2nd to 4th digit ratio and offspring sex ratio

There is evidence that the ratio of the length of the 2nd and 4th digit (2D:4D) is negatively related to prenatal and adult concentrations of testosterone. It has also been reported that high levels of testosterone at conception in both fathers and mothers are associated with an increased sex ratio (proportion of males at birth). It follows from these observations that low values of 2D:4D may be related to high sex ratio. We present evidence from three populations (English, Spanish and Jamaican) that 2D:4D is negatively related to sex ratio, independent of the sex and ethnicity of the parents.     

Low digit ratio 2D:4D in alcohol dependent patients

The ratio of the lengths of the second and fourth finger (2D∶4D) has been described as reflecting the degree of prenatal androgen exposure in humans. 2D∶4D is smaller for males than females and is associated with traits such as left-handedness, physical aggression, attention-deficit-hyperactivity disorder and a genetic polymorphism of the androgen receptor. All of these traits are known to be correlated to the vulnerability for alcohol dependency. We therefore hypothesized low 2D∶4D in patients with alcohol dependency. In the present study on 131 patients suffering from alcohol dependency and 185 healthy volunteers, we found that alcohol dependent patients had smaller 2D∶4D ratios compared to controls with preserved sexual dimorphism but with reduced right-left differences. The detection of alcohol dependency based on 2D∶4D ratios was most accurate using the right hand of males (ROC-analysis: AUC 0.725, sensitivity 0.667, specificity 0.723). These findings provide novel insights into the role of prenatal androgen exposure in the development of alcohol dependency and for the use of 2D∶4D as a possible trait marker in identifying patients with alcohol dependency.     

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.     

Re-examining the Manning hypothesis: androgen receptor polymorphism and the 2D:4D digit ratio

The ‘Manning hypothesis,’ the idea that small differences in the ratio of the lengths of the human second to fourth digits—the 2D:4D ratio—reflect differences in the level of fetal androgen exposure, has been highly influential in the biological and biobehavioral sciences. The ratio is widely used to investigate the involvement of fetal androgens in the differentiation of sexually dimorphic traits. The validity of such studies is based on the premise that individual differences in the size of the 2D:4D ratio mirror differences across individuals in developmental levels of androgen exposure in a dose-dependent manner. Despite its widespread adoption by researchers, clinical evidence has yet to confirm that individual gradation in the ratio denotes differences in testosterone action. Key support for the view that 2D:4D does, in fact, reflect fetal testosterone in a graded fashion is the finding, based on a single small-sample study, that the magnitude of 2D:4D covaries with a polymorphic repeat (CAG) sequence in exon 1 of the gene coding the androgen receptor, AR. In a larger independent sample, we reexamine this genetic association and fail to substantiate a correlation between AR CAG length and 2D:4D. Combined with other recent reports, these data question one of the fundamental pieces of evidence on which the Manning hypothesis rests and raise new issues regarding the extent to which 2D:4D is a valid reflection of differences in fetal testosterone action in normally developing individuals.     

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.     

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.     

Second to fourth digit ratio (2D:4D) and concentrations of circulating sex hormones in adulthood

Background  

The second to fourth digit ratio (2D:4D) is used as a marker of prenatal sex hormone exposure. The objective of this study was to examine whether circulating concentrations of sex hormones and SHBG measured in adulthood was associated with 2D:4D.     

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.     

Unilateral and bilateral hybridization barriers in inter-series crosses of 4x 2EBN Solanum stoloniferum, S. pinnatisectum, S. cardiophyllum, and 2x 2EBN S. tuberosum haploids and haploid-species hybrids

Wild Mexican potato species are an important untapped source of useful variation for potato improvement. Introgression methods such as 2n gametes, chromosome doubling, and crossing with disomic 4x 2 endosperm balance number (EBN) bridge species have been used to overcome post-zygotic endosperm failure according to the EBN hypothesis. Stylar barriers can prevent zygote formation, bilaterally when zygote formation is blocked in both directions of the cross or unilaterally when zygote formation is blocked in self incompatible (SI) × self compatible (SC) crosses. In several Solanaceae species, the S-locus for SI has been implicated in interspecific incompatibility. The objectives of this research were to determine if: (1) disomic 4x 2EBN Solanum stoloniferum can be used as a bridge species for introgression of the Mexican 2x 1EBN species Solanum cardiophyllum and Solanum pinnatisectum, (2) pre- and/or post-zygotic barriers limit hybridization among EBN compatible Solanum inter-series crosses, and (3) reproductive barriers act unilaterally or bilaterally. Fruit formation and seed set was recorded for inter-pollinations of S. stoloniferum, 4x 2EBN chromosome doubled S. cardiophyllum and S. pinnatisectum, and 2x 2EBN S. tuberosum haploids (HAP) or haploid-species hybrids (H-S). In vivo pollen tube growth was analyzed for each cross combination with fluorescence microscopy. Attempts to create bridge hybrids between S. stoloniferum, and S. cardiophyllum or S. pinnatisectum were not successful. Pre- and post-zygotic barriers prevented seed formation in crosses involving S. cardiophyllum and S. pinnatisectum. Self compatibility in S. stoloniferum and S. pinnatisectum suggests that the S-locus does not contribute to the stylar barriers observed with these species. Alternatively, the presence of functional and nonfunctional (SC) S-alleles may explain interspecific incompatibility in intra- and inter-ploidy crosses. A non-stylar unilateral incongruity was discovered in H-S/HAP × S. stoloniferum crosses, indicating either a post-zygotic barrier, or a pre-zygotic barrier acting at or within the ovary. Furthermore, lack of S. stoloniferum pollen rejection may occur through absence of S. stoloniferum pollen-active genes needed to initiate pollen rejection, or through competitive interaction in S-locus heterozygous S. stoloniferum pollen. Introgression strategies using these species would benefit potato breeding by introducing genetic diversity for several traits simultaneously through co-current introgression.     

Evolution of sexual dimorphism in the digit ratio 2D:4D--relationships with body size and microhabitat use in iguanian lizards

The ratio between lengths of digit II and IV (digit ratio 2D:4D) is a morphological feature that likely affects tetrapod locomotor performances in different microhabitats. Modifications of this trait may be triggered by changes in steroids concentrations during embryo development, which might reflect direct selection acting on digit ratio or be solely a consequence of hormonal differences related for example to body size. Here we apply both conventional and phylogenetic analyses on morphological data from 25 lizard species of 3 families of Iguania (Iguanidae, Polychrotidae, and Tropiduridae), in order to verify whether selective pressures related to locomotion in different microhabitats could override the prenatal developmental cues imposed on the digit ratio 2D:4D by differences in body size between males and females. Data suggest that this trait evolved in association with ecological divergence in the species studied, despite the clear effect of body size on the digit ratio 2D:4D. The ecological associations of size-corrected digit ratios were restricted to one sex, and females of species that often use perches exhibited small digit ratios in the front limbs, which translated into larger sexual dimorphism indexes of arboreal species. The results, together with the subsequent discussion, provide outlines for further investigation about possible developmental mechanisms related to the evolution of adaptive changes in digit lengths that may have occurred during the evolution of ecological divergence in squamates.     

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.     

Evidence for assortative mating on digit ratio (2D:4D), a biomarker for prenatal androgen exposure

The second-to-fourth digit ratio (2D:4D) presents an anatomical sex difference in humans. On average, men tend to have lower 2D:4D compared with women. There is fairly strong evidence for a role of the 2D:4D ratio as a biomarker for the organizational (permanent) effects of prenatal testosterone on the brain and behaviour. Recently, an accumulating research programme has shown 2D:4D to be related to a multitude of sex-dependent, hormonally influenced biosocial traits and phenotypes which reach into the domains of ability, behaviour, fertility, health, personality and sexuality. This study investigated the degree of assortative mating (spousal similarity) in a sample of 239 native Austrian couples of parental or grandparental age, all of them having reproduced. Results included: (i) significant spousal correlations of +0.19 and +0.18 for right-hand and left-hand 2D:4D, respectively, and +0.24 for average 2D:4D; (ii) no assortative mating effect on the right-minus-left difference in 2D:4D; (iii) indications consistent with a possible generational decrease of spousal similarity in 2D:4D; (iv) a prevalence of couples with a lower right-hand 2D:4D observed in the husband compared with his wife; and (v) relations of spousal 2D:4D patterns to spousal age differences, such that matings of men with more male-typical trait expressions (namely, a generally low right-hand 2D:4D or showing a lower right-minus-left 2D:4D difference than their wives) implicated larger male-minus-female age differences, i.e. younger wives. It is argued that assortative mating on 2D:4D operates indirectly and may be mediated through the assortment on other, more perceptible, physical traits and psychological phenotypes that entertain associations with 2D:4D and are relevant for courtship and mate choice.     

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.     

Differences in the 2nd to 4th digit length ratio in humans reflect shifts along the common allometric line

Ratios often lead to biased conclusions concerning the actual relationships between examined traits and comparisons of the relative size of traits among groups. Therefore, the use of ratios has been abandoned in most comparative studies. However, ratios such as body mass index and waist-to-hip ratio are widely used in evolutionary biology and medicine. One such, the ratio of the 2nd to the 4th finger (2D : 4D), has been the subject of much recent interest in both humans and animals. Most studies agree that 2D : 4D is sexually dimorphic. In men, the 2nd digit tends to be shorter than the 4th, while in women the 2nd digit tends to be of the same size or slightly longer than the 4th. Nevertheless, here we demonstrate that the sexes do not greatly differ in the scaling between the 2nd and 4th digit. Sexual differences in 2D : 4D are mainly caused by the shift along the common allometric line with non-zero intercept, which means 2D : 4D necessarily decreases with increasing finger length, and the fact that men have longer fingers than women. We conclude that previously published results on the 2D : 4D ratio are biased by its covariation with finger length. We strongly recommend regression-based approaches for comparisons of hand shape among different groups.     

Sex differences, activation level, and bilateral electrodermal activity

The effects of the activation level and subject's sex on bilateral skin conductance measures were studied. Thirty right-handed subjects (15 males and 15 females) were exposed to three types of stimulus conditions: rest-period, verbal task and spatial task. Results showed that no relationship was observed between EDA asymmetry and the increase in the activation level induced by the verbal and the spatial tasks. Males showed both higher SCRs and greater frequency of responses on the left than on the right hand. The direction of electrodermal asymmetry remained constant regardless of the stimulus conditions. It was concluded that sex differences are important in the study of EDA asymmetry and that this asymmetry appeared to depend on peripheral variations.     

Quantitative genetics and behavioural correlates of digit ratio in the zebra finch

A recent study on a captive zebra finch population suggested that variation in digit ratio (i.e. the relative length of the second to the fourth toe) might be an indicator of the action of sex steroids during embryo development, as is widely assumed for human digits. Zebra finch digit ratio was found to vary with offspring sex, laying order of eggs within a clutch, and to predict aspects of female mating behaviour. Hence, it was proposed that the measurement of digit ratio would give insights into how an individual's behaviour is shaped by its maternal environment. Studying 500 individuals of a different zebra finch population I set out to: (1) determine the proximate causes of variation in digit ratio by means of quantitative genetics and (2) to search for phenotypic and genetic correlations between digit ratio, sexual behaviour and aspects of fitness. In contrast to the earlier study, I found no sexual dimorphism in digit ratio and no effect of either laying order or experimentally altered hatching order on digit ratio. Instead, I found that variation in digit ratio was almost entirely additive genetic, with heritability estimates ranging from 71 to 84%. The rearing environment (from egg deposition to independence) explained an additional 5-6% of the variation in digit ratio, but there was no indication of any maternal effects transmitted through the egg. I found highly significant phenotypic correlations (and genetic correlations of similar size) between digit ratio and male song rate (positive correlation) as well as between digit ratio and female hopping activity in a choice chamber (negative correlation). Rather surprisingly, the strength of these correlations differed significantly between subsequent generations of the same population, illustrating how quickly such correlations can appear and disappear probably due to genotype-environment interactions.     

Sex differences and bilateral electrodermal activity: a replication

The present study was aimed at replicating the results of a previous work on sex differences and electrodermal asymmetry from our laboratory (Román, et al. 1987). Skin conductance was bilaterally recorded in a sample of 44 dextral volunteers (22 males and 22 females) during a stimulus-free period, and the performance of two tasks (verbal and spatial). Subjects were grouped into two groups of right-hand and left-hand dominance in their electrodermal responses (EDRs) in accordance with their laterality coefficient scores at rest. During the tasks subjects appeared clearly differentiated by their lateralization in the magnitude and frequency of EDRs independently of gender: right-hand responders showed higher electrodermal activity on their right hand, while left-hand responders showed higher electrodermal activity on their left hand. Sex differences were not observed within each responsiveness pattern. The orientation of phasic electrodermal changes remained constant throughout the different conditions.     

Unilateral and bilateral expression of a quantitative trait: asymmetry and symmetry in coronal craniosynostosis

Bilateral symmetry in vertebrates is imperfect and mild asymmetries are found in normal growth and development. However, abnormal development is often characterized by strong asymmetries. Coronal craniosynostosis, defined here as consisting of premature suture closure and a characteristic skull shape, is a complex trait. The premature fusion of the coronal suture can occur unilaterally associated with skull asymmetry (anterior plagiocephaly) or bilaterally associated with a symmetric but brachycephalic skull. We investigated the relationship between coronal craniosynostosis and skull bilateral symmetry. Three-dimensional landmark coordinates were recorded on preoperative computed tomography images of children diagnosed with coronal nonsyndromic craniosynostosis (N = 40) and that of unaffected individuals (N = 20) and analyzed by geometric morphometrics. Our results showed that the fusion pattern of the coronal suture is similar across individuals and types of coronal craniosynostosis. Shape analysis showed that skulls of bilateral coronal craniosynostosis (BCS) and unaffected individuals display low degrees of asymmetry, whereas right and left unilateral coronal craniosynostosis (UCS) skulls are asymmetric and mirror images of one another. When premature fusion of the coronal suture (without taking into account cranial dysmorphology) is scored as a qualitative trait, the expected relationship between trait frequency and trait unilateral expression (i.e. negative correlation) is confirmed. Overall, we interpret our results as evidence that the same biological processes operate on the two sides in BCS skulls and on the affected side in UCS skulls, and that coronal craniosynostosis is a quantitative trait exhibiting a phenotypic continuum with BCS displaying more intense shape changes than UCS.