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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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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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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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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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Horner, V., Whiten, A., Flynn, E., & de Waal, F. B. M. (2006). Faithful replication of foraging techniques along cultural transmission chains by chimpanzees and children. Proc. Natl. Acad. Sci. U.S.A., 103(37), 13878–13883.
Abstract: Observational studies of wild chimpanzees (Pan troglodytes) have revealed population-specific differences in behavior, thought to represent cultural variation. Field studies have also reported behaviors indicative of cultural learning, such as close observation of adult skills by infants, and the use of similar foraging techniques within a population over many generations. Although experimental studies have shown that chimpanzees are able to learn complex behaviors by observation, it is unclear how closely these studies simulate the learning environment found in the wild. In the present study we have used a diffusion chain paradigm, whereby a behavior is passed from one individual to the next in a linear sequence in an attempt to simulate intergenerational transmission of a foraging skill. Using a powerful three-group, two-action methodology, we found that alternative methods used to obtain food from a foraging device (“lift door” versus “slide door”) were accurately transmitted along two chains of six and five chimpanzees, respectively, such that the last chimpanzee in the chain used the same method as the original trained model. The fidelity of transmission within each chain is remarkable given that several individuals in the no-model control group were able to discover either method by individual exploration. A comparative study with human children revealed similar results. This study is the first to experimentally demonstrate the linear transmission of alternative foraging techniques by non-human primates. Our results show that chimpanzees have a capacity to sustain local traditions across multiple simulated generations.
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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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