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Mercado, E. 3rd, Herman, L. M., & Pack, A. A. (2005). Song copying by humpback whales: themes and variations. Anim. Cogn., 8(2), 93–102.
Abstract: Male humpback whales (Megaptera novaeangliae) produce long, structured sequences of sound underwater, commonly called “songs.” Humpbacks progressively modify their songs over time in ways that suggest that individuals are copying song elements that they hear being used by other singers. Little is known about the factors that determine how whales learn from their auditory experiences. Song learning in birds is better understood and appears to be constrained by stable core attributes such as species-specific sound repertoires and song syntax. To clarify whether similar constraints exist for song learning by humpbacks, we analyzed changes over 14 years in the sounds used by humpback whales singing in Hawaiian waters. We found that although the properties of individual sounds within songs are quite variable over time, the overall distribution of certain acoustic features within the repertoire appears to be stable. In particular, our findings suggest that species-specific constraints on temporal features of song sounds determine song form, whereas spectral variability allows whales to flexibly adapt song elements.
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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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Lonsdorf, E. V. (2006). What is the role of mothers in the acquisition of termite-fishing behaviors in wild chimpanzees (Pan troglodytes schweinfurthii)? Anim. Cogn., 9(1), 36–46.
Abstract: This paper explores the role of maternal influences on the acquisition of a tool-using task in wild chimpanzees (Pan troglodytes schweinfurthii) in order to build on and complement previous work done in captivity. Young chimpanzees show a long period of offspring dependency on mothers and it is during this period that offspring learn several important skills, especially how to and on what to forage. At Gombe National Park, one skill that is acquired during dependency is termite-fishing, a complex behavior that involves inserting a tool made from the surrounding vegetation into a termite mound and extracting the termites that attack and cling to the tool. All chimpanzees observed at Gombe have acquired the termite-fishing skill by the age of 5.5 years. Since the mother is the primary source of information throughout this time period, I investigated the influence of mothers' individual termite-fishing characteristics on their offsprings' speed of acquisition and proficiency at the skill once acquired. Mother's time spent alone or with maternal family members, which is highly correlated to time spent termite-fishing, was positively correlated to offspring's acquisition of critical elements of the skill. I also investigated the specific types of social interactions that occur between mothers and offspring at the termite mound and found that mothers are highly tolerant to offspring, even when the behavior of the offspring may disrupt the termite-fishing attempt. However, no active facilitation by mothers of offsprings' attempts were observed.
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Bard, K. A. (2007). Neonatal imitation in chimpanzees (Pan troglodytes) tested with two paradigms. Anim. Cogn., 10(2), 233–242.
Abstract: Primate species differ in their imitative performance, perhaps reflecting differences in imitative capacity. The developmentally earliest form of imitation in humans, neonatal imitation, occurs in early interactions with social partners, and may be a more accurate index of innate capacity than imitation of actions on objects, which requires more cognitive ability. This study assessed imitative capacity in five neonatal chimpanzees, within a narrow age range (7-15 days of age), by testing responses to facial and vocal actions with two different test paradigms (structured and communicative). Imitation of mouth opening was found in both paradigms. In the communicative paradigm, significant agreement was found between infant actions and demonstrations. Additionally, chimpanzees matched the sequence of three actions of the TC model, but only on the second demonstration. Newborn chimpanzees matched more modeled actions in the communicative test than in the structured paradigm. These performances of chimpanzees, at birth, are in agreement with the literature, supporting a conclusion that imitative capacity is not unique to the human species. Developmental histories must be more fully considered in the cross-species study of imitation, as there is a greater degree of innate imitative capacity than previously known. Socialization practices interact with innate and developing competencies to determine the outcome of imitation tests later in life.
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Heyes, C. M., & Dawson, G. R. (1990). A demonstration of observational learning in rats using a bidirectional control. Q J Exp Psychol B, 42(1), 59–71.
Abstract: Hungry rats observed a conspecific demonstrator pushing a single manipulandum, a joystick, to the right or to the left for food reward and were then allowed access to the joystick from a different orientation. The effects of right-pushing vs left-pushing observation experience on (1) response acquisition, (2) reversal of a left-right discrimination, and (3) responding in extinction, were examined. Rats that had observed left-pushing made more left responses during acquisition than rats that had observed right-pushing, and rats that had observed demonstrators pushing in the direction that had previously been reinforced took longer to reach criterion reversal and made more responses in extinction than rats that had observed demonstrators pushing in the opposite direction to that previously reinforced. These results provide evidence that rats are capable of learning a response, or a response-reinforcer contingency, through conspecific observation.
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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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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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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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Caldwell, C. A., & Whiten, A. (2004). Testing for social learning and imitation in common marmosets, Callithrix jacchus, using an artificial fruit. Anim. Cogn., 7(2), 77–85.
Abstract: We tested for social learning and imitation in common marmosets using an artificial foraging task and trained conspecific demonstrators. We trained a demonstrator marmoset to open an artificial fruit, providing a full demonstration of the task to be learned. Another marmoset provided a partial demonstration, controlling for stimulus enhancement effects, by eating food from the outside of the apparatus. We thus compared three observer groups, each consisting of four animals: those that received the full demonstration, those that received the partial demonstration, and a control group that saw no demonstration prior to testing. Although none of the observer marmosets succeeded in opening the artificial fruit during the test periods, there were clear effects of demonstration type. Those that saw the full demonstration manipulated the apparatus more overall, whereas those from the control group manipulated it the least of the three groups. Those from the full-demonstration group also contacted the particular parts of the artificial fruit that they had seen touched (localised stimulus enhancement) to a greater extent than the other two groups. There was also an interaction between the number of hand and mouth touches made to the artificial fruit for the full- and partial-demonstration groups. Whether or not these data represent evidence for imitation is discussed. We also propose that the clear differences between the groups suggest that social learning mechanisms provide real benefits to these animals in terms of developing novel food-processing skills analogous to the one presented here.
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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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