteristics in the face. Facial attractiveness milffaceness et al. For the male face stimuli, the shape selected by Caucasians as most attractive was significantly feminized for both the Caucasian male face and the Japanese male face continua.

Similarly, Japanese participants also selected significantly feminized versions of the male stimuli for both the Japanese and Caucasian male face continua. Thus, in both cultures it was found that participants showed a preference for feminized male faces. Since then, several studies have also documented preferences for femininity [ 62909293 ], but some similar computer graphic studies have also reported preferences for masculinity [ 9495 ].

Although some of this variation may be attributed to other characteristics of the faces that varied between sets of stimuli [ 96 ], this does not explain the variability in preferences. We discuss the sources of individual differences in preferences for sexually dimorphic shape cues in the latter sections of our article. The face traits discussed so far Facial attractiveness milffaceness often been measured and manipulated but also studied in terms of perception and related to attractiveness.

The reasoning for why traits like symmetry are preferred is often related to underlying health.

Facial attractiveness milffaceness

Thus, it is important to examine perceptions of facial health directly. Perceived health is difficult to relate to any one metric, but people will readily rate faces for perceived health and show very high agreement on such ratings e. In evolutionary terms, there is a large and obvious selective advantage in detecting healthy partners both for social exchange and mate choice. Indeed, while the role of health in mate preferences is clear see belowrecent work has demonstrated that participants are more willing Facial attractiveness milffaceness reciprocate trust from healthy-looking social partners Facial attractiveness milffaceness from social partners who are relatively unhealthy-looking [ 98 ].

Such findings sex training Male slut the importance of health perceptions for social interaction generally. Again, as for previous traits, there may be both direct and indirect benefits to partnering with individuals who are perceived to be healthy. Facial healthiness. High healthiness is associated with higher ratings of attractiveness. There have been several studies Facial attractiveness milffaceness have addressed how facial appearance relates to the healthiness of an individual in humans.

The three traits discussed above are often manipulated by changing only face shape, but health perception appears to be related to facial colour and texture also. Fewer studies have examined how colour and texture of faces influence attractiveness judgements. One study has examined how well ratings of health from small patches of skin of faces are related to overall rated attractiveness when the whole face image is available.

Jones et al. In other research, homogeneity of skin colour Facial attractiveness milffaceness positively related to attractiveness [ 99 ]. Findings have also suggested that more heterozygous men also have healthier appearing skin [ 56 ].

Skin health may be a particularly useful marker of current health condition as it is more changeable than aspects such as symmetry or averageness. Coloration is directly related to the appearance of skin. Coloration Facial attractiveness milffaceness appears to be an important component of sexual selection in many species.

Red coloration is associated with dominance in fish [ ], birds [ ] and non-human primates [] and, consequently, is linked to attracting the opposite sex. It has been noted that primates with trichromatic vision are generally bare-faced [ ] and that, at least in humans, facial flushing is associated with anger and confrontation [ ].

In research on non-human primates, Facial attractiveness milffaceness has been much interest in colour. For example, experimental manipulation of colour shows that female rhesus macaques prefer images of redder male faces [ ], while males prefer images of redder female hindquarters [ ].

In mandrills, red facial colour is related to rank in males [ ], and females sexually present more visit web page to brighter males and also groom them more frequently [ ].

Red coloration also has consequences for behaviour in other species. For example, in bird species, the addition of red to stimuli can increase social dominance [ ]. In humans, it has been shown that wearing red in a variety of physically competitive sports is associated with an increased chance of winning over opponents [ ]. This has been interpreted as natural associations of red with dominance being extended to artificially displayed red in the same way that artificial stimuli can exploit innate responses to natural stimuli [].

One study pitting red versus blue shapes found that red shapes were seen as more aggressive, dominant and more likely to win in physical competitions [ ]. Red does generally seem to have aversive effects on human behaviour. For example, when taking exams, individuals move their body away from tests with red covers more than they do from those with green or grey covers [ ]. While these studies suggest the colour red may be seen as a threatening stimulus in humans, red also appears to enhance Facial attractiveness milffaceness in some instances.

For example, women are seen as more attractive by men when Facial attractiveness milffaceness with Facial attractiveness milffaceness backgrounds or with red clothing, relative to other colours [ ]. This effect appears to be specific to attractiveness judgements; red colour does not influence judgements of other traits such as kindness or intelligence and does not influence women's attractiveness judgements of other women [ ].

Further research has examined red coloration in faces and demonstrated a positive association with perceived health [ ]. The authors suggest that perception of healthy, oxygenated blood may drive associations between red and healthiness. Alongside redness, people also appear to think that skin yellowness Facial attractiveness milffaceness associated with healthy appearance in faces [ ].

Yellowness may advertise health via an association with diet, as carotenoids are associated with skin yellowness and are absorbed via the intake of fruit and vegetables [ ]. In a classic social psychology study, Dion et al. For example, attractive individuals were thought to be able to achieve more prestigious occupations, be more competent spouses with happier marriages and have better prospects for personal fulfilment. There has been a wealth of studies examining this attractiveness stereotype, demonstrating that attractive people are Facial attractiveness milffaceness in a positive light for a wide range of attributes compared with unattractive people.

Studies on attractiveness stereotypes have generally not addressed Facial attractiveness milffaceness particular characteristics of faces that make individuals either attractive or unattractive, or Facial attractiveness milffaceness features that Facial attractiveness milffaceness personality attributions, although Facial attractiveness milffaceness faces reliably elicit the same personality attributions [ ].

Expression certainly has large effects, with, for example, faces shown Facial attractiveness milffaceness smiles rated as more attractive and as having more positive personality traits than Facial attractiveness milffaceness faces e. Such facial expressions are transient, however, and will differ rapidly within individuals over time and across photographs. Despite some findings showing a preference for more masculine and dominant male faces e. Personality traits are reported cross-culturally to be among the most important factors in partner choice by Facial attractiveness milffaceness sexes [ 1].

If desired personality is so important, it Facial attractiveness milffaceness appear likely that personality attributions elicited by a face would affect its attractiveness. For example, women who value cooperation and good parenting may avoid masculine-faced men.

Thus, instead of feminine faces being attractive and Facial attractiveness milffaceness attractiveness driving positive personality attributions, it may be that the personality attributions are driving click the following article attractiveness judgements. Individuals may use personality stereotypes in mate selection to select partners with a personality that they desire.

Facial attractiveness milffaceness perceptual Facial attractiveness milffaceness to facial photographs are somewhat accurate e.

Ciderella Porn Watch Video Hot Kissis. For example, although many studies have demonstrated that women's preferences for the body odours of symmetric men are enhanced around ovulation reviewed in [ ] , evidence for cyclic shifts in women's preferences for symmetry in men's faces is inconsistent. One study has found that women's preferences for symmetric male faces were stronger around ovulation than during other phases of the menstrual cycle, at least among partnered women who were instructed to judge men's attractiveness as short-term mates [ ]. By contrast, other studies have observed no evidence for cyclic shifts in women's preferences for symmetric men's faces e. Although evidence that women's preferences for symmetry in men's faces vary systematically over the menstrual cycle is equivocal, that is not to say that robust cyclic shifts in women's perceptions of faces are only evident in their preferences for facial masculinity. For example, women's aversions to self-resembling faces are enhanced around ovulation and positively correlated with women's estimated progesterone levels during the menstrual cycle [ ]. This variation in attitudes to self-resembling faces may reflect increased inbreeding avoidance around ovulation and increased preferences for caring, supportive and trustworthy individuals when increased progesterone prepares the body for pregnancy [ ]. Moreover, women's aversions to facial cues associated with current illness e. Indeed, pregnant women and women using oral contraceptives which mimic the effects of increased progesterone during pregnancy demonstrate stronger aversions to individuals displaying facial cues of illness than do women with natural menstrual cycles [ 97 ]. These latter findings for aversions to facial cues of illness and progesterone during the menstrual cycle complement other research on increased aversions to possible sources of contagion in women's food preferences during pregnancy [ ], as well as increased sensitivity to facial expressions signalling that sources of threat and contagion are nearby when progesterone levels are raised [ , ]. While our discussion of hormone-mediated face preferences in women has emphasized the positive findings that have been reported in the literature, it is important to note that there have also been unsuccessful replications of cyclic variation in women's face preferences. For example, two recent studies observed no evidence for cyclic variations in women's preferences for masculine versus feminine male faces [ , ]. One possible explanation of these null findings comes from findings that suggest the extent to which women's preferences for masculine men vary over the menstrual cycle vary systematically among women. For example, cyclic variation in women's preferences for masculine characteristics in men's voices is significantly greater among women with high trait i. This pattern of results may occur because varying their sexual strategy during the menstrual cycle may benefit unattractive women more than it benefits attractive women [ ]. More recent research has presented additional evidence that women's family background, prenatal hormone levels and mortality salience might also affect the extent to which they vary their masculinity preferences according to their menstrual cycle phase [ — ]. We also note that there are significant methodological differences between studies examining cycle effects, making direct comparisons e. For example, some studies distinguish between short- and long-term mating contexts, generally with larger cyclic shifts for short-term judgements [ ], while others do not [ ]. Studies also differ in stimuli number, stimuli type and how fertility is defined. A thorough description of methodological differences between studies is not the focus here, but methodology is certainly a factor that could explain differences in findings across studies. It is likely that further research concerning individual differences in cyclic shifts and comparing different methodologies would provide important insights into the motivations, functions and mechanisms behind cyclic shifts in fundamental aspects of face perception. While the previous section discussed research implicating hormone levels and fertility in individual differences in face perception, this section will discuss the relationships between face preferences and indices of own condition and attractiveness. Several studies have reported positive correlations between women's ratings of their own physical attractiveness and the strength of their preferences for masculine characteristics in men's faces [ 92 ]. Other studies have extended this work by demonstrating that more objective measures of women's condition and attractiveness, such as their waist—hip ratio or oestrogen levels, predict their preferences for masculine characteristics in men's faces in the same way [ , ]. Similar correlations between indices of women's own attractiveness and the strength of their preferences for masculine characteristics in other domains, such as men's voices, have also been reported [ , ], and indices of women's own condition and attractiveness are positively correlated with the strength of their preferences for symmetry and healthy-looking skin in men's faces [ 92 , ]. The findings described above appear to be somewhat analogous to condition-dependent preferences observed in other species, in which individuals in good physical condition show stronger preferences for high-quality mates e. Condition-dependent preferences in both humans and non-humans may have a common function and occur because individuals in good physical condition i. Particularly compelling evidence for this proposal comes from one of the few experimental studies of condition-dependent mate preferences. These findings suggest that women recalibrate subjective impressions of their own attractiveness i. While the research described above focused on the relationships between mate preferences and both individuals' own physical characteristics and their subjective evaluations of these physical characteristics, other work on condition-dependent preferences has investigated whether personality traits and other psychological factors predict individual differences in mate preferences in similar ways. For example, individual differences in systemizing and sensation-seeking, both of which are components of male sex-typical psychology, are positively correlated with men's preferences for feminine characteristics in women's, but not men's, faces [ , ]. Among women, individual differences in empathy, a component of female sex-typical psychology, and extraversion, a key predictor of social status that is correlated with women's physical attractiveness, are positively correlated with preferences for masculine characteristics in men's, but not women's, faces [ , ]. These findings not only implicate personality traits in individual differences in face preferences but also raise the intriguing possibility that some personality traits might mediate the relationships between an individual's physical characteristics and their face preferences. While factors such as hormones and own attractiveness can explain differences in face preferences between individuals, the context under which judgements are made can also contribute to variation in standards of beauty. In the following section, we discuss how context affects face preferences in three types of contexts: Information about genetic kinship is available in the face and is perceived somewhat accurately [ — ]. Judgements of facial similarity are highly synonymous with judgements of kinship [ — ], and facial similarity produced by computer-graphic manipulation affects behaviour in ways consistent with inclusive fitness theory e. Therefore, responses to facial resemblance are likely to be affected by prosocial versus sexual contexts. Cues of kinship are predicted to increase preferences in non-sexual, prosocial contexts, owing to the benefits associated with inclusive fitness [ ]. In other words, evolutionary models show that behaviours that benefit other individuals who share genes through common descent will be favoured. Therefore, if physical similarity is a reliable cue of genetic relatedness, we expect individuals to act prosocially towards individuals who appear similar to themselves. However, cues of kinship should have a less positive effect in sexual contexts, because of inbreeding's detrimental effects on offspring quality [ ]. One study investigated this prediction by comparing perceptions of the attractiveness of self-resembling own-sex and opposite-sex faces [ ]. Participants judged self-resemblance to be more attractive in the context of own-sex faces than in the context of opposite-sex faces. However, there was no such opposite-sex bias when the same faces were judged for averageness. This own-sex bias in preferences for self-resemblance indicates that, while self-resemblance is attractive in an exclusively prosocial i. Stronger attraction to cues of kinship in own-sex faces than in opposite-sex faces is likely to promote prosocial behaviour towards own-sex kin, while minimizing occurrences of inbreeding with opposite-sex kin. Transforms of self-similarity. Images are made by using the difference between a composite image of the same sex and an individual participant to make faces more similar to the participant. Self-dissimilar faces can be made by applying the same technique but using images other than the participant. Further evidence for context sensitivity in judgements of self-resembling faces is provided by a study comparing men's and women's preferences for self-resemblance in opposite-sex faces in explicitly prosocial versus sexual contexts [ ]. Participants were shown images of self-resembling opposite-sex faces and asked to judge their trustworthiness i. Consistent with both inclusive fitness and inbreeding avoidance theories, self-resemblance increased perceptions of trustworthiness, decreased attractiveness for short-term relationships and had no significant effect on attractiveness for long-term relationships. The fact that self-resemblance in opposite-sex faces was found to be trustworthy, but not attractive in short-term contexts, emphasizes the context-sensitivity of responses to self-resemblance. Importantly, because familiarity increases judgements of both attractiveness and trustworthiness [ ], this pattern of context-sensitivity strongly suggests that responses to self-resemblance do not occur simply because of familiarity alone i. Another example of social context influencing face preferences comes from research on interactions among the effects of different facial characteristics on preferences. For example, both behavioural and neurobiological evidence suggest that viewers demonstrate stronger attraction to attractive physical cues in faces e. Similarly, behavioural and neurobiological evidence also suggests that viewers demonstrate stronger attraction to cues associated with positive social interest e. Conway et al. Similarly, the putative costs of low investment are much less of a concern in short-term than in long-term relationships and, thus, women may demonstrate stronger masculinity preferences when judging men's attractiveness as possible short-term than long-term partners. Little et al. Women who were not using oral contraceptives made this face more masculine in the context of a short-term relationship than in the context of a long-term relationship. Penton-Voak et al. One potential explanation for this pattern of preference is that attractive women are better able to compete for, retain or replace high-quality, masculine partners and, therefore, do not show as large a shift in their preferences between short-term and long-term contexts. The effects of temporal context on judgements of attractiveness are not limited to faces. Women prefer lower pitched male voices in the context of a short-term relationship than in the context of a long-term relationship [ ]. This same study also found that the effect of relationship context was greatest when women were in the fertile phase of the menstrual cycle, a finding that is consistent with research on cyclic shifts in preferences for facial masculinity [ ]. A strong theoretical prediction of the trade-off account of variability in women's preferences for masculine men is that women in environments where poor health is particularly harmful to survival e. Regional differences in pathogen prevalence have been shown to be positively correlated with the importance placed on physical beauty and health [ , ]. DeBruine et al. This relationship remained significant, even when controlling for regional variation in wealth and women's mating strategies i. Brooks et al. Masculine-faced men may be favoured under such conditions, for example, as they may be better able to compete for resources. A further study of US states, in contrast, has shown that environmental health factors, and not indices of male—male violence such as homicide rates, predicts regional variation in women's masculinity preferences [ ]. Health, wealth and male—male violence are, of course, inter-related. While it is ultimately possible that health, wealth and male—male violence may all individually contribute to variation in preference, it is important to note that all of these analyses show that regional variation in women's masculinity preferences occurs in ways that are highly consistent with trade-off theories of sexual selection. The availability of resources in an environment may also influence face preferences. In low-resource environments, the resources to raise a child may be scarce or difficult to acquire and a preference for an investing partner be adaptive. To test these ideas, Little et al. Both men and women decreased their preferences for high quality mates for long-term relationships in the context of a harsh environment. This is consistent with the logic of trading genetic quality for commitment and investment in environments where resources are scarce. Individuals are confronted with a myriad of faces and social interactions every day. Research has shown that such experience leads to changes in preferences for faces. In the following section, we discuss two aspects of visual experience examining: Familiarity is a powerful determinant of attraction. For many types of stimuli, including faces, exposure increases attraction even when the exposure is unconscious [ — ]. Structural features of the face must be stored and represented in order to determine familiarity. As noted earlier, one idea for why averageness in faces is attractive comes from a link with familiarity—as average faces appear familiar this could positively affect their attractiveness [ 60 , 62 ]. Familiarity, when not paired with aversive stimuli, is thought to be rewarding [ ], and indeed there are obvious benefits to avoiding the unfamiliar. This can then help explain why exposure may cause increases in preference. There may, however, be more to increasing face preference than simple exposure. For example, recent studies have demonstrated that the nature of association positive or negative can affect face preferences, with positive experiences leading to increased attraction and negative experiences to decreased attraction [ ]. Moreover, these effects of valenced exposure are not bound solely to the specific individuals who were encountered and generalize to judgements of novel, physically similar individuals [ ]. Familiarity with parental traits has been implicated in human preferences. The phenomenon of imprinting, whereby individuals are attracted to parental traits, is well-studied in non-human animals [ , ] and there is increasing evidence for similar effects in humans. Following studies of facial similarity, judges have been shown to correctly match wives to their mother-in-law at a significantly higher rate than expected by chance and that wife—mother-in-law similarity is higher than similarity between husbands and their wives [ ]. Such effects are also seen in adopted daughters, controlling for any potential genetic effects, with significant facial resemblance between daughter's husband and her adoptive father [ ]. Other studies have shown that, for hair and eye colour, the best predictors of partner traits are the opposite-sex parent's colour traits [ ] and that individuals are attracted to age in faces consistent with the age of their parents when they were born [ ]. It is worth noting that at least in one study, effects were seen mainly for the opposite-sex parent [ ], which may indicate a more complex mechanism than simple exposure. Another line of argument suggesting imprinting-like effects appear not simply to reflect exposure comes from studies that have shown effects to be dependent on the quality of the relationship to the parent [ , ]. For example, daughters who report that they received greater emotional support from their adoptive fathers are more likely to choose mates who are similar to their father than individuals who report their father provided less emotional support [ ]. Similarly, women who rate their childhood relationships with their father positively show stronger attraction to face proportions similar to their father's face than women who rate their relationships less well [ ]. Imprinting-like effects then appear more complicated than simple exposure being directed more to one parent than the other and showing dependence on the relationship with that parent. Imprinting-like effects may lead to positive assortative mating pairing with similar partners , at least for long-term relationships, and this may have benefits in terms of keeping adaptive suites of genes together [ ] or increasing behaviour compatibility [ ]. There is certainly evidence that couples resemble each other facially [ , ]. Potentially then, a system that learns about known individuals and increases attraction to their face traits could be adaptive. Both familiarity and imprinting posit that exposure affects attractiveness. In recent years, exposure has been thought to have specific effects on our representations of faces via visual adaptation. We are unlikely to have an inbuilt average face and what is average must be calculated from experience. For each class of stimuli, the human visual system encounters may develop an individual representation, or prototype, made up of an average of the characteristics of all the different stimuli of that type that have been seen [ — ]. Computer modelling has revealed that algorithms trained to discriminate different stimuli produce stronger responses to stimuli that represent the average of the training set, even though this average was not previously encountered [ , ]. These findings have been interpreted as evidence that prototype formation is a property of learning to recognize different stimuli as members of a class [ , ]. Studies on category learning have a long history e. Learning studies examine how categorical perception develops using abstract stimuli. In classic studies, it has been shown that exposure to different dot patterns with particular configurations results in abstraction so that the average of each of the patterns, while never previously seen, is recognized as belonging to the set of patterns from which it was derived [ ]. Faces have been the focus of much research regarding recognition and prototype formation. While it has been proposed that faces may be coded as veridical representations of individuals or exemplars [ ], recent neuroimaging and single-cell recording studies have supported a prototype-referenced model of face coding [ , ]. Exposure to faces biases subsequent perceptions of novel faces, causing faces similar to those initially viewed to appear more prototypical than they would otherwise be perceived as, presumably, a prototype or population of exemplars becomes updated [ — ]. For example, adaptation to faces with contracted features causes novel faces with contracted features to be perceived as more normal than prior to this exposure [ , , ]. Such after-effects are thought to reflect changes in the responses of neural mechanisms underlying face processing [ , — ]. These studies may then shed light on how the brain builds an average representation to which the other faces can be compared. Importantly, exposure in the manner described above also influences attractiveness judgements. After exposure to faces possessing certain traits, these traits come to be preferred [ , , , ]. For example, if exposed to faces that look more like one identity, then new faces that resemble that identity are found more attractive than if exposed to the opposite set of face traits. A similar effect has also been observed for judgements of the trustworthiness of faces [ ]. Adaptation then reflects the rapid updating of face norms and can therefore be tied both to the effects of familiarity and imprinting-like effects. We have dealt briefly with some aspects of simple experience on preferences above, but, of course, humans are highly social and much human experience is of what other humans do. Humans can therefore learn about attractiveness from the behaviour of those around them: We have recently reviewed social learning in human face preferences [ ], and so present a brief overview here. Social learning can be adaptive if it allows an individual to assess potential mates more quickly and efficiently than through individual trial and error or allows an individual to use another's expertise. Mate choice copying has been observed among females in a number of different non-human species [ — ], including fish [ — ] and bird species [ — ]. Such studies have generally shown that when females observe another female the model to be paired with one of the two males the targets , they are subsequently more likely to prefer the target male they had seen paired with the model over the male that was not paired with the model. Inspired by work on non-human species, recent research also suggests that social learning may influence human mate preferences. While some research has shown that the presence of wedding rings on men did not increase women's preferences for those men [ ], other studies have found that images of men labelled as married were more attractive than those labelled as single [ ] and that women rate men as more desirable when they are shown surrounded by women than when they are shown alone or with other men [ ]. Another study has shown that women prefer pictures of men that had been previously seen paired with images of other women who were looking at the face with smiling i. Women therefore do appear to mimic the attitude of other women to particular men. Alongside partnership status, simple presence, and expressions of attitude towards the male, the physical traits of the observed model may also play a role in the social transmission of preference. Previous studies have shown that men's and women's attractiveness judgements are influenced by the apparent choice of attractive members of the same sex. Such a phenomenon suggests a more sophisticated form of mate-choice copying, whereby women can use the attractiveness of a partner that a man can acquire in order to judge the man's own attractiveness. Another study using images that were presented with a fictitious partner has shown that both men and women find a face paired with an attractive partner to be more attractive than one paired with an unattractive partner for a long-term but not a short-term relationship [ ]. Being more or less attractive has important social consequences and people do generally agree on who is and who is not attractive. Beauty is not just a simple social construct—attractiveness appears to be ingrained in our biology. While some aspects of face perception might be innate, other aspects are clearly influenced by experience; it seems unlikely that individuals are born with a representation of what a perfect partner looks like. If a trait reliably advertises some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive. It is clear that individual differences in preferences for some traits will prove adaptive and so are consistent with evolutionary theory. Work on facial attractiveness is also integrative, combining theories and methods from behavioural ecology, cognition, cross-cultural research and social psychology. Anthony C. Jones , 2 and Lisa M. DeBruine 2. Benedict C. Lisa M. This article has been cited by other articles in PMC. Abstract Face preferences affect a diverse range of critical social outcomes, from mate choices and decisions about platonic relationships to hiring decisions and decisions about social exchange. The evolutionary basis of attraction: Open in a separate window. Adaptive individual differences In humans, while individuals may share certain basic criteria for finding faces attractive, many factors may influence the specific types of face they find attractive. Summary and conclusions Being more or less attractive has important social consequences and people do generally agree on who is and who is not attractive. References 1. Buss D. Preferences in human mate selection. Langlois J. Maxims or myths of beauty? A meta-analytic and theoretical review. Andersson M. Sexual selection. Princeton, NJ: Princeton University Press [ Google Scholar ]. Bilateral symmetry and sexual selection: Elder G. Appearance and education in marriage mobility. Holmes S. Personal appearance as related to scholastic records and marriage selection in college women. Riggio R. The role of non-verbal and physical attractiveness in the selection of dating partners. Berscheid E. Physical attractiveness and dating choice: Walster E. Importance of physical attractiveness in dating behaviour. Dion K. What is beautiful is good. Eagly A. What is beautiful is good, but …: Feingold A. Good-looking people are not what we think. Cash T. The aye of the beholder: Sigall H. Beautiful but dangerous: Downs A. Natural observations of the links between attractiveness and initial legal judgments. B 17 , — Chiu R. The relative importance of facial attractiveness and gender in Hong Kong selection decisions. Marlowe C. Gender and attractiveness biases in hiring decisions: Wolf N. The beauty myth. New York, NY: Morrow [ Google Scholar ]. Hume D. Four dissertations. Of the standard of taste. London, UK: Millar [ Google Scholar ]. Darwin C. The descent of man, and selection in relation to sex. John Murray [ Google Scholar ]. Physical attractiveness. In Advances in experimental social psychology ed. Berkowitz L. Academic Press [ Google Scholar ]. Ford C. Patterns of sexual behaviour. Cunningham M. Zebrowitz-McArthur L. Toward and ecological approach to social perception. Thornhill R. Facial attractiveness. Trends Cogn. Rhodes G. The evolutionary psychology of facial beauty. Gibson R. Relationships between blood parasites, mating success and phenotypic cues in male sage grouse. Moller A. Developmental stability and fitness: Asymmetry, developmental stability, and evolution. Oxford, UK: Oxford University Press [ Google Scholar ]. Valen L. A study of fluctuating asymmetry. Evolution 16 , — Dufour K. Bilateral symmetry and social dominance in captive male red-winged blackbirds. Manning J. Developmental stability, ejaculate size, and sperm quality in men. Breast asymmetry and phenotypic quality in women. Breast asymmetry, sexual selection, and human reproductive success. Facial sexual dimorphism, developmental stability, and susceptibility to disease in men and women. Gangestad S. Facial masculinity and fluctuating asymmetry. Little A. Symmetry is related to sexual dimorphism in faces: Human fluctuating asymmetry and sexual behaviour. Grammer K. Human Homo sapiens facial attractiveness and sexual selection: Scheib J. Facial attractiveness, symmetry, and cues to good genes. B , — Penton-Voak I. Symmetry, sexual dimorphism in facial proportions, and male facial attractiveness. Jones B. When facial attractiveness is only skin deep. Perception 33 , — Facial symmetry and judgements of apparent health: Mealey L. Symmetry and perceived facial attractiveness. Kowner R. Facial asymmetry and attractiveness judgment in developmental perspective. Human 22 , — Facial symmetry and the perception of beauty. Perrett D. Symmetry and human facial attractiveness. Evidence against perceptual bias views for symmetry preferences in human faces. Attraction independent of detection suggests special mechanisms for symmetry preferences in human face perception. Preferences for symmetry in human faces in two cultures: Waitt C. Preferences for symmetry in conspecific facial shape among Macaca mulatta. Perceived health contributes to the attractiveness of facial symmetry, averageness, and sexual dimorphism. Perception 36 , — Human facial beauty: Mitton J. Associations among proteins heterozygosity, growth rate, and developmental homeostasis. Roberts S. MHC-heterozygosity and human facial attractiveness. Lie H. Genetic diversity revealed in human faces. Evolution 62 , — Do facial averageness and symmetry signal health? Galton F. Composite portraits. Nature 18 , 97— Attractive faces are only average. What is average and what is not average about attractive faces. In evolutionary terms, there is a large and obvious selective advantage in detecting healthy partners both for social exchange and mate choice. Indeed, while the role of health in mate preferences is clear see below , recent work has demonstrated that participants are more willing to reciprocate trust from healthy-looking social partners than from social partners who are relatively unhealthy-looking [ 98 ]. Such findings demonstrate the importance of health perceptions for social interaction generally. Again, as for previous traits, there may be both direct and indirect benefits to partnering with individuals who are perceived to be healthy. Facial healthiness. High healthiness is associated with higher ratings of attractiveness. There have been several studies that have addressed how facial appearance relates to the healthiness of an individual in humans. The three traits discussed above are often manipulated by changing only face shape, but health perception appears to be related to facial colour and texture also. Fewer studies have examined how colour and texture of faces influence attractiveness judgements. One study has examined how well ratings of health from small patches of skin of faces are related to overall rated attractiveness when the whole face image is available. Jones et al. In other research, homogeneity of skin colour was positively related to attractiveness [ 99 ]. Findings have also suggested that more heterozygous men also have healthier appearing skin [ 56 ]. Skin health may be a particularly useful marker of current health condition as it is more changeable than aspects such as symmetry or averageness. Coloration is directly related to the appearance of skin. Coloration also appears to be an important component of sexual selection in many species. Red coloration is associated with dominance in fish [ ], birds [ ] and non-human primates [ , ] and, consequently, is linked to attracting the opposite sex. It has been noted that primates with trichromatic vision are generally bare-faced [ ] and that, at least in humans, facial flushing is associated with anger and confrontation [ ]. In research on non-human primates, there has been much interest in colour. For example, experimental manipulation of colour shows that female rhesus macaques prefer images of redder male faces [ ], while males prefer images of redder female hindquarters [ ]. In mandrills, red facial colour is related to rank in males [ ], and females sexually present more frequently to brighter males and also groom them more frequently [ ]. Red coloration also has consequences for behaviour in other species. For example, in bird species, the addition of red to stimuli can increase social dominance [ ]. In humans, it has been shown that wearing red in a variety of physically competitive sports is associated with an increased chance of winning over opponents [ ]. This has been interpreted as natural associations of red with dominance being extended to artificially displayed red in the same way that artificial stimuli can exploit innate responses to natural stimuli [ , ]. One study pitting red versus blue shapes found that red shapes were seen as more aggressive, dominant and more likely to win in physical competitions [ ]. Red does generally seem to have aversive effects on human behaviour. For example, when taking exams, individuals move their body away from tests with red covers more than they do from those with green or grey covers [ ]. While these studies suggest the colour red may be seen as a threatening stimulus in humans, red also appears to enhance attraction in some instances. For example, women are seen as more attractive by men when presented with red backgrounds or with red clothing, relative to other colours [ ]. This effect appears to be specific to attractiveness judgements; red colour does not influence judgements of other traits such as kindness or intelligence and does not influence women's attractiveness judgements of other women [ ]. Further research has examined red coloration in faces and demonstrated a positive association with perceived health [ ]. The authors suggest that perception of healthy, oxygenated blood may drive associations between red and healthiness. Alongside redness, people also appear to think that skin yellowness is associated with healthy appearance in faces [ ]. Yellowness may advertise health via an association with diet, as carotenoids are associated with skin yellowness and are absorbed via the intake of fruit and vegetables [ ]. In a classic social psychology study, Dion et al. For example, attractive individuals were thought to be able to achieve more prestigious occupations, be more competent spouses with happier marriages and have better prospects for personal fulfilment. There has been a wealth of studies examining this attractiveness stereotype, demonstrating that attractive people are seen in a positive light for a wide range of attributes compared with unattractive people. Studies on attractiveness stereotypes have generally not addressed the particular characteristics of faces that make individuals either attractive or unattractive, or the features that elicit personality attributions, although different faces reliably elicit the same personality attributions [ ]. Expression certainly has large effects, with, for example, faces shown with smiles rated as more attractive and as having more positive personality traits than neutral faces e. Such facial expressions are transient, however, and will differ rapidly within individuals over time and across photographs. Despite some findings showing a preference for more masculine and dominant male faces e. Personality traits are reported cross-culturally to be among the most important factors in partner choice by both sexes [ 1 , ]. If desired personality is so important, it would appear likely that personality attributions elicited by a face would affect its attractiveness. For example, women who value cooperation and good parenting may avoid masculine-faced men. Thus, instead of feminine faces being attractive and this attractiveness driving positive personality attributions, it may be that the personality attributions are driving the attractiveness judgements. Individuals may use personality stereotypes in mate selection to select partners with a personality that they desire. Some perceptual attributions to facial photographs are somewhat accurate e. Attraction to faces based on personality stereotypes may happen regardless of whether attributions are accurate or not, especially as many individuals do believe that face provides an important guide to character [ , ]. One study has indeed demonstrated that a desire for some personality traits influences judgements of facial attractiveness [ ]. Individuals valuing particular personality traits find faces appearing to display these traits attractive. Conversely, those not valuing particular traits find faces attractive that are perceived to possess that trait less. Thus, desired personality influences perceptions of facial attractiveness in opposite sex faces, changing the result to: In terms of benefits to perceivers, it is easy to see why traits such as appearing trustworthy would make a face appear more attractive. For individual-specific traits, the logic is more complicated, but such preferences could be related to behavioural compatibility within couples, as people do tend to desire partners with personalities similar to their own [ ]. One reason for variability in preferences for male facial masculinity may lie in the personality traits that masculine- and feminine-faced men are assumed to possess. Increasing the masculinity of face shape increased perceptions of dominance, masculinity and age but decreased perceptions of warmth, emotionality, honesty, cooperativeness and quality as a parent [ 83 ]. Indeed, recent work has shown that masculine facial characteristics are associated with indices of physical dominance, such as physical strength [ ], and the perception of such traits [ ], and that feminine men show weaker preferences for short-term relationships and stronger preferences for committed, long-term relationships than their masculine peers do [ ]. Women's face preferences may thus represent a trade-off between the desire for good genes and the desire for a cooperative partner. Of course, the five types of trait listed above are not a complete list of factors involved in the judgement of facial attractiveness. While individual traits impact on attractiveness, there is also scope for interaction between them. Certain face traits also appear to interact in generating preferences, however. For example, preferences for masculinity vary as a function of the healthiness of the face [ 96 ] and women's preferences for facial self-similarity are higher when men are more facially masculine [ ]. Such interactions highlight that facial attractiveness judgements are not simple: In humans, while individuals may share certain basic criteria for finding faces attractive, many factors may influence the specific types of face they find attractive. In this section, we review three broad areas leading to individual differences in preferences: Research suggests that internal factors predict individual differences in several aspects of face perception, including attractiveness judgements. Importantly, the nature of these individual differences suggests adaptive design in face perception and face preferences. In the following section, we discuss two broad types of internal factors: The influence of hormones on face perception is an area that has generated a considerable amount of empirical research in recent years. As detailed previously, masculine characteristics in men's faces are associated with measures of long-term medical health [ 35 , 77 ] and indices of developmental stability [ 36 , 37 ], physical strength [ ] and reproductive potential [ ]. By contrast, feminine characteristics in men's faces are associated with cues of investment and stronger preferences for long-term over short-term sexual relationships e. There is now compelling evidence that how women resolve this trade off between the costs and benefits associated with choosing a masculine mate is affected by hormone levels and fertility. Many studies have reported that women demonstrate stronger preferences for men displaying masculine facial characteristics around ovulation, when women are most fertile, than during other phases of the menstrual cycle [ — ]. Some studies have also reported that these cyclic shifts in women's preferences for masculine characteristics in men's faces are greatest among women who already have romantic partners and when women judge men's attractiveness for short-term, extra-pair relationships [ ]. Although the ultimate function of these cyclic shifts remains somewhat controversial, many researchers have interpreted cyclic shifts in women's masculinity preferences as evidence for adaptations that function to increase offspring health via high paternal investment from a long-term partner while promoting attraction to other men displaying cues of heritable immunity to infectious disease when most fertile discussed in [ ]. Women may gain maximal benefits by selecting investing long-term partners and high-quality extra-pair partners. Importantly, other explanations that have been suggested, such as increased attraction to individuals who appear to be likely sources of high-quality care and support during phases of the menstrual cycle when increased progesterone prepares the body for pregnancy i. Increased attraction to masculine men is by no means unique to face preferences; women also demonstrate stronger attraction to masculine men when judging the attractiveness of men's voices [ — ], body shapes [ ] and body odours [ ], as well as when judging the attractiveness of videoclips of male behavioural displays of dominance [ , ]. Furthermore, converging evidence for fertility-related variation in women's preferences for facial masculinity comes from studies investigating circum-pubertal and circum-menopausal variation in women's masculinity preferences; post-menopausal and pre-pubertal women report weaker preferences for masculine facial characteristics than do their pre-menopausal and post-pubertal counterparts, respectively e. The ultimate function of cyclic shifts in women's preferences for masculine facial characteristics is not the only controversial aspect of cyclic shifts in women's masculinity preferences. For example, although some researchers have suggested that cyclic shifts in women's masculinity preferences may be an artefact of the computer graphic methods that are generally used in these studies to experimentally manipulate sexually dimorphic cues in digital face images [ ], this claim is very difficult to reconcile with findings from studies that have demonstrated cyclic shifts in women's preferences for masculinity in real i. While these findings suggest that cyclic shifts in women's masculinity preferences are not an artefact of the stimuli used, an aspect of research on cyclic shifts in women's masculinity preferences that remains controversial is whether the effect of cycle phase on women's face preferences is relatively specific to judgements of men's faces, or also occurs when women judge the attractiveness of other women. To date, evidence is equivocal; some studies have observed cyclic shifts in women's preferences for masculine-faced men, but not masculine-faced women [ ], while others have observed cyclic shifts in women's preferences for masculine faces, irrespective of their sex [ , ]. These latter papers speculate that cyclic shifts in women's preferences for masculine-faced women could represent a low-cost functionless by-product of a mechanism that evolved primarily to increase women's preferences for masculine men around ovulation [ ], or have suggested that higher attractiveness ratings given to masculine women around ovulation could reflect increased derogation of feminine, and therefore attractive, same-sex competitors when women are most fertile [ ] see also [ ]. In addition to the sex-specificity of the effects of cycle phase on face preferences, the mechanisms that underpin cyclic shifts in women's preferences for masculine characteristics in men's faces have also been a topic of considerable interest in recent years. For example, research into the hormonal mechanisms that might underpin these cyclic shifts has variously emphasized the effects of variation in levels of testosterone [ ], oestrogen [ ] and progesterone [ , ], or has suggested, perhaps unsurprisingly, that cyclic shifts in women's masculinity preferences might be best explained by complex interactions among multiple hormones [ , ]. While findings from research into the hormonal mechanisms that might underpin cyclic shifts in women's masculinity preferences have arguably been inconsistent, the findings of corresponding research into the psychological mechanisms have been relatively consistent; various studies have demonstrated that women are quicker to categorize men and access male stereotypes around ovulation e. These findings suggest that cyclic variations in stereotype access and sexual desire might be important psychological mechanisms for regulating facial masculinity preferences during the menstrual cycle. While research on hormone-mediated face perception has generally focused on women's judgements of men's attractiveness, some recent research has investigated hormone-mediated face preferences among men. Men, of course, do not cycle in the same way women do, but levels of testosterone fluctuate within individuals. Research using natural variation in testosterone has shown that men's preferences for feminine characteristics in women's faces are stronger when their testosterone levels are high than when they are relatively low [ ]. This finding suggests that hormones, such as testosterone, can generate within-participant individual differences in face preference in men. As can be seen from the previous paragraphs, there is compelling evidence that women's preferences for masculine men, be they assessed from face preferences or from preferences for male characteristics in other domains, vary systematically over the menstrual cycle. Whether or not preferences for other putative cues of men's long-term health are similarly affected by menstrual cycle is equivocal, however. For example, although many studies have demonstrated that women's preferences for the body odours of symmetric men are enhanced around ovulation reviewed in [ ] , evidence for cyclic shifts in women's preferences for symmetry in men's faces is inconsistent. One study has found that women's preferences for symmetric male faces were stronger around ovulation than during other phases of the menstrual cycle, at least among partnered women who were instructed to judge men's attractiveness as short-term mates [ ]. By contrast, other studies have observed no evidence for cyclic shifts in women's preferences for symmetric men's faces e. Although evidence that women's preferences for symmetry in men's faces vary systematically over the menstrual cycle is equivocal, that is not to say that robust cyclic shifts in women's perceptions of faces are only evident in their preferences for facial masculinity. For example, women's aversions to self-resembling faces are enhanced around ovulation and positively correlated with women's estimated progesterone levels during the menstrual cycle [ ]. This variation in attitudes to self-resembling faces may reflect increased inbreeding avoidance around ovulation and increased preferences for caring, supportive and trustworthy individuals when increased progesterone prepares the body for pregnancy [ ]. Moreover, women's aversions to facial cues associated with current illness e. Indeed, pregnant women and women using oral contraceptives which mimic the effects of increased progesterone during pregnancy demonstrate stronger aversions to individuals displaying facial cues of illness than do women with natural menstrual cycles [ 97 ]. These latter findings for aversions to facial cues of illness and progesterone during the menstrual cycle complement other research on increased aversions to possible sources of contagion in women's food preferences during pregnancy [ ], as well as increased sensitivity to facial expressions signalling that sources of threat and contagion are nearby when progesterone levels are raised [ , ]. While our discussion of hormone-mediated face preferences in women has emphasized the positive findings that have been reported in the literature, it is important to note that there have also been unsuccessful replications of cyclic variation in women's face preferences. For example, two recent studies observed no evidence for cyclic variations in women's preferences for masculine versus feminine male faces [ , ]. One possible explanation of these null findings comes from findings that suggest the extent to which women's preferences for masculine men vary over the menstrual cycle vary systematically among women. For example, cyclic variation in women's preferences for masculine characteristics in men's voices is significantly greater among women with high trait i. This pattern of results may occur because varying their sexual strategy during the menstrual cycle may benefit unattractive women more than it benefits attractive women [ ]. More recent research has presented additional evidence that women's family background, prenatal hormone levels and mortality salience might also affect the extent to which they vary their masculinity preferences according to their menstrual cycle phase [ — ]. We also note that there are significant methodological differences between studies examining cycle effects, making direct comparisons e. For example, some studies distinguish between short- and long-term mating contexts, generally with larger cyclic shifts for short-term judgements [ ], while others do not [ ]. Studies also differ in stimuli number, stimuli type and how fertility is defined. A thorough description of methodological differences between studies is not the focus here, but methodology is certainly a factor that could explain differences in findings across studies. It is likely that further research concerning individual differences in cyclic shifts and comparing different methodologies would provide important insights into the motivations, functions and mechanisms behind cyclic shifts in fundamental aspects of face perception. While the previous section discussed research implicating hormone levels and fertility in individual differences in face perception, this section will discuss the relationships between face preferences and indices of own condition and attractiveness. Several studies have reported positive correlations between women's ratings of their own physical attractiveness and the strength of their preferences for masculine characteristics in men's faces [ 92 ]. Other studies have extended this work by demonstrating that more objective measures of women's condition and attractiveness, such as their waist—hip ratio or oestrogen levels, predict their preferences for masculine characteristics in men's faces in the same way [ , ]. Similar correlations between indices of women's own attractiveness and the strength of their preferences for masculine characteristics in other domains, such as men's voices, have also been reported [ , ], and indices of women's own condition and attractiveness are positively correlated with the strength of their preferences for symmetry and healthy-looking skin in men's faces [ 92 , ]. The findings described above appear to be somewhat analogous to condition-dependent preferences observed in other species, in which individuals in good physical condition show stronger preferences for high-quality mates e. Condition-dependent preferences in both humans and non-humans may have a common function and occur because individuals in good physical condition i. Particularly compelling evidence for this proposal comes from one of the few experimental studies of condition-dependent mate preferences. These findings suggest that women recalibrate subjective impressions of their own attractiveness i. While the research described above focused on the relationships between mate preferences and both individuals' own physical characteristics and their subjective evaluations of these physical characteristics, other work on condition-dependent preferences has investigated whether personality traits and other psychological factors predict individual differences in mate preferences in similar ways. For example, individual differences in systemizing and sensation-seeking, both of which are components of male sex-typical psychology, are positively correlated with men's preferences for feminine characteristics in women's, but not men's, faces [ , ]. Among women, individual differences in empathy, a component of female sex-typical psychology, and extraversion, a key predictor of social status that is correlated with women's physical attractiveness, are positively correlated with preferences for masculine characteristics in men's, but not women's, faces [ , ]. These findings not only implicate personality traits in individual differences in face preferences but also raise the intriguing possibility that some personality traits might mediate the relationships between an individual's physical characteristics and their face preferences. While factors such as hormones and own attractiveness can explain differences in face preferences between individuals, the context under which judgements are made can also contribute to variation in standards of beauty. In the following section, we discuss how context affects face preferences in three types of contexts: Information about genetic kinship is available in the face and is perceived somewhat accurately [ — ]. Judgements of facial similarity are highly synonymous with judgements of kinship [ — ], and facial similarity produced by computer-graphic manipulation affects behaviour in ways consistent with inclusive fitness theory e. Therefore, responses to facial resemblance are likely to be affected by prosocial versus sexual contexts. Cues of kinship are predicted to increase preferences in non-sexual, prosocial contexts, owing to the benefits associated with inclusive fitness [ ]. In other words, evolutionary models show that behaviours that benefit other individuals who share genes through common descent will be favoured. Therefore, if physical similarity is a reliable cue of genetic relatedness, we expect individuals to act prosocially towards individuals who appear similar to themselves. However, cues of kinship should have a less positive effect in sexual contexts, because of inbreeding's detrimental effects on offspring quality [ ]. One study investigated this prediction by comparing perceptions of the attractiveness of self-resembling own-sex and opposite-sex faces [ ]. Participants judged self-resemblance to be more attractive in the context of own-sex faces than in the context of opposite-sex faces. However, there was no such opposite-sex bias when the same faces were judged for averageness. This own-sex bias in preferences for self-resemblance indicates that, while self-resemblance is attractive in an exclusively prosocial i. Stronger attraction to cues of kinship in own-sex faces than in opposite-sex faces is likely to promote prosocial behaviour towards own-sex kin, while minimizing occurrences of inbreeding with opposite-sex kin. Transforms of self-similarity. Images are made by using the difference between a composite image of the same sex and an individual participant to make faces more similar to the participant. Self-dissimilar faces can be made by applying the same technique but using images other than the participant. Further evidence for context sensitivity in judgements of self-resembling faces is provided by a study comparing men's and women's preferences for self-resemblance in opposite-sex faces in explicitly prosocial versus sexual contexts [ ]. Participants were shown images of self-resembling opposite-sex faces and asked to judge their trustworthiness i. Consistent with both inclusive fitness and inbreeding avoidance theories, self-resemblance increased perceptions of trustworthiness, decreased attractiveness for short-term relationships and had no significant effect on attractiveness for long-term relationships. The fact that self-resemblance in opposite-sex faces was found to be trustworthy, but not attractive in short-term contexts, emphasizes the context-sensitivity of responses to self-resemblance. Importantly, because familiarity increases judgements of both attractiveness and trustworthiness [ ], this pattern of context-sensitivity strongly suggests that responses to self-resemblance do not occur simply because of familiarity alone i. Another example of social context influencing face preferences comes from research on interactions among the effects of different facial characteristics on preferences. For example, both behavioural and neurobiological evidence suggest that viewers demonstrate stronger attraction to attractive physical cues in faces e. Similarly, behavioural and neurobiological evidence also suggests that viewers demonstrate stronger attraction to cues associated with positive social interest e. Conway et al. Similarly, the putative costs of low investment are much less of a concern in short-term than in long-term relationships and, thus, women may demonstrate stronger masculinity preferences when judging men's attractiveness as possible short-term than long-term partners. Little et al. Women who were not using oral contraceptives made this face more masculine in the context of a short-term relationship than in the context of a long-term relationship. Penton-Voak et al. One potential explanation for this pattern of preference is that attractive women are better able to compete for, retain or replace high-quality, masculine partners and, therefore, do not show as large a shift in their preferences between short-term and long-term contexts. The effects of temporal context on judgements of attractiveness are not limited to faces. Women prefer lower pitched male voices in the context of a short-term relationship than in the context of a long-term relationship [ ]. This same study also found that the effect of relationship context was greatest when women were in the fertile phase of the menstrual cycle, a finding that is consistent with research on cyclic shifts in preferences for facial masculinity [ ]. A strong theoretical prediction of the trade-off account of variability in women's preferences for masculine men is that women in environments where poor health is particularly harmful to survival e. Regional differences in pathogen prevalence have been shown to be positively correlated with the importance placed on physical beauty and health [ , ]. DeBruine et al. This relationship remained significant, even when controlling for regional variation in wealth and women's mating strategies i. Brooks et al. Masculine-faced men may be favoured under such conditions, for example, as they may be better able to compete for resources. A further study of US states, in contrast, has shown that environmental health factors, and not indices of male—male violence such as homicide rates, predicts regional variation in women's masculinity preferences [ ]. Health, wealth and male—male violence are, of course, inter-related. While it is ultimately possible that health, wealth and male—male violence may all individually contribute to variation in preference, it is important to note that all of these analyses show that regional variation in women's masculinity preferences occurs in ways that are highly consistent with trade-off theories of sexual selection. The availability of resources in an environment may also influence face preferences. In low-resource environments, the resources to raise a child may be scarce or difficult to acquire and a preference for an investing partner be adaptive. To test these ideas, Little et al. Both men and women decreased their preferences for high quality mates for long-term relationships in the context of a harsh environment. This is consistent with the logic of trading genetic quality for commitment and investment in environments where resources are scarce. Individuals are confronted with a myriad of faces and social interactions every day. Research has shown that such experience leads to changes in preferences for faces. In the following section, we discuss two aspects of visual experience examining: Familiarity is a powerful determinant of attraction. For many types of stimuli, including faces, exposure increases attraction even when the exposure is unconscious [ — ]. Structural features of the face must be stored and represented in order to determine familiarity. As noted earlier, one idea for why averageness in faces is attractive comes from a link with familiarity—as average faces appear familiar this could positively affect their attractiveness [ 60 , 62 ]. Familiarity, when not paired with aversive stimuli, is thought to be rewarding [ ], and indeed there are obvious benefits to avoiding the unfamiliar. This can then help explain why exposure may cause increases in preference. There may, however, be more to increasing face preference than simple exposure. For example, recent studies have demonstrated that the nature of association positive or negative can affect face preferences, with positive experiences leading to increased attraction and negative experiences to decreased attraction [ ]. Moreover, these effects of valenced exposure are not bound solely to the specific individuals who were encountered and generalize to judgements of novel, physically similar individuals [ ]. Familiarity with parental traits has been implicated in human preferences. The phenomenon of imprinting, whereby individuals are attracted to parental traits, is well-studied in non-human animals [ , ] and there is increasing evidence for similar effects in humans. Following studies of facial similarity, judges have been shown to correctly match wives to their mother-in-law at a significantly higher rate than expected by chance and that wife—mother-in-law similarity is higher than similarity between husbands and their wives [ ]. Such effects are also seen in adopted daughters, controlling for any potential genetic effects, with significant facial resemblance between daughter's husband and her adoptive father [ ]. Other studies have shown that, for hair and eye colour, the best predictors of partner traits are the opposite-sex parent's colour traits [ ] and that individuals are attracted to age in faces consistent with the age of their parents when they were born [ ]. It is worth noting that at least in one study, effects were seen mainly for the opposite-sex parent [ ], which may indicate a more complex mechanism than simple exposure. Another line of argument suggesting imprinting-like effects appear not simply to reflect exposure comes from studies that have shown effects to be dependent on the quality of the relationship to the parent [ , ]. For example, daughters who report that they received greater emotional support from their adoptive fathers are more likely to choose mates who are similar to their father than individuals who report their father provided less emotional support [ ]. Similarly, women who rate their childhood relationships with their father positively show stronger attraction to face proportions similar to their father's face than women who rate their relationships less well [ ]. Imprinting-like effects then appear more complicated than simple exposure being directed more to one parent than the other and showing dependence on the relationship with that parent. Imprinting-like effects may lead to positive assortative mating pairing with similar partners , at least for long-term relationships, and this may have benefits in terms of keeping adaptive suites of genes together [ ] or increasing behaviour compatibility [ ]. There is certainly evidence that couples resemble each other facially [ , ]. Potentially then, a system that learns about known individuals and increases attraction to their face traits could be adaptive. Both familiarity and imprinting posit that exposure affects attractiveness. In recent years, exposure has been thought to have specific effects on our representations of faces via visual adaptation. We are unlikely to have an inbuilt average face and what is average must be calculated from experience. For each class of stimuli, the human visual system encounters may develop an individual representation, or prototype, made up of an average of the characteristics of all the different stimuli of that type that have been seen [ — ]. Computer modelling has revealed that algorithms trained to discriminate different stimuli produce stronger responses to stimuli that represent the average of the training set, even though this average was not previously encountered [ , ]. Despite a widespread belief that beauty cannot be defined, in fact, there is considerable agreement across individuals and cultures on what is found attractive. By considering that attraction and mate choice are critical components of evolutionary selection, we can better understand the importance of beauty. There are many traits that are linked to facial attractiveness in humans and each may in some way impart benefits to individuals who act on their preferences. If a trait is reliably associated with some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive..

Attraction to faces based on personality stereotypes may happen regardless of whether attributions are accurate or not, especially as many individuals do believe that face provides an important guide to character [].

One study has indeed demonstrated that a desire for some personality traits influences Facial attractiveness milffaceness of facial attractiveness [ ].

Individuals valuing particular personality traits Facial attractiveness milffaceness faces appearing to display these traits attractive. Conversely, those not valuing particular traits find faces attractive that are perceived to possess that trait less. Thus, desired personality influences perceptions of facial attractiveness in opposite sex faces, changing the result Facial attractiveness milffaceness In terms of benefits to perceivers, it is easy to see why traits such as appearing trustworthy would make a face appear more attractive.

For individual-specific traits, the logic is more complicated, but such preferences could be related to behavioural compatibility within couples, as people do tend to desire partners with personalities similar to their own [ ]. One reason for variability in preferences for male facial masculinity may lie in the personality traits that masculine- and feminine-faced men are Facial attractiveness milffaceness to possess.

Increasing the masculinity of face shape increased perceptions of dominance, masculinity and Facial attractiveness milffaceness but decreased perceptions of warmth, emotionality, honesty, cooperativeness and quality as a parent [ 83 ]. Indeed, recent work has shown that masculine facial characteristics are associated with indices of physical dominance, such as physical strength [ ], and the check this out of such traits [ ], and that feminine men show weaker preferences for short-term relationships and stronger preferences for committed, Facial attractiveness milffaceness relationships than their masculine peers do [ ].

Women's face preferences may thus represent a trade-off between the desire for good genes and the desire for a cooperative partner. Of course, the five types of trait listed above are not a complete list of factors involved in the judgement of Facial attractiveness milffaceness attractiveness.

While individual traits impact on attractiveness, there is also scope for interaction between them. Certain face traits also appear to interact in generating preferences, however. For example, preferences for masculinity vary as a function of the healthiness of the face [ 96 ] and women's preferences for facial self-similarity are higher when men are more facially masculine [ ].

See more interactions highlight that facial attractiveness judgements are not simple: In humans, Facial attractiveness milffaceness individuals may share certain basic criteria for finding faces attractive, many factors may influence the specific types of face they find attractive. In this section, we review three broad areas leading to individual differences in preferences: Research suggests that internal factors predict individual differences in several aspects of face perception, including attractiveness judgements.

Importantly, the nature of these individual differences suggests adaptive design Facial attractiveness milffaceness face perception and face preferences.

In the following section, we discuss two broad types of internal factors: The influence of hormones on face perception is an area that has generated a considerable amount of empirical research in recent years. As Facial attractiveness milffaceness previously, masculine characteristics in men's faces are associated with measures of long-term medical health [ 3577 ] and indices of developmental stability [ 3637 ], physical strength [ Facial attractiveness milffaceness and reproductive potential [ ].

Blowjov videos Watch Video Emily Sparxxx. Again, as for previous traits, there may be both direct and indirect benefits to partnering with individuals who are perceived to be healthy. Facial healthiness. High healthiness is associated with higher ratings of attractiveness. There have been several studies that have addressed how facial appearance relates to the healthiness of an individual in humans. The three traits discussed above are often manipulated by changing only face shape, but health perception appears to be related to facial colour and texture also. Fewer studies have examined how colour and texture of faces influence attractiveness judgements. One study has examined how well ratings of health from small patches of skin of faces are related to overall rated attractiveness when the whole face image is available. Jones et al. In other research, homogeneity of skin colour was positively related to attractiveness [ 99 ]. Findings have also suggested that more heterozygous men also have healthier appearing skin [ 56 ]. Skin health may be a particularly useful marker of current health condition as it is more changeable than aspects such as symmetry or averageness. Coloration is directly related to the appearance of skin. Coloration also appears to be an important component of sexual selection in many species. Red coloration is associated with dominance in fish [ ], birds [ ] and non-human primates [ , ] and, consequently, is linked to attracting the opposite sex. It has been noted that primates with trichromatic vision are generally bare-faced [ ] and that, at least in humans, facial flushing is associated with anger and confrontation [ ]. In research on non-human primates, there has been much interest in colour. For example, experimental manipulation of colour shows that female rhesus macaques prefer images of redder male faces [ ], while males prefer images of redder female hindquarters [ ]. In mandrills, red facial colour is related to rank in males [ ], and females sexually present more frequently to brighter males and also groom them more frequently [ ]. Red coloration also has consequences for behaviour in other species. For example, in bird species, the addition of red to stimuli can increase social dominance [ ]. In humans, it has been shown that wearing red in a variety of physically competitive sports is associated with an increased chance of winning over opponents [ ]. This has been interpreted as natural associations of red with dominance being extended to artificially displayed red in the same way that artificial stimuli can exploit innate responses to natural stimuli [ , ]. One study pitting red versus blue shapes found that red shapes were seen as more aggressive, dominant and more likely to win in physical competitions [ ]. Red does generally seem to have aversive effects on human behaviour. For example, when taking exams, individuals move their body away from tests with red covers more than they do from those with green or grey covers [ ]. While these studies suggest the colour red may be seen as a threatening stimulus in humans, red also appears to enhance attraction in some instances. For example, women are seen as more attractive by men when presented with red backgrounds or with red clothing, relative to other colours [ ]. This effect appears to be specific to attractiveness judgements; red colour does not influence judgements of other traits such as kindness or intelligence and does not influence women's attractiveness judgements of other women [ ]. Further research has examined red coloration in faces and demonstrated a positive association with perceived health [ ]. The authors suggest that perception of healthy, oxygenated blood may drive associations between red and healthiness. Alongside redness, people also appear to think that skin yellowness is associated with healthy appearance in faces [ ]. Yellowness may advertise health via an association with diet, as carotenoids are associated with skin yellowness and are absorbed via the intake of fruit and vegetables [ ]. In a classic social psychology study, Dion et al. For example, attractive individuals were thought to be able to achieve more prestigious occupations, be more competent spouses with happier marriages and have better prospects for personal fulfilment. There has been a wealth of studies examining this attractiveness stereotype, demonstrating that attractive people are seen in a positive light for a wide range of attributes compared with unattractive people. Studies on attractiveness stereotypes have generally not addressed the particular characteristics of faces that make individuals either attractive or unattractive, or the features that elicit personality attributions, although different faces reliably elicit the same personality attributions [ ]. Expression certainly has large effects, with, for example, faces shown with smiles rated as more attractive and as having more positive personality traits than neutral faces e. Such facial expressions are transient, however, and will differ rapidly within individuals over time and across photographs. Despite some findings showing a preference for more masculine and dominant male faces e. Personality traits are reported cross-culturally to be among the most important factors in partner choice by both sexes [ 1 , ]. If desired personality is so important, it would appear likely that personality attributions elicited by a face would affect its attractiveness. For example, women who value cooperation and good parenting may avoid masculine-faced men. Thus, instead of feminine faces being attractive and this attractiveness driving positive personality attributions, it may be that the personality attributions are driving the attractiveness judgements. Individuals may use personality stereotypes in mate selection to select partners with a personality that they desire. Some perceptual attributions to facial photographs are somewhat accurate e. Attraction to faces based on personality stereotypes may happen regardless of whether attributions are accurate or not, especially as many individuals do believe that face provides an important guide to character [ , ]. One study has indeed demonstrated that a desire for some personality traits influences judgements of facial attractiveness [ ]. Individuals valuing particular personality traits find faces appearing to display these traits attractive. Conversely, those not valuing particular traits find faces attractive that are perceived to possess that trait less. Thus, desired personality influences perceptions of facial attractiveness in opposite sex faces, changing the result to: In terms of benefits to perceivers, it is easy to see why traits such as appearing trustworthy would make a face appear more attractive. For individual-specific traits, the logic is more complicated, but such preferences could be related to behavioural compatibility within couples, as people do tend to desire partners with personalities similar to their own [ ]. One reason for variability in preferences for male facial masculinity may lie in the personality traits that masculine- and feminine-faced men are assumed to possess. Increasing the masculinity of face shape increased perceptions of dominance, masculinity and age but decreased perceptions of warmth, emotionality, honesty, cooperativeness and quality as a parent [ 83 ]. Indeed, recent work has shown that masculine facial characteristics are associated with indices of physical dominance, such as physical strength [ ], and the perception of such traits [ ], and that feminine men show weaker preferences for short-term relationships and stronger preferences for committed, long-term relationships than their masculine peers do [ ]. Women's face preferences may thus represent a trade-off between the desire for good genes and the desire for a cooperative partner. Of course, the five types of trait listed above are not a complete list of factors involved in the judgement of facial attractiveness. While individual traits impact on attractiveness, there is also scope for interaction between them. Certain face traits also appear to interact in generating preferences, however. For example, preferences for masculinity vary as a function of the healthiness of the face [ 96 ] and women's preferences for facial self-similarity are higher when men are more facially masculine [ ]. Such interactions highlight that facial attractiveness judgements are not simple: In humans, while individuals may share certain basic criteria for finding faces attractive, many factors may influence the specific types of face they find attractive. In this section, we review three broad areas leading to individual differences in preferences: Research suggests that internal factors predict individual differences in several aspects of face perception, including attractiveness judgements. Importantly, the nature of these individual differences suggests adaptive design in face perception and face preferences. In the following section, we discuss two broad types of internal factors: The influence of hormones on face perception is an area that has generated a considerable amount of empirical research in recent years. As detailed previously, masculine characteristics in men's faces are associated with measures of long-term medical health [ 35 , 77 ] and indices of developmental stability [ 36 , 37 ], physical strength [ ] and reproductive potential [ ]. By contrast, feminine characteristics in men's faces are associated with cues of investment and stronger preferences for long-term over short-term sexual relationships e. There is now compelling evidence that how women resolve this trade off between the costs and benefits associated with choosing a masculine mate is affected by hormone levels and fertility. Many studies have reported that women demonstrate stronger preferences for men displaying masculine facial characteristics around ovulation, when women are most fertile, than during other phases of the menstrual cycle [ — ]. Some studies have also reported that these cyclic shifts in women's preferences for masculine characteristics in men's faces are greatest among women who already have romantic partners and when women judge men's attractiveness for short-term, extra-pair relationships [ ]. Although the ultimate function of these cyclic shifts remains somewhat controversial, many researchers have interpreted cyclic shifts in women's masculinity preferences as evidence for adaptations that function to increase offspring health via high paternal investment from a long-term partner while promoting attraction to other men displaying cues of heritable immunity to infectious disease when most fertile discussed in [ ]. Women may gain maximal benefits by selecting investing long-term partners and high-quality extra-pair partners. Importantly, other explanations that have been suggested, such as increased attraction to individuals who appear to be likely sources of high-quality care and support during phases of the menstrual cycle when increased progesterone prepares the body for pregnancy i. Increased attraction to masculine men is by no means unique to face preferences; women also demonstrate stronger attraction to masculine men when judging the attractiveness of men's voices [ — ], body shapes [ ] and body odours [ ], as well as when judging the attractiveness of videoclips of male behavioural displays of dominance [ , ]. Furthermore, converging evidence for fertility-related variation in women's preferences for facial masculinity comes from studies investigating circum-pubertal and circum-menopausal variation in women's masculinity preferences; post-menopausal and pre-pubertal women report weaker preferences for masculine facial characteristics than do their pre-menopausal and post-pubertal counterparts, respectively e. The ultimate function of cyclic shifts in women's preferences for masculine facial characteristics is not the only controversial aspect of cyclic shifts in women's masculinity preferences. For example, although some researchers have suggested that cyclic shifts in women's masculinity preferences may be an artefact of the computer graphic methods that are generally used in these studies to experimentally manipulate sexually dimorphic cues in digital face images [ ], this claim is very difficult to reconcile with findings from studies that have demonstrated cyclic shifts in women's preferences for masculinity in real i. While these findings suggest that cyclic shifts in women's masculinity preferences are not an artefact of the stimuli used, an aspect of research on cyclic shifts in women's masculinity preferences that remains controversial is whether the effect of cycle phase on women's face preferences is relatively specific to judgements of men's faces, or also occurs when women judge the attractiveness of other women. To date, evidence is equivocal; some studies have observed cyclic shifts in women's preferences for masculine-faced men, but not masculine-faced women [ ], while others have observed cyclic shifts in women's preferences for masculine faces, irrespective of their sex [ , ]. These latter papers speculate that cyclic shifts in women's preferences for masculine-faced women could represent a low-cost functionless by-product of a mechanism that evolved primarily to increase women's preferences for masculine men around ovulation [ ], or have suggested that higher attractiveness ratings given to masculine women around ovulation could reflect increased derogation of feminine, and therefore attractive, same-sex competitors when women are most fertile [ ] see also [ ]. In addition to the sex-specificity of the effects of cycle phase on face preferences, the mechanisms that underpin cyclic shifts in women's preferences for masculine characteristics in men's faces have also been a topic of considerable interest in recent years. For example, research into the hormonal mechanisms that might underpin these cyclic shifts has variously emphasized the effects of variation in levels of testosterone [ ], oestrogen [ ] and progesterone [ , ], or has suggested, perhaps unsurprisingly, that cyclic shifts in women's masculinity preferences might be best explained by complex interactions among multiple hormones [ , ]. While findings from research into the hormonal mechanisms that might underpin cyclic shifts in women's masculinity preferences have arguably been inconsistent, the findings of corresponding research into the psychological mechanisms have been relatively consistent; various studies have demonstrated that women are quicker to categorize men and access male stereotypes around ovulation e. These findings suggest that cyclic variations in stereotype access and sexual desire might be important psychological mechanisms for regulating facial masculinity preferences during the menstrual cycle. While research on hormone-mediated face perception has generally focused on women's judgements of men's attractiveness, some recent research has investigated hormone-mediated face preferences among men. Men, of course, do not cycle in the same way women do, but levels of testosterone fluctuate within individuals. Research using natural variation in testosterone has shown that men's preferences for feminine characteristics in women's faces are stronger when their testosterone levels are high than when they are relatively low [ ]. This finding suggests that hormones, such as testosterone, can generate within-participant individual differences in face preference in men. As can be seen from the previous paragraphs, there is compelling evidence that women's preferences for masculine men, be they assessed from face preferences or from preferences for male characteristics in other domains, vary systematically over the menstrual cycle. Whether or not preferences for other putative cues of men's long-term health are similarly affected by menstrual cycle is equivocal, however. For example, although many studies have demonstrated that women's preferences for the body odours of symmetric men are enhanced around ovulation reviewed in [ ] , evidence for cyclic shifts in women's preferences for symmetry in men's faces is inconsistent. One study has found that women's preferences for symmetric male faces were stronger around ovulation than during other phases of the menstrual cycle, at least among partnered women who were instructed to judge men's attractiveness as short-term mates [ ]. By contrast, other studies have observed no evidence for cyclic shifts in women's preferences for symmetric men's faces e. Although evidence that women's preferences for symmetry in men's faces vary systematically over the menstrual cycle is equivocal, that is not to say that robust cyclic shifts in women's perceptions of faces are only evident in their preferences for facial masculinity. For example, women's aversions to self-resembling faces are enhanced around ovulation and positively correlated with women's estimated progesterone levels during the menstrual cycle [ ]. This variation in attitudes to self-resembling faces may reflect increased inbreeding avoidance around ovulation and increased preferences for caring, supportive and trustworthy individuals when increased progesterone prepares the body for pregnancy [ ]. Moreover, women's aversions to facial cues associated with current illness e. Indeed, pregnant women and women using oral contraceptives which mimic the effects of increased progesterone during pregnancy demonstrate stronger aversions to individuals displaying facial cues of illness than do women with natural menstrual cycles [ 97 ]. These latter findings for aversions to facial cues of illness and progesterone during the menstrual cycle complement other research on increased aversions to possible sources of contagion in women's food preferences during pregnancy [ ], as well as increased sensitivity to facial expressions signalling that sources of threat and contagion are nearby when progesterone levels are raised [ , ]. While our discussion of hormone-mediated face preferences in women has emphasized the positive findings that have been reported in the literature, it is important to note that there have also been unsuccessful replications of cyclic variation in women's face preferences. For example, two recent studies observed no evidence for cyclic variations in women's preferences for masculine versus feminine male faces [ , ]. One possible explanation of these null findings comes from findings that suggest the extent to which women's preferences for masculine men vary over the menstrual cycle vary systematically among women. For example, cyclic variation in women's preferences for masculine characteristics in men's voices is significantly greater among women with high trait i. This pattern of results may occur because varying their sexual strategy during the menstrual cycle may benefit unattractive women more than it benefits attractive women [ ]. More recent research has presented additional evidence that women's family background, prenatal hormone levels and mortality salience might also affect the extent to which they vary their masculinity preferences according to their menstrual cycle phase [ — ]. We also note that there are significant methodological differences between studies examining cycle effects, making direct comparisons e. For example, some studies distinguish between short- and long-term mating contexts, generally with larger cyclic shifts for short-term judgements [ ], while others do not [ ]. Studies also differ in stimuli number, stimuli type and how fertility is defined. A thorough description of methodological differences between studies is not the focus here, but methodology is certainly a factor that could explain differences in findings across studies. It is likely that further research concerning individual differences in cyclic shifts and comparing different methodologies would provide important insights into the motivations, functions and mechanisms behind cyclic shifts in fundamental aspects of face perception. While the previous section discussed research implicating hormone levels and fertility in individual differences in face perception, this section will discuss the relationships between face preferences and indices of own condition and attractiveness. Several studies have reported positive correlations between women's ratings of their own physical attractiveness and the strength of their preferences for masculine characteristics in men's faces [ 92 ]. Other studies have extended this work by demonstrating that more objective measures of women's condition and attractiveness, such as their waist—hip ratio or oestrogen levels, predict their preferences for masculine characteristics in men's faces in the same way [ , ]. Similar correlations between indices of women's own attractiveness and the strength of their preferences for masculine characteristics in other domains, such as men's voices, have also been reported [ , ], and indices of women's own condition and attractiveness are positively correlated with the strength of their preferences for symmetry and healthy-looking skin in men's faces [ 92 , ]. The findings described above appear to be somewhat analogous to condition-dependent preferences observed in other species, in which individuals in good physical condition show stronger preferences for high-quality mates e. Condition-dependent preferences in both humans and non-humans may have a common function and occur because individuals in good physical condition i. Particularly compelling evidence for this proposal comes from one of the few experimental studies of condition-dependent mate preferences. These findings suggest that women recalibrate subjective impressions of their own attractiveness i. While the research described above focused on the relationships between mate preferences and both individuals' own physical characteristics and their subjective evaluations of these physical characteristics, other work on condition-dependent preferences has investigated whether personality traits and other psychological factors predict individual differences in mate preferences in similar ways. For example, individual differences in systemizing and sensation-seeking, both of which are components of male sex-typical psychology, are positively correlated with men's preferences for feminine characteristics in women's, but not men's, faces [ , ]. Among women, individual differences in empathy, a component of female sex-typical psychology, and extraversion, a key predictor of social status that is correlated with women's physical attractiveness, are positively correlated with preferences for masculine characteristics in men's, but not women's, faces [ , ]. These findings not only implicate personality traits in individual differences in face preferences but also raise the intriguing possibility that some personality traits might mediate the relationships between an individual's physical characteristics and their face preferences. While factors such as hormones and own attractiveness can explain differences in face preferences between individuals, the context under which judgements are made can also contribute to variation in standards of beauty. In the following section, we discuss how context affects face preferences in three types of contexts: Information about genetic kinship is available in the face and is perceived somewhat accurately [ — ]. Judgements of facial similarity are highly synonymous with judgements of kinship [ — ], and facial similarity produced by computer-graphic manipulation affects behaviour in ways consistent with inclusive fitness theory e. Therefore, responses to facial resemblance are likely to be affected by prosocial versus sexual contexts. Cues of kinship are predicted to increase preferences in non-sexual, prosocial contexts, owing to the benefits associated with inclusive fitness [ ]. In other words, evolutionary models show that behaviours that benefit other individuals who share genes through common descent will be favoured. Therefore, if physical similarity is a reliable cue of genetic relatedness, we expect individuals to act prosocially towards individuals who appear similar to themselves. However, cues of kinship should have a less positive effect in sexual contexts, because of inbreeding's detrimental effects on offspring quality [ ]. One study investigated this prediction by comparing perceptions of the attractiveness of self-resembling own-sex and opposite-sex faces [ ]. Participants judged self-resemblance to be more attractive in the context of own-sex faces than in the context of opposite-sex faces. However, there was no such opposite-sex bias when the same faces were judged for averageness. This own-sex bias in preferences for self-resemblance indicates that, while self-resemblance is attractive in an exclusively prosocial i. Stronger attraction to cues of kinship in own-sex faces than in opposite-sex faces is likely to promote prosocial behaviour towards own-sex kin, while minimizing occurrences of inbreeding with opposite-sex kin. Transforms of self-similarity. Images are made by using the difference between a composite image of the same sex and an individual participant to make faces more similar to the participant. Self-dissimilar faces can be made by applying the same technique but using images other than the participant. Further evidence for context sensitivity in judgements of self-resembling faces is provided by a study comparing men's and women's preferences for self-resemblance in opposite-sex faces in explicitly prosocial versus sexual contexts [ ]. Participants were shown images of self-resembling opposite-sex faces and asked to judge their trustworthiness i. Consistent with both inclusive fitness and inbreeding avoidance theories, self-resemblance increased perceptions of trustworthiness, decreased attractiveness for short-term relationships and had no significant effect on attractiveness for long-term relationships. The fact that self-resemblance in opposite-sex faces was found to be trustworthy, but not attractive in short-term contexts, emphasizes the context-sensitivity of responses to self-resemblance. Importantly, because familiarity increases judgements of both attractiveness and trustworthiness [ ], this pattern of context-sensitivity strongly suggests that responses to self-resemblance do not occur simply because of familiarity alone i. Another example of social context influencing face preferences comes from research on interactions among the effects of different facial characteristics on preferences. For example, both behavioural and neurobiological evidence suggest that viewers demonstrate stronger attraction to attractive physical cues in faces e. Similarly, behavioural and neurobiological evidence also suggests that viewers demonstrate stronger attraction to cues associated with positive social interest e. Conway et al. Similarly, the putative costs of low investment are much less of a concern in short-term than in long-term relationships and, thus, women may demonstrate stronger masculinity preferences when judging men's attractiveness as possible short-term than long-term partners. Little et al. Women who were not using oral contraceptives made this face more masculine in the context of a short-term relationship than in the context of a long-term relationship. Penton-Voak et al. One potential explanation for this pattern of preference is that attractive women are better able to compete for, retain or replace high-quality, masculine partners and, therefore, do not show as large a shift in their preferences between short-term and long-term contexts. The effects of temporal context on judgements of attractiveness are not limited to faces. Women prefer lower pitched male voices in the context of a short-term relationship than in the context of a long-term relationship [ ]. This same study also found that the effect of relationship context was greatest when women were in the fertile phase of the menstrual cycle, a finding that is consistent with research on cyclic shifts in preferences for facial masculinity [ ]. A strong theoretical prediction of the trade-off account of variability in women's preferences for masculine men is that women in environments where poor health is particularly harmful to survival e. Regional differences in pathogen prevalence have been shown to be positively correlated with the importance placed on physical beauty and health [ , ]. DeBruine et al. This relationship remained significant, even when controlling for regional variation in wealth and women's mating strategies i. Brooks et al. Masculine-faced men may be favoured under such conditions, for example, as they may be better able to compete for resources. A further study of US states, in contrast, has shown that environmental health factors, and not indices of male—male violence such as homicide rates, predicts regional variation in women's masculinity preferences [ ]. Health, wealth and male—male violence are, of course, inter-related. While it is ultimately possible that health, wealth and male—male violence may all individually contribute to variation in preference, it is important to note that all of these analyses show that regional variation in women's masculinity preferences occurs in ways that are highly consistent with trade-off theories of sexual selection. The availability of resources in an environment may also influence face preferences. In low-resource environments, the resources to raise a child may be scarce or difficult to acquire and a preference for an investing partner be adaptive. To test these ideas, Little et al. Both men and women decreased their preferences for high quality mates for long-term relationships in the context of a harsh environment. This is consistent with the logic of trading genetic quality for commitment and investment in environments where resources are scarce. Individuals are confronted with a myriad of faces and social interactions every day. Research has shown that such experience leads to changes in preferences for faces. In the following section, we discuss two aspects of visual experience examining: Familiarity is a powerful determinant of attraction. For many types of stimuli, including faces, exposure increases attraction even when the exposure is unconscious [ — ]. Structural features of the face must be stored and represented in order to determine familiarity. As noted earlier, one idea for why averageness in faces is attractive comes from a link with familiarity—as average faces appear familiar this could positively affect their attractiveness [ 60 , 62 ]. Familiarity, when not paired with aversive stimuli, is thought to be rewarding [ ], and indeed there are obvious benefits to avoiding the unfamiliar. This can then help explain why exposure may cause increases in preference. There may, however, be more to increasing face preference than simple exposure. For example, recent studies have demonstrated that the nature of association positive or negative can affect face preferences, with positive experiences leading to increased attraction and negative experiences to decreased attraction [ ]. Moreover, these effects of valenced exposure are not bound solely to the specific individuals who were encountered and generalize to judgements of novel, physically similar individuals [ ]. Familiarity with parental traits has been implicated in human preferences. The phenomenon of imprinting, whereby individuals are attracted to parental traits, is well-studied in non-human animals [ , ] and there is increasing evidence for similar effects in humans. Following studies of facial similarity, judges have been shown to correctly match wives to their mother-in-law at a significantly higher rate than expected by chance and that wife—mother-in-law similarity is higher than similarity between husbands and their wives [ ]. Such effects are also seen in adopted daughters, controlling for any potential genetic effects, with significant facial resemblance between daughter's husband and her adoptive father [ ]. Other studies have shown that, for hair and eye colour, the best predictors of partner traits are the opposite-sex parent's colour traits [ ] and that individuals are attracted to age in faces consistent with the age of their parents when they were born [ ]. It is worth noting that at least in one study, effects were seen mainly for the opposite-sex parent [ ], which may indicate a more complex mechanism than simple exposure. Another line of argument suggesting imprinting-like effects appear not simply to reflect exposure comes from studies that have shown effects to be dependent on the quality of the relationship to the parent [ , ]. For example, daughters who report that they received greater emotional support from their adoptive fathers are more likely to choose mates who are similar to their father than individuals who report their father provided less emotional support [ ]. Similarly, women who rate their childhood relationships with their father positively show stronger attraction to face proportions similar to their father's face than women who rate their relationships less well [ ]. Imprinting-like effects then appear more complicated than simple exposure being directed more to one parent than the other and showing dependence on the relationship with that parent. Imprinting-like effects may lead to positive assortative mating pairing with similar partners , at least for long-term relationships, and this may have benefits in terms of keeping adaptive suites of genes together [ ] or increasing behaviour compatibility [ ]. There is certainly evidence that couples resemble each other facially [ , ]. Potentially then, a system that learns about known individuals and increases attraction to their face traits could be adaptive. Both familiarity and imprinting posit that exposure affects attractiveness. In recent years, exposure has been thought to have specific effects on our representations of faces via visual adaptation. We are unlikely to have an inbuilt average face and what is average must be calculated from experience. For each class of stimuli, the human visual system encounters may develop an individual representation, or prototype, made up of an average of the characteristics of all the different stimuli of that type that have been seen [ — ]. Computer modelling has revealed that algorithms trained to discriminate different stimuli produce stronger responses to stimuli that represent the average of the training set, even though this average was not previously encountered [ , ]. These findings have been interpreted as evidence that prototype formation is a property of learning to recognize different stimuli as members of a class [ , ]. Studies on category learning have a long history e. Learning studies examine how categorical perception develops using abstract stimuli. Such an approach has highlighted face traits such as age, health, symmetry, and averageness, which are proposed to be associated with benefits and so associated with facial attractiveness. This view may postulate that some traits will be universally attractive; however, this does not preclude variation. Indeed, it would be surprising if there existed a template of a perfect face that was not affected by experience, environment, context, or the specific needs of an individual. Research on facial attractiveness has documented how various face traits are associated with attractiveness and various factors that impact on an individual's judgments of facial attractiveness. 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Cyclic variation in women's preferences for masculine traits: Feinberg D. Menstrual cycle, trait estrogen level, and masculinity preferences in the human voice. Mating context and menstrual phase affect women's preferences for male voice pitch. Preferences for masculinity in male bodies change across the menstrual cycle. Havlicek J. Women's preference for dominant male odour: Changes in women's mate preferences across the ovulatory cycle. Women's preferences for male behavioral displays change across the menstrual cycle. Women's preferences for masculinity in male faces are highest during reproductive age range and lower around puberty and post-menopause. Psychoneuroendocrinology 35 , — Vukovic J. Circum-menopausal effects on women's judgements of facial attractiveness. Peters M. Preferences across the menstrual cycle for masculinity and symmetry in photographs of male faces and bodies. Preferences for variation in masculinity in real male faces change across the menstrual cycle: Welling L. Raised salivary testosterone in women is associated with increased attraction to masculine faces. Roney J. Women's estradiol predicts preference for facial cues of men's testosterone. Johnston L. The need for speed: Macrae C. Person perception across the menstrual cycle: Sex drive is positively associated with women's preferences for sexual dimorphism in men's and women's faces. Exposure to sexually attractive men decreases women's preferences for feminine faces. Men report stronger attraction to femininity in women's faces when their testosterone levels are high. Preferences for symmetry in faces change across the menstrual cycle. Oinonen K. Facial symmetry detection ability changes across the menstrual cycle. Symmetry and sexual dimorphism in human faces: Women's attractiveness judgments of self-resembling faces change across the menstrual cycle. Social perception of facial resemblance in humans. Sexual Behav. Pepper G. Rates of nausea and vomiting in pregnancy and dietary characteristics across populations. Derntl B. Emotion recognition accuracy in healthy young females is associated with cycle phase. Conway C. Salience of emotional displays of danger and contagion in faces is enhanced when progesterone levels are raised. Harris C. Menstrual cycle and facial preferences reconsidered. Sex Roles. Scarbrough P. Individual differences in women's facial preferences as a function of digit ratio and mental rotation ability. Father absence, parent—daughter relationships and partner preferences. Vaughn J. The effect of mortality salience on women's judgments of male faces. Waist—hip ratio predicts women's preferences for masculine male faces, but not perceptions of men's trustworthiness. Female condition influence preferences for sexual dimorphism in faces of male humans Homo sapiens. Psychol , — Self-rated attractiveness predicts individual differences in women's preferences for masculine men's voices. Women's own voice pitch predicts their preferences for masculinity in men's voices. Women's physical and psychological condition independently predict their preference for apparent health in faces. Bakker T. Condition-related mate-choice in sticklebacks. Individual differences in empathizing and systemizing predict variation in face preferences. Sensation seeking and men's face preferences. Extraversion predicts individual differences in women's face preferences. Alvergne A. Differential facial resemblance of young children to their parents: Nesse R. Sex-differences in ability to recognize family resemblance. Bressan P. Parental resemblance in 1-year-olds and the Gaussian curve. Talis pater, talis filius: Kaminski G. Human ability to detect kinship in strangers' faces: Dal Martello M. Where are kin recognition signals in the human face. Maloney L. 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Mothers determine male sexual preferences. Lorenz K. The innate forms of potential experience. Zietschrift Tierpsychol. Bereczkei T. Homogamy, genetic similarity, and imprinting; parental influence on mate choice preferences. Sexual imprinting in human mate choice. Facial attractiveness judgements reflect learning of parental age characteristics. Wiszewska A. Father—daughter relationship as a moderator of sexual imprinting: Thiessen D. Human assortative mating and genetic equilibrium: Hill C. Breakups before marriage: Issues 32 , — Assortative mating for perceived facial personality traits. Hinsz V. Facial resemblance in engaged and married couples. Bateson P. Optimal outbreeding. In Mate choice ed. Cambridge University Press [ Google Scholar ]. Enquist M. Symmetry, beauty and evolution. Giese M. Physiologically inspired neural model for the encoding of face spaces. Neurocomputing 65 , 93— Johnstone R. 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Not by genes alone:.

By contrast, feminine characteristics in men's faces are associated with cues of investment and stronger preferences for long-term over short-term sexual relationships e. There is now compelling evidence that how women resolve this trade off between the costs and benefits associated with choosing a masculine mate is affected by hormone levels and fertility.

Many studies have reported that women demonstrate stronger preferences for men displaying masculine facial characteristics around ovulation, when women are most fertile, than during other phases of the menstrual cycle [ — ]. Some studies have also reported that these cyclic shifts in women's preferences for masculine characteristics in men's faces are greatest among women who already have romantic partners and when women judge men's attractiveness for short-term, extra-pair relationships [ ].

Although the ultimate function of these cyclic shifts remains somewhat controversial, many researchers have interpreted cyclic shifts in women's masculinity preferences as evidence for adaptations that function to increase Facial attractiveness milffaceness health via high paternal investment from a long-term partner while promoting attraction to other men displaying cues of heritable immunity to infectious disease when most fertile discussed in [ ].

Women may gain maximal benefits by selecting investing long-term partners and high-quality extra-pair partners. Importantly, other explanations that have been suggested, such as increased attraction to individuals who appear to be likely sources of high-quality care and support during phases of the menstrual cycle when increased progesterone prepares the body for pregnancy i.

Increased attraction to masculine men is by no means unique to face preferences; women Facial attractiveness milffaceness demonstrate stronger attraction to click men when judging the attractiveness of men's voices [ — ], body shapes [ ] and body odours [ Facial attractiveness milffaceness, as well as when judging the attractiveness of videoclips Facial attractiveness milffaceness male behavioural displays of dominance [].

Furthermore, converging evidence for fertility-related variation in women's preferences for facial masculinity comes from studies investigating circum-pubertal and circum-menopausal variation in women's masculinity preferences; post-menopausal and pre-pubertal women report weaker preferences for masculine facial characteristics than do their pre-menopausal and post-pubertal counterparts, respectively e. The ultimate function of cyclic shifts in women's preferences for masculine facial characteristics is not the only controversial aspect of cyclic shifts in women's masculinity preferences.

For example, although some researchers have suggested that cyclic shifts in women's masculinity preferences may be an artefact of the computer graphic methods Facial attractiveness milffaceness are generally used in these studies to experimentally manipulate sexually dimorphic cues in digital face images [ ], this claim is very difficult to reconcile with findings from studies that have Facial attractiveness milffaceness cyclic shifts in women's preferences for masculinity in real i.

While these findings suggest Facial attractiveness milffaceness cyclic shifts in women's masculinity preferences are not an artefact of the stimuli used, an aspect of research on cyclic shifts in women's masculinity preferences that remains controversial is whether the effect of cycle phase on women's face preferences is relatively specific to judgements of men's faces, or also occurs when Facial attractiveness milffaceness judge the attractiveness of other women.

To date, evidence Facial attractiveness milffaceness equivocal; some studies have observed cyclic shifts in women's preferences for masculine-faced men, but not masculine-faced women [ ], while others have observed cyclic shifts in women's preferences for masculine faces, irrespective of their sex []. These latter papers speculate that cyclic shifts in women's preferences for masculine-faced women could represent a low-cost functionless by-product of a mechanism that evolved primarily to increase women's preferences for masculine men around ovulation [ ], or have suggested that higher attractiveness ratings given to masculine women Facial attractiveness milffaceness ovulation could reflect click at this page derogation of feminine, and therefore attractive, same-sex competitors when women are most fertile [ ] see also [ ].

In addition to the sex-specificity of the effects of cycle phase on face preferences, the mechanisms that underpin cyclic Facial attractiveness milffaceness in women's preferences for masculine characteristics in men's faces have also Facial attractiveness milffaceness a topic of considerable interest in recent years.

Facial attractiveness.

For example, research into the hormonal mechanisms that might underpin these cyclic shifts has variously emphasized the effects of Facial attractiveness milffaceness in levels of testosterone [ ], oestrogen [ ] and progesterone [], or has suggested, perhaps unsurprisingly, Nina nude cyclic shifts in women's masculinity preferences might be best explained by complex interactions among multiple hormones [].

While findings from research into the hormonal mechanisms that might underpin cyclic shifts in women's masculinity preferences have arguably been inconsistent, the Facial attractiveness milffaceness of corresponding research into the psychological mechanisms have been relatively consistent; various studies have demonstrated that women are quicker to categorize men and access male stereotypes around ovulation e.

These findings suggest Facial attractiveness milffaceness cyclic variations in stereotype access and sexual desire might be important psychological mechanisms for regulating facial masculinity preferences during the menstrual cycle. While research on Facial attractiveness milffaceness face perception has generally focused on women's judgements of men's attractiveness, some recent research has investigated hormone-mediated face preferences among men. Men, of course, do not cycle in the same way women do, but levels of testosterone fluctuate within individuals.

Research using natural variation in testosterone has shown that men's preferences for feminine characteristics in women's faces are stronger when their testosterone levels are high than when they are relatively low [ ].

Facial attractiveness milffaceness finding suggests that hormones, such as testosterone, can generate within-participant individual differences in face preference in men. Facial attractiveness milffaceness

Xxxxxvif Vidaos Watch Video Profail photo. We note that much research has focused on women's preferences, although most traits are also relevant for men. Symmetry refers to the extent to which one-half of an object image, organism, etc. Individuals differ in their ability to maintain the stable development of their morphology under the prevailing environmental conditions under which that development is taking place [ 28 , 29 ]. The ability of an individual to develop successfully in the face of environmental pressures is therefore one proposed indicator of genetic quality. A character demonstrates fluctuating asymmetry FA when symmetry reflects the normal development, and deviations from this symmetry are randomly distributed with respect to side [ 30 ]. FA is a particularly useful measure of developmental stability because we know that the optimal developmental outcome is symmetry. Therefore, any deviation from perfect symmetry can be considered a sub-optimal solution which will result in performance problems in the future. FA is also a useful measure as it subsumes a huge amount of individual variation in development, being the outcome of differences in genetic e. Preferences for symmetry can then, potentially, provide both direct e. Whether symmetry is actually related to quality in other animals and humans is an issue addressed by a large literature, and a complete review is not the focus of this paper. While the issue is divided, and there is some evidence that symmetry is not associated with quality e. In humans, male body symmetry is positively related to sperm number per ejaculate and sperm speed [ 32 ] and female breast symmetry is positively correlated with fecundity [ 33 , 34 ]. Relating to faces, one study has demonstrated that facial asymmetry is positively related to self-reported number of occurrences of respiratory disease [ 35 ] and some studies have observed positive correlations between symmetry and other putative indices of underlying physical condition i. The relationship between symmetry and quality is not reviewed in detail here, but it should be noted that fitness-related characteristics, such as growth rate, fecundity and survivability, are positively associated with symmetry across a number of species and taxa e. Studies of naturally occurring human facial asymmetries also provide evidence that symmetry is found attractive, though such studies can be confounded by potential correlates. Studies measuring symmetry from unmanipulated faces have reported positive correlations with rated attractiveness [ 40 — 44 ] and one study has even demonstrated that with pairs of monozygotic twins, the twin with more symmetric measurements is seen as more attractive [ 45 ]. Studies using more sophisticated symmetry manipulations have demonstrated that symmetry can have a positive influence on attractiveness [ 47 , 48 ] and have established that the chimeric manipulations used in the early studies introduced unnatural proportions into the symmetric faces see [ 48 ]. Thus, the methodologically superior computer graphic studies [ 47 , 48 ] parallel the findings of investigations into naturally occurring facial asymmetries [ 40 , 41 , 43 — 45 ]. The computer graphic studies demonstrate that increasing symmetry alone is sufficient to increase attractiveness. Subsequently, other studies have replicated preferences for symmetry using manipulated stimuli in different Western samples e. Preferences for symmetry using manipulated faces have been found in African hunter—gatherers [ 51 ], and macaque monkeys gaze longer at symmetrical than at asymmetrical face images of conspecifics [ 52 ]. Symmetry and asymmetry. Symmetric images are usually preferred to asymmetric images. Importantly, recent studies have implicated perceptions of health in attraction to symmetric faces [ 44 , 53 ] and have suggested that the mechanisms underpinning preferences for symmetric faces are different from those that might drive preferences for symmetry in mate-choice-irrelevant stimuli e. Such findings suggest that preferences for symmetric faces reflect, at least in part, adaptations for mate choice. Averageness refers to how closely a face resembles the majority of other faces within a population; non-average faces have more extreme characteristics than the average of a population. Average faces may be attractive because an alignment of features that is close to a population average is linked to genetic diversity [ 54 , 55 ]. Parasites are generally best adapted to proteins that are common in the host population; hence, parasites are adapted to the genes that code for the production of these proteins. A second evolutionary theory for the attractiveness of averageness in faces is that extreme non-average genotypes are more likely to be homozygous for deleterious alleles, that is, to be more likely to possess genes that are detrimental to an individual than those with more average genotypes [ 54 ]. Both of these theories propose evolutionary benefits to mating with individuals possessing average faces. Recent studies have supported the link between averageness, heterozygosity i. Heterozygosity in the major histocompatibility complex MHC genes that code for proteins involved in immune response, is positively associated with facial attractiveness [ 56 ] and facial averageness [ 57 ]. More directly, another study has shown that facial averageness is positively related to medical health as measured from actual medical records in both men and women [ 58 ]. There is good evidence that average faces are indeed found attractive. Galton [ 59 ] first noted that multiple faces blended together were more attractive than the constituent faces. Recent studies have improved upon these techniques using computers to create digitally blended composite faces; generally, the more images in a composite, the more attractive it is found [ 60 — 62 ]. Aside from composite images, Light et al. Average faces are generally more symmetric and symmetry is typically attractive in faces discussed in more detail above. Several studies have controlled for this confound in the original studies. When averageness and symmetry were independently manipulated, one study found that both manipulations positively and independently influenced attractiveness judgements [ 65 ]. Other studies have used perfectly symmetric images manipulated in averageness and still have demonstrated preferences for averageness [ 66 , 67 ]. Indeed, by comparing preferences for averageness when the effects of symmetry were controlled for and were not controlled for, Jones et al. It has also been noted that, in the original composite studies, the more images that are blended together the smoother the skin texture becomes, as imperfections such as lines or blemishes are averaged [ 68 ]. Image c should be more attractive than both of the other images. Composites are made by marking key locations around the main facial features e. The average location of each point of the component faces is then calculated to define the shape of the composite. The images of the individual faces are then warped to the relevant average shape before superimposing the images to produce a photographic quality composite image. While the majority of the work described above has been carried out in North America, Britain and Australia, averageness has also been found to be attractive across different cultures. For example, facial averageness is also found attractive in Japanese participants [ 69 ] and in African hunter—gatherers [ 67 ]. Male and female faces differ in their shape. Mature features in adult human faces reflect the masculinization or feminization of secondary sexual characteristics that occurs at puberty. These face shape differences, in part, arise because of the action of hormones such as testosterone. Larger jawbones, more prominent cheekbones and thinner cheeks are all features of male faces that differentiate them from female faces e. From an evolutionary view, extremes of secondary sexual characteristics more feminine for women, more masculine for men are proposed to be attractive because they advertise the quality of an individual in terms of heritable benefits; they indicate that the owners of such characteristics possess good genes. In other words, such traits advertise the possession of genes that are beneficial to offspring inheriting them in terms of survival or reproduction. One explanation of the importance of these facial traits is that they represent a handicap to an organism [ 71 ] and the costs of growing the trait means that only healthy individuals can afford to produce them. For example, secondary sexual characteristics are proposed to be linked to parasite resistance because the sex hormones that influence their growth, particularly testosterone, lower immunocompetence. Testosterone has been linked to the suppression of immune function in many species [ 72 ], including humans [ 73 , 74 ]. Larger secondary sexual characteristics should be related to a healthier immune system because only healthy organisms can afford the high sex-hormone handicap on the immune system that is necessary to produce these characteristics [ 75 ]. In many non-human animal studies, there is a positive association between secondary sexual trait expression and immunocompetence e. A study by Rhodes et al. No relationship was found between femininity and actual health in female faces, though [ 77 ]. Another study has demonstrated that men's facial masculinity and women's facial femininity are negatively related to self reports of respiratory disease [ 35 ]. If health is heritable, then female preferences for masculinity and male preferences for femininity may indeed also reflect the choice of mates with good genes. There is also a link between hormonal profile and face shape. Women with higher circulating oestrogen have more feminine faces [ 78 ], while men with high testosterone have more masculine faces [ 79 ], but see also [ 80 ]. If women with high oestrogen and men with high testosterone are valued as mates, preferences for cues of hormonal profile could drive preferences for sexually dimorphic face shape. Masculinity is transformed using the difference between male and female face shape as defined by creating a male and female composite. Preferences for masculinity in male faces vary across studies, but feminine female faces are consistently found more attractive than masculine female faces. There is considerable evidence that feminine female faces are considered attractive. Studies measuring facial features from photographs of women [ 40 , 81 , 82 ] and studies manipulating facial composites [ 83 ] all indicate that feminine features increase the attractiveness of female faces across different cultures. If oestrogenized female faces provide cues to fertility and health, then male preferences for such features are potentially adaptive. This reasoning does not require oestrogen to be immunosuppressive or part of a handicap. The link between masculinity and attractiveness in male faces is less clear. Cunningham et al. Despite some findings showing a preference for more masculine and dominant faces, several studies have shown that feminine characteristics and faces of low dominance are of increased attractiveness [ 62 , 83 , 84 , 89 — 91 ]. Many studies have made use of computer graphic techniques to manipulate masculinity. Sexual dimorphism in face shape can be manipulated by taking the geometrical differences between average male and female face shapes and applying this difference to new faces, making more or less masculine or feminine versions [ 83 ]. This process simultaneously changes all dimorphic shape characteristics in the face. Perrett et al. For the male face stimuli, the shape selected by Caucasians as most attractive was significantly feminized for both the Caucasian male face and the Japanese male face continua. Similarly, Japanese participants also selected significantly feminized versions of the male stimuli for both the Japanese and Caucasian male face continua. Thus, in both cultures it was found that participants showed a preference for feminized male faces. Since then, several studies have also documented preferences for femininity [ 62 , 90 , 92 , 93 ], but some similar computer graphic studies have also reported preferences for masculinity [ 94 , 95 ]. Although some of this variation may be attributed to other characteristics of the faces that varied between sets of stimuli [ 96 ], this does not explain the variability in preferences. We discuss the sources of individual differences in preferences for sexually dimorphic shape cues in the latter sections of our article. The face traits discussed so far have often been measured and manipulated but also studied in terms of perception and related to attractiveness. The reasoning for why traits like symmetry are preferred is often related to underlying health. Thus, it is important to examine perceptions of facial health directly. Perceived health is difficult to relate to any one metric, but people will readily rate faces for perceived health and show very high agreement on such ratings e. In evolutionary terms, there is a large and obvious selective advantage in detecting healthy partners both for social exchange and mate choice. Indeed, while the role of health in mate preferences is clear see below , recent work has demonstrated that participants are more willing to reciprocate trust from healthy-looking social partners than from social partners who are relatively unhealthy-looking [ 98 ]. Such findings demonstrate the importance of health perceptions for social interaction generally. Again, as for previous traits, there may be both direct and indirect benefits to partnering with individuals who are perceived to be healthy. Facial healthiness. High healthiness is associated with higher ratings of attractiveness. There have been several studies that have addressed how facial appearance relates to the healthiness of an individual in humans. The three traits discussed above are often manipulated by changing only face shape, but health perception appears to be related to facial colour and texture also. Fewer studies have examined how colour and texture of faces influence attractiveness judgements. One study has examined how well ratings of health from small patches of skin of faces are related to overall rated attractiveness when the whole face image is available. Jones et al. In other research, homogeneity of skin colour was positively related to attractiveness [ 99 ]. Findings have also suggested that more heterozygous men also have healthier appearing skin [ 56 ]. Skin health may be a particularly useful marker of current health condition as it is more changeable than aspects such as symmetry or averageness. Coloration is directly related to the appearance of skin. Coloration also appears to be an important component of sexual selection in many species. Red coloration is associated with dominance in fish [ ], birds [ ] and non-human primates [ , ] and, consequently, is linked to attracting the opposite sex. It has been noted that primates with trichromatic vision are generally bare-faced [ ] and that, at least in humans, facial flushing is associated with anger and confrontation [ ]. In research on non-human primates, there has been much interest in colour. For example, experimental manipulation of colour shows that female rhesus macaques prefer images of redder male faces [ ], while males prefer images of redder female hindquarters [ ]. In mandrills, red facial colour is related to rank in males [ ], and females sexually present more frequently to brighter males and also groom them more frequently [ ]. Red coloration also has consequences for behaviour in other species. For example, in bird species, the addition of red to stimuli can increase social dominance [ ]. In humans, it has been shown that wearing red in a variety of physically competitive sports is associated with an increased chance of winning over opponents [ ]. This has been interpreted as natural associations of red with dominance being extended to artificially displayed red in the same way that artificial stimuli can exploit innate responses to natural stimuli [ , ]. One study pitting red versus blue shapes found that red shapes were seen as more aggressive, dominant and more likely to win in physical competitions [ ]. Red does generally seem to have aversive effects on human behaviour. For example, when taking exams, individuals move their body away from tests with red covers more than they do from those with green or grey covers [ ]. While these studies suggest the colour red may be seen as a threatening stimulus in humans, red also appears to enhance attraction in some instances. For example, women are seen as more attractive by men when presented with red backgrounds or with red clothing, relative to other colours [ ]. This effect appears to be specific to attractiveness judgements; red colour does not influence judgements of other traits such as kindness or intelligence and does not influence women's attractiveness judgements of other women [ ]. Further research has examined red coloration in faces and demonstrated a positive association with perceived health [ ]. The authors suggest that perception of healthy, oxygenated blood may drive associations between red and healthiness. Alongside redness, people also appear to think that skin yellowness is associated with healthy appearance in faces [ ]. Yellowness may advertise health via an association with diet, as carotenoids are associated with skin yellowness and are absorbed via the intake of fruit and vegetables [ ]. In a classic social psychology study, Dion et al. For example, attractive individuals were thought to be able to achieve more prestigious occupations, be more competent spouses with happier marriages and have better prospects for personal fulfilment. There has been a wealth of studies examining this attractiveness stereotype, demonstrating that attractive people are seen in a positive light for a wide range of attributes compared with unattractive people. Studies on attractiveness stereotypes have generally not addressed the particular characteristics of faces that make individuals either attractive or unattractive, or the features that elicit personality attributions, although different faces reliably elicit the same personality attributions [ ]. Expression certainly has large effects, with, for example, faces shown with smiles rated as more attractive and as having more positive personality traits than neutral faces e. Such facial expressions are transient, however, and will differ rapidly within individuals over time and across photographs. Despite some findings showing a preference for more masculine and dominant male faces e. Personality traits are reported cross-culturally to be among the most important factors in partner choice by both sexes [ 1 , ]. If desired personality is so important, it would appear likely that personality attributions elicited by a face would affect its attractiveness. For example, women who value cooperation and good parenting may avoid masculine-faced men. Thus, instead of feminine faces being attractive and this attractiveness driving positive personality attributions, it may be that the personality attributions are driving the attractiveness judgements. Individuals may use personality stereotypes in mate selection to select partners with a personality that they desire. Some perceptual attributions to facial photographs are somewhat accurate e. Attraction to faces based on personality stereotypes may happen regardless of whether attributions are accurate or not, especially as many individuals do believe that face provides an important guide to character [ , ]. One study has indeed demonstrated that a desire for some personality traits influences judgements of facial attractiveness [ ]. Individuals valuing particular personality traits find faces appearing to display these traits attractive. Conversely, those not valuing particular traits find faces attractive that are perceived to possess that trait less. Thus, desired personality influences perceptions of facial attractiveness in opposite sex faces, changing the result to: In terms of benefits to perceivers, it is easy to see why traits such as appearing trustworthy would make a face appear more attractive. For individual-specific traits, the logic is more complicated, but such preferences could be related to behavioural compatibility within couples, as people do tend to desire partners with personalities similar to their own [ ]. One reason for variability in preferences for male facial masculinity may lie in the personality traits that masculine- and feminine-faced men are assumed to possess. Increasing the masculinity of face shape increased perceptions of dominance, masculinity and age but decreased perceptions of warmth, emotionality, honesty, cooperativeness and quality as a parent [ 83 ]. Indeed, recent work has shown that masculine facial characteristics are associated with indices of physical dominance, such as physical strength [ ], and the perception of such traits [ ], and that feminine men show weaker preferences for short-term relationships and stronger preferences for committed, long-term relationships than their masculine peers do [ ]. Women's face preferences may thus represent a trade-off between the desire for good genes and the desire for a cooperative partner. Of course, the five types of trait listed above are not a complete list of factors involved in the judgement of facial attractiveness. While individual traits impact on attractiveness, there is also scope for interaction between them. Certain face traits also appear to interact in generating preferences, however. For example, preferences for masculinity vary as a function of the healthiness of the face [ 96 ] and women's preferences for facial self-similarity are higher when men are more facially masculine [ ]. Such interactions highlight that facial attractiveness judgements are not simple: In humans, while individuals may share certain basic criteria for finding faces attractive, many factors may influence the specific types of face they find attractive. In this section, we review three broad areas leading to individual differences in preferences: Research suggests that internal factors predict individual differences in several aspects of face perception, including attractiveness judgements. Importantly, the nature of these individual differences suggests adaptive design in face perception and face preferences. In the following section, we discuss two broad types of internal factors: The influence of hormones on face perception is an area that has generated a considerable amount of empirical research in recent years. As detailed previously, masculine characteristics in men's faces are associated with measures of long-term medical health [ 35 , 77 ] and indices of developmental stability [ 36 , 37 ], physical strength [ ] and reproductive potential [ ]. By contrast, feminine characteristics in men's faces are associated with cues of investment and stronger preferences for long-term over short-term sexual relationships e. There is now compelling evidence that how women resolve this trade off between the costs and benefits associated with choosing a masculine mate is affected by hormone levels and fertility. Many studies have reported that women demonstrate stronger preferences for men displaying masculine facial characteristics around ovulation, when women are most fertile, than during other phases of the menstrual cycle [ — ]. Some studies have also reported that these cyclic shifts in women's preferences for masculine characteristics in men's faces are greatest among women who already have romantic partners and when women judge men's attractiveness for short-term, extra-pair relationships [ ]. Although the ultimate function of these cyclic shifts remains somewhat controversial, many researchers have interpreted cyclic shifts in women's masculinity preferences as evidence for adaptations that function to increase offspring health via high paternal investment from a long-term partner while promoting attraction to other men displaying cues of heritable immunity to infectious disease when most fertile discussed in [ ]. Women may gain maximal benefits by selecting investing long-term partners and high-quality extra-pair partners. Importantly, other explanations that have been suggested, such as increased attraction to individuals who appear to be likely sources of high-quality care and support during phases of the menstrual cycle when increased progesterone prepares the body for pregnancy i. Increased attraction to masculine men is by no means unique to face preferences; women also demonstrate stronger attraction to masculine men when judging the attractiveness of men's voices [ — ], body shapes [ ] and body odours [ ], as well as when judging the attractiveness of videoclips of male behavioural displays of dominance [ , ]. Furthermore, converging evidence for fertility-related variation in women's preferences for facial masculinity comes from studies investigating circum-pubertal and circum-menopausal variation in women's masculinity preferences; post-menopausal and pre-pubertal women report weaker preferences for masculine facial characteristics than do their pre-menopausal and post-pubertal counterparts, respectively e. The ultimate function of cyclic shifts in women's preferences for masculine facial characteristics is not the only controversial aspect of cyclic shifts in women's masculinity preferences. For example, although some researchers have suggested that cyclic shifts in women's masculinity preferences may be an artefact of the computer graphic methods that are generally used in these studies to experimentally manipulate sexually dimorphic cues in digital face images [ ], this claim is very difficult to reconcile with findings from studies that have demonstrated cyclic shifts in women's preferences for masculinity in real i. While these findings suggest that cyclic shifts in women's masculinity preferences are not an artefact of the stimuli used, an aspect of research on cyclic shifts in women's masculinity preferences that remains controversial is whether the effect of cycle phase on women's face preferences is relatively specific to judgements of men's faces, or also occurs when women judge the attractiveness of other women. To date, evidence is equivocal; some studies have observed cyclic shifts in women's preferences for masculine-faced men, but not masculine-faced women [ ], while others have observed cyclic shifts in women's preferences for masculine faces, irrespective of their sex [ , ]. These latter papers speculate that cyclic shifts in women's preferences for masculine-faced women could represent a low-cost functionless by-product of a mechanism that evolved primarily to increase women's preferences for masculine men around ovulation [ ], or have suggested that higher attractiveness ratings given to masculine women around ovulation could reflect increased derogation of feminine, and therefore attractive, same-sex competitors when women are most fertile [ ] see also [ ]. In addition to the sex-specificity of the effects of cycle phase on face preferences, the mechanisms that underpin cyclic shifts in women's preferences for masculine characteristics in men's faces have also been a topic of considerable interest in recent years. For example, research into the hormonal mechanisms that might underpin these cyclic shifts has variously emphasized the effects of variation in levels of testosterone [ ], oestrogen [ ] and progesterone [ , ], or has suggested, perhaps unsurprisingly, that cyclic shifts in women's masculinity preferences might be best explained by complex interactions among multiple hormones [ , ]. While findings from research into the hormonal mechanisms that might underpin cyclic shifts in women's masculinity preferences have arguably been inconsistent, the findings of corresponding research into the psychological mechanisms have been relatively consistent; various studies have demonstrated that women are quicker to categorize men and access male stereotypes around ovulation e. These findings suggest that cyclic variations in stereotype access and sexual desire might be important psychological mechanisms for regulating facial masculinity preferences during the menstrual cycle. While research on hormone-mediated face perception has generally focused on women's judgements of men's attractiveness, some recent research has investigated hormone-mediated face preferences among men. Men, of course, do not cycle in the same way women do, but levels of testosterone fluctuate within individuals. Research using natural variation in testosterone has shown that men's preferences for feminine characteristics in women's faces are stronger when their testosterone levels are high than when they are relatively low [ ]. This finding suggests that hormones, such as testosterone, can generate within-participant individual differences in face preference in men. As can be seen from the previous paragraphs, there is compelling evidence that women's preferences for masculine men, be they assessed from face preferences or from preferences for male characteristics in other domains, vary systematically over the menstrual cycle. Whether or not preferences for other putative cues of men's long-term health are similarly affected by menstrual cycle is equivocal, however. For example, although many studies have demonstrated that women's preferences for the body odours of symmetric men are enhanced around ovulation reviewed in [ ] , evidence for cyclic shifts in women's preferences for symmetry in men's faces is inconsistent. One study has found that women's preferences for symmetric male faces were stronger around ovulation than during other phases of the menstrual cycle, at least among partnered women who were instructed to judge men's attractiveness as short-term mates [ ]. By contrast, other studies have observed no evidence for cyclic shifts in women's preferences for symmetric men's faces e. Although evidence that women's preferences for symmetry in men's faces vary systematically over the menstrual cycle is equivocal, that is not to say that robust cyclic shifts in women's perceptions of faces are only evident in their preferences for facial masculinity. For example, women's aversions to self-resembling faces are enhanced around ovulation and positively correlated with women's estimated progesterone levels during the menstrual cycle [ ]. This variation in attitudes to self-resembling faces may reflect increased inbreeding avoidance around ovulation and increased preferences for caring, supportive and trustworthy individuals when increased progesterone prepares the body for pregnancy [ ]. Moreover, women's aversions to facial cues associated with current illness e. Indeed, pregnant women and women using oral contraceptives which mimic the effects of increased progesterone during pregnancy demonstrate stronger aversions to individuals displaying facial cues of illness than do women with natural menstrual cycles [ 97 ]. These latter findings for aversions to facial cues of illness and progesterone during the menstrual cycle complement other research on increased aversions to possible sources of contagion in women's food preferences during pregnancy [ ], as well as increased sensitivity to facial expressions signalling that sources of threat and contagion are nearby when progesterone levels are raised [ , ]. While our discussion of hormone-mediated face preferences in women has emphasized the positive findings that have been reported in the literature, it is important to note that there have also been unsuccessful replications of cyclic variation in women's face preferences. For example, two recent studies observed no evidence for cyclic variations in women's preferences for masculine versus feminine male faces [ , ]. One possible explanation of these null findings comes from findings that suggest the extent to which women's preferences for masculine men vary over the menstrual cycle vary systematically among women. For example, cyclic variation in women's preferences for masculine characteristics in men's voices is significantly greater among women with high trait i. This pattern of results may occur because varying their sexual strategy during the menstrual cycle may benefit unattractive women more than it benefits attractive women [ ]. More recent research has presented additional evidence that women's family background, prenatal hormone levels and mortality salience might also affect the extent to which they vary their masculinity preferences according to their menstrual cycle phase [ — ]. We also note that there are significant methodological differences between studies examining cycle effects, making direct comparisons e. For example, some studies distinguish between short- and long-term mating contexts, generally with larger cyclic shifts for short-term judgements [ ], while others do not [ ]. Studies also differ in stimuli number, stimuli type and how fertility is defined. A thorough description of methodological differences between studies is not the focus here, but methodology is certainly a factor that could explain differences in findings across studies. It is likely that further research concerning individual differences in cyclic shifts and comparing different methodologies would provide important insights into the motivations, functions and mechanisms behind cyclic shifts in fundamental aspects of face perception. Despite a widespread belief that beauty cannot be defined, in fact, there is considerable agreement across individuals and cultures on what is found attractive. By considering that attraction and mate choice are critical components of evolutionary selection, we can better understand the importance of beauty. There are many traits that are linked to facial attractiveness in humans and each may in some way impart benefits to individuals who act on their preferences. If a trait is reliably associated with some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive. There have been several studies that have addressed how facial appearance relates to the healthiness of an individual in humans. The three traits discussed above are often manipulated by changing only face shape, but health perception appears to be related to facial colour and texture also. Fewer studies have examined how colour and texture of faces influence attractiveness judgements. One study has examined how well ratings of health from small patches of skin of faces are related to overall rated attractiveness when the whole face image is available. Jones et al. In other research, homogeneity of skin colour was positively related to attractiveness [ 99 ]. Findings have also suggested that more heterozygous men also have healthier appearing skin [ 56 ]. Skin health may be a particularly useful marker of current health condition as it is more changeable than aspects such as symmetry or averageness. Coloration is directly related to the appearance of skin. Coloration also appears to be an important component of sexual selection in many species. Red coloration is associated with dominance in fish [ ], birds [ ] and non-human primates [ , ] and, consequently, is linked to attracting the opposite sex. It has been noted that primates with trichromatic vision are generally bare-faced [ ] and that, at least in humans, facial flushing is associated with anger and confrontation [ ]. In research on non-human primates, there has been much interest in colour. For example, experimental manipulation of colour shows that female rhesus macaques prefer images of redder male faces [ ], while males prefer images of redder female hindquarters [ ]. In mandrills, red facial colour is related to rank in males [ ], and females sexually present more frequently to brighter males and also groom them more frequently [ ]. Red coloration also has consequences for behaviour in other species. For example, in bird species, the addition of red to stimuli can increase social dominance [ ]. In humans, it has been shown that wearing red in a variety of physically competitive sports is associated with an increased chance of winning over opponents [ ]. This has been interpreted as natural associations of red with dominance being extended to artificially displayed red in the same way that artificial stimuli can exploit innate responses to natural stimuli [ , ]. One study pitting red versus blue shapes found that red shapes were seen as more aggressive, dominant and more likely to win in physical competitions [ ]. Red does generally seem to have aversive effects on human behaviour. For example, when taking exams, individuals move their body away from tests with red covers more than they do from those with green or grey covers [ ]. While these studies suggest the colour red may be seen as a threatening stimulus in humans, red also appears to enhance attraction in some instances. For example, women are seen as more attractive by men when presented with red backgrounds or with red clothing, relative to other colours [ ]. This effect appears to be specific to attractiveness judgements; red colour does not influence judgements of other traits such as kindness or intelligence and does not influence women's attractiveness judgements of other women [ ]. Further research has examined red coloration in faces and demonstrated a positive association with perceived health [ ]. The authors suggest that perception of healthy, oxygenated blood may drive associations between red and healthiness. Alongside redness, people also appear to think that skin yellowness is associated with healthy appearance in faces [ ]. Yellowness may advertise health via an association with diet, as carotenoids are associated with skin yellowness and are absorbed via the intake of fruit and vegetables [ ]. In a classic social psychology study, Dion et al. For example, attractive individuals were thought to be able to achieve more prestigious occupations, be more competent spouses with happier marriages and have better prospects for personal fulfilment. There has been a wealth of studies examining this attractiveness stereotype, demonstrating that attractive people are seen in a positive light for a wide range of attributes compared with unattractive people. Studies on attractiveness stereotypes have generally not addressed the particular characteristics of faces that make individuals either attractive or unattractive, or the features that elicit personality attributions, although different faces reliably elicit the same personality attributions [ ]. Expression certainly has large effects, with, for example, faces shown with smiles rated as more attractive and as having more positive personality traits than neutral faces e. Such facial expressions are transient, however, and will differ rapidly within individuals over time and across photographs. Despite some findings showing a preference for more masculine and dominant male faces e. Personality traits are reported cross-culturally to be among the most important factors in partner choice by both sexes [ 1 , ]. If desired personality is so important, it would appear likely that personality attributions elicited by a face would affect its attractiveness. For example, women who value cooperation and good parenting may avoid masculine-faced men. Thus, instead of feminine faces being attractive and this attractiveness driving positive personality attributions, it may be that the personality attributions are driving the attractiveness judgements. Individuals may use personality stereotypes in mate selection to select partners with a personality that they desire. Some perceptual attributions to facial photographs are somewhat accurate e. Attraction to faces based on personality stereotypes may happen regardless of whether attributions are accurate or not, especially as many individuals do believe that face provides an important guide to character [ , ]. One study has indeed demonstrated that a desire for some personality traits influences judgements of facial attractiveness [ ]. Individuals valuing particular personality traits find faces appearing to display these traits attractive. Conversely, those not valuing particular traits find faces attractive that are perceived to possess that trait less. Thus, desired personality influences perceptions of facial attractiveness in opposite sex faces, changing the result to: In terms of benefits to perceivers, it is easy to see why traits such as appearing trustworthy would make a face appear more attractive. For individual-specific traits, the logic is more complicated, but such preferences could be related to behavioural compatibility within couples, as people do tend to desire partners with personalities similar to their own [ ]. One reason for variability in preferences for male facial masculinity may lie in the personality traits that masculine- and feminine-faced men are assumed to possess. Increasing the masculinity of face shape increased perceptions of dominance, masculinity and age but decreased perceptions of warmth, emotionality, honesty, cooperativeness and quality as a parent [ 83 ]. Indeed, recent work has shown that masculine facial characteristics are associated with indices of physical dominance, such as physical strength [ ], and the perception of such traits [ ], and that feminine men show weaker preferences for short-term relationships and stronger preferences for committed, long-term relationships than their masculine peers do [ ]. Women's face preferences may thus represent a trade-off between the desire for good genes and the desire for a cooperative partner. Of course, the five types of trait listed above are not a complete list of factors involved in the judgement of facial attractiveness. While individual traits impact on attractiveness, there is also scope for interaction between them. Certain face traits also appear to interact in generating preferences, however. For example, preferences for masculinity vary as a function of the healthiness of the face [ 96 ] and women's preferences for facial self-similarity are higher when men are more facially masculine [ ]. Such interactions highlight that facial attractiveness judgements are not simple: In humans, while individuals may share certain basic criteria for finding faces attractive, many factors may influence the specific types of face they find attractive. In this section, we review three broad areas leading to individual differences in preferences: Research suggests that internal factors predict individual differences in several aspects of face perception, including attractiveness judgements. Importantly, the nature of these individual differences suggests adaptive design in face perception and face preferences. In the following section, we discuss two broad types of internal factors: The influence of hormones on face perception is an area that has generated a considerable amount of empirical research in recent years. As detailed previously, masculine characteristics in men's faces are associated with measures of long-term medical health [ 35 , 77 ] and indices of developmental stability [ 36 , 37 ], physical strength [ ] and reproductive potential [ ]. By contrast, feminine characteristics in men's faces are associated with cues of investment and stronger preferences for long-term over short-term sexual relationships e. There is now compelling evidence that how women resolve this trade off between the costs and benefits associated with choosing a masculine mate is affected by hormone levels and fertility. Many studies have reported that women demonstrate stronger preferences for men displaying masculine facial characteristics around ovulation, when women are most fertile, than during other phases of the menstrual cycle [ — ]. Some studies have also reported that these cyclic shifts in women's preferences for masculine characteristics in men's faces are greatest among women who already have romantic partners and when women judge men's attractiveness for short-term, extra-pair relationships [ ]. Although the ultimate function of these cyclic shifts remains somewhat controversial, many researchers have interpreted cyclic shifts in women's masculinity preferences as evidence for adaptations that function to increase offspring health via high paternal investment from a long-term partner while promoting attraction to other men displaying cues of heritable immunity to infectious disease when most fertile discussed in [ ]. Women may gain maximal benefits by selecting investing long-term partners and high-quality extra-pair partners. Importantly, other explanations that have been suggested, such as increased attraction to individuals who appear to be likely sources of high-quality care and support during phases of the menstrual cycle when increased progesterone prepares the body for pregnancy i. Increased attraction to masculine men is by no means unique to face preferences; women also demonstrate stronger attraction to masculine men when judging the attractiveness of men's voices [ — ], body shapes [ ] and body odours [ ], as well as when judging the attractiveness of videoclips of male behavioural displays of dominance [ , ]. Furthermore, converging evidence for fertility-related variation in women's preferences for facial masculinity comes from studies investigating circum-pubertal and circum-menopausal variation in women's masculinity preferences; post-menopausal and pre-pubertal women report weaker preferences for masculine facial characteristics than do their pre-menopausal and post-pubertal counterparts, respectively e. The ultimate function of cyclic shifts in women's preferences for masculine facial characteristics is not the only controversial aspect of cyclic shifts in women's masculinity preferences. For example, although some researchers have suggested that cyclic shifts in women's masculinity preferences may be an artefact of the computer graphic methods that are generally used in these studies to experimentally manipulate sexually dimorphic cues in digital face images [ ], this claim is very difficult to reconcile with findings from studies that have demonstrated cyclic shifts in women's preferences for masculinity in real i. While these findings suggest that cyclic shifts in women's masculinity preferences are not an artefact of the stimuli used, an aspect of research on cyclic shifts in women's masculinity preferences that remains controversial is whether the effect of cycle phase on women's face preferences is relatively specific to judgements of men's faces, or also occurs when women judge the attractiveness of other women. To date, evidence is equivocal; some studies have observed cyclic shifts in women's preferences for masculine-faced men, but not masculine-faced women [ ], while others have observed cyclic shifts in women's preferences for masculine faces, irrespective of their sex [ , ]. These latter papers speculate that cyclic shifts in women's preferences for masculine-faced women could represent a low-cost functionless by-product of a mechanism that evolved primarily to increase women's preferences for masculine men around ovulation [ ], or have suggested that higher attractiveness ratings given to masculine women around ovulation could reflect increased derogation of feminine, and therefore attractive, same-sex competitors when women are most fertile [ ] see also [ ]. In addition to the sex-specificity of the effects of cycle phase on face preferences, the mechanisms that underpin cyclic shifts in women's preferences for masculine characteristics in men's faces have also been a topic of considerable interest in recent years. For example, research into the hormonal mechanisms that might underpin these cyclic shifts has variously emphasized the effects of variation in levels of testosterone [ ], oestrogen [ ] and progesterone [ , ], or has suggested, perhaps unsurprisingly, that cyclic shifts in women's masculinity preferences might be best explained by complex interactions among multiple hormones [ , ]. While findings from research into the hormonal mechanisms that might underpin cyclic shifts in women's masculinity preferences have arguably been inconsistent, the findings of corresponding research into the psychological mechanisms have been relatively consistent; various studies have demonstrated that women are quicker to categorize men and access male stereotypes around ovulation e. These findings suggest that cyclic variations in stereotype access and sexual desire might be important psychological mechanisms for regulating facial masculinity preferences during the menstrual cycle. While research on hormone-mediated face perception has generally focused on women's judgements of men's attractiveness, some recent research has investigated hormone-mediated face preferences among men. Men, of course, do not cycle in the same way women do, but levels of testosterone fluctuate within individuals. Research using natural variation in testosterone has shown that men's preferences for feminine characteristics in women's faces are stronger when their testosterone levels are high than when they are relatively low [ ]. This finding suggests that hormones, such as testosterone, can generate within-participant individual differences in face preference in men. As can be seen from the previous paragraphs, there is compelling evidence that women's preferences for masculine men, be they assessed from face preferences or from preferences for male characteristics in other domains, vary systematically over the menstrual cycle. Whether or not preferences for other putative cues of men's long-term health are similarly affected by menstrual cycle is equivocal, however. For example, although many studies have demonstrated that women's preferences for the body odours of symmetric men are enhanced around ovulation reviewed in [ ] , evidence for cyclic shifts in women's preferences for symmetry in men's faces is inconsistent. One study has found that women's preferences for symmetric male faces were stronger around ovulation than during other phases of the menstrual cycle, at least among partnered women who were instructed to judge men's attractiveness as short-term mates [ ]. By contrast, other studies have observed no evidence for cyclic shifts in women's preferences for symmetric men's faces e. Although evidence that women's preferences for symmetry in men's faces vary systematically over the menstrual cycle is equivocal, that is not to say that robust cyclic shifts in women's perceptions of faces are only evident in their preferences for facial masculinity. For example, women's aversions to self-resembling faces are enhanced around ovulation and positively correlated with women's estimated progesterone levels during the menstrual cycle [ ]. This variation in attitudes to self-resembling faces may reflect increased inbreeding avoidance around ovulation and increased preferences for caring, supportive and trustworthy individuals when increased progesterone prepares the body for pregnancy [ ]. Moreover, women's aversions to facial cues associated with current illness e. Indeed, pregnant women and women using oral contraceptives which mimic the effects of increased progesterone during pregnancy demonstrate stronger aversions to individuals displaying facial cues of illness than do women with natural menstrual cycles [ 97 ]. These latter findings for aversions to facial cues of illness and progesterone during the menstrual cycle complement other research on increased aversions to possible sources of contagion in women's food preferences during pregnancy [ ], as well as increased sensitivity to facial expressions signalling that sources of threat and contagion are nearby when progesterone levels are raised [ , ]. While our discussion of hormone-mediated face preferences in women has emphasized the positive findings that have been reported in the literature, it is important to note that there have also been unsuccessful replications of cyclic variation in women's face preferences. For example, two recent studies observed no evidence for cyclic variations in women's preferences for masculine versus feminine male faces [ , ]. One possible explanation of these null findings comes from findings that suggest the extent to which women's preferences for masculine men vary over the menstrual cycle vary systematically among women. For example, cyclic variation in women's preferences for masculine characteristics in men's voices is significantly greater among women with high trait i. This pattern of results may occur because varying their sexual strategy during the menstrual cycle may benefit unattractive women more than it benefits attractive women [ ]. More recent research has presented additional evidence that women's family background, prenatal hormone levels and mortality salience might also affect the extent to which they vary their masculinity preferences according to their menstrual cycle phase [ — ]. We also note that there are significant methodological differences between studies examining cycle effects, making direct comparisons e. For example, some studies distinguish between short- and long-term mating contexts, generally with larger cyclic shifts for short-term judgements [ ], while others do not [ ]. Studies also differ in stimuli number, stimuli type and how fertility is defined. A thorough description of methodological differences between studies is not the focus here, but methodology is certainly a factor that could explain differences in findings across studies. It is likely that further research concerning individual differences in cyclic shifts and comparing different methodologies would provide important insights into the motivations, functions and mechanisms behind cyclic shifts in fundamental aspects of face perception. While the previous section discussed research implicating hormone levels and fertility in individual differences in face perception, this section will discuss the relationships between face preferences and indices of own condition and attractiveness. Several studies have reported positive correlations between women's ratings of their own physical attractiveness and the strength of their preferences for masculine characteristics in men's faces [ 92 ]. Other studies have extended this work by demonstrating that more objective measures of women's condition and attractiveness, such as their waist—hip ratio or oestrogen levels, predict their preferences for masculine characteristics in men's faces in the same way [ , ]. Similar correlations between indices of women's own attractiveness and the strength of their preferences for masculine characteristics in other domains, such as men's voices, have also been reported [ , ], and indices of women's own condition and attractiveness are positively correlated with the strength of their preferences for symmetry and healthy-looking skin in men's faces [ 92 , ]. The findings described above appear to be somewhat analogous to condition-dependent preferences observed in other species, in which individuals in good physical condition show stronger preferences for high-quality mates e. Condition-dependent preferences in both humans and non-humans may have a common function and occur because individuals in good physical condition i. Particularly compelling evidence for this proposal comes from one of the few experimental studies of condition-dependent mate preferences. These findings suggest that women recalibrate subjective impressions of their own attractiveness i. While the research described above focused on the relationships between mate preferences and both individuals' own physical characteristics and their subjective evaluations of these physical characteristics, other work on condition-dependent preferences has investigated whether personality traits and other psychological factors predict individual differences in mate preferences in similar ways. For example, individual differences in systemizing and sensation-seeking, both of which are components of male sex-typical psychology, are positively correlated with men's preferences for feminine characteristics in women's, but not men's, faces [ , ]. Among women, individual differences in empathy, a component of female sex-typical psychology, and extraversion, a key predictor of social status that is correlated with women's physical attractiveness, are positively correlated with preferences for masculine characteristics in men's, but not women's, faces [ , ]. These findings not only implicate personality traits in individual differences in face preferences but also raise the intriguing possibility that some personality traits might mediate the relationships between an individual's physical characteristics and their face preferences. While factors such as hormones and own attractiveness can explain differences in face preferences between individuals, the context under which judgements are made can also contribute to variation in standards of beauty. In the following section, we discuss how context affects face preferences in three types of contexts: Information about genetic kinship is available in the face and is perceived somewhat accurately [ — ]. Judgements of facial similarity are highly synonymous with judgements of kinship [ — ], and facial similarity produced by computer-graphic manipulation affects behaviour in ways consistent with inclusive fitness theory e. Therefore, responses to facial resemblance are likely to be affected by prosocial versus sexual contexts. Cues of kinship are predicted to increase preferences in non-sexual, prosocial contexts, owing to the benefits associated with inclusive fitness [ ]. In other words, evolutionary models show that behaviours that benefit other individuals who share genes through common descent will be favoured. Therefore, if physical similarity is a reliable cue of genetic relatedness, we expect individuals to act prosocially towards individuals who appear similar to themselves. However, cues of kinship should have a less positive effect in sexual contexts, because of inbreeding's detrimental effects on offspring quality [ ]. One study investigated this prediction by comparing perceptions of the attractiveness of self-resembling own-sex and opposite-sex faces [ ]. Participants judged self-resemblance to be more attractive in the context of own-sex faces than in the context of opposite-sex faces. However, there was no such opposite-sex bias when the same faces were judged for averageness. This own-sex bias in preferences for self-resemblance indicates that, while self-resemblance is attractive in an exclusively prosocial i. Stronger attraction to cues of kinship in own-sex faces than in opposite-sex faces is likely to promote prosocial behaviour towards own-sex kin, while minimizing occurrences of inbreeding with opposite-sex kin. Transforms of self-similarity. Images are made by using the difference between a composite image of the same sex and an individual participant to make faces more similar to the participant. Self-dissimilar faces can be made by applying the same technique but using images other than the participant. Further evidence for context sensitivity in judgements of self-resembling faces is provided by a study comparing men's and women's preferences for self-resemblance in opposite-sex faces in explicitly prosocial versus sexual contexts [ ]. Participants were shown images of self-resembling opposite-sex faces and asked to judge their trustworthiness i. Consistent with both inclusive fitness and inbreeding avoidance theories, self-resemblance increased perceptions of trustworthiness, decreased attractiveness for short-term relationships and had no significant effect on attractiveness for long-term relationships. The fact that self-resemblance in opposite-sex faces was found to be trustworthy, but not attractive in short-term contexts, emphasizes the context-sensitivity of responses to self-resemblance. Importantly, because familiarity increases judgements of both attractiveness and trustworthiness [ ], this pattern of context-sensitivity strongly suggests that responses to self-resemblance do not occur simply because of familiarity alone i. Another example of social context influencing face preferences comes from research on interactions among the effects of different facial characteristics on preferences. For example, both behavioural and neurobiological evidence suggest that viewers demonstrate stronger attraction to attractive physical cues in faces e. Similarly, behavioural and neurobiological evidence also suggests that viewers demonstrate stronger attraction to cues associated with positive social interest e. Conway et al. Similarly, the putative costs of low investment are much less of a concern in short-term than in long-term relationships and, thus, women may demonstrate stronger masculinity preferences when judging men's attractiveness as possible short-term than long-term partners. Little et al. Women who were not using oral contraceptives made this face more masculine in the context of a short-term relationship than in the context of a long-term relationship. Penton-Voak et al. One potential explanation for this pattern of preference is that attractive women are better able to compete for, retain or replace high-quality, masculine partners and, therefore, do not show as large a shift in their preferences between short-term and long-term contexts. The effects of temporal context on judgements of attractiveness are not limited to faces. Women prefer lower pitched male voices in the context of a short-term relationship than in the context of a long-term relationship [ ]. This same study also found that the effect of relationship context was greatest when women were in the fertile phase of the menstrual cycle, a finding that is consistent with research on cyclic shifts in preferences for facial masculinity [ ]. A strong theoretical prediction of the trade-off account of variability in women's preferences for masculine men is that women in environments where poor health is particularly harmful to survival e. Regional differences in pathogen prevalence have been shown to be positively correlated with the importance placed on physical beauty and health [ , ]. DeBruine et al. This relationship remained significant, even when controlling for regional variation in wealth and women's mating strategies i. Brooks et al. Masculine-faced men may be favoured under such conditions, for example, as they may be better able to compete for resources. A further study of US states, in contrast, has shown that environmental health factors, and not indices of male—male violence such as homicide rates, predicts regional variation in women's masculinity preferences [ ]. Health, wealth and male—male violence are, of course, inter-related. While it is ultimately possible that health, wealth and male—male violence may all individually contribute to variation in preference, it is important to note that all of these analyses show that regional variation in women's masculinity preferences occurs in ways that are highly consistent with trade-off theories of sexual selection. The availability of resources in an environment may also influence face preferences. In low-resource environments, the resources to raise a child may be scarce or difficult to acquire and a preference for an investing partner be adaptive. To test these ideas, Little et al. Both men and women decreased their preferences for high quality mates for long-term relationships in the context of a harsh environment. This is consistent with the logic of trading genetic quality for commitment and investment in environments where resources are scarce. Individuals are confronted with a myriad of faces and social interactions every day. Research has shown that such experience leads to changes in preferences for faces. In the following section, we discuss two aspects of visual experience examining: Familiarity is a powerful determinant of attraction. For many types of stimuli, including faces, exposure increases attraction even when the exposure is unconscious [ — ]. Structural features of the face must be stored and represented in order to determine familiarity. As noted earlier, one idea for why averageness in faces is attractive comes from a link with familiarity—as average faces appear familiar this could positively affect their attractiveness [ 60 , 62 ]. Familiarity, when not paired with aversive stimuli, is thought to be rewarding [ ], and indeed there are obvious benefits to avoiding the unfamiliar. This can then help explain why exposure may cause increases in preference. There may, however, be more to increasing face preference than simple exposure. For example, recent studies have demonstrated that the nature of association positive or negative can affect face preferences, with positive experiences leading to increased attraction and negative experiences to decreased attraction [ ]. Moreover, these effects of valenced exposure are not bound solely to the specific individuals who were encountered and generalize to judgements of novel, physically similar individuals [ ]. Familiarity with parental traits has been implicated in human preferences. The phenomenon of imprinting, whereby individuals are attracted to parental traits, is well-studied in non-human animals [ , ] and there is increasing evidence for similar effects in humans. Following studies of facial similarity, judges have been shown to correctly match wives to their mother-in-law at a significantly higher rate than expected by chance and that wife—mother-in-law similarity is higher than similarity between husbands and their wives [ ]. Such effects are also seen in adopted daughters, controlling for any potential genetic effects, with significant facial resemblance between daughter's husband and her adoptive father [ ]. Other studies have shown that, for hair and eye colour, the best predictors of partner traits are the opposite-sex parent's colour traits [ ] and that individuals are attracted to age in faces consistent with the age of their parents when they were born [ ]. It is worth noting that at least in one study, effects were seen mainly for the opposite-sex parent [ ], which may indicate a more complex mechanism than simple exposure. Another line of argument suggesting imprinting-like effects appear not simply to reflect exposure comes from studies that have shown effects to be dependent on the quality of the relationship to the parent [ , ]. For example, daughters who report that they received greater emotional support from their adoptive fathers are more likely to choose mates who are similar to their father than individuals who report their father provided less emotional support [ ]. Similarly, women who rate their childhood relationships with their father positively show stronger attraction to face proportions similar to their father's face than women who rate their relationships less well [ ]. Imprinting-like effects then appear more complicated than simple exposure being directed more to one parent than the other and showing dependence on the relationship with that parent. Imprinting-like effects may lead to positive assortative mating pairing with similar partners , at least for long-term relationships, and this may have benefits in terms of keeping adaptive suites of genes together [ ] or increasing behaviour compatibility [ ]. There is certainly evidence that couples resemble each other facially [ , ]. Potentially then, a system that learns about known individuals and increases attraction to their face traits could be adaptive. Both familiarity and imprinting posit that exposure affects attractiveness. In recent years, exposure has been thought to have specific effects on our representations of faces via visual adaptation. We are unlikely to have an inbuilt average face and what is average must be calculated from experience. For each class of stimuli, the human visual system encounters may develop an individual representation, or prototype, made up of an average of the characteristics of all the different stimuli of that type that have been seen [ — ]. Computer modelling has revealed that algorithms trained to discriminate different stimuli produce stronger responses to stimuli that represent the average of the training set, even though this average was not previously encountered [ , ]. These findings have been interpreted as evidence that prototype formation is a property of learning to recognize different stimuli as members of a class [ , ]. Studies on category learning have a long history e. Learning studies examine how categorical perception develops using abstract stimuli. In classic studies, it has been shown that exposure to different dot patterns with particular configurations results in abstraction so that the average of each of the patterns, while never previously seen, is recognized as belonging to the set of patterns from which it was derived [ ]. Faces have been the focus of much research regarding recognition and prototype formation. While it has been proposed that faces may be coded as veridical representations of individuals or exemplars [ ], recent neuroimaging and single-cell recording studies have supported a prototype-referenced model of face coding [ , ]. Exposure to faces biases subsequent perceptions of novel faces, causing faces similar to those initially viewed to appear more prototypical than they would otherwise be perceived as, presumably, a prototype or population of exemplars becomes updated [ — ]. For example, adaptation to faces with contracted features causes novel faces with contracted features to be perceived as more normal than prior to this exposure [ , , ]. Such after-effects are thought to reflect changes in the responses of neural mechanisms underlying face processing [ , — ]. These studies may then shed light on how the brain builds an average representation to which the other faces can be compared. Importantly, exposure in the manner described above also influences attractiveness judgements. After exposure to faces possessing certain traits, these traits come to be preferred [ , , , ]. For example, if exposed to faces that look more like one identity, then new faces that resemble that identity are found more attractive than if exposed to the opposite set of face traits. A similar effect has also been observed for judgements of the trustworthiness of faces [ ]. Adaptation then reflects the rapid updating of face norms and can therefore be tied both to the effects of familiarity and imprinting-like effects. We have dealt briefly with some aspects of simple experience on preferences above, but, of course, humans are highly social and much human experience is of what other humans do. Humans can therefore learn about attractiveness from the behaviour of those around them: We have recently reviewed social learning in human face preferences [ ], and so present a brief overview here. Social learning can be adaptive if it allows an individual to assess potential mates more quickly and efficiently than through individual trial and error or allows an individual to use another's expertise. Mate choice copying has been observed among females in a number of different non-human species [ — ], including fish [ — ] and bird species [ — ]. Such studies have generally shown that when females observe another female the model to be paired with one of the two males the targets , they are subsequently more likely to prefer the target male they had seen paired with the model over the male that was not paired with the model. Inspired by work on non-human species, recent research also suggests that social learning may influence human mate preferences. While some research has shown that the presence of wedding rings on men did not increase women's preferences for those men [ ], other studies have found that images of men labelled as married were more attractive than those labelled as single [ ] and that women rate men as more desirable when they are shown surrounded by women than when they are shown alone or with other men [ ]. Another study has shown that women prefer pictures of men that had been previously seen paired with images of other women who were looking at the face with smiling i. Women therefore do appear to mimic the attitude of other women to particular men. Alongside partnership status, simple presence, and expressions of attitude towards the male, the physical traits of the observed model may also play a role in the social transmission of preference. Previous studies have shown that men's and women's attractiveness judgements are influenced by the apparent choice of attractive members of the same sex. Such a phenomenon suggests a more sophisticated form of mate-choice copying, whereby women can use the attractiveness of a partner that a man can acquire in order to judge the man's own attractiveness. Another study using images that were presented with a fictitious partner has shown that both men and women find a face paired with an attractive partner to be more attractive than one paired with an unattractive partner for a long-term but not a short-term relationship [ ]. Being more or less attractive has important social consequences and people do generally agree on who is and who is not attractive. Beauty is not just a simple social construct—attractiveness appears to be ingrained in our biology. While some aspects of face perception might be innate, other aspects are clearly influenced by experience; it seems unlikely that individuals are born with a representation of what a perfect partner looks like. If a trait reliably advertises some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive. It is clear that individual differences in preferences for some traits will prove adaptive and so are consistent with evolutionary theory. Work on facial attractiveness is also integrative, combining theories and methods from behavioural ecology, cognition, cross-cultural research and social psychology. Anthony C. Jones , 2 and Lisa M. DeBruine 2. Benedict C. Lisa M. This article has been cited by other articles in PMC. Abstract Face preferences affect a diverse range of critical social outcomes, from mate choices and decisions about platonic relationships to hiring decisions and decisions about social exchange. The evolutionary basis of attraction: Open in a separate window. Adaptive individual differences In humans, while individuals may share certain basic criteria for finding faces attractive, many factors may influence the specific types of face they find attractive. Summary and conclusions Being more or less attractive has important social consequences and people do generally agree on who is and who is not attractive. References 1. Buss D. Preferences in human mate selection. Langlois J. Maxims or myths of beauty? A meta-analytic and theoretical review. Andersson M. Sexual selection. Princeton, NJ: Princeton University Press [ Google Scholar ]. Bilateral symmetry and sexual selection: Elder G. Appearance and education in marriage mobility. Holmes S. Personal appearance as related to scholastic records and marriage selection in college women. Riggio R. The role of non-verbal and physical attractiveness in the selection of dating partners. Berscheid E. Physical attractiveness and dating choice: Walster E. Importance of physical attractiveness in dating behaviour. Dion K. What is beautiful is good. Eagly A. What is beautiful is good, but …: Feingold A. Good-looking people are not what we think. Cash T. The aye of the beholder: Sigall H. Beautiful but dangerous: Downs A. Natural observations of the links between attractiveness and initial legal judgments. B 17 , — Chiu R. The relative importance of facial attractiveness and gender in Hong Kong selection decisions. Marlowe C..

As can be seen from the Facial attractiveness milffaceness paragraphs, there is compelling evidence that women's preferences for masculine men, be they assessed from Facial attractiveness milffaceness preferences or from preferences for male characteristics in other domains, vary systematically over the menstrual cycle.

Whether or not preferences for other putative cues of men's long-term health are similarly affected by menstrual cycle is equivocal, however. For example, although many studies have demonstrated that women's preferences for the read more odours of symmetric men are enhanced around ovulation reviewed in [ ]evidence for cyclic shifts in women's preferences for symmetry in men's faces is inconsistent.

One study has found Facial attractiveness milffaceness women's preferences for symmetric male faces were stronger around ovulation than during other phases of the menstrual cycle, at least among partnered women who were instructed to judge men's attractiveness as short-term mates [ ].

By contrast, other studies have observed Facial attractiveness milffaceness evidence for cyclic shifts in women's preferences for symmetric men's faces e. Although evidence that women's preferences for symmetry in men's faces vary systematically over the menstrual cycle is equivocal, that is not to say that robust cyclic shifts in women's perceptions of faces are only evident in their preferences for facial masculinity.

For Facial attractiveness milffaceness, women's aversions to self-resembling faces are enhanced around ovulation and positively correlated with women's estimated progesterone levels during the menstrual cycle [ ]. This variation in attitudes to self-resembling faces may reflect increased inbreeding Facial attractiveness milffaceness around ovulation and increased preferences for caring, supportive and trustworthy individuals when increased progesterone prepares the body for pregnancy [ ].

Moreover, women's aversions to facial cues associated with current illness e. Facial attractiveness milffaceness, pregnant women and women using oral contraceptives which mimic the effects of increased progesterone during pregnancy demonstrate stronger Facial attractiveness milffaceness to individuals displaying facial cues of illness than Female nipple pics women with natural menstrual cycles [ 97 ].

These latter findings for aversions to facial cues of illness and progesterone during the menstrual cycle complement other research on increased aversions to possible sources of contagion in women's food preferences during pregnancy [ read article, as well as increased sensitivity to facial expressions signalling that sources of threat and contagion are Facial attractiveness milffaceness when progesterone levels are raised [].

While our discussion of hormone-mediated face preferences in women has emphasized the positive findings that have been reported in the literature, it is important to note that there have also been unsuccessful replications of cyclic variation in women's face preferences. For example, two recent studies observed no evidence for cyclic variations in women's preferences for masculine versus feminine male faces [].

One possible explanation of these null findings comes from findings that suggest the extent to which women's preferences for masculine men vary over the menstrual cycle vary systematically among women.

For example, cyclic variation in women's preferences for masculine characteristics in men's voices is significantly greater among women with high trait i. This pattern of results may occur because varying their sexual strategy during the menstrual cycle may benefit unattractive women more than it benefits attractive women [ ].

More recent research has presented additional evidence that women's family background, Facial attractiveness milffaceness hormone levels and mortality salience might also affect the extent to which they vary their masculinity preferences according to their menstrual cycle phase [ — Facial attractiveness milffaceness. We Facial attractiveness milffaceness note that there are significant methodological differences between studies examining cycle effects, making direct comparisons e.

For example, some studies distinguish between short- and long-term mating contexts, generally with larger cyclic shifts for short-term judgements [ ], while others do not [ ].

Studies also differ in stimuli number, stimuli type and how fertility is defined. A thorough description of methodological differences between studies is not the focus here, but methodology is certainly a factor that could explain differences in findings across studies.

It is likely that further research concerning individual Facial attractiveness milffaceness in cyclic shifts and comparing different methodologies would provide important insights into the motivations, functions and mechanisms Facial attractiveness milffaceness cyclic shifts in fundamental aspects of face perception.

While the previous section discussed research implicating hormone levels and fertility in individual differences in face perception, this section will discuss the relationships between face preferences and indices of own Facial attractiveness milffaceness and attractiveness.

Facial attractiveness milffaceness studies have reported positive correlations between women's ratings of their own physical attractiveness and the strength of their preferences for masculine characteristics in men's faces [ 92 ].

Other studies have extended this work by demonstrating that more objective measures of women's condition Facial attractiveness milffaceness attractiveness, such as their waist—hip ratio or oestrogen levels, predict their preferences for masculine characteristics in men's faces in the same way Facial attractiveness milffaceness]. Similar correlations between indices of women's own attractiveness and the strength of their preferences for masculine characteristics in other domains, such as men's voices, have also been reported [], and indices of women's own condition and attractiveness are positively correlated with the strength of their preferences Facial attractiveness milffaceness symmetry and healthy-looking skin in men's faces [ 92].

The findings described above appear to be somewhat analogous to condition-dependent preferences observed in other species, in which individuals in good physical condition show stronger preferences for high-quality mates e.

Condition-dependent preferences in both humans and non-humans may have a common Facial attractiveness milffaceness and occur because individuals in good physical condition i. Particularly compelling evidence for this proposal comes from one of the few experimental studies of condition-dependent mate preferences. These findings suggest that women recalibrate subjective impressions of their own attractiveness i.

Nude blonde Watch Video xxxhomesexhot. Being more or less attractive has important social consequences and people do generally agree on who is and who is not attractive. Beauty is not just a simple social construct—attractiveness appears to be ingrained in our biology. While some aspects of face perception might be innate, other aspects are clearly influenced by experience; it seems unlikely that individuals are born with a representation of what a perfect partner looks like. If a trait reliably advertises some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive. It is clear that individual differences in preferences for some traits will prove adaptive and so are consistent with evolutionary theory. Work on facial attractiveness is also integrative, combining theories and methods from behavioural ecology, cognition, cross-cultural research and social psychology. Login to your account. Forgot password? Keep me logged in. New User. Change Password. Old Password. New Password. Create a new account. Returning user. Can't sign in? Forgot your password? Enter your email address below and we will send you the reset instructions. If the address matches an existing account you will receive an email with instructions to reset your password Close. Request Username. Forgot your username? Enter your email address below and we will send you your username. You have access. View PDF. Little Anthony C. Facial attractiveness: Biological Sciences http: You have access Articles. Anthony C. Benedict C. Jones Benedict C. Lisa M. DeBruine Lisa M. Abstract Face preferences affect a diverse range of critical social outcomes, from mate choices and decisions about platonic relationships to hiring decisions and decisions about social exchange. Download figure Open in new tab Download PowerPoint. References 1 Buss D. Langlois J. A meta-analytic and theoretical review. Andersson M. Princeton, NJ: Princeton University Press. Elder G. Holmes S. Riggio R. Berscheid E. Walster E. Dion K. Eagly A. Feingold A. Cash T. Sigall H. Downs A. B 17 , — Chiu R. Marlowe C. Wolf N. New York, NY: Hume D. Of the standard of taste. London, UK: Darwin C. John Murray. Advances in experimental social psychology ed. Academic Press. Ford C. Cunningham M. Zebrowitz-McArthur L. Thornhill R. Trends Cogn. Rhodes G. Gibson R. Moller A. Oxford, UK: Oxford University Press. Valen L. Evolution 16 , — Dufour K. Manning J. Gangestad S. Little A. Grammer K. Scheib J. B , — Penton-Voak I. Jones B. Perception 33 , — Mealey L. Kowner R. Human 22 , — Perrett D. Waitt C. Perception 36 , — Mitton J. Roberts S. Lie H. Evolution 62 , — Galton F. Nature 18 , 97— Light L. B 7 , — Apicella C. Alley T. Perception 30 , — Enlow D. Philadelphia, PA: Zahavi A. Hillgarth N. Parasites and pathogens ed. Kanda N. Yesilova Z. Folstad I. B , S93—S Law-Smith M. Neave N. Jones D. Nature , — Facialmetric assessment of multiple motives in the perception of male facial physical attractiveness. Berry D. Personality correlates of facial babyishness. B 15 , — Keating C. McArthur L. Swaddle J. B , 39— DeBruine L. Smith F. Krupp D. In press. Apparent health encourages reciprocity. Fink B. Millinski M. Pryke S. Setchell J. Ethology , 25— B , S—S Changizi M. Drummond P. Psychophysiology 38 , — Cuthill I. Hill R. Nature , Burley N. Elliot A. B 35 , — Stephen I. Hassin R. Otta E. Motor Skills 82 , — Buss D. Brain Sci. Borkenau P. Liggett J. Botwin M. Boothroyd L. Korthase K. Motor Skills 54 , — Coetzee V. Perception 38 , — Osborn D. Makeup and posture effects on physical attractiveness judgments. Saxton T. Johnston V. Puts D. Feinberg D. Havlicek J. Psychoneuroendocrinology 35 , — Vukovic J. Peters M. Welling L. Roney J. Johnston L. Macrae C. Oinonen K. Sexual Behav. Pepper G. Derntl B. Conway C. Harris C. Menstrual cycle and facial preferences reconsidered. Sex Roles. Scarbrough P. Vaughn J. Psychol , — Bakker T. Alvergne A. Nesse R. Bressan P. Kaminski G. Dal Martello M. Maloney L. Hamilton W. Bittles A. Buckingham G. Zajonc R. O'Doherty J. Neuropsychologia 41 , — B , 63— Kampe K. Main J. Inquiry 17 , — Schmitt D. Brooks R. Geary D. Sex Res. Mace R. If a trait is reliably associated with some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive. Such an approach has highlighted face traits such as age, health, symmetry, and averageness, which are proposed to be associated with benefits and so associated with facial attractiveness. This view may postulate that some traits will be universally attractive; however, this does not preclude variation. Indeed, it would be surprising if there existed a template of a perfect face that was not affected by experience, environment, context, or the specific needs of an individual. Hillgarth N. Testosterone and immunosuppression in vertebrates: In Parasites and pathogens ed. Beckage N. Kanda N. Testosterone inhibits immunoglobulin production by human peripheral blood mononuclear cells. Yesilova Z. The effects of gonadotropin treatment on the immunological features of male patients with idiopathic hypogonadotropic hypogonadism. Folstad I. Parasites, bright males and the immunocompetence handicap. Parasitism, host immune function, and sexual selection. Does sexual dimorphism in human faces signal health? B , S93—S Law-Smith M. Facial appearance is a cue to oestrogen levels in women. High salivary testosterone is linked to masculine male facial appearance in humans. Neave N. Second to fourth digit ratio, testosterone and perceived male dominance. Measuring the physical in physical attractiveness: Jones D. Criteria of facial attractiveness in five populations. Effects of sexual dimorphism on facial attractiveness. Nature , — What do women want? Facialmetric assessment of multiple motives in the perception of male facial physical attractiveness. Berry D. Were the physiognomists right? Personality correlates of facial babyishness. B 15 , — Keating C. Gender and the physiognomy of dominance and attractiveness. McArthur L. Impressions of baby-faced adults. Cross-cultural agreement in perceptions of babyfaced adults. Some components and consequences of a babyface. Sex-typicality and attractiveness: Swaddle J. Testosterone increases perceived dominance but not attractiveness in human males. Self-perceived attractiveness influences human female preferences for sexual dimorphism and symmetry in male faces. B , 39— Partnership status and the temporal context of relationships influence human female preferences for sexual dimorphism in male face shape. DeBruine L. Correlated preferences for facial masculinity and ideal or actual partner's masculinity. Viewing attractive or unattractive same-sex individuals changes self-rated attractiveness and face preferences in women. Smith F. Interactions between masculinity-femininity and apparent health in face preferences. Menstrual cycle, pregnancy and oral contraceptive use alter attraction to apparent health in faces. Krupp D. In press. Apparent health encourages reciprocity. Fink B. Visible skin color distribution plays a role in the perception of age, attractiveness, and health in female faces. Millinski M. Female sticklebacks use male coloration in sticklebacks and therefore avoid parasitised males. Pryke S. Red dominates black: Setchell J. Dominance, status signals and coloration in male mandrills Mandrillus sphinx. Ethology , 25— Evidence from rhesus macaques suggests that male coloration plays a role in female primate mate choice. B , S—S Changizi M. Bare skin, blood and the evolution of primate colour vision. Drummond P. The effect of expressing anger on cardiovascular reactivity and facial blood flow in Chinese and Caucasians. Psychophysiology 38 , — Selective attention toward female secondary sexual color in male rhesus macaques. Do female mandrills prefer brightly colored males? Cuthill I. Colour bands, dominance, and body mass regulation in male zebra finches Taeniopygia guttata. Hill R. Red enhances human performance in contests. Nature , Burley N. Influence of color-banding on the conspecific preferences of zebra finches. Social perception of red suggests special role in dominance signalling. Elliot A. The effect of red on avoidance behavior in achievement contexts. B 35 , — Romantic red: Stephen I. Facial skin coloration affects perceived health of human faces. Hassin R. Facing faces: Otta E. Reading a smiling face: A cross-cultural exploration of physiognomic traits of dominance and happiness. Sex differences in human mate preferences: Brain Sci. Borkenau P. Trait inferences: Liggett J. 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Women's preferences for masculinity in male faces are highest during reproductive age range and lower around puberty and post-menopause. Psychoneuroendocrinology 35 , — Vukovic J. Circum-menopausal effects on women's judgements of facial attractiveness. Peters M. Preferences across the menstrual cycle for masculinity and symmetry in photographs of male faces and bodies. Preferences for variation in masculinity in real male faces change across the menstrual cycle: Welling L. Raised salivary testosterone in women is associated with increased attraction to masculine faces. Roney J. Women's estradiol predicts preference for facial cues of men's testosterone. Johnston L. The need for speed: Macrae C. Person perception across the menstrual cycle: Sex drive is positively associated with women's preferences for sexual dimorphism in men's and women's faces. Exposure to sexually attractive men decreases women's preferences for feminine faces. Men report stronger attraction to femininity in women's faces when their testosterone levels are high. Preferences for symmetry in faces change across the menstrual cycle. Oinonen K. Facial symmetry detection ability changes across the menstrual cycle. Symmetry and sexual dimorphism in human faces: Women's attractiveness judgments of self-resembling faces change across the menstrual cycle. Social perception of facial resemblance in humans. Sexual Behav. Pepper G. Rates of nausea and vomiting in pregnancy and dietary characteristics across populations. Derntl B. Emotion recognition accuracy in healthy young females is associated with cycle phase. Conway C. Salience of emotional displays of danger and contagion in faces is enhanced when progesterone levels are raised. Harris C. Menstrual cycle and facial preferences reconsidered. Sex Roles. Scarbrough P. Individual differences in women's facial preferences as a function of digit ratio and mental rotation ability. Father absence, parent—daughter relationships and partner preferences. Vaughn J. The effect of mortality salience on women's judgments of male faces. Waist—hip ratio predicts women's preferences for masculine male faces, but not perceptions of men's trustworthiness. Female condition influence preferences for sexual dimorphism in faces of male humans Homo sapiens. Psychol , — Self-rated attractiveness predicts individual differences in women's preferences for masculine men's voices. Women's own voice pitch predicts their preferences for masculinity in men's voices. Women's physical and psychological condition independently predict their preference for apparent health in faces. Bakker T. Condition-related mate-choice in sticklebacks. Individual differences in empathizing and systemizing predict variation in face preferences. Sensation seeking and men's face preferences. Extraversion predicts individual differences in women's face preferences. Alvergne A. 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While the research described above focused on the relationships between mate preferences and both individuals' own physical characteristics and their Facial attractiveness milffaceness evaluations of these physical characteristics, other work on condition-dependent preferences has investigated whether personality traits and other psychological factors predict individual differences in mate preferences in similar ways.

For example, individual differences in systemizing and sensation-seeking, both of which are components of male sex-typical psychology, are positively correlated with men's preferences for Facial attractiveness milffaceness characteristics in women's, but not men's, faces []. Among women, individual differences in empathy, a component of female sex-typical psychology, and extraversion, a key predictor of social status that is correlated with women's physical attractiveness, are positively correlated with preferences for masculine characteristics in men's, Facial attractiveness milffaceness not women's, faces [].

These findings not only Facial attractiveness milffaceness personality traits in individual differences in face preferences but also raise the intriguing possibility that some personality traits might mediate the relationships between an individual's physical characteristics and their face preferences.

While factors such as hormones and own attractiveness can Facial attractiveness milffaceness differences in face preferences between individuals, the context under which judgements are made can also contribute to variation in standards of beauty. In the following section, we discuss how context affects face preferences in three check this out of contexts: Information about genetic kinship is available in the face and is perceived somewhat accurately [ — ].

Facial attractiveness milffaceness

Judgements of facial similarity Facial attractiveness milffaceness highly synonymous with judgements of kinship [ — ], and facial Facial attractiveness milffaceness produced by computer-graphic manipulation affects behaviour in ways consistent with inclusive fitness theory e.

Therefore, responses to facial resemblance are likely to be affected by prosocial versus sexual contexts. Cues of kinship are predicted to increase preferences in non-sexual, prosocial contexts, owing to the benefits associated with inclusive fitness [ ]. In other words, evolutionary models show that behaviours that benefit other individuals who share genes through common descent will be favoured. Therefore, if physical similarity is a reliable cue of genetic relatedness, we expect individuals to act prosocially towards individuals who appear similar to themselves.

However, cues of kinship should have a less positive effect in sexual contexts, because Facial attractiveness milffaceness inbreeding's detrimental effects on offspring quality [ ].

One study investigated this prediction by comparing perceptions of the attractiveness of self-resembling own-sex and opposite-sex faces [ ]. Participants judged self-resemblance to be more attractive in the context of own-sex faces Facial attractiveness milffaceness in the context of opposite-sex faces. However, there was no such opposite-sex bias when the same faces were judged for averageness.

This own-sex bias in preferences for self-resemblance indicates that, while self-resemblance is attractive in an exclusively prosocial i. Stronger attraction to cues of kinship in own-sex faces than in opposite-sex faces is likely to promote prosocial behaviour towards own-sex kin, while minimizing occurrences of inbreeding with opposite-sex kin.

Transforms of self-similarity. Images are made by using the difference between a composite image of the same sex and an individual participant to make faces more similar to the participant.

Self-dissimilar faces can be made by applying the same technique but using images Facial attractiveness milffaceness than article source participant.

Further evidence for context sensitivity in judgements of self-resembling faces is provided by a study comparing men's and women's preferences for self-resemblance in opposite-sex faces in explicitly prosocial versus sexual contexts [ ]. Participants were shown images of self-resembling opposite-sex faces and asked to judge their trustworthiness i.

Consistent with both inclusive Joslyn morsel and inbreeding avoidance theories, self-resemblance increased perceptions of trustworthiness, decreased attractiveness for short-term relationships and had no significant effect on attractiveness for long-term relationships.

The fact that self-resemblance in opposite-sex faces was found to be trustworthy, but not attractive in short-term contexts, emphasizes the context-sensitivity of responses to self-resemblance. Importantly, because familiarity increases judgements of both attractiveness and trustworthiness [ ], this pattern of context-sensitivity strongly suggests that responses to self-resemblance do not occur simply because of familiarity alone i.

Another example of social context influencing face preferences comes from research on interactions among the effects of different facial characteristics on preferences. For example, both behavioural and neurobiological evidence suggest that viewers demonstrate stronger attraction to attractive physical cues in faces e. Similarly, behavioural and neurobiological evidence also suggests that viewers demonstrate stronger attraction to cues associated with positive social interest e.

Conway et al. Similarly, the putative costs of low investment are much less of a concern in short-term than in long-term relationships Facial attractiveness milffaceness, thus, women may demonstrate stronger masculinity preferences when judging men's attractiveness as possible short-term than long-term partners. Little et al. Women who were not using oral contraceptives made this face more masculine in the context of a short-term relationship than in the context of a long-term relationship.

Penton-Voak et al. One potential explanation for this pattern of preference is that attractive women are better able to compete for, retain or replace high-quality, masculine partners and, therefore, do not show as large a shift in their preferences between short-term and long-term contexts. The effects of temporal context on judgements of attractiveness are not limited to faces.

Women prefer lower pitched male voices in the context of a short-term relationship than in the context of a long-term relationship [ ]. This same study also found that the effect of relationship context was greatest when women were in the fertile phase of the menstrual cycle, a finding that is consistent with research on cyclic shifts in preferences for facial masculinity [ ]. A strong theoretical prediction of the trade-off account of variability in women's preferences for masculine men is that women in environments where poor health is particularly harmful to survival e.

Regional differences in pathogen prevalence have been shown to be positively correlated with the importance placed on physical beauty and health []. DeBruine et al. This relationship remained significant, even when controlling for regional variation in wealth and women's mating strategies i. Brooks et al. Masculine-faced men may be favoured under such conditions, for example, as they may be better able to compete for resources.

A further study of US states, in contrast, has shown that environmental health factors, and not indices of male—male violence such as homicide rates, predicts regional variation in women's masculinity preferences [ ]. Health, wealth Facial attractiveness milffaceness male—male violence are, of course, Facial attractiveness milffaceness. While it is ultimately Facial attractiveness milffaceness that health, wealth and male—male violence may all individually contribute to variation in preference, it is important to note that all of these analyses show that regional variation in women's masculinity preferences occurs in ways that are highly consistent with trade-off theories of sexual selection.

The availability of resources in an environment may also influence face preferences. In low-resource environments, the resources to raise a child may be scarce or difficult to acquire and a preference for an investing partner be adaptive.

To test these ideas, Little et al. Both men and women decreased their preferences for high quality mates for long-term relationships in the context of a harsh environment.

This is consistent with the logic of trading genetic quality for commitment and Facial attractiveness milffaceness in environments where resources are scarce. Individuals are confronted with a myriad of faces and social interactions every day.

Research has shown that such experience leads to changes in preferences for faces. In the following click at this page, we discuss two aspects of visual experience examining: Familiarity is a powerful determinant of attraction. For many types of stimuli, including faces, exposure increases attraction even when the exposure is unconscious [ — ].

Structural features of the face must be stored and represented in order to determine familiarity. As noted Facial attractiveness milffaceness, one idea for why averageness in faces is attractive comes from a link with familiarity—as average faces appear familiar this could positively affect their attractiveness [ 6062 ]. Familiarity, when not paired with aversive stimuli, is thought to be rewarding [ ], and indeed there are obvious benefits to avoiding the unfamiliar.

This can then help explain why exposure may cause increases in preference. There may, however, be View my area to increasing face preference than simple exposure. For example, recent studies have demonstrated that the nature of association positive or negative can affect face preferences, with positive experiences leading to increased attraction and negative experiences to decreased attraction [ ].

Facial attractiveness milffaceness, these effects of valenced exposure are not bound Facial attractiveness milffaceness to the specific individuals who were encountered and generalize to judgements of novel, physically similar individuals [ ]. Familiarity with parental traits has been implicated in human preferences. The phenomenon of imprinting, whereby individuals are attracted to parental traits, is well-studied in non-human animals [] and there is increasing evidence for similar effects in humans.

Following studies of facial similarity, judges have been shown to correctly match wives to their mother-in-law at a significantly higher rate than expected by chance and that wife—mother-in-law similarity is higher than similarity between husbands and their wives [ ]. Such effects are also seen in adopted daughters, controlling for any potential genetic effects, with significant facial resemblance between daughter's husband and her adoptive father [ ].

Other studies have shown that, for hair and eye colour, the best predictors of partner traits are the opposite-sex parent's colour traits [ ] and that individuals are attracted to age in faces consistent with the age of their parents when they were born [ ]. Visit web page is Facial attractiveness milffaceness noting that at least in one study, effects were seen mainly for the opposite-sex parent [ ], which may indicate a more complex mechanism than simple exposure.

Another line of argument suggesting imprinting-like effects appear not simply to reflect exposure comes from studies that have shown effects to be dependent on the quality of the relationship to the parent []. For example, daughters who report that they received greater emotional support from their adoptive fathers are Facial attractiveness milffaceness likely to choose mates who are similar to their father than Facial attractiveness milffaceness who report their father provided Facial attractiveness milffaceness emotional support [ ].

Similarly, women who rate their childhood relationships with their father positively show stronger attraction to face proportions similar to their father's face than women who rate their relationships less well [ ]. Imprinting-like effects then appear more complicated than Facial attractiveness milffaceness exposure being directed more to one parent than the other and showing dependence on the relationship with that parent.

Imprinting-like effects may lead to positive assortative mating pairing with similar partnersat least for long-term relationships, and this may have benefits in terms of keeping adaptive suites of genes together [ ] or increasing behaviour compatibility [ ]. There is certainly evidence that couples resemble each other facially [].

Potentially then, a system that learns about known individuals and increases attraction to their face traits could be adaptive. Both familiarity and imprinting posit that exposure affects attractiveness. Facial attractiveness milffaceness recent years, exposure has been thought to have specific effects on our representations of faces via visual adaptation.

We are unlikely to have an inbuilt average face and what is average must be calculated from experience. For each class of stimuli, the human visual system encounters may develop an individual representation, or prototype, made up of an average of the characteristics of all Facial attractiveness milffaceness different stimuli of that type that have been seen [ — ].

Computer Facial attractiveness milffaceness has revealed that algorithms trained to discriminate different stimuli produce stronger responses to stimuli that represent the average of the training set, even though this average was not previously encountered [].

These findings have been interpreted as evidence that prototype formation is a property of learning to recognize different stimuli as members of a class []. Studies on category learning have a long history e. Learning studies examine how categorical perception develops using abstract Facial attractiveness milffaceness. In classic here, it has been shown that exposure to different dot patterns with particular configurations results in abstraction so that the average of each of the patterns, while never previously seen, is recognized as belonging to the set of patterns from which it was derived [ ].

Faces have been the focus of much research regarding recognition and prototype formation. While it has been proposed that faces may be coded as Facial attractiveness milffaceness representations of individuals or exemplars [ ], Facial attractiveness milffaceness neuroimaging and single-cell recording studies have supported a prototype-referenced model of face coding [].

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If the address matches an existing account you will receive an email with instructions to retrieve your username. Google Scholar. Facial attractiveness milffaceness this author on PubMed.

Search for more papers by this author. Face preferences affect a diverse range of critical social outcomes, from mate choices and decisions about platonic relationships to hiring decisions and decisions about social exchange. Firstly, we review the facial characteristics that influence attractiveness judgements of faces e.

The research relating to these issues highlights flexible, sophisticated Facial attractiveness milffaceness that support and promote adaptive responses to faces that appear to function to maximize the benefits of both our mate choices and more general decisions about other types of social partners.

Our magazines and television screens are not just filled with any faces—they are filled with attractive faces, and both women and men are highly concerned with good looks in a potential partner [ 1 ]. Physical appearance is important to humans and certain features appear Facial attractiveness milffaceness be found attractive across individuals and cultures [ 2 ].

The same holds true across the animal kingdom; most non-human species Facial attractiveness milffaceness on external traits, such as the size, shape and colour of adornments e. Research on animals has focused on individual traits that are attractive across individuals, and even species, Facial attractiveness milffaceness as symmetry [ 4 ]. Physical attractiveness has important social consequences. For example, beauty Facial attractiveness milffaceness associated with upward economic mobility, especially for women [ 56 ], attractive people have more dates than less attractive people [ 7 ], and people who have dated more attractive individuals report being more satisfied with their dates [ 89 ].

In mock interviews, attractive people are more likely to be hired than less attractive individuals [ 13 ] and attractiveness can also influence judgements about the seriousness of committed crimes [ 14 ]. Outside the laboratory, attractive people also appear to lead favourable lives; attractive individuals pay lower bail [ 15 ] and are more likely to be hired for jobs [ 16Facial attractiveness milffaceness ] than less attractive individuals.

Despite research on social consequences, exactly what it is that makes a face beautiful remains poorly defined. Facial attractiveness milffaceness of the major deterrents in determining the features of an attractive face lies in Facial attractiveness milffaceness widespread Facial attractiveness milffaceness that standards of attractiveness are learned gradually through exposure to culturally presented ideals e. If this were true, it would mean that attractiveness is arbitrary and what is beautiful now could, in Facial attractiveness milffaceness different time or place, be considered unattractive.

While individual and cross-cultural differences exist see laterthis politically correct view of beauty is to some extent false. In Facial attractiveness milffaceness, agreement between individuals is one of the best-documented and most robust findings in facial attractiveness research since the s. Across many studies it has been found that there is a high degree of agreement from individuals within a particular culture and also high agreement Facial attractiveness milffaceness individuals from different cultures see [ 2 ] for a meta-analytical review.

If different people can agree on Facial attractiveness milffaceness faces are attractive and which are not attractive when judging faces of varying ethnic background e. Cross-cultural agreement on attractiveness is evidence against the notion that attractiveness ideals are slowly Facial attractiveness milffaceness by those growing up within a particular culture and this suggests that there is something universal about attractive continue reading and unattractive faces that is recognized both across individuals and cultures.

In the next section, we discuss traits that are proposed to be generally attractive by reasoning based on evolutionary theories, but we return Facial attractiveness milffaceness the notion of individual variation later. An evolutionary view assumes that perception and preferences serve an adaptive function: Theoretically then, preferences guide us to choose mates who will provide the best chance of our genes surviving.

In many studies, this evolutionary view of attractiveness has been used to predict the specific characteristics of attractive faces see [ 25 ] for review. Facial attractiveness milffaceness selection Facial attractiveness milffaceness the theoretical framework for much work and Facial attractiveness milffaceness thorough discussion of this topic in general is beyond the current review.

Khristian Sex Watch Video Sirvent Sex. A character demonstrates fluctuating asymmetry FA when symmetry reflects the normal development, and deviations from this symmetry are randomly distributed with respect to side [ 30 ]. FA is a particularly useful measure of developmental stability because we know that the optimal developmental outcome is symmetry. Therefore, any deviation from perfect symmetry can be considered a sub-optimal solution which will result in performance problems in the future. FA is also a useful measure as it subsumes a huge amount of individual variation in development, being the outcome of differences in genetic e. Preferences for symmetry can then, potentially, provide both direct e. Whether symmetry is actually related to quality in other animals and humans is an issue addressed by a large literature, and a complete review is not the focus of this paper. While the issue is divided, and there is some evidence that symmetry is not associated with quality e. In humans, male body symmetry is positively related to sperm number per ejaculate and sperm speed [ 32 ] and female breast symmetry is positively correlated with fecundity [ 33 , 34 ]. Relating to faces, one study has demonstrated that facial asymmetry is positively related to self-reported number of occurrences of respiratory disease [ 35 ] and some studies have observed positive correlations between symmetry and other putative indices of underlying physical condition i. The relationship between symmetry and quality is not reviewed in detail here, but it should be noted that fitness-related characteristics, such as growth rate, fecundity and survivability, are positively associated with symmetry across a number of species and taxa e. Studies of naturally occurring human facial asymmetries also provide evidence that symmetry is found attractive, though such studies can be confounded by potential correlates. Studies measuring symmetry from unmanipulated faces have reported positive correlations with rated attractiveness [ 40 — 44 ] and one study has even demonstrated that with pairs of monozygotic twins, the twin with more symmetric measurements is seen as more attractive [ 45 ]. Studies using more sophisticated symmetry manipulations have demonstrated that symmetry can have a positive influence on attractiveness [ 47 , 48 ] and have established that the chimeric manipulations used in the early studies introduced unnatural proportions into the symmetric faces see [ 48 ]. Thus, the methodologically superior computer graphic studies [ 47 , 48 ] parallel the findings of investigations into naturally occurring facial asymmetries [ 40 , 41 , 43 — 45 ]. The computer graphic studies demonstrate that increasing symmetry alone is sufficient to increase attractiveness. Subsequently, other studies have replicated preferences for symmetry using manipulated stimuli in different Western samples e. Preferences for symmetry using manipulated faces have been found in African hunter—gatherers [ 51 ], and macaque monkeys gaze longer at symmetrical than at asymmetrical face images of conspecifics [ 52 ]. Symmetry and asymmetry. Symmetric images are usually preferred to asymmetric images. Importantly, recent studies have implicated perceptions of health in attraction to symmetric faces [ 44 , 53 ] and have suggested that the mechanisms underpinning preferences for symmetric faces are different from those that might drive preferences for symmetry in mate-choice-irrelevant stimuli e. Such findings suggest that preferences for symmetric faces reflect, at least in part, adaptations for mate choice. Averageness refers to how closely a face resembles the majority of other faces within a population; non-average faces have more extreme characteristics than the average of a population. Average faces may be attractive because an alignment of features that is close to a population average is linked to genetic diversity [ 54 , 55 ]. Parasites are generally best adapted to proteins that are common in the host population; hence, parasites are adapted to the genes that code for the production of these proteins. A second evolutionary theory for the attractiveness of averageness in faces is that extreme non-average genotypes are more likely to be homozygous for deleterious alleles, that is, to be more likely to possess genes that are detrimental to an individual than those with more average genotypes [ 54 ]. Both of these theories propose evolutionary benefits to mating with individuals possessing average faces. Recent studies have supported the link between averageness, heterozygosity i. Heterozygosity in the major histocompatibility complex MHC genes that code for proteins involved in immune response, is positively associated with facial attractiveness [ 56 ] and facial averageness [ 57 ]. More directly, another study has shown that facial averageness is positively related to medical health as measured from actual medical records in both men and women [ 58 ]. There is good evidence that average faces are indeed found attractive. Galton [ 59 ] first noted that multiple faces blended together were more attractive than the constituent faces. Recent studies have improved upon these techniques using computers to create digitally blended composite faces; generally, the more images in a composite, the more attractive it is found [ 60 — 62 ]. Aside from composite images, Light et al. Average faces are generally more symmetric and symmetry is typically attractive in faces discussed in more detail above. Several studies have controlled for this confound in the original studies. When averageness and symmetry were independently manipulated, one study found that both manipulations positively and independently influenced attractiveness judgements [ 65 ]. Other studies have used perfectly symmetric images manipulated in averageness and still have demonstrated preferences for averageness [ 66 , 67 ]. Indeed, by comparing preferences for averageness when the effects of symmetry were controlled for and were not controlled for, Jones et al. It has also been noted that, in the original composite studies, the more images that are blended together the smoother the skin texture becomes, as imperfections such as lines or blemishes are averaged [ 68 ]. Image c should be more attractive than both of the other images. Composites are made by marking key locations around the main facial features e. The average location of each point of the component faces is then calculated to define the shape of the composite. The images of the individual faces are then warped to the relevant average shape before superimposing the images to produce a photographic quality composite image. While the majority of the work described above has been carried out in North America, Britain and Australia, averageness has also been found to be attractive across different cultures. For example, facial averageness is also found attractive in Japanese participants [ 69 ] and in African hunter—gatherers [ 67 ]. Male and female faces differ in their shape. Mature features in adult human faces reflect the masculinization or feminization of secondary sexual characteristics that occurs at puberty. These face shape differences, in part, arise because of the action of hormones such as testosterone. Larger jawbones, more prominent cheekbones and thinner cheeks are all features of male faces that differentiate them from female faces e. From an evolutionary view, extremes of secondary sexual characteristics more feminine for women, more masculine for men are proposed to be attractive because they advertise the quality of an individual in terms of heritable benefits; they indicate that the owners of such characteristics possess good genes. In other words, such traits advertise the possession of genes that are beneficial to offspring inheriting them in terms of survival or reproduction. One explanation of the importance of these facial traits is that they represent a handicap to an organism [ 71 ] and the costs of growing the trait means that only healthy individuals can afford to produce them. For example, secondary sexual characteristics are proposed to be linked to parasite resistance because the sex hormones that influence their growth, particularly testosterone, lower immunocompetence. Testosterone has been linked to the suppression of immune function in many species [ 72 ], including humans [ 73 , 74 ]. Larger secondary sexual characteristics should be related to a healthier immune system because only healthy organisms can afford the high sex-hormone handicap on the immune system that is necessary to produce these characteristics [ 75 ]. In many non-human animal studies, there is a positive association between secondary sexual trait expression and immunocompetence e. A study by Rhodes et al. No relationship was found between femininity and actual health in female faces, though [ 77 ]. Another study has demonstrated that men's facial masculinity and women's facial femininity are negatively related to self reports of respiratory disease [ 35 ]. If health is heritable, then female preferences for masculinity and male preferences for femininity may indeed also reflect the choice of mates with good genes. There is also a link between hormonal profile and face shape. Women with higher circulating oestrogen have more feminine faces [ 78 ], while men with high testosterone have more masculine faces [ 79 ], but see also [ 80 ]. If women with high oestrogen and men with high testosterone are valued as mates, preferences for cues of hormonal profile could drive preferences for sexually dimorphic face shape. Masculinity is transformed using the difference between male and female face shape as defined by creating a male and female composite. Preferences for masculinity in male faces vary across studies, but feminine female faces are consistently found more attractive than masculine female faces. There is considerable evidence that feminine female faces are considered attractive. Studies measuring facial features from photographs of women [ 40 , 81 , 82 ] and studies manipulating facial composites [ 83 ] all indicate that feminine features increase the attractiveness of female faces across different cultures. If oestrogenized female faces provide cues to fertility and health, then male preferences for such features are potentially adaptive. This reasoning does not require oestrogen to be immunosuppressive or part of a handicap. The link between masculinity and attractiveness in male faces is less clear. Cunningham et al. Despite some findings showing a preference for more masculine and dominant faces, several studies have shown that feminine characteristics and faces of low dominance are of increased attractiveness [ 62 , 83 , 84 , 89 — 91 ]. Many studies have made use of computer graphic techniques to manipulate masculinity. Sexual dimorphism in face shape can be manipulated by taking the geometrical differences between average male and female face shapes and applying this difference to new faces, making more or less masculine or feminine versions [ 83 ]. This process simultaneously changes all dimorphic shape characteristics in the face. Perrett et al. For the male face stimuli, the shape selected by Caucasians as most attractive was significantly feminized for both the Caucasian male face and the Japanese male face continua. Similarly, Japanese participants also selected significantly feminized versions of the male stimuli for both the Japanese and Caucasian male face continua. Thus, in both cultures it was found that participants showed a preference for feminized male faces. Since then, several studies have also documented preferences for femininity [ 62 , 90 , 92 , 93 ], but some similar computer graphic studies have also reported preferences for masculinity [ 94 , 95 ]. Although some of this variation may be attributed to other characteristics of the faces that varied between sets of stimuli [ 96 ], this does not explain the variability in preferences. We discuss the sources of individual differences in preferences for sexually dimorphic shape cues in the latter sections of our article. The face traits discussed so far have often been measured and manipulated but also studied in terms of perception and related to attractiveness. The reasoning for why traits like symmetry are preferred is often related to underlying health. Thus, it is important to examine perceptions of facial health directly. Perceived health is difficult to relate to any one metric, but people will readily rate faces for perceived health and show very high agreement on such ratings e. In evolutionary terms, there is a large and obvious selective advantage in detecting healthy partners both for social exchange and mate choice. Indeed, while the role of health in mate preferences is clear see below , recent work has demonstrated that participants are more willing to reciprocate trust from healthy-looking social partners than from social partners who are relatively unhealthy-looking [ 98 ]. Such findings demonstrate the importance of health perceptions for social interaction generally. Again, as for previous traits, there may be both direct and indirect benefits to partnering with individuals who are perceived to be healthy. Facial healthiness. High healthiness is associated with higher ratings of attractiveness. There have been several studies that have addressed how facial appearance relates to the healthiness of an individual in humans. The three traits discussed above are often manipulated by changing only face shape, but health perception appears to be related to facial colour and texture also. Fewer studies have examined how colour and texture of faces influence attractiveness judgements. One study has examined how well ratings of health from small patches of skin of faces are related to overall rated attractiveness when the whole face image is available. Jones et al. In other research, homogeneity of skin colour was positively related to attractiveness [ 99 ]. Findings have also suggested that more heterozygous men also have healthier appearing skin [ 56 ]. Skin health may be a particularly useful marker of current health condition as it is more changeable than aspects such as symmetry or averageness. Coloration is directly related to the appearance of skin. Coloration also appears to be an important component of sexual selection in many species. Red coloration is associated with dominance in fish [ ], birds [ ] and non-human primates [ , ] and, consequently, is linked to attracting the opposite sex. It has been noted that primates with trichromatic vision are generally bare-faced [ ] and that, at least in humans, facial flushing is associated with anger and confrontation [ ]. In research on non-human primates, there has been much interest in colour. For example, experimental manipulation of colour shows that female rhesus macaques prefer images of redder male faces [ ], while males prefer images of redder female hindquarters [ ]. In mandrills, red facial colour is related to rank in males [ ], and females sexually present more frequently to brighter males and also groom them more frequently [ ]. Red coloration also has consequences for behaviour in other species. For example, in bird species, the addition of red to stimuli can increase social dominance [ ]. In humans, it has been shown that wearing red in a variety of physically competitive sports is associated with an increased chance of winning over opponents [ ]. This has been interpreted as natural associations of red with dominance being extended to artificially displayed red in the same way that artificial stimuli can exploit innate responses to natural stimuli [ , ]. One study pitting red versus blue shapes found that red shapes were seen as more aggressive, dominant and more likely to win in physical competitions [ ]. Red does generally seem to have aversive effects on human behaviour. For example, when taking exams, individuals move their body away from tests with red covers more than they do from those with green or grey covers [ ]. While these studies suggest the colour red may be seen as a threatening stimulus in humans, red also appears to enhance attraction in some instances. For example, women are seen as more attractive by men when presented with red backgrounds or with red clothing, relative to other colours [ ]. This effect appears to be specific to attractiveness judgements; red colour does not influence judgements of other traits such as kindness or intelligence and does not influence women's attractiveness judgements of other women [ ]. Further research has examined red coloration in faces and demonstrated a positive association with perceived health [ ]. The authors suggest that perception of healthy, oxygenated blood may drive associations between red and healthiness. Alongside redness, people also appear to think that skin yellowness is associated with healthy appearance in faces [ ]. Yellowness may advertise health via an association with diet, as carotenoids are associated with skin yellowness and are absorbed via the intake of fruit and vegetables [ ]. In a classic social psychology study, Dion et al. For example, attractive individuals were thought to be able to achieve more prestigious occupations, be more competent spouses with happier marriages and have better prospects for personal fulfilment. There has been a wealth of studies examining this attractiveness stereotype, demonstrating that attractive people are seen in a positive light for a wide range of attributes compared with unattractive people. Studies on attractiveness stereotypes have generally not addressed the particular characteristics of faces that make individuals either attractive or unattractive, or the features that elicit personality attributions, although different faces reliably elicit the same personality attributions [ ]. Expression certainly has large effects, with, for example, faces shown with smiles rated as more attractive and as having more positive personality traits than neutral faces e. Such facial expressions are transient, however, and will differ rapidly within individuals over time and across photographs. Despite some findings showing a preference for more masculine and dominant male faces e. Personality traits are reported cross-culturally to be among the most important factors in partner choice by both sexes [ 1 , ]. If desired personality is so important, it would appear likely that personality attributions elicited by a face would affect its attractiveness. For example, women who value cooperation and good parenting may avoid masculine-faced men. Thus, instead of feminine faces being attractive and this attractiveness driving positive personality attributions, it may be that the personality attributions are driving the attractiveness judgements. Individuals may use personality stereotypes in mate selection to select partners with a personality that they desire. Some perceptual attributions to facial photographs are somewhat accurate e. Attraction to faces based on personality stereotypes may happen regardless of whether attributions are accurate or not, especially as many individuals do believe that face provides an important guide to character [ , ]. One study has indeed demonstrated that a desire for some personality traits influences judgements of facial attractiveness [ ]. Individuals valuing particular personality traits find faces appearing to display these traits attractive. Conversely, those not valuing particular traits find faces attractive that are perceived to possess that trait less. Thus, desired personality influences perceptions of facial attractiveness in opposite sex faces, changing the result to: In terms of benefits to perceivers, it is easy to see why traits such as appearing trustworthy would make a face appear more attractive. For individual-specific traits, the logic is more complicated, but such preferences could be related to behavioural compatibility within couples, as people do tend to desire partners with personalities similar to their own [ ]. One reason for variability in preferences for male facial masculinity may lie in the personality traits that masculine- and feminine-faced men are assumed to possess. Increasing the masculinity of face shape increased perceptions of dominance, masculinity and age but decreased perceptions of warmth, emotionality, honesty, cooperativeness and quality as a parent [ 83 ]. Indeed, recent work has shown that masculine facial characteristics are associated with indices of physical dominance, such as physical strength [ ], and the perception of such traits [ ], and that feminine men show weaker preferences for short-term relationships and stronger preferences for committed, long-term relationships than their masculine peers do [ ]. Women's face preferences may thus represent a trade-off between the desire for good genes and the desire for a cooperative partner. Of course, the five types of trait listed above are not a complete list of factors involved in the judgement of facial attractiveness. While individual traits impact on attractiveness, there is also scope for interaction between them. Certain face traits also appear to interact in generating preferences, however. For example, preferences for masculinity vary as a function of the healthiness of the face [ 96 ] and women's preferences for facial self-similarity are higher when men are more facially masculine [ ]. Such interactions highlight that facial attractiveness judgements are not simple: In humans, while individuals may share certain basic criteria for finding faces attractive, many factors may influence the specific types of face they find attractive. In this section, we review three broad areas leading to individual differences in preferences: Research suggests that internal factors predict individual differences in several aspects of face perception, including attractiveness judgements. Importantly, the nature of these individual differences suggests adaptive design in face perception and face preferences. In the following section, we discuss two broad types of internal factors: The influence of hormones on face perception is an area that has generated a considerable amount of empirical research in recent years. As detailed previously, masculine characteristics in men's faces are associated with measures of long-term medical health [ 35 , 77 ] and indices of developmental stability [ 36 , 37 ], physical strength [ ] and reproductive potential [ ]. By contrast, feminine characteristics in men's faces are associated with cues of investment and stronger preferences for long-term over short-term sexual relationships e. There is now compelling evidence that how women resolve this trade off between the costs and benefits associated with choosing a masculine mate is affected by hormone levels and fertility. Many studies have reported that women demonstrate stronger preferences for men displaying masculine facial characteristics around ovulation, when women are most fertile, than during other phases of the menstrual cycle [ — ]. Some studies have also reported that these cyclic shifts in women's preferences for masculine characteristics in men's faces are greatest among women who already have romantic partners and when women judge men's attractiveness for short-term, extra-pair relationships [ ]. Although the ultimate function of these cyclic shifts remains somewhat controversial, many researchers have interpreted cyclic shifts in women's masculinity preferences as evidence for adaptations that function to increase offspring health via high paternal investment from a long-term partner while promoting attraction to other men displaying cues of heritable immunity to infectious disease when most fertile discussed in [ ]. Women may gain maximal benefits by selecting investing long-term partners and high-quality extra-pair partners. Importantly, other explanations that have been suggested, such as increased attraction to individuals who appear to be likely sources of high-quality care and support during phases of the menstrual cycle when increased progesterone prepares the body for pregnancy i. Increased attraction to masculine men is by no means unique to face preferences; women also demonstrate stronger attraction to masculine men when judging the attractiveness of men's voices [ — ], body shapes [ ] and body odours [ ], as well as when judging the attractiveness of videoclips of male behavioural displays of dominance [ , ]. Furthermore, converging evidence for fertility-related variation in women's preferences for facial masculinity comes from studies investigating circum-pubertal and circum-menopausal variation in women's masculinity preferences; post-menopausal and pre-pubertal women report weaker preferences for masculine facial characteristics than do their pre-menopausal and post-pubertal counterparts, respectively e. The ultimate function of cyclic shifts in women's preferences for masculine facial characteristics is not the only controversial aspect of cyclic shifts in women's masculinity preferences. For example, although some researchers have suggested that cyclic shifts in women's masculinity preferences may be an artefact of the computer graphic methods that are generally used in these studies to experimentally manipulate sexually dimorphic cues in digital face images [ ], this claim is very difficult to reconcile with findings from studies that have demonstrated cyclic shifts in women's preferences for masculinity in real i. While these findings suggest that cyclic shifts in women's masculinity preferences are not an artefact of the stimuli used, an aspect of research on cyclic shifts in women's masculinity preferences that remains controversial is whether the effect of cycle phase on women's face preferences is relatively specific to judgements of men's faces, or also occurs when women judge the attractiveness of other women. To date, evidence is equivocal; some studies have observed cyclic shifts in women's preferences for masculine-faced men, but not masculine-faced women [ ], while others have observed cyclic shifts in women's preferences for masculine faces, irrespective of their sex [ , ]. These latter papers speculate that cyclic shifts in women's preferences for masculine-faced women could represent a low-cost functionless by-product of a mechanism that evolved primarily to increase women's preferences for masculine men around ovulation [ ], or have suggested that higher attractiveness ratings given to masculine women around ovulation could reflect increased derogation of feminine, and therefore attractive, same-sex competitors when women are most fertile [ ] see also [ ]. In addition to the sex-specificity of the effects of cycle phase on face preferences, the mechanisms that underpin cyclic shifts in women's preferences for masculine characteristics in men's faces have also been a topic of considerable interest in recent years. For example, research into the hormonal mechanisms that might underpin these cyclic shifts has variously emphasized the effects of variation in levels of testosterone [ ], oestrogen [ ] and progesterone [ , ], or has suggested, perhaps unsurprisingly, that cyclic shifts in women's masculinity preferences might be best explained by complex interactions among multiple hormones [ , ]. While findings from research into the hormonal mechanisms that might underpin cyclic shifts in women's masculinity preferences have arguably been inconsistent, the findings of corresponding research into the psychological mechanisms have been relatively consistent; various studies have demonstrated that women are quicker to categorize men and access male stereotypes around ovulation e. These findings suggest that cyclic variations in stereotype access and sexual desire might be important psychological mechanisms for regulating facial masculinity preferences during the menstrual cycle. While research on hormone-mediated face perception has generally focused on women's judgements of men's attractiveness, some recent research has investigated hormone-mediated face preferences among men. Men, of course, do not cycle in the same way women do, but levels of testosterone fluctuate within individuals. Research using natural variation in testosterone has shown that men's preferences for feminine characteristics in women's faces are stronger when their testosterone levels are high than when they are relatively low [ ]. This finding suggests that hormones, such as testosterone, can generate within-participant individual differences in face preference in men. As can be seen from the previous paragraphs, there is compelling evidence that women's preferences for masculine men, be they assessed from face preferences or from preferences for male characteristics in other domains, vary systematically over the menstrual cycle. Whether or not preferences for other putative cues of men's long-term health are similarly affected by menstrual cycle is equivocal, however. For example, although many studies have demonstrated that women's preferences for the body odours of symmetric men are enhanced around ovulation reviewed in [ ] , evidence for cyclic shifts in women's preferences for symmetry in men's faces is inconsistent. One study has found that women's preferences for symmetric male faces were stronger around ovulation than during other phases of the menstrual cycle, at least among partnered women who were instructed to judge men's attractiveness as short-term mates [ ]. By contrast, other studies have observed no evidence for cyclic shifts in women's preferences for symmetric men's faces e. Although evidence that women's preferences for symmetry in men's faces vary systematically over the menstrual cycle is equivocal, that is not to say that robust cyclic shifts in women's perceptions of faces are only evident in their preferences for facial masculinity. For example, women's aversions to self-resembling faces are enhanced around ovulation and positively correlated with women's estimated progesterone levels during the menstrual cycle [ ]. This variation in attitudes to self-resembling faces may reflect increased inbreeding avoidance around ovulation and increased preferences for caring, supportive and trustworthy individuals when increased progesterone prepares the body for pregnancy [ ]. Moreover, women's aversions to facial cues associated with current illness e. Indeed, pregnant women and women using oral contraceptives which mimic the effects of increased progesterone during pregnancy demonstrate stronger aversions to individuals displaying facial cues of illness than do women with natural menstrual cycles [ 97 ]. These latter findings for aversions to facial cues of illness and progesterone during the menstrual cycle complement other research on increased aversions to possible sources of contagion in women's food preferences during pregnancy [ ], as well as increased sensitivity to facial expressions signalling that sources of threat and contagion are nearby when progesterone levels are raised [ , ]. While our discussion of hormone-mediated face preferences in women has emphasized the positive findings that have been reported in the literature, it is important to note that there have also been unsuccessful replications of cyclic variation in women's face preferences. For example, two recent studies observed no evidence for cyclic variations in women's preferences for masculine versus feminine male faces [ , ]. One possible explanation of these null findings comes from findings that suggest the extent to which women's preferences for masculine men vary over the menstrual cycle vary systematically among women. For example, cyclic variation in women's preferences for masculine characteristics in men's voices is significantly greater among women with high trait i. This pattern of results may occur because varying their sexual strategy during the menstrual cycle may benefit unattractive women more than it benefits attractive women [ ]. More recent research has presented additional evidence that women's family background, prenatal hormone levels and mortality salience might also affect the extent to which they vary their masculinity preferences according to their menstrual cycle phase [ — ]. We also note that there are significant methodological differences between studies examining cycle effects, making direct comparisons e. For example, some studies distinguish between short- and long-term mating contexts, generally with larger cyclic shifts for short-term judgements [ ], while others do not [ ]. Studies also differ in stimuli number, stimuli type and how fertility is defined. A thorough description of methodological differences between studies is not the focus here, but methodology is certainly a factor that could explain differences in findings across studies. It is likely that further research concerning individual differences in cyclic shifts and comparing different methodologies would provide important insights into the motivations, functions and mechanisms behind cyclic shifts in fundamental aspects of face perception. While the previous section discussed research implicating hormone levels and fertility in individual differences in face perception, this section will discuss the relationships between face preferences and indices of own condition and attractiveness. Several studies have reported positive correlations between women's ratings of their own physical attractiveness and the strength of their preferences for masculine characteristics in men's faces [ 92 ]. Other studies have extended this work by demonstrating that more objective measures of women's condition and attractiveness, such as their waist—hip ratio or oestrogen levels, predict their preferences for masculine characteristics in men's faces in the same way [ , ]. Similar correlations between indices of women's own attractiveness and the strength of their preferences for masculine characteristics in other domains, such as men's voices, have also been reported [ , ], and indices of women's own condition and attractiveness are positively correlated with the strength of their preferences for symmetry and healthy-looking skin in men's faces [ 92 , ]. This view may postulate that some traits will be universally attractive; however, this does not preclude variation. Indeed, it would be surprising if there existed a template of a perfect face that was not affected by experience, environment, context, or the specific needs of an individual. Research on facial attractiveness has documented how various face traits are associated with attractiveness and various factors that impact on an individual's judgments of facial attractiveness. Overall, facial attractiveness is complex, both in the number of traits that determine attraction and in the large number of factors that can alter attraction to particular faces. Perrett D. Symmetry and human facial attractiveness. Evidence against perceptual bias views for symmetry preferences in human faces. Attraction independent of detection suggests special mechanisms for symmetry preferences in human face perception. Preferences for symmetry in human faces in two cultures: Waitt C. Preferences for symmetry in conspecific facial shape among Macaca mulatta. Perceived health contributes to the attractiveness of facial symmetry, averageness, and sexual dimorphism. Perception 36 , — Human facial beauty: Mitton J. Associations among proteins heterozygosity, growth rate, and developmental homeostasis. Roberts S. MHC-heterozygosity and human facial attractiveness. Lie H. Genetic diversity revealed in human faces. Evolution 62 , — Do facial averageness and symmetry signal health? Galton F. Composite portraits. Nature 18 , 97— Attractive faces are only average. What is average and what is not average about attractive faces. The role of masculinity and distinctiveness in judgments of human male facial attractiveness. Light L. Why attractive people are harder to remember. B 7 , — Averageness, exaggeration, and facial attractiveness. Are average facial configurations attractive only because of their symmetry? The role of symmetry in attraction to average faces. Apicella C. Facial averageness and attractiveness in an isolated population of hunter-gatherers. Alley T. Averaged faces are attractive, but very attractive faces are not average. Attractiveness of facial averageness and symmetry in non-Western populations: Perception 30 , — Enlow D. Handbook of facial growth , 2nd edn. Philadelphia, PA: Saunders [ Google Scholar ]. Zahavi A. Mate selection: Hillgarth N. Testosterone and immunosuppression in vertebrates: In Parasites and pathogens ed. Beckage N. Kanda N. Testosterone inhibits immunoglobulin production by human peripheral blood mononuclear cells. Yesilova Z. The effects of gonadotropin treatment on the immunological features of male patients with idiopathic hypogonadotropic hypogonadism. Folstad I. Parasites, bright males and the immunocompetence handicap. Parasitism, host immune function, and sexual selection. Does sexual dimorphism in human faces signal health? B , S93—S Law-Smith M. Facial appearance is a cue to oestrogen levels in women. High salivary testosterone is linked to masculine male facial appearance in humans. Neave N. Second to fourth digit ratio, testosterone and perceived male dominance. Measuring the physical in physical attractiveness: Jones D. Criteria of facial attractiveness in five populations. Effects of sexual dimorphism on facial attractiveness. Nature , — What do women want? Facialmetric assessment of multiple motives in the perception of male facial physical attractiveness. Berry D. Were the physiognomists right? Personality correlates of facial babyishness. B 15 , — Keating C. Gender and the physiognomy of dominance and attractiveness. McArthur L. Impressions of baby-faced adults. Cross-cultural agreement in perceptions of babyfaced adults. Some components and consequences of a babyface. Sex-typicality and attractiveness: Swaddle J. Testosterone increases perceived dominance but not attractiveness in human males. Self-perceived attractiveness influences human female preferences for sexual dimorphism and symmetry in male faces. B , 39— Partnership status and the temporal context of relationships influence human female preferences for sexual dimorphism in male face shape. DeBruine L. Correlated preferences for facial masculinity and ideal or actual partner's masculinity. Viewing attractive or unattractive same-sex individuals changes self-rated attractiveness and face preferences in women. Smith F. Interactions between masculinity-femininity and apparent health in face preferences. Menstrual cycle, pregnancy and oral contraceptive use alter attraction to apparent health in faces. Krupp D. In press. Apparent health encourages reciprocity. Fink B. Visible skin color distribution plays a role in the perception of age, attractiveness, and health in female faces. Millinski M. Female sticklebacks use male coloration in sticklebacks and therefore avoid parasitised males. Pryke S. Red dominates black: Setchell J. Dominance, status signals and coloration in male mandrills Mandrillus sphinx. Ethology , 25— Evidence from rhesus macaques suggests that male coloration plays a role in female primate mate choice. B , S—S Changizi M. Bare skin, blood and the evolution of primate colour vision. Drummond P. The effect of expressing anger on cardiovascular reactivity and facial blood flow in Chinese and Caucasians. Psychophysiology 38 , — Selective attention toward female secondary sexual color in male rhesus macaques. Do female mandrills prefer brightly colored males? Cuthill I. Colour bands, dominance, and body mass regulation in male zebra finches Taeniopygia guttata. Hill R. Red enhances human performance in contests. Nature , Burley N. Influence of color-banding on the conspecific preferences of zebra finches. Social perception of red suggests special role in dominance signalling. Elliot A. The effect of red on avoidance behavior in achievement contexts. B 35 , — Romantic red: Stephen I. Facial skin coloration affects perceived health of human faces. Hassin R. Facing faces: Otta E. Reading a smiling face: A cross-cultural exploration of physiognomic traits of dominance and happiness. Sex differences in human mate preferences: Brain Sci. Borkenau P. Trait inferences: Liggett J. The human face. Constable [ Google Scholar ]. What is good is beautiful: Botwin M. Personality and mate preferences: Male facial appearance signals physical strength to women. Facial cues of dominance modulate the short-term gaze-cuing effect in human observers. Boothroyd L. Facial correlates of sociosexuality. Korthase K. Perceived age and perceived physical attractiveness. Motor Skills 54 , — [ Google Scholar ]. Coetzee V. Facial adiposity: Perception 38 , — Investigating an imprinting-like phenomenon in humans: The effects of facial hair manipulation on female perceptions of attractiveness, masculinity, and dominance in male faces. Osborn D. Beauty is as beauty does? Makeup and posture effects on physical attractiveness judgments. Facial resemblance increases the attractiveness of same-sex faces more than other-sex faces. Trustworthy but not lust-worthy: Saxton T. Trade-offs between markers of absolute and relative quality in human facial preferences. Attractiveness and sexual behavior: Johnston V. Male facial attractiveness: Commitment to relationships and preferences for femininity and apparent health in faces are strongest on days of the menstrual cycle when progesterone level is high. Female preference for male faces changes cyclically: Menstrual cycle alters face preference. Human oestrus. Effects of menstrual cycle phase on face preferences. Puts D. Cyclic variation in women's preferences for masculine traits: Feinberg D. Menstrual cycle, trait estrogen level, and masculinity preferences in the human voice. Mating context and menstrual phase affect women's preferences for male voice pitch. Preferences for masculinity in male bodies change across the menstrual cycle. Havlicek J. Women's preference for dominant male odour: Changes in women's mate preferences across the ovulatory cycle. Women's preferences for male behavioral displays change across the menstrual cycle. Women's preferences for masculinity in male faces are highest during reproductive age range and lower around puberty and post-menopause. Psychoneuroendocrinology 35 , — Vukovic J. Circum-menopausal effects on women's judgements of facial attractiveness. Peters M. Preferences across the menstrual cycle for masculinity and symmetry in photographs of male faces and bodies. Preferences for variation in masculinity in real male faces change across the menstrual cycle: Welling L. Raised salivary testosterone in women is associated with increased attraction to masculine faces. Roney J. Women's estradiol predicts preference for facial cues of men's testosterone. Johnston L. The need for speed: Macrae C. Person perception across the menstrual cycle: Sex drive is positively associated with women's preferences for sexual dimorphism in men's and women's faces. Exposure to sexually attractive men decreases women's preferences for feminine faces. Men report stronger attraction to femininity in women's faces when their testosterone levels are high. Preferences for symmetry in faces change across the menstrual cycle. Oinonen K. Facial symmetry detection ability changes across the menstrual cycle. Symmetry and sexual dimorphism in human faces: Women's attractiveness judgments of self-resembling faces change across the menstrual cycle. Social perception of facial resemblance in humans. Sexual Behav. Pepper G. Rates of nausea and vomiting in pregnancy and dietary characteristics across populations. Derntl B. Emotion recognition accuracy in healthy young females is associated with cycle phase. Conway C. Salience of emotional displays of danger and contagion in faces is enhanced when progesterone levels are raised. Harris C. Menstrual cycle and facial preferences reconsidered. Sex Roles. Scarbrough P. Individual differences in women's facial preferences as a function of digit ratio and mental rotation ability. Father absence, parent—daughter relationships and partner preferences. Vaughn J. The effect of mortality salience on women's judgments of male faces. Waist—hip ratio predicts women's preferences for masculine male faces, but not perceptions of men's trustworthiness. Female condition influence preferences for sexual dimorphism in faces of male humans Homo sapiens. Psychol , — Self-rated attractiveness predicts individual differences in women's preferences for masculine men's voices. Women's own voice pitch predicts their preferences for masculinity in men's voices. Women's physical and psychological condition independently predict their preference for apparent health in faces. Bakker T. Condition-related mate-choice in sticklebacks. Individual differences in empathizing and systemizing predict variation in face preferences. Sensation seeking and men's face preferences. Extraversion predicts individual differences in women's face preferences. Alvergne A. Differential facial resemblance of young children to their parents: Nesse R. Sex-differences in ability to recognize family resemblance. Bressan P. Parental resemblance in 1-year-olds and the Gaussian curve. Talis pater, talis filius: Kaminski G. Human ability to detect kinship in strangers' faces: Dal Martello M. Where are kin recognition signals in the human face. Maloney L. Kin recognition and the perceived facial similarity of children. Kin recognition signals in adult faces. A cue of kinship promotes cooperation for the public good. Facial resemblance enhances trust. Hamilton W. Bittles A. The costs of human inbreeding and their implications for variations at the DNA level. Buckingham G. SepVisual adaptation to masculine and feminine faces influences generalized preferences and perceptions of trustworthiness. Zajonc R. Attitudinal effects of mere exposure. Integrating gaze direction and expression in preferences for attractive faces. O'Doherty J. Beauty in a smile: Neuropsychologia 41 , — Evidence for adaptive design in human gaze preference. B , 63— Integrating physical and social cues when forming face preferences: Kampe K. Reward value of attractiveness and gaze: Main J. Integrating gaze direction and sexual dimorphism of face shape when perceiving the dominance of others. The evolution of human physical attractiveness. The evolution of human mating: Evolutionary psychology of facial attractiveness. SepFemale condition influences preferences for sexual dimorphism in faces of male humans Homo sapiens. Pathogen prevalence and human mate preferences. Toward an integrative understanding of evoked and transmitted culture: Inquiry 17 , — The health of a nation predicts their mate preferences: Schmitt D. Sociosexuality from Argentina to Zimbabwe: Brooks R. National income inequality predicts women's preferences for masculinized faces better than health does. Further evidence for regional variation in women's masculinity preferences. Geary D. Evolution of human mate choice. Sex Res. Mace R. Evolutionary ecology of human life history. Human preferences for facial masculinity change with relationship type and environmental harshness. Bornstein Exposure and effect: Mere exposure:.

Interested readers can see Andersson [ 3 ] for a thorough review, including issues relating to how preferences may arise in populations. Although we can say whether a Facial attractiveness milffaceness is attractive or unattractive, it is extremely difficult to articulate the specific features that determine this attraction. There are, however, several facial traits that have been proposed to advertise the biological Facial attractiveness milffaceness of an individual in human faces, and hence to influence Facial attractiveness milffaceness as a mate: The former is relevant to both same- and opposite-sex attractiveness judgements, whereas the latter has consequences for reproductive Facial attractiveness milffaceness.

For example, avoiding a parasitized mate has obvious direct advantages whether parasite resistance is heritable or not [ 27 ] as there are direct benefits to choosing a parasite-free mate. Preferences for facial traits that are associated with parasite resistance may be adaptive because this can lead individuals to associate with those who are not carrying contagious parasites which may be passed on to the individual or to the offspring and who are able to act as good parents providing material benefits or care.

Individuals who are attracted to those having face traits associated with parasite resistance may also increase the chances of passing on heritable parasite-resistant genes to their offspring.

In other words, there are several reasons Facial attractiveness milffaceness avoiding a parasitized mate is advantageous.

Ultimately it may be unnecessary to consider the relative weights of indirect and direct benefits; both indirect and direct benefits are likely to be important in evolution and their contributions to attractiveness are difficult to tease apart. We note that much research has focused on women's preferences, although most traits are also relevant for men.

Symmetry refers to the extent to which see more of an object image, organism, etc. Individuals differ in their ability to maintain the stable development of their morphology under the prevailing environmental conditions under which that development is Facial attractiveness milffaceness place [ Facial attractiveness milffaceness29 ].

The ability of an individual more info develop successfully in the face of environmental pressures is therefore one proposed indicator of genetic quality. Read article character demonstrates fluctuating asymmetry FA when symmetry reflects the normal development, and deviations from this symmetry are randomly distributed Facial attractiveness milffaceness respect to side [ 30 ].

Facial attractiveness milffaceness is a particularly useful measure of developmental stability because we know that the optimal developmental outcome is symmetry. Therefore, any deviation from perfect Facial attractiveness milffaceness can be considered a sub-optimal solution which will result in performance problems in the future. FA Facial attractiveness milffaceness also a useful measure as it subsumes a huge amount of individual variation in development, being the outcome of differences in genetic e.

Preferences for symmetry can then, potentially, provide both Facial attractiveness milffaceness e. Whether symmetry is actually related to quality in other animals and humans is an issue addressed by a large literature, and a complete review is not the focus of this paper. While the issue is divided, and there is some evidence that symmetry is not associated with quality e.

In humans, male body symmetry is positively related to sperm number per ejaculate and sperm speed [ 32 ] and female breast symmetry is positively correlated with fecundity [ 3334 ]. Relating to faces, one Facial attractiveness milffaceness has demonstrated that facial asymmetry is positively related to self-reported number of occurrences of respiratory disease [ 35 ] and some studies have observed positive correlations between symmetry and other putative indices of underlying physical condition i.

The relationship between symmetry and quality is not reviewed in detail here, but it should be noted that fitness-related characteristics, such as growth rate, fecundity and survivability, are positively associated with symmetry across a number of species and taxa e.

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Studies of naturally occurring human facial asymmetries also provide evidence that symmetry is found attractive, though such studies can be confounded by potential Facial attractiveness milffaceness.

Studies measuring symmetry from unmanipulated faces source reported positive Facial attractiveness milffaceness with rated attractiveness [ more info — 44 ] and one study has even demonstrated that with pairs of monozygotic twins, Facial attractiveness milffaceness twin with more symmetric measurements is seen as more attractive [ 45 ].

Studies using more sophisticated symmetry manipulations have demonstrated that symmetry can have a positive influence on attractiveness [ 4748 ] and have established that the chimeric manipulations used in the early studies introduced unnatural proportions into the symmetric faces see [ 48 ]. Thus, the methodologically superior computer graphic studies [ 4748 ] parallel the findings of investigations into naturally occurring facial asymmetries Facial attractiveness milffaceness 404143 — 45 ].

The computer graphic studies demonstrate that increasing symmetry alone is sufficient to increase attractiveness. Subsequently, other studies have replicated preferences for symmetry using manipulated stimuli in different Western samples e. Preferences for symmetry using manipulated faces have been Facial attractiveness milffaceness in African hunter—gatherers [ 51 ], and macaque monkeys gaze longer at symmetrical than at asymmetrical face images of conspecifics [ 52 ].

Symmetry and asymmetry. Symmetric images are usually preferred to asymmetric images. Importantly, recent studies have implicated perceptions of health in attraction to symmetric faces [ 4453 ] and read more suggested that the mechanisms underpinning preferences for symmetric faces are different from those that might "Facial attractiveness milffaceness" preferences for symmetry in mate-choice-irrelevant stimuli e.

Such findings suggest that preferences for symmetric faces reflect, at least in part, adaptations for mate choice. Averageness refers to how closely a face resembles the majority of other faces within a population; non-average faces have more Facial attractiveness milffaceness characteristics than the average of a population. Average faces may be attractive because an alignment of features that is close to a population average is linked to genetic diversity [ 5455 ].

Parasites are generally best adapted to proteins that are common in the host population; hence, parasites are adapted to the genes that code for the production of these proteins.

A second evolutionary theory for the attractiveness of averageness in faces is that extreme non-average genotypes are more likely to be homozygous for deleterious alleles, that is, to be more likely to possess genes that are detrimental to an individual than Facial attractiveness milffaceness with more average genotypes [ 54 ].

Both of these theories propose evolutionary benefits to mating with individuals possessing average faces. Recent studies have supported the link between averageness, heterozygosity i. Heterozygosity in the major histocompatibility complex MHC genes that code for proteins involved in immune response, is positively associated with Facial attractiveness milffaceness attractiveness [ 56 ] and facial averageness [ 57 ].

More directly, another study has shown that Facial attractiveness milffaceness averageness is positively related to medical health as measured from actual medical records in both men and women Facial attractiveness milffaceness 58 ]. There is good evidence that average faces are indeed found attractive. Galton [ 59 ] first noted that multiple faces blended together were more attractive than the constituent faces. Recent studies have improved upon these techniques using computers to create digitally blended composite faces; generally, the more images in a composite, the more attractive it Facial attractiveness milffaceness found [ 60 — 62 ].

Aside from composite images, Light et al. Average faces are generally more symmetric and symmetry is typically attractive in faces discussed in more detail above. Several studies have controlled for this confound in the original studies. When Facial attractiveness milffaceness and symmetry were independently manipulated, one study found that both manipulations positively and independently influenced Facial attractiveness milffaceness judgements [ 65 ].

Other studies have used perfectly symmetric images manipulated in averageness and still have demonstrated preferences for averageness [ 6667 ].

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Indeed, by comparing preferences for averageness when the effects of symmetry were controlled for and were not controlled for, Jones et al. It has also been noted that, in the original Facial attractiveness milffaceness studies, the more images that are blended together the smoother Facial attractiveness milffaceness skin texture becomes, as imperfections such as lines or blemishes are averaged [ 68 ]. Image c should be more attractive than both of the other images.

Composites are made by marking key locations around the main facial features e. The average location of each point of the component faces is then calculated to define the shape of the composite.

The images of the individual faces are then warped to Facial attractiveness milffaceness relevant average shape before superimposing the images to visit web page a photographic quality composite image. While the majority of the work described above has been carried out in Facial attractiveness milffaceness America, Britain and Australia, averageness has also been found to be attractive across different cultures.

Hotels Mms Watch Video Pamelamix porn. Importantly, exposure in the manner described above also influences attractiveness judgements. After exposure to faces possessing certain traits, these traits come to be preferred [ , , , ]. For example, if exposed to faces that look more like one identity, then new faces that resemble that identity are found more attractive than if exposed to the opposite set of face traits. A similar effect has also been observed for judgements of the trustworthiness of faces [ ]. Adaptation then reflects the rapid updating of face norms and can therefore be tied both to the effects of familiarity and imprinting-like effects. We have dealt briefly with some aspects of simple experience on preferences above, but, of course, humans are highly social and much human experience is of what other humans do. Humans can therefore learn about attractiveness from the behaviour of those around them: We have recently reviewed social learning in human face preferences [ ], and so present a brief overview here. Social learning can be adaptive if it allows an individual to assess potential mates more quickly and efficiently than through individual trial and error or allows an individual to use another's expertise. Mate choice copying has been observed among females in a number of different non-human species [ — ], including fish [ — ] and bird species [ — ]. Such studies have generally shown that when females observe another female the model to be paired with one of the two males the targets , they are subsequently more likely to prefer the target male they had seen paired with the model over the male that was not paired with the model. Inspired by work on non-human species, recent research also suggests that social learning may influence human mate preferences. While some research has shown that the presence of wedding rings on men did not increase women's preferences for those men [ ], other studies have found that images of men labelled as married were more attractive than those labelled as single [ ] and that women rate men as more desirable when they are shown surrounded by women than when they are shown alone or with other men [ ]. Another study has shown that women prefer pictures of men that had been previously seen paired with images of other women who were looking at the face with smiling i. Women therefore do appear to mimic the attitude of other women to particular men. Alongside partnership status, simple presence, and expressions of attitude towards the male, the physical traits of the observed model may also play a role in the social transmission of preference. Previous studies have shown that men's and women's attractiveness judgements are influenced by the apparent choice of attractive members of the same sex. Such a phenomenon suggests a more sophisticated form of mate-choice copying, whereby women can use the attractiveness of a partner that a man can acquire in order to judge the man's own attractiveness. Another study using images that were presented with a fictitious partner has shown that both men and women find a face paired with an attractive partner to be more attractive than one paired with an unattractive partner for a long-term but not a short-term relationship [ ]. Being more or less attractive has important social consequences and people do generally agree on who is and who is not attractive. Beauty is not just a simple social construct—attractiveness appears to be ingrained in our biology. While some aspects of face perception might be innate, other aspects are clearly influenced by experience; it seems unlikely that individuals are born with a representation of what a perfect partner looks like. If a trait reliably advertises some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive. It is clear that individual differences in preferences for some traits will prove adaptive and so are consistent with evolutionary theory. Work on facial attractiveness is also integrative, combining theories and methods from behavioural ecology, cognition, cross-cultural research and social psychology. Login to your account. Forgot password? Keep me logged in. New User. Change Password. Old Password. New Password. Create a new account. Returning user. Can't sign in? Forgot your password? Enter your email address below and we will send you the reset instructions. If the address matches an existing account you will receive an email with instructions to reset your password Close. Request Username. Forgot your username? Enter your email address below and we will send you your username. You have access. View PDF. Little Anthony C. Facial attractiveness: Biological Sciences http: You have access Articles. Anthony C. Benedict C. Jones Benedict C. Lisa M. DeBruine Lisa M. Abstract Face preferences affect a diverse range of critical social outcomes, from mate choices and decisions about platonic relationships to hiring decisions and decisions about social exchange. Download figure Open in new tab Download PowerPoint. References 1 Buss D. Langlois J. A meta-analytic and theoretical review. Andersson M. Princeton, NJ: Princeton University Press. Elder G. Holmes S. Riggio R. Berscheid E. Walster E. Dion K. Eagly A. Feingold A. Cash T. Sigall H. Downs A. B 17 , — Chiu R. Marlowe C. Wolf N. New York, NY: Hume D. Of the standard of taste. London, UK: Darwin C. John Murray. Advances in experimental social psychology ed. Academic Press. Ford C. Cunningham M. Zebrowitz-McArthur L. Thornhill R. Trends Cogn. Rhodes G. Gibson R. Moller A. Oxford, UK: Oxford University Press. Valen L. Evolution 16 , — Dufour K. Manning J. Gangestad S. Little A. Grammer K. Scheib J. B , — Penton-Voak I. Jones B. Perception 33 , — Mealey L. Kowner R. Human 22 , — Perrett D. Waitt C. Perception 36 , — Mitton J. Roberts S. Lie H. Evolution 62 , — Galton F. Nature 18 , 97— Light L. B 7 , — Apicella C. Alley T. Perception 30 , — Enlow D. Philadelphia, PA: Zahavi A. Hillgarth N. Parasites and pathogens ed. Kanda N. Yesilova Z. Folstad I. B , S93—S Law-Smith M. Neave N. Jones D. Nature , — Facialmetric assessment of multiple motives in the perception of male facial physical attractiveness. Berry D. Personality correlates of facial babyishness. B 15 , — Keating C. McArthur L. Swaddle J. B , 39— DeBruine L. Smith F. Krupp D. In press. Apparent health encourages reciprocity. Fink B. Millinski M. Pryke S. Setchell J. Ethology , 25— B , S—S Changizi M. Drummond P. Psychophysiology 38 , — Cuthill I. Hill R. Nature , Burley N. Elliot A. B 35 , — Stephen I. Hassin R. Otta E. Motor Skills 82 , — Buss D. Brain Sci. Borkenau P. Liggett J. Botwin M. Boothroyd L. Korthase K. Motor Skills 54 , — Coetzee V. Perception 38 , — Osborn D. Makeup and posture effects on physical attractiveness judgments. Saxton T. Johnston V. Puts D. Feinberg D. Havlicek J. Psychoneuroendocrinology 35 , — Vukovic J. Peters M. Welling L. Roney J. Johnston L. Macrae C. Oinonen K. Sexual Behav. Pepper G. Derntl B. Conway C. Harris C. Menstrual cycle and facial preferences reconsidered. Sex Roles. Scarbrough P. Vaughn J. Psychol , — Bakker T. Alvergne A. Nesse R. There are many traits that are linked to facial attractiveness in humans and each may in some way impart benefits to individuals who act on their preferences. If a trait is reliably associated with some benefit to the perceiver, then we would expect individuals in a population to find that trait attractive. Such an approach has highlighted face traits such as age, health, symmetry, and averageness, which are proposed to be associated with benefits and so associated with facial attractiveness. This view may postulate that some traits will be universally attractive; however, this does not preclude variation. MHC-heterozygosity and human facial attractiveness. Lie H. Genetic diversity revealed in human faces. Evolution 62 , — Do facial averageness and symmetry signal health? Galton F. Composite portraits. Nature 18 , 97— Attractive faces are only average. What is average and what is not average about attractive faces. The role of masculinity and distinctiveness in judgments of human male facial attractiveness. Light L. Why attractive people are harder to remember. B 7 , — Averageness, exaggeration, and facial attractiveness. Are average facial configurations attractive only because of their symmetry? The role of symmetry in attraction to average faces. Apicella C. Facial averageness and attractiveness in an isolated population of hunter-gatherers. Alley T. Averaged faces are attractive, but very attractive faces are not average. Attractiveness of facial averageness and symmetry in non-Western populations: Perception 30 , — Enlow D. Handbook of facial growth , 2nd edn. Philadelphia, PA: Saunders [ Google Scholar ]. Zahavi A. Mate selection: Hillgarth N. Testosterone and immunosuppression in vertebrates: In Parasites and pathogens ed. Beckage N. Kanda N. Testosterone inhibits immunoglobulin production by human peripheral blood mononuclear cells. Yesilova Z. The effects of gonadotropin treatment on the immunological features of male patients with idiopathic hypogonadotropic hypogonadism. Folstad I. Parasites, bright males and the immunocompetence handicap. Parasitism, host immune function, and sexual selection. Does sexual dimorphism in human faces signal health? B , S93—S Law-Smith M. Facial appearance is a cue to oestrogen levels in women. High salivary testosterone is linked to masculine male facial appearance in humans. Neave N. Second to fourth digit ratio, testosterone and perceived male dominance. Measuring the physical in physical attractiveness: Jones D. Criteria of facial attractiveness in five populations. Effects of sexual dimorphism on facial attractiveness. Nature , — What do women want? Facialmetric assessment of multiple motives in the perception of male facial physical attractiveness. Berry D. Were the physiognomists right? Personality correlates of facial babyishness. B 15 , — Keating C. Gender and the physiognomy of dominance and attractiveness. McArthur L. Impressions of baby-faced adults. Cross-cultural agreement in perceptions of babyfaced adults. Some components and consequences of a babyface. Sex-typicality and attractiveness: Swaddle J. Testosterone increases perceived dominance but not attractiveness in human males. Self-perceived attractiveness influences human female preferences for sexual dimorphism and symmetry in male faces. B , 39— Partnership status and the temporal context of relationships influence human female preferences for sexual dimorphism in male face shape. DeBruine L. Correlated preferences for facial masculinity and ideal or actual partner's masculinity. Viewing attractive or unattractive same-sex individuals changes self-rated attractiveness and face preferences in women. Smith F. Interactions between masculinity-femininity and apparent health in face preferences. Menstrual cycle, pregnancy and oral contraceptive use alter attraction to apparent health in faces. Krupp D. In press. Apparent health encourages reciprocity. Fink B. Visible skin color distribution plays a role in the perception of age, attractiveness, and health in female faces. Millinski M. Female sticklebacks use male coloration in sticklebacks and therefore avoid parasitised males. Pryke S. Red dominates black: Setchell J. Dominance, status signals and coloration in male mandrills Mandrillus sphinx. Ethology , 25— Evidence from rhesus macaques suggests that male coloration plays a role in female primate mate choice. B , S—S Changizi M. Bare skin, blood and the evolution of primate colour vision. Drummond P. The effect of expressing anger on cardiovascular reactivity and facial blood flow in Chinese and Caucasians. Psychophysiology 38 , — Selective attention toward female secondary sexual color in male rhesus macaques. Do female mandrills prefer brightly colored males? Cuthill I. Colour bands, dominance, and body mass regulation in male zebra finches Taeniopygia guttata. Hill R. Red enhances human performance in contests. Nature , Burley N. Influence of color-banding on the conspecific preferences of zebra finches. Social perception of red suggests special role in dominance signalling. Elliot A. The effect of red on avoidance behavior in achievement contexts. B 35 , — Romantic red: Stephen I. Facial skin coloration affects perceived health of human faces. Hassin R. Facing faces: Otta E. Reading a smiling face: A cross-cultural exploration of physiognomic traits of dominance and happiness. Sex differences in human mate preferences: Brain Sci. Borkenau P. Trait inferences: Liggett J. The human face. Constable [ Google Scholar ]. What is good is beautiful: Botwin M. Personality and mate preferences: Male facial appearance signals physical strength to women. Facial cues of dominance modulate the short-term gaze-cuing effect in human observers. Boothroyd L. Facial correlates of sociosexuality. Korthase K. Perceived age and perceived physical attractiveness. Motor Skills 54 , — [ Google Scholar ]. Coetzee V. Facial adiposity: Perception 38 , — Investigating an imprinting-like phenomenon in humans: The effects of facial hair manipulation on female perceptions of attractiveness, masculinity, and dominance in male faces. Osborn D. 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For example, facial Facial attractiveness milffaceness is also found attractive in Japanese participants [ 69 ] and in African hunter—gatherers [ 67 ]. Male and female faces differ in their shape. Mature features in adult human faces reflect the masculinization or feminization of secondary sexual characteristics that Facial attractiveness milffaceness at puberty. These face shape differences, in part, arise because of the action of hormones such as testosterone.

Larger jawbones, more prominent cheekbones and thinner cheeks are Facial attractiveness milffaceness features of male faces that differentiate them from female faces e. From an evolutionary view, extremes of secondary sexual characteristics more Facial attractiveness milffaceness for women, more masculine for men are proposed to be attractive because they advertise the quality of an individual in terms of heritable benefits; they indicate that the owners of such characteristics possess good genes.

In other words, such traits advertise the possession of genes that are beneficial to offspring inheriting them continue reading terms of survival or reproduction. One explanation of the importance of these facial traits is that they represent a handicap to an organism [ 71 ] and the Facial attractiveness milffaceness of growing the trait means that only healthy individuals can afford to produce them.

For example, secondary sexual characteristics are proposed to be linked to parasite resistance because the sex hormones that influence their growth, particularly testosterone, lower immunocompetence. Testosterone has been linked to the suppression of immune function in many species [ 72 ], including Crystal steal [ 7374 ].

Larger secondary sexual characteristics should be related to a healthier immune system because only healthy organisms can afford the high sex-hormone handicap on the immune system that is necessary to produce these characteristics [ 75 ]. In many non-human animal studies, there is a positive Facial attractiveness milffaceness between secondary sexual trait expression and immunocompetence e. A study by Rhodes et al. Large anus photos.

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