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Eye gaze is an important source of information for animals, implicated in communication, cooperation, hunting and antipredator behaviour. Gaze perception and its cognitive underpinnings are much studied in primates, but the specific features that are used to estimate gaze can be difficult to isolate behaviourally. We photographed 13 laboratory-housed tufted capuchin monkeys ( Sapajus [Cebus] apella ) to quantify chromatic and achromatic contrasts between their iris, pupil, sclera and skin. We used colour vision models to quantify the degree to which capuchin eye gaze is discriminable to capuchins, their predators and their prey. We found that capuchins, regardless of their colour vision phenotype, as well as their predators, were capable of effectively discriminating capuchin gaze across ecologically relevant distances. Their prey, in contrast, were not capable of discriminating capuchin gaze, even under relatively ideal conditions. These results suggest that specific features of primate eyes can influence gaze perception, both within and across species.

Once considered a hallmark of human uniqueness, brain asymmetry has emerged as a feature shared with several other species, including chimpanzees, one of our closest living relatives. Most notable has been the discovery of asymmetries in homologues of cortical language areas in apes, particularly in the planum temporale (PT), considered a central node of the human language network. Several lines of evidence indicate a role for genetic mechanisms in the emergence of PT asymmetry; however, the genetic determinants of cerebral asymmetries have remained elusive. Studies in humans suggest that there is heritability of brain asymmetries of the PT, but this has not been explored to any extent in chimpanzees. Furthermore, the potential influence of non-genetic factors has raised questions about the reproducibility of earlier observations of PT asymmetry reported in chimpanzees. As such, the present study was aimed at examining both the heritability of phenotypic asymmetries in PT morphology, as well as their reproducibility. Using magnetic resonance imaging, we evaluated morphological asymmetries of PT surface area (mm 2 ) and mean depth (mm) in captive chimpanzees ( n = 291) derived from two genetically isolated populations. Our results confirm that chimpanzees exhibit a significant population-level leftward asymmetry for PT surface area, as well as significant heritability in the surface area and mean depth of the PT. These results conclusively demonstrate the existence of a leftward bias in PT asymmetry in chimpanzees and suggest that genetic mechanisms play a key role in the emergence of anatomical asymmetry in this region.

Across the animal kingdom, males tend to exhibit more behavioural and morphological variability than females, consistent with the 'greater male variability hypothesis'. This may reflect multiple mechanisms operating at different levels, including selective mechanisms that produce and maintain variation, extended male development, and X chromosome effects. Interestingly, human neuroanatomy shows greater male variability, but this pattern has not been demonstrated in any other species. To address this issue, we investigated sex-specific neuroanatomical variability in chimpanzees by examining relative and absolute surface areas of 23 cortical sulci across 226 individuals (135F/91M), using permutation tests of the male-to-female variance ratio of residuals from MCMC generalized linear mixed models controlling for relatedness. We used these models to estimate sulcal size heritability, simulations to assess the significance of heritability, and Pearson correlations to examine inter-sulcal correlations. Our results show that: (i) male brain structure is relatively more variable; (ii) sulcal surface areas are heritable and therefore potentially subject to selection; (iii) males exhibit lower heritability values, possibly reflecting longer development; and (iv) males exhibit stronger inter-sulcal correlations, providing indirect support for sex chromosome effects. These results provide evidence that greater male neuroanatomical variability extends beyond humans, and suggest both evolutionary and developmental explanations for this phenomenon.

Various non-human animal species have been shown to exhibit behavioural traditions. Importantly, this research has been guided by what we know of human culture, and the question of whether animal cultures may be homologous or analogous to our own culture. In this paper, we assess whether models of human cultural transmission are relevant to understanding biological fundamentals by investigating whether accounts of human payoff-biased social learning are relevant to chimpanzees ( Pan troglodytes ). We submitted 4- and 5-year-old children ( N = 90) and captive chimpanzees ( N = 69) to a token-reward exchange task. The results revealed different forms of payoff-biased learning across species and contexts. Specifically, following personal and social exposure to different tokens, children's exchange behaviour was consistent with proportional imitation, where choice is affected by both prior personally acquired and socially demonstrated token-reward information. However, when the socially derived information regarding token value was novel, children's behaviour was consistent with proportional observation; paying attention to socially derived information and ignoring their prior personal experience. By contrast, chimpanzees' token choice was governed by their own prior experience only, with no effect of social demonstration on token choice, conforming to proportional reservation. We also find evidence for individual- and group-level differences in behaviour in both species. Despite the difference in payoff strategies used, both chimpanzees and children adopted beneficial traits when available. However, the strategies of the children are expected to be the most beneficial in promoting flexible behaviour by enabling existing behaviours to be updated or replaced with new and often superior ones., (© 2017 The Authors.)

Human brains are markedly asymmetric in structure and lateralized in function, which suggests a relationship between these two properties. The brains of other closely related primates, such as chimpanzees, show similar patterns of asymmetry, but to a lesser degree, indicating an increase in anatomical and functional asymmetry during hominin evolution. We analysed the heritability of cerebral asymmetry in chimpanzees and humans using classic morphometrics, geometric morphometrics, and quantitative genetic techniques. In our analyses, we separated directional asymmetry and fluctuating asymmetry (FA), which is indicative of environmental influences during development. We show that directional patterns of asymmetry, those that are consistently present in most individuals in a population, do not have significant heritability when measured through simple linear metrics, but they have marginally significant heritability in humans when assessed through three-dimensional configurations of landmarks that reflect variation in the size, position, and orientation of different cortical regions with respect to each other. Furthermore, genetic correlations between left and right hemispheres are substantially lower in humans than in chimpanzees, which points to a relatively stronger environmental influence on left-right differences in humans. We also show that the level of FA has significant heritability in both species in some regions of the cerebral cortex. This suggests that brain responsiveness to environmental influences, which may reflect neural plasticity, has genetic bases in both species. These results have implications for the evolvability of brain asymmetry and plasticity among humans and our close relatives., (© 2016 The Author(s).)

Chimpanzees are well known for their tool using abilities. Numerous studies have documented variability in tool use among chimpanzees and the role that social learning and other factors play in their development. There are also findings on hand use in both captive and wild chimpanzees; however, less understood are the potential roles of genetic and non-genetic mechanisms in determining individual differences in tool use skill and laterality. Here, we examined heritability in tool use skill and handedness for a probing task in a sample of 243 captive chimpanzees. Quantitative genetic analysis, based on the extant pedigrees, showed that overall both tool use skill and handedness were significantly heritable. Significant heritability in motor skill was evident in two genetically distinct populations of apes, and between two cohorts that received different early social rearing experiences. We further found that motor skill decreased with age and that males were more commonly left-handed than females. Collectively, these data suggest that though non-genetic factors do influence tool use performance and handedness in chimpanzees, genetic factors also play a significant role, as has been reported in humans., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

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.

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.

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.