Temporal encoding in trace conditioning

Conditioned lick suppression in rats was used to explore the role of timing in trace conditioning. In Experiment 1, two groups of rats were exposed to pairings of a CS (CS1) with a US, under conditions in which the interstimulus interval (ISI) that separated CS1 offset and US onset was either 0 or 5 sec. Two additional groups were also exposed to the same CS1→US pairings with either a 0 or a 5-sec ISI, and then received “backward” second-order conditioning in which CS1 was immediately followed by a novel CS2 (i.e., CS1→CS2). A trace conditioning deficit was observed in that the CS1 conditioned with the 5-sec gap supported less excitatory responding than the CS1 conditioned with the 0-sec gap. However, CS2 elicited more conditioned responding in the group trained with the 5-sec CS1-US gap than in the group trained with the 0-sec CS1-US gap. Thus, the CS1-US interval had inverse effects on first- and second-order conditioned responding. Experiment 2 was conducted as a sensory preconditioning analogue to Experiment 1. In Experiment 2, rats received the CS1?CS2 pairings prior to the CS1→US pairings (in which CS1 was again conditioned with either a 0 or a 5-sec ISI). Experiment 2 showed a dissociation between first- and second-order conditioned responding similar to that observed in Experiment 1. These outcomes are not compatible with the view that differences in responding to CSs conditioned with different ISIs are mediated exclusively by differences in associative value. The results are discussed in the framework of the temporal coding hypothesis, according to which temporal relationships between events are encoded in elementary associations.     

Context effects on conditioning,extinction, and reinstatement in an appetitive conditioning preparation

Three experiments with rat subjects examined the effects of contextual stimuli on performance in appetitive conditioning. A 10-sec tone conditioned stimulus (CS) was paired with a food-pellet unconditioned stimulus (US); conditioning was indexed by the observation of headjerking, a response of the rat to auditory stimuli associated with food. In Experiment 1, a context switch following initial conditioning did not affect conditioned responding to the tone; however, when the response was extinguished in the different context, a return to the original conditioning context “renewed” extinguished responding. These results were replicated in Experiments 2 and 3 after equating exposure to the two contexts (Experiment 2) and massing the conditioning and extinction trials (Experiment 3). The results of Experiment 1 also demonstrated that separate exposure to the US following extinction reinstates extinguished responding to the tone; this effect was further shown to depend at least partly on presenting the US in the context in which testing is to occur (Experiments 2 and 3). Overall, the results are consistent with previous data from aversive conditioning procedures. In either appetitive or aversive conditioning, the context may be especially important in affecting performance after extinction.     

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.     

Backward conditioning to a CS following an expected vs. a surprising UCS

Rabbits were trained in eyelid conditioning with a “backward” arrangement of unconditioned stimulus (UCS) followed by conditioned stimulus (CS). When such a CS was tested alone it was observed to produce substantial conditioned responding if the UCS had been arranged to be “surprising” during the backward pairings, but not if it had been arranged to be “expected.” The comparisons were made in a within-subjects design where the surprisingness of the UCS on the different pairing occasions was manipulated by preceding the UCS by discriminative CSs which were otherwise either never followed by the UCS (CS?) or consistently followed by the UCS (CS+). The results may have implications for the nonmonotonic course of responding seen during backward conditioning, as a UCS is at first surprising, but then expected on the basis of contextual cues.     

Conditioned suppression of a positively reinforced shuttle response

Rats were trained to run up and down an alleyway for sucrose reinforcement on a variable interval schedule. Differential aversive classical conditioning with auditory CSs was then conducted in a separate apparatus (“off the baseline”) prior to those CSs being presented while the subjects were responding for sucrose in the alleyway. Once the effects of the CSs had extinguished, shock was reintroduced following one CS but not the other (“on the baseline” differential aversive classical conditioning). Both “off the baseline” and “on the baseline” conditioning resulted in conditioned suppression to the CS followed by shock, but little effect of the CS followed by no shock was found. In the “on the baseline” phase, total suppression of baseline responding occurred at moderate US intensities, and this appeared to result from the subject avoiding the location at which he was last shocked. At lower values, both baseline response rate and relative suppression ratio were functions of US intensity. The results are discussed in relation to the effects found in similar experiments using avoidance baselines.     

Contextual associations in trace conditioning

Rats were given tone-footshock pairings with a 0-, 10-, or 30-sec trace interval between tone offset and shock onset. Half the rats within each trace interval were tested for their conditioned fear of the tone through a lick suppression procedure; the remaining rats were evaluated for their fear of the background or contextual cues through their avoidance of the compartment in which conditioning had occurred. Less conditioning was observed to the tone with increasing trace intervals. However, conditioned fear of the context increased with increases in the trace duration. The ability of the more predictive stimulus, the tone, to overshadow the contextual cues was determined by the tone’s temporal contiguity with the footshock. The need to incorporate temporal parameters within current theories of conditioning is discussed.     

The role of signaled periods of nonreinforcement in responding on a random schedule in autoshaping

Little responding develops to a conditioned stimulus (CS) that is placed in a random relation to an unconditioned stimulus (US). However, if the USs not preceded by that CS are themselves signaled by another stimulus, then the CS does come to elicit responding. This result has been attributed (e.g., by Durlach, 1983) to the signal’s blocking of conditioning to background cues that otherwise would prevent conditioning of the CS. However, Goddard and Jenkins (1987) have suggested the alternative that signaling the USs promotes responding due to the adventitious creation of periods of signaled nonreinforcement. Two experiments were conducted to assess this alternative, involving an autoshaping preparation in pigeons. In Experiment 1, little responding to a keylight CS presented in a random relation to a food US occurred, despite the explicit presentation of a discrete noise signaling periods of no food in the intertrial interval (ITI). Experiment 2 was designed to replicate the procedure of Goddard and Jenkins, in which an auditory stimulus extended throughout the ITI of a random schedule, terminating only prior to extra USs and during the CS. Contrary to their findings, little responding developed to the target CS. However, responding did develop when the sound-free period occurred only prior to the extra USs. These results offer little support for the hypothesis that signaled periods of nonreinforcement promote responding on random schedules. However, they are consistent with the view that signaling of ITI USs acts by preventing conditioning of potentially competitive background cues.     

Facilitation of appetitive conditioning with naturalistic conditioned stimuli: CS and US factors

Adding limited female cues to a conditioned stimulus (CS) facilitates conditioned male sexual responding. In two experiments, we examined the mechanisms of this facilitation effect. The color of the female cues on the CS was varied in Experiment 1. Similarity between the CS plumage color and the color of the live female (the unconditioned stimulus [US]) could only partially account for the results. The extent to which the facilitation effect represents a specialization of sexual behavior was examined in Experiment 2 by comparing conditioning with either food or copulation as the US. The CSs with female cues elicited more approach and grab responses regardless of which US was used. However, uniquely sexual conditioned responses (mounts and cloacal contacts) were enhanced only when sexual reinforcement served as the US. These findings suggest that the facilitation effect of female cues represents a general feature of appetitive behavior systems.     

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.     

A within-event analysis of taste-potentiated odor and contextual aversions

A series of experiments was conducted to examine the phenomenon of potentiation. Experiment 1 demonstrated potentiation of odor aversions by taste when morphine served as the unconditioned stimulus (US). Experiment 2 provided evidence that the observed potentiation was due to a within-event association between odor and taste stimuli, rather than reflecting an enhanced odor-morphine association. In Experiment 3, morphine supported place conditioning to contextual cues and aversive conditioning to a taste cue, but potentiation of place conditioning by a taste cue was not obtained. Apparently the absence of potentiation was due to the dual nature of the morphine US, as potentiation of a contextual aversion by taste was obtained in Experiment 4 when a strictly aversive US (lithium) was used. These data suggest that potentiation depends on (1) an initially weak association between the to-be-potentiated conditioned stimulus (CS) element and the US, and (2) the elicitation of qualitatively similar responses by the individual elements of the CS compound. Collectively, these results support an explanation of potentiation based on within-event learning.     

Transient variations in responding to Pavlovian conditioned stimuli have implications for the mechanisms of “priming”

Conditioned responding to a well-trained conditioned stimulus (CS) was assessed in an eyelid conditioning situation under three conditions of prior stimulation: when preceded by recent presentation of the same CS, when preceded by recent presentation of a different CS, and when preceded by no recent stimulation. As compared to the latter condition, prior presentation of the same CS depressed responding, while prior presentation of a different CS augmented responding. Both effects diminished as the interval separating the test stimulus from the preceding CS was lengthened over the range of 2.5 to 10 sec. The results were discussed in terms of the interaction of short-term memory representations with subsequent stimulation, and a specific theory concerning the effects of “priming.”     

Fear-motivated vicious-circle behavior maintained through secondary punishment

Two experiments attempted to establish vicious-circle behavior through fear motivation combined with secondary punishment. In Experiment 1, rats were trained with two CSs, a tone and a buzzer, paired with shock in different contexts. Secondary punishment based on delay and trace conditioning procedures facilitated running in fear-motivated rats, relative to four control groups. In Experiment 2, rats were given pairings of a tone CS with shock, and a buzzer CS with a drop into a water tank. Fear-motivated rats which received secondary punishment during either 33% or 100% of test trials exhibited self-punitive running relative to a nonpunished (0%) group and a backward-conditioning control group. Results indicate that “all secondary” vicious-circle behavior can be established through Pavlovian conditioning, thus supporting a conditioned fear interpretation.     

Context excitation and modulation of conditioned sexual behavior

Three experiments were conducted to investigate direct and modulatory influences of context in the conditioned sexual behavior of male Japanese quail. A preference test procedure was used to assess the acquisition of contextual excitation. In Experiment 1, following direct context-unconditioned stimulus (US) pairings, male quail shifted their contextual preference from an initially preferred context to one in which they received copulatory opportunity with a female quail (US). Unpaired control group subjects did not demonstrate this shift in preference. This place preference procedure was used in Experiments 2 and 3 to assess contextual excitation when context was trained in the presence of a discrete conditioned stimulus (CS). Experiment 2 provided evidence that context can modulate responding to a discrete CS. In Experiment 3, we varied the spatial contiguity between the context and the US. Some subjects received the US directly in the training context, whereas other subjects received the US in an alternate context. Contextual excitation was evident only in subjects that received the former. Thus, there is a dissociation between the modulatory and excitatory properties of context in sexual conditioning that may depend on the context-US spatial contiguity.     

Context sensitivity of conditioned suppression following preexposure to the conditioned stimulus

Conditioned suppression in rats is often unaffected when the context (or set of background stimuli) is changed following conditioning. This suggests that responding to the conditioned stimulus (CS) can be relatively insensitive to the context in which the CS is presented. In two experiments, we examined whether sensitivity to contextual stimuli is affected by preexposure to the CS. In Experiment 1, when the CS was novel at the outset of conditioning, conditioned suppression was not affected when the context was changed following conditioning. However, when the CS had been preexposed, responding was weaker when extinction occurred outside of the conditioning context. In Experiment 2, responding was again sensitive to the test context, regardless of whether preexposure occurred in the conditioning context or in an alternate context. These results suggest that the extent to which responding is sensitive to context can depend on the conditioning history of the CS.     

Concurrent extinction does not render appetitive conditioning context specific

In an experiment with rats, an appetitive conditioning method was used to investigate the generality of the hypothesis that extinction should arouse attention to contextual cues, resulting in all learning in that context becoming context specific. Rats received appetitive conditioning with a tone either while extinction of a flasher occurred (Group With Extinction) or while it did not (Group No Extinction). Half of each group was subsequently tested in extinction in the context in which training had taken place or in a different context. The results revealed a three-way interaction of extinction and context with trials, in a direction opposite to the one the hypothesis would suggest. When rats were tested in a different context, there was generally better responding in Group With Extinction than in Group No Extinction. In the same context, there was generally lower responding in Group With Extinction than in Group No Extinction. Subsequent testing showed an ABA recovery effect. Results are discussed in terms of the challenges they pose for the revised retrieval theory presented by Callejas-Aguilera and Rosas (2011).     

The erasure of reinstated fear

Three experiments investigated the reinstatement of fear to a previously conditioned and extinguished CS as a result of separate presentation of the original US. That reinstatement was found to be sharply attenuated by nonreinforcement of a second fear elicitor between presentations of the US and testing of the CS. This “erasure” of reinstatement depended upon the fear-eliciting power of the intervening stimulus and, under some circumstances, was essentially complete. Moreover, erasure reduced not only the response to the CS but also the extinction it underwent as a result of subsequent nonreinforcement. It is argued that neither the conditioning of background stimuli nor stimulus generalization among explicit CSs provides an adequate account of these reinstatement and erasure results. Rather, they are interpreted in terms of the construction and destruction of a nonassociative representation of the US during conditioning, extinction, reinstatement, and erasure. In that context, some inferences can be made about the rules governing these nonassociative changes and the ways in which they interact with modifications in associations.     

Stimulus-reinforcer interactions in Pavlovian conditioning of pigeons: Implications for selective associations

In a Pavlovian procedure, groups of pigeons were presented with a compound auditory-visual stimulus that terminated with either response-independent electric shock or food. In a subsequent test, the tone CS was dominant in aversive conditioning, reliably eliciting conditioned head raising and prancing. The red light CS was dominant in appetitive conditioning, reliably eliciting pecking. This result was replicated in a second experiment, in which trials were widely spaced. Pour additional groups of pigeons received pairings of the separate element CSs with the USs. Red light, but not tone, was an effective CS in appetitive conditioning, whereas tone, but not red light, was effective in aversive conditioning. There was no discriminative responding in zero-contingency control groups. Several theoretical accounts of these data are discussed.     

CS modality,context conditioning,and conditioned freezing

Two studies used a one-trial-a-day aversive conditioning procedure with rats as subjects to investigate the effects of a noise versus a light CS on conditioned freezing. Experiment 1 demonstrated that less conditioned freezing was elicited by the light, although the two CSs led to similar levels of freezing to the contextual cues of the conditioning chamber. Experiment 2 replicated these outcomes and showed that the manipulation of CS intensity produced results similar to those of modality, with the more intense CSs eliciting less freezing. The second experiment also determined that freezing to contextual cues resulted from context conditioning. According to the Rescorla-Wagner model, CSs that condition poorly should generate little competition with context conditioning. Since neither the modality nor intensity factor reliably influenced context conditioning, as measured by context-evoked freezing, the studies provide no support for the view that the effects on CS-evoked freezing represent differences in the strength of conditioning to the various stimuli. This finding raises the possibility that all of the CSs conditioned well but varied in their abilities to elicit freezing because they differed in terms of the form of defensive behavior under their control.     

Stimulus specificity of concurrent recovery in the rabbit nictitating membrane response

Three experiments demonstrated that, following the extinction of an established conditioned stimulus (CS; e.g., tone), the pairing of an orthogonal stimulus from another modality (e.g., light) with the unconditioned stimulus (US) results in strong recovery of responding to the extinguished CS. This recovery occurred to about an equal degree regardless of whether or not initial training contained unambiguous stimulus—reinforcer relationships—that is, consistent CS—US pairings—or some degree of ambiguity, including intramodal discrimination training, partial reinforcement, or even cross-modal discrimination training (tone vs. light). Experiments 1 and 2 demonstrated that this recovery of responding was largely specific to the extinguished CS, but moderate generalization to other stimuli from the same modality did appear. The results are discussed with reference to alternative mechanisms applicable to learning-dependent generalization between otherwise distinct CSs. These models assume that such generalization is mediated by either a shared response, shared reinforcer, shared context, or shared hidden units within a layered neural network. A specific layered network is proposed to explain the present results as well as other types of savings seen previously in conditioning of the rabbit nictitating membrane response.     

Effects of context exposure during conditioning on conditioned taste aversions

Rats were used in a conditioned taste aversion procedure in order to examine the effects of context exposure duration during the conditioning sessions on conditioned responding. One flavor was paired with lithium chloride during a long session in one context, whereas another flavor was conditioned during a short session in another context. Testing occurred in the home cage. The results showed that conditioning during short sessions produced strong conditioned taste aversions. Conditioning during long sessions produced strong conditioned taste aversions when the conditioned-stimulus-unconditionedstimulus (CS-US) pairing occurred at the end of the lengthy session. Other results showed that context-US associations were formed during the short duration sessions and that these associations supported conditioned responding to the CS trained in that context. The results are discussed with respect to the different influences that contextual cues can exert on conditioned responding.