Masaki Tomonaga

Group differences in the mutual gaze of chimpanzees (Pan troglodytes).

A comparative developmental framework was used to determine whether mutual gaze is unique to humans and, if not, whether common mechanisms support the development of mutual gaze in chimpanzees and humans. Mother-infant chimpanzees engaged in approximately 17 instances of mutual gaze per hour. Mutual gaze occurred in positive, nonagonistic contexts. Mother-infant chimpanzees at a Japanese center exhibited significantly more mutual gaze than those at a center in the United States. Cradling and motor stimulation varied across groups. Time spent cradling infants was inversely related to mutual gaze. It is suggested that in primates, mutual engagement is supported via an interchangeability of tactile and visual modalities. The importance of mutual gaze is best understood within a perspective that embraces both cross-species and cross-cultural data.

How chimpanzees look at pictures: a comparative eye-tracking study

Surprisingly little is known about the eye movements of chimpanzees, despite the potential contribution of such knowledge to comparative cognition studies. Here, we present the first examination of eye tracking in chimpanzees. We recorded the eye movements of chimpanzees as they viewed naturalistic pictures containing a full-body image of a chimpanzee, a human or another mammal; results were compared with those from humans. We found a striking similarity in viewing patterns between the two species. Both chimpanzees and humans looked at the animal figures for longer than at the background and at the face region for longer than at other parts of the body. The face region was detected at first sight by both species when they were shown pictures of chimpanzees and of humans. However, the eye movements of chimpanzees also exhibited distinct differences from those of humans; the former shifted the fixation location more quickly and more broadly than the latter. In addition, the average duration of fixation on the face region was shorter in chimpanzees than in humans. Overall, our results clearly demonstrate the eye-movement strategies common to the two primate species and also suggest several notable differences manifested during the observation of pictures of scenes and body forms.

Spontaneous synchronized tapping to an auditory rhythm in a chimpanzee

Humans actively use behavioral synchrony such as dancing and singing when they intend to make affiliative relationships. Such advanced synchronous movement occurs even unconsciously when we hear rhythmically complex music. A foundation for this tendency may be an evolutionary adaptation for group living but evolutionary origins of human synchronous activity is unclear. Here we show the first evidence that a member of our closest living relatives, a chimpanzee, spontaneously synchronizes her movement with an auditory rhythm: After a training to tap illuminated keys on an electric keyboard, one chimpanzee spontaneously aligned her tapping with the sound when she heard an isochronous distractor sound. This result indicates that sensitivity to, and tendency toward synchronous movement with an auditory rhythm exist in chimpanzees, although humans may have expanded it to unique forms of auditory and visual communication during the course of human evolution.

Emergence of symmetry in a visual conditional discrimination by chimpanzees (Pan troglodytes)

In Exp. 1, three young chimpanzees were trained to match red to a cross and green to a circle in an arbitrary matching-to-sample task. After acquisition of this task, they were tested for the emergence of associative symmetry of these conditional relations using the trials on which shapes were presented as samples and colors as comparisons. One of the three chimpanzees showed statistically significant accuracy on these test trials. This successful subject served in Exp. 2, in which an auditory-visual stimulus appeared contingent upon red and a cross while another auditory-visual stimulus was contingent upon green and a circle. This subject showed higher accuracies in symmetry tests than in Exp. 1, which suggested the facilitative effects of these events common to sample and correct comparison on the development of symmetry. In Exp. 3, subsequent tests in which only these stimuli were presented as samples indicated that these stimuli might have become the members of equivalence classes.

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.

Relative numerosity discrimination by chimpanzees ( Pan troglodytes ): evidence for approximate numerical representations

Two adult chimpanzees were trained on a relative “numerosity” discrimination task. In each trial, two arrays containing different numbers of red dots were presented on a CRT monitor. The subjects were required to choose the array containing the larger number of dots. In Experiment 1, using numerosities between 1 and 8, 28 different pairs were presented repeatedly, and accuracy scores were analyzed to explore which cues the chimpanzee subjects utilized to perform the task. Multiple regression analyses revealed that the subjects’ performance was (1) not simply controlled by the “numerical” difference between arrays, but that it was (2) best described by Fechner’s Law–that is accuracy increased linearly with the logarithmic value of the numerical difference between arrays divided by the number in the larger of the two arrays. This relationship was maintained when using much larger numerosities (Experiment 3). In Experiment 2, the chimpanzees were tested on the effects of total area and density by manipulating dot size and presentation area. The results revealed that these factors clearly affected the subjects’ performance but that they could not alone explain the results, suggesting that the chimpanzees did use relative numerosity difference as a discriminative cue.

Enumeration of briefly presented items by the chimpanzee (Pan troglodytes) and humans (Homo sapiens)

In this study, we compared the performances on an enumeration task (numerical labeling task) of 1 chimpanzee (Pan troglodytes) and 4 humans. In this task, two types of trials, with different exposure durations of the sample that was to be enumerated, were used. In the unlimited-exposure trials, the sample remained on until the subject made a choice. In the brief-exposure trials, the sample was presented for 100 msec and then was masked. The results show clear differences between the different species. The main differences had to do with accuracy during the unlimited trials and response times during the brief trials. Detailed analyses of the pattern of response times for the chimpanzee and of looking-back behavior during the task suggests that the enumeration process underlying the subject’s performance was not counting but estimation.

How laboratory-raised Japanese monkeys (Macaca fuscata) perceive rotated photographs of monkeys: Evidence for an inversion effect in face perception

Five laboratory-raised Japanese monkeys (Macaca fuscata) were presented various types of photographs of Japanese and rhesus monkeys (Macaca mulatta) in upright, horizontal, and inverted orientations in a sensory-reinforcement experiment. The ratio of the duration of potential viewing time for the photographs which the subjects controlled to the interval between subject-controlled presentations of them (the D/I score) was used as a measure of preference for the photographs. When inverted photographs were presented, the D/I scores were lower than for upright photographs. The difference in D/I scores between photographs of the two species, which indicated discriminability between them, also diminished when the photographs were inverted. The results obtained suggest an inversion effect in face perception in macaque monkeys.

Differential Prefrontal White Matter Development in Chimpanzees and Humans

A comparison of developmental patterns of white matter (WM) within the prefrontal region between humans and nonhuman primates is key to understanding human brain evolution. WM mediates complex cognitive processes and has reciprocal connections with posterior processing regions [1, 2]. Although the developmental pattern of prefrontal WM in macaques differs markedly from that in humans [3], this has not been explored in our closest evolutionary relative, the chimpanzee. The present longitudinal study of magnetic resonance imaging scans demonstrated that the prefrontal WM volume in chimpanzees was immature and had not reached the adult value during prepuberty, as observed in humans but not in macaques. However, the rate of prefrontal WM volume increase during infancy was slower in chimpanzees than in humans. These results suggest that a less mature and more protracted elaboration of neuronal connections in the prefrontal portion of the developing brain existed in the last common ancestor of chimpanzees and humans, and that this served to enhance the impact of postnatal experiences on neuronal connectivity. Furthermore, the rapid development of the human prefrontal WM during infancy may help the development of complex social interactions, as well as the acquisition of experience-dependent knowledge and skills to shape neuronal connectivity.

Face scanning in chimpanzees and humans: continuity and discontinuity

How do chimpanzees, Pan troglodytes, the species with the closest evolutionary connection to humans, view faces? This study is the first to use the eye-tracking method to perform direct comparisons between humans and chimpanzees with regard to face scanning. Members of both species viewed the same sets of photographs representing conspecific and nonconspecific faces under the same experimental conditions. Chimpanzees and humans showed systematic and similar patterns of face scanning, including intensely viewing main facial features (i.e. eyes, nose and mouth) and inspecting the eyes and mouth, in that order. However, several differences between the species were also evident. For example, humans were more likely to show sequential refixations on the eye regions than were chimpanzees, whereas chimpanzees were more likely to engage in quick, vertical scanning over the eyes and mouth. Such species similarities and differences were consistent across conspecific and nonconspecific faces and were thus independent of the external morphologies of species-specific faces. Furthermore, when presented with facial expressions, chimpanzees changed their scanning patterns in response to those facial actions, whereas humans maintained intense eye viewing across the expressions. Finally, we discuss how these face-scanning patterns are related to species-specific forms of facial communication in chimpanzees and humans, and suggest that both species have unique eye movement strategies for interactions with conspecifics.