Effect of signaling extra unconditioned stimuli on autoshaping

Acquisition to a target conditioned stimulus (CS) is prevented when extra, unsignaled unconditioned stimuli (USs) are presented with sufficient frequency to remove contingency between target CS and US. Acquisition occurs, however, when the extra USs are signaled by another CS. According to the Rescorla-Wagner theory, signaling reduces contextual conditioning, which otherwise prevents acquisition. Results of Experiment 1 led to the rejection of a rival explanation derived from scalar expectancy theory by showing that acquisition does not occur when only half of the extra USs are signaled. The results of Experiment 2 were, however, contrary to the Rescorla-Wagner theory because they showed equivalent acquisition when the stimulus used to signal the extra USs was also present concurrently with the target CS. Signaling may exert its effect by converting the intertriai interval to CS?.     

Some determinants of second-order conditioning

In a Pavlovian conditioning situation, an initially neutral stimulus may be made excitatory by nonreinforced presentations in compound with an established conditioned excitor [i.e., second-order conditioning (SOC)]. The established excitor may be either a punctate cue or the training context. In four conditioned suppression experiments using rats, we investigated whether SOC phenomena parallel other cue interaction effects. In Experiment 1, we found that the response potential of a target stimulus was directly related to the intertrial interval when SOC was mediated by a punctate cue, and inversely related to the intertrial interval when SOC was mediated by the training context. Experiment 2 demonstrated that punctate- and context-mediated SOC are oppositely affected by posttraining context extinction, and Experiments 3 and 4 demonstrated that context- and punctate-mediated SOC are differentially affected by conditioned stimulus (Experiment 3) and unconditioned stimulus (Experiment 4) preexposure treatments. These findings parallel phenomena in conditioned inhibition and cue competition situations.     

Nonassociative habituation,US preexposure,and backward inhibitory conditioning with signaled and unsignaled USs

In a conditioned suppression experiment, rats received a single, massed session of conditioning in which one backward conditioned inhibitory stimulus (CS-) followed shocks that were signaled by a visual cue, and a second backward CS-followed shocks that were unsignaled. Conditioning was preceded by a preexposure phase in which some groups of rats were preexposed to unsignaled shock, while others were not preexposed and remained in the experimental apparatus in the absence of shock. The groups were further distinguished by whether US preexposure and conditioning occurred in the same or different contexts, and by whether conditioning began immediately or after a 24-h rest period in the home cage. Although the conditioning itself was effective in establishing the visual cue as a conditioned excitor in the nonpreexposed groups, it was not effective in establishing the two backward cues as reliable inhibitors with either signaled or unsignaled USs. After 210 US preexposures, however, the same conditioning sessions did yield conditioned inhibition to both CS-s. A 24-h rest period in the home cage reduced the magnitude of, but did not completely abolish, the facilitative effect of US preexposure on inhibitory conditioning. Other tests demonstrated that US preexposure had retarded excitatory conditioning to the visual cue. This interference with excitatory conditioning was unchanged in magnitude after the 24-h rest period. The facilitative effect of US preexposure on backward inhibitory conditioning, and the interference effect on excitatory conditioning, were both eliminated by a change in context between US preexposure and conditioning. These observations encourage predominantly associative accounts of the effects, but allow for a small nonassociative habituation component.     

Overshadowing of situational cues with variable but not fixed intertrial intervals

In two experiments, we examined the conditions under which signaling an unconditioned stimulus (US) with a nominal conditioned stimulus (CS) interferes with the conditioning of situational cues in defensive freezing in the rat. Subjects received footshock USs that were (1) either signaled or unsignaled and (2) either varied or fixed in their temporal location within the conditioning session. Experiment 1, with only one trial per session, yielded no evidence that signaling affected pretrial freezing using either a fixed or variable interval between placement in the context and shock onset. In a test in which no CSs or footshocks were presented, groups that previously had received footshock at a fixed temporal location showed greatest freezing at around that same time. For groups that had received footshocks at various times, freezing declined across the test session. Experiment 2 showed overshadowing of pretrial freezing after more extensive conditioning with many trials per session, but only if the intershock intervals were variable rather than fixed.     

Encoding of the unconditioned stimulus in Pavlovian conditioning

Three experiments, using rats, demonstrated the encoding of a food unconditioned stimulus (US) in a simple Pavlovian conditioning paradigm. In all three studies, one stimulus was used to signal the delivery of pellets and a different stimulus was used to signal the delivery of sucrose. In Experiment 1, postconditioning devaluation of one of the food USs selectively reduced the frequency of conditioned magazine-directed behavior during the stimulus trained with that US. In Experiment 2, transfer of the stimuli to instrumental responses resulted in selective depression of the response trained with a different outcome. In Experiment 3, acquisition of stimulus-outcome learning was impaired by unsignaled intertrial presentations of the same outcome but not of a different outcome. These results indicate that a detailed representation of the outcome is encoded in the normal course of Pavlovian conditioning.     

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.     

On the generality of conditioned bradycardia in rabbits: Assessment of CS and US modality

To determine whether the magnitude of heart rate (HR) slowing induced by classical conditioning contingencies is comparable under a broad range of stimulus conditions, experiments were conducted in which rabbits were exposed to tones, increases in illumination, or vibratory stimuli as conditioned stimuli (CSs) and in which paraorbital electric shocks, corneal airpuffs, or intraoral pulses of water served as unconditioned stimuli (USs). The results indicated that conditioned bradycardia was elicited by all three CSs. Moreover, when a corneal airpuff served as the US, small but reliable CS-evoked HR decelerations also occurred. Finally, CS-evoked HR slowing also occurred in response to a tone CS employed in an appetitive task, in which water was the US. These findings suggest that HR slowing is a general phenomenon that occurs when rabbits are exposed to signals that systematically predict aversive or appetitive consequences according to a Pavlovian conditioning paradigm.     

Competition between ethanol-induced reward and aversion in place conditioning

Previous place conditioning studies in mice have shown that injection of ethanol immediately before a conditioned stimulus (CS+) produces conditioned preference, whereas injection of ethanol immediately after CS+ produces conditioned aversion. In the present experiments, we examined the learning that occurs when ethanol is injected in “ambiguous“ procedures that provide the opportunity for both types of conditioning. When ethanol was given midway through the CS (Experiments 1 and 2) or both before and after the CS (Experiment 3), the direction of place conditioning was the same as when mice were exposed only to whichever contingency occurred first (a primacy effect). That is, injection of ethanol in the middle of the CS conditioned aversion, whereas injection both before and after the CS conditioned preference. Because these results support the idea that ethanol elicits both aversive and rewarding effects, they are most consistent with conditioning theories that conceptualize unconditioned stimuli (USs) as events that can activate multiple representational components.     

Retention of conditioned autonomic and behavioral responses in preweanling rats: Forgetting and reinstatement

Preweanling rats, 16 days of age, responded to an olfactory conditioned stimulus (CS) paired with a shock unconditioned stimulus (US) with increases in heart rate and behavioral activation. In two experiments this finding was replicated and, in addition, it was found that the form of these conditioned responses (CRs) changed after a retention interval. When tested 24 h after CS-US pairings, the subjects displayed a decrease in heart rate accompanied by CS-elicited freezing. Giving two unsignaled shocks prior to the delayed test effectively reinstated the tachycardia and behavioral arousal CRs. The results are discussed in terms of contextual influences on the form of the CR and how changes in the magnitude of context fear may alter responding to an olfactory CS.     

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.     

Associative foundation of causal learning in rats

Are humans unique in their ability to interpret exogenous events as causes? We addressed this question by observing the behavior of rats for indications of causal learning. Within an operant motor–sensory preconditioning paradigm, associative surgical techniques revealed that rats attempted to control an outcome (i.e., a potential effect) by manipulating a potential exogenous cause (i.e., an intervention). Rats were able to generate an innocuous auditory stimulus. This stimulus was then paired with an aversive stimulus. The animals subsequently avoided potential generation of the predictive cue, but not if the aversive stimulus was subsequently devalued or the predictive cue was extinguished (Exp. 1). In Experiment 2, we demonstrated that the aversive stimulus we used was in fact aversive, that it was subject to devaluation, that the cue–aversive stimulus pairings did make the cue a conditioned stimulus, and that the cue was subject to extinction. In Experiments 3 and 4, we established that the decrease in leverpressing observed in Experiment 1 was goal-directed instrumental behavior rather than purely a product of Pavlovian conditioning. To the extent that interventions suggest causal reasoning, it appears that causal reasoning can be based on associations between contiguous exogenous events. Thus, contiguity appears capable of establishing causal relationships between exogenous events. Our results challenge the widely held view that causal learning is uniquely human, and suggest that causal learning is explicable in an associative framework.     

Conditioning-specific reflex modification of the rabbit (Oryctolagus cuniculus) nictitating membrane response is sensitive to context

Conditioning-specific reflex modification occurs when an unconditioned response is modified in theabsence of the conditioned stimulus as a result of pairings of the conditioned stimulus and an unconditioned stimulus. In two experiments, we assessed conditioning-specific reflex modification in either a novel context (Experiment 1) or a context different from, but equally familiar in relation to, the training context (Experiment 2). Conditioning-specific reflex modification did not demonstrate sensitivity to a novel context but did demonstrate sensitivity to a change in familiar context. The data cannot be explained by unconditioned stimulus preexposure, overtraining, or context insensitivity. The results suggest that conditioning-specific reflex modification models normal stress and may be used to evaluate theories of and treatments for posttraumatic stress disorder.     

Evaluating conditioning of related and unrelated stimuli using a compound test

Three experiments used a compound test procedure to evaluate whether superior conditioning results from the pairing of stimuli that are related to each other. In each case, a stimulus compound was tested after its component conditioned stimuli (CSs) had been conditioned by the same unconditioned stimuli (USs) arranged such that either related or unrelated CSs and USs were paired. Experiment 1 explored auditory and gustatory stimuli conditioned by LiCl or shock, using rats. Experiments 2 and 3 used second-order conditioning in pigeons to pair stimuli that were similar by virtue either of qualitative features or of shared physical location. In each case, the compound test provided clear evidence that pairing related stimuli produces superior associative learning.     

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.     

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.     

Chemosensory conditioning in molluscs: II. A critical review

We critically review chemosensory conditioning studies with molluscs and find that, in many studies, the influence of nonassociative processes complicates, obscures, and renders ambiguous the unique contribution of associative learning. These nonassociative processes include sensory adaptation, habituation, sensitization, and changes in feeding motivation. They arise from both the food extracts that have often been used as conditioned stimuli and the aversive stimuli that have been used as unconditioned stimuli.     

Devaluation of Pavlovian conditioning in the 10-day-old rat

This experiment established that for the infant rat, 10 days postpartum, a preference conditioned to an olfactory stimulus (conditioned stimulus) could be substantially-decreased-by subsequently lowering the value of the unconditioned stimulus (heat). This devaluation effect disappeared when a sufficiently long interval elapsed between the devaluation treatment and the test, despite maintained retention of the original conditioned preference over this same interval. This suggests that devaluation in infant rats does not permanently change the animal’s original representation of the unconditioned stimulus, but instead may replace it temporarily with a conflicting representation.     

The effects of signaling the US in backward conditioning: A shift from excitatory to inhibitory learning

The associative effects of “backward” US-CS pairings were compared when the pairing occasions were either consistently preceded by a well-trained CS+ or were unannounced. The investigation employed a conditioned emotional response procedure with rats, under conditions in which all subjects received the same schedule of shock USs, some signaled and others not, and the back-ward CS was arranged to follow either the former or the latter, in separate groups. The major finding was that although the backward CS became excitatory when it followed unsignaled USs, it became inhibitory when it followed signaled USs. This outcome, which is in line with prior findings of Wagner and Terry (1975), is in accordance with a “sometimes-opponent-process” model proposed by Wagner (1981). It is contrary to data reported by Fowler, Kleiman, and Lysle (1984) that may have resulted from a confounding of the different circumstances of backward conditioning with differences in the predictability of the US in the experimental context.     

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.     

On the nature of CS and US representations in Pavlovian learning

A significant problem in the study of Pavlovian conditioning is characterizing the nature of the representations of events that enter into learning. This issue has been explored extensively with regard to the question of what features of the unconditioned stimulus enter into learning, but considerably less work has been directed to the question of characterizing the nature of the conditioned stimulus. This article introduces a multilayered connectionist network approach to understanding how “perceptual” or “conceptual” representations of the conditioned stimulus might emerge from conditioning and participate in various learning phenomena. The model is applied to acquired equivalence/distinctiveness of cue effects, as well as a variety of conditional discrimination learning tasks (patterning, biconditional, ambiguous occasion setting, feature discriminations). In addition, studies that have examined what aspects of the unconditioned stimulus enter into learning are also reviewed. Ultimately, it is concluded that adopting a multilayered connectionist network perspective of Pavlovian learning provides us with a richer way in which to view basic learning processes, but a number of key theoretical problems remain to be solved, particularly as they relate to the integration of what we know about the nature of the representations of conditioned and unconditioned stimuli.