Abstract: We investigated the behavioural and cognitive development of a captive male infant chimpanzee, Ayumu, raised by his mother, Ai. Here we report Ayumu's achievements up to the age of 2 years and 3 months, in the context of complex computer-controlled tasks. From soon after birth, Ayumu had been present during an experiment performed by his mother. The task consisted of two phases, a matching-to-sample task in which she received token rewards, and the insertion of these tokens into a vending machine to obtain food rewards. Ayumu himself received no reward or encouragement from humans for any of the actions he exhibited during the experiment. At the age of 9 months and 3 weeks, Ayumu performed his first matching-to-sample trial. At around 1 year and 3 months, he began to perform them consistently. Also during this period, he frequently stole food rewards from his mother. At 2 years and 3 months, Ayumu succeeded for the first time in inserting a token into the vending machine. Once he had succeeded in using a token, he performed both phases of the task in sequence 20 times consecutively. The infant's behaviour was not shaped by food rewards but by a strong motivation to copy his mother's behaviour. Our observations of Ayumu thus mirror the learning processes shown by wild chimpanzees.

Abstract: This paper aims to review a long-term research project exploring the chimpanzee mind within historical and ecological contexts. The Ai project began in 1978 and was directly inspired by preceding ape-language studies conducted in Western countries. However, in contrast with the latter, it has focused on the perceptual and cognitive capabilities of chimpanzees rather than communicative skills between humans and chimpanzees. In the original setting, a single chimpanzee faced a computer-controlled apparatus and performed various kinds of matching-to-sample discrimination tasks. Questions regarding the chimpanzee mind can be traced back to Wolfgang Koehler's work in the early part of the 20th century. Yet, Japan has its unique natural and cultural background: it is home to an indigenous primate species, the Japanese snow monkey. This fact has contributed to the emergence of two previous projects in the wild led by the late Kinji Imanishi and his students. First, the Koshima monkey project began in 1948 and became famous for its discovery of the cultural propagation of sweet-potato washing behavior. Second, pioneering work in Africa, starting in 1958, aimed to study great apes in their natural habitat. Thanks to the influence of these intellectual ancestors, the present author also undertook the field study of chimpanzees in the wild, focusing on tool manufacture and use. This work has demonstrated the importance of social and ecological perspectives even for the study of the mind. Combining experimental approaches with a field setting, the Ai project continues to explore cognition and behavior in chimpanzees, while its focus has shifted from the study of a single subject toward that of the community as a whole.

Abstract: We investigated the ability to perceive depth from shading, one of the pictorial depth cues, in three chimpanzee infants aged 4-10 months old, using a preferential reaching task commonly used to study pictorial depth perception in human infants. The chimpanzee infants reached significantly more to three-dimensional toys than to pictures thereof and more to the three-dimensional convex than to the concave. Furthermore, two of the three infants reached significantly more to the photographic convex than to the photographic concave. These infants also looked longer at the photographic convex than the concave. Our results suggest that chimpanzees perceive, at least as early as the latter half of the first year of life, pictorial depth defined by shading information. Photographic convexes contain richer information about pictorial depth (e.g., attached shadow, cast shadow, highlighted area, and global difference in brightness) than simple computer-graphic graded patterns. These cues together might facilitate the infants' perception of depth from shading.

Abstract: The ability of organisms to discriminate social signals, such as affective displays, using different sensory modalities is important for social communication. However, a major problem for understanding the evolution and integration of multimodal signals is determining how humans and animals attend to different sensory modalities, and these different modalities contribute to the perception and categorization of social signals. Using a matching-to-sample procedure, chimpanzees discriminated videos of conspecifics' facial expressions that contained only auditory or only visual cues by selecting one of two facial expression photographs that matched the expression category represented by the sample. Other videos were edited to contain incongruent sensory cues, i.e., visual features of one expression but auditory features of another. In these cases, subjects were free to select the expression that matched either the auditory or visual modality, whichever was more salient for that expression type. Results showed that chimpanzees were able to discriminate facial expressions using only auditory or visual cues, and when these modalities were mixed. However, in these latter trials, depending on the expression category, clear preferences for either the visual or auditory modality emerged. Pant-hoots and play faces were discriminated preferentially using the auditory modality, while screams were discriminated preferentially using the visual modality. Therefore, depending on the type of expressive display, the auditory and visual modalities were differentially salient in ways that appear consistent with the ethological importance of that display's social function.

Abstract: Auditory-visual processing of species-specific vocalizations was investigated in a female chimpanzee named Pan. The basic task was auditory-visual matching-to-sample, where Pan was required to choose the vocalizer from two test movies in response to a chimpanzee's vocalization. In experiment 1, movies of vocalizing and silent faces were paired as the test movies. The results revealed that Pan recognized the status of other chimpanzees whether they vocalized or not. In experiment 2, two different types of vocalizing faces of an identical individual were prepared as the test movies. Pan recognized the correspondence between vocalization types and faces. These results suggested that chimpanzees possess crossmodal representations of their vocalizations, as do humans. Together with the ability of vocal individual recognition, this ability might reflect chimpanzees' profound understanding of the status of other individuals.