Analysis of a trial-spacing effect with relatively long intertrial intervals

In three experiments with rat subjects, we examined the effects of trial spacing in appetitive conditioning. Previous research in this preparation suggests that self-generated priming of the conditional stimulus (CS) and/or unconditional stimulus (US) in short-term memory is a cause of the trial-spacing effect that occurs with intertrial intervals (ITIs) of less than 240 sec. Experiment 1 nonetheless showed that a trial-spacing effect still occurs when ITIs are increased beyond 240 sec, and that the effect of ITI over 60–1,920 sec on conditioned responding is best described as a linear function. In Experiment 2, some subjects were removed from the context during the ITIs, preventing extinction of the context. Removal abolished the advantage of the long ITI, suggesting the importance of exposure to the context during the long ITI. Experiment 3 still produced a trial-spacing effect in a within-subjects design that controlled for the level of context conditioning and reinforcement rate in the absence of the CS. Overall, the results are most consistent with the idea that adding time to the ITI above 240 sec facilitates conditioning by extinguishing context-CS associations—and possibly context-US associations—that otherwise interfere with CS-US learning through retrieval-generated priming (see, e.g., Wagner, 1981).     

Sign tracking in domesticated quail with one trial a day: Generality across CS and US parameters

When the conditioned stimulus (CS) is located some distance from the unconditioned stimulus (US), pairings of the CS and US can yield either conditioned approach to the CS (sign tracking) or conditioned approach to the US (goal tracking). However, goal tracking is the more common outcome, and, because of that, goal tracking has come to serve as a “standard” measure of associative learning in several laboratories. In contrast, in previous studies of sexual conditioning with domesticated quail, only sign tracking was observed. In the present study, quail continued to show sign tracking rather than goal tracking whether or not the US was highly localized (Experiment 1), whether food or a sexual US was used (Experiment 2), whether the CS was mobile or immobile (Experiment 3), and whether the CS was 91 or 233 cm from the US compartment (Experiment 4). The present findings encourage caution in the routine use of goal tracking as a measure of learning. The possible mechanisms of goal tracking are discussed in terms of occasion setting and behavior systems theory.     

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.     

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.     

One-trial simultaneous and backward fear conditioning as reflected in conditioned suppression of licking in rats

In three experiments, groups of albino rats received one strictly simultaneous pairing of a 4-sec auditory conditioned stimulus (CS) and a 4-sec 1-mA shock unconditioned stimulus (US). Other groups received a backward pairing, in which the US began before the CS, or a forward pairing, in which the CS began before the US. Control groups received only the US or received both the CS and the US but widely separated in time. Later, the CS was presented while the rats licked a drinking tube for water, and CS-elicited suppression of licking was taken as an index of the Pavlovian conditioned response (CR). It was found that groups receiving a single forward or a single simultaneous pairing suppressed more than groups that had received a backward pairing; and the backward groups, in turn, suppressed more than the control groups. It appears, then, that excitatory fear conditioning, as reflected in conditioned suppression of licking in rats, can be produced in a single trial by both backward and simultaneous conditioning procedures.     

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.”     

CS and US duration effects in one-trial simultaneous fear conditioning as assessed by conditioned suppression of licking in rats

In three experiments, rats received a single presentation of an auditory conditioned stimulus (CS) beginning simultaneously with an electric grid-shock unconditioned stimulus (US). Later, the CS was presented while the rats licked a drinking tube for water, and CS-elicited suppression of licking was taken as an index of the excitation conditioned to the CS. It was found that conditioning increased as a joint function of the duration of CS-US overlap and US duration. The evidence suggested that weak conditioning due to a brief CS-US overlap could be increased by extending the US beyond CS termination. Extending CS duration beyond US termination, however, did not strengthen conditioning; indeed, extending the CS 60 sec beyond US termination weakened conditioning significantly. It is suggested that these results shed light on a discrepancy in the recent literature on simultaneous conditioning.     

The CS− effect in simple conditioning and stimulus selection during development

Changes in affect toward a particular stimulus can take place very rapidly through Pavlovian conditioning, if presentation of the conditioned stimulus (CS+) paired with the unconditioned stimulus (US) is accompanied by presentation of a “CS?,” another value of the same dimension as the CS+ but not paired with a US. This effect has considerable generality. It has been observed in terms of both olfactory and visual CSs, in terms of appetitive as well as aversive conditioning, and for adult as well as infant rats. The CS? effect has seemed especially important for infants, which may be related to the general tendency for infants to exhibit less stimulus selection than older animals. Finally, the CS? effect has enabled the development of a simple test of short-term retention that can quite effectively assess memory for either incidental or target events. These tests so far have indicated a clear ontogenetic decrease in rate of forgetting over short intervals, corresponding to the well-known development-related decrease in forgetting over long intervals (infantile amnesia). The tests also have shown that short-term forgetting of intentional and target events is surprisingly similar, with some indication of more rapid forgetting for the incidental events. Alternative interpretations of the CS? effect and some preliminary tests of these interpretations are discussed.     

Temporal integration in second-order conditioning and sensory preconditioning

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

Divergence of conditioned eyeblink and conditioned fear in backward Pavlovian training

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

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.     

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

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

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.     

Conditioning of tentacle lowering in the snail (Helix aspersa): Acquisition, latent inhibition, overshadowing, second-order conditioning, and sensory preconditioning

In a series of related experiments, we studied associative phenomena in snails (Helix aspersa), using the conditioning procedure of tentacle lowering. Experiments 1A and 1B demonstrated a basic conditioning effect in which the pairing of an odor (apple) as the conditioned stimulus (CS) with the opportunity to feed on carrot as the unconditioned stimulus (US) made snails exhibit increased levels of tentacle lowering in the presence of the CS. Experiments 2 and 3 showed that the magnitude of the conditioning was reduced when snails were exposed to the CS prior to the conditioning trial (a latent inhibition effect). Experiment 4 examined the effects produced by pairing a compound CS (apple—pear) with food presentations and demonstrated the existence of an overshadowing effect between the two odors. Experiment 5 revealed that pairing one CS with another previously conditioned stimulus increased tentacle lowering to the new CS (a second-order conditioning effect). Finally, Experiment 6 showed that pairing two odors prior to conditioning of one of them promoted an increase in tentacle lowering in response to the other (a sensory preconditioning effect). The results are discussed in terms of an associative analysis of conditioning and its implications for the study of cognition in invertebrates.     

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

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

Contrasting AAC and ABC renewal: the role of context associations

Rats were used in a lick suppression preparation to assess the contribution of conditioned-stimulus (CS)–context and context–unconditioned-stimulus (US) associations to experimental extinction. Experiment 1 investigated whether strengthening the CS–acquisition context association enhances extinction by determining whether stronger extinction is observed when CS-alone trials (i.e., extinction treatment) are administered in the acquisition context (AAC renewal), relative to a context that is neutral with respect to the US (ABC renewal). Less recovery of responding to the CS was observed in the former than in the latter case, extending the finding that AAC renewal is weaker than ABC renewal to our lick suppression preparation. Experiment 2 assessed the contribution of the acquisition context–US association to extinction of a CS by examining the effect of postextinction exposure to the acquisition context on responding to the extinguished CS. This manipulation enhanced responding to the extinguished CS in AAC, but not ABC, renewal. Experiment 3 addressed the contribution of the CS–acquisition context association by examining the potential of a neutral stimulus, presented in compound with the target CS during extinction treatment, to overshadow the CS–acquisition context association. This manipulation enhanced responding to the extinguished CS in AAC, but not ABC, renewal. The results stress the important role of contextual association in extinction and renewal.     

Temporal discrimination using different feature-target intervals in classical conditioning of the rabbit’s nictitating membrane response

In a typical conditional discrimination, a target stimulus (X) is reinforced during one feature cue (A→X+), but not during another feature cue (B→X−). The present experiments used only a single “feature” cue (a 66-sec tone). On half of the trials, the target stimulus (a 400-msec light) was paired with the reinforcer when the feature-target interval was one duration (e.g., 5 sec). On the remaining trials, the interval was different (e.g., 45 sec), and the target stimulus was presented without the reinforcer. All the animals acquired this temporal discrimination, and subsequent testing with other feature-target intervals yielded generalization-like gradients. These results provide solid evidence that each portion of a feature cue is encoded in a distinctive fashion. Had temporal encoding not occurred, the feature cue would have been just as ambiguous a predictor of the reinforcer as was the target stimulus, and discrimination would not have been possible. The integration of real-time temporal encoding mechanisms into models of conditional discrimination is discussed.     

Effects of varying trial distribution,intra- and extramaze cues,and amount of reward on proactive interference in the radial maze

Rats are typically less accurate in their arm selections in the radial maze over successive trials in a session (Roberts & Dale, 1981). In the present study, rats’ choice accuracy declined when such trials were separated by 2-min (massed) but not by 2-h (spaced) intertriai intervals. Changing intramaze visual/tactile arm stimuli (Experiments 1 and 3) or extramaze landmark stimuli (Experiment 4) between trials weakened the massed-trials effect, but changing the number of food pellets per arm, either alone or in conjunction with changes in intramaze cues (Experiments 2 and 3), did not. The rats also tended to avoid the spatial locations of their last four choices on a previous trial during their first four choices on a current trial, and more so with massed than with spaced trials. These findings indicate that intertriai proactive interference (PI) occurred only with massed trials and was weakened by changing intra- and extramaze cues between such trials.     

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.     

Temporal coding in conditioned inhibition: Retardation tests

Two lick suppression experiments with rats were conducted in order to determine the nature of the temporal information that is encoded as a result of Pavlovian conditioned inhibition training (conditioned stimulus {CS} A→unconditioned stimulus {US}/AX→noUS). After inhibition training, the conditioned inhibitor (X) was paired with the US in order to measure inhibition, as assessed through retarded behavioral control by CS X. Three temporal relationships were manipulated: the A-US interval, the X-A interval of inhibition training, and the X-US interval of the retardation test pairings. Retardation was greatest when the X-US temporal relationship matched the time at which the US was expected (but not delivered) on the X-A inhibition training trials. Thus, the present experiments provide evidence with retardation tests that, during conditioned inhibition training, subjects encode the temporal location of the omitted US relative to the inhibitory CS. These findings complement those of previous studies using summation tests of conditioned inhibition (Barnet & Miller, 1996; Denniston, Blaisdell, á Miller, 1998; Denniston, Cole, & Miller, 1998).