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Imitation in Animals

Imitation in Animals

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Published by Nicholay Atanassov

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Published by: Nicholay Atanassov on Sep 24, 2013
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Imitation in Animals:Evidence, Function, and Mechanisms
Tom ZentallandChana AkinsUniversity of Kentucky
The term imitation has been used descriptively and theoretically to characterize a broad range of animabehavior from physical antipredatory adaptations such as eye spots, which are totally under genetic control, to thehuman capacity for the exaggeration of individual characteristics, know as caricature, which are largely under cognitive control. In the present review many behaviors that have been called imitative are described, and aredistinguished from "true imitation." It is suggested that at much of the ambiguity in the literature as to what should be called imitation can be attributed to the distinction between the function of imitation and the mechanismsresponsible for it. Finally, the various mechanisms that have been proposed to account for true imitation are presented and an attempt is made to evaluate them.
I. Introduction
When the term imitation is used by psychologists it typically implies more than the simplereproduction of behavior. Imitation carries with it the implication of intentionality or purposiveness. Whenhumans use the term to describe their own behavior, it implies that there is an understanding of therelation between that behavior and the behavior being modeled (or demonstrated). For example, when achild imitates the behavior of an adult, one assumes that the child understands that the two behaviorsmatch (see photo to left). According to Freud (1933), imitation forms an integral part of the process of identification, and it is motivated by the need to identify (e.g., with the same-sex parent). But when welook for imitative learning in animals we must first pare away the mentalistic terms often used to describeand explain it, and then identify and rule out simpler mechanisms that might be involved in reports of itsoccurrence.The purpose of this chapter is first to classify the various cases of social influence depending onwhat mechanisms are thought to underlie them. For example, among the simplest kinds of socialinfluence are those social behaviors that are typical of the species and that happen to occur in unison(e.g., schooling, flocking, and herding). These behaviors can be thought of as genetically predisposed.Other behaviors may be made more likely to occur whenever there is an change in the motivation of theobserver, and the mere presence of another animal may have a motivational effect on the observer. Stillother behaviors may occur because the demonstrator or model can serve as a salient discriminativestimulus that predicts the appearance of food and allows for an account based on simple associativelearning. It is also possible for the behavior of a demonstrator to draw the observer’s attention to alocation or to a stimulus that is the focus of the target behavior (e.g., a bar that must be manipulated)and the observer’s mere proximity to that location or stimulus may make the target behavior moreprobable. Finally, it is possible for the behavior of the observer to have an effect on the environment suchthat the observer can learn how objects in the environment work. For example, seeing a door openoutward rather than inward. In some cases more than one simpler mechanism may be involved. In manycases it may not be possible to detect the presence of true imitative learning because of the potentialpresence of other simpler accounts. After addressing each of these classes of social influence in turn, wewill identify procedures that may separate them from what has come to be called true imitation.
A second purpose of this chapter is to distinguish between the function of imitation for the survivalof the organism and mechanisms involved in its use. The implication of this analysis is that social learningin general and imitation in particular may be more wide spread than previously thought. Finally, anattempt will be made to identify the mechanisms by which organisms are able to imitate.
II. Social Influence
The broad use of the term imitation extends to any influence that an organism may have onanother that results is a similarity of behavior or appearance between the two. Because biologists typicallytake a genetic approach to the study of organisms, for them, both physical appearance and behavior haveevolutionary bases and thus, are closely related.
Species Typical Factors
. When imitation involves copying of the physical appearance of one species by another, itis generally referred to as mimicry.When a relatively defenseless animals takes on the appearance of an animal that has betterdefenses, it is known as Batesian (or Mertensian) mimicry. The best known case of Batesian mimicry isthat of the palatable viceroy butterfly (left photo) mimicking the unpalatable monarch butterfly (rightphoto, Turner, 1984).A special case of mimicry involving behavior is the broken-wing display of the ground-nestingkilldeer and avocet (Sordahl, 1981). When the female bird is near thenest and a predator approaches, the bird flies away from the nestwhile mimicking the erratic flight pattern that might be shown by abird with a broken wing.
. When two or more animals engage in similar behaviorand that behavior is species typical, the term contagion is often used(Thorpe, 1963; also called mimesis, Armstrong, 1951, or responsefacilitation, Byrne, 1994). Contagion can be used to describe certaincourtship displays when they involve coordinated movementsbetween the male and female that are can sometimes appear to bevirtual mirror images (Tinbergen, 1960). Also, antipredatory behaviorcan be considered contagious when it involves the coordinatedmovement of a group of animals for defensive purposes. Suchbehavior occurs in certain mammalian (herding) and avian (flocking)species. When this coordinated behavior is aggressive (i.e., directedtoward rather than away from danger), it is known as mobbing(Hoogland & Sherman, 1976). Contagion can also be shown in anappetitive context. A satiated animal in the presence of food will oftenresume eating upon the introduction of a hungry animal which beginseating (Tolman, 1964). In the case of contagion, the behavior of oneanimal appears to serve as a releaser for the unlearned behavior of others (Thorpe, 1963). For thisreason, most research on animal imitation has used arbitrary, novel, to-be-acquired behavior as the focusresearch.
Motivational Factors
When imitative behavior is typically studied, an observing animal (e.g., a rat) is exposed to ademonstrator (rat) that is performing a novel response such as bar pressing and the rate of acquisition of the novel response is taken as a measure of imitation. But because rate of acquisition is a relativemeasure (it will depend on a variety of procedural factors unrelated to the conditions of observation), itmust be compared with that of a control group. The choice of control group often depends on the way theexperimental questions are asked. For example, early research appeared to assume that rats could learnthrough observation of the performance of a demonstrator, and the question was, did such learning occurmore quickly than the more typically used experimenter-trained shaping procedures (i.e., by successiveapproximations; Corson, 1967; Jacoby & Dawson, 1969; Powell, 1968; Powell & Burns, 1970; Powell,Saunders, & Thompson, 1969). The experimental question asked appears to have been a practical one:Would observation of the demonstration of a response be a faster means of training rats to bar press. Theproblem with the use of a “shaped” control group is that the procedures used to shape animals to barpress are difficult to specify in ways that can be accurately reproduced. Furthermore, comparison of therate of bar-press acquisition by an imitation group with that of a group of shaped animals may indicatelittle about the ability of animals to imitate, especially if the imitation group acquires the response at thesame rate as, or slower than, the control group. To ask about the ability of animals to acquire a responsethrough observation of a demonstrator, a more appropriate control group would be a “trial-and-error” control (i.e., a group of animals that acquires the response on its own).
Social facilitation
. The implication of imitative learning is that information transmitted from thedemonstrator to the observer has led to facilitated acquisition. There is evidence, however, that the merepresence of an animal (of the same species, i.e., a conspecific) can have effects on the motivational stateof an observer (Zajonc, 1965), and that changes in the observer’s motivational state can affect the2
acquisition and performance of a response (Zentall & Levine, 1972; Levine & Zentall, 1974). Zajonc hasreferred to this effect of the “mere presence” of a conspecific on motivation as social facilitation, and hehas proposed a version of Hull’s (1943) theory to account for the variety of findings of behavioralfacilitation and inhibition that have been reported in humans and animals. According to Zajonc, thepresence of a conspecific leads to an increase in arousal which can actually lead to the retardation of acquisition of a novel (to-be-learned) response. Others have suggested that the mere presence of aconspecific will facilitate the acquisition of a new response for the same reason (Gardner & Engel, 1971)or because the conspecific may have the ability toreduce fearin the observer (Davitz & Mason, 1955;Morrison & Hill, 1967). In any case, experiments concerned with true imitative learning must include acontrol for the possibility of facilitation or retardation of response acquisition resulting from mere presenceeffects.One further potential source of demonstrator-provided motivation should be mentioned here.Although the mere presence of a conspecific may contribute to the motivational state of an observer,general (nonspecific) activity may make an additional contribution. Being in the presence of an activeconspecific (i.e., one that is working for food but that is responding in a way that is irrelevant to thetarget response) might constitute an even better control than mere presence. We will return to this pointlater.
 Incentive motivation
. Reinforcement provided to the observer during the demonstration of anovel response (i.e., incentive motivation) may also play a role in the rate at which a novel response isacquired. Although Del Russo (1971) did not find significant evidence for imitative learning by rats thatobserved a demonstrator bar pressing for food (relative to a trial-and-error control) he did find significantfacilitation of bar pressing by a group of observers that got fed whenever their bar-pressingdemonstrators got fed. This facilitation may have involved a general increase in arousal on the part of thereinforced observer or a more specific association of the apparatus context with reinforcement. In eithercase, the observers would likely have been more active following observation than the nonreinforcedcomparison groups, and more active animals would be more likely to make accidental contact with thebar.
Observation of aversive conditioning
. The imitation of a novel response being acquired orbeing performed by a demonstrator that is motivated by the avoidance of painful stimulation (e.g.,electric shock) presents the need for special control. Emotional cues provided by a conspecific eitherescaping from or avoiding shock may provide emotional cues of pain or fear that could instill fear in anobserver. For example, John, Chesler, Bartlett, and Victor (1968) found that cats that had observed ademonstrator being trained to jump over a hurdle to avoid foot shock learned the hurdle-jumpingresponse faster than controls that did not observe the demonstrators. It may be, however, that being inthe presence of a cat being shocked was sufficient to increase the observers’ fear (motivation) associatedwith the conditioning context. Under such conditions, a change in motivation may facilitate acquisition.Under different conditions, however, although rats that observed a trained demonstrator that hadacquired a discriminated shuttle avoidance response acquired that response faster than rats that observeda merely present demonstrator, rats merely exposed to the empty shuttle box acquired the shuttleresponse fastest (Sanavio & Savardi, 1980).Trying to sort out mere presence and emotional motivational effects from learning effects can bequite complex. Although using well-trained demonstrators can reduce the likelihood of pain-produced cuesbeing transmitted to the observers (Del Russo, 1975), it may not be possible to avoid the effects of demonstrator-provided, fear-produced cues.One way to avoid problems associated with differential motivational cues encountered withobservation of aversively motivated conditioning is to include an observation control group exposed toperforming demonstrators but with the observer's view of a critical component of the demonstrator'sresponse blocked. Such a control was included in an experiment by Bunch and Zentall (1980) who used acandle flame avoidance task originally developed by Lore, Blanc, and Suedfeld (1971). Bunch and Zentallfound that rats learned the candle-flame-avoidance task faster after having seen a demonstrator acquirethe task, as compared with (1) a group for which a small barrier was placed in front of the candle suchthat the observer's view of the rat's contact with the candle was blocked and (2) a social facilitationcontrol group. Thus, although a variety of auditory cues (a potential by-product of the demonstrator'spain), olfactory cues (e.g., potentially produced by singed whiskers, defecation, and urination), and visualcues (e.g. seeing the demonstrator approach and then rapidly withdraw from something directly behindthe barrier) associated with the task should have provided comparable motivational cues to these controlobservers, task acquisition was not facilitated as much as for observers that could also observe thedemonstrator's contact with the candle.Another means of controlling for potential motivational cues provided by the demonstratorperforming a pain- or fear-motivated task, is to expose the observers to demonstrators performing adiscrimination (Kohn, 1976; Kohn & Dennis, 1972). In this research, rats that observed a demonstratorperforming a relevant shock-avoidance discrimination acquired that task faster than controls for which the3

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