Digit length ratios predict reactive aggression in women, but not in men

Considerable evidence suggests that digit length and dermatoglyphic asymmetry patterns in the hand are affected by early exposure to androgens. Because androgens play an important role in sexual differentiation of morphological and behavioral traits, digit length patterns often display sex differences. When present, sex differences in digit lengths are more pronounced on the right side as compared to the left side. Moreover, the ratio of the second to fourth digit length (2D:4D) in the right hand is inversely correlated with testosterone (T) in men. Because T is implicated in agonistic behavior, 2D:4D may be used as a marker of androgen exposure and subsequent behavioral variation in adulthood. Consequently, we investigated the relationships among 2D:4D, directional asymmetry of 2D:4D (left hand 2D:4D minus right hand 2D:4D) as a variant of T, and human reactive aggression. One hundred young men (n = 51) and women (n = 49) participated in our experiments (mean age = 20.1 years). Participants called two noncompliant confederates to solicit donations for a fictitious charity organization and selected follow-up letters after the calls. The force exerted when hanging up the phone and the "tone" of the follow-up letters were used to assess reactive aggression. High aggression scores were associated with high directional asymmetry of 2D:4D and masculinized (low) right hand 2D:4D, only in females and under high provocation. Directional asymmetry of 2D:4D was positively correlated with T in males (pooled data, n = 97). Taken together, these data confirm the predominantly right-sided influence of androgens on digit length and suggest that digit length ratios may be associated with female reactive aggression when sufficient provocation is present.     

2D:4D finger-length ratios in children and adults with gender identity disorder

Previous research suggests that prenatal testosterone affects the 2D:4D finger ratio in humans, and it has been speculated that prenatal testosterone also affects gender identity differentiation. If both things are true, then one would expect to find an association between the 2D:4D ratio and gender identity. We measured 2D:4D in two samples of patients with gender identity disorder (GID). In Study 1, we compared the 2D:4D ratios of 96 adult male and 51 female patients with GID to that of 90 heterosexual male and 112 heterosexual female controls. In Study 2, we compared the 2D:4D ratios of 67 boys and 34 girls with GID to that of 74 control boys and 72 control girls. In the sample of adults with GID, we classified their sexual orientation as either homosexual or non-homosexual (in relation to their birth sex) to examine whether or not there were any within-group differences as a function of sexual orientation. In the sample of adult men with GID (both homosexual and non-homosexual) and children with GID, we found no evidence of an altered 2D:4D ratio relative to same-sex controls. However, women with GID had a significantly more masculinized ratio compared to the control women. This last finding was consistent with the prediction that a variance in prenatal hormone exposure contributes to a departure from a sex-typical gender identity in women.     

A longitudinal study of digit ratio (2D:4D) and other finger ratios in Jamaican children

It has been hypothesised that the ratio between the length of the 2nd and 4th digits (2D:4D) is a correlate of prenatal sex steroids, and this relationship is strongest for the right hand. Furthermore, it has been suggested that 2D:4D is sexually dimorphic, the dimorphism is determined early, and 2D:4D among children is stable with growth. Here, we present the first longitudinal study of right and left hand 2D:4D. Our sample was 108 (54 males) Jamaican children. The first measurements were made in 1998 when mean age was 9.68 +/- 1.39 years, and a second set of measurements were made in 2002. We found that: (i) there was a small increase in 2D:4D with age which was lowest in the right hand; (ii) 2D:4D was sexually dimorphic, the means for males and females differed in the same direction in the 1998 and 2002 samples, and the sex difference was significant in the 1998 but not in the 2002 sample; (iii) the correlation between the 1998 and 2002 measurements of 2D:4D was high, indicating that rank order of the ratio was stable across year groups; and (iv) the rate of change in 2D:4D did not differ significantly across year groups. We conclude that 2D:4D increases slightly with age in children with the effect less marked for the right hand (i.e. the hand which is likely to show the strongest association with prenatal steroids), 2D:4D is sexually dimorphic from an early age, and the rank order of 2D:4D is stable in children. We discuss the implications of our findings for the status of 2D:4D as a correlate of prenatal sex steroids. The patterns of change in other finger ratios are also considered.     

2D:4D ratios in the first 2 years of life: Stability and relation to testosterone exposure and sensitivity

The relative lengths of the 2nd and 4th digits (2D:4D) may provide an easily measurable and stable anthropometric index of prenatal androgen exposure, but no study has examined the development of 2D:4D in infancy and the potential impact of neonatal testosterone levels. We collected 2D:4D ratios from 364 children between 0 and 2 years of age. Saliva samples were collected from 236 of these children 3 months after birth and analyzed for testosterone. In addition, 259 children provided DNA samples which were genotyped for the CAG repeat polymorphism in the androgen receptor. There was substantial variability across age in 2D:4D. Sex differences were small compared to adults and did not consistently reach statistical significance. This suggests that 2D:4D may not function well as a proxy measure of prenatal testosterone exposure in infancy. In addition, the interaction of salivary T and CAG repeats predicted right hand digit ratio at 12 months and left hand digit ratio at 12 months and 24 months in males. The interaction of salivary testosterone and CAG repeat length also predicted change in left hand 2D:4D from 2 weeks to 12 months in males. This suggests that 2D:4D in adults may reflect, in part, neonatal testosterone exposure. No significant relationships were observed within females. No significant relationships were observed when salivary testosterone and CAG repeats were examined independent of each other. Results have important implications for the design and interpretation of studies which use 2D:4D as a proxy measure of prenatal testosterone exposure.     

Testing the prenatal androgen hypothesis: measuring digit ratios, sexual orientation, and spatial abilities in adults

The present study examined whether the following variables putatively associated with prenatal androgens are inter-related in women: spatial abilities, sexual orientation, and 2nd to 4th finger (digit) length ratio (2D:4D). Participants were 99 healthy premenopausal women tested in the menstrual phase of the ovarian cycle between 0800 and 0930 hr. Women completed the Kinsey scales of sexual orientation, and were either strictly heterosexual (HS; N=79) or not-strictly heterosexual (NHS; N=20). Photocopies of the two hands were collected, and participants completed the revised Vandenberg Mental Rotations test, the Paper Folding test, and a short version of the Guilford-Zimmerman Spatial Orientation Test. Results showed that NHS women exhibited superior spatial ability relative to HS women. No significant difference was found between the HS and NHS women in the 2D:4D digit ratio. There was no association between the digit ratio and spatial performance. These results support an association between increased spatial abilities and heteroflexible sexual orientation, which may possibly be mediated by high prenatal androgens.     

Digit ratio (2D:4D) and physical fitness in males and females: Evidence for effects of prenatal androgens on sexually selected traits

It has been suggested that male achievement in sports and athletics is correlated with a putative measure of prenatal testosterone the 2nd to 4th digit ratio (2D:4D). It is not known whether this association also extends to females, or whether the association results from an effect of testosterone on behavior (such as exercise frequency) or on physical fitness. Here, we report for the first time data from two studies which consider associations between 2D:4D and physical fitness in females in addition to males: Study I--in a sample of teenage boys (n = 114) and girls (n = 175), their 'physical education grade' was negatively associated with 2D:4D of the right hand (boys), and right and left hand (girls), and Study II-among a sample of young men (n = 102) and women (n = 77), a composite measure of physical fitness was negatively related to right hand 2D:4D in men and left hand 2D:4D in women. We conclude that 2D:4D is negatively related to physical fitness in both men and women. In Study II, there was evidence that the relationship between physical fitness and 2D:4D in men was mediated through an association with exercise frequency. Thus, 2D:4D in males may be a negative correlate of frequent exercise which then relates to achievement in sports and athletics.     

Sex differences in the relative lengths of metacarpals and metatarsals in gorillas and chimpanzees

Sex differences other than the simple dimorphism in size were documented for the metapodials of two primate species. Lengths of metacarpals and metatarsals were obtained from the skeletons of 64 gorillas and 42 chimpanzees. Length ratios were constructed for all possible pairings of the five bones in each individual hand and foot. For both species, several of these length ratios exhibited substantial differences between the sexes. Body size was not the basis for these sex differences; when specimens of similar size were compared, the sex differences remained. In humans, length ratios for the fingers and toes also have previously been demonstrated to exhibit sex differences, and the length ratio for the index and ring fingers (the 2D:4D ratio) has been shown to correlate with various medical conditions. Various facts suggest that length ratios in human digits are associated with androgen exposure, probably during prenatal development. For gorillas, the metacarpal length ratio showing the largest sex difference was 4Mc:5Mc in both hands, and the metatarsal length ratio showing the largest sex difference was 1Mt:2Mt in the left foot. Sex differences in length ratios also existed for chimpanzees, but they were generally smaller than for gorillas. Apparently, both gorillas and chimpanzees are affected by developmental mechanisms, possibly androgenic mechanisms, similar to those in humans. Analyses of previous measurements [Susman, R.L., 1979 Comparative and functional morphology of hominoid fingers. Am. J. Phys. Anthropol. 50, 215-236] revealed that all components of the rays are not affected equally by whatever mechanisms are responsible for the sex differences in length ratios.     

Relationship between the 2D:4D and prenatal testosterone,adult level testosterone,and testosterone change: Meta-analysis of 54 studies

The ratio between the hands' second to the fourth finger (2D:4D) is commonly hypothesized to result from prenatal testosterone. The 2D:4D has also been hypothesized to relate to adult-level testosterone and, more recently, to the testosterone response to a challenging situation. In the present work, we tested these core assumptions. Drawing from, in total, 54 studies and 8077 participants, we investigated whether the 2D:4D is related to adult level testosterone (44 studies), testosterone change (6 studies), and prenatal testosterone (10 studies). We found no evidence of the relationship between the above testosterone types and digit ratios. Furthermore, there was no relationship between testosterone and the right and left 2D:4D, male and female 2D:4D, and the 2D:4D and testosterone measurement (i.e., measured in blood or saliva). However, we found some evidence that prenatal testosterone measured in amniotic fluid (but not cord blood) might be related to the digit ratios—further studies are necessary to validate this observation. In summary, considering the current state of knowledge, any conclusions drawn from the assumption of the digit ratios as the proxy for testosterone (prenatal, adult level, or testosterone change under a challenging situation) warrant great caution.     

Early prenatal attainment of adult metacarpal-phalangeal rankings and proportions.

As shown in 56 human embryos and fetuses between 15 and 104 mm in crown-rump length, "adult" metacarpal-phalangeal length rankings are attained by the seventh intrauterine week and near-adult bone-to-bone ratios or proportions by the theirteenth week. Micrometric measurements of optically-projected histological hand sections show relative elongation of the distals between the 15-29 mm and 30-44 mm crown-rump range, and relative reduction to radiogrammetrically-determined adult proportions by the 90-104 mm CRL.     

Bonobos have a more human-like second-to-fourth finger length ratio (2D:4D) than chimpanzees: a hypothesized indication of lower prenatal androgens

The ratio of the second-to-fourth finger lengths (2D:4D) has been proposed as an indicator of prenatal sex differentiation. However, 2D:4D has not been studied in the closest living human relatives, chimpanzees (Pan troglodytes) and bonobos (Pan paniscus). We report the results from 79 chimpanzees and 39 bonobos of both sexes, including infants, juveniles, and adults. We observed the expected sex difference in 2D:4D, and substantially higher, more human-like, 2D:4D in bonobos than chimpanzees. Previous research indicates that sex differences in 2D:4D result from differences in prenatal sex hormone levels. We hypothesize that the species difference in 2D:4D between bonobos and chimpanzees suggests a possible role for early exposure to sex hormones in the development of behavioral differences between the two species.     

The effects of prenatal testosterone on wages: Evidence from Russia

Is in utero exposure to testosterone correlated with earnings? The question matters for understanding determinants of wage differences that have attracted so much attention among economists in the past decade. Evidence indicates that markers for early testosterone exposure are correlated with traits like risk-taking and aggressiveness. But it is not at all clear how such findings might map into labor market success. We combine unique data from the Russian Longitudinal Monitoring Survey with measured markers (2D:4D ratios) for testosterone exposure and find that lower digit ratios (higher T) correlate with higher wages for women and for men, when controlling for age, education and occupation. There is also some evidence of a potential non-linear, inverse U-effect of digit ratios on wages but this is sensitive to choice of specification. These findings are consistent with earlier work on prenatal T and success in careers (Coates et al., 2009) but inconsistent with the work of Gielen et al. (2016) who find differing effects for men and women.     

The relative lengths and weights of metacarpals and metatarsals in baboons (papio hamadryas)

The lengths and weights of metacarpals and metatarsals were measured in two collections of baboon skeletons-33 animals, all born and raised in the wild, and 60 animals, nearly all born and raised in captivity. For both length and weight, ratios were constructed for all possible pairings of the five bones in each individual hand and foot. The pattern of results was not identical for the two collections, but substantial sex differences existed in both collections for many of the ratios calculated. Nearly all of the large sex differences were in the direction of the length or weight ratio being smaller for males than for females. For the length ratios showing the largest sex differences, those differences were larger for the right hand than for the left, and larger for the left foot than for the right, but this pattern was less evident for the weight ratios. Some length ratios for human fingers show sex differences of the sort seen in the metacarpals and metatarsals of these baboons, and the human differences in relative length exist early in life. The implication is that the marked sex difference in androgen exposure during prenatal development may effect the relative size of the individual bones of the extremities in male and female humans and baboons. The relative sizes of the bones of the hand and foot may provide helpful supplemental information on the relations between species, thus making the study of hand and foot bones in other primates and mammals worthwhile.     

The relationships between otoacoustic emissions and relative lengths of fingers and toes in humans

Both otoacoustic emissions (OAEs) and the relative length of the index and ring fingers (the 2D:4D ratio) exhibit large sex differences, and both exhibit masculinization effects in female homosexuals and bisexuals. Because these sex differences exist in young children, the implication is that both types of measure are affected by prenatal androgen exposure, but it has been unknown to what degree these two types of measure are related. Accordingly, OAEs and the relative lengths of the fingers and the toes were measured in 59 heterosexual females, 55 heterosexual males, 29 homosexual females, and 33 homosexual males. The correlations between the two types of measure were unexpectedly quite low in both the heterosexual and nonheterosexual groups. For example, the correlation between number of spontaneous OAEs per ear and 2D:4D was less than 0.25, for both sexes and both sexual orientations. One interpretation of these results is that the prenatal hormonal mechanisms producing the sex differences in OAEs differ in quality, degree, or timing from those producing the sex differences in relative finger length. That is, OAEs and 2D:4D may be windows onto slightly different prenatal processes or times during prenatal development. Measures of mental-rotation ability also were obtained on these participants, and those correlations with relative finger length also were small.     

Environmental levels of oestrogenic and antiandrogenic compounds feminize digit ratios in male rats and their unexposed male progeny

Digit length ratios, especially the second-to-fourth digit ratio (2D : 4D), are associated with various pathological and behavioural conditions in many species including humans and are dependent upon prenatal androgen to oestrogen balance. It is unknown whether digit ratios are modified by environmental exposure to ubiquitous endocrine disruptors. We studied the effect on adult male Wistar rat digit ratios of a gestational exposure to the oestrogenic and antiandrogenic compounds bisphenol A (BPA), genistein and vinclozolin, in low doses, and in combination with investigating in parallel a possible sexual dimorphism of this trait. We also investigated the effects on the male progeny not exposed during gestation. X-rays were taken of the left and right forepaws, and 2D–5D proximal to distal phalanx distances were measured by a standardized procedure based on semi-automatic image analysis. We provide evidence that there is a sexual dimorphism of digit ratios in the Wistar rat, and we found that BPA alone or in combination with genistein and vinclozolin significantly feminized digit ratios in male rats. Intriguingly, significant feminization of digit ratios was also found in the unexposed male progeny of males that had been exposed to compound mixtures. In conclusion, prenatal environmental levels of endocrine-active substances permanently disrupt digit ratios. Digit ratio measurement in adults is thus a promising biomarker of prenatal exposure to low-dose endocrine disruptors in rodents, with potential implications for future studies in humans.     

Individual differences in women''s facial preferences as a function of digit ratio and mental rotation ability

The second-to-fourth digit ratio (2D:4D) and mental rotation (MR) ability are sexually dimorphic traits that appear early in development and have been correlated with exposure to prenatal androgens (Grimshaw, Sitarenios, & Finegan, 1995; Lutchmaya, Baron-Cohen, Raggatt, Knickmeyer, & Manning, 2004). The current study examined how 2D:4D and MR differences among women of European descent (N=41) were related to their (a) preferences for male faces: attractive (ATM), short-term (STM), and long-term mate (LTM), and (b) psychological femininity and masculinity, as measured by the Bem Sex-Role Inventory (BSRI) (Bem, 1981). To examine potential changes in facial preferences over their menstrual cycle, participants' preferences were measured during two experimental sessions separated by 2 weeks. The results indicated that (a) femininity scores decreased with decreasing 2D:4D, (b) masculinity scores increased with faster MR, (c) women preferred a more masculine male face for an STM than for an LTM, and (d) preference changes over the menstrual cycle varied systematically with 2D:4D. When compared with women with high 2D:4D ratios, low 2D:4D women (e) preferred a more masculine LTM, (f) recalled less parental bonding, (g) had shorter intimate relationships, and (h) reported more menstrual irregularity. The results are interpreted as support for an interactive hormonal theory of physical attraction.     

Sex dimorphism in digital formulae of children

This paper presents results of a study designed to: 1) test for a sex difference in the relative lengths of the finger bones, including the second-to-fourth digit ratio (2D:4D), using left-hand radiographs taken in young children, 2) test whether sex differences can be explained by sex differences in fetal growth, and 3) test the serial stability of sex differences in relative digit lengths, including 2D:4D. Results are presented from 1,060 subjects of the California Child Health and Development Studies. One serial replication at about 9 years old is available from 271 subjects. Results indicate that relative digit lengths are sex-dimorphic in children (Manning et al. [1998] Hum. Reprod. 13:3000-3004, [2004] Early Hum. Dev. 80:161-168). Sex differences in digit length ratios are more pronounced within sibships, where shared family factors are controlled, and are not strongly associated with gross measures of fetal growth, like birth length or weight. Thus, sex differences in the fetal growth of the body are not implicated in sex differences in digital formulae, leaving open the possibility of more direct hormonal and/or genetic causation. However, 2D:4D declined between ages 6-8 in a longitudinal sample, and was a less consistent sex-dimorphic marker than 3D:4D across ethnic groups, suggesting that 3D:4D may be a better marker of perinatal sex differentiation. Prior conflicting findings about 2D:4D may be partly explained by variations in age and ethnicity of populations studied.     

Digit ratio (2D:4D) and behavioral differences between inbred mouse strains

Digit ratio (2D:4D) is a trait, which is sexually differentiated in a variety of species. In humans, males typically have shorter second digits (2Ds) (index fingers) compared to fourth digits (4Ds) (ring fingers) whereas females' fingers are more equal in length. Smaller, more masculine, digit ratios are thought to be associated with higher prenatal testosterone levels, greater sensitivity to prenatal androgens or both. Men with more masculine digit ratios have shown increased ability, achievement and speed in sports and tend to report that they are more physically aggressive. Previous research has shown the same sexually differentiated pattern in the hind paws of laboratory mice as in human hands, males have lower 2D:4D than females. We measured hind paw digit ratio in mice of eight inbred strains. These measurements were made while blind to strain, sex and whether the paw was from the left or right side. We found large differences in digit ratio between the strains and suggest that inbred mice are a promising system for investigating the correlation between digit ratio and behavioral traits.