Recovery from the overexpectation effect: Contrasting performance-focused and acquisition-focused models of retrospective revaluation

In four Pavlovian conditioned lick-suppression experiments, rats had two conditioned stimuli (CSs X and A) independently paired with footshock, followed by pairings of a compound of A and X with the footshock. On subsequent tests with CS X, less conditioned suppression was observed than in control subjects that lacked the compound AX→footshock trials. Thisoverexpectation effect was reversed through posttraining extinction of CS A, a result consistent with both performance- and acquisition-focused models of retrospective revaluation. However, only performance-focused models could account for how posttraining increases or decreases in the A-footshock temporal interval attenuate the overexpectation effect.     

Overshadowing of explicitly unpaired conditioned inhibition is disrupted by preexposure to the overshadowed inhibitor

In two conditioned lick-suppression experiments with rats, the interaction of preexposure to a target stimulus and subsequent overshadowing of conditioned inhibition treatment was examined. Blaisdell, Bristol, Gunther, and Miller (1998) have demonstrated that stimulus preexposure and overshadowing counteract each other in their effects on Pavlovian excitation, producing strong excitatory responding to the target stimulus. In the present experiments, a conditioned inhibition analogue of this effect was examined. Overshadowing of conditioned inhibition (i.e., attenuation of inhibitory control of behavior by the target stimulus) was demonstrated when a more salient stimulus was compounded with the target inhibitor during conditioned inhibition training. However, the overshadowing deficit was attenuated by preexposing the target stimulus prior to overshadowing treatment. To account for this phenomenon, we contrast the comparator hypothesis (Miller & Matzel, 1988) with contemporary models that focus on associative acquisition.     

When more is less: Extending training of the blocking association following compound training attenuates the blocking effect

Three conditioned lick suppression experiments with rats were performed to assess the influence, following compound training of two stimuli (A and X) with the same outcome (AX-O trials), of extending training of the blocking association (i.e., A-O) on responding to the target stimulus (X) at test. In Experiment 1, backward blocking was attenuated when the blocking association was extensively trained. Experiment 2 showed that forward blocking was also attenuated by extensive further training of the blocking association following the AX-O trials. Experiment 3 contrasted candidate explanations of the results of Experiments 1 and 2 and demonstrated that these results are consistent with the framework of the extended comparator hypothesis (Denniston, Savastano, & Miller, 2001).     

Associative learning phenomena in the snail (Helix aspersa): conditioned inhibition

Two experiments using garden snails (Helix aspersa) showed conditioned inhibition using both retardation and summation tests. Conditioned inhibition is a procedure by which a stimulus becomes a predictor of the absence of a relevant event—the unconditioned stimulus (US). Typically, conditioned inhibition consists of pairings between an initially neutral conditioned stimulus, CS₂, and an effective excitatory conditioned stimulus, CS₁, in the absence of the US. Retardation and summation tests are required in order to confirm that CS₂ has acquired inhibitory properties. Conditioned inhibition has previously been found in invertebrates; however, these demonstrations did not use the retardation and summation tests required for an unambiguous demonstration of inhibition, allowing for alternative explanations. The implications of our results for the fields of comparative cognition and invertebrate physiological models of learning are discussed.     

Retardation of conditioned excitation following operational inhibitory blocking

Water-deprived adult rats were used in a conditioned-suppression-of-licking procedure to determine the effect of inhibitory training with a novel stimulus trained in simultaneous compound with a previously established conditioned inhibitor. This procedure constitutes an inhibitory analogue to the excitatory blocking procedure in classical conditioning. The conditioned-inhibition training consisted of either explicitly unpaired CS and US presentations or negative contingency training, in which the likelihood of the US was greater in the absence than in the presence of the CS, but the CS and the US were occasionally paired. To assess conditioned inhibition, a retardation test was used, and comparable retardation was obtained for subjects that were administered the blocking treatment and control subjects given similar conditioned-inhibition training with a compound stimulus in which the nontarget element was not previously established as a conditioned inhibitor.     

Divergence of conditioned eyeblink and conditioned fear in backward Pavlovian training

Two experiments of Pavlovian conditioning with rabbits evaluated the effects of initiating or continuing a conditioned stimulus (CS) after a paraorbital unconditioned stimulus (US). In Experiment 1, backward pairings, in which a CS came on after the US, produced a CS that appeared inhibitory on a measure of eyeblink conditioning but excitatory on a potentiated-startle measure of conditioned fear. In Experiment 2, extending the duration of a CS that came on prior to the US, so that it continued after the US, decreased eyeblink conditioned responses, whereas it increased conditioned fear. The data from the two experiments confirm and extend those of Tait and Saladin (1986), supporting the suppositions of AESOP (Wagner & Brandon, 1989) that conditioned eyeblink and conditioned fear can be dissociated under various temporal relationships between the CS and US.     

Conditioned inhibitory effects of discriminated Pavlovian training with food in rats depend on interactions of search modes, related repertoires, and response measures

Like other accounts of conditioned inhibition, behavior systems predicts (and Experiment 1 shows) that during summation and retardation tests, presentation of a negative conditioned stimulus (a CS−) created by discriminative Pavlovian food conditioning will interfere with a focal search response, such as nosing in the feeder. Unlike most other views, behavior systems predicts (and Experiment 2 shows) that the same CS− can potentiate a general search response, like attending to a moving artificial prey stimulus. Contacting the prey stimulus in extinction increased over baseline when a CS- but not a CS Novel preceded it. Experiment 3 showed this effect was not due to unconditioned qualities of the CS−. It appears that the effects of a discriminative CS-depend on the interaction of the training contingency with search modes related to the unconditioned stimulus (US), their perceptual-motor repertoires and environmental support, and the choice of response measure.     

Reminder-induced attenuation of the effect of relative stimulus validity

The roles of deficient acquisition and deficient expression of learned information in the effect of relative stimulus validity were examined using rats in a conditioned lick suppression paradigm. Recovery from the effect without further pairings of the conditioned stimulus (CS) and the unconditioned stimulus (US) would favor an interpretation of the relative validity effect based on a latent CS-US association as distinct from a failure to acquire the CS-US association. As a potential recovery manipulation, “reminder” treatments, consisting of the US alone (Experiment 1) or the CS alone (Experiment 2), were administered following relative validity training. In both cases, subjects for which the CS target was of low relative predictive validity exhibited enhanced responding relative to appropriate controls. Additionally, Experiment 2 showed that the amelioration of the relative validity deficit was stimulus specific. Thus, the results of these experiments support previous suggestions that the performance deficit resulting from low relative stimulus validity is due, at least in part, to a failure to express acquired information (Cole, Barnet, & Miller, 1995a).     

Conditioned inhibition training of the competing cue after compound conditioning does not reduce cue competition

In each of two experiments, we studied Pavlovian fear conditioning (as assessed by barpress conditioned suppression) in 32 albino rats. Following a two-stage cue-competition procedure (A+ then AX+), we subjected the competing cue (A) to conditioned inhibition training (B+, BA?) before testing the target cue (X). Conditioned inhibition training was designed to weaken the putative A-unconditioned stimulus (US) association, perhaps changing it to an A-no-US association. Performance-deficit theories of cue competition, such as comparator theory and retrieval-interference theory, predict that such procedures should weaken cue competition, causing Conditioned Stimulus X (CS X) to evoke strong responding. The same prediction can be deduced from recent acquisition-focused models (Dickinson &; Burke, 1996; Van Hamme &; Wasserman, 1994). In opposition to this prediction, however, we found in both experiments that conditioned inhibition training had no detectable effect on cue competition even though it successfully abolished conditioned responding to CS A. In Experiment 2, moreover, we found evidence against the hypothesis that the weak response to CS X was due to generalization decrement rather than to cue competition. Results favor early learning-deficit theories of cue competition over performance-deficit theories and over the recent acquisition-focused models.     

Conditioned avoidance responses survive contingency degradation in the garden slug,Lehmannia valentiana

Joint presentations of a conditioned stimulus (CS) and an unconditioned stimulus (US) strengthen the contingency between them, whereas presentations of one stimulus without the other degrade this contingency. For example, the CS can be presented without the US either before conditioning (CS–no US and then CS–US; latent inhibition) or after conditioning (CS–US and then CS–no US; extinction). In vertebrate subjects and several invertebrate species, a time lapse usually results in a return of the conditioned response, or spontaneous recovery. However, in land mollusks, spontaneous recovery from extinction has only recently been reported, and response recovery after latent inhibition has not been reported. In two experiments, using conditioned aversion to a food odor as a measure of learning and memory retention, we observed contingency degradation via latent inhibition (Experiment 1) and extinction (Experiment 2) in the common garden slug, Lehmannia valentiana. In both situations, the contingency degradation procedure successfully attenuated conditioned responding, and delaying testing by several days resulted in recovery of the conditioned response. This suggests that the CS–US association survived the degradation manipulation and was retained over an interval of several days.     

Overshadowing as a function of trial number: Dynamics of first- and second-order comparator effects

In two conditioned lick suppression experiments with rats, we examined the permanence of the overshadowing effect as a function of the number of compound reinforced training trials. In Experiment 1, robust overshadowing was observed following 4 compound-US pairings but dissipated with 36 pairings. Overshadowing decreased because responding to the overshadowed stimulus increased, not because responding by the control group decreased. This dissipation was stimulus specific and not attributable to a response ceiling. Experiment 2 extended the generality of the effect to a sensory preconditioning design and further demonstrated that overshadowing lost through many compound-US pairings was restored by posttraining extinction of the training context. The results are explicable in terms of the extended comparator hypothesis (Denniston, Savastano, & Miller, 2001) under the assumption that the impacts of first- and second-order comparator processes grow differentially as a function of number of trials.     

Latent inhibition as a performance deficit resulting from CS—context associations

Treatments that attenuate latent inhibition (LI) were examined using conditioned suppression in rats. In Experiment 1, retarded conditioned responding was produced by nonreinforced exposure to the CS prior to the CS-US pairings used to assess retardation (i.e., conventional LI). In Experiment la, retarded conditioned responding was induced by preexposure to pairings of the CS and a weak US prior to retardation-test pairings of the CS with a strong US (i.e., Hall-Pearce [1979] LI). Both types of LI were attenuated by extensive exposure to the training context (i.e., context extinction) following the CS-US pairings of the retardation test. Experiment 2 examined the specificity of the attenuated LI effect observed in Experiment 1. After preexposure to two different CSs in two different contexts, each CS was paired with a US in its respective preexposure context. One of the two contexts was then extinguished. This attenuated LI to a greater degree for the CS that had been trained in the extinguished context. Experiment 3 differentiated the roles in LI of CS-context associations and context-US associations. Following preexposure to the CS in the training context, LI was reduced by further exposure to the CS outside the training context. This observation was interpreted as implicating the CS-context association as a factor in LI. Thus, the results of these experiments suggest that LI is a performance deficit mediated by unusually strong CS-context associations. Implications for Wagner’s (1981) SOP model and Miller and Matzel’s (1988) comparator hypothesis are discussed.     

Renewal of Pavlovian conditioned inhibition

Prior research has demonstrated renewal, which is the ability of contextual cues to modulate excitatory responding to a Pavlovian conditioned stimulus (CS). In the present research, conditioned lick suppression in rats was used to examine similar contextual modulation of Pavlovian conditioned inhibition. After Pavlovian conditioned inhibition training with a CS in one context, subjects were exposed to pairings of the CS with an unconditioned stimulus (US) either in the same or in a second context. Results indicated that, when the CS was paired with the US in the second context, the CS retained its inhibitory control over behavior, provided that testing occurred in the context used for inhibition training. However, when the CS-US pairings occurred in the inhibition training context, the CS subsequently proved to be excitatory regardless of where testing occurred. These observations indicate that conditioned inhibition is subject to renewal.     

Local context and the comparator hypothesis

“Comparator” accounts of associative conditioning (e.g., Gibbon & Balsam, 1981; Miller & Matzel, 1988) suggest that performance to a Pavlovian CS is determined, by a comparison of the US expectancy of the CS with the US expectancy of general background cues. Recent research indicates that variation in the excitatory value of cues in the local temporal context of a CS may have a profound impact on conditioned responding to the CS (e.g., Kaplan & Hearst, 1982), implicating US expectancy based on local, rather than overall, background cues as the critical comparator term for a CS. In two experiments, an excitatory training context attenuated responding to a target CS. In Experiment 1, the context was made excitatory by interspersing unsignaled USs with target CS-US trials. In this case, posttraining extinction of the conditioning context restored responding to the target CS. In Experiment 2, the target CS’s local context was made excitatory by the placement of excitatory “cover” stimuli in the immediate temporal proximity of each target CS-US trial. In this experiment, posttraining extinction of the proximal cover stimuli, not extinction of the conditioning context alone, restored responding to the target CS. An observation from both experiments was that signaling the otherwise unsignaled USs did not appear to influence the associative value of the conditioning context. The results are discussed in relation to a local context version of the comparator hypothesis and serve to emphasize the importance of local context cues in the modulation of acquired behavior. Taken together with other recent reports (e.g., Cooper, Aronson, Balsam, & Gibbon, 1990; Schachtman & Reilly, 1987), the present observations encourage contemporary comparator theories to reevaluate which aspects of the conditioning situation comprise the CS’s comparator term.     

Intertrial unconditioned stimuli differentially impact trace conditioning

Three experiments assessed how appetitive conditioning in rats changes over the duration of a trace conditioned stimulus (CS) when unsignaled unconditioned stimuli (USs) are introduced into the intertrial interval. In Experiment 1, a target US occurred at a fixed time either shortly before (embedded), shortly after (trace), or at the same time (delay) as the offset of a 120-s CS. During the CS, responding was most suppressed by intertrial USs in the trace group, less so in the delay group, and least in the embedded group. Unreinforced probe trials revealed a bell-shaped curve centered on the normal US arrival time during the trace interval, suggesting that temporally specific learning occurred both with and without intertrial USs. Experiments 2a and 2b confirmed that the bulk of the trace CS became inhibitory when intertrial USs were scheduled, as measured by summation and retardation tests, even though CS offset evoked a temporally precise conditioned response. Thus, an inhibitory CS may give rise to new stimuli specifically linked to its termination, which are excitatory. A modification to the microstimulus temporal difference model is offered to account for the data.     

CS-duration and partial-reinforcement effects counteract overshadowing in select situations

Two experiments used rats in a conditioned lick suppression preparation to investigate how the conditioned stimulus (CS)-duration and partial-reinforcement effects (i.e., weakened responding due to conditioning with a CS of longer duration and presenting nonreinforced CSs intermingled with CS—unconditioned stimulus [US] pairings, respectively) interact with overshadowing. Experiment 1 found that when overshadowing treatment was combined with either extended CS duration or partial reinforcement, the response deficit was weaker than when either of these three treatments was administered alone. In Experiment 2, the generality of the findings in Experiment 1 was investigated by replicating it with various US—US intervals. This time counteraction was observed only when both the absolute duration of total CS exposure and the US—US interval were short. The results support neither the view that the ratio between the total CS exposure and total time in the context determines the CS-duration and the partial-reinforcement effects nor the view that these two effects arise from a loss of effectiveness of the excitatory CS—US association during CS-alone exposures in partial reinforcement or early periods of CS exposure with long CSs.     

Temporal integration in Pavlovian appetitive conditioning in rats

We used an appetitive sensory preconditioning procedure to investigate temporal integration in rats in two experiments. In Phase 1, rats were presented with simultaneous compound trials on which a 10-sec conditioned stimulus (CS) X was embedded within a 60-sec CS A. In Group Early, CS X occurred during the early portion of CS A, whereas in Group Late, CS X occurred during the latter portion of CS A. In Phase 2, CS X was paired simultaneously with sucrose. On a subsequent test with CS A, the rate of magazine entries peaked during the early portions of the stimulus in Group Early and during the latter portions of the stimulus in Group Late (Experiments 1 and 2). Similar response peaks were not observed on tests with a control stimulus that had been presented in compound with a stimulus that did not signal reward (Experiment 2).     

Posttraining flavor exposure in hungry rats after simultaneous conditioning with a nutrient converts the CS into a conditioned inhibitor

The present study investigated the decrement in nutrient-based conditioned flavor preference found in hungry rats exposed to a flavor following simultaneous flavor–sucrose conditioning while thirsty. Although a significant decrease in preference was found in the experimental group in each experiment, there was no evidence of either spontaneous recovery (Experiment 1) or reinstatement (Experiment 2). In addition, posttraining flavor exposure weakened the original flavor–sucrose association (Experiment 3). These results suggested that the flavor–US association might have been impaired after posttraining flavor exposure. Two further experiments assessed whether the flavor acquired the properties of a net inhibitor, using the retardation and summation tests for conditioned inhibition. Experiment 4 revealed that the flavor suffered retardation when retraining was conducted after the exposure phase. In Experiment 5, the target flavor decreased the preference shown for a different flavor previously paired simultaneously with sucrose when both were presented forming an unreinforced compound in the summation tests. None of these effects was found in a control group, which had received serial flavor → nutrient presentations during training. Together, these results suggest that a flavor simultaneously paired with sucrose acquires the properties of a net inhibitor when it is subsequently presented outside the compound to hungry animals.     

Temporal integration in second-order conditioning and sensory preconditioning

Lick suppression experiments with rats revealed that the magnitude of both second-order conditioning (Experiment 1) and sensory preconditioning (Experiment 2) was superior when that conditioning was based on backward (US→CS) relative to forward (CS→US) first-order pairings of a CS and US. The superiority of backward relative to forward first-order conditioning on suppression to the higher order cues can be understood by assuming that the magnitude of higher order conditioning was determined by a memory representation of the higher order cues that provided information about the expected temporal location of the US. The results suggest that temporal information such as order between paired CSs and USs was encoded, preserved, and integrated with memory for the higher order stimuli. The relevance of these findings to memory integration in Pavlovian learning, the temporal coding hypothesis (Barnet, Arnold, & Miller, 1991; Matzel, Held, & Miller, 1988), backward excitatory conditioning, and the associative structure that underlies second-order Pavlovian fear conditioning are discussed.     

The role of temporal variables in inhibition produced through extinction

In two experiments with rats as subjects, the temporal characteristics of inhibition produced through extinction were investigated. Each experiment established two independent signals for unconditioned stimulus presentation, one trace and one delay. Following initial training, either the trace or the delay conditioned stimulus (CS) was massively extinguished. In Experiment 1, a summation test established that an extinguished delay CS (but not a neutral CS) passed a summation test with a delay, but not with a trace, transfer excitor, and an extinguished trace CS (but not a neutral CS) passed a summation test with a trace, but not with a delay, transfer excitor. In Experiment 2, a retardation test showed retarded behavioral control by an extinguished delay CS when the CS was retrained as a delay CS, but not as a trace CS, and by an extinguished trace CS when the CS was retrained as a trace CS, but not as a delay CS. The results are discussed in terms of contemporary theories of extinction.