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.     

Relative lengths of fingers and toes in human males and females

Digital scans of the hands and feet were obtained from 62 heterosexual females and 60 heterosexual males. Scans only of the hands were obtained from 29 homosexual females and 35 homosexual males. The lengths of the individual fingers and toes were estimated from those images by two experienced judges, and length ratios were constructed for all possible pairs of fingers (or toes) on each hand (or foot). Thumbs were not measured, but the great toe was measured and used to construct length ratios. Past research had concentrated on the relative lengths of the index and ring fingers (the 2D:4D ratio). This ratio is close to 1.0 in females and smaller than 1.0 in males. Here 2D:4D did exhibit the largest sex difference, for both hands, followed by 2D:5D and 3D:4D. The sex differences were larger for the right hand than for the left. For both homosexual females and homosexual males, nearly all of the length ratios for fingers were intermediate to those for heterosexual females and heterosexual males; that is, the ratios of homosexual females were masculinized and those of homosexual males were hypomasculinized, but few of these differences were significant. Because many toes were substantially arched, acceptable estimates of length often could not be obtained from the two-dimensional scans, meaning that conclusions about toes are much less certain than those for fingers. Nevertheless, the length ratios were generally larger for toes than for fingers, and the sex differences were generally smaller for toes.     

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.     

Consistent sex ratio bias of individual female dragon lizards

Sex ratio evolution relies on genetic variation in either the phenotypic traits that influence sex ratios or sex-determining mechanisms. However, consistent variation among females in offspring sex ratio is rarely investigated. Here, we show that female painted dragons (Ctenophorus pictus) have highly repeatable sex ratios among clutches within years. A consistent effect of female identity could represent stable phenotypic differences among females or genetic variation in sex-determining mechanisms. Sex ratios were not correlated with female size, body condition or coloration. Furthermore, sex ratios were not influenced by incubation temperature. However, the variation among females resulted in female-biased mean population sex ratios at hatching both within and among years.     

The development of sex differences in digital formula from infancy in the Fels Longitudinal Study

Relative finger lengths, especially the second-to-fourth finger length ratio, have been proposed as useful markers for prenatal testosterone action. This claim partly depends on an association of relative finger lengths in adults with related sex differences in children and infants. This paper reports the results of a study using serial radiographs to test for both sex differences in the fingers of infants and children and for a relationship between sex differences in the children and infant finger and adult finger length ratios. This is the first study using long-term serial data to evaluate the validity of finger length ratios as markers. We found not only that sex differences in finger length ratios arise prior to puberty, but that sex differences in the fingers of children are highly correlated with adult finger length ratios. Our results strongly encourage the further use of finger length ratios as markers of perinatal testosterone action.     

Food Preferences Among Captive Western Gorillas (Gorilla gorilla gorilla) and Chimpanzees (Pan troglodytes)

I used a zoological park setting to address food preferences among gorillas (Gorilla gorilla gorill) and chimpanzees (Pan troglodytes). Gorillas and chimpanzees are different sizes, and consequently, have been traditionally viewed as ecologically distinct. Sympatric western gorillas and chimpanzees have proved difficult to study in the wild. Limited field data have provided conflicting information about whether gorillas are fundamentally different from chimpanzees in diet and behavior. Fruit eating shapes the behavior of most apes, but it is unclear whether the large-bodied gorillas are an exception to this rule, specifically whether they are less selective and more opportunistic fruit eaters than chimpanzees are. My research provides experimental observational data to complement field data and to better characterize the diets and food preferences of the African apes. During laboratory research at the San Francisco Zoological Gardens, I examined individual and specific differences in food preferences of captive gorillas and chimpanzees via experimental paired-choice food trials with foods that varied in nutritional content. During the study, I offered 2500 paired-food choices to 6 individual gorillas and 2000 additional pairs to them as a group. I also proffered 600 food pairs to 4 individual chimpanzees. Despite expectations of the implications of body size differences for diet, gorillas and chimpanzees exhibited similar food preferences. Both species preferred foods high in non-starch sugars and sugar-to-fiber ratios, and low in total dietary fiber. Neither species avoided foods containing tannins. These data support other suggestions of African apes sharing a frugivorous adaptation.     

Hypoglossal canal size in living hominoids and the evolution of human speech

The relative size of the hypoglossal canal has been proposed as a useful diagnostic tool for the identification of human-like speech capabilities in the hominid fossil record. Relatively large hypoglossal canals (standardized to oral cavity size) were observed in humans and assumed to correspond to relatively large hypoglossal nerves, the cranial nerve that controls motor function of the tongue. It was suggested that the human pattern of tongue motor innervation and associated speech potential are very different from those of African apes and australopithecines; the modern human condition apparently appeared by the time of Middle Pleistocene Homo. A broader interspecific analysis of hypoglossal canal size in primates conducted in 1999 has rejected this diagnostic and inferences based upon it. In an attempt to resolve these differences of opinion, which we believe are based in part on biased size-adjustments and/or unwarranted assumptions, a new data set was collected and analyzed from 298 extant hominoid skulls, including orangutans, gorillas, chimpanzees, bonobos, siamang, gibbons, and modern humans. Data on the absolute size of the hypoglossal nerve itself were also gathered from a small sample of humans and chimpanzee cadavers. A scale-free index of relative hypoglossal canal size (RHCS) was computed as 100 x (hypoglossal canal area(0.5)/oral cavity volume(0.333)). No significant sexual dimorphism in RHCS was discovered in any species of living hominoid, but there are significant interspecific differences in both absolute and relative sizes of the hypoglossal canal. In absolute terms, humans possess significantly larger canals than any other species except gorillas, but there is considerable overlap with chimpanzees. Humans are also characterized by large values of RHCS, but gibbons possess an even larger average mean for this index; siamang and bonobos overlap appreciably with humans in RHCS. The value of RHCS in Australopithecus afarensis is well within both human and gibbon ranges, as are the indices computed for selected representatives of fossil Homo. Furthermore, the size of the hypoglossal nerve itself, expressed as the mass of nerve per millimeter of length, does not distinguish chimpanzees from modern humans. We conclude, therefore, that the relative size of the hypoglossal canal is neither a reliable nor sufficient predictor of human-like speech capabilities, and paleoanthropology still lacks a quantifiable, morphological diagnostic for when this capability finally emerged in the human career.     

Dehydroepiandrosterone-sulfate (DHEA-S), sex,and age in zoo-housed western lowland gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>)

Among humans, dehydroepiandrosterone-sulfate (DHEA-S) declines with age and is hypothesized to be involved in somatic maintenance and healthy aging. Men have significantly higher DHEA-S than women, contradicting longer lifespans in the latter. Declines of DHEA-S with age also are observed in chimpanzees. In both chimpanzees and bonobos, males and females show no differences in DHEA-S production. Based on human and chimpanzee data, gorillas were predicted to show declining DHEA-S with age. Similar to chimpanzees and bonobos, it also was predicted DHEA-S would not be significantly different between males and females. DHEA-S was assayed from serum banked during physical examinations of gorillas housed at three North American zoos (n = 63). Gorillas ranged from 6 to 52 years of age. Differences between males and females were examined using t tests. Linear regression was used to determine the relationship of DHEA-S with age. There was no significant difference in DHEA-S between males and females. Additionally, there was no significant relationship between DHEA-S and age. As predicted, there were no sex-based differences in DHEA-S in gorillas, which is similar to chimpanzees and bonobos but different from modern humans. Unlike chimpanzees and humans, there was no significant relationship between DHEA-S and age in gorillas. The absence of a relationship between age and DHEA-S may be due to the lack of gorillas under age 6 years in this sample as declines in chimpanzees occur prior to age 5 years, more rapid growth and development among gorillas compared with other African hominoids, or a unique pattern of DHEA-S production.     

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.     

Plantigrady and foot adaptation in African apes: implications for hominid origins.

In living primates, except the great apes and humans, the foot is placed in a heel-elevated or semi-plantigrade position when these animals move upon arboreal or terrestrial substrates. Heel placement and bone positions in the non-great ape primate foot are designed to increase mobility and flexibility in the arboreal environment. Orangutans have further enhanced foot mobility by adapting their feet for suspension and thus similarly utilize foot positions where the heel does not touch the substrate. Chimpanzees and gorillas represent an alternative pattern (plantigrady), in which the heel contacts the surface of the support at the end of swing phase, especially during terrestrial locomotion. Thus, chimpanzees and gorillas possess feet adapted for both arboreal and terrestrial substrates. African apes also share several osteological features related to plantigrady and terrestrial locomotion with early hominids. From this analysis, it is apparent that hominid locomotor evolution passed through a quadrupedal terrestrial phase.     

The role of taste in food selection by African apes: implications for niche separation and overlap in tropical forests

Ripe fruit eating shapes the behavior of most of the apes. Gorillas (Gorilla gorilla) and chimpanzees (Pan troglodytes) are very different sizes and, consequently, have been traditionally viewed as ecologically distinct, but few studies have explored the behavioral and physiological foundations of their diets. Debate continues on the extent that large-bodied gorillas may be less selective and more opportunistic fruit eaters than chimpanzees. Taste responses have been predicted to relate to body size and digestive strategies. This study employs laboratory research on taste perception and discrimination among captive zoo-housed chimpanzees and relates it to previous work on gorillas to better characterize diets and niche separation among these apes. During the captive trials, differences were recorded in consumption patterns of water and varying concentrations of dilute aqueous fructose (sweet) and tannic acid solutions (astringent), compounds commonly found in wild foods. The chimpanzees exhibited similar preference thresholds for fructose (50 mM) to other primates studied. They exhibited slightly lower inhibition thresholds for tannic acid solutions than gorillas, but higher than smaller primates studied to date. These preliminary findings suggest that tannin tolerance may well be mediated by body size, though possible species differences in salivary proteins or other sensory differences remain to be explored. This research furthers our efforts to understand the roles of body size and physiological adaptations in shaping diet and niche separation of chimpanzees and gorillas.     

Genetic sex identification in orangutans

To date, no established protocol for genetic sex identification in orangutans (Pongo pygmaeus) exists. In nearly all apes (gibbons, gorillas, chimpanzees, and humans), genetic sex identification is possible using the amelogenin gene because copies located on X and Y chromosomes have different sizes. Here we report that orangutan sex identification can be resolved through multiplex polymerase chain reaction (PCR) of the Y-linked SRY locus and the amelogenin locus. PCR amplifications of orangutan amelogenin produces one fragment size in both sexes, while SRY amplifies only in males. This protocol will allow primatologists to identify the sex of orangutans through genetic analysis.     

Socioecological factors influencing population structure of gorillas and chimpanzees

Differences in distribution and density between gorillas and chimpanzees are reconsidered with special reference to population structure. Both ecological and social factors influencing population structure are compared between species and between habitats within species. Gorillas and chimpanzees respond differently to a decline in food quality, such as fruit scarcity: gorillas change diet and decrease range, while chimpanzees do not change diet but may expand range. These responses result in different effects on their grouping patterns. For gorillas the dispersed distribution and reduction of range size decreases the rate of inter-unit encounters and female transfer. The concentration of social units increases the rate of aggressive contact between units and stimulates female transfer. Social units of gorillas may crowd or disperse in order to attain the optimal density. This tendency may result in similar densities of gorillas across habitats. By contrast, the distribution patterns or range size may not affect inter-unit relationships of chimpanzees. Within a single unit-group, various reproductive strategies are adopted by both sexes. Independent travel of females and flexible grouping patterns enable them to survive at very low density in extraordinary large ranges. Density and inter-unit relationships are good criteria for a healthy population of gorillas, while the size of unit-group and inter-individual relationships are good criteria for chimpanzees. Conservation planners should consider these differences for sympatric and allopatric survival in these species.     

Renin angiotensin aldosterone axis, including aldosterone binding globulin and blood pressure in three species of nonhuman primates

Variables of renin-angiotensin-aldosterone axis with inclusion of protein binding to specific plasma globulin (ABG), plasma cortisol, and the blood pressure (BP) were measured in 24 chimpanzees, 4 gorillas, and 16 cynomolgus monkeys. ABG activity was readily detected in plasma from the primates. In chimpanzees and gorillas, all the variables under baseline conditions were similar to those in humans. In cynomolgus (Macaca fascicularis), both the ABG binding capacity for aldosterone and the diastolic or systolic BP were significantly higher (p less than 0.001 and p less than 0.01 respectively) than in chimpanzees and gorillas.     

Programming of offspring sex ratios by maternal stress in humans: assessment of physiological mechanisms using a comparative approach

Sex ratio adjustment has become a hot topic in ecology and evolutionary biology, as documentations of sex ratio skews are numerous, and include examples in diverse animal species. Over the past several decades, scientists have repeatedly debated whether human sex ratios also significantly deviate toward one sex or the other based on environmental or social conditions. An increasing number of studies supports the idea that exposure to stressful conditions can influence the sexes of offspring produced by humans, a majority of which document significantly fewer males after exposure to adverse conditions such as severe life events, economic disruption, or natural disasters. From a comparative standpoint, these findings are similar to studies in non-human mammals and other vertebrate species showing a bias toward females during times of stress. However, the mechanisms by which stress-related biases in the offspring sex ratio may occur remain elusive, and the involvement of glucocorticoids indicating a true influence of stress itself remains unstudied. Here, I review the evidence that stressful events induce sex ratio adjustment in humans. Additionally, I discuss the possibility for glucocorticoid mediation of sex ratio adjustment and the potential reproductive stages during which stress-induced sex ratio adjustment may occur in humans and other mammals.     

The human SDF1 gene polymorphism is located on a mutational hot spot that was identified by the hominoid genome study.

Nucleotide sequences of a part of the stromal cell-derived factor-1 (SDF-1) gene 3' untranslated region were studied among hominoids (chimpanzees, gorillas, orangutans and gibbons). An identical sequence to the human SDF1-3'G allele was found in chimpanzees and gibbons, whereas that to the 3'A allele was found in gorillas. Based on the sequence data and the hominoid phylogenetic relation, it was suggested that an adenine nucleotide at nucleotide position (np) 801 in humans and gorillas was independently introduced into each lineage after the specific divergence and an ancestral hominoid sequence of this site (np 799-802) was deduced as CCGG. The present data showing a mutational hot spot on this site suggest the possible presence of multiple origins of the worldwide distribution of the SDF1-3'A allele in humans.     

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.     

Rh gene evolution in primates: study of intron sequences

By amplification and sequencing of RH gene intron 4 of various primates we demonstrate that an Alu-Sx-like element has been inserted in the RH gene of the common ancestor of humans, apes, Old World monkeys, and New World monkeys. The study of mouse and lemur intron 4 sequences allowed us to precisely define the insertion point of the Alu-Sx element in intron 4 of the RH gene ancestor common to Anthropoidea. Like humans, chimpanzees and gorillas possess two types of RH intron 4, characterized by the presence (human RHCE and ape RHCE-like genes) or absence (human RHD and ape RHD-like genes) of the Alu-Sx element. This led us to conclude that in the RH common ancestor of humans, chimpanzees, and gorillas, a duplication of the common ancestor gene gave rise to two genes, one differing from the other by a 654-bp deletion encompassing an Alu-Sx element. Moreover, most of chimpanzees and some gorillas posses two types of RHD-like intron 4. The introns 4 of type 1 have a length similar to that of human RHD intron 4, whereas introns 4 of type 2 display an insertion of 12 bp. The latest insertion was not found in the human genome (72 individuals tested). The study of RH intron 3 length polymorphism confirmed that, like humans, chimpanzees and gorillas possess two types of intron 3, with the RHD-type intron 3 being 289 bases shorter than the RHCE intron 3. By amplification and sequencing of regions encompassing introns 3 and 4, we demonstrated that chimpanzee and gorilla RH-like genes displayed associations of introns 3 and 4 distinct to those found in man. Altogether, the results demonstrate that, as in humans, chimpanzee and gorilla RH genes experienced intergenic exchanges.     

Corpus callosum morphology in capuchin monkeys is influenced by sex and handedness

Sex differences have been reported in both overall corpus callosum area and its regional subdivisions in humans. Some have suggested this reflects a unique adaptation in humans, as similar sex differences in corpus callosum morphology have not been reported in any other species of primate examined to date. Furthermore, an association between various measurements of corpus callosum morphology and handedness has been found in humans and chimpanzees. In the current study, we report measurements of corpus callosum cross-sectional area from midsagittal MR images collected in vivo from 14 adult capuchin monkeys, 9 of which were also characterized for hand preference on a coordinated bimanual task. Adult females were found to have a significantly larger corpus callosum: brain volume ratio, rostral body, posterior midbody, isthmus, and splenium than adult males. Left-handed individuals had a larger relative overall corpus callosum area than did right-handed individuals. Additionally, a significant sex and handedness interaction was found for anterior midbody, with right-handed males having a significantly smaller area than right-handed females. These results suggest that sex and handedness influences on corpus callosum morphology are not restricted to Homo sapiens.