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	Author	Biro, D.; Inoue-Nakamura, N.; Tonooka, R.; Yamakoshi, G.; Sousa, C.; Matsuzawa, T.
	Title	Cultural innovation and transmission of tool use in wild chimpanzees: evidence from field experiments			Type	Journal Article
	Year	2003	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	6	Issue	4	Pages	213-223
	Keywords	Animals; Cooking and Eating Utensils; Culture; Diffusion of Innovation; Feeding Behavior/psychology; Female; Functional Laterality; Imitative Behavior; Male; Motor Skills; Pan troglodytes/psychology; Social Environment
	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.
	Address	Primate Research Institute, Kyoto University, Japan. dora.biro@zoology.oxford.ac.uk
	Corporate Author				Thesis
	Publisher	Springer-Verlag	Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:12898285			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2560
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	Author	Horner, V.; Whiten, A.; Flynn, E.; de Waal, F.B.M.
	Title	Faithful replication of foraging techniques along cultural transmission chains by chimpanzees and children			Type	Journal Article
	Year	2006	Publication	Proceedings of the National Academy of Sciences of the United States of America	Abbreviated Journal	Proc. Natl. Acad. Sci. U.S.A.
	Volume	103	Issue	37	Pages	13878-13883
	Keywords	Animals; Child, Preschool; Humans; Imitative Behavior; Pan troglodytes/psychology
	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.
	Address	Centre for Social Learning and Cognitive Evolution, School of Psychology, University of St. Andrews, Fife KY16 9JP, United Kingdom
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0027-8424	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16938863			Approved	no
	Call Number	refbase @ user @			Serial	159
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	Author	Whiten, A.; Custance, D.M.; Gomez, J.C.; Teixidor, P.; Bard, K.A.
	Title	Imitative learning of artificial fruit processing in children (Homo sapiens) and chimpanzees (Pan troglodytes)			Type	Journal Article
	Year	1996	Publication	Journal of comparative psychology (Washington, D.C. : 1983)	Abbreviated Journal	J Comp Psychol
	Volume	110	Issue	1	Pages	3-14
	Keywords	Animals; Child, Preschool; Discrimination Learning; Female; Food Preferences/psychology; Fruit; Humans; Imitative Behavior; Male; Mental Recall; Pan troglodytes/psychology; Social Environment
	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.
	Address	Scottish Primate Research Group, University of St. Andrews, Fife, Scotland. aw2@st-andrews.ac.uk
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0735-7036	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:8851548			Approved	no
	Call Number	refbase @ user @			Serial	744
Permanent link to this record



	Author	Call, J.; Carpenter, M.; Tomasello, M.
	Title	Copying results and copying actions in the process of social learning: chimpanzees (Pan troglodytes) and human children (Homo sapiens)			Type	Journal Article
	Year	2005	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	8	Issue	3	Pages	151-163
	Keywords	Animals; Child Behavior; Child, Preschool; Concept Formation; Female; Humans; Imitative Behavior; Learning; Male; Pan troglodytes; Problem Solving; Psychomotor Performance; Random Allocation; *Social Environment; Species Specificity
	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.
	Address	Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103, Leipzig, Germany. call@eva.mpg.de
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15490290			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2504
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	Author	Horner, V.; Whiten, A.
	Title	Causal knowledge and imitation/emulation switching in chimpanzees (Pan troglodytes) and children (Homo sapiens)			Type	Journal Article
	Year	2005	Publication	Animal cognition	Abbreviated Journal	Anim. Cogn.
	Volume	8	Issue	3	Pages	164-181
	Keywords	Animals; Child Behavior; Child, Preschool; Concept Formation; Female; Humans; Imitative Behavior; Learning; Male; Pan troglodytes; Problem Solving; Psychomotor Performance; *Social Environment; Species Specificity
	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.
	Address	Centre for Social Learning and Cognitive Evolution, School of Psychology, University of St Andrews, St Andrews, KY16 9JU, UK. vkh1@st-andrews.ac.uk
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15549502			Approved	no
	Call Number	refbase @ user @			Serial	732
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	Author	Nguyen, N.H.; Klein, E.D.; Zentall, T.R.
	Title	Imitation of a two-action sequence by pigeons			Type	Journal Article
	Year	2005	Publication	Psychonomic bulletin & review	Abbreviated Journal	Psychon Bull Rev
	Volume	12	Issue	3	Pages	514-518
	Keywords	Animals; Behavior, Animal; Cognition; Columbidae; Imitative Behavior; *Learning
	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).
	Address	University of Kentucky, Lexington, Kentucky 40506, USA
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1069-9384	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16235638			Approved	no
	Call Number	refbase @ user @			Serial	221
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	Author	Dorrance, B.R.; Zentall, T.R.
	Title	Imitation of conditional discriminations in pigeons (Columba livia)			Type	Journal Article
	Year	2002	Publication	Journal of comparative psychology (Washington, D.C. : 1983)	Abbreviated Journal	J Comp Psychol
	Volume	116	Issue	3	Pages	277-285
	Keywords	Animals; Behavior, Animal/physiology; Columbidae; Conditioning (Psychology)/physiology; Discrimination Learning/physiology; Habituation, Psychophysiologic; *Imitative Behavior; Light; Reinforcement (Psychology)
	Abstract	In the present experiments, the 2-action method was used to determine whether pigeons could learn to imitate a conditional discrimination. Demonstrator pigeons (Columba livia) stepped on a treadle in the presence of 1 light and pecked at the treadle in the presence of another light. Demonstration did not seem to affect acquisition of the conditional discrimination (Experiment 1) but did facilitate its reversal of the conditional discrimination (Experiments 2 and 3). The results suggest that pigeons are not only able to learn a specific behavior by observing another pigeon, but they can also learn under which circumstances to perform that behavior. The results have implications for proposed mechanisms of imitation in animals.
	Address	Department of Psychology, Augustana College, Rock Island, Illinois 61201, USA. psdorrance@augustana.edu
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0735-7036	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:12234078			Approved	no
	Call Number	refbase @ user @			Serial	240
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	Author	Stoinski, T.S.; Wrate, J.L.; Ure, N.; Whiten, A.
	Title	Imitative learning by captive western lowland gorillas (Gorilla gorilla gorilla) in a simulated food-processing task			Type	Journal Article
	Year	2001	Publication	Journal of comparative psychology (Washington, D.C. : 1983)	Abbreviated Journal	J Comp Psychol
	Volume	115	Issue	3	Pages	272-281
	Keywords	Animals; Behavior, Animal/physiology; Discrimination Learning; Feeding Behavior/physiology; Gorilla gorilla; *Imitative Behavior; Social Behavior
	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.
	Address	TECHlab, Zoo Atlanta, Georgia 30315, USA. stoinskit@mindspring.com
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0735-7036	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:11594496			Approved	no
	Call Number	refbase @ user @			Serial	738
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	Author	Biederman, G.B.; Robertson, H.A.; Vanayan, M.
	Title	Observational learning of two visual discriminations by pigeons: a within-subjects design			Type	Journal Article
	Year	1986	Publication	Journal of the experimental analysis of behavior	Abbreviated Journal	J Exp Anal Behav
	Volume	46	Issue	1	Pages	45-49
	Keywords	Animals; Attention; Columbidae; Conditioning, Operant; Cues; Discrimination Learning; Imitative Behavior; Male; Visual Perception
	Abstract	Pigeon's observational learning of successive visual discrimination was studied using within-subject comparisons of data from three experimental conditions. Two pairs of discriminative stimuli were used; each bird was exposed to two of the three experimental conditions, with different pairs of stimuli used in a given bird's two conditions. In one condition, observers were exposed to visual discriminative stimuli only. In a second condition, subjects were exposed to a randomly alternating sequence of two stimuli where the one that would subsequently be used as S+ was paired with the operation of the grain magazine. In a third experimental condition, subjects were exposed to the performance of a conspecific in the operant discrimination procedure. After exposures to conspecific performances, there was facilitation of discriminative learning, relative to that which followed exposures to stimulus and reinforcement sequences or exposures to stimulus sequences alone. Exposure to stimulus and food-delivery sequences enhanced performance relative to exposure to stimulus sequences alone. The differential effects of these three types of exposure were not attributable to order effects or to task difficulty; rather, they clearly were due to the type of exposure.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0022-5002	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:3746187			Approved	no
	Call Number	refbase @ user @			Serial	853
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	Author	Akins, C.K.; Klein, E.D.; Zentall, T.R.
	Title	Imitative learning in Japanese quail (Coturnix japonica) using the bidirectional control procedure			Type	Journal Article
	Year	2002	Publication	Animal learning & behavior	Abbreviated Journal	Anim Learn Behav
	Volume	30	Issue	3	Pages	275-281
	Keywords	Animals; Attention; Behavior, Animal; Coturnix; Discrimination Learning; Imitative Behavior; Male; Smell
	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.
	Address	University of Kentucky, Lexington, Kentucky 40506-0044, USA
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0090-4996	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:12391793			Approved	no
	Call Number	refbase @ user @			Serial	239
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