Extinction and renewal of Pavlovian modulation in human sequential Feature Positive discrimination learning

Using a conditioned suppression task, we investigated extinction and renewal of Pavlovian modulation in human sequential Feature Positive (FP) discrimination learning. In Experiment 1, in context a participants were first trained on two FP discriminations, X-->A+/A- and Y-->B+/B-. Extinction treatment was administered in the acquisition context a (aaa group) or in a new context b (aba group), and comprised X-->A- extinction and Y- control trials. Discriminative X-->A/A responding was lost in both groups when tested in the extinction context, but partially recovered in the aba and not in the aaa group when tested in the acquisition context, suggesting extinction and renewal of extinguished modulation. The same was observed for the Y-->B/B control pair, however, questioning whether the loss of discriminative X-->A/A responding represented genuine extinction of modulation. In Experiment 2, including only aba groups, participants were trained in context a on two FP discriminations, X-->A+/A- and Y-->B+/B-, after which the group "Extinction" was exposed to X-->A- extinction trials in context b, whereas the group "Control" was exposed to X- control trials; concurrently, both groups received further Y-->B+/B- training. In the group Control, differential Y-->B/B and X-->A/A responding were acquired and maintained throughout the experiment. In the group Extinction, while Y-->B/B responding was also maintained throughout, differential X-->A/A responding disappeared because of X-->A- extinction treatment when tested in the extinction context b, but partially reappeared when tested in the acquisition context a. This evidences aba-renewal of extinguished modulation.     

Associative effects of US preexposure: modulation of conditioned responding by an excitatory training context

In two experiments we examined factors that contribute to retarded emergence of conditioned responding to a conditioned stimulus (CS) trained in a context in which unsignaled unconditioned stimuli (USs) had previously been administered. In both experiments water-deprived rats were used in a conditioned lick suppression task to measure the conditioned response elicitation potential of the CS and the training context. From Experiment 1 we determined that nonreinforced exposure to the excitatory context after US preexposure and prior to CS-US pairings in that context eliminated the conditioned response deficit observed on a subsequent test of the CS. The recovery from the US preexposure deficit was nearly as great in animals that received nonreinforced exposure to the excitatory training context after the CS-US pairings but prior to the ultimate test of the CS. From Experiment 2 we determined that the recovery induced by contextual deflation after CS training was specific to deflation of the context in which the CS was trained as opposed to another excitatory context. In total, these experiments suggest that context-US associations partially mask the expression of a learned CS-US association. These results are discussed in terms of recent models of conditioned response generation.     

Associations in Pavlovian conditioned inhibition

Two experiments with rat subjects used a variety of transfer tests to examine the associations learned when Pavlovian inhibition is established by an A+, AX− paradigm. Experiment 1 found in a conditioned suppression situation that inhibition conditioned to X with one exciter (A) readily transferred to another exciter (B) which had been paired with the same shock US. Transfer occurred even when the response to A had been extinguished prior to testing with B. However, X did not inhibit a general activity response produced by a B which had been subsequently paired with a food US. Experiment 2 employed a Pavlovian conditioning situation in which A and B, when separately paired with the same food US, evoked dissimilar responses. Nevertheless, an inhibitor trained in an A+, AX− paradigm successfully inhibited the different response evoked by B. However, such an X did not inhibit the behaviors acquired by A or B when they were subsequently paired with a shock US. The transfer of Pavlovian inhibition across conditioned stimuli and responses but not across unconditioned stimuli is consistent with the notion that a conditioned inhibitor acts to prevent activation of a US representation which would normally be activated by conditioned exciters.     

Blocking of occasion setting in feature-positive discriminations

In two experiments rats received feature-positive discrimination training in which brief conditioned stimuli (CSs) were paired with food during presentations of an extended feature stimulus, and non-reinforced in its absence. In Experiment 1 a novel feature was trained in compound with a second, pretrained feature. Acquisition of control over responding to the CS by the novel feature was blocked if the pretrained feature had also been trained in a feature-positive discrimination, compared to a group for whom the pretrained feature had been accompanied by uncorrelated presentations of CSs and food. Experiment 2 employed a within-subjects design. It demonstrated that the feature from a feature-positive discrimination with a particular CS, x, blocked acquisition of control by an added, novel feature over responding to x, compared to the control acquired by the same novel feature over a novel, CS y.     

Conditioned Inhibition Produced by Extinction of a Conditioned Stimulus

Four experiments used a conditioned taste aversion procedure to examine the potential for CS-alone extinction treatment to produce a conditioned stimulus that possesses inhibitory properties. In Experiment 1, saccharin was paired with LiCl, and then saccharin was presented alone for several trials to produce extensive behavioral extinction. Animals receiving this treatment were retarded in reacquiring conditioned responding to saccharin relative to control subjects receiving conditioning to the flavor for the first time. In Experiment 2, the extinguished saccharin stimulus was shown to decrease conditioned responding to a known excitor when the two stimuli were presented in compound as a summation test. Experiments 3A and 3B replicated the findings of Experiments 1 and 2 while providing evidence that the effects were not due to the differential effects of neophobia during testing. These three experiments revealed that an extinguished conditioned excitor passes retardation and summation tests for conditioned inhibition. Experiment 4 found that extinction of a known excitor was slowed when the excitor was extinguished in compound with a previously extinguished conditioned stimulus. That is, an extinguished CS provided protection from extinction to another CS, a finding also consistent with the view that extinction produces conditioned inhibition.     

Interactions between conditioned stimuli for food and water in the rat

In two experiments rats were initially trained with one conditioned stimulus (CS) signalling food and another CS signalling water. The main purpose of the experiments was to examine the effects of presenting these stimuli in compound. Experiment 1 demonstrated that the level of conditioned responding to this compound was equal to that of a group for which the compound was composed of elements that had separately signalled the same reinforcer. Moreover, the level of this responding was greater than for a group that had received only compound conditioning. In the second experiment it was found that after the initial training, repeatedly pairing the compound with a single reinforcer resulted in an equivalent loss of conditioned responding to both elements. These findings suggest that the interaction between conditioned stimuli for food and water is facilitatory rather than inhibitory.     

Sequential and simultaneous feature positive discriminations: occasion setting and configural learning in human Pavlovian conditioning

Using a conditioned suppression preparation, the authors investigated sequential (X --> A+/A-) versus simultaneous (XA+/A-) feature positive (FP) discrimination learning in humans. The sequential discrimination was expected to be resolved by means of a Feature X Modulated Target A-US association and the simultaneous discrimination by a feature X-US association. After sequential FP training, extinction of Feature X did not affect discriminative X --> A/A responding (Experiment 1), and X transferred its modulatory ability only to new targets, B, that had also been modulated (Experiment 2). This suggests that the sequential FP discrimination indeed resulted in occasion setting. Unlike expected, Feature X Extinction did not affect discriminative XA/A responding after simultaneous FP training (Experiment 3), while at the same time Feature X did show the predicted nonselective transfer to new targets, B (Experiment 4). J. M. Pearce's (1987) configural learning theory can account for most but not all findings of Experiments 3 and 4.     

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.     

Overshadowing and associability change

Three appetitive Pavlovian conditioning experiments with rats examined the associability of stimuli A and B that had a history of compound conditioning (AB+), relative to stimuli X and Y that had a history of conditioning in isolation (X+, Y+). Following this training, Experiment 1 revealed that conditioned responding was higher to X and Y than to A and B (overshadowing). In a subsequent AY+, AX-, BY- test discrimination, the AY/BY discrimination was solved more readily than the AY/AX discrimination. In Experiment 2, following AB+, X+, Y+ training, A and Y were presented as a compound and signaled the availability of reinforcement upon the performance of an instrumental response. Test trials in which A and Y were presented alone, and in extinction, revealed that A acquired greater control of instrumental responding than Y. Experiment 3 revealed that following AB+, X+, Y+ training, A and B served as more effective discriminative stimuli for instrumental responding than X and Y. Overall, these results imply that the associability of stimuli conditioned in compound is higher than stimuli conditioned in isolation. These results are discussed in terms of attentional theories of associative learning.     

The effects of physical context changes and multiple extinction contexts on two forms of renewal in a conditioned suppression task with humans

Three experiments examined the effects of physical context changes and multiple extinction contexts on the renewal of conditioned suppression in humans. A conditioned suppression task used an undesirable event as the unconditional stimulus (US). One conditional stimulus (CS+) predicted the occurrence of the US and another (CS−) predicted US absence. In Experiment 1 (N = 32), conditioned suppression was acquired to the CS+ in one context and extinguished in a different context. An increase in suppression was found for the CS+ and not for the CS− when subsequent test trials were conducted in the acquisition context (ABA renewal). Experiment 2 (N = 32) tested for ABC Renewal and showed increased suppression to both the CS+ and CS− when test was conducted in a novel context. Experiment 3 (N = 80) showed that these two effects were abolished when extinction was conducted in multiple contexts. The experiments extend the ABA renewal of conditioned suppression found with non-human animal subjects and the reduction of renewal by extinction in multiple contexts. Context changes may also facilitate cue competition effects after training with elementary stimuli, as shown by the effects of US omission in the ABA and ABC renewal groups.     

Effects of contextual conditioning and unconditional stimulus presentation on performance in appetitive conditioning

Four experiments with rat subjects examined the effects of contextual conditioning on conditioned appetitive performance. Experiment 1 compared the effects of contextual conditioning on performance to conditioned stimuli (CSs) with different conditioning histories. Contextual conditioning enhanced performance to the CS if the CS had first been conditioned and then extinguished, but had no effect on performance when the CS had been merely paired or unpaired with food. Experiments 2 and 3 then asked whether the effect on the extinguished CS was due to contextual conditioning acting as a cue for conditioning. In Experiment 2, extinction procedures in which extra unconditioned stimuli (USs) were presented during the intertrial intervals were found to reduce the CS's sensitivity to enhancement by contextual conditioning, but had no effect on spontaneous recovery. In Experiment 3, USs added to conditioning or extinction acquired the ability to cue the corresponding performance. Under some conditions, USs added to conditioning could suppress performance (Experiment 4). The results suggest that contextual conditioning has complex effects that can be better understood by recognizing that contextual conditioning, as well as the USs that create it,Mayacquire discriminative control over conditioned responding.     

The comparator hypothesis of conditioned response generation: manifest conditioned excitation and inhibition as a function of relative excitatory strengths of CS and conditioning context at the time of testing

In the present research water-deprived rats were used in a conditioned lick suppression paradigm to test and further develop Rescorla's (1968) contingency theory, which posits that excitatory associations are formed when a conditioned stimulus (CS) signals an increase in unconditioned stimulus (US) likelihood and that inhibitory associations develop when the CS signals a decrease in US likelihood. In Experiment 1 we found that responding to a CS varied inversely with the associative status of the context in which the CS was trained and that this response was unaltered when testing occurred in a distinctively dissimilar context with a different conditioning history, provided associative summation with the test context was minimized. These results suggest that manifest excitatory and inhibitory conditioned responding is modulated by the associative value of the training context rather than that of the test context. In Experiment 2 it was demonstrated that postconditioning decreases in the associative value of the CS training context reduced the effective inhibitory value of the CS even when testing occurred outside of the training context. Moreover, this contextual deflation effect was specific to the CS training context as opposed to any other excitatory context. Collectively, these studies support the comparator hypothesis, which states that conditioned responding is determined by a comparison of the associative strengths of the CS and its training context that occurs at the time of testing rather than at the time of conditioning. This implies that all associations are excitatory and that responding indicative of conditioned inhibition reflects a CS-US association that is below (or near) the associative strength of its comparator stimulus. It is suggested that response rules which go beyond a monotonic relation between associative value and response strength can partially relieve learning theories of their explanatory burdens, thereby allowing for simpler models of acquisition.     

Effects of contingency violations on the extinction of a conditioned fear inhibitor and a conditioned fear excitor

Rats were used in a conditioned-suppression paradigm to assess the effects of contingency variations on responding to a conditioned inhibitor (CS-) and a conditioned excitor (CS+). In Experiment 1, various unconditioned stimulus (US) frequencies were equated across the presence and absence of a CS- in the context of either background cues (continuous-trial procedure) or an explicit neutral event (discrete-trial procedure). With both procedures, a CS-alone treatment enhanced inhibition, whereas treatments involving 50% or 100% reinforcement for the CS- eliminated inhibition without conditioning excitation to that CS. The latter outcome also occurred in Experiment 2, with discrete-trial training equating considerably reduced US frequencies for the presence and absence of the CS-. In further evidence that inhibition was eliminated without conditioning excitation to the CS-, Experiment 3 showed that a novel CS did not acquire excitation when 25%, 50%, or 100% reinforcement was equated across the presence and absence of that CS in the context of a discrete-trial event. Using the procedures of Experiment 1, Experiment 4 showed that a CS+ was extinguished by a CS-alone treatment but was substantially maintained by treatments involving 50% or 100% uncorrelated reinforcement. These effects for a CS+ and a CS- implicate CS-US contiguity, rather than contingency, as the factor determining the extinction of a CS.     

The influence of associability changes in negative patterning and other discriminations

Normal rats showed faster learning of a serial negative patterning (NP) discrimination (X+, A+, X-->A-) than of a comparable feature negative (FN) discrimination (A+, X-->A-). This advantage was absent in rats with lesions of the amygdala central nucleus. Earlier data indicated that this brain lesion interferes with surprise-induced increases in attention specified by the Pearce-Hall model (J. M. Pearce & G. Hall, 1980). In the NP task, but not the FN task, omission of the reinforcer after X on X-->A- trials was surprising. A variation of the NP task (NPX), in which X was reinforced on both X+ and X-->A- trials, was learned more rapidly than the NP task. Lesioned rats were unimpaired in learning the NPX task. Evaluation of the lesion effects and the results of posttraining transfer tests suggested that the NP advantage involved attentional processes, whereas the NPX advantage was based on the acquisition of inhibitory control by aspects of excitation conditioned to X.     

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).     

Facilitation of instrumental behavior by a Pavlovian appetitive conditioned stimulus

Three experiments examined appetitive Pavlovian-instrumental interactions by presenting separately trained conditioned stimuli (CSs) during reinforced instrumental responding in rabbits. Intra-oral reinforcement was used to minimize interference from peripheral responses such as magazine approach. In experiment 1, the rabbits were first trained to perform an instrumental head-raising response for sucrose reward. A conditioned jaw movement response was then established to a 2-sec CS by pairing it with sucrose; a control stimulus was unpaired with sucrose. Instrumental responding maintained by a variable-interval 40-sec schedule was enhanced during 10-sec presentations of the paired, but not the unpaired, CS. Responding on a variable-ratio 15 schedule was unaffected except on trials on which the pre-CS baseline response rate was low; in such cases the paired CS caused a long-lasting acceleration of responding. Noncontingent presentation of the sucrose reinforcer itself briefly suppressed responding but had no long-term effect. In Experiment 2, a CS that had been conditioned at a 10-sec duration produced the same pattern of effects as in the first study, indicating that facilitation resulted from CS presentation rather than from the frustrative effects of non-reinforcement of the CS. In Experiment 3 an inhibitory CS blocked facilitation by the excitatory CS but did not itself affect instrumental responding. These results support the view that Pavlovian processes play a positive role in instrumental performance and suggest that previous findings of suppression by a short-duration CS reflect peripheral interference. The dependence of facilitation on the baseline level of responding is discussed in terms of associative and motivational theories of Pavlovian mediation.     

The impact of arbitrarily applicable relational responding on evaluative learning about hypothetical money and shock outcomes

Evaluative learning comprises changes in preferences after co-occurrences between conditioned stimuli (CSs) and an unconditioned stimulus (US) of affective value. Co-occurrences may involve relational responding. Two experiments examined the impact of arbitrary relational responding on evaluative preferences for hypothetical money and shock outcomes. In Experiment 1, participants were trained to make arbitrary relational responses by placing CSs of the same size but different colours into boxes and were then instructed that these CSs represented different intensities of hypothetical USs (money or shock). Liking ratings of the CSs were altered in accordance with the underlying bigger/smaller than relations. A reversal of preference was also observed: the CS associated with the smallest hypothetical shock was rated more positively than the CS associated with the smallest amount of hypothetical money. In Experiment 2, procedures from Relational Frame Theory (RFT) established a relational network of more than/less than relations consisting of five CSs (A-B-C-D-E). Overall, evaluative preferences were altered, but not reversed, depending on (a) how stimuli had been related to one another during the learning phase and (b) whether those stimuli referred to money or shocks. The contribution of RFT to evaluative learning research is discussed.     

Transfer and Counterconditioning of Conditional Control in the Rabbit Nictitating Membrane Response

Two experiments using the rabbit nictitating membrane response investigated whether training in one conditional discrimination (A X+, B X-), enabled the feature cues (A and B) to modulate responding to another CS (Y) trained as a target stimulus in a second conditional discrimination (C Y+, D Y-). There was near-complete transfer of the feature cue's conditional control, indicating that the feature cue's ability to modulate responding is not based on an association specific to the training target. Experiment 2 also revealed that the role of a stimulus to act as a conditional cue is affected by its ability to act as a simple conditioned excitor or inhibitor. Following initial acquisition of two conditional discriminations, two feature cues were reinforced in a pattern consistent with the initial conditional discrimination (A +, B-), whereas the other two feature cues were reinforced in the reverse pattern to that of the original conditional discrimination (C-, D +). Subsequent tests revealed that the reversed training of the feature cues interfered with the original conditional discriminations. The results are consistent with theories that the feature cue gains an association with a representation of the emotional attributes of the US, which acts to modulate responding to the target stimulus through a diffuse change in motivational level. However, hierarchical theories of conditional discriminations that assume a lack of CS-specificity may also be able to explain the findings.     

Temporal specificity of extinction in autoshaping

Three experiments investigated the effects of varying the conditioned stimulus (CS) duration between training and extinction. Ring doves (Streptopelia risoria) were autoshaped on a fixed CS-unconditioned stimulus (US) interval and extinguished with CS presentations that were longer, shorter, or the same as the training duration. During a subsequent test session, the training CS duration was reintroduced. Results suggest that the cessation of responding during an extinction session is controlled by generalization of excitation between the training and extinction CSs and by the number of nonreinforced CS presentations. Transfer of extinction to the training CS is controlled by the similarity between the extinction and training CSs. Extinction learning is temporally specific.     

Inhibition as a "slave" process: deactivation of conditioned inhibition through extinction of conditioned excitation

Rats were used in a conditioned-suppression paradigm to investigate why a conditioned inhibition (CS-) does not extinguish when presented alone. Experiment 1 assessed the role of blocking by excitatory contextual cues and/or an evoked representation of the conditioned excitor (CS+), which had been nonreinforced in compound with the CS-. When the CS+ and context were extinguished prior to presentations of the CS- alone, the CS- showed a retardation effect, evidently reflecting latent inhibition, because no inhibition was detected in controls for which presentations of the CS- alone had been omitted. Experiment 2 showed that the loss of conditioned inhibition (CI) was due to excitatory extinction and not to time since conditioning. Furthermore, when excitation was reconditioned to the extinguished CS+ (Experiment 1), or to a novel CS in the same context (Experiment 2), CI was restored. Two other experiments evaluated whether the maintenance of CI depended upon excitation that was generic in form or associatively tied to the training context. They showed no loss of CI when groups received CS+ extinction in that context, along with concomitant presentations in a different context of the US by itself, for a novel CS, or correlated either positively or negatively with the original CS+. Collectively, the findings argue that CI is a "slave" to excitation, for when excitation is extinguished, CI is deactivated; and yet when excitation is reconditioned to the original or a new CS+ in the same or a different context, CI is restored.