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Chimpanzees confer benefits on group members, both in the wild and in captive populations. Experimental studies of how animals allocate resources can provide useful insights about the motivations underlying prosocial behavior, and understanding the relationship between task design and prosocial behavior provides an important foundation for future research exploring these animals' social preferences. A number of studies have been designed to assess chimpanzees' preferences for outcomes that benefit others (prosocial preferences), but these studies vary greatly in both the results obtained and the methods used, and in most cases employ procedures that reduce critical features of naturalistic social interactions, such as partner choice. The focus of the current study is on understanding the link between experimental methodology and prosocial behavior in captive chimpanzees, rather than on describing these animals' social motivations themselves. We introduce a task design that avoids isolating subjects and allows them to freely decide whether to participate in the experiment. We explore key elements of the methods utilized in previous experiments in an effort to evaluate two possibilities that have been offered to explain why different experimental designs produce different results: (a) chimpanzees are less likely to deliver food to others when they obtain food for themselves, and (b) evidence of prosociality may be obscured by more "complex" experimental apparatuses (e.g., those including more components or alternative choices). Our results suggest that the complexity of laboratory tasks may generate observed variation in prosocial behavior in laboratory experiments, and highlights the need for more naturalistic research designs while also providing one example of such a paradigm.

In a retrospective analysis of data from three studies using a delayed response task in cynomolgus monkeys, we examined the subjects' search patterns and success rates. Twenty-seven monkeys of both sexes, divided into three age groups, were tasked with retrieving two food items hidden in an array of six identical opaque cups. Although the task was challenging for all subjects, generating a high level of guesswork, evidence of common behaviors when approaching the spatial memory test were found. The search patterns employed by the monkeys suggest the use of landmark cues, adaption in response to failure and chronological memory recall. These strategies appeared to be shared by most subjects, however, the overall success rate appeared to also depend on individual characteristics including age, gender and whether the subject had been born in caged captivity or not. By elucidating some of the underlying cognitive mechanisms, these findings may serve to refine interpretation of future studies using similar delayed response tasks in non-human primates., (Copyright © 2014 International Institute of Anticancer Research (Dr. John G. Delinassios), All rights reserved.)

Despite the importance of individual problem solvers for group- and individual-level fitness, the correlates of individual problem-solving success are still an open topic of investigation. In addition to demographic factors, such as age or sex, certain personality dimensions have also been revealed as reliable correlates of problem-solving by animals. Such correlates, however, have been little-studied in chimpanzees. To empirically test the influence of age, sex, estrous state, and different personality factors on chimpanzee problem-solving, we individually tested 36 captive chimpanzees with two novel foraging puzzles. We included both female (N=24) and male (N=12) adult chimpanzees (aged 14-47 years) in our sample. We also controlled for the females' estrous state-a potential influence on cognitive reasoning-by testing cycling females both when their sexual swelling was maximally tumescent (associated with the luteinizing hormone surge of a female's estrous cycle) and again when it was detumescent. Although we found no correlation between the chimpanzees' success with either puzzle and their age or sex, the chimpanzees' personality ratings did correlate with responses to the novel foraging puzzles. Specifically, male chimpanzees that were rated highly on the factors Methodical, Openness (to experience), and Dominance spent longer interacting with the puzzles. There was also a positive relationship between the latency of females to begin interacting with the two tasks and their rating on the factor Reactivity/Undependability. No other significant correlations were found, but we report tentative evidence for increased problem-solving success by the females when they had detumescent estrous swellings.

Type 2 diabetes can be a problem for captive chimpanzees. Accurate blood glucose (BG) readings are necessary to monitor and treat this disease. Thus, obtaining voluntary samples from primates through positive reinforcement training (PRT) is critical. The current study assessed the voluntary participation of 123 chimpanzees in BG sampling and investigated factors that may contribute to individual success. All subjects participate in regular PRT sessions as part of a comprehensive behavioral management program. Basic steps involved in obtaining BG values include: voluntarily presenting a finger/toe; allowing digit disinfection; holding for the lancet device; and allowing blood collection onto a glucometer test strip for analysis. We recorded the level of participation (none, partial, or complete) when each chimpanzee was first asked to perform the testing procedure. Nearly 30% of subjects allowed the entire procedure in one session, without any prior specific training for the target behavior. Factors that affected this initial successful BG testing included sex, personality (chimpanzees rated higher on the factor "openness" were more likely to participate with BG testing), and past training performance for "present-for-injection" (chimpanzees that presented for their most recent anesthetic injection were more likely to participate). Neither age, rearing history, time since most recent anesthetic event nor social group size significantly affected initial training success. These results have important implications for captive management and training program success, underlining individual differences in training aptitude and the need for developing individual management plans in order to provide optimal care and treatment for diabetic chimpanzees in captivity., (© 2014 Wiley Periodicals, Inc.)

The discernment of resource quality is pertinent to many daily decisions faced by animals. Public information is a critical information source that promotes quality assessments, attained by monitoring others' performance. Here we provide the first evidence, to our knowledge, that chimpanzees (Pan troglodytes) use public information to guide resource selection. Thirty-two chimpanzees were presented with two simultaneous video demonstrations depicting a conspecific acquiring resources at a fast (resource-rich) or slow (resource-poor) rate. Subsequently, subjects selected the resource-rich site above chance expectation. As a comparison, we report evidence of public information use in young children. Investigation of public information use in primates is pertinent, as it can enhance foraging success and potentially facilitate payoff-biased social learning., (©2014 APA, all rights reserved.)

We compared delayed response task performance in young, middle-aged, and old cynomolgus monkeys using three memory tests that have been used with non-human primates. Eighteen cynomolgus monkeys--6 young (4-9 years), 6 middle-aged (10-19 years), and 6 old (above 20 years)--were tested. In general, the old monkeys scored significantly worse than did the animals in the two other age groups. Longer delays between stimulus presentation and response increased the performance differences between the old and younger monkeys. The old monkeys in particular showed signs of impaired visuo-spatial memory and deteriorated memory consolidation and executive functioning. These results add to the body of evidence supporting the utility of Macaca fascicularis in studies of cognition and as a potential translational model for age-associated memory impairment/dementia-related disorders.

Consequences of rearing history in chimpanzees (Pan troglodytes) have been explored in relation to behavioral abnormalities and cognition; however, little is known about the effects of rearing conditions on anatomical brain development. Human studies have revealed that experiences of maltreatment and neglect during infancy and childhood can have detrimental effects on brain development and cognition. In this study, we evaluated the effects of early rearing experience on brain morphology in 92 captive chimpanzees (ages 11-43) who were either reared by their mothers (n = 46) or in a nursery (n = 46) with age-group peers. Magnetic resonance brain images were analyzed with a processing program (BrainVISA) that extracts cortical sulci. We obtained various measurements from 11 sulci located throughout the brain, as well as whole brain gyrification and white and grey matter volumes. We found that mother-reared chimpanzees have greater global white-to-grey matter volume, more cortical folding and thinner grey matter within the cortical folds than nursery-reared animals. The findings reported here are the first to demonstrate that differences in early rearing conditions have significant consequences on brain morphology in chimpanzees and suggests potential differences in the development of white matter expansion and myelination., (© 2013 John Wiley & Sons Ltd.)

Background: Due to their similarities in behavior and disease pathology to humans, non-human primate models are desirable to complement small animals as models for the study of age-related dementia., Materials and Methods: Based on their performance on delayed response task (DRT) tests of memory, aged cynomolgus monkeys were divided into two groups to compare high-performing (n=6) and low-performing (n=6) subjects. Both groups were tested for biomarkers related to Alzheimer's disease and their brains were scanned using structural magnetic resonance imaging., Results: The subjects with poor DRT performance had evidence of atrophy in the hippocampus and cortical areas, significantly lower cerebrospinal fluid levels of amyloid beta amino acid 1-42 (p<0.001) and higher cerebrospinal fluid total tau levels (p<0.05) compared to the group performing well on the DRT tests., Conclusion: Old, memory-impaired Cynomolgus monkeys may be useful as a spontaneous non-human primate model for investigations of age-related neurodegenerative diseases.

Clinical and experimental data have implicated the posterior superior temporal gyrus as an important cortical region in the processing of socially relevant stimuli such as gaze following, eye direction, and head orientation. Gaze following and responding to different socio-communicative signals is an important and highly adaptive skill in primates, including humans. Here, we examined whether individual differences in responding to socio-communicative cues was associated with variation in either gray matter (GM) volume and asymmetry in a sample of chimpanzees. Magnetic resonance image scans and behavioral data on receptive joint attention (RJA) was obtained from a sample of 191 chimpanzees. We found that chimpanzees that performed poorly on the RJA task had less GM in the right compared to left hemisphere in the posterior but not anterior superior temporal gyrus. We further found that middle-aged and elderly chimpanzee performed more poorly on the RJA task and had significantly less GM than young-adult and sub-adult chimpanzees. The results are consistent with previous studies implicating the posterior temporal gyrus in the processing of socially relevant information.

Despite their genetic similarity to humans, our understanding of the role of genes on cognitive traits in chimpanzees remains virtually unexplored. Here, we examined the relationship between genetic variation in the arginine vasopressin V1a receptor gene (AVPR1A) and social cognition in chimpanzees. Studies have shown that chimpanzees are polymorphic for a deletion in a sequence in the 5' flanking region of the AVPR1A, DupB, which contains the variable RS3 repetitive element, which has been associated with variation in social behavior in humans. Results revealed that performance on the social cognition task was significantly heritable. Furthermore, males with one DupB(+) allele performed significantly better and were more responsive to socio-communicative cues than males homozygous for the DupB- deletion. Performance on a non-social cognition task was not associated with the AVPR1A genotype. The collective findings show that AVPR1A polymorphisms are associated with individual differences in performance on a receptive joint attention task in chimpanzees.

In adult humans the prefrontal cortex possesses wider minicolumns and more neuropil space than other cortical regions. These aspects of prefrontal cortex architecture, furthermore, are increased in comparison to chimpanzees and other great apes. In order to determine the developmental appearance of this human cortical specialization, we examined the spatial organization of neurons in four cortical regions (frontal pole [Brodmann's area 10], primary motor [area 4], primary somatosensory [area 3b], and prestriate visual cortex [area 18]) in chimpanzees and humans from birth to approximately the time of adolescence (11 years of age). Horizontal spacing distance (HSD) and gray level ratio (GLR) of layer III neurons were measured in Nissl-stained sections. In both human and chimpanzee area 10, HSD was significantly higher in the postweaning specimens compared to the preweaning ones. No significant age-related differences were seen in the other regions in either species. In concert with other recent studies, the current findings suggest that there is a relatively slower maturation of area 10 in both humans and chimpanzees as compared to other cortical regions, and that further refinement of the spatial organization of neurons within this prefrontal area in humans takes place after the postweaning periods included here., (Copyright © 2013 Wiley Periodicals, Inc.)

There remain considerable questions regarding the evidence for population-level handedness in nonhuman primates when compared with humans. One challenge in comparing human and nonhuman primate handedness involves the procedures used to characterize individual handedness. Studies of human handedness use consistency in hand use within and between tasks as a basis for hand preference classification. In contrast, studies of handedness in nonhuman primates use statistical criteria for classifying handedness. In this study, we examined within- and between-task consistency in hand use as a means of characterizing individual handedness in a sample of 300 captive chimpanzees (Pan troglodytes). Chimpanzees showed population-level right-handedness for both within- and between-tasks consistency, though the proportion of right-handed chimpanzees was lower than what has typically been reported for humans. We further found that there were small, but significant, associations in hand use between measures. There were no significant sex or colony effects on the distribution of handedness. The results are discussed in the context of theories on the evolution of handedness in nonhuman primates.

One effective method for measuring personality in primates is to use personality trait ratings to distill the experience of people familiar with the individual animals. Previous rating instruments were created using either top-down or bottom-up approaches. Top-down approaches, which essentially adapt instruments originally designed for use with another species, can unfortunately lead to the inclusion of traits irrelevant to chimpanzees or fail to include all relevant aspects of chimpanzee personality. Conversely, because bottom-up approaches derive traits specifically for chimpanzees, their unique items may impede comparisons with findings in other studies and other species. To address the limitations of each approach, we developed a new personality rating scale using a combined top-down/bottom-up design. Seventeen raters rated 99 chimpanzees on the new 41-item scale, with all but one item being rated reliably. Principal components analysis, using both varimax and direct oblimin rotations, identified six broad factors. Strong evidence was found for five of the factors (Reactivity/Undependability, Dominance, Openness, Extraversion, and Agreeableness). A sixth factor (Methodical) was offered provisionally until more data are collected. We validated the factors against behavioral data collected independently on the chimpanzees. The five factors demonstrated good evidence for convergent and predictive validity, thereby underscoring the robustness of the factors. Our combined top-down/bottom-up approach provides the most extensive data to date to support the universal existence of these five personality factors in chimpanzees. This framework, which facilitates cross-species comparisons, can also play a vital role in understanding the evolution of personality and can assist with husbandry and welfare efforts., (© 2013 Wiley Periodicals, Inc.)

The primate cerebral cortex is characterized by regional variation in the structure of pyramidal neurons, with more complex dendritic arbors and greater spine density observed in prefrontal compared with sensory and motor cortices. Although there are several investigations in humans and other primates, virtually nothing is known about regional variation in the morphology of pyramidal neurons in the cerebral cortex of great apes, humans' closest living relatives. The current study uses the rapid Golgi stain to quantify the dendritic structure of layer III pyramidal neurons in 4 areas of the chimpanzee cerebral cortex: Primary somatosensory (area 3b), primary motor (area 4), prestriate visual (area 18), and prefrontal (area 10) cortex. Consistent with previous studies in humans and macaque monkeys, pyramidal neurons in the prefrontal cortex of chimpanzees exhibit greater dendritic complexity than those in other cortical regions, suggesting that prefrontal cortical evolution in primates is characterized by increased potential for integrative connectivity. Compared with chimpanzees, the pyramidal neurons of humans had significantly longer and more branched dendritic arbors in all cortical regions.

We provided chimpanzees (Pan troglodytes) with the ability to improve the quality of food rewards they received in a dyadic test of inequity. We were interested to see if this provision influenced their responses and, if so, whether it was mediated by a social partner's outcomes. We tested eight dyads using an exchange paradigm in which, depending on the condition, the chimpanzees were rewarded with either high-value (a grape) or low-value (a piece of celery) food rewards for each completed exchange. We included four conditions. In the first, "Different" condition, the subject received different, less-preferred, rewards than their partner for each exchange made (a test of inequity). In the "Unavailable" condition, high-value rewards were shown, but not given, to both chimpanzees prior to each exchange and the chimpanzees were rewarded equally with low-value rewards (a test of individual contrast). The final two conditions created equity. In these High-value and Low-value "Same" conditions both chimpanzees received the same food rewards for each exchange. Within each condition, the chimpanzees first completed ten trials in the Baseline Phase, in which the experimenter determined the rewards they received, and then ten trials in the Test Phase. In the Test Phase, the chimpanzees could exchange tokens through the aperture of a small wooden picture frame hung on their cage mesh in order to receive the high-value reward. Thus, in the Test Phase, the chimpanzees were provided with an opportunity to improve the quality of the rewards they received, either absolutely or relative to what their partner received. The chimpanzees responded in a targeted manner; in the Test Phase they attempted to maximize their returns in all conditions in which they had received low-value rewards during the Baseline Phase. Thus, the chimpanzees were apparently motivated to increase their reward regardless of their partners', but they only used the mechanism provided when it afforded the opportunity for them to increase their rewards. We also found evidence that the chimpanzees' responses were enhanced by social facilitation. Specifically, the chimpanzees were more likely to exchange their tokens through the frame when their test partner also did so, even in circumstances in which their reward value could not be improved. Our paradigm provided the chimpanzees with the possibility to improve the quality of rewards they received in the Test Phase. We found that refusals - to exchange tokens or to eat rewards - decreased significantly in the Test Phase compared to the Baseline Phase, where no such opportunity for improvement of outcomes existed. Thus, the chimpanzees participated more when they could improve the rewards they received.

Acupuncture is an ancient practice that is currently used to treat disorders ranging from osteoarthritis to cardiomyopathy. Acupuncture involves the insertion of thin, sterile needles into defined acupuncture points that stimulate physiologic processes through neural signaling. Numerous scientific studies have proven the benefits of acupuncture, and given this scientific support, we hypothesized that acupuncture could benefit the nonhuman primates at our facility. As our chimpanzee colony ages, we are observing an increase in osteoarthritis and have focused our initial acupuncture treatments on this condition. We successfully trained 3 chimpanzees, by using positive-reinforcement training techniques, to voluntarily participate in acupuncture treatments for stifle osteoarthritis. We used 3 acupuncture points that correlate with alleviation of stifle pain and inflammation in humans. A mobility scoring system was used to assess improvements in mobility as a function of the acupuncture treatments. The 2 chimpanzees with the most severe osteoarthritis showed significant improvement in mobility after acupuncture treatments. Acupuncture therapy not only resulted in improved mobility, but the training sessions also served as enrichment for the animals, as demonstrated by their voluntary participation in the training and treatment sessions. Acupuncture is an innovative treatment technique that our data show to be safe, inexpensive, and, most importantly, effective for chimpanzees.

Neocortical development in humans is characterized by an extended period of synaptic proliferation that peaks in mid-childhood, with subsequent pruning through early adulthood, as well as relatively delayed maturation of neuronal arborization in the prefrontal cortex compared with sensorimotor areas. In macaque monkeys, cortical synaptogenesis peaks during early infancy and developmental changes in synapse density and dendritic spines occur synchronously across cortical regions. Thus, relatively prolonged synapse and neuronal maturation in humans might contribute to enhancement of social learning during development and transmission of cultural practices, including language. However, because macaques, which share a last common ancestor with humans ≈ 25 million years ago, have served as the predominant comparative primate model in neurodevelopmental research, the paucity of data from more closely related great apes leaves unresolved when these evolutionary changes in the timing of cortical development became established in the human lineage. To address this question, we used immunohistochemistry, electron microscopy, and Golgi staining to characterize synaptic density and dendritic morphology of pyramidal neurons in primary somatosensory (area 3b), primary motor (area 4), prestriate visual (area 18), and prefrontal (area 10) cortices of developing chimpanzees (Pan troglodytes). We found that synaptogenesis occurs synchronously across cortical areas, with a peak of synapse density during the juvenile period (3-5 y). Moreover, similar to findings in humans, dendrites of prefrontal pyramidal neurons developed later than sensorimotor areas. These results suggest that evolutionary changes to neocortical development promoting greater neuronal plasticity early in postnatal life preceded the divergence of the human and chimpanzee lineages.

Background: In an attempt to explore cynomolgus monkeys as an animal model for Alzheimer's disease, the present study focused on the Alzheimer's biomarkers beta amyloid 1-42 (Aβ42 ) in serum, and total tau (t-tau) and phosphorylated tau (p-tau) levels in cerebrospinal fluid., Methods: We measured biomarker levels in Young and Aged cynomolgus monkeys and correlated these with performance on three delayed response tasks., Results: The Aβ42 concentration of the Aged monkeys was significantly lower than in the Young subjects, while the t-tau and p-tau did not significantly differ between the groups. The Young subjects performed significantly better than the Aged individuals on the memory tests. Only Aβ42 levels were significantly correlated with performance in the three delayed response tasks., Conclusions: Circulating Aβ42 levels were lower in Aged monkeys and were correlated with inferior performance on delayed response tasks in Aged animals; therefore, both measures may be useful in establishing cynomolgus monkeys as models for studies of AD., (© 2013 John Wiley & Sons A/S. Published by John Wiley & Sons Ltd.)

Nerve myelination facilitates saltatory action potential conduction and exhibits spatiotemporal variation during development associated with the acquisition of behavioral and cognitive maturity. Although human cognitive development is unique, it is not known whether the ontogenetic progression of myelination in the human neocortex is evolutionarily exceptional. In this study, we quantified myelinated axon fiber length density and the expression of myelin-related proteins throughout postnatal life in the somatosensory (areas 3b/3a/1/2), motor (area 4), frontopolar (prefrontal area 10), and visual (areas 17/18) neocortex of chimpanzees (N = 20) and humans (N = 33). Our examination revealed that neocortical myelination is developmentally protracted in humans compared with chimpanzees. In chimpanzees, the density of myelinated axons increased steadily until adult-like levels were achieved at approximately the time of sexual maturity. In contrast, humans displayed slower myelination during childhood, characterized by a delayed period of maturation that extended beyond late adolescence. This comparative research contributes evidence crucial to understanding the evolution of human cognition and behavior, which arises from the unfolding of nervous system development within the context of an enriched cultural environment. Perturbations of normal developmental processes and the decreased expression of myelin-related molecules have been related to psychiatric disorders such as schizophrenia. Thus, these species differences suggest that the human-specific shift in the timing of cortical maturation during adolescence may have implications for vulnerability to certain psychiatric disorders.

Studies involving oro-facial asymmetries in nonhuman primates have largely demonstrated a right hemispheric dominance for communicative signals and conveyance of emotional information. A recent study on chimpanzee reported the first evidence of significant left-hemispheric dominance when using attention-getting sounds and rightward bias for species-typical vocalizations (Losin, Russell, Freeman, Meguerditchian, Hopkins & Fitch, 2008). The current study sought to extend the findings from Losin et al. (2008) with additional oro-facial assessment in a new colony of chimpanzees. When combining the two populations, the results indicated a consistent leftward bias for attention-getting sounds and a right lateralization for species-typical vocalizations. Collectively, the results suggest that both voluntary-controlled oro-facial and gestural communication might share the same left-hemispheric specialization and might have coevolved into a single integrated system present in a common hominid ancestor., (Copyright © 2012 Elsevier Inc. All rights reserved.)

Over the last century, the issue of brain lateralization in primates has been extensively investigated and debated, yet no previous study has reported eye preference in great apes. This study examined eye preference in 45 captive chimpanzees (Pan troglodytes) in response to various stimuli. Eye preference was assessed when animals looked through a hole that only accommodated one eye at an empty box, a mirror, a picture of a dog, a rubber snake, food biscuits, bananas, a rubber duck, and a video camera. Main effects of stimulus type were found for direction of eye preference, number of looks, and looking duration, but not for strength of eye preference. A left-eye bias was found for viewing the rubber snake and a right-eye bias was found for viewing the bananas, supporting theories that emotional valence may affect lateralized behaviors. In addition, a significant shift in eye preference took place from the initial look to subsequent looks when viewing the snake. These results are not consistent with previous reports of human eye preference and may reflect lateralization differences for emotional processing. No relationship between eye preference and previously recorded hand preference was found.

The acquisition of linguistic competency from more experienced social partners is a fundamental aspect of human language. However, there is little evidence that non-human primates learn to use their vocalizations from social partners. Captive chimpanzees (Pan troglodytes) produce idiosyncratic vocal signals that are used intentionally to capture the attention of a human experimenter. Interestingly, not all apes produce these sounds, and it is unclear what factors explain this difference. We tested the hypothesis that these attention-getting (AG) sounds are socially learned via transmission between mothers and their offspring. We assessed 158 chimpanzees to determine if they produced AG sounds. A significant association was found between mother and offspring sound production. This association was attributable to individuals who were raised by their biological mother-as opposed to those raised by humans in a nursery environment. These data support the hypothesis that social learning plays a role in the acquisition and use of communicative vocal signals in chimpanzees.

Functional and neuroanatomical asymmetries are an important characteristic of the human brain. The evolution of such specializations in the human cortex has provoked great interest in primate brain evolution. Most research on cortical sulci has revolved around linear measurements, which represent only one dimension of sulci organization. Here, we used a software program (BrainVISA) to quantify asymmetries in cortical depth and surface area from magnetic resonance images in a sample of 127 chimpanzees and 49 macaques. Population brain asymmetries were determined from 11 sulci in chimpanzees and seven sulci in macaques. Sulci were taken from the frontal, temporal, parietal, and occipital lobes. Population-level asymmetries were evident in chimpanzees for several sulci, including the fronto-orbital, superior precentral, and sylvian fissure sulci. The macaque population did not reveal significant population-level asymmetries, except for surface area of the superior temporal sulcus. The overall results are discussed within the context of the evolution of higher order cognition and motor functions., (Copyright © 2012 Elsevier Inc. All rights reserved.)

Cognitive and behavioral biases, which are widespread among humans, have recently been demonstrated in other primates, suggesting a common origin. Here we examine whether the expression of one shared bias, the endowment effect, varies as a function of context. We tested whether objects lacking inherent value elicited a stronger endowment effect (or preference for keeping the object) in a context in which the objects had immediate instrumental value for obtaining valuable resources (food). Chimpanzee subjects had opportunities to trade tools when food was not present, visible but unobtainable, and obtainable using the tools. We found that the endowment effect for these tools existed only when they were immediately useful, showing that the effect varies as a function of context-specific utility. Such context-specific variation suggests that the variation seen in some human biases may trace predictably to behaviors that evolved to maximize gains in specific circumstances.

In captive research environments for nonhuman primates (NHP), social housing strategies are often in conflict with protocols designed to minimize disease transmission. This is particularly true in breeding colonies, and is especially relevant when attempting to eliminate specific pathogens from a population of primates. Numerous strategies have been used to establish such specific pathogen free (SPF) breeding colonies (primarily of macaques), ranging from nursery rearing of neonates to single housing of socially reared yearlings to the rearing of infants in large social groups. All these strategies attempt to balance the effects of the chosen socialization strategy on parameters related to disease transmission, including the ultimate elimination of the target pathogens. Such strategies may affect the overall disease states of NHP breeding colonies through selective breeding processes. This can occur either by creating subpopulations of animals that do not have target diseases (SPF colonies), but may have other issues; or by creating situations in which the "best" animals are sold and the breeding colony is stocked with animals that may be more disease susceptible than those that were sold. The disease states of NHP research colonies also may be affected by selective utilization programs, in which animals removed from the breeding colony for health/behavior reasons, are preferentially chosen for use in scientific investigations. Such utilization criteria raise the question of whether ideal subjects are being chosen for use in research. Finally, captive primate colonies, where both socialization and disease states are intensely managed, may provide opportunities for those testing predictions from models of the interactions of socialization and disease transmission in the evolution of wild populations of NHP. This would be especially true for some extreme conditions of these disease ecology models, given the exceedingly high social densities and levels of pathogen control that exist in many captive nonhuman primate colonies., (© 2011 Wiley Periodicals, Inc.)

Whether lateralization of communicative signalling in non-human primates might constitute prerequisites of hemispheric specialization for language is unclear. In the present study, we examined (i) hand preference for a communicative gesture (clapping in 94 captive chimpanzees from two research facilities) and (ii) the in vivo magnetic resonance imaging brain scans of 40 of these individuals. The preferred hand for clapping was defined as the one in the upper position when the two hands came together. Using computer manual tracing of regions of interest, we measured the neuroanatomical asymmetries for the homologues of key language areas, including the inferior frontal gyrus (IFG) and planum temporale (PT). When considering the entire sample, there was a predominance of right-handedness for clapping and the distribution of right- and left-handed individuals did not differ between the two facilities. The direction of hand preference (right- versus left-handed subjects) for clapping explained a significant portion of variability in asymmetries of the PT and IFG. The results are consistent with the view that gestural communication in the common ancestor may have been a precursor of language and its cerebral substrates in modern humans.

Video displays for behavioral research lend themselves particularly well to studies with chimpanzees (Pan troglodytes), as their vision is comparable to humans', yet there has been no formal test of the efficacy of video displays as a form of social information for chimpanzees. To address this, we compared the learning success of chimpanzees shown video footage of a conspecific compared to chimpanzees shown a live conspecific performing the same novel task. Footage of an unfamiliar chimpanzee operating a bidirectional apparatus was presented to 24 chimpanzees (12 males, 12 females), and their responses were compared to those of a further 12 chimpanzees given the same task but with no form of information. Secondly, we also compared the responses of the chimpanzees in the video display condition to responses of eight chimpanzees from a previously published study of ours, in which chimpanzees observed live models. Chimpanzees shown a video display were more successful than those in the control condition and showed comparable success to those that saw a live model. Regarding fine-grained copying (i.e. the direction that the door was pushed), only chimpanzees that observed a live model showed significant matching to the model's methods with their first response. Yet, when all the responses made by the chimpanzees were considered, comparable levels of matching were shown by chimpanzees in both the live and video conditions., (© 2012 Wiley Periodicals, Inc.)

The remarkable ecological and demographic success of humanity is largely attributed to our capacity for cumulative culture, with knowledge and technology accumulating over time, yet the social and cognitive capabilities that have enabled cumulative culture remain unclear. In a comparative study of sequential problem solving, we provided groups of capuchin monkeys, chimpanzees, and children with an experimental puzzlebox that could be solved in three stages to retrieve rewards of increasing desirability. The success of the children, but not of the chimpanzees or capuchins, in reaching higher-level solutions was strongly associated with a package of sociocognitive processes-including teaching through verbal instruction, imitation, and prosociality-that were observed only in the children and covaried with performance.

Manipulations of the environments of captive nonhuman primates often have welfare consequences to the animals, including behavioral effects, and for certain manipulations, physiological effects as well. The processes of transporting, relocating, and acclimatizing nonhuman primates across facilities represent manipulations that are likely to have welfare, behavioral, and physiological consequences to the relocated animals. Seventy-two chimpanzees were relocated from the Primate Foundation of Arizona (PFA) in Arizona to the Keeling Center (KCCMR) in Texas. Animals were transported for approximately 21 h in single cages in a USDA-approved, climate-controlled trailer. Chimpanzees were weighed, anesthetized, and blood samples were collected 1) immediately prior to departure from PFA, 2) immediately upon arrival at the KCCMR, and 3) at additional time point(s) between 3 and 12 weeks after arrival at the KCCMR. Chimpanzees were quarantined in familiar pairs or social groups for 60-90 days at the KCCMR. Blood samples were analyzed for hematological and clinical chemistry parameters and compared across time points. In addition, samples from a subset of animals were assayed for cell-mediated immune parameters. Comparisons of the data obtained just prior to transport, to the data obtained immediately upon arrival, revealed numerous statistically significant differences in hematological, clinical chemistry, and immunological parameters. Some of these were indicative of stress, and thus, changes in welfare state, although many remained within the published normal ranges for chimpanzees. Additional analyses showed that many of the clinical chemistry values collected 3 to 12 weeks after arrival at the KCCMR had returned to pre-transport values. In contrast, of the cell-mediated immune parameters that were affected by transport and relocation, few had returned to pre-transport levels 8 weeks after transport, and three of the four hematology variables analyzed had not returned to pre-transport levels 12 weeks after transport. Comparisons of body weights before and immediately after transport revealed that animals lost an average of 2.5 kg during the 21-h transport, a statistically significant reduction that some animals never regained. These results demonstrate that transport and relocation affect a variety of physiological parameters with potential welfare implications and that some of these effects last as long as 3 months. These findings have important implications for the welfare and use of recently transported nonhuman primates, especially chimpanzees, in biomedical research. In order to allow animals to adapt to their new surroundings and to prevent unwanted confounds from influencing experiments, sufficient time must be provided after transport for chimpanzees to acclimatize.

Copulation preferences in our closest living relative, the chimpanzee, suggest that males prefer older females who have had previous offspring. However, this finding is counter to some behavioral models, which predict that chimpanzee males, as promiscuous breeders with minimal costs to mating, should show little or no preference when choosing mating partners (e.g. should mate indiscriminately). To determine if the preferences indicated by copulations appear in other contexts as well as how they interact, we examined how male chimpanzees' grooming patterns varied amongst females. We found that males' preferences were based on interactions among females' fertility status, age, and parity. First, grooming increased with increasing female parity. We further found an effect of the estrous cycle on grooming; when females were at the lowest point of their cycle, males preferentially groomed parous females at peak reproductive age, but during maximal tumescence, males preferred the oldest multiparous females. Nulliparous females received relatively little grooming regardless of age or fertility. Thus, male chimpanzees apparently chose grooming partners based on both female's experience and fertility, possibly indicating a two-pronged social investment strategy. Male selectivity seems to have evolved to effectively distribute costly social resources in a pattern which may increase their overall reproductive success., (© 2011 Wiley-Liss, Inc.)

Chimpanzees remain fixed on a single strategy, even if a novel, more efficient, strategy is introduced. Previous studies reporting such findings have incorporated paradigms in which chimpanzees learn one behavioural method and then are shown a new one that the chimpanzees invariably do not adopt. This study provides the first evidence that chimpanzees show such conservatism even when the new method employs the identical required behaviour as the first, but for a different reward. Groups of chimpanzees could choose to exchange one of two types of inedible tokens, with each token type being associated with a different food reward: one type was rewarded with a highly preferred food (grape) and the other type was rewarded with a less preferred food (carrot). Individuals first observed a model chimpanzee from their social group trained to choose one of the two types of tokens. In one group, this token earned a carrot, while in the other, control, group the token earned a grape. In both groups, chimpanzees conformed to the trained model's choice. This was especially striking for those gaining the pieces of carrot, the less favoured reward. This resulted in a population-level trend of food choices, even when counter to their original, individual, preferences. Moreover, the chimpanzees' food preferences did not change over time, demonstrating that these results were not due to a simple shift in preferences. We discuss social factors apparent in the interactions and suggest that, despite seeming to be inefficient, in chimpanzees, conformity may benefit them, possibly by assisting with the maintenance of group relations.

Whether or not nonhuman primates exhibit population-level handedness remains a topic of considerable scientific debate. Here, we examined handedness for coordinated bimanual actions in a sample of 777 great apes including chimpanzees, bonobos, gorillas, and orangutans. We found population-level right-handedness in chimpanzees, bonobos and gorillas, but left-handedness in orangutans. Directional biases in handedness were consistent across independent samples of apes within each genus. We suggest that, contrary to previous claims, population-level handedness is evident in great apes but differs among species as a result of ecological adaptations associated with posture and locomotion. We further suggest that historical views of nonhuman primate handedness have been too anthropocentric, and we advocate for a larger evolutionary framework for the consideration of handedness and other aspects of hemispheric specialization among primates., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

There is great interest in the evolution of economic behavior. In typical studies, species are asked to play one of a series of economic games, derived from game theory, and their responses are compared. The advantage of this approach is the relative level of consistency and control that emerges from the games themselves; however, in the typical experiment, procedures and conditions differ widely, particularly between humans and other species. Thus, in the current study, we investigated how three primate species, capuchin monkeys, chimpanzees, and humans, played the Assurance (or Stag Hunt) game using procedures that were, to the best of our ability, the same across species, particularly with respect to training and pretesting. Our goal was to determine what, if any, differences existed in the ways in which these species made decisions in this game. We hypothesized differences along phylogenetic lines, which we found. However, the species were more similar than might be expected. In particular, humans who played using "nonhuman primate-friendly" rules did not behave as is typical. Thus, we find evidence for similarity in decision-making processes across the order Primates. These results indicate that such comparative studies are possible and, moreover, that in any comparison rating species' relative abilities, extreme care must be taken in ensuring that one species does not have an advantage over the others due to methodological procedures.

Understanding the relationship between physical environments and nonhuman primate behavior is a key element for effective care and management in a range of settings. The physical features of the captive environment, including not only gross useable space but also environmental complexity, can have a significant influence on primate behavior and ultimately, animal welfare. But despite this connection, there remains relatively little conclusive data on how captive primates, especially great apes, use the spaces provided to them, especially in modern, indoor-outdoor enclosures that have become more prevalent in recent years. In this study, we used four years of detailed data on where 23 great apes (chimpanzees and gorillas) positioned themselves within a modern, indoor-outdoor zoo enclosure to determine not only how the apes utilized their space but also how access to outdoor areas affected their spatial selectivity. We found that both species used relatively little of their available space: chimpanzees and gorillas spent half their time in only 3.2 and 1.5% of their useable three-dimensional space, respectively. Chimpanzees utilized the outdoor space more than gorillas, but access to the outdoors did not affect space selectivity in the indoor area for either species. Although both species of ape were highly selective in their space use, consideration should be given to the importance of providing the choice to locate in a variety of spaces, including outdoor areas. These data represent an extremely detailed account of space selectivity by great apes in an indoor-outdoor environment and have substantial implications for future facility design and captive primate management., (© 2010 Wiley-Liss, Inc.)

In many facilities, primates are voluntarily transferred between different enclosures on a daily basis to facilitate animal husbandry and exhibit maintenance. This procedure is particularly relevant in the management of great apes living in zoos, where the requirements of functional management must be balanced with the desire to maintain enriching and naturalistic exhibit enclosures that benefit ape residents and attract the visiting public. In these settings, examinations of ape behavior and welfare typically focus exclusively on activity in the primary exhibit area. However, physical, social and sensory experiences unique to each area may shape different patterns of behavior. In the current study, zoo-living chimpanzees and gorillas were moved each day from exhibit areas to off-exhibit holding areas for a short duration as a part of regular management procedures. Behavioral data indicated species-specific reactions to the holding area, including increased aggression and self-directed behavior by chimpanzees and increased activity and prosocial behavior among gorilla subjects. Both species showed more feeding-foraging behavior while in the exhibit enclosure. Results suggest that holding areas may not meet all behavior needs of captive great apes and demonstrate the importance of including all components of the captive enclosure in comprehensive analyses of great ape behavior and welfare., (© 2010 Wiley-Liss, Inc.)

Several species of nonhuman primates respond negatively to inequitable outcomes, a trait shared with humans. Despite previous research, questions regarding the response to inequity remain. In this study, we replicated the methodology from previous studies to address four questions related to inequity. First, we explored the impact of basic social factors. Second, we addressed whether negative responses to inequity require a task, or exist when rewards are given for 'free'. Third, we addressed whether differences in the experimental procedure or the level of effort required to obtain a reward affected responses. Finally, we explored the interaction between 'individual' expectations (based on one's own previous experience) and 'social' expectations (based on the partner's experience). These questions were investigated in 16 socially housed adult chimpanzees using eight conditions that varied across the dimensions of reward, effort and procedure. Subjects did respond to inequity, but only in the context of a task. Differences in procedure and level of effort required did not cause individuals to change their behaviour. Males were more sensitive to social than to individual expectation, while females were more sensitive to individual expectation. Finally, subjects also increased refusals when they received a better reward than their partner, which has not been documented previously. These results indicate that chimpanzees are more sensitive to reward inequity than procedures, and that there is interaction between social and individual expectations that depends upon social factors.

Although behavioral lateralization is known to correlate with certain aspects of brain asymmetry in primates, there are limited data concerning hemispheric biases in the microstructure of the neocortex. In the present study, we investigated whether there is asymmetry in synaptophysin-immunoreactive puncta density and protein expression levels in the region of hand representation of the primary motor cortex in chimpanzees (Pan troglodytes). Synaptophysin is a presynaptic vesicle-associated protein found in nearly all synapses of the central nervous system. We also tested whether there is a relationship between hand preference on a coordinated bimanual task and the interhemispheric distribution of synaptophysin as measured by both stereologic counts of immunoreactive puncta and by Western blotting. Our results demonstrated that synaptophysin-immunoreactive puncta density is not asymmetric at the population level, whereas synaptophysin protein expression levels are significantly higher in the right hemisphere. Handedness was correlated with interindividual variation in synaptophysin-immunoreactive puncta density. As a group, left-handed and ambidextrous chimpanzees showed a rightward bias in puncta density. In contrast, puncta densities were symmetrical in right-handed chimpanzees. These findings support the conclusion that synapse asymmetry is modulated by lateralization of skilled motor behavior in chimpanzees.

Although tool use occurs in diverse species, its complexity may mark an important distinction between humans and other animals. Chimpanzee tool use has many similarities to that seen in humans, yet evidence of the cumulatively complex and constructive technologies common in human populations remains absent in free-ranging chimpanzees. Here we provide the first evidence that chimpanzees have a latent capacity to socially learn to construct a composite tool. Fifty chimpanzees were assigned to one of five demonstration conditions that varied in the amount and type of information available in video footage of a conspecific. Chimpanzees exposed to complete footage of a chimpanzee combining the two components to retrieve a reward learned to combine the tools significantly more than those exposed to more restricted information. In a follow-up test, chimpanzees that constructed tools after watching the complete demonstration tended to do so even when the reward was within reach of the unmodified components, whereas those that spontaneously solved the task (without seeing the modification process) combined only when necessary. Social learning, therefore, had a powerful effect in instilling a marked persistence in the use of a complex technique at the cost of efficiency, inhibiting insightful tool use.

Chimpanzees provide help to unrelated individuals in a broad range of situations. The pattern of helping within pairs suggests that contingent reciprocity may have been an important mechanism in the evolution of altruism in chimpanzees. However, correlational analyses of the cumulative pattern of interactions over time do not demonstrate that helping is contingent upon previous acts of altruism, as required by the theory of reciprocal altruism. Experimental studies provide a controlled approach to examine the importance of contingency in helping interactions. In this study, we evaluated whether chimpanzees would be more likely to provide food to a social partner from their home group if their partner had previously provided food for them. The chimpanzees manipulated a barpull apparatus in which actors could deliver rewards either to themselves and their partners or only to themselves. Our findings indicate that the chimpanzees' responses were not consistently influenced by the behavior of their partners in previous rounds. Only one of the 11 dyads that we tested demonstrated positive reciprocity. We conclude that contingent reciprocity does not spontaneously arise in experimental settings, despite the fact that patterns of behavior in the field indicate that individuals cooperate preferentially with reciprocating partners.

There is continued debate over the factors influencing handedness in captive and wild primates, notably chimpanzees. Previous studies in wild chimpanzees have revealed population-level left handedness for termite fishing. Here we examined hand preferences and performance on a tool use task designed to simulate termite fishing in a sample of 190 captive chimpanzees to evaluate whether patterns of hand use in captive chimpanzees differed from those observed for wild apes. No population-level handedness was found for this task; however, significant sex differences in preference and performance were found, with males showing greater left handedness and poorer performance compared to females. We also found that the hand preferences of offspring were significantly positively correlated with the hand preferences of their mothers. Lastly, older females performed more slowly on the task compared to younger individuals. The overall results neither confirm nor reject previous hypotheses claiming that raising chimpanzees in captivity induces right-handedness, but rather suggest that other factors may account for differences in hand preferences for tool use seen in wild and captive chimpanzees.

The stress associated with transportation of non-human primates used in scientific research is an important but almost unexplored part of laboratory animal husbandry. The procedures and routines concerning transport are not only important for the animals' physical health but also for their mental health as well. The transport stress in cynomolgus monkeys (Macaca fascicularis) was studied in two experiments. In Experiment 1, 25 adult female cynomolgus monkeys were divided into five groups of five animals each that received different diets during the transport phase of the experiment. All animals were transported in conventional single animal transport cages with no visual or tactile contact with conspecifics. The animals were transported by lorry for 24 h at ambient temperatures ranging between 20 degrees C and 35 degrees C. Urine produced before, during and after transport was collected and analysed for cortisol by enzyme-linked immunosorbent assay (ELISA). All monkeys exhibited a significant increase in cortisol excretion per time unit during the transport and on the first day following transport.Although anecdotal reports concerning diet during transport, including the provision of fruits and/or a tranquiliser, was thought likely to influence stress responses, these were not corrobated by the present study. In Experiment 2, behavioural data were collected from 18 cynomolgus macaques before and after transfer from group cages to either single or pair housing, and also before and after a simulated transport, in which the animals were housed in transport cages. The single housed monkeys were confined to single transport cages and the pair housed monkeys were kept in their pairs in double size cages. Both pair housed and singly housed monkeys showed clear behavioural signs of stress soon after their transfer out of their group cages.However, stress-associated behaviours were more prevalent in singly housed animals than in pair housed animals, and these behaviours persisted for a longer time after the simulated transport housing event than in the pair housed monkeys. Our data confirm that the transport of cynomolgus monkeys is stressful and suggest that it would be beneficial for the cynomolgus monkeys to be housed and transported in compatible pairs from the time they leave their group cages at the source country breeding facility until they arrive at their final laboratory destination in the country of use.

We conducted experiments on two populations of chimpanzees, Pan troglodytes , to determine whether they would take advantage of opportunities to provide food rewards to familiar group members at little cost to themselves. In both of the experiments described here, chimpanzees were able to deliver identical rewards to themselves and to other members of their social groups. We compared the chimpanzees' behaviour when they were paired with another chimpanzee and when they were alone. If chimpanzees are motivated to provide benefits to others, they are expected to consistently deliver rewards to others and to distinguish between the partner-present and partner-absent conditions. Results from both experiments indicate that our subjects were largely indifferent to the benefits they could provide to others. They were less likely to provide rewards to potential recipients as the experiment progressed, and all but one of the 18 subjects were as likely to deliver rewards to an empty enclosure as to an enclosure housing another chimpanzee. These results, in conjunction with similar results obtained in previous experiments, suggest that chimpanzees are not motivated by prosocial sentiments to provide food rewards to other group members.

Emulation has been distinguished from imitation as a form of observational learning because it focuses not on the model's actions but on the action's environmental results. Whether a species emulates, imitates or displays only simpler observational learning is expected to have profound implications for its capacity for cultural transmission. Chimpanzees' observational learning has been suggested to be primarily emulative, but this is an inference largely based upon low fidelity copying in experiments when comparing chimpanzees with humans rather than direct testing. Here we test directly for emulation learning by chimpanzees and children using a 'ghost' condition in which a sliding door obscuring a reward was moved to left or right with no agent visible, a context associated with the only published evidence for emulation learning in a non-human species (pigeons). Both children and chimpanzees matched the observed direction of ghost door movement on their first test trial. This is the first evidence for emulation in a non-human primate in the restricted context of a ghost condition. However, only the children continued to match in later trials. Individuals of both species continued to match with 99% or better fidelity when viewing a conspecific model operates the door. We conclude that chimpanzees can and will display emulation learning when the task is as simple as the present one, which contrasts with a failure to do so in a more complex manipulative task tested earlier. However, even with a simple task, emulation alone creates only fleeting fidelity compared with the opportunity to copy a conspecific, when considerable conformity is displayed.

Background: Economists believe that barter is the ultimate cause of social wealth--and even much of our human culture--yet little is known about the evolution and development of such behavior. It is useful to examine the circumstances under which other species will or will not barter to more fully understand the phenomenon. Chimpanzees (Pan troglodytes) are an interesting test case as they are an intelligent species, closely related to humans, and known to participate in reciprocal interactions and token economies with humans, yet they have not spontaneously developed costly barter., Methodology/principle Findings: Although chimpanzees do engage in noncostly barter, in which otherwise value-less tokens are exchanged for food, this lack of risk is not typical of human barter. Thus, we systematically examined barter in chimpanzees to ascertain under what circumstances chimpanzees will engage in costly barter of commodities, that is, trading food items for other food items with a human experimenter. We found that chimpanzees do barter, relinquishing lower value items to obtain higher value items (and not the reverse). However, they do not trade in all beneficial situations, maintaining possession of less preferred items when the relative gains they stand to make are small., Conclusions/significance: Two potential explanations for this puzzling behavior are that chimpanzees lack ownership norms, and thus have limited opportunity to benefit from the gains of trade, and that chimpanzees' risk of defection is sufficiently high that large gains must be imminent to justify the risk. Understanding the conditions that support barter in chimpanzees may increase understanding of situations in which humans, too, do not maximize their gains.

As more emphasis is placed on enhancing the psychological well-being of nonhuman primates, many research facilities have started using positive reinforcement training (PRT) techniques to train primates to voluntarily participate in husbandry and research procedures. PRT increases the animal's control over its environment and desensitizes the animal to stressful stimuli. Blood draw is a common husbandry and research procedure that can be particularly stressful for nonhuman primate subjects. Although studies have demonstrated that chimpanzees can be trained for in-cage venipuncture using PRT only, fewer studies have demonstrated success using similar techniques to train macaques. It is often assumed that macaques cannot be trained in the same manner as apes. In this study, we compare PRT data from singly housed adult rhesus macaques (Macaca mulatta; n = 8) with data from group-housed adult chimpanzees (Pan troglodytes; n = 4). All subjects were trained to place an arm in a 'blood sleeve' and remain stationary for venipuncture. Both facilities used similar PRT techniques. We were able to obtain repeated blood samples from 75% of the macaques and all of the chimpanzees. The training time did not differ significantly between the 2 species. These data demonstrate that macaques can be trained for venipuncture in a manner similar to that used for chimpanzees.

Human behavior is not always consistent with standard rational choice predictions. Apparent deviations from rational choice predictions provide a promising arena for the merger of economics and biology [1-6]. Although little is known about the extent to which other species exhibit these seemingly irrational patterns [7-9], similarities across species would suggest a common evolutionary root to the phenomena. The present study investigated whether chimpanzees exhibit an endowment effect, a seemingly paradoxical behavior in which humans tend to value a good they have just come to possess more than they would have only a moment before [10-13]. We show the first evidence that chimpanzees do exhibit an endowment effect, by favoring items they just received more than their preferred items that could be acquired through exchange. Moreover, the effect is stronger for food than for less evolutionarily salient objects, perhaps because of historically greater risks associated with keeping a valuable item versus attempting to exchange it for another [14, 15]. These findings suggest that many seeming deviations from rational choice predictions may be common to humans and chimpanzees and that the evaluation of these through a lens of evolutionary relevance may yield further insights in humans and other species.

Field reports provide increasing evidence for local behavioral traditions among fish, birds, and mammals. These findings are significant for evolutionary biology because social learning affords faster adaptation than genetic change and has generated new (cultural) forms of evolution. Orangutan and chimpanzee field studies suggest that like humans, these apes are distinctive among animals in each exhibiting over 30 local traditions. However, direct evidence is lacking in apes and, with the exception of vocal dialects, in animals generally for the intergroup transmission that would allow innovations to spread widely and become evolutionarily significant phenomena. Here, we provide robust experimental evidence that alternative foraging techniques seeded in different groups of chimpanzees spread differentially not only within groups but serially across two further groups with substantial fidelity. Combining these results with those from recent social-diffusion studies in two larger groups offers the first experimental evidence that a nonhuman species can sustain unique local cultures, each constituted by multiple traditions. The convergence of these results with those from the wild implies a richness in chimpanzees' capacity for culture, a richness that parsimony suggests was shared with our common ancestor.

Many people have devoted considerable effort to enhancing the environments of nonhuman primates in captivity. There is substantial motivation to develop experimental, analytical, and interpretational frameworks to enable objective measurements of the value of environmental enrichment/behavioral management efforts. The consumer-demand approach is a framework not frequently implemented in studies of nonhuman primate welfare but profitably used in studies of the welfare of nonhuman animals in agriculture. Preference studies, in which primates can voluntarily choose to socialize or to participate in training, may be the best current examples of a consumer-demand-like approach to assessing the effects of captive management strategies on primate welfare. Additional work in this area would be beneficial; however, there are potential ethical constraints on purposefully subjecting primates to adverse circumstances to measure their demand for a resource. Primate welfare researchers need to design consumer-demand studies with obstacles that will help measure the relative value of resources to captive primates without compromising the welfare they are attempting to evaluate and enhance.

It has been proposed that human right handedness is determined by genetic factors associated with the emergence of language, whereas non-human primate handedness is determined by random, non-genetic factors. These different mechanisms account for differences in the distribution of handedness between human and non-human primates. Here we report evidence that genetic factors play a role in the determination of handedness in chimpanzees. We further report that differential rearing has no influence on the expression of handedness in related individuals. Contrary to many theories of the origin of handedness, these results indicate that genetic factors have a significant influence on handedness in chimpanzees.