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Whiten, A., Custance, D. M., Gomez, J. C., Teixidor, P., & Bard, K. A. (1996). Imitative learning of artificial fruit processing in children (Homo sapiens) and chimpanzees (Pan troglodytes). J Comp Psychol, 110(1), 3–14.
Abstract: Observational learning in chimpanzees and young children was investigated using an artificial fruit designed as an analog of natural foraging problems faced by primates. Each of 3 principal components could be removed in 2 alternative ways, demonstration of only one of which was watched by each subject. This permitted subsequent imitation by subjects to be distinguished from stimulus enhancement. Children aged 2-4 years evidenced imitation for 2 components, but also achieved demonstrated outcomes through their own techniques. Chimpanzees relied even more on their own techniques, but they did imitate elements of 1 component of the task. To our knowledge, this is the first experimental evidence of chimpanzee imitation in a functional task designed to simulate foraging behavior hypothesized to be transmitted culturally in the wild.
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Hayashi, M., & Matsuzawa, T. (2003). Cognitive development in object manipulation by infant chimpanzees. Anim. Cogn., 6(4), 225–233.
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.
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Matsuzawa, T. (2003). The Ai project: historical and ecological contexts. Anim. Cogn., 6(4), 199–211.
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.
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Xitco, M. J. J., Gory, J. D., & Kuczaj, S. A. 2nd. (2004). Dolphin pointing is linked to the attentional behavior of a receiver. Anim. Cogn., 7(4), 231–238.
Abstract: In 2001, Xitco et al. (Anim Cogn 4:115-123) described spontaneous behaviors in two bottlenose dolphins (Tursiops truncatus) that resembled pointing and gaze alternation. The dolphins' spontaneous behavior was influenced by the presence of a potential receiver, and the distance between the dolphin and the receiver. The present study adapted the technique of Call and Tomasello [(1994) J Comp Psychol 108:307-317], used with orangutans to test the effect of the receiver's orientation on pointing in these same dolphins. The dolphins directed more points and monitoring behavior at receivers whose orientation was consistent with attending to the dolphins. The results demonstrated that the dolphins' pointing and monitoring behavior, like that of apes and infants, was linked to the attentional behavior of the receiver.
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Stoinski, T. S., Wrate, J. L., Ure, N., & Whiten, A. (2001). Imitative learning by captive western lowland gorillas (Gorilla gorilla gorilla) in a simulated food-processing task. J Comp Psychol, 115(3), 272–281.
Abstract: Although field studies have suggested the existence of cultural transmission of foraging techniques in primates, identification of transmission mechanisms has remained elusive. To test experimentally for evidence of imitation in the current study, we exposed gorillas (Gorilla gorilla gorilla) to an artificial fruit foraging task designed by A. Whiten and D. M. Custance (1996). Gorillas (n=6) watched a human model remove a series of 3 defenses around a fruit. Each of the defenses was removed using 1 of 2 alternative techniques. Subsequent video analysis of gorillas' behavior showed a significant tendency to copy the observed technique on 1 of the individual defenses and the direction of removal on another defense. This is the first statistically reliable evidence of imitation in gorillas. Sequence of defense removal was not replicated. The gorillas' responses were most similar to those of chimpanzees.
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Heyes, C. M., & Dawson, G. R. (1990). A demonstration of observational learning in rats using a bidirectional control. Q J Exp Psychol B, 42(1), 59–71.
Abstract: Hungry rats observed a conspecific demonstrator pushing a single manipulandum, a joystick, to the right or to the left for food reward and were then allowed access to the joystick from a different orientation. The effects of right-pushing vs left-pushing observation experience on (1) response acquisition, (2) reversal of a left-right discrimination, and (3) responding in extinction, were examined. Rats that had observed left-pushing made more left responses during acquisition than rats that had observed right-pushing, and rats that had observed demonstrators pushing in the direction that had previously been reinforced took longer to reach criterion reversal and made more responses in extinction than rats that had observed demonstrators pushing in the opposite direction to that previously reinforced. These results provide evidence that rats are capable of learning a response, or a response-reinforcer contingency, through conspecific observation.
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Nguyen, N. H., Klein, E. D., & Zentall, T. R. (2005). Imitation of a two-action sequence by pigeons. Psychon Bull Rev, 12(3), 514–518.
Abstract: Developmental psychologists have described imitation as a process that suggests perspective-taking abilities. However, imitative behavior has been found in animals, which are generally not considered capable of taking the perspective of another. Previous studies with birds have demonstrated the imitation of a single response (sometimes referred to as action-level imitation). In the present experiment, we examined the extent to which pigeons would imitate an unfamiliar sequence of two behaviors (sometimes referred to as program-level imitation). Our results indicate that, although there are individual differences, pigeons show a significant tendency to match a demonstrated sequence of behavior involving, first, a response to a treadle (pecking at it or stepping on it) and, second, pushing aside a screen that blocks access to food (a left-vs.-right push).
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Akins, C. K., Klein, E. D., & Zentall, T. R. (2002). Imitative learning in Japanese quail (Coturnix japonica) using the bidirectional control procedure. Anim Learn Behav, 30(3), 275–281.
Abstract: In the bidirectional control procedure, observers are exposed to a conspecific demonstrator responding to a manipulandum in one of two directions (e.g., left vs. right). This procedure controls for socially mediated effects (the mere presence of a conspecific) and stimulus enhancement (attention drawn to a manipulandum by its movement), and it has the added advantage of being symmetrical (the two different responses are similar in topography). Imitative learning is demonstrated when the observers make the response in the direction that they observed it being made. Recently, however, it has been suggested that when such evidence is found with a predominantly olfactory animal, such as the rat, it may result artifactually from odor cues left on one side of the manipulandum by the demonstrator. In the present experiment, we found that Japanese quail, for which odor cues are not likely to play a role, also showed significant correspondence between the direction in which the demonstrator and the observer push a screen to gain access to reward. Furthermore, control quail that observed the screen move, when the movement of the screen was not produced by the demonstrator, did not show similar correspondence between the direction of screen movement observed and that performed by the observer. Thus, with the appropriate control, the bidirectional procedure appears to be useful for studying imitation in avian species.
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