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.     

Finger drawing by infant chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

We introduced a new technique to investigate the development of scribbling in very young infants. We tested three infant chimpanzees to compare the developmental processes of scribbling between humans and chimpanzees. While human infants start to scribble on paper at around the age of 18 months, our 13- to 23-month-old infant chimpanzees had never been observed scribbling prior to this study. We used a notebook computer with a touch-sensitive screen. This apparatus was able to record the location of the subjects touches on the screen. Each touch generated a fingertip-sized dot at the corresponding on-screen location. During spontaneous interactions with this apparatus, all three infants and two mother chimpanzees left scribbles with their fingers on the screen. The scribbles contained not only simple dots or short lines, but also curves and hook-like lines or loops, most of which were observed in the instrumental drawings of adult chimpanzees. The results suggest that perceptual-motor control for finger drawing develops in infant chimpanzees. Two of the infants performed their first scribble with a marker on paper at the age of 20–23 months. Just prior to this, they showed a rapid increase in combinatory manipulation of objects. These findings suggest that the development of combinatory manipulation of objects as well as that of perceptual-motor control may be necessary for the emergence of instrumental drawing on paper.     

Cultural innovation and transmission of tool use in wild chimpanzees: evidence from field experiments

Chimpanzees (Pan troglodytes) are the most proficient and versatile users of tools in the wild. How such skills become integrated into the behavioural repertoire of wild chimpanzee communities is investigated here by drawing together evidence from three complementary approaches in a group of oil-palm nut- (Elaeis guineensis) cracking chimpanzees at Bossou, Guinea. First, extensive surveys of communities adjacent to Bossou have shown that population-specific details of tool use, such as the selection of species of nuts as targets for cracking, cannot be explained purely on the basis of ecological differences. Second, a 16-year longitudinal record tracing the development of nut-cracking in individual chimpanzees has highlighted the importance of a critical period for learning (3–5 years of age), while the similar learning contexts experienced by siblings have been found to result in near-perfect (13 out of 14 dyads) inter-sibling correspondence in laterality. Third, novel data from field experiments involving the introduction of unfamiliar species of nuts to the Bossou group illuminates key aspects of both cultural innovation and transmission. We show that responses of individuals toward the novel items differ markedly with age, with juveniles being the most likely to explore. Furthermore, subjects are highly specific in their selection of conspecifics as models for observation, attending to the nut-cracking activities of individuals in the same age group or older, but not younger than themselves. Together with the phenomenon of inter-community migration, these results demonstrate a mechanism for the emergence of culture in wild chimpanzees.     

The use of tokens as rewards and tools by chimpanzees (Pan troglodytes)

This paper explores the effectiveness of token rewards in maintaining chimpanzees (Pan troglodytes) in working at intellectually costly tasks, and studies the "saving" behavior of the subjects, investigating the factors that can condition it. Two experiments were run. Tokens were introduced as rewards in a matching-to-sample task and used as exchange tools for food by three adult female chimpanzees. Subjects' performances were maintained at constant high levels of accuracy, suggesting that the tokens were almost equivalent to direct food rewards. The results also showed the emergence of saving behavior. The subjects spontaneously saved the tokens during the matching-to-sample task before exchanging them for food. The chimpanzees also learned a new symbolic discrimination task, with tokens as the reward. During this learning process a rarely reported phenomenon emerged: one of the subjects showed symmetry, a form of stimulus equivalence. Accepted after revision: 7 August 2001 Electronic Publication     

Spatial construction skills of chimpanzees (Pan troglodytes) and young human children (Homo sapiens sapiens)

Spatial construction tasks are basic tests of visual‐spatial processing. Two studies have assessed spatial construction skills in chimpanzees (Pan troglodytes) and young children (Homo sapiens sapiens) with a block modelling task. Study 1a subjects were three young chimpanzees and five adult chimpanzees. Study 1b subjects were 30 human children belonging to five age groups (24, 30, 36, 42, 48 months). Subjects were given three model constructions to reproduce: Line, Cross‐Stack and Arch, which differed in type and number of spatial relations and dimensions, but required comparable configurational understanding. Subjects’ constructions were rated for accuracy. Our results show that: (1) chimpanzees are relatively advanced in constructing in the vertical dimension; (2) Among chimpanzees only adults make accurate copies of constructions; (3) Chimpanzees do not develop in the direction of constructing in two dimensions as human children do starting from age 30 months. The pattern of development of construction skills in chimpanzees partially diverges from that of human children and indicates that spatial analysis and spatial representation are partially different in the two species.     

Plasticity of ability to form cross‐modal representations in infant Japanese macaques

In a previous study, Adachi, Kuwahata, Fujita, Tomonaga & Matsuzawa demonstrated that infant Japanese macaques (Macaca fuscata) form cross‐modal representations of conspecifics but not of humans. However, because the subjects in the experiment were raised in a large social group and had considerably less exposure to humans than to conspecifics, it was an open question whether their lack of cross‐modal representation of humans simply reflected their lower levels of exposure to humans or was caused by some innate restrictions on the ability. To answer the question, we used the same procedure but tested infant Japanese macaques with more extensive experience of humans in daily life. Briefly, we presented monkeys with a photograph of either a monkey or a human face on an LCD monitor after playing a vocalization of one of these two species. The subjects looked at the monitor longer when a voice and a face were mismatched than when they were matched, irrespective of whether the preceding vocalization was a monkey's or a human's. This suggests that once monkeys have extensive experience with humans, they will form a cross‐modal representation of humans as well as of conspecifics.     

Tool-composite reuse in wild chimpanzees (Pan troglodytes): archaeologically invisible steps in the technological evolution of early hominins?

Recent etho-archaeological studies of stone-tool use by wild chimpanzees have contributed valuable data towards elucidating the variables that influenced the emergence and development of the first lithic industries among Plio-Pleistocene hominins. Such data help to identify potential behaviours entailed in the first percussive technologies that are invisible in archaeological records. The long-term research site of Bossou in Guinea features a unique chimpanzee community whose members systematically use portable stones as hammers and anvils to crack open nuts in natural as well as in field experimental settings. Here we present the first analysis of repeated reuse of the same tool-composites in wild chimpanzees. Data collected over 5 years of experimental nut-cracking sessions at an “outdoor laboratory” site were assessed for the existence of systematic patterns in the selection of tool-composites, at group and at individual levels. Chimpanzees combined certain stones as hammer and anvil more often than expected by chance, even when taking into account preferences for individual stones by themselves. This may reflect an ability to recognise the nut-cracker as a single tool (composed of two elements, but functional only as a whole), as well as discrimination of tool quality-effectiveness. Through repeatedly combining the same pairs of stones—whether due to preferences for particular composites or for the two elements independently—tool-users may amplify use-wear traces and increase the likelihood of fracturing the stones, and thus of detaching pieces by battering.     

Social influences on ant-dipping acquisition in the wild chimpanzees (Pan troglodytes verus) of Bossou,Guinea, West Africa

We currently have little understanding of the influence of learning opportunity, whether social or environmental, and maternal role on tool-use acquisition in young wild chimpanzees. This study aims to fill this gap by focusing on the acquisition of ant-dipping among chimpanzees of Bossou, Guinea. Ant-dipping is a hazardous tool-use behaviour aimed at army ants (Dorylus spp.). Bossou chimpanzees target these ants both at nests (high risk) and trails (low risk) and employ two techniques to consume them: direct mouthing and pull-through. We present data for 13 mother–offspring pairs (1–10 years old). Mothers with young ≤5 years old dipped significantly more often at trails than at nests, thus minimizing the risk posed to themselves and their young. Infants thus benefited from better conditions to observe and practice ant-dipping. Mothers also varied greatly in their percent time spent ant-dipping and offspring differed in their learning opportunity. Our results suggest that high opportunity young started to observe and perform ant-dipping sooner and were better at ant-dipping than low opportunity young. Although mothers and weaned offspring correlated positively in their percent time spent dipping and proficiency, they did not match in technique used or tool length. Finally, we propose that the learning trajectory of young may predict individual and sex differences in adulthood. This study demonstrates the important role of mothers and learning opportunity in the acquisition of a hazardous tool-use behaviour and suggests that chimpanzee material culture is a product of a complex interaction between social processes and ecological factors.     

Development of saccade target selection in infants.

To investigate the development of the target selection process for saccade generation, saccade latencies toward one of two targets were compared with those toward a target presented alone, under conditions where the timing of fixation stimulus offset was varied. 12 2-mo.-olds (7 boys and 5 girls), 12 3-mo.-olds (6 boys and 6 girls), and 12 5-mo.-olds (8 boys and 4 girls) participated. The latencies were longer when two saccade targets were presented on the left and right sides of the display (Double-target condition) than when one target was presented either on the left or right side of the display (Single-target condition). The difference between the Single and Double-target conditions tended to be larger in 2-mo. old infants than in 5-mo.-old infants when a central fixation target remained on after the onset of peripheral stimuli (Overlap condition). However, there was no difference between the 2-mo.-olds and the 5-mo.-olds when the fixation target was turned off before the peripheral stimuli were presented (Gap condition). The results provided evidence of the maturational development of a target selection process for saccade generation during 2 to 5 mo. of age.     

Correlation between menstrual cycle and cognitive performance in a chimpanzee (Pan troglodytes)

Extensive research on human subjects has tried to investigate whether there is a correlation between cognitive performance and the menstrual cycle. Less is known about the relationship between the menstrual cycle and task performance in other cognitive animals. We test whether the secretion of a sex hormone [luteinizing hormone(LH)] influences the performance of cognitive tasks by a female chimpanzee (Pan troglodytes) who is part of a long-term cognition research program. We focus on two cognitive tasks: an "easy task," which consists of simple numerical ordering, and a "difficult task," which combines numerical ordering with memorizing the numerals' spatial location. Data on the performance of these cognitive tasks, urine samples, and sexual swelling over six menstrual cycles showed that the chimpanzee's performance accuracy decreased and that the intertrial interval was longer during the LH-surge of the menstrual cycle, but only for the performance of the difficult task. These performance attributes seem to reflect a decrease in attention or motivation during ovulation. In summary, the cognitive performance of a chimpanzee was disturbed by hormonal changes despite her long-term experience in the tasks.