The neurobiology and evolutionary foundations of the perception of beauty
Ertugrul Esel, Gulustan Polat Esel
Article No: 12   Article Type :  Review
Beauty in human beings can be defined as physical attractiveness to the opposite sex. Although the perception of attractiveness varies between cultures and individuals to a certain extent, it is established that most of the criteria for attractiveness are common among many cultures. According to evolutionary psychologists, facial and body-related features that people find attractive reflect the adaptations determined by sexual selection, which is one of the driving forces of evolution. These adaptations evolved to explore the mate value and reproductive success of a potential partner. Being attractive provides many social advantages to a person, and it is known that people make some positive attributions about other characteristics of such a person as well. Among humans, features such as facial beauty, youth, body shape, behaviors, voice tone, and ornamentation are important factors in the evaluation of attractiveness of the opposite sex.
Keywords : Attractiveness, evolutionary psychology, mate selection, neurobiology
Dusunen Adam : The Journal of Psychiatry and Neurological Sciences : 2017;30:368-388
Full Text:


The question of what people find beautiful and why has yet to be given a satisfying answer. Throughout history, the topic of how beauty should be defined has been a matter of debate among thinkers, with the notion of ideal beauty in various fields of art changing and developing throughout the centuries. In this review article, instead of examining the general notion of beauty in the opposite sex, perceptions and elements of beauty in the opposite sex, in other words, sexual attractiveness, will be examined while looking into the evolutionary and neurobiological basis of this perception.

The notion of beauty in humankind can be defined as the attribute of being found attractive by the opposite sex; hence, beauty and sexual attractiveness are terms that can be used interchangeably (1). Sexual attractiveness is the product of the interplay between facial or bodily beauty of a person and the brain of the onlooker (2). However, the definition and the elements that make up sexual attractiveness are a matter of debate (3). Even though the perception of beauty may differ to a certain extent according to a person and culture, it is asserted that measures of beauty in different cultures, for the most part, share significant commonalities (1,3).

One important argument regarding the perception of beauty claims that human beings internalize and slowly gain a notion of beauty based on the attributes that are considered ideal by the cultures they grew up in from their childhood onwards and thus, the notion of beauty will change according to culture (3). That being said, as as result of studies that have been made since the 1970s, a “universalist” view seems to have gained ascendancy over a “relativist” perception that had argued that “beauty is in the eye of the beholder”. The universalist view asserts that beauty is similar in all cultures (3,4). Evolutionary scientists who hold this view maintain that the interest that people have in the facial or bodily features of individuals and the social messages that these features convey are neither dependent on a specific culture nor arbitrary; rather, they reflect adaptations shaped by sexual selection, one of the driving forces in evolution (3). These adaptations have evolved with a view to seeking for the quality desired in a potential romantic partner (5).

Although it is known that men place more emphasis on beauty in the opposite sex, it is also known that women pay more attention to beauty in other women than to abstract attributes such as personality traits (5-7). It is reported that the leading topic of conversation among women is “outer appearance,” and women are more curious about the perception of their attractiveness by other women than by men (8,9). These findings give rise to the idea that the main impulse for women wanting to appear beautiful, rather than attracting men, is to get the better of the competitors of their own sex.

It has been reported that when people are evaluating their own level of attractiveness, they labor under a personal bias, and often find themselves more attractive than they are in the evaluation by others (10,11). Studies indicate that people find themselves, on average, 10% more attractive than they actually are (10).

The Advantages of Being Beautiful in Social


Studies reveal that physical attractiveness provides individuals with many advantages in social life (12). In addition to the fact that people who are found beautiful attain an advantageous status when compared to their counterparts in many social arenas, it is known that the stereotype that “what is beautiful is good,” which has been around since Ancient Greece, is still maintained in many societies: That is to say, people who are found beautiful are also attributed with positive personal characteristics (12). For instance, being found beautiful and being beautiful are found to play an important role in the development of confidence and the shaping of the social life of every individual (13). It is said that people who are beautiful are more desired as friends and more successful in social relations, with these positive effects being more evident in the female sex (14). Other findings include that people who are beautiful are more preferred sexually and throughout life as partners and those who marry more attractive people are happier with their lives (15,16).

It is very telling for the advantages that beauty provides in life that beautiful people receive better treatment from others throughout their lives, are at an advantage in job promotions, have a greater chance of success at imaginary or actual job interviews, and beautiful people may even have an advantage regarding the punishment received in court (17-22). It is also believed that beautiful children receive more care from their parents and the photos of children who are not beautiful arouse negative feelings in adults, leading to physiological changes in these adults subsequently. These findings lead us to believe that the advantages of being beautiful begin at birth (22,24).

Another reality is that attractive people are also perceived positive in terms of non-esthetic personal attributes (Halo Effect). For example, people who have more attractive faces are perceived by others as being happier (25). Similarly, studies and meta-analyses assert that there is widely-held judgment that people who are beautiful are also more morally upright (26). When subjects were shown a photograph of a person and asked questions about the pictured individual, it was revealed that those who are attractive were assumed to be “more giving, helpful, intelligent, friendly” and these stereotypes exist even in children as young as 7-9 years (27).

The Evolutionary Foundation of the

Perception of Beauty

For researchers who look at human behavior from an evolutionary perspective, attractiveness in a human being needs to be evaluated as a concept being directly related to their mate value and their reproductive success (28,29). Thus, “attractiveness” research, while on the one hand examining which physical attributes are found attractive in human beings, on the other hand needs to investigate the evolutionary adaptive value of features that are found attractive.

Selecting a partner based on elements that are found “beautiful” constitutes one of the basic mechanism of “sexual selection,” which has been known since Charles Darwin. In sexual selection, the individual to be chosen (generally the male) can attract the attention of the choosing partner (generally the female) and thus gain a selective advantage in two ways (30):

1. Develop weapons such as horns, teeth, etc. that will increase the likelihood of being the winner in a contest with their competitors of their own

2. Equip themselves with various extensions (peacock tail, lion’s mane), beautiful colors, or attractive songs, as in the case of birds, to display high genetic quality.

Beauty and the perception of beauty must have developed as a product of this second mechanism in sexual selection, considering that the members of a species give three messages to the opposite sex with the attributes that make up their attractiveness (31):

1. “I am healthy,” in other words “I have good genes.”

2. “I have an abundance of estrogen (or testosterone),” that is to say, “I have great reproductive potential.”

3. “I would be a good mother or father.”

Thus, individuals from the opposite sex who correctly evaluate this type of information, demonstrating the carrier’s genetic quality, reproductive power, partnership capacity, and social value, will have a great evolutionary advantage and leave behind more genes (3). Therfore, signs indicating partnerşhip quality will be perceived positively (thus attractive) by the other sex (13). High-quality individuals will also increase the success of their partners in reproduction and survival, which will give them a higher preference. As a result of this mechanism, in the evolutionary process members of both sexes have been carefully attuned to signs that demonstrate “high spousal quality” (13). In sum, the current human standards of beauty are a product of our past and people select their partners so as to maximize their reproductive powers.

Attributes Found “Beautiful” in Humans

Which features are more important than others for humans to find someone attractive or not is a research topic that has gained momentum in recent years. It appears that in finding someone from the opposite sex attractive, some of the physical and behavioral attributes of the person being viewed (facial beauty, appearance of youth, body shape and weight, tone of voice, adornments and attire, behavior, etc.) as well as certain features of the viewer (such as their culture, values, personal attributes, in the case of women: the time of their menstrual cycle) are of importance. Due to lack of space, in this article only the attributes relevant to the perception of beauty of the person being viewed will be examined.

Facial Beauty

In human beings, facial perception occurs much faster than the perception of other objects (32). The evaluation of facial characteristics gives us information not only about a person’s sex, age, race, etc., but also finer details of relevance in social life such as their emotions, reliability, attractiveness, and intentions (33). In humans and monkeys, the facial processing area is essentially in the lateral fusiform gyrus (the fusiform face area), which is part of the occipitotemporal cortex (34,35). This fusiform facial area has an effective connection to the orbitofrontal cortex (OFC) that also plays an important role in facial processing (36).

The perception and processing of whether or not a face is beautiful in the human mind is incredibly efficient. It is reported that it takes 100 milliseconds to understand whether a face is attractive or not (37). This shows that the perception of the attractiveness of the face is based on strong biological foundations. The beauty detectors which fulfill this cognitive function are believed to exist in every human being from the moment of birth. Studies have found that even three-month-old babies look more at faces that adults find attractive (38,39).

Whether or not a face is considered attractive is very important in human spouse selection behavior, which is determined by neural pathways that have been shaped as a result of thousands of years of natural selection in the human brain: A beautiful face provides the onlooker valuable information about the other person’s being healthy, having a high level of the right gonadal steroids, and being resistant to parasites, as well as, particularly in women, being fertile (40,41).

In addition, it can be said that facial beauty is a sign indicating that the whole body is healthier/more attractive. When male subjects were shown cropped photographs of women which showed either only their faces or their bathing suit-clad bodies and asked “which one is more attractive?”, they gave scores similar to those they would give when the complete women – face and body – were visible (42). In other words, it appears that the messages carried by each of the different parts of the body that create attractiveness are consistent with one another to a significant degree, indicating each by itself general attractiveness and health.

A study which supporting the perception that the facial attractiveness of a person is related to their being healthy found that women assessed the facial photographs of men whose handshakes were considered “strong” according to a handshake measurement as being more handsome and attractive (43). Based on this, it has been asserted that for a man, having an attractive face carries information about him being particularly strong and healthy, in addition to giving information about his testosterone level. Also, the finding that individuals who have an attractive face live longer and have more children supports the positive corrrelation between facial attractiveness, health, and reproductivity (44,45). Furthermore, the findings that people with attractive faces were found to have lower levels of cortisol under stressful circumstances, while women with beautiful faces were found to have higher estrogen levels (therefore have a higher potential of becoming pregnant), support this belief (46, 47).

It is Reported that the Following Factors are

Important in the Evaluation of Facial Beauty:


Being average

Masculine or feminine features

Smooth skin

Youthful appearance


Facial expression of feelings

Facial Beauty: Symmetry

What is symmetrical is considered more beautiful in many animal species and in human beings. The brains of animals are programmed to perceive symmetry quickly (48). In the animal world, symmetry is found to be related to gene quality; asymmetry may reflect factors such as inbreeding (which causes poor quality genes), mutations, homozygosity, or environmental factors such as an unhealthy development in the first few months or years of life, inadequate nutrition, or parasitic infection, which may give very valuable information to the one who is evaluating the individual as a potential mate (3,12). On this basis, from an evolutionary perspective it can be assumed that facial symmetry in human beings indicates similar attributes and humans unwittingly prefer symmetrical faces for this reason.

As expected, studies show that humans, too, find symmetrical faces to be more attractive (1,49,50). In many studies where the levels of symmetry of different faces were altered on a computer, facial symmetry was found to increase attractiveness (51).

In studies conducted on humans, it is demonstrated that symmetry of the face is somehow associated with good health. For example, the perception in human beings that those with symmetrical faces are healthier is supported by the examination of actual medical sources where a corrrelation is found between facial symmetry in men and women and their being healthy. These findings demonstrate that there may be a real relationship between facial symmetry and being healthy, or that at the very least, this relationship is perceived as such (52-54). Similarly, findings that facial symmetry in humans and level of intelligence are directly proportional as well as facial attractiveness, even when controlled for symmetry, is directly proportional to the person’s socio-economic status may make us think that symmetry is an indicator that a person is generally healthy and “has good genes” (55-56).

Facial Beauty: Being Average

It is established that there is a relationship between a human face (and generally speaking forms and objects) being close to average and being perceived as attractive. Up to a certain point, men and women tend to find faces that are close to average more attractive; however, particularly for certain personality traits and periods, in both sexes the accentuation of masculine of feminine characteristics is found more attractive than being average (3).

The fact that human beings find average faces more attractive can be explained as follows: The human mind forms “an average for the face” (prototype) based on the faces it has been exposed to from the moment of birth. In fact, our visual system creates an internal prototype for every stimulus, and this prototype is the average of all of the stimuli that one has been exposed to so far. Thus, when our mind is faced with a new stimulus, it compares it with the prototype and builds a feeling of familiarity with the prototype. Hence, the reason for average faces seeming more attractive to us is that we find them more familiar (57).

In support of this view, in studies where facial photographs are shown — some of them twice — individuals on photographs that had been seen before were perceived as more attractive by both sexes (58). However, this effect (finding what is familiar more attractive) was more evident in women and less apparent in men. Similarly, it was found that women find faces resembling that of their partner more attractive, while this effect does not exist in men (58). In summary, the phenomenon of finding what is familiar more attractive is more evident in women, while men are more prone to new relationships and thus also find what is new attractive. This finding can be interpreted as information that confirms men’s tendencies towards multiple and short-term relations (59,60).

The phenomenon of average faces being found more attractive has been demonstrated in studies conducted in different cultures (11,54,61). In addition it has been demonstrated that children of the ages of five and nine years, just like adults, find average faces more attractive than others (62).

Facial Beauty: Masculine-Feminine Features:

In non-human species, exaggerated gender-specific features (ex: the large antlers of a deer, the tail of a peacock) have the function of attracting the opposite sex and intimidating rivals of the same sex (63,64). Based on this knowledge, the effect of gender-specific morphological attributes (the dimorphism of facial and bodily features) on the perception of beauty is increasingly becoming a research topic in human beings, too.

In the male face, features that are perceived as masculine are a broad chin, thick eyebrows, a nose that is broader than a woman’s, deep eyes that are close together, and ears that are close to the head. In women, feminine features are listed as a narrow chin, full lips, slender eyebrows, large eyes, small nose and a short distance between mouth and tip of the chin.

In many studies, it is shown that men find feminine features more attractive in female faces and women find masculine features more attractive in male faces; however, there is a limit to this perception (65,66). Contrarily, there are studies which do not find a relationship between facial femininity or masculinity and attractiveness (67).

The leading biological factors related to a human face being feminine or masculine can be listed as: healthy genes, the effect of hormones in the intrauterine period, the effect of surging hormones during puberty and the level of hormones in adult life (3).

The Effect of Genes

From an evolutionary perspective, it can be said that extreme secondary sexual characteristics (in other words, more feminine in women, more masculine in men) point to the person having good genes, which makes them more desired (3). Since in animals, a gross exaggeration of gender-specific attributes leads to the consumption of resources, the individual’s capacity to cope with this burden indicates its being healthy. Even though there are some findings in humans suggesting that features differentiating men and women demonstrate gene quality, this correlation is not very clear and certain (3).

Support for the positive relationship between masculine attributes in men and gene quality comes from studies which assert that there is a correlation between masculine features and the strength of the immune system (18). The “immunocompetence hypothesis” makes the following claim: as is known, testosterone is a steroid that suppresses immunity (68). Thus, only men with a very strong immune system can remain healthy despite their high levels of testosterone. Men who have masculine attributes in their faces have been able to remain healthy despite their high levels of testosterone, which means that they must be men with very strong immune systems (69). Thus it is asserted that exaggerated gender-specific features are indicative of a hereditary immunity and thus “good genes.” It needs to be pointed out that this subject is still at the level of a hypothesis and there are very few findings to confirm it (69).

The Effect of Intrauterine Hormones

It is maintained that the level of pre-natal testosterone the fetus has been subjected to is important in the development of masculine or feminine features. In other words, prenatal testosterone has an effect not only on the body and brain developing as a male, but also may play an organizational role in the masculine or feminine development of the face. A finding supporting this view comes from research that examines the index finger to ring finger ratio in men and women (2D:4D). According to most of these studies, the 2D:4D level in men is lower than in women. The digit ratio drops as the prenatal testosterone level increases (70). There are studies that indicate that as the 2D:4D level decreases (in other words, as the exposure to pre-natal testosterone increases),men’s attractiveness, physical/athletic capacity, and the number of spouses and children increases (71,72). However, we must add that a newly conducted meta-analysis does not confirm this argument and that there was no relationship found between 2D:4D ratios and male attractiveness.

The Effect of Rising Hormones in Puberty

Many of the gender-specific different facial features of men and women form during puberty based on the masculinization or feminization of secondary sexual characteristics. Hormones such as testosterone and estrogen which rise during puberty lead to this differentiation. The masculinization of the face in men as well as the increase in testosterone are changes that unfold during puberty (74). The features of the female face that men find attractive indicate rising levels of estrogen and decreased androgen women are exposed to during puberty. The increasing levels of estrogen inhibiting the effect of testosterone in women during puberty lead to the formation of feminine features such as wide eyes and full lips (2,75).

The Effect of Circulating Hormones

There are studies which indicate a positive correlation in adulthood between circulating testosterone in men and facial masculinity, and high estrogen levels and female facial attributes in women (47,76,77). For example, there are studies to indicate that the attractiveness of the female face is positively correlated to the level of estrogen in the blood (47). Therefore, faces of ovulating women are found to be more feminine and attractive.

As the sexually dimorphic features in the human face are partly hormone-controlled signs, they may be indicators of their reproductive quality or their ability to compete with same-sex competitors (78). Even though it has been demonstrated many times in animals that masculine traits increase procreative power and success, this remains a matter of debate where human beings are concerned (79-81). Some studies have found a positive correlation between masculine features of the face and attractiveness and the quality of sperm in humans, but others have not (82-84). It has been reported that there is a positive correlation between the attractiveness of the face and its masculinity and physical strength as measured by the strength of the handshake (43). The finding that men who have a high level of testosterone have a greater number of sexual partners throughout their lives and more children indirectly supports this idea (85,86).

Male Preference of Feminine Faces:

All studies carried out using either real women’s faces or modifying female facial features on the computer demonstrated that a feminine female face is found more attractive by men (87-89).

It is said that the feminine attributes of the female face are also an indication of youth; thus the preference for femininity in women by men might partially be due to the preference for young women (88). And in fact, many of the features which men find attractive and feminine are features that resemble those of babies (small chin, small nose, large eyes, bright and smooth skin, etc.). Make-up applied by women, which is a universal phenomenon, is used to emphasize these attributes (making the eyes appear larger and deeper, smoothing the skin, concealing lesions, and making the lips to appear fuller) (90).

Another finding is that men are most strongly attracted by feminine faces when in their 30s, while with increasing age, their level of finding feminine faces attractive decreases (91). Consequently, it is thought that the preference for feminine faces in men may be related to testosterone levels and the decrease in the preference for femininity may be a reflection of decreasing levels of testosterone. This may have an evolutionary adaptive advantage as well: the decrease in the preference for femininity as they age may protect men from the risky behavior of having to fight with younger and stronger men for highly fertile women (91).

Female Preference of Masculine Faces:

The subject of masculine faces appearing attractive to women is highly debated (3). Just as there are studies that indicate an increasing female attraction to male faces with increased masculine features, there are also studies suggesting that exaggerated masculine features in male faces as indicators of dominance are not found attractive (50,65,92). While the female attraction towards certain male features such as a wide chin is a more consistent finding (93, 94), this is not the case for all masculine features (13). An increase in masculine facial and vocal features in men, while consistently being perceived as an indicator of dominance, is not always evaluated as a phenomenon to increase attractiveness (96,97).

When we assess the studies in their entirety, it can be said that masculine attributes demonstrating dominance in men increase their attractiveness up to a certain point, beyond which it decreases. Another finding on this matter indicates that women close to ovulation or looking for a short-term relationship find masculine faces attractive, while during a large part of their menstrual cycles and particularly when looking for a long-term relationship, they prefer men with faces that show low dominance (98,99). Thus the level of masculine face preference in women was found to be associated with their blood estrogen levels (99). It is also reported that married women (particularly during ovulation) find extremely masculine-faced men more attractive than single women and that in both sexes the level of preference for exaggerated masculine or feminine features in the opposite sex is directly correlated to the intensity of their sexual desire (65,89,100).

The preference of women towards masculine faces is also found to be associated with oxytocin levels. It is reported that when women are administered oxytocin through their noses, their preferences shift towards masculine faces (101). The reason for this is stated to be that oxytocin, as a hormone that “supports socializing and spousal relationships” covers (makes invisible) the negative personality traits that masculine faces are normally associated with.

The decrease in women’s preference for masculine facial features, when they are in search of long-term partners is a consistent finding. This is said to be due to the fact that extreme masculinity is a disadvantage in long-term relationships (in other words, parenthood). Extremely masculine characteristics are associated with certain negative attributes such as aggressiveness, controlling behavior and compulsiveness. It has been shown that women perceive men with increased masculine features in their faces as being more dominant, more aloof, less emotional, less honest, less cooperative and being worse parents (88). There are findings that indeed demonstrate that with increasing masculinity, men’s parenting quality decreases. For example, men who have more masculine attributes are noted to be less loyal to their families and spouses, taking less time to care for their children (18). Similarly, observations such as men with high levels of testosterone being less likely to be married, with a higher rate of divorce, and if married spending less time with their partners and children are supporting this suggestion (102-104).

In conclusion, it can be said that from a female perspective, there are both advantages (good genes) and disadvantages (bad parenthood) is a man is “very male.” Thus, some authors suggest that women are solving this dilemma (masculine and dominant or feminine and soft, good spouse and parent) by choosing a spouse whose face combines both attributes, reflecting “multiple instincts”(98). In other words, the facial selection in women is based on a balance between good genes and a desire for a cooperative spouse. Therefore, masculine male faces will be found attractive under certain circumstances and when the women are in certain periods. The evolutionary explanation of this balancing strategy can be done in the following way: From the perspective of evolution, it is appropriate for the woman to select a mate who has the potential to be a “good father” in the long run; however, if during her fertile period she is to encounter a male with better genes, i.e. someone more masculine than her spouse, she will engage in a sexual escapade (105).

Facial Beauty: The Skin

Just as skin color distribution and its homogeneity affect the attractiveness perception of the face, they also give us information about the person’s age and healthiness (106-108). A smooth face and homogeneous skin color increase female beauty in particular. In women, a skin that is free of lesions, acne, tumors, and facial hair, in short, good skin, is a feature that is universally desired by men (106).

One of the most important reasons for smooth skin increasing attractiveness is that it gives the individual a “youthful” appearance (109). Decreasing homogeneity in facial skin coloration in both sexes leads to the perception of being older, less healthy, and less attractive (10). An important reason for the use of cosmetics in women is to appear to have spotless, flawless skin and an overall young and healthy appearance.

Smooth skin in a woman, in addition to being an indicator of youth, may also carry the message that a female is healthy and productive. For example, illnesses that destroy the homogeneity of the skin such as acne and hirsutism may point to a hormonal abnormality (109,111). In women, a smooth skin tone may give information about the power of her fertility, because smooth skin is found to be associated with low levels of androgen and high levels of estrogen (112). Another message which smooth appearance of the skin may relay is that the person has a good immune system, because it is known that the skin of persons with a weak immune systems has a high chance of being attacked by micro- and macroparasites (112).

Another factor that is associated with an increased attractiveness and healthy appearance is a rosy color of the cheeks. In both sexes, rosy cheeks point to adequate oxygenation and thus good health (108,113) In addition a high level of gonadal hormones in women is known to increase the vascularity of the skin and thus the redness of the cheeks (114). Some associate the beautifying effect that small doses of alcohol has on a person to this as well. Studies show that the face of a person who has consumed a small dosage of alcohol (such as 250 cc of wine) is evaluated as being more attractive by members of the opposite sex (115).

Contrasting colors on facial skin are also known to increase the attractiveness of women. Contrasting colors on the skin normally occur more in women than men and thus are found attractive by men. Increased contrast in the face (an increased color difference between the skin and organs such as the lips and eyes) work to increase the attractiveness of a woman while they decrease the attractiveness of a man (116); The skin color of women is on average lighter than that of men and this is what is preferred (109). It has been demonstrated in many cultures that men prefer women who have skin colors that are lighter than the average (117,118). The two most important substances that generate the color of the skin are melanin and hemoglobin, and both are found in smaller quantities in women. In sum, the fact that the woman has more contrast in her face is found attractive by men because it is an attribute that demonstrates sexual dimorphism (being a feminine attribute) (109).

Another finding related to contrast in women increasing attractiveness is that red lips increase the attractiveness of a woman in almost every culture (119). As most likely reasons for this has been asserted that red lips are considered an imitation of the vasodilation that takes place during sexual arousal, pointing to blood with high levels of oxygen and thus good health, in addition to high estrogen levels (119).

Facial Beauty: Youthful Appearance

Even though youthful-appearing faces are found more attractive compared to older-looking faces in both sexes, it is known that a young appearance of women is more important for men (107,120,121). In many cultures, men prefer women who are younger than them while women prefer men who are slightly older as partners (120). From an evolutionary perspective, this is expected, because youth means strength and health, and furthermore, a person who is young has more time to have children and be a parent. Because the period of fertility in women is shorter, the advantages of being young are more evident. In other words, due to the effects of aging on fertility and health the decrease in mate value is more evident in women (122).

In men, given that aging goes along with an increase in status and material resources, the effects of decreasing attractiveness due to age are not as evident as in women (121). In other words, the decreased attractiveness of the male face after middle age is partially redeemed by status and resource-related increases, because the wealth and status value of men is important for women with regard to raising children and giving them a good upbringing. And the attractiveness evaluation of post-menopausal women is lower than their male counterparts’ who are in the same age range (121).

Facial Beauty: Eyes

The eyes are the body part that reveals most about a person’s intention. Generally speaking, people evaluate faces that look directly at them as being “well intentioned” or “more attractive” (123). Deep and big eyes stand out as a factor that increases attractiveness, particularly in women. It is reported that among the sexually dimorphic attributes of the face, the most important area is that surrounding the eyes (124). One study found that even children younger than five years of age look longer at faces that have big eyes (125).

Furthermore, it has been demonstrated that men find women on photos where their pupils have been made larger than normal (without the men noticing) more attractive and feminine (126). As a reason for this, it has been proposed that dilated pupils are a sign of sexual arousal in women, which men may realize through the unconscious-automatic portion of their minds.

Additionally, a pronounced medial canthus of the eyes makes them appear to be tilted inward and slanted downward, increasing attractiveness particularly in women (124). This is because inward- and downward-tilted eyes are a prominent feature in babies, also demonstrating sexual dimorphism (more pronounced in women). Thus, women with pronounced medial canthus are considered more attractive by men because they create the illusion that their eyes are slanted inwardly-downwardly (124).

Facial Beauty: Feeling in the Face

One of the most important elements that affect the attractiveness of the face is the “expression of feeling” (25). Generally, the abundant use of facial movements (facial mimics) when expressing feelings is found more attractive by men (127). The finding that woman who are not in a romantic relationship use more facial expressions when engaging with men compared to women who are may be revealing that they, too are aware of the attractiveness of using facial mimics (128). An increase in facial expression during ovulation also confirms this idea (127).

The smile, which indicates happiness, holds an important place in the relationship between attractiveness and the expression of feelings. When individuals were shown pictures of humans, a smiling face was found more attractive than a neutral face in both sexes (25,129,130). Furthermore, there is a positive correlation between attractiveness score and the width and intensity of the smile (94). Pictures of smiling and attractive faces are noted to cause a higher activity increase in the brain reward system and the medial prefrontal cortex (mPFC) (130). It is held that the attractiveness-increasing phenomenon of the smile is more valid in women than men (131).

Brain Activity of a Person Looking at a

Beautiful Face:

In brain activity studies conducted with fMRI, it was found that when humans are shown the picture of an attractive face from the opposite sex, the brain – in comparison to looking at faces that are not attractive – shows an increase in activity in the regions of OFC, mPFC, nucleus accumbens, and anterior and posterior cingulate regions (33,130,132-135). These areas are generally associated with the brain’s reward system and thus it is believed that looking at the attractive face of the opposite sex has a rewarding effect on a person. It is asserted that the mOFC in particular is the part of the human brain that determines and appreciates beauty in all modalities (136). As the attractiveness of the face being viewed increases, OFC activity increases in a linear fashion (132). When male participants were shown beautiful female and male faces, those of both sexes were correctly evaluated as attractive; however, probands only showed increased activity in the right orbifrontal and bilateral nucleus accumbens when looking at pictures of women (133,137). In summary, it can be asserted that the OFC in particular is the area automatically evaluating whether a face is beautiful or not, and thus it is likely the essential area for choosing a potential spouse (133).

Bodily Features that are Found Attractive

When people are evaluating whether the opposite sex is attractive or not, they reach a decision after processing information coming from the most different sources. Thus, in addition to the face, the shape of the body, physical attributes and movements, too, are important in determining attractiveness and thus the quality of a potential mate (138).

As variables that are important in determining whether a body is beautiful or attractive, its symmetry, height, weight, in women the waist-to-hip ratio, the size of breasts, and the hips, and in men broad shoulders can be considered relevant. It can be argued that all of these physical attributes may increase a person’s attractiveness because they are related to being healthy or more reproductive.

Bodily Beauty: Symmetry

Symmetry is an important element in bodily shape and beauty, just as it is for the face. Humans who have a symmetrical skeletal structure are found more attractive, while those who have a body structure that is symmetrical are reported to have more sexual partners (139). The finding that there is a positive correlation between bodily symmetry and sperm count in ejaculation and the speed of sperm (140) demonstrates that there may be a positive relationship between symmetry and reproductive power.

Bodily Beauty: Slimness

It is shown that being overweight decreases attractiveness for both sexes. The two most commonly used measures in studies on this topic are Body Mass Index (BMI) and Waist-to-Hip Ratio (WHR). It appears as though lower figures for both measures are preferred by males and females alike (141,142).

In the current century, particularly for women the relation between slimness and beauty is increasingly exaggerated; thus the assumption that a slimmer person is found more attractive is gaining force. The media have been pointed out as the most effective institution creating the stereotypical notion that “slim is good” regarding the ideal weight (143,144). We see that today, women who are presented by the media as being ideal tend to be below normal weight (145). A study examining the bodies of 559 playboy beauties from the years 1950-2000 found that over time, ever taller and thinner models were featured on the pages of the magazine (146). Consequently, women who follow fashion magazines have more deeply internalized the idea that “being slim is ideal,” and therefore, they more often suffer from eating disorders (147,148). Some studies have found that women subjects who are exposed to the photographs of slim-bodied women for a short period of time change their own body image and ideal body notions, increasing their dissatisfaction with their own physiques (149,150). It is established that the subject of thinness is deemed more important by women who view themselves as more attractive and beautiful, and thus young girls who have this type of a hang-up regarding beauty are at an increased risk of suffering from anorexia nervosa (151).

Studies indicate that men find women who are found beautiful by other women to be too thin, while in reality finding women who carry slightly more weight to be more beautiful (152). In women, BMI and general degree of health are to a certain extent proportionally related. It is established that a rise in BMI increases fertility and that women with greater weight have more children (153). Extreme thinness or obesity decrease attractiveness and may directly cause amenorrhea, thus preventing genes from being passed on to the next generation (154). This subject appears to be significantly influenced by cultural perspectives. For example, we know that rural men prefer women that are heavier compared to those chosen by men living in industrialized cities (155).

One of the variables related to slimness is the BMI. Studies indicate that women with a BMI of 20 or thereabout are found more attractive by men (141). Another of the variables in this matter is the WHR. It is reported that there is an inverse relationship between WHR and attractiveness in women (6,156). Women generally have a smaller WHR compared to men, and this difference is preferred by men. A preference for the hour-glass figure is seen in all cultures (157,158). Generally speaking, women with a WHR of 0.7 are found ideal by men (159). When the silhouettes of women are placed before men in a very speedy fashion, men are still able to find women with the ideal waist-to-hip ratio (159). Furthermore, it has been established in studies by fMRI that when men are looking at images of ideal women with a 0.7 WHR, the parts of their brain that deal with perceiving beauty and reward processing, such as the OFC, are more active (160).

The WHR is an indicator of fat storage and is controlled by sex hormones (6). While estrogen triggers fat storage in the hips and thighs, it represses fat storage in the internal organs. Therefore, the female body shape found attractive by men (an hourglass figure: big breasts, low waist/hip ratio) is actually displaying attributes which are the effects of estrogen (161). A low WHR is known to correlate with a high level of reproductive hormones (estrogen), being healthy, and high conception and reproduction levels (162). Furthermore, findings that women who have low levels of WHR experience an earlier menarche, with more regular menstrual and ovular cycles during adulthood, confirm the idea that this attribute is directly related to the sexual competence and health of a women (163,164). In line with these findings, it can be ascertained that there is an adaptive quality to the male preference of a low WHR.

Bodily Beauty: Height:

Tallness affects the qualities attributed to a person by others in both sexes. Particularly in man, having a tall stature is reported to increase attractivity (165).Tall men are found to have an advantage in being chosen as a flirt partner (166,167). It appears though being tall affects perceptions other than attractiveness as well. For example, men who are taller are attributed more positive personality traits by women, while tall women are perceived by men as being smarter, wealthier, in addition to having more initiative and ambitions (168,169). These perceptions may partly express a reality: Some studies have found a positive correlation between height and professional success and annual income (170). Furthermore, taller men are known to have more children (171).

Behaviors and Attitudes:

While masculine behavior in men and feminine behavior in women increase their attractiveness, the reverse decreases their allure to the other sex. It is noted that women are particularly attracted to men who have a high level of self-confidence, but at the same time are compassionate and protective. Furthermore, it is suggested that the way of walking, too, is important for attractiveness: a coordinated and symmetrical gait increases attractiveness. It has been found that women move in a more coordinated and symmetrical fashion when they are in their mid-luteal phase (the period of optimal estrogen levels) compared to their menstrual period (172).

A number of studies indicate that courting or flirting women more frequently engage in behaviors such as head-nodding, leaning forward and towards their potential partner, touching the person before them and themselves, playing with their hair and tossing their hair back (173). Narcissistic behaviors, too, are reported to increase attractiveness, especially as a short-term romantic or sexual partner (174).

In recent years, an increasing number of studies has assessed the attractiveness of individuals who dance. Men who were found attractive by women while dancing reportedly have a greater level of physical power measured by the strength of their handshake (175). In other words, it can be said that dance movements in particular give information about the physical strength of men and women admire the dancing of strong men.

In one study, facial photographs were shared with participants along with a number of sentences evoking moral judgments (this person is a human rights defender, works as a volunteer nurse in Africa, or a drug-dealer, this person was proven to have stolen their thesis from someone else, etc.), in order to see if these individuals’ moral attitudes influence their “attractiveness” scores (176). This study determined that while in both sexes the perception of attractiveness was influenced by moral evaluations, this effect was more pronounced in women and less so in men. In the end, men were found to place less significance on the moral status of their potential partners in the assessment of their attractiveness. This finding matches the classical evolutionary view that men are less selective in choosing a partner as they have to invest less in the process of gene transmission (176).

Tone of Voice:

Voice frequency in men is inversely proportional to their level of testosterone during puberty (177). A deep voice of men is found more attractive by women (178-180). In addition, it is known that a deep voice is considered an indication of dominance in both men and women (181).

The voice of a man may carry clues about the size of his body and his hormonal state. In a study where the voice of male participants was recorded and played to female participants, male voices with a lower frequency were rated as more masculine; these repective men were actually taller than the others and their spit testosterone was higher than their counterparts’ (182). Thus, it can be asserted that women can recognize the signs of male dominance through hearing their voice alone.

Furthermore, some studies find that men who have a more attractive voice have a higher rate of reproductive success (183). There is a negative correlation between the frequency of voice (its high pitch) and the body size and shoulder width (179). It has been found that women find men with deep voices more attractive when they are in their fertile period (late follicular phase); it has been found that the level of finding a deep voice attractive and being affected by this voice is positively related to estrogen in the saliva or the level of estrogen metabolites in urine (180,184). In other words, there is an increase in the preference of masculine attributes (face, behavior, etc.) in men as the level of estrogen increases in women during their fertile period (99), and it appears as though the preference for a deep voice, too, increases (180).

Adornments and Attire:

The main effect of adornment in animals is to show the gene quality and good health of the owner, consequently conveying to the other sex the message that they are “a superior partner” (185). In animals, the sex that adorns itself more and thus becomes increasingly beautiful in order to be chosen by the female is the male. However, females are seen to be adorned in many animal species as well, a fact that is more related to genes from male ancestors who increasingly adorned themselves in the evolutionary process (shared genetic architecture) (185,186). Because intra-sex competition is less forceful in monogamous animals, both the males and females resemble one another, while male ornamentation is more emphasized in polygamous animals. It is reported that ornamentation in males is an honest indication of quality (the more the ornamentation the better the gene) (185).

In many animal species, males adorn themselves under selection pressure; at the same time, males also have energy to spend on ornamentation as they do not invest in offspring. Because the female is focused on the offspring, she cannot set aside energy for adorning herself; furthermore, after securing pregnancy, she no longer has the need to adorn herself (187). In one species of animals where females are more adorned, it has been found that they invest less in their children (188). In the end, there appears to be an inverse relationship between investing in offspring and ornamentation in animals.

Contrary to most animal species, in human beings it is the female who adorns herself. The application of makeup by women is an almost universal phenomenon that is known to have existed throughout history (189). Cosmetic products produced for use by women, as well as cosmetic surgery procedures, aim to exaggerate features which are normally found to be beautiful (1). It has been proven over and over again that make-up increases the attractiveness of women (190,191). Moreover, when participants were shown photographs of women with and without make-up, it was the faces with cosmetics that triggered a higher level of activity in the left OFC and the right hippocampus (with made-up faces being perceived as more attractive and found more rewarding) (191).

Some studies in humans have proven that a red background or a red outfit increase the allure of a woman (192). In restorants, men were found to pay female waitresses wearing a read outfit a more generous tip (193). It has also been found that men will more often perceive a woman donning red as being more “sexually eager” (194). Interestingly, red creates the same perception of women with other women as well: Other women will evaluate a female in red as having “high sexual receptivity and low sexual fidelity” (195). The same study found that women will exhibit the tendency to protect their partners from “women wearing red” in particular.

It is reported that women’s preference of formal attire in men, such as a suit, over “casual” (sporty) wear may have to do due with an estimation that a man who wears a suit is “of a higher status” (196).


People are perceptive to what is beautiful, and beauty is an important factor in mate selection. While being found beautiful has many psychological advantages such as boosting an individual’s self-confidence and self-satisfaction, it also provides many social advantages. Even though the perception of beauty differs depending on culture and person, many studies prove that there are common measures of beauty found in different cultures. Studies even assert that these common beauty measures have existed in the human mind since birth: In other words, people may have a tendency to differentiate what is beautiful and what is not based on a biological skill that has been, to a large extent, embedded in their genes. It is observed that traits that are found automatically attractive in the human mind by the opposite sex are of the those relating to fertility, health, youth; in short: gene quality and being a good partner or parent in particular, as human beings subconsciously choose partners in a way that maximizes their chances of reproduction/gene transfer. Thus, when thinking of the assessment of beauty and the evolutionary advantages of choosing the right partner, it becomes better understood why the perception of beauty has developed as a strong and automatic action of the human mind and why this subject is given so much importance in virtually every society. Researching the evolutionary and biological foundations of beauty and sexual attractiveness serves a highly insightful and informative function in understanding how the human mind works, in addition to comprehending the strong skills that have developed through the evolutionary process.


1. Campbell A. Female competition: causes, constrains, contents, and contexts. J Sex Res 2004; 41:16-26. [CrossRef]

2. Johnston VS. Mate choice decisions: the role of facial beauty. Trends Cogn Sci 2006; 10:9-13. [CrossRef]

3. Little AC, Caldwell CA, Jones BC, DeBruine LM. Effects of partner beauty on opposite-sex attractiveness judgments. Arch Sex Behav 2011; 40:1119-1127. [CrossRef]

4. Laeng B, Vermeer O, Sulutvedt U. Is beauty in the face of the beholder? PLoS One 2013; 8:e68395. [CrossRef]

5. Buss DM. Conflict between the sexes: strategic interference and the evocation of anger and upset. J Pers Soc Psychol 1989; 56:735-747. [CrossRef]

6. Fisher ML, Voracek M. The shape of beauty: determinants of female physical attractiveness. J Cosmet Dermatol 2006; 5:190-194. [CrossRef]

7. Fisher ML. Female intrasexual competition decreases female facial attractiveness. Proc Biol Sci 2004; 271(Suppl.5):283-285. [CrossRef]

8. Martin R. Girls don’t talk about garages: perceptions of conversation in same-sex and cross-sex friendships. Pers Relatsh 1997; 4:115-130. [CrossRef]

9. Graziano WG, Jensen-Campbell LA, Shebilske LJ, Lundgren SR. Social influence, sex differences and judgements of beauty: putting the interpersonal back into interpersonal attraction. J Pers Soc Psychol 1993; 65:522-531. [CrossRef]

10. Epley N, Whitchurch E. Mirror, mirror on the wall: enhancement in self-recognition. Pers Soc Psychol Bull 2008; 34:1159-1170. [CrossRef]

11. Mu-oz-Reyes JA, Iglesias-Julios M, Pita M, Turiegano E. Facial features: what women perceive as attractive and what men consider attractive. PLoS One. 2015; 10:e0132979. [CrossRef]

12. Buggio L, Vercellini P, Somigliana E, Viganò P, Frattaruolo MP, Fedele L. “You are so beautiful”: behind women’s attractiveness towards the biology of reproduction: a narrative review. Gynecol Endocrinol 2012; 28:753-757. [CrossRef]

13. Little AC. Facial attractiveness. Wiley Interdiscip Rev Cogn Sci 2014; 5:621-634. [CrossRef]

14. Senna A, Abbenante D, Tremolizzo L, Campus G, Strohmenger L. The relationship between facial skeletal class and expert-rated interpersonal skill: an epidemiological survey on young Italian adults. BMC Psychiatry 2006; 6:41. [CrossRef]

15. Riggio R, Woll S. The role of non-verbal and physical attractiveness in the selection of dating partners. J Soc Pers Relat 1984; 1:347-357. [CrossRef]

16. Berscheid E, Dion KK, Walster E, Walster GW. Physical attractiveness and dating choice: A test of the matching hypothesis. J Exp Soc Psychol 1971; 7:173-189. [CrossRef]

17. Eagly AH, Ashmore RD, Makhijani MG, Longo LC. What is beutiful is good, but …: a meta-analytic review of research on the physical attractiveness streotype. Psychol Bull 1991; 110:109-128. [CrossRef]

18. Thornhill R, Gangestad SW. Facial attractiveness. Trends Cogn Sci 1999; 3:452-460. [CrossRef]

19. Cash TF, Kilcullen RN. The eye of the beholder: susceptibility to sexism and beautyism in the evaluation of managerial applicants. J Appl Soc Psychol 1985; 15:591-605. [CrossRef]

20. Chiu RK ve Babcock RD. The relative importance of facial attractiveness and gender in Hong Kong selection decisions. International Journal of Human Resource Management 2002; 13:141-155. [CrossRef]

21. Izzett RR, Legiski W. Group discussion and the influence of defendant characteristics in a smulated jury setting. J Soc Psychol 1974; 93:271-279. [CrossRef]

22. Ahola AS, Christianson SÅ, Hellström Å. Justice needs a blindfold: effects of gender and attractiveness on prison sentences and attributions of personal characteristics in a judicial process. Psychiatr Psychol Law 2009; 16(Suppl.):90-100. [CrossRef]

23. Ritter JM, Casey RJ, Langlois JH. Adults’ responses to infants varying in appearance of age and attractiveness. Child Dev 1991; 62:68-82. [CrossRef]

24. Schein SS, Langlois JH. Unattractive infant faces elicit negative affect from adults. Infant Behav Dev 2015; 38:130-134. [CrossRef]

25. Golle J, Mast FW, Lobmaier JS. Something to smile about: the interrelationship between attractiveness and emotional expression. Cogn Emot 2014; 28:298-310. [CrossRef]

26. Langlois JH, Kalakanis L, Rubenstein AJ, Larson A, Hallam M, Smoot M. Maxims or myths of beauty? A meta-analytic and theoretical review. Psychol Bull 2000; 126:390-423. [CrossRef]

27. Griffin AM, Langlois JH. Stereotype directionality and attractiveness stereotyping: is beauty good or is ugly bad? Soc Cogn 2006; 24:187-206. [CrossRef]

28. Buss DM. Desires in human mating. Ann N Y Acad Sci 2000; 907:39-49. [CrossRef]

29. Rhodes G. The evolutionary psychology of facial beauty. Annu Rev Psychol 2006; 57:199-226. [CrossRef]

30. Grammer K, Fink B, Moller AP, Thornhill R. Darwinian aesthetics: sexual selection and the biology of beauty. Biol Rev Camb Philos Soc 2003; 78:385-407. [CrossRef]

31. Cornwell RE, Boothroyd L, Burt DM, Feinberg DR, Jones BC, Little AC, Pitman R, Whiten S, Perrett DI. Concordant preferences for opposite-sex signals? Human pheromones and facial characteristics. Proc Biol Sci 2004; 271:635-640. [CrossRef]

32. Tottenham N, Leon AC, Casey BJ. The face behind the mask: a developmental study. Dev Sci 2006; 9:288-294. [CrossRef]

33. Hahn AC, Perrett DI. Neural and behavioral responses to attractiveness in adult and infant faces. Neurosci Biobehav Rev 2014; 46:591-603. [CrossRef]

34. Halgren E, Dale AM, Sereno MI, Tootell RB, Marinkovic K, Rosen BR. Location of human face-selective cortex with respect to retinotopic areas. Hum Brain Mapp 1999; 7:29-37. [CrossRef]

35. Freiwald WA, Tsao DY. Functional compartmentalization and viewpoint generalization within the macaque face-processing system. Science 2010; 330:845-851. [CrossRef]

36. Fairhall SL, Ishai A. Effective connectivity within the distributed cortical network for face perception. Cereb Cortex 2007; 17:2400-2406. [CrossRef]

37. Willis J, Todorov A. First impressions: making up your mind after a 100-ms exposure to a face. Psychol Sci 2006; 17:592-598. [CrossRef]

38. Ramsey JL, Langlois JH, Hoss RA, Rubenstein AJ, Griffin AM. Origins of a stereotype: categorization of facial attractiveness by 6-month-old infants. Dev Sci 2004; 7:201-211. [CrossRef]

39. Cellerino A. Psychobiology of facial attractiveness. J Endocrinol Invest 2003; 26(Suppl. 3):45-48.

40. Eisenthal Y, Dror G, Ruppin E. Facial attractiveness: beauty and the machine. Neural Comput 2006; 18:119-142. [CrossRef]

41. Valenzano DR, Mennucci A, Tartarelli G, Cellerino A. Shape analysis of female facial attractiveness. Vision Res 2006; 46:1282-1291. [CrossRef]

42. Bleske-Rechek A, Kolb CM, Stern AS, Quigley K, Nelson LA. Face and body: independent predictors of women’s attractiveness. Arch Sex Behav 2014; 43:1355-1365. [CrossRef]

43. Fink B, Neave N, Seydel H. Male facial appearance signals physical strength to women. Am J Hum Biol 2007; 19:82-87. [CrossRef]

44. Henderson JJA, Anglin JM. Facial attractiveness predicts longevity. Evolution and Human Behavior 2003; 24:351-356. [CrossRef]

45. Jokela M. Physical attractiveness and reproductive success in humans: evidence from the late 20 century United States. Evol Hum Behav 2009; 30:342-350. [CrossRef]

46. Rantala MJ, Coetzee V, Moore FR, Skrinda I, Kecko S, Krama T, Kivleniece I, Krams I. Facial attractiveness is related to women’s cortisol and body fat, but not with immune responsiveness. Biol Lett 2013; 9:20130255. [CrossRef]

47. Law Smith MJ, Perrett DI, Jones BC, Cornwell RE, Moore FR, Feinberg DR, Boothroyd LG, Durrani SJ, Stirrat MR, Whiten S, Pitman RM, Hillier SG. Facial appearance is a cue to oestrogen levels in women. Proc Biol Sci 2006; 273:135-140. [CrossRef]

48. Zaidel DW, Cohen JA. The face, beauty, and symmetry: perceiving asymmetry in beautiful faces. Int J Neurosci. 2005; 115:1165-1173. [CrossRef]

49. Scheib JE, Gangestad SW, Thornhill R. Facial attractiveness, symmetry and cues of good genes. Proc Biol Sci 1999; 266:1913-1917. [CrossRef]

50. Penton-Voak IS, Jones BC, Little AC, Baker S, Tiddeman B, Burt DM, Perrett DI. Symmetry, sexual dimorphism in facial proportions and male facial attractiveness. Proc Biol Sci 2001; 268:1617-1623. [CrossRef]

51. Perrett DI, Burt DM, Penton-Voak IS, Lee KJ, Rowland DA, Edwards R. Symmetry and human facial attractiveness. Evol Hum Behav 1999; 20:295-307. [CrossRef]

52. Mealey L, Bridgstock R, Townsend GC. Symmetry and perceived facial attractiveness: a monozygotic co-twin comparison. J Pers Soc Psychol 1999; 76:151-158. [CrossRef]

53. Rhodes G, Yoshikawa S, Palermo R, Simmons LW, Peters M, Lee K, Halberstadt J, Crawford JR. Perceived health contributes to the attractiveness of facial symmetry, averageness, and sexual dimorphism. Perception 2007; 36:1244-1252. [CrossRef]

54. Rhodes G, Zebrowitz LA, Clark A, Kalick SM, Hightower A, McKay R. Do facial averageness and symmetry signal health? Evol Hum Behav 2001; 22:31-46. [CrossRef]

55. Furlow FB, Armijo-Prewitt T, Gangestad SW, Thornhill R. Fluctuating asymmetry and psychometric intelligence. Proc Biol Sci 1997; 264:823-829. [CrossRef]

56. Hume DK, Montgomerie R. Facial attractiveness signals different aspects of “quality” in women and men. Evol Hum Behav 2001; 22:93-112. [CrossRef]

57. Halberstadt J, Rhodes G. The attractiveness of nonface averages: implications for an evolutionary explanation of the attractiveness of average faces. Psychol Sci 2000; 11:285-289. [CrossRef]

58. Little AC, DeBruine LM, Jones BC. Sex differences in attraction to familiar and unfamiliar opposite-sex faces: men prefer novelty and women prefer familiarity. Arch Sex Behav 2014; 43:973-981. [CrossRef]

59. Buss DM, Schmitt DP. Sexual strategies theory: an evolutionary perspective on human mating. Psychol Rev 1993; 100:204-232. [CrossRef]

60. Simpson JA, Gangestad SW. Individual differences in sociosexuality: evidence for convergent and discriminant validity. J Pers Soc Psychol 1991; 60:870-883. [CrossRef]

61. Apicella CL, Little AC, Marlowe FW. Facial averageness and attractiveness in an isolated population of hunter-gatherers. Perception 2007; 36:1813-1820. [CrossRef]

62. Vingilis-Jaremko L, Maurer D. The influence of averageness on children’s judgments of facial attractiveness. J Exp Child Psychol 2013; 115:624-639. [CrossRef]

63. Ryan MJ, Keddy-Hector A. Directional patterns of female mate choice and the role of sensory biases. Am Nat 1992; 139(Suppl.):4-35. [CrossRef]

64. Folstad I, Karter AJ. Parasites, bright males, and immunocompetence handicap. Am Nat 1992; 139:603-622. [CrossRef]

65. Little AC, Jones BC, Penton-Voak IS, Burt DM, Perrett DI. Partnership status and the temporal context of relationships influence human female preferences for sexual dimorphism in male face shape. Proc Biol Sci 2002; 269:1095-1100. [CrossRef]

66. Penton-Voak IS, Little AC, Jones BC, Burt DM, Perrett DI. Measures of human female condition predict preferences for sexually dimorphic characteristics in men’s faces. J Comp Psychol 2003; 117:264-271. [CrossRef]

67. Scott IM, Pound N, Stephen ID, Clark AP, Penton-Voak IS. Does masculinity matter? The contribution of masculine face shape to male attractiveness in humans. PLoS One 2010; 5:e13585. [CrossRef]

68. Angele MK, Schwacha MG, Ayala A, Chaudry IH. Effect of gender and sex hormones on immune responses following shock. Shock 2000; 14:81-90. [CrossRef]

69. Scott IM, Clark AP, Boothroyd LG, Penton-Voak IS. Do men’s faces really signal heritable immunocompetence? Behav Ecol 2013; 24:579-589. [CrossRef]

70. Voracek M, Manning JT, Ponocny I. Digit ratio (2D:4D) in homosexual and heterosexual men from Austria. Arch Sex Behav 2005; 34:335-340. [CrossRef]

71. Manning JT, Henzi P, Venkatramana P, Martin S, Singh D. Second to fourth digit ratio: ethnic differences and family size in English, Indian and South African populations. Ann Hum Biol 2003; 30:579-588. [CrossRef]

72. Hönekopp J, Voracek M, Manning JT. 2nd to 4th digit ratio (2D:4D) and number of sex partners: evidence for effects of prenatal testosterone in men. Psychoneuroendocrinology 2006; 31:30-37. [CrossRef]

73. Hönekopp J. Digit ratio (2D:4D) and male facial attractiveness: new data and a meta-analysis. Evol Psychol 2013; 11:944-952. [CrossRef]

74. Enlow DH, Hans MG. Essentials of facial growth. Philadelphia: Saunders, 1996.

75. Thornhill R, Møller AP. Developmental stability, disease and medicine. Biol Rev Camb Philos Soc 1997; 72:497-548. [CrossRef]

76. Penton-Voak IS, Chen JY. High salivary testosterone is linked to masculine male facial appearance in humans. Evol Hum Behav 2004; 25:229-241. [CrossRef]

77. Pound N, Penton-Voak IS, Surridge AK. Testosterone responses to competition in men are related to facial masculinity. Proc Biol Sci 2009; 276:153-159. [CrossRef]

78. Wells T, Baguley T, Sergeant M, Dunn A. Perceptions of human attractiveness comprising face and voice cues. Arch Sex Behav 2013; 42:805-811. [CrossRef]

79. Kortet R, Vainikka A, Rantala MJ, Myntti J, Taskinen J. In vitro embryo survival and early viability of larvae in relation to male sexual ornaments and parasite resistance in roach, Rutilus rutilus L. J Evol Biol 2004; 17:1337-1344. [CrossRef]

80. Locatello L, Rasotto MB, Evans JP, Pilastro A. Colourful male guppies produce faster and more viable sperm. J Evol Biol 2006; 19:1595-1602. [CrossRef]

81. Rogers DW, Denniff M, Chapman T, Fowler K, Pomiankowski A. Male sexual ornament size is positively associated with reproductive morphology and enhanced fertility in the stalk-eyed fly Teleopsis dalmanni. BMC Evol Biol 2008; 8:236. [CrossRef]

82. Soler C, de Monserrat JJ, Gutiérrez R, Nu-ez J, Nu-ez M, Sancho M, Pérez-Sánchez F, Cooper TG. Use of the Sperm-Class Analyser for objective assessment of human sperm morphology. Int J Androl 2003; 26:262-270. [CrossRef]

83. Peters M, Rhodes G, Simmons LW. Does attractiveness in men provide clues to semen quality? J Evol Biol 2008; 21:572-579. [CrossRef]

84. Soler C, Kekäläinen J, Nú-ez M, Sancho M, Álvarez JG, Nú-ez J, Yaber I, Gutiérrez R. Male facial attractiveness and masculinity may provide sex- and culture-independent cues to semen quality. J Evol Biol 2014; 27:1930-1938. [CrossRef]

85. Pollet TV, van der Meij L, Cobey KD, Buunk AP. Testosterone levels and their associations with lifetime number of opposite sex partners and remarriage in a large sample of American elderly men and women. Horm Behav 2011; 60:72-77. [CrossRef]

86. Pollet TV, Cobey KD, van der Meij L. Testosterone levels are negatively associated with fatherhood in males, but positively related to offspring count in fathers. PLoS One 2013; 8:e60018. [CrossRef]

87. Jones BC, DeBruine LM, Perrett DI, Little AC, Feinberg DR, Law Smith MJ. Effects of menstrual cycle phase on face preferences. Arch Sex Behav 2008; 37:78-84. [CrossRef]

88. Perrett DI, Lee KJ, Penton-Voak I, Rowland D, Yoshikawa S, Burt DM, Henzi SP, Castles DL, Akamatsu S. Effects of sexual dimorphism on facial attractiveness. Nature 1998; 394:884-887. [CrossRef]

89. Jones BC, Little AC, Watkins CD, Welling LL, DeBruine LM. Reported sexual desire predicts men’s preferences for sexually dimorphic cues in women’s faces. Arch Sex Behav 2011; 40:1281-1285. [CrossRef]

90. Cotterill J. What is really true about the cosmetic industry? Int J Dermatol 1988; 27:682-683. [CrossRef]

91. Marcinkowska UM, Dixson BJ, Kozlov MV, Prasai K, Rantala MJ. Men’s preferences for female facial femininity decline with age. J Gerontol B Psychol Sci Soc Sci 2017; 72:180-186. [CrossRef]

92. Berry DS, Brownlow S (1989) Were the physiognomists right? Personality correlates of facial babyishness. Pers Soc Psychol Bull 1989; 15:266-279. [CrossRef]

93. Grammer K, Thornhill R. Human (Homo sapiens) facial attractiveness and sexual selection: the role of symmetry and averageness. J Comp Psychol 1994; 108:233-242. [CrossRef]

94. Cunningham MR, Barbee AP, Pike CL. What do women want? Facialmetric assessment of multiple motives in the perception of male facial physical attractiveness. J Pers Soc Psychol 1990; 59:61-72. [CrossRef]

95. Saxton TK, Mackey LL, McCarty K, Neave N. A lover or a fighter? Opposing sexual selection pressures on men’s vocal pitch and facial hair. Behav Ecol 2016; 27:512-519. [CrossRef]

96. Rhodes G, Hickford C, Jeffery L. Sex-typicality and attractiveness: are supermale and superfemale faces super-attractive? Br J Psychol 2000; 91:125-140. [CrossRef]

97. Swaddle JP, Reierson GW. Testosterone increases perceived dominance but not attractiveness in human males. Proc Biol Sci 2002; 269:2285-2289. [CrossRef]

98. Conway CA, Jones BC, DeBruine LM, Little AC. Sexual dimorphism of male face shape, partnership status and the temporal context of relationship sought modulate women’s preferences for direct gaze. Br J Psychol 2010; 101:109-121. [CrossRef]

99. Roney JR, Simmons ZL, Gray PB. Changes in estradiol predict within-women shifts in attraction to facial cues of men’s testosterone. Psychoneuroendocrinology 2011; 36:742-749. [CrossRef]

100. Lippa RA. The relation between sex drive and sexual attraction to men and women: a cross-national study of heterosexual, bisexual, and homosexual men and women. Arch Sex Behav 2007; 36:209-222. [CrossRef]

101. Theodoridou A, Rowe AC, Rogers PJ, Penton-Voak IS. Oxytocin administration leads to a preference for masculinized male faces. Psychoneuroendocrinology 2011; 36:1257-1260. [CrossRef]

102. Booth A, Dabbs JM. Testosterone and men’s marriage. Soc Forces 1993; 72:463-477. [CrossRef]

103. Gray PB, Kahlenberg SM, Barrett ES, Lipson SF, Ellison PT. Marriage and fatherhood are associated with lower testosterone in males. Evol Hum Behav 2002; 23:193-201. [CrossRef]

104. Gettler LT, McDade TW, Feranil AB, Kuzawa CW. Longitudinal evidence that fatherhood decreases testosterone in human males. Proc Natl Acad Sci USA 2011; 108:16194-16199. [CrossRef]

105. Haselton MG, Gangestad SW. Conditional expression of women’s desires and men’s mate guarding across the ovulatory cycle. Horm Behav 2006; 49:509-518. [CrossRef]

106. Fink B, Grammer K, Matts PJ. Visual skin color distribution plays a role in the perception of age, attractiveness, and health of female faces. Evol Hum Behav 2006; 27:433-442. [CrossRef]

107. Matts PJ, Fink B, Grammer K, Burquest M. Color homogeneity and visual perception of age, health, and attractiveness of female facial skin. J Am Acad Dermatol 2007; 57:977-984. [CrossRef]

108. Stephen ID, Law Smith MJ, Stirrat MR, Perrett DI. Facial skin coloration affects perceived health of human faces. Int J Primatol 2009; 30:845-857. [CrossRef]

109. Samson N, Fink B, Matts PJ. Visible skin condition and perception of human facial appearance. Int J Cosmet Sci 2010; 32:167-184. [CrossRef]

110. Fink B, Bunse L, Matts PJ, D’Emiliano D. Visible skin colouration predicts perception of male facial age, health and attractiveness. Int J Cosmet Sci 2012; 34:307-310. [CrossRef]

111. Slayden SM, Moran C, Sams WM Jr, Boots LR, Azziz R. Hyperandrogenemia in patients presenting with acne. Fertil Steril 2001; 75:889-892. [CrossRef]

112. Fink B, Neave N. The biology of facial beauty. Int J Cosmetic Sci 2005; 27:317-325. [CrossRef]

113. Elliot AJ, Kayser DN, Greitemeyer T, Lichtenfeld S, Gramzow RH, Maier MA, Liu H. Red, rank, and romance in women viewing men. J Exp Psychol Gen 2010; 139:399-417. [CrossRef]

114. Thornton MJ. The biological actions of estrogens on skin. Exp Dermatol 2002; 11:487-502. [CrossRef]

115. Van Den Abbeele J, Penton-Voak IS, Attwood AS, Stephen ID, Munafò MR. Increased facial attractiveness following moderate, but not high, alcohol consumption. Alcohol Alcohol 2015; 50:296-301. [CrossRef]

116. Russell R. Sex, beauty, and the relative luminance of facial features. Perception 2003; 32:1093-1107. [CrossRef]

117. van den Berghe PL, Frost P. Skin color preference, sexual dimorphism and sexual selection: a case of gene culture evolution? Ethn Racial Stud 1986; 9:87-113. [CrossRef]

118. Swami V, Furnham A, Joshi K. The influence of skin tone, hair length, and hair colour on ratings of women’s physical attractiveness, health and fertility. Scand J Psychol 2008; 49:429-437. [CrossRef]

119. Stephen ID, McKeegan AM. Lip colour affects perceived sex typicality and attractiveness of human faces. Perception 2010; 39:1104-1110. [CrossRef]

120. Kenrick DT, Keefe RC. Age preferences in mates reflect sex differences in human reproductive strategies. Behav Brain Sci 1992; 15:75-133. [CrossRef]

121. Maestripieri D, Klimczuk AC, Traficonte DM, Wilson MC. A greater decline in female facial attractiveness during middle age reflects women’s loss of reproductive value. Front Psychol 2014; 5:179. [CrossRef]

122. Pawłowski B, Dunbar RI. Impact of market value on human mate choice decisions. Proc Biol Sci 1999; 266:281-285. [CrossRef]

123. Mason MF, Tatkow EP, Macrae CN. The look of love: gaze shifts and person perception. Psychol Sci 2005; 16:236-239. [CrossRef]

124. Bashour M, Geist C. Is medial canthal tilt a powerful cue for facial attractiveness? Ophthal Plast Reconstr Surg 2007; 23:52-56. [CrossRef]

125. Geldart S, Maurer D, Carney K. Effects of eye size on adults’ aesthetic ratings of faces and 5-month-olds’ looking times. Perception 1999; 28:361-374. [CrossRef]

126. Hess EH. The role of pupil size in communication. Sci Am 1975; 233:110-112. [CrossRef]

127. Guéguen N. The receptivity of women to courtship solicitation across the menstrual cycle: a field experiment. Biol Psychol 2009; 80:321-324. [CrossRef]

128. Karremans JC, Verwijmeren T. Mimicking attractive opposite-sex others: the role of romantic relationship status. Pers Soc Psychol Bull 2008; 34:939-950. [CrossRef]

129. Otta E, Folladore Abrosio F, Hoshino RL. Reading a smiling face: messages conveyed by various forms of smiling. Percept Mot Skills 1996; 82:1111-1121. [CrossRef]

130. O’Doherty J, Winston J, Critchley H, Perrett D, Burt DM, Dolan RJ. Beauty in a smile: the role of medial orbitofrontal cortex in facial attractiveness. Neuropsychologia 2003; 41:147-155. [CrossRef]

131. Tatarunaite E, Playle R, Hood K, Shaw W, Richmond S. Facial attractiveness: a longitudinal study. Am J Orthod Dentofacial Orthop 2005; 127:676-682. [CrossRef]

132. Tsukiura T, Cabeza R. Shared brain activity for aesthetic and moral judgments: implications for the Beauty-is-Good stereotype. Soc Cogn Affect Neurosci 2011; 6:138-148. [CrossRef]

133. Ishai A. Sex, beauty and the orbitofrontal cortex. Int J Psychophysiol 2007; 63:181-185. [CrossRef]

134. Bzdok D, Langner R, Caspers S, Kurth F, Habel U, Zilles K, Laird A, Eickhoff SB. ALE meta-analysis on facial judgments of trustworthiness and attractiveness. Brain Struct Funct 2011; 215:209-223. [CrossRef]

135. Mende-Siedlecki P, Said CP, Todorov A. The social evaluation of faces: a meta-analysis of functional neuroimaging studies. Soc Cogn Affect Neurosci 2013; 8:285-299. [CrossRef]

136. Ishizu T, Zeki S. Toward a brain-based theory of beauty. PLoS One 2011; 6:e21852. [CrossRef]

137. Aharon I, Etcoff N, Ariely D, Chabris CF, O’Connor E, Bretier HC. Beautiful faces have variable reward value: fMRI and behavioral evidence. Neuron 2001; 32:537-551. [CrossRef]

138. Fink B, Weege B, Neave N, Pham MN, Shackelford TK. Integrating body movement into attractiveness research. Front Psychol 2015; 6:220. [CrossRef]

139. Thornhill R, Gangestad SW. Human fluctuating asymmetry and human sexual behavior. Psychol Sci 1994; 5:297-302. [CrossRef]

140. Manning JT, Scutt D, Lewis-Jones DI. Developmental stability, ejaculate size and sperm quality in men. Evol Hum Behav 1998; 19:273-282. [CrossRef]

141. Tovée MJ, Reinhardt S, Emery JL, Cornelissen PL. Optimum body-mass index and maximum sexual attractiveness. Lancet 1998; 352:548. [CrossRef]

142. Wade TJ, Fuller L, Bresnan I, Schaefer S, Mlynarski L. Weight halo effects: individual differences in personality evaluations and perceived life success of men as a function of weight? Pers Individ Dif 2007; 42:317-324. [CrossRef]

143. Blowers LC, Loxton NJ, Grady-Flesser M, Occhipinti S, Dawe S. The relationship between sociocultural pressure to be thin and body dissatisfaction in preadolescent girls. Eat Behav 2003; 4:229-244. [CrossRef]

144. Myers TA, Crowther JH. Sociocultural pressures, thin-ideal internalization, self-objectification, and body dissatisfaction: could feminist beliefs be a moderating factor? Body Image 2007; 4:296-308. [CrossRef]

145. Grogan S. Body Image: Understanding Body Dissatisfaction in Men, Women and Children. Second ed., London: Routledge, 2008.

146. Seifert T. Anthropomorphic characteristics of centerfold models: trends towards slender figures over time. Int J Eat Disord 2005; 37:271-274. [CrossRef]

147. Stice E, Schupak-Neuberg E, Shaw HE, Stein RI. Relation of media exposure to eating disorder symptomatology: an examination of mediating mechanisms. J Abnorm Psychol 1994; 103:836-840. [CrossRef]

148. Field AE, Camargo CA Jr, Taylor CB, Berkey CS, Colditz GA. Relation of peer and media influences to the development of purging behaviors among preadolescent and adolescent girls. Arch Pediatr Adolesc Med 1999; 153:1184-1189. [CrossRef]

149. Glauert R, Rhodes G, Byrne S, Fink B, Grammer K. Body dissatisfaction and the effects of perceptual exposure on body norms and ideals. Int J Eat Disord 2009; 42:443-452. [CrossRef]

150. Stephen ID, Perera AT. Judging the difference between attractiveness and health: does exposure to model images influence the judgments made by men and women? PLoS One 2014; 9:e86302. [CrossRef]

151. Colabianchi N, Ievers-Landis CE, Borawski EA. Weight preoccupation as a function of observed physical attractiveness: ethnic differences among normal-weight adolescent females. J Pediatr Psychol 2006; 31:803-812. [CrossRef]

152. Furnham A, Radley S. Sex differences in the perception of male and female body shapes. Pers Individ Dif 1989; 10:653-662. [CrossRef]

153. Singh D, Zambarano RJ. Offspring sex ratio in women with android body fat distribution. Hum Biol 1997; 69:545-556.

154. Hartz AJ, Barboriak PN, Wong A, Katayama KP, Rimm AA. The association of obesity with infertility and related menstural abnormalities in women. Int J Obes 1979; 3:57-73.

155. Swami V, Tovée MJ. Female physical attractiveness in Britain and Malaysia: a cross-cultural study. Body Image 2005; 2:115-128. [CrossRef]

156. Singh D. Female mate value at a glance: relationship of waist-to-hip ratio to health, fecundity and attractiveness. Neuro Endocrinol Lett 2002; 23(Suppl.4):81-91.

157. Yu DW, Shepard GH Jr. Is beauty in the eye of the beholder? Nature 1998; 396:321-322. [CrossRef]

158. Westman A, Marlowe F. How universal are preferences for female waist-hip ratios? Evidence from the Hadza of Tanzania. Evol Hum Behav 1999; 20:219-228. [CrossRef]

159. Schutzwohl A. Judging female figures: a new methodological approach to male attractiveness judgments of female waist-to-hip ratio. Biol Psychol 2006; 71:223-229. [CrossRef]

160. Platek SM, Singh D. Optimal waist-to-hip ratios in women activate neural reward centers in men. PLoS One 2010; 5:e9042. [CrossRef]

161. Jasienska G, Ziomkiewicz A, Ellison PT, Lipson SF, Thune I. Large breasts and narrow waists indicate high reproductive potential in women. Proc Biol Sci 2004; 271:1213-1217. [CrossRef]

162. Zaadstra BM, Seidell JC, Van Noord PA, te Velde ER, Habbema JD, Vrieswijk B, Karbaat J. Fat and female fecundity: prospective study of effect of body fat distribution on conception rates. BMJ 1993; 306:484-487. [CrossRef]

163. Lassek WD, Gaulin SJ. Brief communication: menarche is related to fat distribution. Am J Phys Anthropol 2007; 133:1147-1151. [CrossRef]

164. van Hooff MH, Voorhorst FJ, Kaptein MB, Hirasing RA, Koppenaal C, Schoemaker J. Insulin, androgen, and gonadotropin concentrations, body mass index, and waist to hip ratio in the first years after menarche in girls with regular menstrual cycles, irregular menstrual cycles, or oligomenorrhea. J Clin Endocrinol Metab 2000; 85:1394-1400.

165. Little AC, Roberts CS. Evolution, appearance, and occupational success. Evol Psychol 2012; 10:782-801. [CrossRef]

166. Shepherd JA, Strathman AJ. Attractiveness and height: the role of stature in dating preference, frequency of dating and perceptions of attractiveness. Pers Soc Psychol Bull 1989; 15:617-627. [CrossRef]

167. Hensley WE. Height as a basis for interpersonal attraction. Adolescence 1994; 29:469-474.

168. Jackson LA, Ervin KS. Height stereotypes of women and men: the liability of shortness for both sexes. J Soc Psychol 1992; 132:433-445. [CrossRef]

169. Chu S, Geary K. Physical stature influences character perception in women. Pers Individ Dif 2005; 38:1927-1934. [CrossRef]

170. Judge TA, Cable DM. The effect of physical height on workplace success and income: preliminary test of a theoretical model. J Appl Psychol 2004; 89:428-441. [CrossRef]

171. Pawlowski B, Dunbar RI, Lipowicz A. Tall men have more reproductive success. Nature 2000; 403:156. [CrossRef]

172. Hampson E, Kimura D. Reciprocal effects of hormonal fluctuations on human motor and perceptual-spatial skills. Behav Neurosci 1988; 102:456-459. [CrossRef]

173. Walsh DG, Hewitt J. Giving men the come on: effect of eye contact and smiling in a bar environment. Percept Mot Skills 1985; 61:873-874. [CrossRef]

174. Dufner M, Rauthmann JF, Czarna AZ, Denissen JJ. Are narcissists sexy? Zeroing in on the effect of narcissism on short-term mate appeal. Pers Soc Psychol Bull 2013; 39:870-882. [CrossRef]

175. McCarty K, Hönekopp J, Neave N, Caplan N, Fink B. Male body movements as possible cues to physical strength: a biomechanical analysis. Am J Hum Biol 2013; 25:307-312. [CrossRef]

176. González-Álvarez J. Men dissociate sexual attraction from moral judgement more than women. Int J Psychol 2015; 52:381-388. [CrossRef]

177. Harries ML, Walker JM, Williams DM, Hawkins S, Hughes IA. Changes in the male voice at puberty. Arch Dis Child 1997; 77:445-447. [CrossRef]

178. Collins SA. Men’s voices and women’s choices. Anim Behav 2000; 60:773-780. [CrossRef]

179. Evans S, Neave N, Wakelin D. Relationships between vocal characteristics and body size and shape in human males: an evolutionary explanation for a deep male voice. Biol Psychol 2006; 72:160-163. [CrossRef]

180. Pisanski K, Fraccaro PJ, Tigue CC, O’Connor JJ, Feinberg DR. Return to Oz: voice pitch facilitates assessments of men’s body size. J Exp Psychol Hum Percept Perform 2014; 40:1316-1331. [CrossRef]

181. Tusing KJ, Dillard JP. The sounds of dominance: vocal precursors of perceived dominance during interpersonal influence. Hum Commun Res 2000; 26:148-171. [CrossRef]

182. Cartei V, Bond R, Reby D. What makes a voice masculine: physiological and acoustical correlates of women’s ratings of men’s vocal masculinity. Horm Behav 2014; 66:569-576. [CrossRef]

183. Hughes SM, Dispenza F, Gallup GG Jr. Rating of voice attractiveness predict sexual behavior and body configuration. Evol Hum Behav 2004; 25:295-304. [CrossRef]

184. Feinberg DR, Jones BC, Law Smith MJ, Moore FR, DeBruine LM, Cornwell RE, Hillier SG, Perrett DI. Menstrual cycle, trait estrogen level, and masculinity preferences in the human voice. Horm Behav 2006; 49:215-222. [CrossRef]

185. Tobias JA, Montgomerie R, Lyon BE. The evolution of female ornaments and weaponry: social selection, sexual selection and ecological competition. Philos Trans R Soc Lond B Biol Sci 2012; 367:2274-2293. [CrossRef]

186. Kraaijeveld K. Degree of mutual ornamentation in birds is related to divorce rate. Proc Biol Sci 2003; 270:1785-1791. [CrossRef]

187. LeBas NR, Hockham LR, Ritchie MG. Nonlinear and correlational sexual selection on ‘honest’ female ornamentation. Proc Biol Sci 2003; 270:2159-2165. [CrossRef]

188. Berglund A, Rosenqvist G, Bernet P. Ornamentation predicts reproductive success in female pipefish. Behav Ecol Sociobiol 1997; 40:145-150. [CrossRef]

189. Ueda S, Koyama T. Influence of eye make-up on the perception of gaze direction. Int J Cosmet Sci 2011; 33:514-518. [CrossRef]

190. Mulhern R, Fieldman G, Hussey T, Lévêque JL, Pineau P. Do cosmetics enhance female Caucasian facial attractiveness? Int J Cosmet Sci 2003; 25:199-205. [CrossRef]

191. Ueno A, Ito A, Kawasaki I, Kawachi Y, Yoshida K, Murakami Y, Sakai S, Iijima T, Matsue Y, Fujii T. Neural activity associated with enhanced facial attractiveness by cosmetics use. Neurosci Lett 2014; 566:142-146. [CrossRef]

192. Elliot AJ, Niesta D. Romantic red: red enhances men’s attraction to women. J Pers Soc Psychol 2008; 95:1150-1164. [CrossRef]

193. Guéguen N, Jacob C. Clothing color and tipping: Gentlemen patrons give more tips to waitresses with red clothes. Journal of Hospitality and Tourism Research 2012; 38:275-280. [CrossRef]

194. Guéguen N. Color and women attractiveness: when red clothed women are perceived to have more intense sexual intent. J Soc Psychol 2012; 152:261-265. [CrossRef]

195. Pazda AD, Prokop P, Elliot AJ. Red and romantic rivalry: viewing another woman in red increases perceptions of sexual receptivity, derogation, and intentions to mate-guard. Pers Soc Psychol Bull 2014; 40:1260-1269. [CrossRef]

196. Hesslinger VM, Goldbach L, Carbon CC. Men in red: A reexamination of the red-attractiveness effect. Psychon Bull Rev 2015; 22:1142-1148. [CrossRef]

11th National Alcohol and Substance Abuse Congress
Creative Commons Lisansı

Dusunen Adam: The Journal of Psychiatry and Neurological Sciences is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License.
Düşünen Adam - Psikiyatri ve Nörolojik Bilimler Dergisi
Bakırköy Prof. Dr. Mazhar Osman Ruh Sağlığı ve Sinir Hastalıkları Eğitim ve Araştırma Hastanesi
Yerküre Tanıtım ve Yayıncılık Hizmetleri A.Ş.