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Nicol, C. J. (2004). Development, direction, and damage limitation: social learning in domestic fowl. Learn Behav, 32(1), 72–81.
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.
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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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Whiten, A., Horner, V., Litchfield, C. A., & Marshall-Pescini, S. (2004). How do apes ape? Learn. Behav., 32(1), 36–52.
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.
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Fragaszy, D., & Visalberghi, E. (2004). Socially biased learning in monkeys. Learn Behav, 32(1), 24–35.
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.
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Biederman, G. B., Robertson, H. A., & Vanayan, M. (1986). Observational learning of two visual discriminations by pigeons: a within-subjects design. J Exp Anal Behav, 46(1), 45–49.
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.
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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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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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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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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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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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