A componential view of configural cues in generalization and discrimination in Pavlovian conditioning

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1 Behavioural Brain Research 110 (2000) A componential view of configural cues in generalization and discrimination in Pavlovian conditioning Susan E. Brandon *, Edgar H. Vogel, Allan R. Wagner Department of Psychology, Yale Uni ersity, P.O. Box Yale Station, New Ha en, CT 06520, USA Accepted 25 November 1999 Abstract This paper describes three theoretical approaches to the representation of configural cues in generalization and discrimination in Pavlovian conditioning: that of the Rescorla Wagner model, the Pearce model, and the authors replaced elements model. We summarize the results of a generalization experiment using the rabbit Pavlovian conditioned eyeblink response where animals were trained with cues A, AB, or ABC, and tested with A, AB, and ABC. The pattern of generalization decrement in testing supported the replaced elements model Elsevier Science B.V. All rights reserved. Keywords: Pavlovian; Conditioning; Generalization; Configural; Rabbits 1. Introduction 1.1. Three componential models Any theory of Pavlovian conditioning must address the fact that, with appropriate schedules of reinforcement, conditioned responding can be brought under the control of particular conjunctions of stimuli, as well as under the control of separable stimuli. For example, Saavedra [12] demonstrated that rabbits can learn a biconditional discrimination of the form, AX+/BX / AY /BY+ (the compounds AX and BY reinforced, and the compounds BX and AY nonreinforced). In this case, the animals came to respond to the combinations AX and BY but not to the same stimuli in the other combinations, BX and AY. These data, and related data on negative patterning [17,18], caused Rescorla and Wagner [11,16] to assume, like Spence [13], that there are configural elements that represent conjunctions of stimuli, in addition to those elements that represent the separate stimuli. This has sometimes been called the unique stimulus hypothesis [6]. We refer to it here as the added elements notion. * Corresponding author. Tel.: ; fax: address: susan brandon@yale.edu (S.E. Brandon) The essence of the added elements notion is depicted in panel a of Fig. 1, drawn in a manner that will facilitate subsequent comparison with other models. Any CS i is assumed to activate one set of context independent elements, and another CS j, is assumed to activate another set of context independent elements, which are assumed, for simplicity, to be nonoverlapping. The configural assumption is that CS i and CS j, when presented in conjunction, also activate an additional, unique, context dependent element. Thus, if the four stimuli, A, B, C, and D, were each represented in isolation by the sets of elements shown in panel b, the compounds AC, AD, BC, and BD would be represented by the sets indicated in panel c. The unique elements are ac, ad, bc, and bd. The Saavedra biconditional discrimination study [12] included a comparison group that was given the discrimination problem, AC+, BC, AD+, BD, using the same compounds. According to the added elements view, the biconditional discrimination should be possible because of the configural elements ac, ad, bc, and bd that are unique to each compound, but it should be more difficult than the comparison component problem, which has more unique elements. This is what Saavedra observed. Pearce [8,9] proposed that data such as those reported by Saavedra should encourage us to give up our /00/$ - see front matter 2000 Elsevier Science B.V. All rights reserved. PII: S (99)

2 68 S.E. Brandon et al. / Beha ioural Brain Research 110 (2000) elementistic view of the stimulus and suppose that association is always between the stimulus configurations that represent the CS and US on each trial. He presented an articulate, quantitative model of how such association occurs and how generalization between stimulus configurations can be conceptualized. Pearce made two essential assumptions. One is that the salience of all configurations is equal. The other is that the similarity between any pair of configurations, i and j, which dictates the degree of generalization between them, is equal to the ratio of the number of common cues to total cues in one configuration times the ratio of the number of common cues to total cues in the other configuration. For example, by this similarity rule, stimulus A has a similarity of 1/2 with both AC and AD (1/1 1/2), but AC and AD have only a similarity of 1/4 with each other (1/2 1/2). We have devised a componential model that approximates the calculations of Pearce s configural model to an arbitrarily close degree ([15], see also [1]). In componential terms, the assumption that the salience of all configurations is equivalent to assuming that the total number of theoretical elements in any compound is a Fig. 2. The inhibited-elements model. Panel a shows how CS i and CS j are assumed by this model to form a compound consisting of the original and inhibited elements. Panel b shows the elements which are assumed to comprise the CSs A, B, C, and D, for purposes of illustration of a component and biconditional discrimination, as shown in Panel c. Excitatory connections are depicted by the arrow; inhibitory connections by the square. See text for details. Fig. 1. The added-elements model. Panel a shows how CS i and CS j are assumed by this model to form a compound consisting of the original and added elements. Panel b shows the elements which are assumed to comprise the CSs A, B, C, and D, for purposes of illustration of a component and biconditional discrimination, as shown in Panel c.. Excitatory connections are depicted by the arrow; the and gate symbol is used for elements that are activated only when two or more components are activated. See text for details. constant. In this case, the Pearce similarity rule follows from assuming that the subset of theoretical elements representing any stimulus A that is active when A is in compound with one stimulus is statistically independent of the subset that is active when A is in compound with a different stimulus. Fig. 2 illustrates how a componential version of Pearce s theory works in simple diagrammatic form and in application to the biconditional versus component comparisons, as previously shown for the added elements view. What is depicted in Panel a is that CS i alone is represented by a number of elements (here, two), CS j alone by the same number (two), and CS i in compound with CS j, also by two elements. The componential application of Pearce s rule specifies that if CS i is coactive with CS j, it effectively inhibits half of the elements otherwise activated by CS i. Likewise, if CS j is coactive with CS i, it inhibits half of the elements otherwise activated by CS i. Illustrating the application of this model to the data on biconditional discriminations requires that A, B, C, and D alone (Panel b) each be represented by multiples of four elements. One can then see in Panel c how the assumption of statistical independence produces the appropriate generalization. In

3 S.E. Brandon et al. / Beha ioural Brain Research 110 (2000) the case of each compound, the elements activated by each stimulus alone that are not active in the compound are represented as darkened. Note that AC and AD each share half of their active theoretical elements with A alone, in agreement with Pearce s computation of 50% generalization between them. However, because a1 is the only active element common to both AC and AD, AC and AD share only 1/4 of their theoretical elements with each other. This is also consonant with Pearce s computation of 25% generalization between the two. This inhibited elements model accounts for the data just as well as the added elements model: The biconditional discrimination can be learned because there are elements (a2, c2, b2 and d2) that are unique to the reinforced AC and BD compounds. But, it should not be learned as rapidly as the component discrimination, which has more elements (a1, a2, a3, d1 and d3) that are unique to the reinforced AC and AD compounds. The added elements view was that representation of a compound stimulus includes the addition of unique configural elements. The componential interpretation of Fig. 3. The replaced-elements model. Panel a shows how CS i and CS j are assumed by this model to form a compound consisting of the original and replaced elements. Panel b shows the elements which are assumed to comprise the CSs A, B, C, and D, for purposes of illustration of a component and biconditional discrimination, as shown in Panel c. Excitatory connections are depicted by the arrow; inhibitory connections by the square. The and gate symbol is used for elements that are activated only when two or more components are activated. See text for details. Pearce s theory corresponds to the view that representation of a compound stimulus involves the inhibition of elements. The view considered here is that the representation of a compound stimulus involves both the addition of unique configural elements and the inhibition of elements otherwise contributed by the constituent stimuli. In effect, this replaced elements view is that representation of a compound stimulus involves the replacement of some components otherwise contributed by the constituent stimuli. Fig. 3 is a diagram of the replaced elements notion and illustration of its application to the biconditional and component discrimination problems. What is shown in Panel a is that CS i alone is represented by two elements, CS j alone by two elements, and CS i in compound with CS j by four elements. As can be seen, although the number of elements activated by the compound is the sum of the number of elements activated by the constituents, the identity of the elements activated by the compound is not the same as that of the elements activated by the constituents. We suggest that the presentation of CS j along with CS i both activates an element that is not activated by CS i alone, and at the same time inhibits the activation of an element that is activated by CS i alone. Likewise, the presentation of CS i along with CS j activates an element that is not activated by CS j alone, and inhibits an element that is activated by CS j alone. We view this conception of stimulus compounding as a way to make computational Hull s historically influential notion of afferent neural interaction ([3 6]), according to which all afferent neural impulses (s) active in the nervous system at any given instant, interact with each other in such a way as to change each into something partially different (š) in a manner which varies with every concurrent associated impulse or combination of such impulses ([3], p. 47). That is, the replaced elements model expresses how the presence of one stimulus can make the representation of a second stimulus different than it would be if it were presented alone: the representation of CS i is changed by the afferent neural interaction with CS j to include one element (or set of elements) rather than another. In implementing this view, the assumption is that each stimulus with which a CS can be compounded engenders the replacement of different elements (and that such pairwise replacement process is adequate to produce the necessary discriminable representations in any n-term compounds). In brief, there are context-specific components in the representation of constituent stimuli as well as in the representation of compound stimuli. Panel c of Fig. 3 shows how this works in the case of the biconditional problem. Again, the components otherwise activated by a stimulus that are replaced when that stimulus is in compound with another stimulus are represented as darkened. As may be seen, the compounding of A with C leads to the replacement of a4 by

4 70 S.E. Brandon et al. / Beha ioural Brain Research 110 (2000) ac, whereas the compounding of A with D leads to the replacement of a3 by ad. With these and the other indicated results of compounding, animals should learn the biconditional discrimination because there are elements that are unique to the reinforced (a3, ac, c3, ca, b3, bd, d3 and db) and nonreinforced (a4, ad, da, d4, b4, bc, cb, c4) compounds. But, they should not learn the task as quickly as the component discrimination problem, where there are more elements that are present in the reinforced versus the nonreinforced compounds Differential predictions The experiment described here provided one test of these models. The experiment focused on the differential predictions that the three models make when we train a subject in Pavlovian conditioning and then test for generalization to a new stimulus that is formed by either removing one or more of the nominal trained stimuli or from adding comparable stimuli to the trained stimulus. With reference again to the Rescorla Wagner reasoning as shown in Panel c of Fig. 1, training on AC and then testing on A should lead to a considerable generalization decrement: whatever associative strength had developed to the c elements and the ac configural element during training should be lost in the test. In contrast, training on A and then testing on AC should lead to no decrement. Whatever associative strength had developed to the a elements during training should be accessible in test: the added c and ac configural elements should be associatively neutral and have no predicted effect. But this is clearly erroneous: Pavlov [7] reported a decrement as a result of the adding of novel stimuli in test; the term that is commonly used to describe this effect, external inhibition, was suggested by Pavlov. Pearce [8,9] claimed as one of the advantages of his theoretical conception that it avoided this erroneous Rescorla Wagner prediction. His similarity rule embodies the premise that the generalization decrement from adding a cue, e.g. from A to AC, is identical to the decrement from removing a cue, e.g. from AC to A. In our componential rendition of Pearce (see again Panel c of Fig. 2), training on AC and then testing on A should lead to a generalization decrement: whatever associative strength had developed to the c elements should be lost in the test. This is the same as the Rescorla Wagner added elements prediction. The notable difference from the Rescorla Wagner view is that training on A and then testing of AC should lead to an equivalent decrement: whatever associative strength had been developed to the A components that are active during A alone but are inhibited during the AC compound should be lost in the test. If the distinguishing prediction of the added elements view is that there should be no generalization decrement from AC to A, the distinguishing prediction of the Pearce view is that there should be an equal decrement from A to AC as from AC to A. According to the replaced element view, and with reference to Panel c of Fig. 3, if a subject were trained on the AC compound and then tested on A, there should be considerable generalization decrement due to the loss of the c, ac, and ca elements that are in the AC representation but not in the A representation. If a subject were trained on the A stimulus and then tested on the AC compound, there should also be a generalization decrement due to the loss of the a4 element that is in the A representation but is replaced by the ac element in the AC compound. Our replaced unit view, unlike the Rescorla-Wagner view, but like the Pearce view, predicts that there will be a generalization decrement as a result of either the addition or the withdrawal of a stimulus. On the other hand, like the Rescorla Wagner view, but unlike the Pearce view, it predicts that the generalization decrement as a result of adding a stimulus should always be less than that as a result of removing a stimulus An experimental e aluation These differential predictions were tested in an experiment in which three groups of rabbits were trained with Pavlovian conditioning procedures with a single cue, A, a double compound cue, AB, or a triple compound cue, ABC. Following acquisition of a conditioned eyelid closure to its training cue, each animal was tested with each of the cues A, AB, and ABC. The prediction of the added elements model is that there should be no decrement in response for animals tested with elements added, but there should be a decrement in response for animals tested with elements lost, and the degree of decrement should be proportional to the number of elements lost. The prediction of the inhibited elements model is that there should be an equal amount of decrement in responding in each instance where one or two elements are lost as when they are added. And, the prediction of the replaced elements model is that there should be decrement both when elements are lost and when they are added, but the decrement should be greater in the former case than in the latter. 2. Materials and methods 2.1. Animals The animals were 18 experimentally naive, male, kg New Zealand white rabbits, individually

5 S.E. Brandon et al. / Beha ioural Brain Research 110 (2000) Following preparation, the animals were trained for seven daily sessions, where each session contained trials with the assigned CS, with an intertrial interval of 120 s. Two test sessions followed, where each consisted of 12 reinforced and 12 unreinforced trials with each of the three test compounds A, AB, and ABC, in an order of presentation that equated for first order sequential probabilities. A conditioned response was scored when the record indicated an eyelid closure of 0.5 mm or more, relative to the pre-cs baseline, initiated from 100 to 1000 ms after the onset of the CS and prior to the presentation of the US. 3. Results Fig. 4. The mean percentage eyelid CRs to each of the test stimuli, A, AB, and ABC by each of the training groups A, AB, and ABC. The brackets represent standard errors. housed and maintained with ad libitum food and water, except during experimental sessions. The experiment was run in three replications of six animals each, two from each experimental group Training and testing Training and testing were conducted in six identical cm isolation chambers. During experimental sessions the rabbit was loosely restrained within the chamber in a cm Plexiglas box, from which its head protruded. Three different stimuli served as CSs: an interrupted light (eight flashes/s), a 3000-Hz, 85-dB tone, and a vibrotactual stimulus supplied by a 60-Hz hand massager device applied to the animal s chest. All CSs were 1050 ms in duration. The assignment of a stimulus as A, B, and C was such that each stimulus was equally often designated A, B, and C within each group. The US, which coterminated with the CS, was a 50-ms train of 100/s, 3-mA, square-wave shock pulses, delivered to the right paraorbital region through stainless steel electrodes, one implanted approx. 5 mm ventral to the extreme nasal extent, and the other approx. 5 mm caudal to the extreme lateral extent, of each eye. Closure of the rabbit s eye was monitored with an adaptation of the photoresistive, rotary transducer described by Gormezano and Gibbs [2]. There were no notable differences among the groups across the acquisition sessions (data not shown). The test data are shown in Fig. 4. The bar graphs display separately for each of the three groups the mean percentage of eyeblink CRs to the training stimulus and the two generalization test stimuli during the test sessions. As may be seen, there was an equivalent level of responding among the three groups to their training cues. Also, there was less responding to the two novel stimuli than to the training stimulus in each group. However, in all of the direct comparisons, there was a reliably greater decrement produced by the withdrawal of a stimulus than by its addition: there was more of a decrement when B was withdrawn from the trained AB compound than when B was added to the trained A cue; there was more of a decrement when C was removed from the trained ABC compound that when C was added to the trained AB. And, there was more of a decrement when BC was removed from the trained ABC compound than when BC was added to the trained A cue. These comparisons were supported by parametric statistical analyses (Ps 0.05). 4. Discussion That there was a decrement due to the adding of stimuli is clearly contrary to the Rescorla Wagner view. As also may be seen, however, the effects of adding or withdrawing a stimulus were not as predicted by Pearce. The pattern of data supports the replaced elements model: there was a greater decrement in response with the removal of a cue than with the addition. When Pavlov offered a description of external inhibition, that is, the decrement in CRs that he observed when a well-trained CS was tested in compound with some novel stimulus, he postulated that there were competing, orienting responses, and that these might account for at least some of the response decrement:

6 72 S.E. Brandon et al. / Beha ioural Brain Research 110 (2000) The dog and experimenter would be isolated in the experimental room, all the conditions remaining for a while constant. Suddenly some disturbing factor would arise a sound would penetrate into the room; some quick change in illumination would occur, the sun going behind a cloud; or a draught would get in underneath the door, and maybe bring some odour with it. If any one of these extra stimuli happened to be introduced just at the time of application of the conditioned stimulus, it would inevitably bring about a more or less pronounced weakening or even a complete disappearance of the reflex response...the appearance of any new stimulus immediately evokes the investigatory reflex and the animal fixes all its appropriate receptor organs upon the source of the disturbance... ([7], p. 44). Although the models proposed in the present paper provide associative accounts of response generalization as a function of cue addition and/or cue removal, one might consider that competing, orienting responses contributed to the outcome we observed. For example, using the added element model, which does not predict a response decrement with the addition of a cue, one might argue that the reason for such decrement as observed is a nonassociative orienting response to the added cue. However, the inhibited elements model, which predicts equal generalization decrement from the removal and addition of cues, might equally allow one to argue that there is known to be orienting responses to the withdrawal of a usual cue as well as to the addition of cues [14], and perhaps the asymmetry in decrement observed here was due to a greater orienting response when a cue is withdrawn than when added. The contribution of antagonistic orienting responses to response decrement under various conditions of stimulus change certainly deserves investigation. We are, however, impressed that Reiss and Wagner [10] observed a response decrement when a stimulus was added to a well-trained CS, even though the added stimulus was preexposed over 1000 time prior to CS conditioning; i.e. when it was unlikely to evoke an investigatory reflex. Our theoretical analysis portrays context-dependent components as of more general behavioral significance than generally acknowledged [11,13,16]. As the reported experiment shows, they need to be recognized in order to account for basic facts of generalization following simple reinforced training. What is conceptually pleasing is the way in which a componential analysis allows one to relate what otherwise might appear to be quite disparate views. Pearce s configural theory [8,9] has been one of the most stimulating additions to Pavlovian theory in the last decade. Hull s theory of afferent neural interaction [3,4] was one of the most respected theories of his day. Both can be seen as componential theories, Pearce s theory as one in which elements otherwise activated by constituent stimuli are inhibited in compound, Hull s theory as one in which elements otherwise activated by constituent stimuli are replaced in compound. References [1] Bahçekapili HG. An evaluation of Rescorla and Wagner s elementistic model versus Pearce s configural model in discrimination learning. In: Unpublished Doctoral Dissertation. New Haven: Yale University, [2] Gormezano I, Gibbs CM. Transduction of the rabbit s nictitating membrane response. Behav Res Meth Instr Comput 1988;20: [3] Hull CL. Principles of Behavior. Appleton-Century-Crofts, 1943:422. [4] Hull CL. The discrimination of stimulus configuration and the hypothesis of afferent neural interaction. Psychol Rev 1945;52: [5] Hull CL. A Behavior System. New Haven: Yale University Press, 1927:372. [6] Kehoe EJ. A layered network model of associative learning: learning to learn and configuration. Psychol Rev 1988;95: [7] Pavlov IP. Conditioned Reflexes. Oxford: Oxford University Press, [8] Pearce JM. A model for stimulus generalization in Pavlovian conditioning. Psychol Rev 1987;94: [9] Pearce JM. Similarity and discrimination: a selective review and a connectionist model. Psychol Rev 1994;101: [10] Reiss S, Wagner AR. CS habituation produces a latent inhibition effect but no active conditioned inhibition. Learn Motiv 1972;3: [11] Rescorla RA, Wagner AR. A theory of Pavlovian conditioning: variations in the effectiveness of reinforcement and nonreinforcement. In: Black AH, Prokasy WF, editors. Classical Conditioning II. New York: Appleton-Century-Crofts, 1972: [12] Saavedra MA. Pavlovian compound conditioning in the rabbit. Learn Motiv 1975;6: [13] Spence KW. The nature of the response in discrimination learning. Psychol Rev 1952;59: [14] Sokolov EN. The neuronal mechanisms of the orienting reflex. In: Sokolov EN, Vinogradova OS, editors. Neuronal Mechanisms of the Orienting Reflex. Hillsdale, NJ: Erlbaum, 1975: [15] Wagner AR, Brandon SE. A componential theory of associative learning. In: Mowrer RR, Klein SB, editors. Contemporary Learning: Theory and Application. Hillsdale, NJ:Erlbaum, in press. [16] Wagner AR, Rescorla RA. Inhibition in Pavlovian conditioning: application of a theory. In: Boakes RA, Haliday MS, editors. Inhibition and Learning. New York: Academic Press, 1972: [17] Whitlow JW Jr, Wagner AR. Negative patterning in classical conditioning: summation of response tendencies to isolable and configural components. Psychon Sci 1972;27: [18] Woodbury CB. The learning of stimulus patterns by dogs. J Comp Psychol 1943;35:

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