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Horowitz, A. C. (2003). Do humans ape? Or do apes human? Imitation and intention in humans (Homo sapiens) and other animals. J Comp Psychol, 117(3), 325–336.
Abstract: A. Whiten, D. M. Custance, J.-C. Gomez, P. Teixidor, and K. A. Bard (1996) tested chimpanzees' (Pan troglodytes) and human children's (Homo sapiens) skills at imitation with a 2-action test on an “artificial fruit.” Chimpanzees imitated to a restricted degree; children were more thoroughly imitative. Such results prompted some to assert that the difference in imitation indicates a difference in the subjects' understanding of the intentions of the demonstrator (M. Tomasello, 1996). In this experiment, 37 adult human subjects were tested with the artificial fruit. Far from being perfect imitators, the adults were less imitative than the children. These results cast doubt on the inference from imitative performance to an ability to understand others' intentions. The results also demonstrate how any test of imitation requires a control group and attention to the level of behavioral analysis.
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Whiten, A. (1998). Imitation of the sequential structure of actions by chimpanzees (Pan troglodytes). J Comp Psychol, 112(3), 270–281.
Abstract: Imitation was studied experimentally by allowing chimpanzees (Pan troglodytes) to observe alternative patterns of actions for opening a specially designed “artificial fruit.” Like problematic foods primates deal with naturally, with the test fruit several defenses had to be removed to gain access to an edible core, but the sequential order and method of defense removal could be systematically varied. Each subject repeatedly observed 1 of 2 alternative techniques for removing each defense and 1 of 2 alternative sequential patterns of defense removal. Imitation of sequential organization emerged after repeated cycles of demonstration and attempts at opening the fruit. Imitation in chimpanzees may thus have some power to produce cultural convergence, counter to the supposition that individual learning processes corrupt copied actions. Imitation of sequential organization was accompanied by imitation of some aspects of the techniques that made up the sequence.
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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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Stoinski, T. S., & Whiten, A. (2003). Social learning by orangutans (Pongo abelii and Pongo pygmaeus) in a simulated food-processing task. J Comp Psychol, 117(3), 272–282.
Abstract: Increasing evidence for behavioral differences between populations of primates has created a resurgence of interest in examining mechanisms of information transfer between individuals. The authors examined the social transmission of information in 15 captive orangutans (Pongo abelii and Pongo pygmaeus) using a simulated food-processing task. Experimental subjects were shown 1 of 2 methods for removing a suite of defenses on an “artificial fruit.” Control subjects were given no prior exposure before interacting with the fruit. Observing a model provided a functional advantage in the task, as significantly more experimental than control subjects opened the fruit. Within the experimental groups, the authors found a trend toward differences in the actual behaviors used to remove 1 of the defenses. Results support observations from the wild implying horizontal transfer of information in orangutans and show that a number of social learning processes are likely to be involved in the transfer of knowledge in this species.
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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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Whiten, A. (2005). The second inheritance system of chimpanzees and humans. Nature, 437(7055), 52–55.
Abstract: Half a century of dedicated field research has brought us from ignorance of our closest relatives to the discovery that chimpanzee communities resemble human cultures in possessing suites of local traditions that uniquely identify them. The collaborative effort required to establish this picture parallels the one set up to sequence the chimpanzee genome, and has revealed a complex social inheritance system that complements the genetic picture we are now developing.
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Janik, V. M. (2000). Whistle matching in wild bottlenose dolphins (Tursiops truncatus). Science, 289(5483), 1355–1357.
Abstract: Dolphin communication is suspected to be complex, on the basis of their call repertoires, cognitive abilities, and ability to modify signals through vocal learning. Because of the difficulties involved in observing and recording individual cetaceans, very little is known about how they use their calls. This report shows that wild, unrestrained bottlenose dolphins use their learned whistles in matching interactions, in which an individual responds to a whistle of a conspecific by emitting the same whistle type. Vocal matching occurred over distances of up to 580 meters and is indicative of animals addressing each other individually.
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Henning, J. M., & Zentall, T. R. (1981). Imitation, social facilitation, and the effects of ACTH 4-10 on rats' bar-pressing behavior. Am J Psychol, 94(1), 125–134.
Abstract: The effects of ACTH 4-10 on rats' imitation learning was examined during the acquisition and extinction of a bar-press response for water reinforcement. Rats were exposed to either a bar-pressing conspecific (OB), an experimentally naive conspecific (ON), or an empty box (OE) during bar-press acquisition. In a factorial design, each rat was then exposed to one of the same three conditions during extinction. An 80 mcg dose of ACTH 4-10 was administered to half of the rats in each group prior to observation. Performance differences during acquisition were generally small, but significant performance differences during extinction were found. Social facilitation was indicated by the finding that rats extinguished in the presence of a conspecific exhibited significantly greater resistance to extinction than rats extinguished in the presence of an empty box. An imitation effect was also found. Rats that observed a bar-pressing conspecific during both acquisition and extinction (group OB-OB) showed significantly greater resistance top extinction than did groups OB-ON, CB-OE, or OE-OE. There were no significant effects of the hormone, however, relative to saline controls.
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Akins, C. K., & Zentall, T. R. (1996). Imitative learning in male Japanese quail (Coturnix japonica) using the two-action method. J Comp Psychol, 110(3), 316–320.
Abstract: The study of imitative learning in animals has suffered from the presence of a number of confounding motivational and attentional factors (e.g., social facilitation and stimulus enhancement). The two-action method avoids these problems by exposing observers to demonstrators performing a response (e.g., operating a treadle) using 1 of 2 distinctive topographies (e.g., by pecking or by stepping). Japanese quail (Coturnix japonica) observers exposed to conspecific demonstrators showed a high correlation between the topography of the response they observed and the response they performed. These data provide strong evidence for the existence of true imitative learning in an active, precocious bird under conditions that control for alternative accounts.
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Dorrance, B. R., & Zentall, T. R. (2001). Imitative learning in Japanese quail (Coturnix japonica) depends on the motivational state of the observer quail at the time of observation. J Comp Psychol, 115(1), 62–67.
Abstract: The 2-action method was used to examine whether imitative learning in Japanese quail (Coturnix japonica) depends on the motivational state of the observer quail at the time of observation of the demonstrated behavior. Two groups of observers were fed before observation (satiated groups), whereas 2 other groups of observers were deprived of food before observation (hungry groups). Quail were tested either immediately following observation or after a 30-min delay. Results indicated that quail in the hungry groups imitated, whereas those in the satiated groups did not, regardless of whether their test was immediate or delayed. The results suggest that observer quail may not learn (through observation) behavior that leads to a reinforcer for which they are unmotivated at the time of test. In addition, the results show that quail are able to delay the performance of a response acquired through observation (i.e., they show deferred imitation).
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