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Zentall, T. R. (2006). Imitation: definitions, evidence, and mechanisms. Anim. Cogn., 9(4), 335–353.
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.
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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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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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Akins, C. K., Klein, E. D., & Zentall, T. R. (2002). Imitative learning in Japanese quail (Coturnix japonica) using the bidirectional control procedure. Anim Learn Behav, 30(3), 275–281.
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.
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Nocera, J. J., Forbes, G. J., & Giraldeau, L. - A. (2006). Inadvertent social information in breeding site selection of natal dispersing birds. Proc Biol Sci, 273(1584), 349–355.
Abstract: Several species use the number of young produced as public information (PI) to assess breeding site quality. PI is inaccessible for synchronously breeding birds because nests are empty by the time the young can collect this information. We investigate if location cues are the next best source of inadvertent social information (ISI) used by young prospectors during breeding site choice. We experimentally deployed ISI as decoys and song playbacks of breeding males in suitable and sub-optimal habitats during pre- and post-breeding periods, and monitored territory establishment during the subsequent breeding season for a social, bobolink (Dolichonyx oryzivorus), and a more solitary species, Nelson's sharp-tailed sparrow (Ammodramus nelsoni). The sparrows did not respond to treatments, but bobolinks responded strongly to post-breeding location cues, irrespective of habitat quality. The following year, 17/20 sub-optimal plots to which bobolink males were recruited were defended for at least two weeks, indicating that song heard the previous year could exert a “carry-over attraction” effect on conspecifics the following year. Sixteen recruited males were natal dispersers, as expected when animals have little opportunity to directly sample their natal habitat quality. We suggest that differences in breeding synchronicity may induce an equivalent clinal distribution of ISI use.
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Range, F., Bugnyar, T., Schlogl, C., & Kotrschal, K. (2006). Individual and sex differences in learning abilities of ravens. Behav. Process., 73(1), 100–106.
Abstract: Behavioral and physiological characteristics of individuals within the same species have been found to be stable across time and contexts. In this study, we investigated individual differences in learning abilities and object and social manipulation to test for consistency within individuals across different tasks. Individual ravens (Corvus corax) were tested in simple color and position discrimination tasks to establish their learning abilities. We found that males were significantly better in the acquisition of the first discrimination task and the object manipulation task, but not in any of the other tasks. Furthermore, faster learners engaged less often in manipulations of conspecifics and exploration of objects to get access to food. No relationship between object and social manipulation and reversal training were found. Our results suggest that individual differences in regard to the acquisition of new tasks may be related to personalities or at least object manipulation in ravens.
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Chase, I. D., Tovey, C., Spangler-Martin, D., & Manfredonia, M. (2002). Individual differences versus social dynamics in the formation of animal dominance hierarchies. Proc. Natl. Acad. Sci. U.S.A., 99(8), 5744–5749.
Abstract: Linear hierarchies, the classical pecking-order structures, are formed readily in both nature and the laboratory in a great range of species including humans. However, the probability of getting linear structures by chance alone is quite low. In this paper we investigate the two hypotheses that are proposed most often to explain linear hierarchies: they are predetermined by differences in the attributes of animals, or they are produced by the dynamics of social interaction, i.e., they are self-organizing. We evaluate these hypotheses using cichlid fish as model animals, and although differences in attributes play a significant part, we find that social interaction is necessary for high proportions of groups with linear hierarchies. Our results suggest that dominance hierarchy formation is a much richer and more complex phenomenon than previously thought, and we explore the implications of these results for evolutionary biology, the social sciences, and the use of animal models in understanding human social organization.
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Fricke, H. W. (1973). Individual partner recognition in fish: field studies on Amphiprion bicinctus. Naturwissenschaften, 60(4), 204–205.
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Hemelrijk, C. K., & Wantia, J. (2005). Individual variation by self-organisation. Neurosci Biobehav Rev, 29(1), 125–136.
Abstract: In this paper, we show that differences in dominance and spatial centrality of individuals in a group may arise through self-organisation. Our instrument is a model, called DomWorld, that represents two traits that are often found in animals, namely grouping and competing. In this model individual differences grow under the following conditions: (1) when the intensity of aggression increases and grouping becomes denser, (2) when the degree of sexual dimorphism in fighting power increases. In this case the differences among females compared to males grow too, (3) when, upon encountering another individual, the tendency to attack is 'obligate' and not conditional, namely 'sensitive to risks'. Results resemble phenomena described for societies of primates, mice, birds and pigs.
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Kralj-Fiser, S., Scheiber, I. B. R., Blejec, A., Moestl, E., & Kotrschal, K. (2007). Individualities in a flock of free-roaming greylag geese: behavioral and physiological consistency over time and across situations. Horm Behav, 51(2), 239–248.
Abstract: The concept of personality implies individual differences in behavior and physiology that show some degree of repeatability/consistency over time and across contexts. Most studies of animal personality, particularly studies of individuals' variation in physiological mechanisms, have been conducted on selected individuals in controlled conditions. We attempted to detect consistent behaviors as well as physiological patterns in greylag ganders (Anser anser) from a free-roaming flock living in semi-natural conditions. We tested 10 individuals repeatedly, in a handling trial, resembling tests for characterization of “temperaments” in captive animals. We recorded the behavior of the same 10 individuals during four situations in the socially intact flock: (1) a “low density feeding condition”, (2) a “high density feeding condition”, (3) a “low density post-feeding situation” and (4) while the geese rested. We collected fecal samples for determination of excreted immuno-reactive corticosterone (BM) and testosterone metabolites (TM) after handling trials, as well as the “low density feeding” and the “high density feeding” conditions. BM levels were very highly consistent over the repeats of handling trials, and the “low density feeding condition” and tended to be consistent over the first two repeats of the “high density feeding condition”. Also, BM responses tended to be consistent across contexts. Despite seasonal variation, there tended to be inter-test consistency of TM, which pointed to some individual differences in TM as well. Aggressiveness turned out to be a highly repeatable trait, which was consistent across social situations, and tended to correlate with an individual's resistance during handling trials. Also, “proximity to the female partner” and “sociability” – the average number of neighboring geese in a close distance while resting – were consistent. We conclude that aggressiveness, “affiliative tendencies” and levels of excreted corticosterone and testosterone metabolites may be crucial factors of personality in geese.
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