Symmetry and Human Facial Attractiveness

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

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.     

Facial attractiveness, symmetry and cues of good genes.

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

Symmetrical decorations enhance the attractiveness of faces and abstract designs

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

Evidence against perceptual bias views for symmetry preferences in human faces

Symmetrical human faces are attractive. Two explanations have been proposed to account for symmetry preferences: (i) the evolutionary advantage view, which posits that symmetry advertises mate quality and (ii) the perceptual bias view, which posits that symmetry preferences are a consequence of greater ease of processing symmetrical images in the visual system. Here, we show that symmetry preferences are greater when face images are upright than when inverted. This is evidence against a simple perceptual bias view, which suggests symmetry preference should be constant across orientation about a vertical axis. We also show that symmetry is preferred even in familiar faces, a finding that is unexpected by perceptual bias views positing that symmetry is only attractive because it represents a familiar prototype of that particular class of stimuli.     

Higher-level mechanisms detect facial symmetry

The role of symmetry detection in early visual processing and the sensitivity of biological visual systems to symmetry across a wide range of organisms suggest that symmetry can be detected by low-level visual mechanisms. However, computational and functional considerations suggest that higher-level mechanisms may also play a role in facial symmetry detection. We tested this hypothesis by examining whether symmetry detection is better for faces than comparable patterns, which share low-level properties with faces. Symmetry detection was better for upright faces than for inverted faces (experiment 1) and contrast-reversed faces (experiment 2), implicating high-level mechanisms in facial symmetry detection. In addition, facial symmetry detection was sensitive to spatial scale, unlike low-level symmetry detection mechanisms (experiment 3), and showed greater sensitivity to a 45 degrees deviation from vertical than is found for other aspects of face perception (experiment 4). These results implicate specialized, higher-level mechanisms in the detection of facial symmetry. This specialization may reflect perceptual learning resulting from extensive experience detecting symmetry in faces or evolutionary selection pressures associated with the important role of facial symmetry in mate choice and 'mind-reading' or both.     

Symmetry, sexual dimorphism in facial proportions and male facial attractiveness

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

Preferences for Symmetry in Conspecific Facial Shape Among Macaca mulatta

In human males and females, bilateral symmetry of facial shape influences assessments of attractiveness. It is possible, however, that other primate species also possess preferences for conspecific facial symmetry. To assess this experimentally, we presented 13 adult rhesus macaques (8 females, 5 males) with computer-manipulated images of symmetrical and asymmetrical versions of opposite-sexed conspecific faces. We utilized looking behavior to assess visual preferences for these factors. We found significant preferences for symmetry, raising the possibility that human preferences for facial symmetry are more deeply rooted in our evolutionary history than previously realized. Our results also have implications for the use of facial shape as a mechanism for attractiveness appraisals across the Primates.     

Facial masculinity and fluctuating asymmetry

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

Attraction independent of detection suggests special mechanisms for symmetry preferences in human face perception

Symmetrical human faces are attractive and it has been proposed that humans have a specialized mechanism for detecting symmetry in faces and that sensitivity to symmetry determines symmetry preferences. Here, we show that symmetry preferences are influenced by inversion, whereas symmetry detection is not and that within individuals the ability to detect facial symmetry is not related to preferences for facial symmetry. Taken together, these findings suggest that symmetry preferences are indeed driven by a mechanism that is independent of conscious detection. A specialized mechanism for symmetry preference independent of detection may be the result of specific pressures faced by human ancestors to select high-quality mates and could support a modular view of mate choice. Unconscious mechanisms determining face preferences may explain why the reasons behind attraction are often difficult to articulate and demonstrate that detection alone cannot explain symmetry preferences.     

Are faces of different species perceived categorically by human observers?

What are the species boundaries of face processing? Using a face-feature morphing algorithm, image series intermediate between human, monkey (macaque), and bovine faces were constructed. Forced-choice judgement of these images showed sharply bounded categories for upright face images of each species. These predicted the perceptual discrimination boundaries for upright monkey-cow and cow-human images, but not human-monkey images. Species categories were also well-judged for inverted face images, but these did not give sharpened discrimination (categorical perception) at the category boundaries. While categorical species judgements are made reliably, only the distinction between primate faces and cow faces appears to be categorically perceived, and only in upright faces. One inference is that humans may judge monkey faces in terms of human characteristics, albeit distinctive ones.     

A Cross‐Cultural Comparison of Population‐Specific Face Shape Preferences (Homo sapiens)

A substantial number of studies indicate that human facial features such as symmetry and averageness are preferred in human mate choice. Little is known, however, about face preferences in intrapopulation or between‐cultural settings as only a few studies have addressed this topic. One hypothesis is that people prefer faces that resemble faces from their own population as a result of visual adaptation. Here, we present results of an internet‐based experiment in which male and female participants from two European nations, Poland and Scotland, were asked to choose between ‘Polish‐modified’ and ‘Scottish‐modified’ faces (i.e. face portraits modified along one of ten different dimensions to resemble faces of the respective populations) of both sexes while being unaware of the origins of the portraits. The results show that the preferences of Polish judges for Polish‐shaped faces were significantly stronger than the preferences of Scottish judges for the Scottish‐shaped faces. Moreover, both Polish and Scottish judges preferred Polish‐shaped faces to Scottish‐shaped faces. These results suggest that natural visual experience acquired in everyday interpersonal interactions may have limited influence on aesthetic preferences in these particular intrapopulation settings.     

Preferences for Infant Facial Features in Pet Dogs and Cats

A set of infant features (large forehead, large and low‐lying eyes, and bulging cheeks), were described in classical ethology as social releasers, simple stimuli that evoke a stereotyped response, in this case nurturing. We assessed the attractiveness of such features in the faces of dogs or cats (adults and young) or teddy bears or human infants, and also related these preferences to the degree of attachment to a pet. Overall, faces with the infant features were rated as more attractive than those without. Human infant faces were no more attractive than those of kittens or puppies. Pet faces were rated as more attractive by pet owners than non‐pet owners, regardless of whether the faces had infant features. A preference was also found for infant features in teddy bear faces. Women showed higher ratings than men for pets with infant features, but not for human infants or pets without infant features. Parents found human infants’ faces more attractive than did non‐parents, but there were no differences for other faces with infant features. Preferences were to some extent specific to the participant’s preferred pet species. Owners who were more strongly attached to their pets showed stronger preferences for photographs with infant features. The findings are discussed in terms of the concept of social releaser, and its part in the development of attachment to a pet species.     

Faces in places: humans and machines make similar face detection errors

The human visual system seems to be particularly efficient at detecting faces. This efficiency sometimes comes at the cost of wrongfully seeing faces in arbitrary patterns, including famous examples such as a rock configuration on Mars or a toast''s roast patterns. In machine vision, face detection has made considerable progress and has become a standard feature of many digital cameras. The arguably most wide-spread algorithm for such applications (“Viola-Jones” algorithm) achieves high detection rates at high computational efficiency. To what extent do the patterns that the algorithm mistakenly classifies as faces also fool humans? We selected three kinds of stimuli from real-life, first-person perspective movies based on the algorithm''s output: correct detections (“real faces”), false positives (“illusory faces”) and correctly rejected locations (“non faces”). Observers were shown pairs of these for 20 ms and had to direct their gaze to the location of the face. We found that illusory faces were mistaken for faces more frequently than non faces. In addition, rotation of the real face yielded more errors, while rotation of the illusory face yielded fewer errors. Using colored stimuli increases overall performance, but does not change the pattern of results. When replacing the eye movement by a manual response, however, the preference for illusory faces over non faces disappeared. Taken together, our data show that humans make similar face-detection errors as the Viola-Jones algorithm, when directing their gaze to briefly presented stimuli. In particular, the relative spatial arrangement of oriented filters seems of relevance. This suggests that efficient face detection in humans is likely to be pre-attentive and based on rather simple features as those encoded in the early visual system.     

Single-unit responses selective for whole faces in the human amygdala

The human amygdala is critical for social cognition from faces, as borne out by impairments in recognizing facial emotion following amygdala lesions [1] and differential activation of the amygdala by faces [2-5]. Single-unit recordings in the primate amygdala have documented responses selective for faces, their identity, or emotional expression [6, 7], yet how the amygdala represents face information remains unknown. Does it encode specific features of faces that are particularly critical for recognizing emotions (such as the eyes), or does it encode the whole face, a level of representation that might be the proximal substrate for subsequent social cognition? We investigated this question by recording from over 200 single neurons in the amygdalae of seven neurosurgical patients with implanted depth electrodes [8]. We found that approximately half of all neurons responded to faces or parts of faces. Approximately 20% of all neurons responded selectively only to the whole face. Although responding most to whole faces, these neurons paradoxically responded more when only a small part of the face was shown compared to when almost the entire face was shown. We suggest that the human amygdala plays a predominant role in representing global information about faces, possibly achieved through inhibition between individual facial features.     

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

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

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

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

Automatic recognition and measurement of butterfly eyespot patterns

A favorite wing pattern element in butterflies that has been the focus of intense study in evolutionary and developmental biology, as well as in behavioral ecology, is the eyespot. Because the pace of research on these bull's eye patterns is accelerating we sought to develop a tool to automatically detect and measure butterfly eyespot patterns in digital images of the wings. We used a machine learning algorithm with features based on circularity and symmetry to detect eyespots on the images. The algorithm is first trained with examples from a database of images with two different labels (eyespot and non-eyespot), and subsequently is able to provide classification for a new image. After an eyespot is detected the radius measurements of its color rings are performed by a 1D Hough Transform which corresponds to histogramming. We trained software to recognize eyespot patterns of the nymphalid butterfly Bicyclus anynana but eyespots of other butterfly species were also successfully detected by the software.     

Are human preferences for facial symmetry focused on signals of developmental instability?

Humans find symmetrical faces more attractive than are asymmetricalfaces. Evolutionary psychologists claim that our preferencefor symmetry can be explained in the context of mate choicebecause symmetry is an honest indicator of the genetic qualityof potential mates. These arguments assume that asymmetry inhuman faces is fluctuating asymmetry (FA), because this formof asymmetry can be revealing of developmental instability.However, no study has yet examined the characteristics of facialasymmetry. Here we provide the first detailed study of the patternsof asymmetry in human faces. We measured asymmetry in 35 facialtraits. Although some traits had distributions characteristicof FA, many had distributions that characterize directionalasymmetry (DA); on average, both men and women had right hemi-facedominance. For DA traits we used deviations from the mean asymmetryas a measure of developmental instability. Our measures of asymmetryaccounted for a moderate proportion of the variance in perceivedsymmetry. Importantly, only FAs and random deviations from DAcontributed to people's perception of symmetry. DA was not importantin symmetry judgments. Faces rated as symmetrical were alsorated as attractive. Random deviations from DA were weakly relatedto women's attractiveness judgments of men's faces. DAs didnot influence attractiveness judgments. Our data suggest thatpeople focus on aspects of facial asymmetry that may be revealingof developmental instability. Further studies that isolate FAfrom other forms of asymmetry are required to accurately assessthe influence of developmental instability on the quality ofindividuals and its potential role in mate preferences.     

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.