Digit ratio (2D:4D) and rowing ergometer performance in males and females

Fetal and adult testosterone may be vital in the establishment and maintenance of sex-dependent abilities associated with male physical competitiveness. It has been shown that digit ratio (2D:4D) is negatively associated with prenatal testosterone, and it is also negatively associated with ability in sports such as football, skiing, middle distance running, and endurance running, which are dependent upon an efficient cardiovascular system. The relationship between digit ratio and sports requiring high power (physical strength) output in addition to well-developed cardiovascular systems has not been defined. This study investigated this association in male and female young adult rowers. Participants (77 male and 70 female) were student rowers encompassing a range of abilities from the University of Cambridge. Bilateral digit measurements were taken blind from each subject using Mitutoyo vernier calipers. Rowing performance over 2,000 m was assessed using the Concept 2 rowing ergometer. Significant negative correlations were observed between 2,000 m ergometer performance and male digit ratios, which persisted following adjustment for rowing experience and height. However, no such significant association was found in females despite a comparable sample size. Our data indicate that digit ratio is a predictor of ability in rowing, a sport which requires both cardiovascular efficiency and high power output, in males but not females. This in turn suggests that fetal testosterone exposure has long-term effects on traits associated with physical power in males but not females, suggesting a sex-difference in the capacity to respond to such exposures.     

Second to fourth digit ratio and the sporting success of sumo wrestlers

The second (index finger) to fourth (ring finger) digit length ratio (2D:4D) is known to be a putative marker of prenatal exposure to testosterone. It has been reported that fetal and adult testosterone may be critical for development of physical and mental traits such as cardiovascular system, reaction time, aggressiveness and masculinity. Testosterone-driven attributes are associated with success in male-to-male physical competition, which may be proxied by ability in sports. Many researchers have found that 2D:4D is sexually dimorphic and is a negative correlate of athletic performance. This study aims to investigate the associations of 2D:4D with measures of power as another possible testosterone-associated trait using ability in sumo wrestling as a proxy for male physical competitiveness. The measures of sumo performance comprised the sumo ranks and winning percentages of 142 Japanese professional sumo wrestlers. We found that sumo wrestlers with low 2D:4D had higher sumo ranks and better winning records. The significant negative associations between 2D:4D and the athletic prowess of sumo wrestlers provide further evidence of the possible link between high testosterone levels and muscle strength. The relatively small effect sizes found in this study, however, imply that 2D:4D may be a weaker predictor for sports requiring explosive power than for those requiring endurance.     

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.     

Testosterone and intrasexual competition in men: is there any relation with digit ratio (2D:4D)?

Digit ratio 2D:4D is a sexually dimorphic characteristic and it is believed that this difference is related to high levels of prenatal testosterone and circulating testosterone in men. High levels of testosterone are also associated with traits related to competitiveness. Therefore, this study aimed to examine the relation between intrasexual competition and 2D:4D in men. One hundred thirteen college men answered a questionnaire to measure their scores of intrasexual competition and donated a saliva sample to measure their testosterone levels; finally, the finger length from both hands was measured. It was found a positive correlation between testosterone levels and intrasexual competition scores, and a negative correlation between testosterone levels and left 2D:4D. Finally, we did not find a significant association between digit ratios 2D:4D and intrasexual competition scores. Our study shows that men with higher testosterone levels also have higher intrasexual competition scores and lower values of left digit ratio 2D:4D. Further studies will have to take into account fluctuations in testosterone over the time to observe if the relation between competitiveness scores and digit ratios 2D:4D becomes significant.     

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.     

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.     

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.     

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 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.     

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.     

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.     

Digit ratio (2D:4D) and performance in male surfers

The relative length of the second and fourth digits (2D:4D) is thought to be negatively related to prenatal testosterone and positively related to prenatal estrogen. Low 2D:4D has been linked to various measures of performance in a range of sports (e.g., soccer, rugby). In this study, we consider the relationship between 2D:4D and performance among male surfers. Our sample comprised 46 competitors in the Men's 5-star Professional World Qualifying Series surfing competition in Newquay, United Kingdom, in 2009. Three experienced surfing coaches rated the participants for overall surfing ability. The coach's ratings were significantly correlated with one another and an overall measure of surfing performance was obtained by calculating the mean of the 3 ratings. In addition, the final placing of the Newquay competition was used as an additional performance measure. Mean 2D:4D (SD) was as follows: right 0.994 (0.023) and left 0.976 (0.028). We found that right 2D:4D (but not left 2D:4D or right-left 2D:4D) was significantly negatively correlated with coaches' ratings (r(s) = 0.58) and the competition result (r(s) = 0.30). It appears that in line with other sports that low right 2D:4D (high prenatal testosterone and low prenatal estrogen) correlates to high surfing ability in men.     

Visualizing facial shape regression upon 2nd to 4th digit ratio and testosterone

Sex steroids are supposed to moderate the differences between male and female facial characteristics. Studies on women's preferences for male faces reported increased preferences for facial architecture developed under the influence of testosterone as this may indicate masculinity, dominance and social status. Recent research demonstrates that facial sexual dimorphism does not only develop at puberty but may be organized much earlier in ontogeny. However, the actual cause and timing of variation in facial shape due to sex-steroids remains speculative. This study uses data from Neave and colleagues who measured digit ratio (2D:4D) as a proxy to prenatal testosterone and also salivary testosterone samples in order to study differential effects of androgens on perceived male facial shape. Male facial shape was regressed upon 2D:4D ratio and circulating levels of testosterone by means of geometric morphometric methods. We found some evidence for opposite effects of early androgen action (via 2D:4D ratio) on the upper and the lower face respectively (i.e. low 2D:4D ratio results in a relatively robust and prominent lower face), whereas circulating testosterone seems to cause a rather uniform elongation of the face. Local deformations primarily show pronounced and medially tailed eyebrows for the shapes associated with increasing salivary testosterone. These preliminary results suggest that prenatal and pubertal testosterone have differential effects on male facial shape that should be considered in future studies on women's preferences towards male facial appearance.     

Digit ratio (2D:4D) predicts facial, but not voice or body odour, attractiveness in men

There is growing evidence that human second-to-fourth digit ratio (or 2D:4D) is related to facial features involved in attractiveness, mediated by in utero hormonal effects. The present study extends the investigation to other phenotypic, hormone-related determinants of human attractiveness: voice and body odour. Pictures of faces with a neutral expression, recordings of voices pronouncing vowels and axillary odour samples captured on cotton pads worn for 24 h were provided by 49 adult male donors. These stimuli were rated on attractiveness and masculinity scales by two groups of 49 and 35 females, approximately half of these in each sample using hormonal contraception. Multivariate regression analyses showed that males' lower (more masculine) right 2D:4D and lower right-minus-left 2D:4D (Dr-l) were associated with a more attractive (and in some cases more symmetrical), but not more masculine, face. However, 2D:4D and Dr-l did not predict voice and body odour masculinity or attractiveness. The results were interpreted in terms of differential effects of prenatal and circulating testosterone, male facial shape being supposedly more dependent on foetal levels (reflected by 2D:4D ratio), whereas body odour and vocal characteristics could be more dependent on variation in adult circulating testosterone levels.     

Second-to-fourth digit ratio and facial shape in boys: the lower the digit ratio, the more robust the face

During human ontogeny, testosterone has powerful organizational and activational effects on the male organism. This has led to the hypothesis that the prenatal environment (as studied through the second-to-fourth digit ratio, 2D : 4D) is not only associated with robust adult male faces that are perceived as dominant and masculine, but also that there is an activational step during puberty. To test the latter, we collected digit ratios and frontal photographs of right-handed Caucasian boys (aged 4-11 years) along with age, body height and body weight. Using geometric morphometrics, we show a significant relationship between facial shape and 2D : 4D before the onset of puberty (explaining 14.5% of shape variation; p = 0.014 after 10 000 permutations, n = 17). Regression analyses depict the same shape patterns as in adults, namely that the lower the 2D : 4D, the smaller and shorter the forehead, the thicker the eyebrows, the wider and shorter the nose, and the larger the lower face. Our findings add to previous evidence that certain adult male facial characteristics that elicit attributions of masculinity and dominance are determined very early in ontogeny. This has implications for future studies in various fields ranging from social perception to life-history strategies.     

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.     

Finger length ratios (2D:4D) in anthropoids implicate reduced prenatal androgens in social bonding

The second‐to‐fourth digit ratio (2D:4D) has been proposed as a biomarker reflecting prenatal androgen effects (PAE), such that individuals with lower ratios have experienced higher PAE than those with higher ratios. 2D:4D has been correlated with a number of sex‐linked traits in humans such as aggression, promiscuity, and competitiveness. In addition, polygynous societies reportedly have lower 2D:4D (higher PAE) than more monogamous populations. This evidence suggests that PAE may be implicated in the development of sexually selected behaviors in humans. To place 2D:4D research into a broader context, we test the relationship between digit ratios and behavior across nonhuman anthropoids; polygynous species, with higher levels of intrasexual competition, should have more pronounced markers of PAE (lower 2D:4D) than pair‐bonded species. Our results accord with those found in humans: 2D:4D is lower in polygynous species and higher (lower PAE) in pair‐bonded species. Old World monkeys have low, and relatively invariant 2D:4D (high PAE), which is coupled with high levels of intrasexual competition. This contrasts with higher and more variable ratios in both great apes and New World monkeys. In addition, both male and female ratios decrease with increasing levels of intrasexual competition. Human ratios are intermediate between pair‐bonded and more promiscuous hominoids. We propose that PAE may be involved in promoting species characteristic social behavior in anthropoids. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

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 male's perceived status rather than his physical attraction to women.     

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.     

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.