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

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 CAG's 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.     

Brief Communication: Latent toxoplasmosis and salivary testosterone concentration--important confounding factors in second to fourth digit ratio studies

A sexually dimorphic characteristic, the second to fourth digit ratio (2D:4D ratio), has been shown to reflect the prenatal concentration of sex steroid hormones and to correlate with many personality, physiological, and life history traits. The correlations are usually stronger for the right than the left hand. Most studies have shown that the 2D:4D ratio does not vary with age or postnatal concentration of sex steroid hormones. Recently, a strong association between left hand 2D:4D ratio and infection with a common human parasite Toxoplasma has been reported. We hypothesized that the confounding effect of Toxoplasma infection on left hand 2D:4D ratio could be responsible for the stronger association between different traits and right hand rather than left hand 2D:4D ratio. This confounding effect of toxoplasmosis could also be responsible for the difficulty in finding an association between 2D:4D ratio and age or postnatal steroid hormone concentration. To test this hypothesis, we analyzed the association between sex and age and 2D:4D ratio in a population of 194 female and 106 male students with and without controlling for the confounding variables of Toxoplasma infection and testosterone concentration. Our results showed that the relationship between age and sex and 2D:4D ratio increased sharply when Toxoplasma infection and testosterone concentration were controlled. These results suggest that left hand 2D:4D ratio is more susceptible to postnatal influences and that the confounding factors of Toxoplasma infection, testosterone concentration and possibly also age, should be controlled in future 2D:4D ratio studies. Because of a stronger 2D:4D dimorphism in Toxoplasma-infected than Toxoplasma-free subjects, we predict that 2D:4D ratio dimorphism as well as right hand/left hand 2D:4D ratio dimorphism will be higher in countries with a high prevalence of Toxoplasma infection than in those with a low prevalence.     

2D:4D ratios predict hand grip strength (but not hand grip endurance) in men (but not in women)

In humans, the ratio of the second digit to the fourth digit — the 2D:4D ratio — is a sexually dimorphic trait (men, on average, exhibit lower 2D:4D ratios than do women) that is influenced by prenatal testosterone exposure, but not by circulating testosterone levels in adulthood. Consequently, 2D:4D ratios are commonly used as indirect measures of prenatal testosterone exposure. Many studies have examined the associations of 2D:4D ratios with sexually dimorphic adaptations that are thought to be influenced by such exposure, including physical prowess. The existing literature, however, remains unclear as to (1) whether 2D:4D ratios are more closely linked to strength or to endurance; and (2) whether 2D:4D ratios are linked with physical prowess for both men and women. In 100 men and 122 women, the relationship of 2D:4D ratios with maximum voluntary contraction (MVC) scores (hand grip strength) and maximum endurance time (MET) scores (local muscular endurance) using a hand dynamometer was examined. Controlling for age, height, weight, and average digit length, we found that 2D:4D ratios significantly predicted MVC scores in men, but not in women. 2D:4D ratios did not significantly predict MET scores for either sex. These results suggest that prenatal testosterone exposure in this sample is significantly related to hand grip strength in men, but not in women (and to local muscular endurance in neither sex), and, therefore, that strength, rather than local muscular endurance, potentially drives the relationship between 2D:4D ratios and physical prowess.     

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.     

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.     

Sex differences in 2D:4D and aggression in children and adolescents from five regions of Russia

The aim of this study was to consider digit ratio (2D:4D: a putative marker of prenatal testosterone and estrogen levels) and aggression in a sample of 1,452 children and adolescents (mean age 13.6 years) from five regions of Russia. The 2D:4D was calculated from direct measurements of the fingers, and aggression scores were obtained from completed Buss and Perry (J Pers Soc Psychol 63 (1992) 452–459) aggression questionnaires. The 2D:4D demonstrated significant sexual dimorphism, with lower 2D:4D in boys in all regions. Physical aggression scores were highest in boys, but verbal aggression, anger and hostility were highest in girls. The highest right hand 2D:4D in boys was found in the most northerly population (Central Russia Region). Our data revealed small, but highly significant negative correlations between right 2D:4D, right–left 2D:4D (D_R‐L) and self‐ratings on physical aggression in boys, but not in girls. These relationships remained after considering Russian ethnics only, and controlling for region. We suggest that the associations may be due to sex differences in prenatal androgen secretion. Am J Phys Anthropol 152:130–139, 2013. © 2013 Wiley Periodicals, Inc.     

Brief communication: Familial resemblance in digit ratio (2D:4D)

Familial resemblance in the second‐to‐fourth digit ratio (2D:4D), a proxy for prenatal androgen action, was studied in 1,260 individuals from 235 Austrian families. In agreement with findings from twin studies of 2D:4D, heritability estimates based on parent–child and full‐sib dyad similarity indicated substantial genetic contributions to trait expression (57% for right hand, 48% for left hand 2D:4D). Because twin studies have found nonadditive genetic as well as shared environmental effects on 2D:4D to be negligible or nil, these family‐based estimates in all likelihood reflect the narrow‐sense (additive genetic) heritability of the trait. Directional (right‐minus‐left) asymmetry in 2D:4D was only weakly heritable (6%). The pattern of same‐sex and different‐sex parent–child and full‐sib correlations yielded no evidence for X‐linked inheritance. This is surprising, considering evidence for associations of male 2D:4D with sensitivity to testosterone (functional variants of the X‐linked androgen receptor gene). 2D:4D was particularly strongly heritable through male lines (father–son and brother–brother correlations), thus raising the possibility that Y‐linked genes (such as the sex‐determining region SRY) might influence 2D:4D expression. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Digit ratio (2D:4D) and handgrip strength in subjects of Han ethnicity: Impact of sex and age

The ratio of the length of the second finger to the fourth finger (2D:4D) is considered to be a putative proxy of prenatal exposure to testosterone, and has been increasingly used as a promising tool to evaluate the impact of prenatal androgenization in humans in such traits as physical performance. In this study, for the first time, we present 2D:4D data on adult participants of Han ethnicity. We consider the sexual dimorphism of 2D:4D and handgrip strength, and also report the relationship between 2D:4D and handgrip strength of males and females. The sample consisted of 54 males and 55 females recruited from a remote village in the Qinling Mountains, China. We found sexual dimorphism of both 2D:4D and handgrip strength, i.e., males had lower 2D:4D and right‐left 2D:4D than females and greater handgrip strength than females. There was a sex‐specific correlation between 2D:4D and handgrip strength, i.e., 2D:4D in the right hand was negatively correlated with handgrip strength in males but not in females. This relationship may be driven by sexual selection operating on fetal programming. Am J Phys Anthropol 149:266–271, 2012. © 2012 Wiley Periodicals, Inc.     

Interpreting digit ratio (2D:4D)–behavior correlations: 2D:4D sex difference,stability, and behavioral correlates and their replicability in young children

The popularity of using the ratio of the second to the fourth digit (2D:4D) to study influences of early androgen exposure on human behavior relies, in part, on a report that the ratio is sex-dimorphic and stable from age 2 years (Manning etal., 1998). However, subsequent research has rarely replicated this finding. Moreover, although 2D:4D has been correlated with many behaviors, these correlations are often inconsistent. Young children's 2D:4D–behavior correlations may be more consistent than those of older individuals, because young children have experienced fewer postnatal influences. To evaluate the usefulness of 2D:4D as a biomarker of prenatal androgen exposure in studies of 2D:4D–behavior correlations, we assessed its sex difference, temporal stability, and behavioral correlates over a 6- to 8-month period in 126, 2- to 3-year-old children, providing a rare same-sample replicability test. We found a moderate sex difference on both hands and high temporal stability. However, between-sex overlap and within-sex variability were also large. Only 3 of 24 correlations with sex-typed behaviors—scores on the Preschool Activities Inventory (PSAI), preference for a boy-typical toy, preference for a girl-typical toy, were significant and in the predicted direction, all of which involved the PSAI, partially confirming findings from another study. Correlation coefficients were larger for behaviors that showed larger sex differences. But, as in older samples, the overall pattern showed inconsistency across time, sex, and hand. Therefore, although sex-dimorphic and stable, 2D:4D–behavior correlations are no more consistent for young children than for older samples. Theoretical and methodological implications are discussed.     

The Relationship between the Second-to-Fourth Digit Ratio and Behavioral Sexual Dimorphism in School-Aged Children

Sexually dimorphic brain development and behavior are known to be influenced by sex hormones exposure in prenatal periods. On the other hand, second-to forth digit ratio (2D/4D) has been used as an indirect method to investigate the putative effects of prenatal exposure to androgen. In the present study, we herein investigated the relationship between gender-role play behavior and the second-to-fourth digit ratio (2D/4D), which has been used as an indirect method to investigate the putative effects of prenatal exposure to androgens, in school-aged children. Among 4981 children who became 8 years old by November 2014 and were contactable for this survey by The Hokkaido Study of Environment and Children''s Health, 1631 (32.7%), who had data for 2D/4D and Pre-school Activities Inventory (PSAI) as well as data for the survey at baseline, were available for analysis. Parents sent reports of PSAI on the sex-typical characteristics, preferred toys, and play activities of children, and black and white photocopies of the left and right hand palms via mail. PSAI consisted of 12 masculine items and 12 feminine items, and a composite score was created by subtracting the feminine score from the masculine score, with higher scores representing masculine-typical behavior. While composite scores in PSAI were significantly higher in boys than in girls, 2D/4D was significantly lower in boys than in girls. Although the presence or absence of brothers or sisters affected the composite, masculine, and feminine scored of PSAI, a multivariate regression model revealed that 2D/4D negatively correlated with the composite scores of PSAI in boys, whereas no correlation was found in girls. Although 2D/4D negatively correlated with the masculine score in boys and girls, no correlation was observed between 2D/4D and the feminine score. In conclusion, although social factors, such as the existence of brother or sisters, affect dimorphic brain development and behavior in childhood, the present study revealed that the prenatal hormonal environment was an important factor influencing masculine-typical dimorphic brain development and behavior in school-aged children.     

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.     

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.     

Testosterone enhancement during pregnancy influences the 2D:4D ratio and open field motor activity of rat siblings in adulthood

In humans, the relationship between the prenatal testosterone exposure and the ratio of the second and the fourth digits (2D:4D) has been extensively studied. Surprisingly, data on this relationship have thus far been lacking in experimental animals such as rats. We studied the effect of maternal testosterone enhancement during pregnancy on the digit ratio and open field activity of adult progeny in Wistar rats. Elevated levels of maternal testosterone resulted in lower 2D:4D ratios and an elongated 4D on the left and right forepaws in both males and females. We found no sex difference in 2D:4D in control animals. In the open field test, control females were more active than control males and testosterone females, while the activity of testosterone females did not differ from that of control males. We found a positive correlation between motor activity and the right forepaw 2D:4D ratio of control males and females. Prenatal exposure to testosterone resulted in the disappearance of this correlation in both males and females. Our results show that elevated levels of testosterone during the prenatal period can influence forepaw 4D length, 2D:4D ratio, and open field motor activity of rats, and that these variables are positively correlated. Thus, this approach represents a noninvasive and robust method for evaluating the effects of prenatal testosterone enhancement on anatomical and physiological parameters.     

Digit Ratio (2D:4D) and Handgrip Strength in Hani Ethnicity

Introduction

The ratio of the length of the second finger to the fourth finger (2D:4D) in humans is considered as a putative marker of prenatal exposure to testosterone, and has been progressively adopted as one useful tool to evaluate the effect of prenatal hormones in some traits such as physical ability. Handgrip strength is one authentic measure of physical ability and is generally used on the anthropological research within an evolutionary viewpoint.

Methods

Here we present the first evidence on 2D:4D and handgrip strength on adult participants of Hani ethnicity and explore the relationship between digit ratio (2D:4D) and handgrip strength. We examined 2D:4D and handgrip strength of 80 males and 60 females at Bubeng village, in the Yunnan province of China.

Results

The mean 2D:4D in females was higher than that in males for each hand. Females showed significantly higher 2D:4D than males in the right hand rather than in the left hand. Males displayed significantly higher handgrip strength than females for both hands. Handgrip strength decreased with age for both sexes. A significant negative correlation between 2D:4D and handgrip strength was found in the right hand of males.

Conclusion

The relationship between 2D:4D and handgrip strength may be attributed to evolutionary drive of sexual selection operating on fetal programming.     

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 2nd:4th digit ratio,sexual dimorphism,population differences,and reproductive success: evidence for sexually antagonistic genes?

The ratio between the length of the 2nd and 4th digit (2D:4D) is sexually dimorphic, with mean male 2D:4D lower than mean female 2D:4D. It recently was suggested that 2D:4D is negatively correlated with prenatal testosterone and positively correlated with prenatal estrogen. It is argued that high prenatal testosterone and low estrogen (indicated by low 2D:4D) favors the male fetus and low prenatal testosterone and high estrogen (indicated by high 2D:4D) favors the female fetus. The patterns of expression of 2D:4D are interpreted in terms of sexually antagonistic genes.We report data on the following. (a) reproductive success and 2D:4D from England, Germany, Spain, Hungary (ethnic Hungarians and Gypsy subjects), Poland, and Jamaica (women only). Significant negative associations were found between 2D:4D in men and reproductive success in the English and Spanish samples and significant positive relationships between 2D:4D in women and reproductive success in the English, German, and Hungarian samples. The English sample also showed that married women had higher 2D:4D ratios than unmarried women, suggesting male choice for a correlate of high ratio in women, and that a female 2D:4D ratio greater than male 2D:4D predicted high reproductive success within couples. Comparison of 2D:4D ratios of 62 father:child pairs gave a significant positive relationship. This suggested that genes inherited from the father had some influence on the formation of the 2D:4D ratio. Waist:hip ratio in a sample of English and Jamaican women was negatively related to 2D:4D. (b) Sex and population differences in mean 2D:4D in samples from England, Germany, Spain, Hungary (including ethnic Hungarians and Gypsy subjects), Poland, Jamaica, Finland, and South Africa (a Zulu sample). Significant sex and population differences in mean 2D:4D were apparent.     

The Left Hand Second to Fourth Digit Ratio (2D:4D) Is Not Related to Any Physical Fitness Component in Adolescent Girls

Introduction

The second to fourth-digit-ratio (2D:4D), a putative marker of prenatal androgen action and a sexually dimorphic trait, has been suggested to be related with fitness and sports performance, although results are not univocal. Most studies however focus on a single aspect of physical fitness or one sports discipline.

Methods

In this study the 2D:4D ratio of 178 adolescent girls (age 13,5–18 y) was measured on X-rays of the left hand. The relation between 2D:4D digit ratio and multiple aspects of physical fitness (balance, speed of limb movement, flexibility, explosive strength, static strength, trunk strength, functional strength, running speed/agility, and endurance) was studied by correlation analyses and stepwise multiple regression. For comparison the relation between these physical fitness components and a selected number of objectively measured anthropometric traits (stature, mass, BMI, somatotype components and the Bayer & Bailey androgyny index) are presented alongside the results of 2D:4D digit ratio.

Results

Left hand 2D:4D digit ratio (0.925±0.019) was not significantly correlated with any of the physical fitness components nor any of the anthropometric variables included in the present study. 2D:4D did not enter the multiple stepwise regression for any of the physical fitness components in which other anthropometric traits explained between 9,2% (flexibility) and 33,9% (static strength) of variance.

Conclusion

Unlike other anthropometric traits the 2D:4D digit ratio does not seem to be related to any physical fitness component in adolescent girls and therefore most likely should not be considered in talent detection programs for sporting ability in girls.     

Relative digit lengths and testosterone levels in Guinea baboons

A growing body of literature suggests that the ratio of the lengths of the second to fourth digits (2D:4D) on human hands is sexually dimorphic and associated with prenatal exposure to gonadal hormones, circulating serum testosterone, and a number of psychological and behavioral measures. Little research has investigated digit ratios in nonhuman species. In the present study, we investigated sex differences in digit ratios and their possible association with serum testosterone in a captive group of Guinea baboons (Papio papio). Contrary to the sex difference typically reported in humans, male baboons exhibited a substantially larger 2D:4D than did female baboons. Consistent with the human data, however, lower 2D:4D was associated with higher serum testosterone among the males. The present findings suggest that the relationship between digit ratios and male gonadal hormones may be phylogenetically well-conserved, although they also suggest possible species differences in the causal relationships between developmental mechanisms and sex-differentiated digit length patterns.     

Is sexual dimorphism affected by the combined action of prenatal stress and sex ratio?

During embryonic development, offspring are exposed to hormones of both maternal and sibling origin. Maternal stress increases offspring exposure to corticosterone, and, in polytocous animals, the sex ratio or intrauterine position can influence the levels of androgens and estrogens experienced by the offspring. Such hormone exposure has the potential to influence many important morphological and behavioural aspects of offspring, in particular sexually dimorphic traits. Although well known in rodents, the impact of prenatal hormone exposure in other vertebrates is poorly documented. We experimentally investigated the relationship between maternal stress, population density, sex ratio (a surrogate for the degree of exposure to steroids produced by siblings), and sexual dimorphism in a viviparous lizard, Lacerta vivipara. Our results show that prenatal sex ratios have consequences for sexually dimorphic morphology (ventral scale count) in both sexes, but with no effect of maternal stress or any interaction between the two. Embryonic steroid exposure can potentially be an important factor in generating individual variation in natural populations of viviparous animals.     

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.