John T. Manning

The second to fourth digit ratio and variation in the androgen receptor gene

Abstract The second to fourth digit ratio (2D:4D) is sexually dimorphic, with lower mean values in males compared to females. It has been suggested that the sex difference in 2D:4D is determined prenatally, 2D:4D is negatively related to prenatal testosterone and positively to prenatal oestrogen, and that 2D:4D is a marker for levels of sex steroids during brain organisation. There is growing evidence that many sex-dependent behaviours are correlated with 2D:4D. However, there is no direct evidence for an effect of prenatal sex steroids on the digit ratio. The response to prenatal testosterone is dependent on the amount produced and the foetal sensitivity to the hormone. Variation in the X-linked androgen receptor gene (AR) determines sensitivity to testosterone. Alleles of AR with low numbers of CAG triplets respond to testosterone with high transactivational activity, while high numbers of CAGs are associated with increased insensitivity to testosterone. We show in a sample of 50 men (49 Caucasian subjects, 1 Caucasian/Chinese subject) that 2D:4D is a phenotypic correlate of AR structure. Right-hand 2D:4D was positively correlated with CAG number and individuals with low 2D:4D in their right hand compared to left hand had AR alleles with low CAG numbers. We discuss the implications of our findings for our understanding of the aetiology of 2D:4D, its relationships with sex-dependent behaviours, and the evolutionary implications of variation in 2D:4D and AR.

Second to fourth digit ratio, testosterone and perceived male dominance.

Previous studies have shown that male faces with extreme features associated with testosterone are perceived as dominant and masculine. Women have been reported to prefer more masculinized male faces as they may consider testosterone markers to be an ‘honest’ indication of good health, and such considerations may underlie their aesthetic preferences. However, pronounced testosterone facial markers are also associated with dominance, and several negative personality traits. This suggests that female aesthetic preferences may be an adaptive compromise between positive attributes associated with higher than average testosterone, and negative attributes associated with more extreme masculinization. This current study attempts to clarify the role of hormone markers in female perceptions of dominance, masculinity and attractiveness, in male facial images. Recent evidence suggests that the relative length of the 2nd to 4th finger (2D : 4D ratio) is a pointer to prenatal testosterone levels and may thus serve as a window to the prenatal hormonal environment. We measured 2D : 4D in a sample of male college students and took salivary samples to analyse circulating levels of testosterone. Women rated facial images of these males for dominance, masculinity and attractiveness. Our results show that male 2D : 4D was significantly negatively related to perceived dominance and masculinity but not attractiveness. Circulating testosterone levels were not related to dominance, masculinity or attractiveness. These findings suggest that: (i) high prenatal levels of testosterone serve to ‘organize’ male facial features to subsequently reflect dominance and masculine characteristics presumably activated during puberty; and (ii) attractiveness is not directly related to testosterone levels. We conclude that facial dominance and masculinity reflect a males perceived status rather than his physical attraction to women.

Choosy females and correlates of male age

The evolution of female choice is controversial. The males of many polygynous species have bizarre features such as long and elaborate tails which seem to be maladaptive. If females choose to mate with such males how could choice genes increase in frequency? It is argued that (a) male age may be an indicator of fitness with older males having a higher average fitness than young males, (b) the size and complexity of many male ornaments and weapons are positively correlated with age. Females that choose males with, say, a longer than average tail are choosing males who are older than average. The choosy females therefore have male and female offspring with a higher fitness than the progeny of randomly mating females. As female choice genes spread the size or complexity of the age-dependent ornament increases. Mutations increasing size or complexity of such ornaments will also spread. A stable situation will be reached when the advantage of choosing older males is balanced by the disadvantage of, say, very long tails.

The ratio of 2nd to 4th digit length and male homosexuality

Sexual orientation may be influenced by prenatal levels of testosterone and oestrogen. There is evidence that the ratio of the length of 2nd and 4th digits (2D:4D) is negatively related to prenatal testosterone and positively to oestrogen. We report that (a) 2D:4D was lower in a sample of 88 homosexual men than in 88 sex- and age-matched controls recruited without regard to sexual orientation, (b) within the homosexual sample, there was a significant positive relationship between mean 2D:4D ratio and exclusive homosexuality, (c) overall, there was a decrease in 2D:4D from controls to homosexual men to bisexual men and (d) fraternal birth order, a positive predictor of male homosexuality, was not associated with 2D:4D in a sample of 240 Caucasian men recruited without regard to sexual orientation and 45 homosexual men.Further work is needed to confirm the relationships between 2D:4D and sexual orientation. However, these and other recent data tend to support an association between male homosexuality and high fetal testosterone. Very high testosterone levels may be associated with a sexual preference for both men and women.

A preliminary investigation of the associations between personality, cognitive ability and digit ratio

Abstract The ratio between second and fourth digit lengths (digit ratio) is known to be sexually dimorphic, with males having lower values. It is believed that digit ratio acts as a marker of pre-natal testosterone exposure and a number of studies of its correlates support this idea. In the present study, associations between digit ratio and a number of cognitive and personality tests which are known to show male–female score differences were investigated. Evidence for associations in the expected direction was found for sensation seeking, psychoticism and neuroticism, in all cases for females only. No association was found between digit ratio and cognitive test scores.

The 2nd:4th digit ratio and asymmetry of hand performance in Jamaican children

Testosterone, particularly prenatal testosterone, has been implicated in the aetiology of many extragenital sexually dimorphic traits. It is difficult to test directly for the effect of prenatal testosterone in humans. However, Manning, Scutt, Wilson, and Lewis-Jones (1998b) have recently shown that the ratio of the length of the 2nd and 4th digits (2D:4D) in right hands negatively predicts testosterone levels in men. As digit ratios are fixed in utero it may be that the 2D:4D ratio is associated with many prenatally determined sexually dimorphic traits. We tested this for one case by examining the relationship between lateralised hand performance (LHP), as measured by an Annett peg board, and 2D:4D ratio in rural Jamaican children. 2D:4D ratio was measured from photocopies and X rays of hands. A low 2D:4D ratio in the right hand of boys and girls (photocopies) and the right hand of boys only (X rays) was associated with a reduction in rightward performance asymmetry. In both samples the difference in 2D:4D ratio between the hands (2D:4D left hand-2D:4D right hand) showed the strongest relationship with LHP i.e. high ratio in the left and low in the right correlated with a tendency towards a fast performance with the left hand. It is suggested that the 2D:4D ratio may be associated with the expression of other sexually dimorphic behavioural traits.

Fluctuating asymmetry and body weight in men and women: Implications for sexual selection

Abstract Fluctuating asymmetry (FA), small random fluctuations from perfect bilateral symmetry, is an indicator of developmental stability and therefore of good genes. Sexually selected weapons, such as the canine teeth of some primates, often show a negative relationship between FA and size of weapon. This suggests that it is only the males with the best genes who are able to produce large symmetrical weapons. Male body mass, which may improve fighting success, is not bilaterally symmetrical, and little is known of its relationship with FA. It is shown that for human males the body weight in adults is negatively related to FA; this suggests that male body weight is condition-dependent in that it is only individuals with the best genes who are able to develop and maintain large size. There is no relationship between weight and FA in children. For human females, there is a positive correlation between body weight and FA in adults but not in children; the implications of this finding for the understanding of the selection pressures that operate on womens weight are discussed.

Digit ratio (2D:4D), dominance, reproductive success, asymmetry, and sociosexuality in the BBC Internet Study

Digit ratio (2D:4D) may be a correlate of prenatal sex steroids, and has been linked to traits, which are influenced by fetal testosterone and estrogen. Here we consider such links in a large Internet study of sex differences (the BBC Internet Study) in which finger lengths were self‐measured. Consistent with lab‐based findings the 2D:4D in this study shows sexual dimorphism, ethnic differences and higher dimorphism of right 2D:4D than left, thereby indicating that 2D:4D does measure real between‐participant variation. High error in self‐measurement of fingers reduces effect sizes. However, the large sample size gives assurance that significant effects are likely to be real. We controlled for ethnicity and sexual orientation by considering White heterosexuals only (153,429 participants). Sexual dimorphism was confirmed in 2D:4D and for the difference of right–left 2D:4D. After Bonferroni correction we found highly significant relationships with low effect sizes as follows. In males and females there were negative associations between 2D:4D and dominance. In males there were negative associations between 2D:4D and family size and factors associated with reproductive success. For females these associations were positive. For asymmetry we found U‐shaped relationships with 2D:4D in both males and females. We found no relationship between 2D:4D and promiscuity (sociosexuality). In total, we considered 48 relationships and found 29 to be significant. We compare our findings with a similar study reported by Putz et al. (2004), which found only 2 out of 57 correlations to be significant and discuss possible reasons for the discrepancies between the studies. Am. J. Hum. Biol., 2008.

Second to fourth digit ratio and face shape

The average human male face differs from the average female face in size and shape of the jaws, cheek-bones, lips, eyes and nose. It is possible that this dimorphism is determined by sex steroids such as testosterone (T) and oestrogen (E), and several studies on the perception of such characteristics have been based on this assumption, but those studies focussed mainly on the relationship of male faces with circulating hormone levels; the corresponding biology of the female face remains mainly speculative. This paper is concerned with the relative importance of prenatal T and E levels (assessed via the 2D : 4D finger length ratio, a proxy for the ratio of T/E) and sex in the determination of facial form as characterized by 64 landmark points on facial photographs of 106 Austrians of college age. We found that (i) prenatal sex steroid ratios (in terms of 2D : 4D) and actual chromosomal sex dimorphism operate differently on faces, (ii) 2D : 4D affects male and female face shape by similar patterns, but (iii) is three times more intense in men than in women. There was no evidence that these effects were confounded by allometry or facial asymmetry. Our results suggest that studies on the perception of facial characteristics need to consider differential effects of prenatal hormone exposure and actual chromosomal gender in order to understand how characteristics have come to be rated ‘masculine’ or ‘feminine’ and the consequences of these perceptions in terms of mate preferences.

Finger and toe ratios in humans and mice: implications for the aetiology of diseases influenced by HOX genes

The differentiation of gonads, fingers, and toes is influenced by HOXA and HOXD genes. Therefore variation in the development of the gonads, and their fetal products such as testosterone, may be reflected in the morphology of the fingers and toes. One trait, the relative length of the second and fourth digits (2D:4D), shows sex differences (lower values in males) which are determined early, and 2D:4D has been found to correlate with fetal growth, sperm counts, family size, autism, myocardial infarction, and breast cancer. HOX genes are highly conserved in mammals and they influence the differentiation of all the fingers and toes. We suggest that (a) 2D:4D and other ratios of finger and toe length show sex differences throughout the mammals including humans and mice, (b) finger and toe ratios correlate with sex determination, the fetal production of sex steroids, and fetal programming of disease, and (c) HOX gene influences on sex determination, the morphogenesis of the urinogenital system, fertility, haematopoiesis, and breast cancer suggests that finger and toe ratios in humans and mice may correlate with many sex dependent diseases.