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.     

Social learning of nut-cracking behavior in East African sanctuary-living chimpanzees (Pan troglodytes schweinfurthii)

Nut cracking is restricted to communities of wild chimpanzees living in West Africa, suggesting it is an example of a socially transmitted tradition. Detailed study of the acquisition of nut cracking in wild chimpanzees is consistent with this conclusion. However, only 2, small-scale experiments have been carried out in captivity to explore the role of social transmission in the acquisition of this behavior. The study presented here does this with a comparatively larger, statistically viable sample of 11 sanctuary-living chimpanzees (Pan troglodytes schweinfurthii), permitting both between- and within-subjects experimental manipulations. Results confirmed that nut cracking can be acquired in a matter of days by social learning, but only in chimpanzees 3 to 4 years old and older. Direct comparisons are made with a study carried out in the wild, revealing striking similarities in developmental profiles.     

Imitation in neonatal chimpanzees (Pan troglodytes)

This paper provides evidence for imitative abilities in neonatal chimpanzees (Pan troglodytes), our closest relatives. Two chimpanzees were reared from birth by their biological mothers. At less than 7 days of age the chimpanzees could discriminate between, and imitate, human facial gestures (tongue protrusion and mouth opening). By the time they were 2 months old, however, the chimpanzees no longer imitated the gestures. They began to perform mouth opening frequently in response to any of the three facial gestures presented to them. These findings suggest that neonatal facial imitation is most likely an innate ability, developed through natural selection in humans and in chimpanzees. The relationship between the disappearance of neonatal imitation and the development of social communicative behavior is discussed from an evolutionary perspective.     

Dominance assertion in male chimpanzees (Pan troglodytes)

The social relations of three adult males in a captive chimpanzee group were evaluated for 12 months. The observations encompassed a period of transition in the dominance hierarchy providing an opportunity to assess behavioral patterns involved in the initiation of status changes. This study shows that dominance in the chimpanzee is expressed primarily through ritualized and non-contact behavior rather than by overt aggression. In particular, display behavior is important for indicating dominance rank, in communicating aggressive intent, and as a prelude to status struggles. The functional relationships between status, displays and other socio-sexual behavior patterns are presented.     

Lateral bias in infant chimpanzees (Pan troglodytes)

This study documents the presence, strength, and direction of lateralization in chimpanzees (Pan troglodytes) over the first 3 months of life. Nursery-reared chimpanzees (7 males and 5 females) were repeatedly assessed on a behavioral scale. Lateral bias was measured for 4 behaviors: hand-to-mouth, hand-to-hand, defensive grasp, and first step. Hand-to-mouth was significantly lateralized for the sample. Eight of the 10 chimpanzees that showed hand-to-mouth used the right hand. Lateral bias for defensive grasp was positively related to lateral bias both of first step and of hand-to-mouth. Lateral bias in hand-to-mouth was inversely related to lateral bias in hand-to-hand. Strength of lateralization increased as chimpanzees matured. These laterality effects in infant chimpanzees were expressed under conditions of emotional arousal. Moreover, degree of laterality may be a predictor of responsivity to stress.     

A group-specific arbitrary tradition in chimpanzees (Pan troglodytes)

Social learning in chimpanzees has been studied extensively and it is now widely accepted that chimpanzees have the capacity to learn from conspecifics through a multitude of mechanisms. Very few studies, however, have documented the existence of spontaneously emerged traditions in chimpanzee communities. While the rigour of experimental studies is helpful to investigate social learning mechanisms, documentation of naturally occurring traditions is necessary to understand the relevance of social learning in the real lives of animals. In this study, we report on chimpanzees spontaneously copying a seemingly non-adaptive behaviour (“grass-in-ear behaviour”). The behaviour entailed chimpanzees selecting a stiff, straw-like blade of grass, inserting the grass into one of their own ears, adjusting the position, and then leaving it in their ear during subsequent activities. Using a daily focal follow procedure, over the course of 1 year, we observed 8 (out of 12) group members engaging in this peculiar behaviour. Importantly, in the three neighbouring groups of chimpanzees (n = 82), this behaviour was only observed once, indicating that ecological factors were not determiners of the prevalence of this behaviour. These observations show that chimpanzees have a tendency to copy each other’s behaviour, even when the adaptive value of the behaviour is presumably absent.     

Development of face recognition in infant chimpanzees (Pan troglodytes)

In this paper, we assessed the developmental changes in face recognition by three infant chimpanzees aged 1–18 weeks, using preferential-looking procedures that measured the infants’ eye- and head-tracking of moving stimuli. In Experiment 1, we prepared photographs of the mother of each infant and an “average” chimpanzee face using computer-graphics technology. Prior to 4 weeks of age, the infants showed few tracking responses and no differential responses. Between 4 and 8 weeks of age, they paid greater attention to their mother's face. From 8 weeks onward, they again showed no differences, but exhibited frequent tracking responses. Experiment 2 investigated the infants’ tracking responses between a familiar human's and an “average” human face. The infants did not show any evidence of recognizing the human faces. We discuss the development of face recognition in relation to the effects of other species’ faces and postnatal visual experience.     

Imitation of intentional manipulatory actions in chimpanzees (Pan troglodytes)

In this study, the authors investigated the understanding of other's actions in 5 adult chimpanzees (Pan troglodytes). A human demonstrated an attempt to open different containers. Each container required a different motor pattern to open it. Along with the container, a 2nd object was made available. After a free play period in which the chimpanzees' natural behaviors toward the objects were recorded, the authors tested the following 2 phases: The demonstrator (a) tried but failed to open and (b) opened the container successfully, with 1 of 2 alternative strategies, either using an "irrelevant tool" or by hand. The chimpanzees did not reproduce the demonstrator's motor patterns precisely but did reproduce the demonstrated strategies in both phases. These results suggest that chimpanzees anticipate the intentions of others by perceiving the directionality and causality of object(s) as available cues.     

Inferences about guessing and knowing by chimpanzees (Pan troglodytes)

The visual perspective-taking ability of 4 chimpanzees (Pan troglodytes) was investigated. The subjects chose between information about the location of hidden food provided by 2 experimenters who randomly alternated between two roles (the guesser and the knower). The knower baited 1 of 4 obscured cups so that the subjects could watch the process but could not see which of the cups contained the reward. The guesser waited outside the room until the food was hidden. Finally, the knower pointed to the correct cup while the guesser pointed to an incorrect one. The chimpanzees quickly learned to respond to the knower. They also showed transfer to a novel variation of the task, in which the guesser remained inside the room and covered his head while the knower stood next to him and watched a third experimenter bait the cups. The results are consistent with the hypothesis that chimpanzees are capable of modeling the visual perspectives of others.     

How cross-fostered chimpanzees (Pan troglodytes) initiate and maintain conversations

This study systematically sampled typical attention-getting sounds and sign language conversations between each of 4 originally cross-fostered chimpanzees (Pan troglodytes), still living freely, but now in a laboratory setting, and a familiar human interlocutor. Videotape records showed that when they encountered a human interlocutor sitting alone at his desk with his back turned to them, the cross-fosterlings either left the scene or made attention-getting sounds. The only signs they made to the interlocutor's back were noisy signs. When the human turned and faced them, the chimpanzees promptly signed to him (98% of the time) and rarely made any sounds during the ensuing signed conversations. Under systematic experimental conditions, the signed responses of the chimpanzees were appropriate to the conversational styles of the human interlocutor, confirming daily field observations.     

Constructive and deconstructive tool modification by chimpanzees (Pan troglodytes)

Nine chimpanzees (Pan troglodytes) were tested for their ability to assemble or disassemble the appropriate tool to obtain a food reward from two different apparatus. In its deconstructed form, the tool functioned as a probe for one apparatus. In its constructed form, the tool functioned as a hook, appropriate for a second apparatus. Each subject completed four test trials with each apparatus type. Tool types were randomized and counter-balanced between the two forms. Results demonstrated that adult and juvenile chimpanzees (N = 7) were successful with both tool types, while two infant chimpanzees performed near chance. Off-line video analyses revealed that tool modifications followed by attempted solutions by the adults and juveniles were typically correct on the first attempt. Neither infant was successful in modifying tools correctly on the first attempt over all eight trials. The older chimpanzees’ ability to modify the appropriate tool consistently prior to use indicates an immediate recognition of the functional attributes necessary for the successful use of tool types on each apparatus, and represents a non-replication of a previously reported study by Povinelli.

Development of self-recognition in chimpanzees (Pan troglodytes).

Chimpanzees (Pan troglodytes) demonstrate the ability to recognize themselves in mirrors, yet investigations of the development of self-recognition in chimpanzees are sparse. Twelve young chimpanzees, grouped by age, were given mirror exposure and tested for self-recognition and contingent movement. All 6 juveniles, 4 and 5 years old, exhibited mirror-guided, mark-directed behavior and clear evidence of self-recognition. In contrast, among the infants, only the oldest group of 2 1/2-year-olds exhibited clear evidence of self-recognition. All chimpanzees exhibited both self-directed behaviors and contingent movements. These results suggest that self-recognition occurs at a slightly older age in chimpanzees than in human infants. In humans, self-recognition is linked with other cognitive abilities. The results conform to the general pattern that great apes exhibit many cognitive skills comparable to those of 2-year-old humans.     

Visual kin recognition and family resemblance in chimpanzees (Pan troglodytes)

The male-offspring biased visual kin recognition in chimpanzees (Pan troglodytes) reported by L. A. Parr and F. B. M. de Waal (1999) was replicated with human (Homo sapiens) participants and a principal components analysis (PCA) of pixel maps of the chimpanzee face photos. With the same original materials and methods, both humans and the PCA produced the same asymmetry in kin recognition as found with the chimpanzees. The PCA suggested that the asymmetry was a function of differences in the distribution of global characteristics associated with the framing of the faces in the son and daughter test sets. Eliminating potential framing biases, either by cropping the photos tightly to the faces or by rebalancing the recognition foils, eliminated the asymmetry but not human participants' ability to recognize chimpanzee kin.     

The generalization of deferred imitation in enculturated chimpanzees (Pan troglodytes)

Deferred imitation of object-related actions and generalization of imitation to similar but not identical tasks was assessed in three human-reared (enculturated) chimpanzees, ranging in age from 5 to 9 years. Each ape displayed high levels of deferred imitation and only slightly lower levels of generalization of imitation. The youngest two chimpanzees were more apt to generalize the model's actions when they had displayed portions of the target behaviors at baseline, consistent with the idea that learning is more likely to occur when working within the "zone of proximal development." We argue that generalization of imitation is the best evidence to date of imitative learning in chimpanzees. Electronic Publication     

Spontaneous discrimination of natural stimuli by chimpanzees (Pan troglodytes)

Six chimpanzees (Pan troglodytes) were presented with pairs of color photographic images of 5 different categories of animals (cat, chimp, gorilla, tiger, fish). The subjects responded to each pair using symbols for "same" and "different." Both within- and between-category discriminations were tested, and all chimpanzees classified the image pairs in accordance with the 5 experimenter-defined categories under conditions of nondifferential reinforcement. Although previous studies have demonstrated identification or discrimination of natural categories by nonhuman animals, subjects were typically differentially reinforced for their responses. The present findings demonstrate that chimpanzees can classify natural objects spontaneously and that such classifications may be similar to those that would be observed in human subjects.     

Spontaneous class grouping behavior by bonobos (Pan paniscus) and common chimpanzees (P. troglodytes)

Two experiments investigated spontaneous class grouping behavior by human-enculturated and language-reared bonobos (Pan paniscus) and common chimpanzees (P. troglodytes). In experiment 1, three chimpanzees ranging in age from 6 to 18 years were presented with six objects. The objects embodied three conditions: additive, multiplicative and disjoint classes. All chimpanzees spontaneously produced single- and two-category classifying. In experiment 2, six chimpanzees ranging in age from 6 to 21 years were presented with 12 objects in the same class conditions. Chimpanzees mainly produced single-category classifying. Their two-category classifying was more rudimentary than that found in experiment 1. Chimpanzees did not produce any three-category classifying which would be necessary to construct the hierarchies that humans begin to construct during early childhood. Received: 1 February 1999 / Accepted after revision: 9 August 1999     

Spontaneous transfer of matching by infant chimpanzees (Pan troglodytes)

Four infant chimpanzees learned a matching-to-sample task when only two training stimuli were used. They then spontaneously transferred the matching concept to novel items, including three-dimensional objects and fabric swatches, without any experimenter-provided differential feedback. These results support the view that the matching concept is broadly construed by chimpanzees from the beginning and does not depend upon explicit training.     

Cognitive and physiological markers of emotional awareness in chimpanzees (Pan troglodytes)

The ability to understand emotion in others is one of the most important factors involved in regulating social interactions in primates. Such emotional awareness functions to coordinate activity among group members, enable the formation of long-lasting individual relationships, and facilitate the pursuit of shared interests. Despite these important evolutionary implications, comparative studies of emotional processing in humans and great apes are practically nonexistent, constituting a major gap in our understanding of the extent to which emotional awareness has played an important role in shaping human behavior and societies. This paper presents the results of two experiments that examine chimpanzees' responses to emotional stimuli. First, changes in peripheral skin temperature were measured while subjects viewed three categories of emotionally negative video scenes; conspecifics being injected with needles (INJ), darts and needles alone (DART), and conspecific directing agonism towards the veterinarians (CHASE). Second, chimpanzees were required to use facial expressions to categorize emotional video scenes, i.e., favorite food and objects and veterinarian procedures, according to their positive and negative valence. With no prior training, subjects spontaneously matched the emotional videos to conspecific facial expressions according to their shared emotional meaning, indicating that chimpanzee facial expressions are processed emotionally, as are human expressions. Decreases in peripheral skin temperature, indicative of negative sympathetic arousal, were significantly lower when subjects viewed the INJ and DART videos, compared to the CHASE videos, indicating greater negative arousal when viewing conspecifics being injected with needles, and needles themselves, than when viewing conspecifics engaged in general agonism. Accepted after revision: 8 April 2001 ❚ Electronic Publication     

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.