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The role of the chimpanzee in the evaluation of the risk of foreign chemicals to man.

Various species of laboratory animals are used to evaluate the efficacy and safety of drugs in man. However, the extrapolation of data from animals to man is often complicated by species differences in the disposition of foreign chemicals. The findings of comparative metabolism studies are used to illustrate species differences in metabolic pathways, rates of biotransformation, kinetics, and excretion routes. Biochemical and structural consequences of the significant differences in enzyme induction between rodents and primate species are discussed. Among the primates, the chimpanzee has been shown to be the most closely related to man not only in the disposition of xenobiotics, but also in the aspects of endocrinology, serology, and immunology. It would, therefore, be the best possible model to predict the fate and effects of foreign chemicals in man. Due to the limited availability of chimpanzees, however, they can only be used for the evaluation of the most critical chemicals and drugs. Comparative metabolism and disposition studies, e.g., of compounds representative of classes of chemicals in chimpanzees and other animals. Only when the pertinent differences between the selected test species and man are known can correct extrapolations to man be made.

Animals↗

Ultrasound guided spleen biopsy in chimpanzees (pan troglodytes).

To the best of our knowledge, this is the first published investigation of the size and position of chimpanzee spleens as measured by ultrasound, and that a fine-needle biopsy of the chimpanzee spleen, guided by ultrasound, has been performed. In general, the chimpanzee spleen is smaller than the human spleen (mean length of 8.9 cm, SD 1.17 cm; mean thickness 2.4 cm, SD 0.35 cm; n = 14) and its position may be different due to different numbers of thoracic vertebrae. Fine-needle biopsy is a safe and simple method to obtain lymphoid tissue sufficient for PCR testing of HIV-1.

Animals↗

Variations in the serum esterases of chimpanzees.

Seventy chimpanzees, representing Pan paniscus (the pygmy chimp) and all four races of the common chimpanzee (Pan troglodytes), were examined for electrophoretic variation in their serum esterases. Only three variant electromorphs were observed at these six loci. One of these was at the albumin-associated esterase locus, and the other two were at esterase 4. This is the first report of any variation in an esterase or albumin in chimpanzees.

Animals↗

Forensic study on stains of blood and saliva in a chimpanzee bite case.

A case of a boy bitten by a chimpanzee is reported. Forensic investigations were carried out on his T-shirt which had been torn open at the right armpit. By presumptive tests using chemical reagents and definitive tests using antisera against human hemoglobin A (HbAo), human serum and human saliva amylase, it was demonstrated that stains scattered around the torn portion of the shirt consisted of saliva or both blood and saliva. Though the blood was immunologically assumed to be human, the saliva could not be identified as human or chimpanzee. The mixed stains and the saliva stains were grouped A and secretor. By treatment of the mixed stains with a mixture of two volumes of chloroform and one volume of methanol (CM solution), the water-insoluble blood substance derived from blood was typed as O and the water-soluble substance derived from saliva was typed as A and secretor. The former was the victim's blood type and the latter were the chimpanzee saliva types.

Animals↗

Response to novel food in infant chimpanzees. Do infants refer to mothers before ingesting food on their own?

We investigated infant response toward novel food in captive chimpanzees under the condition in which they can explore such items freely together with their mother. Infants first approached novel foods rather than familiar ones when presented simultaneously. However, they did not ingest novel food immediately, but always sniff-licked it first. Infants tended to pay attention to their mothers before mouthing or ingesting novel foods themselves, but never did so with familiar ones. In response to the infant's activity, mother chimpanzees were tolerant rather than actively interfering. Those results imply that chimpanzee infants respond to novel foods in a neophobic way and refer to their mother for some kind of cue before attempting to ingest them.

Animals↗

Maintenance of delay of gratification by four chimpanzees (Pan troglodytes): the effects of delayed reward visibility, experimenter presence, and extended delay intervals.

Previous research in our laboratory has demonstrated that chimpanzees can delay gratification by inhibiting consumption of available food items for as long as 3 min as an experimenter transfers additional food items from a transparent container to a bowl placed in front of the subject. In this study, we examined the influence of the visibility of the food source, as well as the presence of the experimenter, on four chimpanzees' self-control in this paradigm. In Experiment 1 an experimenter transferred 15 preferred food items between a distant opaque container and a bowl placed in front of the subject. In Experiment 2 we tested the chimpanzees with an automated system that (in the absence of the experimenter) transferred up to 36 highly preferred food items from a universal food dispenser to a container located either inside or outside of the subject's enclosure. There were no differences in self-directed behaviors or attentiveness to the food items between the self-imposed and externally imposed delay conditions. A final experiment with the automated paradigm indicated that individuals could delay gratification for up to 11min in order to obtain all 36 food items.

Animals↗

Locomotor behavior and long bone morphology in individual free-ranging chimpanzees.

We combine structural limb data and behavioral data for free-ranging chimpanzees from Taï (Ivory Coast) and Mahale National Parks (Tanzania) to begin to consider the relationship between individual variation in locomotor activity and morphology. Femoral and humeral cross sections of ten individuals were acquired via computed tomography. Locomotor profiles of seven individuals were constructed from 3387 instantaneous time-point observations (87.4 hours). Within the limited number of suitable chimpanzees, individual variation in locomotor profiles displayed neither clear nor consistent trends with diaphyseal cross-sectional shapes. The percentages of specific locomotor modes did not relate well to diaphyseal shapes since neither infrequent nor frequent locomotor modes varied consistently with shapes. The percentage of arboreal locomotion, rather than estimated body mass, apparently had comparatively greater biological relevance to variation in diaphyseal shape. The mechanical consequences of locomotor modes on femoral and humeral diaphyseal shapes (e.g., orientation of bending strains) may overlap between naturalistic modes more than currently is recognized. Alternatively, diaphyseal shape may be unresponsive to mechanical demands of these specific locomotor modes. More data are needed in order to discern between these possibilities. Increasing the sample to include additional free-ranging chimpanzees, or primates in general, as well as devoting more attention to the mechanics of a greater variety of naturalistic locomotor modes would be fruitful to understanding the behavioral basis of diaphyseal shapes.

Anatomy, Cross-Sectional↗

Descent of the hyoid in chimpanzees: evolution of face flattening and speech.

The human supralaryngeal vocal tract develops to form a unique two-tube configuration with equally long horizontal and vertical cavities. This anatomy contributes greatly to the morphological foundations of human speech. It is believed to depend on the reduced growth of the palate and on the developmental descent of the larynx relative to the palate. Anatomically, the descent of the larynx is accomplished through both the descent of the laryngeal skeleton relative to the hyoid and the descent of the hyoid relative to the palate. We have studied the development of three living chimpanzees using magnetic resonance imaging. Our previous study showed that, as in humans, chimpanzees show rapid laryngeal descent, with changes in the relative proportion of the vocal tract during early infancy. However, this is not accompanied by the descent of the hyoid relative to the palate, although it is achieved with the descent of the laryngeal skeleton relative to the hyoid. Here, we show that subsequently the chimpanzee hyoid also descends to maintain the rapid descent of the larynx, similarly to humans. We argue that the descent of the larynx probably evolved in a common ancestor of extant hominoids, originally to confer an advantage via a function unrelated to speech. Thus, the descent of the larynx per se is not unique to humans, and facial flattening was probably the major factor that paved the way for speech in the human lineage.

Animals↗

Lagrangian frame diffeomorphic image registration: Morphometric comparison of human and chimpanzee cortex.

We develop a novel Lagrangian reference frame diffeomorphic image and landmark registration method. The algorithm uses the fixed Langrangian reference frame to define the map between coordinate systems, but also generates and stores the inverse map from the Eulerian to the Lagrangian frame. Computing both maps allows facile computation of both Eulerian and Langrangian quantities. We apply this algorithm to estimating a putative evolutionary change of coordinates between a population of chimpanzee and human cortices. Inter-species functional homologues fix the map explicitly, where they are known, while image similarities guide the alignment elsewhere. This map allows detailed study of the volumetric change between chimp and human cortex. Instead of basing the inter-species study on a single species atlas, we diffeomorphically connect the mean shape and intensity templates for each group. The human statistics then map diffeomorphically into the space of the chimpanzee cortex providing a comparison between species. The population statistics show a significant doubling of the relative prefrontal lobe size in humans, as compared to chimpanzees.

Algorithms↗

Variation of the prion gene in chimpanzees and its implication for prion diseases.

In humans, familial prion diseases are linked to mutations in the PRNP gene. We have sequenced part of this gene in a large sample of common chimpanzee, Pan troglodytes (n=130 chromosomes). No variation in codons 129 and 219 has been observed: all chimpanzees were homozygous for the Met allele, which in humans increases susceptibility to Creutzfeldt-Jakob disease. We found two sequence variants: one is a synonymous polymorphism unique to the chimpanzee at codon 226, TAC to TAT (Y), with a TAC allele frequency of 80.6%; the other is a non-synonymous change at codon 148 (R148H) that falls in the target epitope for some common commercial antibodies used for prion diagnostics, and is highly conserved across species. The pathogenicity of this mutation is still unknown.

Amino Acid Sequence↗

Further evidence of a right hand advantage in motor skill by chimpanzees (Pan troglodytes).

Asymmetries in motor skill when grasping small food items was evaluated in a sample of captive chimpanzees. In two experiments, error rates in grasping food were assessed when controlling for individual differences in grip morphology. In both experiments, chimpanzees were found to make more errors with the left compared to the right hand. Male chimpanzees were also found to make more errors compared to females. These results are discussed in the context of a presumed disassociation between hand preference and performance as indicators of cerebral dominance in motor functions.

Analysis of Variance↗

Further evidence of an association between handedness and neuroanatomical asymmetries in the primary motor cortex of chimpanzees (Pan troglodytes).

The neurobiology of handedness is still poorly understood in nonhuman primates. Recently, an association between hand preference and precentral gyrus morphology in chimpanzees was reported. The aim of this study was to further evaluate the association between handedness and asymmetries in the precentral gyrus of chimpanzees (Pan troglodytes) and to evaluate the association between hand preference and brain asymmetry using a different approach to the classification of handedness in chimpanzees. The overall results suggest that differences in handedness groups are specific to a region of the precentral gyrus commonly known as the "knob" and that subjects that show different hand preferences differ in brain asymmetries for specific regions of the primary motor cortex. Moreover, using a continuous scale of measurement rather than discrete classification of handedness, significant associations were found between hand use and asymmetries within the precentral gyrus.

Animals↗

Sex differences in the relative lengths of metacarpals and metatarsals in gorillas and chimpanzees.

Sex differences other than the simple dimorphism in size were documented for the metapodials of two primate species. Lengths of metacarpals and metatarsals were obtained from the skeletons of 64 gorillas and 42 chimpanzees. Length ratios were constructed for all possible pairings of the five bones in each individual hand and foot. For both species, several of these length ratios exhibited substantial differences between the sexes. Body size was not the basis for these sex differences; when specimens of similar size were compared, the sex differences remained. In humans, length ratios for the fingers and toes also have previously been demonstrated to exhibit sex differences, and the length ratio for the index and ring fingers (the 2D:4D ratio) has been shown to correlate with various medical conditions. Various facts suggest that length ratios in human digits are associated with androgen exposure, probably during prenatal development. For gorillas, the metacarpal length ratio showing the largest sex difference was 4Mc:5Mc in both hands, and the metatarsal length ratio showing the largest sex difference was 1Mt:2Mt in the left foot. Sex differences in length ratios also existed for chimpanzees, but they were generally smaller than for gorillas. Apparently, both gorillas and chimpanzees are affected by developmental mechanisms, possibly androgenic mechanisms, similar to those in humans. Analyses of previous measurements [Susman, R.L., 1979 Comparative and functional morphology of hominoid fingers. Am. J. Phys. Anthropol. 50, 215-236] revealed that all components of the rays are not affected equally by whatever mechanisms are responsible for the sex differences in length ratios.

Animals↗

Preference for human direct gaze in infant chimpanzees (Pan troglodytes).

We studied gaze perception in three infant chimpanzees (Pan troglodytes), aged 10-32 weeks, using a two-choice preferential-looking paradigm. The infants were presented with two photographs of a human face: (a) with the eyes open or closed, and (b) with a direct or an averted gaze. We found that the chimpanzees preferred looking at the direct-gaze face. However, in the context of scrambled faces, the infants showed no difference in gaze discrimination between direct and averted gazes. These findings suggest that gaze perception by chimpanzees may be influenced by the surrounding facial context. The relationship between gaze perception, face processing, and the adaptive significance of gaze perception are discussed from an evolutionary perspective.

Animals↗

Birth order and left-handedness revisited: some recent findings in chimpanzees (Pan troglodytes) and their implications for developmental and evolutionary models of human handedness.

This paper examines the relationship between parity, pregnancy outcome, and handedness in a sample of captive chimpanzees (Pan troglodytes). The relation between parity, maternal age and positive or negative pregnancy outcome was assessed from life history data for 536 chimpanzees housed at the Yerkes Regional Primate Research Center. The incidences of negative pregnancy outcome (notably spontaneous abortions and stillbirths) were significantly higher in parities of 8 or higher compared to all other parities. In a sub-sample of 165 chimpanzees, the relation between parity, maternal age and handedness was assessed to determine whether left handedness may serve as a marker of prenatal pathology. These analyses indicated that left-handedness was more prevalent in 1st and 8 or higher parities compared to parities between 2 and 7, respectively. Possible prenatal hormonal and periparturitional factors are discussed as possible mechanisms for the observed findings.

Animals↗

Asymmetry in facial expression of emotions by chimpanzees.

Asymmetries in human facial expressions have long been documented and traditionally interpreted as evidence of brain laterality in emotional behavior. Recent findings in nonhuman primates suggest that this hemispheric specialization for emotional behavior may have precursors in primate evolution. In this study, we present the first data collected on our closest living relative, the chimpanzee. Objective measures (hemimouth length and area) and subjective measures (human judgements of chimeric stimuli) indicate that chimpanzees' facial expressions are asymmetric, with a greater involvement of the left side of the face in the production of emotional responses. No effect of expression type (positive versus negative) on facial asymmetry was found. Thus, chimpanzees, like humans and some other nonhuman primates, show a right hemisphere specialization for facial expression of emotions.

Animals↗

An 8-year longitudinal study of mirror self-recognition in chimpanzees (Pan troglodytes).

In a previous cross-sectional study of mirror self-recognition involving 92 chimpanzees, Povinelli et al. [Journal of Comparative Psychology 107 (1993) 347] reported a peak in the proportion of animals exhibiting self-recognition in the adolescent/young adult sample (8-15 years), with 75% being classified as positive. In contrast, only 26% of the older animals (16-39 years) were classified as positive, suggesting a marked decline in self-recognition in middle to late adulthood. In the present study, all of the chimpanzees from the 8-15-year-old group in the Povinelli et al. study (n=12) were again tested for self-recognition, 8 years later. Using the same criteria, 67% of the animals were classified the same. Although a higher proportion of the adult animals in this study (50%) exhibited self-recognition than would be inferred on the basis of the previous study (25%), all changes in self-recognition status were in the negative direction. These results show that mirror self-recognition is a highly stable trait in many chimpanzees, but may be subject to decline with age. Connections with human research are briefly discussed.

Aging↗

Brain temperature asymmetries and emotional perception in chimpanzees, Pan troglodytes.

The lateralization of emotion has received a great deal of attention over the last few decades, resulting in two main theories. The Right Hemisphere Theory states that the right hemisphere is primarily responsible for emotional processes, while the Valence Theory suggests that the right hemisphere regulates negative emotion and the left hemisphere regulates positive emotion. Despite the important implications of these theories for the evolution of emotion processes, few studies have attempted to assess the lateralization of emotion in non-human primates. This study uses the novel technique of measuring tympanic membrane temperature (Tty) to assess asymmetries in the perception of emotional stimuli in chimpanzees. The tympanic membrane is an indirect, but reliable, site from which to measure brain temperature, and is strongly influenced by autonomic and behavioral activity. Six chimpanzees were shown positive, neutral, and negative emotional videos depicting scenes of play, scenery, and severe aggression, respectively. During the negative emotion condition, right Tty was significantly higher than the baseline temperature. This effect was relatively stable, long lasting, and consistent across subjects. Temperatures did not change significantly from baseline in the neutral or positive emotion condition, although a significant number of measurements showed increased left Tty during the neutral emotion condition. These data suggest that viewing emotional stimuli results in asymmetrical changes in brain temperature, in particular increased right Tty during the negative emotion condition, evidence of emotional arousal in chimpanzees, and in providing partial support of both the Right Hemisphere and Valence Theories of emotional lateralization in our closest living ancestor.

Animals↗