Interaction between preexposure and overshadowing: Further analysis of the extended comparator hypothesis

Three experiments with rats used conditioned suppression of barpress to test predictions of the extended comparator hypothesis, which assumes that the effectiveness of (first-order) comparator stimuli in modulating responding to a target conditioned stimulus (CS) is itself modulated by other (second-order) comparator stimuli. Experiment 1 demonstrated that both pretraining exposure to the target CS alone (i.e., CS-preexposure effect, also known as latent inhibition) and pretraining exposure to a compound of the target CS and nontarget CS (i.e., compound-CS-preexposure effect) counteract overshadowing, and that posttraining deflation (i.e., extinction) of the overshadowing stimulus attenuates responding to the target CS when overshadowing is preceded by a CS-preexposure treatment (i.e., yields a CS-preexposure effect), but not when overshadowing is preceded by a compound-CS-preexposure treatment. Experiment 2 examined the consequences of posttraining associative inflation of the overshadowing stimulus or the preexposure companion stimulus following conjoint compound-CS-preexposure and overshadowing treatment. Experiment 3 examined the consequences of posttraining inflation of the overshadowing stimulus or the context following conjoint CS-alone preexposure and overshadowing treatment. The results support the expression-focused comparator view in contrast to recent acquisition-focused models of retrospective revaluation.     

Responding to a conditioned stimulus depends on the current associative status of other cues present during training of that specific stimulus.

The comparator hypothesis is a response rule stating that responding to a Pavlovian conditioned stimulus (CS) reflects the associative strength of the CS relative to that of other cues (comparator stimuli) that were present during CS training. Thus, modulation of the associative strength of a CS's comparator stimulus should alter responding to that CS. These studies examined the stimulus specificity of this effect using within-subjects designs. Rats were trained on 2 CSs, each with a unique comparator stimulus, to determine the degree to which posttraining extinction of the comparator stimulus for one CS influences responding to the other CS. Using negative contingency (Experiments 1 and 2), overshadowing (Experiment 3), and local context (Experiment 4) preparations, stimulus specificity was observed. In each case, posttraining extinction of the comparator stimulus for one CS had greater impact on responding to that CS than on responding to the alternate CS.     

Degraded contingency revisited: posttraining extinction of a cover stimulus attenuates a target cue's behavioral control

In a Pavlovian conditioning situation, unsignaled outcome presentations interspersed among cue-outcome pairings attenuate conditioned responding to the cue (i.e., the degraded contingency effect). However, if a nontarget cue signals these added outcomes, responding to the target cue is partially restored (i.e., the cover stimulus effect). In 2 conditioned suppression experiments using rats, the effect of posttraining extinction of the cover stimulus was examined. Experiment 1 found that this treatment yielded reduced responding to the target cue. Experiment 2 replicated this finding, while demonstrating that this basic effect was not due to acquired equivalence between the target cue and the cover stimulus. These results are consistent with the extended comparator hypothesis interpretation of the degraded contingency and cover stimulus effects.     

Trial spacing is a determinant of cue interaction

Four conditioned lick suppression experiments with rats examined the effect of trial spacing on cue interaction. Experiments 1 and 2 found overshadowing to be eliminated with massed compound stimulus-outcome pairings and the usual trial spacing effect to be reversed with compound acquisition trials. Experiment 3 found that whether acquisition compound-outcome pairings were massed or spaced determined the effect of posttraining extinction treatment. Extinction of the overshadowing cue reduced responding following massed training and increased responding following spaced training. Extinction of the context decreased responding following massed training. Experiment 4 found the conditioning and devaluation results to be associative and stimulus specific. These results are in accord with the extended comparator hypothesis (J. C. Denniston, H. I. Savastano, & R. R. Miller, 2001).     

Overshadowing of subsequent events and recovery thereafter

Four experiments using a conditioned lick suppression preparation with rats were conducted to examine whether overshadowing of subsequent events could be obtained in Pavlovian backward conditioning (i.e. unconditioned stimulus [US] before conditioned stimulus [CS]), and to determine whether such overshadowing could be reversed without further training with the overshadowed CS, as has been reported in overshadowing of antecedent events. In Experiment 1, a backward-conditioned CS overshadowed a second backward-conditioned CS. Two posttraining manipulations, extinction of the overshadowing CS (Experiment 2) and shifting of the temporal relationship of the overshadowing CS to the US (Experiment 3), increased responding to the overshadowed CS. These results constitute the first unambiguous demonstration of stimulus competition between subsequent events using first-order conditioning, and they show that, like overshadowing with forward conditioning, such overshadowing is due, at least in part if not completely, to a failure to express information that had been acquired.     

A comparator view of Pavlovian and differential inhibition

In 3 experiments using rats as subjects, the authors varied trial spacing to investigate the conditions under which Pavlovian and differential inhibition are observed. Experiment 1 compared Pavlovian and differential inhibition with spaced training trials. Spaced trials resulted in only the Pavlovian inhibitor passing both summation and retardation tests. Conversely, Experiment 2 compared these 2 types of inhibition with massed training trials. This training resulted in only the differential inhibitor passing both tests for conditioned inhibition. Finally, in Experiment 3 all subjects experienced Pavlovian inhibition training with massed trials. Although this training by itself did not result in behavior indicative of inhibition, subjects that also experienced posttraining extinction of the training context did pass both tests for inhibition. Overall, these results are anticipated by the extended comparator hypothesis (Denniston, Savastano, & Miller, 2001) but are problematic for most contemporary associative learning theories.     

An inhibitory within-compound association attenuates overshadowing

According to the comparator hypothesis (Miller & Matzel, 1988), cue competition depends on the association between a target stimulus (X) and a competing cue (e.g., an overshadowing cue [A]). Thus, it was expected that overshadowing would be reduced by establishing an inhibitory-like relationship between X and A before compound conditioning. In three lever press suppression experiments with rats, this expectation was supported. Experiment 1 showed that establishing an inhibitory X-A relationship reduced overshadowing. In Experiment 2, degrading the inhibitory-like relationship before conditioning allowed reinforced AX compound trials to result in overshadowing. Experiment 3 replicated the results of Experiment 2 when the inhibitory relationship was degraded after compound conditioning. The results support the view that within-compound associations are necessary not only for retrospective revaluation, but also for conventional cue competition.     

Conditioned inhibition produced by extinction-mediated recovery from the relative stimulus validity effect: a test of acquisition and performance models of empirical retrospective revaluation

Empirical retrospective revaluation is a phenomenon of Pavlovian conditioning and human causal judgment in which posttraining changes in the conditioned response (Pavlovian task) or causal rating (causal judgment task) of a cue occurs in the absence of further training with that cue. Two experiments tested the contrasting predictions made by 2 families of models concerning retrospective revaluation effects. In a conditioned lick-suppression task, rats were given relative stimulus validity training, consisting of reinforcing a compound of conditioned stimuli (CSs) A and X and nonreinforcement of a compound of CSs B and X, which resulted in low conditioned responding to CS X. Massive posttraining extinction of CS A not only enhanced excitatory responding to CS X, but caused CS B to pass both summation (Experiment 1) and retardation (Experiment 2) tests for conditioned inhibition. The inhibitory status of CS B is predicted by the performance-focused extended comparator hypothesis (J. C. Denniston, H. I. Savastano, & R. R. Miller, 2001), but not by acquisition-focused models of empirical retrospective revaluation (e.g., A. Dickinson & J. Burke, 1996; L. J. Van Hamme & E. A. Wasserman, 1994).     

Mechanisms underlying retarded emergence of conditioned responding following inhibitory training: evidence for the comparator hypothesis

The comparator hypothesis posits that conditioned responding is determined by a comparison at the time of testing between the associative strengths of the conditioned stimulus (CS) and stimuli proximal to the CS at the time of conditioning. The hypothesis treats all associations as being excitatory and treats conditioned inhibition as the behavioral consequence of a CS that is less excitatory than its comparator stimuli. Conditioned lick suppression by rats was used to differentiate four possible sources of retarded responding to an inhibitory CS. These include habituation to the unconditioned stimulus (US), latent inhibition to the CS, blocking of the CS-US association by the conditioning context, and enhanced excitatory associations to the comparator stimuli. Prior research has demonstrated the first three phenomena. Therefore, we employed parameters expected to highlight the fourth one--the comparator process. In Experiment 1, our negative contingency training was shown to produce a conditioned inhibitor that passed inhibitory summation and retardation tests. In Experiment 2 we found transfer of retardation from an inhibitory CS to a novel stimulus when the location where retardation-test training occurred was excitatory, which is indicative of contextual blocking and/or comparator effects. In Experiment 3, extinction of the conditioning context was found to attenuate retardation regardless of whether extinction occurred before or after the CS-US pairings of the retardation test. This indicates that much of the present retardation was due to the comparator process rather than to contextual blocking. Experiment 4 demonstrated that habituation to the US did not contribute to retardation in the present case. Collectively, these studies suggest that retardation following inhibitory training can be explained without recourse to any of the traditional mechanisms of conditioned inhibition.     

Recovery of an overshadowed association achieved by extinction of the overshadowing stimulus

Three experiments are reported in which conditioned lick suppression by water-deprived rats was used as an index of associative strength. In Experiment 1, overshadowing of a light by a tone was observed when the light-tone compound stimulus was paired with foot shock. After initial compound pairings, the tone-shock association was extinguished in one group of subjects. Subsequently, these animals demonstrated significantly higher levels of suppression to the light relative to a control group in which the tone had not been extinguished. Experiment 2 replicated this effect while failing to find evidence to support the possibilities that extinction presentations of the overshadowing tone act as retrieval cues for the light-shock association, or that, via second-order conditioning, the light-shock association is actually formed during extinction of the tone. Experiment 3 determined that the recovery from overshadowing observed in Experiments 1 and 2 was specific to the extinction of the overshadowing stimulus rather than the extinction of any excitatory cue. Collectively, these results suggest that the debilitated response to an overshadowed stimulus does not represent an acquisition failure, but rather the failure of an acquired association to be manifest in behavior.     

The restoration of extinguished avoidance responses

Following extensive signaled shuttlebox avoidance training, Fischer 344 rats were given extinction trials until they reliably stopped responding and, thereafter, they received occasional classical CS-UCS pairings during the course of restoration test trials in which shock was absent. Experiment I demonstrated that the classical contingency was both necessary and sufficient for the complete recovery of such behavior. Experiment II revealed that the degree of recovery was not differentially affected by the frequency of such CS-UCS pairings. However, in Experiment III recovery in the presence of this contingency was shown to depend on the number of previous extinction trials, although even after a large number of such trials, a considerable degree of recovery occurred. These findings demonstrated the central role of the classical CS-UCS contingency in restoring a previously extinguished avoidance response.     

Blocking and overshadowing in two species of fish.

In Experiment 1, a mixed blocking-overshadowing effect of color on an auditory discrimination was demonstrated in goldfish. In Experiment 2 (with lines differing in color and angle), blocking of angle by color and of color by angle was demonstrated in goldfish, In Experiment 3 (again with lines differing in color and angle), overshadowing of angle by color was demonstrated in carp, but the same animals (like goldfish in a previous study) failed to show greater intradimensional than extradimensional transfer. These results are consistent with the hypothesis that blocking and overshadowing are general phenomena of vertebrate learning. They suggest also that the processes responsible for blocking and overshadowing are different from those which produce the dimensional transfer effect.     

Mediated conditioning versus retrospective revaluation in humans: The influence of physical and functional similarity of cues

Two experiments assessed whether similarity between the two elements of a compound would influence the degree of mediated extinction versus recovery from overshadowing in human causal judgements. In both Experiments 1 and 2, we assessed the influence of extinguishing one element of a two-element compound on judgements about the other element. In Experiment 1 we manipulated the physical similarity of the two elements of the compound; in Experiment 2, we used equivalence and distinctiveness pretraining in order to vary their functional similarity. We found that these procedures influenced mediated extinction and recovery from overshadowing as a function of both physical and acquired similarity and distinctiveness, respectively. The implications of these results for previously reported differences between humans and nonprimate animals are discussed.     

Mediated conditioning versus retrospective revaluation in humans: the influence of physical and functional similarity of cues

Two experiments assessed whether similarity between the two elements of a compound would influence the degree of mediated extinction versus recovery from overshadowing in human causal judgements. In both Experiments 1 and 2, we assessed the influence of extinguishing one element of a two-element compound on judgements about the other element. In Experiment 1 we manipulated the physical similarity of the two elements of the compound; in Experiment 2, we used equivalence and distinctiveness pretraining in order to vary their functional similarity. We found that these procedures influenced mediated extinction and recovery from overshadowing as a function of both physical and acquired similarity and distinctiveness, respectively. The implications of these results for previously reported differences between humans and nonprimate animals are discussed.     

Extinction of conditioned taste aversions: Effects of concentration and overshadowing

Extinction of conditioned taste aversions was examined as a function of taste concentration and of the presence of an additional taste. The results of Experiment 1 were consistent with previous evidence in that a conditioned aversion to high concentration saline was more persistent in extinction than an aversion to a low concentration. However, when floor effects were avoided the rate of extinction was faster for the higher (1%) concentration than for 0.2% saline (Experiment 2), a result consistent with accounts of extinction in other preparations. Three further experiments examined extinction of a conditioned sucrose aversion. The addition of 1% saline, but not of 0.2% saline, to sucrose during extinction produced overshadowing ("protection from extinction"; Experiment 3). Such overshadowing by saline was detected after two, but not after a single extinction trial (Experiment 4). This last finding suggests that under the conditions of the present experiments sweet and salty tastes function as elemental stimuli competing for loss of associative strength. No overshadowing was found when almond (an aqueous odour) was used in place of saline as the added stimulus, even when high concentrations of almond were used that produced observable neophobia (Experiments 5A and 5B).     

Failure of serial taste-taste compound presentations to produce overshadowing of extinction of conditioned taste aversion

Two experiments were conducted to study overshadowing of extinction in a conditioned taste aversion preparation. In both experiments, aversive conditioning with sucrose was followed by extinction treatment with either sucrose alone or in compound with another taste, citric acid. Experiment 1 employed a simultaneous compound extinction treatment and found results indicative of overshadowing of extinction. By contrast, Experiment 2, in which extinction treatment involved serial compound presentations, failed to obtain evidence of overshadowing of extinction. The results of Experiment 2 indicate that the serial presentation of two tastes was processed as equivalent to the separate presentation of the tastes. The results are discussed in relation to: (1) Convergent evidence from research on latent inhibition, (2) competing theories of learning and, (3) their possible adaptive value in food-selection learning.     

Reciprocal overshadowing in flavour-aversion learning

The present experiments were concerned with the overshadowing by a novel flavour which preceded the pairing of a target flavour and toxicosis. This effect was demonstrated in Experiment I as was reciprocal overshadowing by a novel flavour which was interpolated between the target flavour and illness. Experiment 2 indicated that such overshadowing only occurred if the novel flavour was presented immediately prior to the target flavour; an interval of 10 min between the two abolished the effect. Experiment 3 indicated that the failure to observe overshadowing with a delay of 10 min between the two flavours was not due to a lack of association between the interfering flavour and illness. These data were interpreted as demonstrating that the introduction of the interfering flavour on the training day resulted in a perceptual generalisation decrement and a subsequent reduction in the aversion displayed to the target flavour.     

The role of contextual associations in producing the partial reinforcement acquisition deficit

Three conditioned suppression experiments with rats as subjects assessed the contributions of the conditioned stimulus (CS)-context and context-unconditioned stimulus (US) associations to the degraded stimulus control by the CS that is observed following partial reinforcement relative to continuous reinforcement training. In Experiment 1, posttraining associative deflation (i.e., extinction) of the training context after partial reinforcement restored responding to a level comparable to the one produced by continuous reinforcement. In Experiment 2, posttraining associative inflation of the context (achieved by administering unsignaled outcome presentations in the context) enhanced the detrimental effect of partial reinforcement. Experiment 3 found that the training context must be an effective competitor to produce the partial reinforcement acquisition deficit. When the context was down-modulated, the target regained behavioral control thereby demonstrating higher-order retrospective revaluation. The results are discussed in terms of retrospective revaluation, and are used to contrast the predictions of a performance-focused model with those of an acquisition-focused model.     

Backward conditioning: mediation by the context

The information acquired in backward conditioning (i.e., outcome-->cue) was assessed in 3 Pavlovian lick-suppression experiments with water-deprived rats as subjects. Experiment 1 confirmed previous research that few outcome-->cue pairings made the cue into a conditioned excitor and additionally showed that massive posttraining extinction of the training context attenuated a backward-trained cue's excitatory value. Experiment 2 found that many outcome-->cue pairings made the cue into a conditioned inhibitor and that the same context manipulation attenuated this inhibitory value. Experiment 3 confirmed the observations of Experiments 1 and 2 and demonstrated that these effects of context extinction were specific to backward-trained cues conditioned in the extinguished context. These results are interpreted in terms of cue-->context and context-->outcome associations.     

The effects of verbal instruction on affective and expectancy learning

The current research assessed the effects of verbal instruction on affective and expectancy learning during repeated contingency reversals (Experiment 1) and during extinction (Experiment 2) in a picture-picture paradigm. Affective and expectancy learning displayed contingency reversal and extinction, but changes were slower for affective learning. Instructions facilitated reversal and extinction of expectancy learning but did not impact on affective learning. These findings suggest a differential susceptibility of affective and expectancy learning to verbal instruction and question previous reports that verbal instructions can accelerate the extinction of non-prepared fear learning in humans.