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Bering, J. M. (2004). A critical review of the “enculturation hypothesis”: the effects of human rearing on great ape social cognition. Anim. Cogn., 7(4), 201–212.
Abstract: Numerous investigators have argued that early ontogenetic immersion in sociocultural environments facilitates cognitive developmental change in human-reared great apes more characteristic of Homo sapiens than of their own species. Such revamping of core, species-typical psychological systems might be manifest, according to this argument, in the emergence of mental representational competencies, a set of social cognitive skills theoretically consigned to humans alone. Human-reared great apes' capacity to engage in “true imitation,” in which both the means and ends of demonstrated actions are reproduced with fairly high rates of fidelity, and laboratory great apes' failure to do so, has frequently been interpreted as reflecting an emergent understanding of intentionality in the former. Although this epigenetic model of the effects of enculturation on social cognitive systems may be well-founded and theoretically justified in the biological literature, alternative models stressing behavioral as opposed to representational change have been largely overlooked. Here I review some of the controversy surrounding enculturation in great apes, and present an alternative nonmentalistic version of the enculturation hypothesis that can also account for enhanced imitative performance on object-oriented problem-solving tasks in human-reared animals.
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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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Zentall, T. R. (2004). Action imitation in birds. Learn Behav, 32(1), 15–23.
Abstract: Action imitation, once thought to be a behavior almost exclusively limited to humans and the great apes, surprisingly also has been found in a number of bird species. Because imitation has been viewed by some psychologists as a form of intelligent behavior, there has been interest in how it is distributed among animal species. Although the mechanisms responsible for action imitation are not clear, we are now at least beginning to understand the conditions under which it occurs. In this article, I try to identify and differentiate the various forms of socially influenced behavior (species-typical social reactions, social effects on motivation, social effects on perception, socially influenced learning, and action imitation) and explain why it is important to differentiate imitation from other forms of social influence. I also examine some of the variables that appear to be involved in the occurrence of imitation. Finally, I speculate about why a number of bird species, but few mammal species, appear to imitate.
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Sousa, C., Okamoto, S., & Matsuzawa, T. (2003). Behavioural development in a matching-to-sample task and token use by an infant chimpanzee reared by his mother. Anim. Cogn., 6(4), 259–267.
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.
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Fripp, D., Owen, C., Quintana-Rizzo, E., Shapiro, A., Buckstaff, K., Jankowski, K., et al. (2005). Bottlenose dolphin (Tursiops truncatus) calves appear to model their signature whistles on the signature whistles of community members. Anim. Cogn., 8(1), 17–26.
Abstract: Bottlenose dolphins are unusual among non-human mammals in their ability to learn new sounds. This study investigates the importance of vocal learning in the development of dolphin signature whistles and the influence of social interactions on that process. We used focal animal behavioral follows to observe six calves in Sarasota Bay, Fla., recording their social associations during their first summer, and their signature whistles during their second. The signature whistles of five calves were determined. Using dynamic time warping (DTW) of frequency contours, the calves' signature whistles were compared to the signature whistles of several sets of dolphins: their own associates, the other calves' associates, Tampa Bay dolphins, and captive dolphins. Whistles were considered similar if their DTW similarity score was greater than those of 95% of the whistle comparisons. Association was defined primarily in terms of time within 50 m of the mother/calf pair. On average, there were six dolphins with signature whistles similar to the signature whistles of each of the calves. These were significantly more likely to be Sarasota Bay resident dolphins than non-Sarasota dolphins, and (though not significantly) more likely to be dolphins that were within 50 m of the mother and calf less than 5% of the time. These results suggest that calves may model their signature whistles on the signature whistles of members of their community, possibly community members with whom they associate only rarely.
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Horner, V., & Whiten, A. (2005). Causal knowledge and imitation/emulation switching in chimpanzees (Pan troglodytes) and children (Homo sapiens). Anim. Cogn., 8(3), 164–181.
Abstract: This study explored whether the tendency of chimpanzees and children to use emulation or imitation to solve a tool-using task was a response to the availability of causal information. Young wild-born chimpanzees from an African sanctuary and 3- to 4-year-old children observed a human demonstrator use a tool to retrieve a reward from a puzzle-box. The demonstration involved both causally relevant and irrelevant actions, and the box was presented in each of two conditions: opaque and clear. In the opaque condition, causal information about the effect of the tool inside the box was not available, and hence it was impossible to differentiate between the relevant and irrelevant parts of the demonstration. However, in the clear condition causal information was available, and subjects could potentially determine which actions were necessary. When chimpanzees were presented with the opaque box, they reproduced both the relevant and irrelevant actions, thus imitating the overall structure of the task. When the box was presented in the clear condition they instead ignored the irrelevant actions in favour of a more efficient, emulative technique. These results suggest that emulation is the favoured strategy of chimpanzees when sufficient causal information is available. However, if such information is not available, chimpanzees are prone to employ a more comprehensive copy of an observed action. In contrast to the chimpanzees, children employed imitation to solve the task in both conditions, at the expense of efficiency. We suggest that the difference in performance of chimpanzees and children may be due to a greater susceptibility of children to cultural conventions, perhaps combined with a differential focus on the results, actions and goals of the demonstrator.
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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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Subiaul, F., Cantlon, J. F., Holloway, R. L., & Terrace, H. S. (2004). Cognitive imitation in rhesus macaques. Science, 305(5682), 407–410.
Abstract: Experiments on imitation typically evaluate a student's ability to copy some feature of an expert's motor behavior. Here, we describe a type of observational learning in which a student copies a cognitive rule rather than a specific motor action. Two rhesus macaques were trained to respond, in a prescribed order, to different sets of photographs that were displayed on a touch-sensitive monitor. Because the position of the photographs varied randomly from trial to trial, sequences could not be learned by motor imitation. Both monkeys learned new sequences more rapidly after observing an expert execute those sequences than when they had to learn new sequences entirely by trial and error.
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Call, J., Carpenter, M., & Tomasello, M. (2005). Copying results and copying actions in the process of social learning: chimpanzees (Pan troglodytes) and human children (Homo sapiens). Anim. Cogn., 8(3), 151–163.
Abstract: There is currently much debate about the nature of social learning in chimpanzees. The main question is whether they can copy others' actions, as opposed to reproducing the environmental effects of these actions using their own preexisting behavioral strategies. In the current study, chimpanzees (Pan troglodytes) and human children (Homo sapiens) were shown different demonstrations of how to open a tube-in both cases by a conspecific. In different experimental conditions, demonstrations consisted of (1) action only (the actions necessary to open the tube without actually opening it); (2) end state only (the open tube, without showing any actions); (3) both of these components (in a full demonstration); or (4) neither of these components (in a baseline condition). In the first three conditions subjects saw one of two different ways that the tube could open (break in middle; caps off ends). Subjects' behavior in each condition was assessed for how often they opened the tube, how often they opened it in the same location as the demonstrator, and how often they copied the demonstrator's actions or style of opening the tube. Whereas chimpanzees reproduced mainly the environmental results of the demonstrations (emulation), human children often reproduced the demonstrator's actions (imitation). Because the procedure used was similar in many ways to the procedure that Meltzoff (Dev Psych 31:1, 1995) used to study the understanding of others' unfulfilled intentions, the implications of these findings with regard to chimpanzees' understanding of others' intentions are also discussed.
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Biro, D., Inoue-Nakamura, N., Tonooka, R., Yamakoshi, G., Sousa, C., & Matsuzawa, T. (2003). Cultural innovation and transmission of tool use in wild chimpanzees: evidence from field experiments. Anim. Cogn., 6(4), 213–223.
Abstract: 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.
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