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.     

Aging and sex affect soluble alpha klotho levels in bonobos and chimpanzees

Background

Throughout life, physiological homeostasis is challenged and the capacity to cope with such challenges declines with increasing age. In many species, sex differences exist in life expectancy. Sex-specific differences have been related to extrinsic factors like mate competition and/or intrinsic proximate mechanisms such as hormonal changes. In humans, an intrinsic factor related to aging is soluble alpha klotho (α-Kl). Both sexes show an age-related decline in α-Kl, but throughout life women have higher levels than men of the same age. Sex differences in α-Kl have been linked to a shorter lifespan, as well as to specific morbidity factors such as atherosclerosis and arteries calcifications. In non-human animals, information on α-Kl levels is rare and restricted to experimental work. Our cross-sectional study is the first on α-Kl levels in two long-lived species: bonobos (Pan paniscus) and chimpanzees (Pan troglodytes). As in most mammals, female bonobos and chimpanzees have longer life expectancy than males.

Methods

We measured serum α-Kl levels of 140 subjects from 16 zoos with an ELISA to examine if α-Kl levels reflect this difference in life expectancy.

Results

In both species and in both sexes, α-Kl levels declined with age suggesting that this marker has potential for aging studies beyond humans. We also found species-specific differences. Adult female bonobos had higher α-Kl levels than males, a difference that corresponds to the pattern found in humans. In chimpanzees, we found the opposite: males had higher α-Kl levels than females.

Conclusion

We suggest that contrasting sex differences in adult α-Kl levels mirror the dominance relations between females and males of the two Pan species; and that this might be related to corresponding sex differences in their exposure to stress. In humans, higher cortisol levels were found to be related to lower α-Kl levels. We conclude that there is great potential for studying aging processes in hominoids, and perhaps also in other non-human primates, by measuring α-Kl levels. To better understand the causes for sex differences in this aging marker, consideration of behavioural parameters such as competition and stress exposure will be required as well as other physiological markers.

     

Aging and fertility patterns in wild chimpanzees provide insights into the evolution of menopause

Human menopause is remarkable in that reproductive senescence is markedly accelerated relative to somatic aging, leaving an extended postreproductive period for a large proportion of women. Functional explanations for this are debated, in part because comparative data from closely related species are inadequate. Existing studies of chimpanzees are based on very small samples and have not provided clear conclusions about the reproductive function of aging females. These studies have not examined whether reproductive senescence in chimpanzees exceeds the pace of general aging, as in humans, or occurs in parallel with declines in overall health, as in many other animals. In order to remedy these problems, we examined fertility and mortality patterns in six free-living chimpanzee populations. Chimpanzee and human birth rates show similar patterns of decline beginning in the fourth decade, suggesting that the physiology of reproductive senescence was relatively conserved in human evolution. However, in contrast to humans, chimpanzee fertility declines are consistent with declines in survivorship, and healthy females maintain high birth rates late into life. Thus, in contrast to recent claims, we find no evidence that menopause is a typical characteristic of chimpanzee life histories.     

Evolution of protamine P1 genes in primates

Protamine P1 genes have been sequenced by PCR amplification and direct DNA sequencing from 9 primates representing 5 major families, Cebidae (new world monkeys), Cercopithecidae (old world monkeys), Hylobatidae (gibbons), Pongidae (gorilla, orangutan, and chimpanzee), and Hominidae (human). In this recently diverged group of primates these genes are clearly orthologous but very variable, both at the DNA level and in their expressed amino acid sequences. The rate of variation amongst the protamine Pls indicates that they are amongst the most rapidly diverging polypeptides studied. However, some regions are conserved both in primates and generally in other placental mammals. These are the 13 N-terminal residues (including a region of alternating serine and arginine residues (the motif SRSR, res. 10–13) susceptible to Ser phosphorylation), a tract of six Arg residues (res. 24–29) in the center of the molecule, and a six-residue region (RCCRRR, res. 39–44), consisting of a pair of cysteines flanked by arginines. Detailed consideration of nearest neighbor matrices and trees based on maximum parsimony indicates that PI genes from humans, gorillas, and chimpanzees are very similar. The amino acid and nucleotide differences between humans and gorillas. are fewer than those between humans and chimpanzees. This finding is at variance with data from DNA-DNA hybridization and extensive globin and mitochondrial DNA sequences which place human and chimpanzee as closest relatives in the super family, Hominoidea. This may be related to the fact that protamine Pls are expressed in germ line rather than somatic cells. In contrast to the variability of the exon regions of the protamine P1 genes, the sequence of the single intron is highly conserved.     

Brain Evolution: Mammals,Primates, Chimpanzees,and Humans

Though many modern techniques are available for studying brains, they are difficult to use in evolutionary contexts that require examination of large numbers of specimens and species, and all major parts of the brain. Thus, evolutionary studies of many species and of whole brains still tend to be based upon simpler data such as sizes of brains and brain components. Such investigations, carried out over many decades, have usually employed univariate and bivariate analyses, though a few investigators used early multivariate methods. In mammals, these studies generally show the primacy of the relationship between brain and brain-part sizes with overall body size. More recent multivariate applications have confirmed this (Finlay, B. L., and Darlington, R. B. (1995). Science 268: 1578–1584) and some have also separated the highest level phylogenetic groups: strepsirrhines and haplorrhines (Barton, R. A., and Harvey, P. H. (2000). Nature 405: 1055–1058). Both findings were, in fact, evident in earlier multivariate studies (Holloway, R. L. (1979). In Hahn, M. E., Jensen C., and Dudek, B. C. (eds.), Development and Evolution of Brain Size: Behavioral Implications, Academic Press, New York, pp. 59–88; Sacher, G. A. (1970). In Noback, C. R., and Montagna, W. (eds.), The Primate Brain: Advances in Primatology. Vol. 1, Appleton-Century-Crofts, Educational Division, Meredith Corporation, New York, pp. 245–287). However, new studies employing proportional data aimed at conveying input/output relationships between brain components show further groupings of species that share convergences in lifestyles (de Winter, W., and Oxnard, C. E. (2001). Nature 409: 710–714). The convergences are brought about by combinations of brain variables that seem to be associated with brain functions implied by the specific lifestyles. Our most recent results demonstrate that chimpanzees and humans are especially different from one another, and the difference is not due to size alone. Part of this difference is merely a continuation, from chimpanzees towards humans, of a trend already present across all other primates that relates mainly to neocortical increase. But several other large and independent differences are not in the direction of the overall primate trend, but are differences of humans from all other mammals examined including all nonhuman primates. The combinations of brain variables associated with the latter differences are not related simply to enhancement of the neocortex, but seem to reflect other internal relationships. The overall separation of humans and chimpanzees is so large that it goes far beyond the conventional 98.6% commonality in their DNAs. It fits better with more recent molecular, developmental and evolutionary studies implying a considerably greater difference between chimpanzees and humans than usually recognized.     

Amino acids in the plasma of fasting Japanese monkeys (Macaca fuscata fuscata andM. f. yakui)

The plasma levels of 26 amino acids and related compounds were determined in five male and five female adult members of each of the two subspecies of Japanese monkeys,Macaca fuscata fuscata andM. f. yakui. Activities of L-asparaginase and histaminase in plasma were also measured.Numerous differences in amino acid levels between the sexes in the subspeciesfuscata were noted, with the female consistently exhibiting lower values. Few differences were observed between the sexes of the subspeciesyakui or between the two subspecies of Japanese monkeys. These monkeys were similar to other previously studied nonhuman primates in exhibiting measurable levels of 3-methylhistidine in plasma. There were numerous quantitative differences among Japanese monkeys and stump-tailed macaques, rhesus monkeys, chimpanzees, and man, with the Japanese monkeys usually exhibiting higher plasma levels.L-asparaginase activity was not detectable in these Japanese monkeys. Histaminase activity was similar to that previously measured in pig-tailed macaques and chimpanzees, lower than that in rhesus monkeys and stump-tailed macaques, and higher than that in man.     

Chimpanzee variation facilitates the interpretation of the incisive suture closure in South African Plio-Pleistocene hominids

For a better understanding of early hominid growth patterns, we need to compare skeletal maturation among humans and chimpanzees. This study provides new data on variation of the incisive suture closure in extant species to facilitate the understanding of growth patterns among South African Plio-Pleistocene hominids. The complete anterior closure of the incisive suture occurs early during human life, mostly before birth. In contrast, in chimpanzees a complete anterior closure occurs mostly after the eruption of either the first permanent molars (pygmy chimpanzees) or the third molars (common chimpanzees). The first aim of this study is to test whether the patterns of closure of both the anterior and palatal components of the incisive suture in chimpanzees accurately mirror their polytypism by investigating 720 museum specimens of known geographical origin. Then we use the data gleaned from the incisive suture closure in chimpanzees to determine whether there are different growth patterns among South African Plio-Pleistocene hominids and to interpret them. Results about the pattern of incisive suture closure are consistent with the differences among chimpanzees as revealed by molecular data. Thus, the variation in chimpanzee patterns of incisive suture closure facilitates the interpretation of morphology in South African fossil hominids. In Australopithecus (Paranthropus) robustus as compared to Australopithecus africanus, the complete anterior closure and, probably, the complete palatal closure of the incisive suture occurs during early life in the same way as they occur in humans. Moreover, the closure pattern observed on Stw 53, a supposed early Homo from Sterkfontein Member 5, is similar to that seen in A. africanus and in chimpanzees. Thus, with respect to the anterior component of the incisive suture, A. africanus and Stw 53 retain the primitive feature for which A. (P.) robustus and Homo share the derived character state. Finally, it is worth noting that the Taung child does not show the robust condition. Am J Phys Anthropol 105:121–135, 1998. © 1998 Wiley-Liss, Inc.     

Comparative rates of violence in chimpanzees and humans

This paper tests the proposal that chimpanzees (Pan troglodytes) and humans have similar rates of death from intraspecific aggression, whereas chimpanzees have higher rates of non-lethal physical attack (Boehm 1999, Hierarchy in the forest: the evolution of egalitarian behavior. Harvard University Press). First, we assembled data on lethal aggression from long-term studies of nine communities of chimpanzees living in five populations. We calculated rates of death from intraspecific aggression both within and between communities. Variation among communities in mortality rates from aggression was high, and rates of death from intercommunity and intracommunity aggression were not correlated. Estimates for average rates of lethal violence for chimpanzees proved to be similar to average rates for subsistence societies of hunter–gatherers and farmers. Second, we compared rates of non-lethal physical aggression for two populations of chimpanzees and one population of recently settled hunter–gatherers. Chimpanzees had rates of aggression between two and three orders of magnitude higher than humans. These preliminary data support Boehms hypothesis.     

Human-like adrenal development in wild chimpanzees: A longitudinal study of urinary dehydroepiandrosterone-sulfate and cortisol

The development of the adrenal cortex varies considerably across primates, being most conspicuous in humans, where a functional zona reticularis–the site of dehydroepiandrosterone-sulfate (DHEA/S) production–does not develop until middle childhood (5–8 years). Prior reports suggest that a human-like adrenarche, associated with a sharp prepubertal increase in DHEA/S, may only occur in the genus Pan. However, the timing and variability in adrenarche in chimpanzees remain poorly described, owing to the lack of longitudinal data, or data from wild populations. Here, we use urine samples from East African chimpanzees (Pan troglodytes schweinfurthii) collected over 20 years at Kanyawara in Kibale National Park, Uganda, to trace the developmental trajectories of DHEAS (n = 1,385 samples, 53 individuals) and cortisol (n = 12,726 samples, 68 individuals). We used generalized additive models (GAM) to investigate the relationship between age, sex, and hormone levels. Adrenarche began earlier in chimpanzees (~2–3 years) compared with what has been reported in humans (6–8 years) and, unlike humans, male and female chimpanzees did not differ significantly in the timing of adrenarche nor in DHEAS concentrations overall. Similar to what has been reported in humans, cortisol production decreased through early life, reaching a nadir around puberty (8–11 years), and a sex difference emerged with males exhibiting higher urinary cortisol levels compared with females by early adulthood (15–16 years). Our study establishes that wild chimpanzees exhibit a human-like pattern of cortisol production during development and corroborates prior reports from captive chimpanzees of a human-like adrenarche, accompanied by significant developmental increases in DHEAS. While the role of these developmental hormone shifts are as yet unclear, they have been implicated in stages of rapid behavioral development once thought unique to humans, especially in regard to explaining the divergence of female and male social behavior before pubertal increases in gonadal hormones.     

Lymphocyte maintenance during healthy aging requires no substantial alterations in cellular turnover

In healthy humans, lymphocyte populations are maintained at a relatively constant size throughout life, reflecting a balance between lymphocyte production and loss. Given the profound immunological changes that occur during healthy aging, including a significant decline in T‐cell production by the thymus, lymphocyte maintenance in the elderly is generally thought to require homeostatic alterations in lymphocyte dynamics. Surprisingly, using in vivo ²H₂O labeling, we find similar dynamics of most lymphocyte subsets between young adult and elderly healthy individuals. As the contribution of thymic output to T‐cell production is only minor from young adulthood onward, compensatory increases in peripheral T‐cell division rates are not required to maintain the T‐cell pool, despite a tenfold decline in thymic output. These fundamental insights will aid the interpretation of further research into aging and clinical conditions related to disturbed lymphocyte dynamics.     

Diet and seasonal changes in sympatric gorillas and chimpanzees at Kahuzi–Biega National Park

Based on 8 years of observations of a group of western lowland gorillas (Gorilla beringei graueri) and a unit-group of chimpanzees (Pan troglodytes schweinfurthii) living sympatrically in the montane forest at Kahuzi–Biega National Park, we compared their diet and analyzed dietary overlap between them in relation to fruit phenology. Data on fruit consumption were collected mainly from fecal samples, and phenology of preferred ape fruits was estimated by monitoring. Totals of 231 plant foods (116 species) and 137 plant foods (104 species) were recorded for gorillas and chimpanzees, respectively. Among these, 38% of gorilla foods and 64% of chimpanzee foods were eaten by both apes. Fruits accounted for the largest overlap between them (77% for gorillas and 59% for chimpanzees). Gorillas consumed more species of vegetative foods (especially bark) exclusively whereas chimpanzees consumed more species of fruits and animal foods exclusively. Although the number of fruit species available in the montane forest of Kahuzi is much lower than that in lowland forest, the number of fruit species per chimpanzee fecal sample (average 2.7 species) was similar to that for chimpanzees in the lowland habitats. By contrast, the number of fruit species per gorilla fecal sample (average 0.8 species) was much lower than that for gorillas in the lowland habitats. Fruit consumption by both apes tended to increase during the dry season when ripe fruits were more abundant in their habitat. However, the number of fruit species consumed by chimpanzees did not change according to ripe fruit abundance. The species differences in fruit consumption may be attributed to the wide ranging of gorillas and repeated usage of a small range by chimpanzees and/or to avoidance of inter-specific contact by chimpanzees. The different staple foods (leaves and bark for gorillas and fig fruits for chimpanzees) characterize the dietary divergence between them in the montane forest of Kahuzi, where fruit is usually scarce. Gorillas rarely fed on insects, but chimpanzees occasionally fed on bees with honey, which possibly compensate for fruit scarcity. A comparison of dietary overlap between gorillas and chimpanzees across habitats suggests that sympatry may not influence dietary overlap in fruit consumed but may stimulate behavioral divergence to reduce feeding competition between them.     

Selenium-dependent Glutathione Peroxidase Activity is Increased in Healthy Post-menopausal Women

Selenium helps protect against peroxidation during aging as part of the glutathione peroxidase (GPx) antioxidant system. Selenium status, however, is often low in elderly persons who have low selenium intake, live in institutions, and have certain chronic diseases. In addition, a relationship has been observed between the female reproductive hormone, estrogen, and selenium status, with blood selenium and GPx activity coinciding with fluctuations in estrogen during the menstrual cycle. These findings suggest that the decrease in estrogen following menopause may cause a decrease in selenium status, and thus accelerate the process of aging and increase the risk of certain diseases. The current study compared selenium status in healthy premenopausal (n = 13, 21 to 43 years) and postmenopausal (n = 10, 57 to 86 years) women. Selenium intakes of both groups were similar and greater than the recommended dietary allowance (RDA) of 55 μg/day for adult women. Although neither plasma nor RBC selenium concentrations were significantly different between groups, postmenopausal women had significantly greater plasma (p < 0.02), and RBC (p < 0.05) GPx activities compared to premenopausal women possibly in response to oxidative processes associated with aging. These results indicate that the selenium status of healthy postmenopausal women did not decline with menopause and that their antioxidant capability, as measured by GPx activity, was preserved with dietary intake of selenium greater than the RDA. Presented in part at the Experimental Biology 2000, April 2000, San Diego, CA [Smith AM, Ha EJ, Medeiros LC. Selenium-dependent glutathione peroxidase activity is increased in healthy post-menopausal women. FASEB J 2000;14:A513.].     

Brief Communication: Quantitative‐ and molecular‐genetic differentiation in humans and chimpanzees: Implications for the evolutionary processes underlying cranial diversification

Estimates of the amount of genetic differentiation in humans among major geographic regions (e.g., Eastern Asia vs. Europe) from quantitative‐genetic analyses of cranial measurements closely match those from classical‐ and molecular‐genetic markers. Typically, among‐region differences account for ～10% of the total variation. This correspondence is generally interpreted as evidence for the importance of neutral evolutionary processes (e.g., genetic drift) in generating among‐region differences in human cranial form, but it was initially surprising because human cranial diversity was frequently assumed to show a strong signature of natural selection. Is the human degree of similarity of cranial and DNA‐sequence estimates of among‐region genetic differentiation unusual? How do comparisons with other taxa illuminate the evolutionary processes underlying cranial diversification? Chimpanzees provide a useful starting point for placing the human results in a broader comparative context, because common chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) are the extant species most closely related to humans. To address these questions, I used 27 cranial measurements collected on a sample of 861 humans and 263 chimpanzees to estimate the amount of genetic differentiation between pairs of groups (between regions for humans and between species or subspecies for chimpanzees). Consistent with previous results, the human cranial estimates are quite similar to published DNA‐sequence estimates. In contrast, the chimpanzee cranial estimates are much smaller than published DNA‐sequence estimates. It appears that cranial differentiation has been limited in chimpanzees relative to humans. Am J Phys Anthropol 154:615–620, 2014. © 2014 Wiley Periodicals, Inc.     

Reproductive Parameters of Female<Emphasis Type="Italic">Pan paniscus</Emphasis> and <Emphasis Type="Italic">P. troglodytes</Emphasis>: Quality versus Quantity

We investigated intra- and interspecific differences in life history and reproductive parameters in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes). We compare the parameters of wild and captive females in order to shed light on the influence of habitat or specific differences or both on reproduction. We present new and additional information on reproductive parameters from captive bonobos and chimpanzees. Captive chimpanzees birth more live offspring and have a shorter interbirth interval, but experience higher infant mortality than captive bonobos. Although captive bonobo females tend to start reproduction at a younger age than chimpanzees, this is effectively only so for wild-born females of both species. Ultimately both species reach the same rate of production of offspring surviving to 5 yr. These results contrast with data from the wild. Wild bonobos tend to have higher reproductive success, a higher fertility rate and a shorter interbirth interval than wild chimpanzees. Reproduction is similar for wild and captive bonobos, which suggests that they are producing at their maximum under both conditions. Overall captive chimpanzees perform better than their wild conspecifics, probably because of lower feeding competition. Infant survival is the only specific difference not affected by captivity. Bonobo infants survive better, which suggests that chimpanzee infants are more at risk. We argue that the interspecific variation in reproductive parameters in captivity is related to the different influence of captivity on reproduction and different pressures of external sources of infant and juvenile mortality.     

The critical period hypothesis of estrogen effects on cognition: Insights from basic research

Background

In addition to its primary role in reproduction estrogen impacts brain areas important for cognition, including the hippocampus and prefrontal cortex. It has been hypothesized that decline in estrogen levels in women following menopause is associated with, or can exacerbate, age-related cognitive decline. However, clinical evidence to support a role for estrogen in preventing cognitive decline in women as they age is equivocal. The critical period hypothesis of estrogen effects on cognition, which proposes that estrogen administration has to be initiated within a critical time period following the loss of ovarian function in order for it to exert positive effects on the central nervous system, is offered as one explanation for inconsistencies across studies.

Scope of review

This review details results from basic research using rodent models investigating the effects of estrogen on cognition in the aging female. Emphasis is placed on work investigating effects of timing of initiation of estrogen administration on its subsequent efficacy.

Major conclusions

Results of basic research provide support for the critical period hypothesis. Furthermore, results of work in rodent models suggest mechanisms by which the response to estrogen is altered if treatment is initiated following long-term ovarian hormone deprivation.

General significance

Understanding if and under what conditions hormone administration following the loss of ovarian function positively affects the brain and behavior could have important implications with regard to female cognitive aging. Results of basic research can contribute to this understanding and provide insight into the complex mechanisms by which estrogen affects cognition.     

Muscle architecture of the common chimpanzee (Pan troglodytes): perspectives for investigating chimpanzee behavior

Thorpe et al. (Am J Phys Anthropol 110:179–199, 1999) quantified chimpanzee (Pan troglodytes) muscle architecture and joint moment arms to determine whether they functionally compensated for structural differences between chimpanzees and humans. They observed enough distinction to conclude that musculoskeletal properties were not compensatory and suggested that chimpanzees and humans do not exhibit dynamically similar movements. These investigators based their assessment on unilateral limb musculatures from three male chimpanzees, of which they called one non-adult representative. Factors such as age, sex, and behavioral lateralization may be responsible for variation in chimpanzee muscle architecture, but this is presently unknown. While the full extent of variation in chimpanzee muscle architecture due to such factors cannot be evaluated with data presently available, the present study expands the chimpanzee dataset and provides a preliminary glimpse of the potential relevance of these factors. Thirty-seven forelimb and 36 hind limb muscles were assessed in two chimpanzee cadavers: one unilaterally (right limbs), and one bilaterally. Mass, fiber length, and physiological cross-sectional area (PCSA) are reported for individual muscles and muscle groups. The musculature of an adult female is more similar in architectural patterns to a young male chimpanzee than to humans, particularly when comparing muscle groups. Age- and sex-related intraspecific differences do not obscure chimpanzee-human interspecific differences. Side asymmetry in one chimpanzee, despite consistent forelimb directional asymmetry, also does not exceed the magnitude of chimpanzee-human differences. Left forelimb muscles, on average, usually had higher masses and longer fiber lengths than right, while right forelimb muscles, on average, usually had greater PCSAs than left. Most muscle groups from the left forelimb exhibited greater masses than right groups, but group asymmetry was significant only for the manual digital muscles. The hind limb exhibited less asymmetry than the forelimb in most comparisons. Examination of additional chimpanzees would clarify the full range of inter- and intra-individual variation.     

Vocal-auditory functions in the chimpanzee: Vowel perception

The perception of vowels was studied in chimpanzees and humans, using a reaction time task in which reaction times for discrimination of vowels were taken as an index of similarity between vowels. Vowels used were five synthetic and natural Japanese vowels and eight natural French vowels. The chimpanzees required long reaction times for discrimination of synthetic [i] from [u] and [e] from [o], that is, they need long latencies for discrimination between vowels based on differences in frequency of the second formant. A similar tendency was observed for discrimination of natural [i] from [u]. The human subject required long reaction times for discrimination between vowels along the first formant axis. These differences can be explained by differences in auditory sensitivity between the two species and the motor theory of speech perception. A vowel, which is pronounced by different speakers, has different acoustic properties. However, humans can perceive these speech sounds as the same vowel. The phenomenon of perceptual constancy in speech perception was studied in chimpanzees using natural vowels and a synthetic [o]- [a] continuum. The chimpanzees ignored the difference in the sex of the speakers and showed a capacity for vocal tract normalization.     

Oxytocin and Vasopressin Receptor Gene Variation as a Proximate Base for Inter- and Intraspecific Behavioral Differences in Bonobos and Chimpanzees

Recent literature has revealed the importance of variation in neuropeptide receptor gene sequences in the regulation of behavioral phenotypic variation. Here we focus on polymorphisms in the oxytocin receptor gene (OXTR) and vasopressin receptor gene 1a (Avpr1a) in chimpanzees and bonobos. In humans, a single nucleotide polymorphism (SNP) in the third intron of OXTR (rs53576 SNP (A/G)) is linked with social behavior, with the risk allele (A) carriers showing reduced levels of empathy and prosociality. Bonobos and chimpanzees differ in these same traits, therefore we hypothesized that these differences might be reflected in variation at the rs53576 position. We sequenced a 320 bp region surrounding rs53576 but found no indications of this SNP in the genus Pan. However, we identified previously unreported SNP variation in the chimpanzee OXTR sequence that differs from both humans and bonobos. Humans and bonobos have previously been shown to have a more similar 5′ promoter region of Avpr1a when compared to chimpanzees, who are polymorphic for the deletion of ∼360 bp in this region (+/− DupB) which includes a microsatellite (RS3). RS3 has been linked with variation in levels of social bonding, potentially explaining part of the interspecies behavioral differences found in bonobos, chimpanzees and humans. To date, results for bonobos have been based on small sample sizes. Our results confirmed that there is no DupB deletion in bonobos with a sample size comprising approximately 90% of the captive founder population, whereas in chimpanzees the deletion of DupB had the highest frequency. Because of the higher frequency of DupB alleles in our bonobo population, we suggest that the presence of this microsatellite may partly reflect documented differences in levels of sociability found in bonobos and chimpanzees.     

Direct somatic embryogenesis fromBegonia gracilis explants

Direct somatic embryogenesis ofBegonia gracilis was achieved from microcultured laminar segments and petioles on Murashige and Skoog medium with 0.5 mg 1^–1 kinetin and 2% coconut water. Somatic embryos were obtained with greater frequency from petiole explants than from leaf blade sections. Under red light (45 mol m^–2 s^–1), approximately 80% of the petiole explants successfully produced somatic embryos but only 30% of the leaf blade sections responded. However, somatic embryos were significantly more abundant on responding lamina explants (60–70 embryos/leaf section) than on petioles (40–50 embryos/petiole). These trends were similar for explants kept in the dark, but overall production was lower. Somatic embryos were produced more quickly (5 weeks) from petioles than from lamina explants (8 weeks). The somatic embryos germinated to produce plantlets and subsequently shoot cultures with the same appearance as the parental clone.Abbreviations BA benzyladenine - MS Murashige and Skoog (1962) - NAA naphthalene acetic acid - SE somatic embryo     

Reproductive aging in captive and wild common chimpanzees: factors influencing the rate of follicular depletion

We examine and discuss evidence of contrasting differences in fertility patterns between captive and wild female chimpanzees, Pan troglodytes, as they age; in the wild females reproduce in their 40s, but captive studies suggest that menopause occurs around that time. Thus, despite the increased longevity generally observed in captive populations reproductive life span is shortened. We outline a hypothesis to explain the apparent differential pace of reproductive decline observed between wild and captive populations. The breeding schedules of captive primates may contribute to accelerated reproductive senescence because continuous cycling in captive animals results in early depletion of the ovarian stock and premature senescence. Available evidence supports the hypothesis that women with patterns of high oocyte loss experience earlier menopause. Chimpanzees in captivity live longer, and thus, similar to humans, they may experience follicular depletion that precedes death by many years. In captivity, chimpanzees typically have an early age at menarche and first birth, shorter interbirth intervals associated with short lactational periods as young mature faster, and nursery rearing, which allows mothers to begin cycling earlier. Variables typical of wild chimpanzee populations, including late age at menarche and first birth, long interbirth intervals associated with prolonged lactational periods, and a long period of female infertility after immigration, spare ovulations and may be responsible for the later age at reproductive termination. Finally, we describe and discuss the timing of specific reproductive landmarks that occur as female chimpanzees age, distinguishing between functional menopause (age at last birth) and operational menopause (end of cycling). Am. J. Primatol. 71:271–282, 2009. © 2008 Wiley‐Liss, Inc.