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).     

Effects of D-cycloserine on extinction of learned fear to an olfactory cue

D-cycloserine (DCS), a partial NMDA receptor agonist, facilitates extinction of learned fear in rats and has been used to treat anxiety disorders in clinical populations. However, research into the effects of DCS on extinction is still in its infancy, with visual cues being the primary fear-eliciting stimuli under investigation. In both human and animal subjects odors have been found to associate strongly with aversive events. Therefore, this study examined the generality of the effects of DCS on extinction by testing odor cues. Sprague-Dawley rats were conditioned and extinguished to an odor using varying parameters, injected with either saline or DCS (15 mg/kg) following extinction, and then tested for a freezing response 24 h later. Experiment 1 demonstrated that after 3 odor-shock pairings, rats did not display short-term extinction and DCS had no effect on long-term extinction. Experiment 2 demonstrated that after 3 odor-noise pairings, rats displayed significant short-term extinction and DCS significantly facilitated long-term extinction. Following 2 odor-shock pairings in Experiment 3, half the rats displayed short-term extinction ("extinguishers") and half did not ("non-extinguishers"). DCS facilitated long-term extinction in the "extinguishers" condition but not in the "non-extinguishers" condition. In Experiment 4, following 2 odor-shock pairings and an extra extinction session, DCS had a significant facilitatory effect on long-term extinction. Thus, extinction of freezing to an odor cue was facilitated by systemic injections of DCS, but only when some amount of within-session extinction occurred prior to injection.     

Associative status of the training context determines the effectiveness of compound extinction

Studies of extinction of a target cue in compound with another excitor have produced evidence of both deepened and attenuated extinction relative to elemental extinction. The present experiments sought to resolve this discrepancy by assessing the effect of the extinction context-unconditioned stimulus (US) association on compound extinction. In an ABC renewal situation with rats, Experiment 1 replicated the observation that enhanced extinction (i.e., reduced conditioned suppression) occurs as a result of nonreinforcement in compound with a concurrent excitor. In Experiment 2, inflation of the extinction context-US association through unsignaled, intertrial US presentations reversed the effect of a concurrent excitor (i.e., extinction with a concurrent excitor was less effective than elemental extinction). Experiment 3 compared compound extinction and second-order conditioning with respect to the effect of context inflation, and produced results incompatible with the view that second-order conditioning contributed to the results of Experiment 2. In Experiment 4, a concurrent excitor enhanced extinction in ABC renewal and attenuated extinction in AAC renewal. The present results suggest that the effect of a concurrent excitor during extinction depends on the strength of the extinction context-US association, thereby confirming predictions of the sometimes -competing retrieval model of associative acquisition and extinction.     

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.     

Representation-mediated extinction of conditioned flavor aversions

Conditioned flavor aversions were extinguished by presenting without consequence auditory stimuli that had been previously paired with the aversive flavor. In Experiment 1, rats that received tone-sucrose pairings, then sucrose-lithium chloride (LiCl) pairings, and finally repeated tone-alone presentations showed greater sucrose consumption in subsequent testing than rats that received similar sucrose-LiCl pairings and tone-alone presentations but no initial tone-sucrose pairings. Experiment 2 demonstrated the stimulus specificity of the mediated extinction observed in Experiment 1. In Experiment 3, rats that received first-order light-food and second-order tone-light pairings prior to sucrose-LiCl pairings did not show greater subsequent sucrose consumption when extinction of the second-order tone intervened. These results suggest that conditioned stimulus (CS)-evoked representations of events can substitute for those events themselves in the extinction of previously established associations.     

Automaintenance,contrast, and contingencies: Effects of local vs. overall and prior vs. impending reinforcement context

The pigeon's keypeck was investigated in a variety of multiple schedules of response-independent or -dependent reinforcement. Experiments 1 and 2 found that keypecking developed to a reinforcement-associated cue that signaled an increase in local rate of reinforcement, but signaled either no change or a decrease in the overall rate of reinforcement, defined as that prevailing in the cue's absence. In Experiment 3 a greater rate of responding to a target stimulus was observed when it was preceded by a second signal—associated with the same, or a lower, rate of reinforcement—and followed by extinction than when this temporal sequence was reversed. In Experiment 4 responding to a target cue increased when either a temporally prior or a subsequent reinforcement-associated cue was changed to signal extinction. Experiment 5 examined the conditioned reinforcing effectiveness of target cues in two types of situations varying the local context of reinforcement. Stimuli associated with selected target components of a response-dependent multiple schedule of reinforcement could appear as the terminal-link consequences in a two-link chain schedule. Enhanced responding during a target cue which accompanied the introduction of an extinction period following this cue was paralleled by an increase in the conditioned reinforcement effects of this cue. No such increase was found for target cues in which an enhancement of responding had been produced by the interposition of an extinction period prior to the cue.     

Prevention of the degraded-contingency effect by signalling training trials

Presentation of unsignalled unconditioned stimuli (USs) interspersed among Pavlovian excitatory conditioning trials weakens conditioned responding to a target conditioned stimulus (CS; Rescorla, 1968). However, signalling these intertrial USs with another cue (a cover stimulus) has been shown to alleviate this degraded-contingency effect (e.g. Durlach, 1982, 1983). In contrast to signalling the inter-trial USs, the present experiments examined the effect on the degraded-contingency effect of signalling the target CS-US pairings. Experiment 1, using parameters selected to avoid overshadowing, found that consistently presenting a cover stimulus immediately prior to the target CS-US pairings during degraded-contingency training alleviated the degraded-contingency effect. Experiment 2 examined the underlying mechanism responsible for this cover-stimulus effect through posttraining associative inflation of the cover stimulus or the context, and found that inflation of the cover stimulus attenuated responding to the target CS (i.e. empirical retrospective revaluation). The results are discussed in terms of various acquisition- and expression-focused models of acquired responding.     

Contrasting reduced overshadowing and blocking

Preexposure of a cue without an outcome (X-) prior to compound pairings with the outcome (XZ-->O) can reduce overshadowing of a target cue (Z). Moreover, pairing a cue with an outcome (X-->O) before compound training can enhance its ability to compete with another cue (i.e., blocking). Four experiments were conducted in a conditioned bar-press suppression preparation with rats to determine whether spacing of the X- or X-->O trials would differentially affect reduced overshadowing and blocking. Experiment 1a showed that reduced overshadowing was larger with massed trials than with spaced trials. Experiment 1b found that blocking was larger with spaced trials than with massed trials. Experiments 2a and 2b indicated that these effects of trial spacing were both mediated by the associative status of the context at test. The results are interpreted in the framework of contemporary learning theories.     

The resistance of renewal to instructions that devalue the role of contextual cues in a conditioned suppression task with humans

The renewal of extinguished conditioned behaviour appears to reflect context-dependent learning. The present research used a conditioned suppression task with humans to examine whether instructions concerning the context could influence renewal. Pairings of a conditional stimulus (CS) and unconditional stimulus (US) were made in one context, followed by extinction trials of CS alone in a second context, and test trials of CS alone upon return to the original learning context. Four experiments tested whether the renewal of conditioned suppression observed during test would be attenuated if participants were instructed that the context changes were irrelevant to the predictive relationship between the CS and US. Using a differential conditioning design, no attenuation was found when the instructions were given prior to conditioning (Experiment 1) or immediately prior to the test trials (Experiment 2). The latter result was replicated with a single-cue conditioning design and further controls for exposure to the extinction contexts (Experiment 3). The collection of on-line ratings about the relationship between the contextual changes and the predictive nature of the CS indicated that participants did attend to and believe the instructions (Experiment 4). The results point to the resistance of renewal to explicit instructions that attempt to devalue the role of the contextual cues.     

The hide-and-seek of retrospective revaluation: recovery from blocking is context dependent in human causal learning

In a typical blocking procedure, pairings of a compound consisting of 2 stimuli, A and X, with the outcome are preceded by pairings of only A with the outcome (i.e., A+ then AX+). This procedure is known to diminish responding to the target cue (X) relative to a control group that does not receive the preceding training with blocking cue A. We report 2 experiments that investigated the effect of extinguishing a blocking cue on responding to the target cue in a human causal learning paradigm (i.e., A+ and AX+ training followed by A- training). The results indicate that extinguishing a blocking cue increases conditioned responding to the target cue. Moreover, this increase appears to be context dependent, such that increased responding to the target is limited to the context in which extinction of the blocking cue took place. We discuss these findings in the light of associative and propositional learning theories.     

Counterconditioning of an overshadowed cue attenuates overshadowing

In 3 Pavlovian conditioned lick-suppression experiments, rats received overshadowing treatment with a footshock unconditioned stimulus such that Conditioned Stimulus (CS) A overshadowed CS X. Subjects that subsequently received CS X paired with an established signal for saccharin (CS B) exhibited less overshadowing of the X-footshock association than subjects that did not receive the X-B pairings (Experiment 1). Experiment 2 replicated this effect and controlled for some additional alternative accounts of the phenomenon. In Experiment 3, this recovery from overshadowing produced by counterconditioning CS X was attenuated if CS B was massively extinguished prior to counterconditioning. These results are more compatible with models of cue competition that emphasize differences in the expression of associations than those that emphasize differences in associative acquisition.     

The effect of an extinction cue on ABA-renewal: Does valence matter?

The present human fear conditioning study examined whether the valence of an extinction cue has a differential effect on attenuating renewal that is induced by removal of the extinction context. Additionally, the study aimed to assess whether such attenuating effect is based on a modulatory or safety-signal role of the cue. In acquisition, extinction, and test stages of the experiment, human participants received pairings of human faces, presented against a particular background color, with the presence or absence of an aversive auditory stimulus. Experimental groups differed in the valence of a cue that was presented during extinction, in whether or not extinction took place against a different background color than present during acquisition and testing, and in whether the extinction cue was present or absent at test. The conditioned response consisted of auditory-stimulus expectancy ratings. It was hypothesized that a positively valued extinction cue yields faster extinction, stronger attenuation of renewal, and better transfer of its inhibitory power to non-extinguished stimuli than a negatively valued cue. All three hypotheses were confirmed, suggesting that the positive, but not the negative, cue had become a safety signal. The results were discussed in the framework of extinction-based exposure treatments.     

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.     

All we need is a cue to remember: the effect of an extinction cue on renewal

In the present study, it was investigated by employing a human fear conditioning paradigm whether an extinction retrieval cue can attenuate renewal of conditioned responding after an extinction treatment procedure, and if so, what the precise role of such an extinction cue comprises. It was hypothesized that such a cue can attenuate renewal and would function as a safety signal capable of directly inhibiting the expectancy of an aversive outcome and conditioned skin conductance responding to a conditioned stimulus. The results demonstrated that the extinguished expectancy of an aversive outcome was renewed when the CS was presented outside the extinction context and that an extinction cue attenuated this effect. This extinction cue, however, only transferred its inhibitory properties to other, non-extinguished stimuli when there was no contextual switch. This safety signal function was not observed after a switch in context. Possible functions of the extinction cue and its application in extinction-based exposure treatments are discussed.     

No sex difference in contextual control over the expression of latent inhibition and extinction in Pavlovian fear conditioning in rats

Three experiments with Wistar rats searched for a sex difference in contextual control over the expression of latent inhibition and extinction. Experiment 1 used a latent inhibition procedure; Experiments 2 and 3 employed an extinction preparation. All experiments used a shock as the unconditioned stimulus, a tone as the conditioned stimulus, and suppression of food magazine visits as the measure of conditioned responding to the tone. Each experiment revealed a reliable context effect on conditioned responding to the tone; after conditioning in a separate context, conditioned responding in the former latent inhibition or extinction context was attenuated relative to conditioned responding in a control context. There was no sex difference in the magnitude of this effect. These results are discussed in the framework of sex differences in the hippocampus and of the putative role of this structure in various instances of contextual learning.     

Mechanisms of second-order conditioning with a backward conditioned stimulus

Five conditioned suppression experiments with rats examined the conditions under which backward pairings endow a first-order conditioned stimulus (CS1) with the ability to serve as a secondary reinforcer. Experiments 2-5B found evidence for excitatory second-order conditioning (SOC) if, during first-order pairings, the US-CS1 interval was 0 s rather than 3 s. Levels of SOC were comparable after forward and backward pairings (Experiments 1-3), and were unaffected by extinction of CS1 after SOC (Experiment 3). These results suggest that forward and backward CS1s support SOC for the same reason, and they call into question the need to invoke any special mechanism such as memory integration.     

Simultaneous and backward fear conditioning as a function of number of CS-UCS pairings.

Four experiments examined the possibility that the outcome of simultaneous and backward fear conditioning procedures might depend upon the number of CS-UCS pairings. A punishment procedure will rats as subjects and shock as the UCS was used; the amount of suppression produced by response-contingent CS presentations indexed the strength of acquired fear. Experiments 1, 3, and 4 examined the suppressive tendencies of simultaneous-and backward-trained CSs after 0, 10, 20, 40, 80, and 160 pairings. The pattern of data suggested that initial pairings have the effect of increasing suppressive tendencies of the CS, while subsequent pairings decrease them. In addition, evidence of fear inhibition was found after 160 pairings in the case of the backward paradigm. Experiment 2 examined several nonassociative accounts based upon differential shock exposure. Groups given 10 pairings or 80 pairings were compared to ggroups given 10 pairings plus 70 shock-alone presentations. The results indicated that number of pairings, rather than number of UCS occurrences, is the important factor in decreasing the initial suppression. The evidence for eventual inhibition in the backward paradigm suggests that this occurs through the acquisition of inhibitory tendencies which are antagonistic to the previously conditioned excitation.     

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.     

Comparing excitatory backward and forward conditioning

Three Pavlovian lick suppression studies with rats were conducted to compare the role of the conditioning context in excitatory backward and forward conditioning. The experiments explored the possibility that excitatory backward conditioning, but not forward conditioning, is mediated by the context. That is, in excitatory backward conditioning, the conditioning context may function as an excitatory mediator, which supports second-order conditioning of the target cue. This possibility contrasts with traditional accounts, which suggests that common processes underlie excitatory backward and forward conditioning. Experiment 1 found that conditioned responding following backward conditioning was attenuated as a result of posttraining extinction of the training context, but the same manipulation elevated responding after forward conditioning. Experiments 2 and 3 found that posttraining and pretraining associative inflation of the context (presenting unsignalled USs) increased conditioned responding to the target of a backward conditioning procedure but either had no effect or reduced responding to the target of a forward conditioning procedure. Thus, excitatory backward and forward conditioning appear to differ in their dependence on the status of the conditioning context.     

Effects of multiple exposures to D-cycloserine on extinction of conditioned fear in rats

Previous research has shown that an acute, post-training injection of D-cycloserine (DCS) facilitates extinction of conditioned fear in rats; however, the effects of multiple exposures to DCS in this situation are not known. In Experiment 1, rats were conditioned (light-shock pairings) and 24 h later given six extinction (light-alone) trials followed by an injection of DCS (15 mg/kg) or saline. The next day, all rats were tested for light-elicited freezing. In Experiment 2, the effect of DCS on extinction was tested in the same manner, except that rats were pre-exposed to DCS (0, 1, or 5 injections) just prior to conditioning. In Experiment 3, rats received five pre-exposures of DCS but conditioning occurred either 2 or 28 days after the last pre-exposure. The results showed that DCS facilitated extinction of conditioned freezing to the light CS when no drug pre-exposure had occurred, but pre-exposure to DCS just prior to conditioning disrupted the facilitation of extinction effect. When 28 days were interposed between pre-exposure and conditioning, the facilitatory effects of DCS on extinction were restored. These findings suggest that DCS has significant clinical value but that behavioral desensitization may occur with multiple exposures; however, desensitization is not permanent and is reduced by the passage of time.