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 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 study focuses on the development of spontaneous object manipulation in three infant chimpanzees during their first 2 years of life. The three infants were raised by their biological mothers who lived among a group of chimpanzees. A human tester conducted a series of cognitive tests in a triadic situation where mothers collaborated with the researcher during the testing of the infants. Four tasks were presented, taken from normative studies of cognitive development of Japanese infants: inserting objects into corresponding holes in a box, seriating nesting cups, inserting variously shaped objects into corresponding holes in a template, and stacking up wooden blocks. The mothers had already acquired skills to perform these manipulation tasks. The infants were free to observe the mothers' manipulative behavior from immediately after birth. We focused on object-object combinations that were made spontaneously by the infant chimpanzees, without providing food reinforcement for any specific behavior that the infants performed. The three main findings can be summarized as follows. First, there was precocious appearance of object-object combination in infant chimpanzees: the age of onset (8-11 months) was comparable to that in humans (around 10 months old). Second, object-object combinations in chimpanzees remained at a low frequency between 11 and 16 months, then increased dramatically at the age of approximately 1.5 years. At the same time, the accuracy of these object-object combinations also increased. Third, chimpanzee infants showed inserting behavior frequently and from an early age but they did not exhibit stacking behavior during their first 2 years of life, in clear contrast to human data.

Abstract: The experiments investigated how two adult captive chimpanzees learned to navigate in an automated interception task. They had to capture a visual target that moved predictably on a touch monitor. The aim of the study was to determine the learning stages that led to an efficient strategy of intercepting the target. The chimpanzees had prior training in moving a finger on a touch monitor and were exposed to the interception task without any explicit training. With a finger the subject could move a small “ball” at any speed on the screen toward a visual target that moved at a fixed speed either back and forth in a linear path or around the edge of the screen in a rectangular pattern. Initial ball and target locations varied from trial to trial. The subjects received a small fruit reinforcement when they hit the target with the ball. The speed of target movement was increased across training stages up to 38 cm/s. Learning progressed from merely chasing the target to intercepting the target by moving the ball to a point on the screen that coincided with arrival of the target at that point. Performance improvement consisted of reduction in redundancy of the movement path and reduction in the time to target interception. Analysis of the finger's movement path showed that the subjects anticipated the target's movement even before it began to move. Thus, the subjects learned to use the target's initial resting location at trial onset as a predictive signal for where the target would later be when it began moving. During probe trials, where the target unpredictably remained stationary throughout the trial, the subjects first moved the ball in anticipation of expected target movement and then corrected the movement to steer the ball to the resting target. Anticipatory ball movement in probe trials with novel ball and target locations (tested for one subject) showed generalized interception beyond the trained ball and target locations. The experiments illustrate in a laboratory setting the development of a highly complex and adaptive motor performance that resembles navigational skills seen in natural settings where predators intercept the path of moving prey.

Abstract: 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.