The evolutionary psychology of physical attractiveness: Sexual selection and human morphology

Psychological evidence suggests that sex differences in morphology have been modified by sexual selection so as to attract mates (intersexual selection) or intimidate rivals (intrasexual selection). Women compete with each other for high quality husbands by advertising reproductive value in terms of the distribution of fat reserves and by exaggerating morphological indicators of youthfulness such as a small nose and small feet and pale, hairless skin. Men's physical appearance tends to communicate social dominance, which has the combined effects of intimidating reproductive rivals and attracting mates. In addition to their attractiveness and intimidatory effects, human secondary sexual characters also provide cues to hormonal status and phenotypic quality consistent with the good genes model of sexual selection (which includes parasite resistance). Low waist-hip ratio is sexually attractive in women and indicates a high estrogen/testosterone ratio (which favors reproductive function). Facial attractiveness provides honest cues to health and mate value. The permanently enlarged female breast appears to have evolved under the influence of both the good genes and the runaway selection mechanisms. The male beard is not obviously related to phenotypic quality and may have evolved through a process of runaway intersexual selection.     

Cross-cultural consensus for waist–hip ratio and women's attractiveness

In women of reproductive age, a gynoid body fat distribution as measured by the size of waist–hip ratio (WHR) is a reliable indicator of their sex hormone profile, greater success in pregnancy and less risk for major diseases. According to evolutionary mate selection theory, such indicators of health and fertility should be judged as attractive. Previous research has confirmed this prediction. In this current research, we use the same stimulus for diverse racial groups (Bakossiland, Cameroon, Africa; Komodo Island, Indonesia; Samoa; and New Zealand) to examine the universality of relationships between WHR and attractiveness. As WHR is positively correlated with body mass index (BMI), we controlled BMI by using photographs of women who have gone through micrograft surgery for cosmetic reasons. Results show that in each culture participants selected women with low WHR as attractive, regardless of increases or decreases in BMI. This cross-cultural consensus suggests that the link between WHR and female attractiveness is due to adaptation shaped by the selection process.     

Human facial beauty

It is hypothesized that human faces judged to be attractive by people possess two features—averageness and symmetry—that promoted adaptive mate selection in human evolutionary history by way of production of offspring with parasite resistance. Facial composites made by combining individual faces are judged to be attractive, and more attractive than the majority of individual faces. The composites possess both symmetry and averageness of features. Facial averageness may reflect high individual protein heterozygosity and thus an array of proteins to which parasites must adapt. Heterozygosity may be an important defense of long-lived hosts against parasites when it occurs in portions of the genome that do not code for the essential features of complex adaptations. In this case heterozygosity can create a hostile microenvironment for parasites without disrupting adaptation. Facial bilateral symmetry is hypothesized to affect positive beauty judgments because symmetry is a certification of overall phenotypic quality and developmental health, which may be importantly influenced by parasites. Certain secondary sexual traits are influenced by testosterone, a hormone that reduces immunocompetence. Symmetry and size of the secondary sexual traits of the face (e.g., cheek bones) are expected to correlate positively and advertise immunocompetence honestly and therefore to affect positive beauty judgments. Facial attractiveness is predicted to correlate with attractive, nonfacial secondary sexual traits; other predictions from the view that parasite-driven selection led to the evolution of psychological adaptations of human beauty perception are discussed. The view that human physical attractiveness and judgments about human physical attractiveness evolved in the context of parasite-driven selection leads to the hypothesis that both adults and children have a species-typical adaptation to the problem of identifying and favoring healthy individuals and avoiding parasite-susceptible individuals. It is proposed that this adaptation guides human decisions about nepotism and reciprocity in relation to physical attractiveness.     

Visible skin color distribution plays a role in the perception of age, attractiveness, and health in female faces

Evolutionary psychologists have proposed that preferences for facial characteristics, such as symmetry, averageness, and sexual dimorphism, may reflect adaptations for mate choice because they signal aspects of mate quality. Here, we show that facial skin color distribution significantly influences the perception of age and attractiveness of female faces, independent of facial form and skin surface topography. A set of three-dimensional shape-standardized stimulus faces—varying only in terms of skin color distribution due to variation in biological age and cumulative photodamage—was rated by a panel of naive judges for a variety of perceptual endpoints relating to age, health, and beauty. Shape- and topography-standardized stimulus faces with the homogeneous skin color distribution of young people were perceived as younger and received significantly higher ratings for attractiveness and health than analogous stimuli with the relatively inhomogeneous skin color distribution of more elderly people. Thus, skin color distribution, independent of facial form and skin surface topography, seems to have a major influence on the perception of female facial age and judgments of attractiveness and health as they may signal aspects of underlying physiological condition of an individual relevant for mate choice. We suggest that studies on human physical attractiveness and its perception need to consider the influence of visible skin condition driven by color distribution and differentiate between such effects and beauty-related traits due to facial shape and skin topography.     

Genetic diversity revealed in human faces

From an evolutionary perspective, human facial attractiveness is proposed to signal mate quality. Using a novel approach to the study of the genetic basis of human preferences for facial features, we investigated whether attractiveness signals mate quality in terms of genetic diversity. Genetic diversity in general has been linked to fitness and reproductive success, and genetic diversity within the major histocompatibility complex (MHC) has been linked to immunocompetence and mate preferences. We asked whether any preference for genetic diversity is specific to MHC diversity or reflects a more general preference for overall genetic diversity. We photographed and genotyped 160 participants using microsatellite markers situated within and outside the MHC, and calculated two measures of genetic diversity: mean heterozygosity and standardized mean d(2). Our results suggest a special role for the MHC in female preferences for male faces. MHC heterozygosity positively predicted male attractiveness, and specifically facial averageness, with averageness mediating the MHC-attractiveness relationship. For females, standardized mean d(2) at non-MHC loci predicted facial symmetry. Thus, attractive facial characteristics appear to provide visual cues to genetic quality in both males and females, supporting the view that face preferences have been shaped by selection pressures to identify high-quality mates.     

Patterns of subcutaneous fat deposition and the relationship between body mass index and waist-to-hip ratio: Implications for models of physical attractiveness

Body mass index (BMI) and waist-to-hip ratio (WHR) are two widely used anthropometric indices of body shape argued to convey different information about health and fertility. Both indices have also been shown to affect attractiveness ratings of female bodies. However, BMI and WHR are naturally positively correlated, complicating studies designed to identify their relative importance in predicting health and attractiveness outcomes. We show that the correlation between BMI and WHR depends on the assumed model of subcutaneous fat deposition. An additive model, whereby fat is added to the waist and hips at a constant rate, predicts a correlation between BMI and WHR because with increasing fat, the difference between the waist and hips becomes smaller relative to total width. This model is supported by longitudinal and cross-sectional data. We parameterised the function relating WHR to BMI for white UK females of reproductive age, and used this function to statistically decompose body shape into two independent components. We show that judgements of the attractiveness of female bodies are well explained by the component of curvaceousness related to BMI but not by residual curvaceousness. Our findings resolve a long-standing dispute in the attractiveness literature by confirming that although WHR appears to be an important predictor of attractiveness, this is largely explained by the direct effect of total body fat on WHR, thus reinforcing the conclusion that total body fat is the primary determinant of female body shape attractiveness.     

Female health,attractiveness, and desirability for relationships: Role of breast asymmetry and waist-to-hip ratio

Fluctuating asymmetry (FA) of ordinarily bilaterally symmetrical traits in humans has been proposed to indicate developmental anomaly. Recent research has shown that individuals with minimal FA are judged to be attractive, and preferred as sexual partners (Thornhill and Gangestad 1993). Waist-to-hip ratios (WHR) have been also shown to reflect health and reproductive capability of woman and those with low WHRs are judged more attractive and healthy (Singh 1993a,b). The present study examines the relative roles of WHR and FA in female breasts in judgments of female attractiveness, health, and desirability for short-term and long-term relationships. Male college students were asked to judge attractiveness of female figures that differed in WHR (high and low) and breast symmetry (none, low, or high). In the first test, paired comparison method was used in which each figure was paired one at a time with all other figures. In the second test, subjects examined all figures simultaneously, estimated their age, and rated each figure for attractiveness, feminine looks, health, overall degree of body symmetry, and willingness to engage in short- and long-term relationship. Results from both tests show that figures with low WHRs were judged to be more attractive than figures with high WHRs, regardless of their degree of breast asymmetry. The figure with low WHR and symmetrical breasts was judged to be most attractive and youngest of all other figures. It appears that men use both WHR and breast asymmetry in judging attractiveness and being willing to develop romantic relationships. It is proposed that WHR and breast asymmetry may signal different aspects of overall female mate quality.     

African and European perception of African female attractiveness

Majority of research on attractiveness is restricted to faces of European origin. The perception of attractiveness may, however, vary across communities due to variations in both facial morphology and local standards of beauty. We investigated the relative contribution of four facial markers of attractiveness based on 101 female facial portraits (standardized, non-manipulated) from Cameroon and Namibia, which were assessed by local male raters and by raters from a distant European population, the Czech Republic. Images from Cameroon include only women of Bantu origin, while Namibians are represented by women of both Bantu (Owambo/Herero) and Nama origin. While controlling for age and BMI, we explored the relationship between female attractiveness and a set of facial traits: fluctuating asymmetry, averageness, shape sexual dimorphism, and skin color (rated and measured in CIELab color space).In the Cameroonian sample, local male raters favored lighter-skinned female faces with morphology closer to average. The attractiveness of Nama women as rated by Nama men positively correlated with lighter complexion, but this did not extend to rating by Cameroonian men. The attractiveness of Namibian Owambo/Herero women was positively associated with facial femininity and lighter complexion when judged by both Cameroonian and Nama male raters. In all samples, the attractiveness as rated by Czech men was predicted by age and BMI, but not by skin color. We found no significant association between attractiveness and fluctuating asymmetry in any of the tested samples. When controlling for age, the effect of skin color on attractiveness turned to be non-significant in the Owambo/Herrero and Nama sample, but remained significant in the Cameroonian sample. Variations in skin color thus represent an important factor of African female attractiveness within the African context, but they do not seem to affect judgements made by European raters. Sensitivity to some facial markers of female attractiveness thus seems to be restricted to regional populations and/or constrained by shared ethnicity.     

Visual cues to female physical attractiveness

Evolutionary psychology suggests that a woman's sexual attractiveness is based on cues of health and reproductive potential. In recent years, research has focused on the ratio of the width of the waist to the width of the hips (the waist-to-hip ratio (WHR). A low WHR (i.e. a curvaceous body) is believed to correspond to the optimal fat distribution for high fertility, and so this shape should be highly attractive. In this paper we present evidence that weight scaled for height (the body mass index (BMI)) is the primary determinant of sexual attractiveness rather than WHR. BMI is also strongly linked to health and reproductive potential. Furthermore, we show how covariation of apparent BMI and WHR in previous studies led to the overestimation of the importance of WHR in the perception of female attractiveness. Finally, we show how visual cues, such as the perimeter-area ratio (PAR), can provide an accurate and reliable index of an individual's BMI and could be used by an observer to differentiate between potential partners.     

Female mate value at a glance: relationship of waist-to-hip ratio to health,fecundity and attractiveness

A fundamental assumption of adaptive explanations of female attractiveness is that bodily features that males judge as attractive reliably signal youthfulness, healthiness, and fertility or female mate value. One of the bodily features, waist-to-hip ratio (WHR), is a reliable indicator of a female's reproductive age, sex hormone profile, parity and risk for various diseases. Systematic variation in the size of WHR also systematically affects the judgment of female attractiveness, healthiness, and youthfulness. This article summarizes recent findings about the relationship between female's WHR and various factors affecting reproductive capability and risk for diseases. Research on the relationship between attractiveness and WHR is discussed in light of some methodological objections to previous research. Finally, cross-cultural and historical data are presented that suggest that the relationship between WHR and female attractiveness is not culture-specific and not inculcated by modern Western fashion dictates or media.     

The relative importance of intra- and intersexual selection on human male sexually dimorphic traits

Recent evidence suggests that in sexual selection on human males, intrasexual competition plays a larger role than female choice. In a sample of men (N?=?164), we sought to provide further evidence on the effects of men's physical dominance and sexual attractiveness on mating success and hence in sexual selection. Objective measures and subjective ratings of male sexually dimorphic traits purportedly under sexual selection (height, vocal and facial masculinity, upper body size from 3D scans, physical strength, and baseline testosterone) and observer perceptions of physical dominance and sexual attractiveness based on self-presentation video recordings were assessed and associated with mating success (sociosexual behaviour and number of potential conceptions) in a partly longitudinal design. Results from structural equation models and selection analyses revealed that physical dominance, but not sexual attractiveness, predicted mating success. Physical dominance mediated associations of upper body size, physical strength, as well as vocal and facial physical dominance and attractiveness with mating success. These findings thus suggest a greater importance of intrasexual competition than female choice in human male sexual selection.     

BMI not WHR modulates BOLD fMRI responses in a sub-cortical reward network when participants judge the attractiveness of human female bodies

In perceptual terms, the human body is a complex 3d shape which has to be interpreted by the observer to judge its attractiveness. Both body mass and shape have been suggested as strong predictors of female attractiveness. Normally body mass and shape co-vary, and it is difficult to differentiate their separate effects. A recent study suggested that altering body mass does not modulate activity in the reward mechanisms of the brain, but shape does. However, using computer generated female body-shaped greyscale images, based on a Principal Component Analysis of female bodies, we were able to construct images which covary with real female body mass (indexed with BMI) and not with body shape (indexed with WHR), and vice versa. Twelve observers (6 male and 6 female) rated these images for attractiveness during an fMRI study. The attractiveness ratings were correlated with changes in BMI and not WHR. Our primary fMRI results demonstrated that in addition to activation in higher visual areas (such as the extrastriate body area), changing BMI also modulated activity in the caudate nucleus, and other parts of the brain reward system. This shows that BMI, not WHR, modulates reward mechanisms in the brain and we infer that this may have important implications for judgements of ideal body size in eating disordered individuals.     

Body shape and women’s attractiveness

This paper examines the role of body fat distribution as measured by waist-to-hip ratio (WHR) on the judgment of women’s physical attractiveness. It presents evidence that WHR is correlated with a woman’s reproductive endocrinological status and long-term health risk. Three studies were conducted to investigate whether humans have perceptual and cognitive mechanisms to utilize the WHR to infer attributes of women’s health, youthfulness, attractiveness, and reproductive capacity. College-age as well as older subjects of both sexes rank female figures with normal weight and low WHR as attractive and assign to them higher reproductive capability. The study concludes that WHR is a reliable and honest signal of a woman’s reproductive potential. The adaptive significance of body fat distribution and its role in mate selection is also discussed.     

No evidence that facial attractiveness,femininity, averageness,or coloration are cues to susceptibility to infectious illnesses in a university sample of young adult women

Previous reports that women with attractive faces are healthier have been widely cited as evidence that sexual selection has shaped human mate preferences. However, evidence for correlations between women's physical health and facial attractiveness is equivocal. Moreover, positive results on this issue have generally come from studies of self-reported health in small samples. The current study took standardized face photographs of women who completed four different health questionnaires assessing susceptibility to infectious illnesses (N?=?590). Of these women, 221 also provided a saliva sample that was assayed for immunoglobulin A (a marker of immune function). Analyses showed no significant correlations between rated facial attractiveness and either scores on any of the health questionnaires or salivary immunoglobulin A. Furthermore there was no compelling evidence that objective measures of sexual dimorphism of face shape, averageness of face shape, or facial coloration were correlated with any of our health measures. While other measures of health may yet reveal robust associations with facial appearance, these null results do not support the prominent and influential assumption that women's facial attractiveness is a cue of young adult women's susceptibility to infectious illnesses, at least in our study population.     

Offspring sex ratio produced by female guppies in the wild correlates with sexual ornaments of their sons

The attractiveness hypothesis predicts that females produce broods with male-biased sex ratios when they mate with attractive males. This hypothesis presumes that sons in broods with male-biased sex ratios sired by attractive males have high reproductive success, whereas the reproductive success of daughters is relatively constant, regardless of the attractiveness of their sires. However, there is little direct evidence for this assumption. We have examined the relationships between offspring sex ratios and (1) sexual ornamentation of sons and (2) body size of daughters in broods from wild female guppies Poecilia reticulata. Wild pregnant females were collected and allowed to give birth in the laboratory. Body size and sexual ornamentation of offspring were measured at maturity. Our analysis revealed a significant positive correlation between offspring sex ratios (the proportion of sons per brood) and the total length as well as the area of orange spots of sons, two attributes that influence female mating preferences in guppies. The sex ratio was not associated with the body size of daughters. These results suggest that by performing adaptive sex allocation according to the expected reproductive success of sons and daughters, female guppies can enhance the overall fitness of their offspring.     

Visual perception of female physical attractiveness

On the basis of visual assessment of figure drawings and front/profile images, past researchers believed that the waist-hip ratio (WHR) and the body mass index (BMI) were two putative cues to female physical attractiveness. However, this view was not tested on three-dimensional (3D) female images. In the present study, 3D images of 31 Caucasian females having varying body weights (BMI ranged from 16 to 35) were shown to 29 male and 25 female viewers, who were asked to rate the physical attractiveness. The results showed that the body volume divided by the square of the height, defined as volume height index (VHI), is the most important and direct visual determinant of female physical attractiveness. In determining the female attractiveness, human observers may first use VHI as a visual cue, which is also a key indicator of health and fertility owing to its strong linear relation to BMI. To fine-tune the judgement, observers may then use body proportions, the most important of which are the ratio of waist height over the chin height (WHC) (a measure of the length of legs over total tallness) and the deviation of WHR from the ideal ratio. It also appears that the effect of the body's physical parameters on the perception of female physical attractiveness conforms to Stevens' power law of psychophysics.     

Facial attractiveness and fertility in populations with low levels of modern birth control

Evolutionary models of human mate choice generally assume that physical attractiveness has evolved through sexual selection, i.e., it has been associated with higher mating opportunities and subsequent reproductive success across our evolutionary history. Here we investigate whether facial attractiveness is related to fertility in order to understand the extent to which selection can operate on attractive traits in modern populations. We used data from two populations where the prevalence of modern birth control methods is low and thus unlikely to disconnect mating opportunities from reproductive success: men and women from contemporary rural Senegal and men from the West Point Military Academy in the USA who graduated in 1950. We found that facial attractiveness negatively predicts age-specific reproduction in both sexes in Senegal and is independent from lifetime reproductive success in men from the USA. Overall, the results suggest that facial attractiveness is not under positive selection and raise questions about methodological approaches currently used to assess attractiveness.     

Hormonal and morphological predictors of women’s body attractiveness

Does women’s body attractiveness predict indices of reproductive capacity? Prior research has provided evidence that large breast size and low waist-to-hip ratio (WHR) are positively associated with women’s estrogen and progesterone concentrations, but no previous studies appear to have directly tested whether ratings of women's body attractiveness are predicted by higher concentrations of ovarian hormones measured across broad regions of the menstrual cycle. Here, we collected daily saliva samples across 1–2 menstrual cycles from a sample of young women; assayed the samples for estradiol, progesterone, and testosterone; obtained anthropometric measurements of the women’s bodies; and also obtained attractiveness ratings of the women’s bodies from photographs of them taken in standardized clothing with faces obscured. Contrary to previous research, mean hormone concentrations were uncorrelated with breast size and WHR. Body mass index (BMI) was a very strong negative predictor of body attractiveness ratings, similar to previous findings. Zero-order associations between women’s mean hormone concentrations and mean attractiveness ratings were not significant; however, after controlling for BMI, attractiveness ratings were independently and positively associated with both estradiol and testosterone concentrations. Discussion focuses on the implications of these findings for whether attractiveness assessment mechanisms are specialized for the detection of cues of differential fecundity in young women’s bodies.