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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	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	Nicol, C.J.
	Title	Development, direction, and damage limitation: social learning in domestic fowl			Type	Journal Article
	Year	2004	Publication	Learning & behavior : a Psychonomic Society publication	Abbreviated Journal	Learn Behav
	Volume	32	Issue	1	Pages	72-81
	Keywords	Adaptation, Psychological; Age Factors; Animals; Behavior, Animal; Chickens; Feeding Behavior; Food Preferences; Imitative Behavior; Imprinting (Psychology); Learning; Maternal Behavior; Reinforcement (Psychology); Social Environment; *Social Facilitation
	Abstract	This review highlights two areas of particular interest in the study of social learning in fowl. First, the role of social learning in the development of feeding and foraging behavior in young chicks and older birds is described. The role of the hen as a demonstrator and possible teacher is considered, and the subsequent social influence of brood mates and other companions on food avoidance and food preference learning is discussed. Second, the way in which work on domestic fowl has contributed to an understanding of the importance of directed social learning is examined. The well-characterized hierarchical social organization of small chicken flocks has been used to design studies which demonstrate that the probability of social transmission is strongly influenced by social relationships between birds. The practical implications of understanding the role of social learning in the spread of injurious behaviors in this economically important species are briefly considered.
	Address	Department of Clinical Veterinary Science, University of Bristol, Langford, Bristol, England. c.j.nicol@bristol.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	1543-4494	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15161142			Approved	no
	Call Number	refbase @ user @			Serial	75
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	Author	Zentall, T.R.
	Title	Action imitation in birds			Type	Journal Article
	Year	2004	Publication	Learning & behavior : a Psychonomic Society publication	Abbreviated Journal	Learn Behav
	Volume	32	Issue	1	Pages	15-23
	Keywords	Adaptation, Psychological; Animals; Birds; Imitative Behavior; Imprinting (Psychology); Learning; Motivation; Psychological Theory; Social Environment; *Social Facilitation; Vocalization, Animal
	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.
	Address	Department of Psychology, University of Kentucky, Lexington, Kentucky 40506, USA. zentall@uky.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	1543-4494	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15161137			Approved	no
	Call Number	refbase @ user @			Serial	230
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	Author	Whiten, A.; Horner, V.; Litchfield, C.A.; Marshall-Pescini, S.
	Title	How do apes ape?			Type	Journal Article
	Year	2004	Publication	Learning & Behavior	Abbreviated Journal	Learn. Behav.
	Volume	32	Issue	1	Pages	36-52
	Keywords	Adaptation, Psychological; Animals; Behavior, Animal; Hominidae/psychology; Imitative Behavior; Imprinting (Psychology); Learning; Psychological Theory; Social Environment; *Social Facilitation
	Abstract	In the wake of telling critiques of the foundations on which earlier conclusions were based, the last 15 years have witnessed a renaissance in the study of social learning in apes. As a result, we are able to review 31 experimental studies from this period in which social learning in chimpanzees, gorillas, and orangutans has been investigated. The principal question framed at the beginning of this era, Do apes ape? has been answered in the affirmative, at least in certain conditions. The more interesting question now is, thus, How do apes ape? Answering this question has engendered richer taxonomies of the range of social-learning processes at work and new methodologies to uncover them. Together, these studies suggest that apes ape by employing a portfolio of alternative social-learning processes in flexibly adaptive ways, in conjunction with nonsocial learning. We conclude by sketching the kind of decision tree that appears to underlie the deployment of these alternatives.
	Address	Centre for Social Learning and Cognitive Evolution, Scottish Primate Research Group, School of Psychology, University of St. Andrews, St. Andrews, Fife, Scotland. a.whiten@st-and.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	1543-4494	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15161139			Approved	no
	Call Number	refbase @ user @			Serial	734
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	Author	Fragaszy, D.; Visalberghi, E.
	Title	Socially biased learning in monkeys			Type	Journal Article
	Year	2004	Publication	Learning & behavior : a Psychonomic Society publication	Abbreviated Journal	Learn Behav
	Volume	32	Issue	1	Pages	24-35
	Keywords	Adaptation, Psychological; Animal Communication; Animals; Behavior, Animal; Feeding Behavior/psychology; Food Preferences/psychology; Haplorhini/psychology; Imitative Behavior; Imprinting (Psychology); Learning; Social Environment; Social Facilitation
	Abstract	We review socially biased learning about food and problem solving in monkeys, relying especially on studies with tufted capuchin monkeys (Cebus apella) and callitrichid monkeys. Capuchin monkeys most effectively learn to solve a new problem when they can act jointly with an experienced partner in a socially tolerant setting and when the problem can be solved by direct action on an object or substrate, but they do not learn by imitation. Capuchin monkeys are motivated to eat foods, whether familiar or novel, when they are with others that are eating, regardless of what the others are eating. Thus, social bias in learning about foods is indirect and mediated by facilitation of feeding. In most respects, social biases in learning are similar in capuchins and callitrichids, except that callitrichids provide more specific behavioral cues to others about the availability and palatability of foods. Callitrichids generally are more tolerant toward group members and coordinate their activity in space and time more closely than capuchins do. These characteristics support stronger social biases in learning in callitrichids than in capuchins in some situations. On the other hand, callitrichids' more limited range of manipulative behaviors, greater neophobia, and greater sensitivity to the risk of predation restricts what these monkeys learn in comparison with capuchins. We suggest that socially biased learning is always the collective outcome of interacting physical, social, and individual factors, and that differences across populations and species in social bias in learning reflect variations in all these dimensions. Progress in understanding socially biased learning in nonhuman species will be aided by the development of appropriately detailed models of the richly interconnected processes affecting learning.
	Address	Psychology Department, University of Georgia, Athens, Georgia 30602, USA. doree@uga.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	1543-4494	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15161138			Approved	no
	Call Number	refbase @ user @			Serial	828
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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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	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	Zentall, T.R.
	Title	Imitation: definitions, evidence, and mechanisms			Type	Journal Article
	Year	2006	Publication	Animal cognition	Abbreviated Journal	Anim. Cogn.
	Volume	9	Issue	4	Pages	335-353
	Keywords	Adaptation, Psychological; Animals; Behavior, Animal; Imitative Behavior; Learning; Motivation; Social Environment; Transfer (Psychology)
	Abstract	Imitation can be defined as the copying of behavior. To a biologist, interest in imitation is focused on its adaptive value for the survival of the organism, but to a psychologist, the mechanisms responsible for imitation are the most interesting. For psychologists, the most important cases of imitation are those that involve demonstrated behavior that the imitator cannot see when it performs the behavior (e.g., scratching one's head). Such examples of imitation are sometimes referred to as opaque imitation because they are difficult to account for without positing cognitive mechanisms, such as perspective taking, that most animals have not been acknowledged to have. The present review first identifies various forms of social influence and social learning that do not qualify as opaque imitation, including species-typical mechanisms (e.g., mimicry and contagion), motivational mechanisms (e.g., social facilitation, incentive motivation, transfer of fear), attentional mechanisms (e.g., local enhancement, stimulus enhancement), imprinting, following, observational conditioning, and learning how the environment works (affordance learning). It then presents evidence for different forms of opaque imitation in animals, and identifies characteristics of human imitation that have been proposed to distinguish it from animal imitation. Finally, it examines the role played in opaque imitation by demonstrator reinforcement and observer motivation. Although accounts of imitation have been proposed that vary in their level of analysis from neural to cognitive, at present no theory of imitation appears to be adequate to account for the varied results that have been found.
	Address	Department of Psychology, University of Kentucky, Lexington, KY 40506-0044, USA. Zentall@uky.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	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:17024510			Approved	no
	Call Number	refbase @ user @			Serial	217
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	Author	Topál, J.; Byrne, R.W.; Miklósi, Á.; Csányi, V.
	Title	Reproducing human actions and action sequences: “Do as I Do!” in a dog			Type	Journal Article
	Year	2006	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	9	Issue	4	Pages	355-367
	Keywords	Animals; Comprehension; Conditioning, Operant; Discrimination Learning; Dogs/psychology; Humans; Imitative Behavior; Male; *Serial Learning
	Abstract	We present evidence that a dog (Philip, a 4-year-old tervueren) was able to use different human actions as samples against which to match his own behaviour. First, Philip was trained to repeat nine human-demonstrated actions on command ('Do it!'). When his performance was markedly over chance in response to demonstration by one person, testing with untrained action sequences and other demonstrators showed some ability to generalise his understanding of copying. In a second study, we presented Philip with a sequence of human actions, again using the 'Do as I do' paradigm. All demonstrated actions had basically the same structure: the owner picked up a bottle from one of six places; transferred it to one of the five other places and then commanded the dog ('Do it!'). We found that Philip duplicated the entire sequence of moving a specific object from one particular place to another more often than expected by chance. Although results point to significant limitations in his imitative abilities, it seems that the dog could have recognized the action sequence, on the basis of observation alone, in terms of the initial state, the means, and the goal. This suggests that dogs might acquire abilities by observation that enhance their success in complex socio-behavioural situations.
	Address	Comparative Ethology Research Group, Hungarian Academy of Sciences, Budapest, Pazmany, P. 1/c H-1117, Hungary. kea@t-online.hu
	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:17024511			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2434
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