Changes in the retrievability of associations to elements of the compound CS determine the expression of overshadowing

In two experiments, rats were presented with a taste conditioned stimulus (CS) alone, an odor CS alone, or an odor-taste compound followed by lithium chloride injection. When tested 1 day following conditioning, there was evidence that the odor cue overshadowed conditioning to the taste; however, there was no indication of overshadowing following a longer (21-day) retention interval, despite undiminished strength of the aversion in animals conditioned with only the single element (taste). The overshadowing observed at the 1-day retention interval was not reciprocal. Rats conditioned with the odor CS alone or with the compound CS expressed odor aversions of comparable strength—that is, no overshadowing. However, in contrast to the taste aversion, overshadowing of conditioning to the odor by taste was evident following a 21-day retention interval. Rather than reflecting a failure of the overshadowed stimulus to acquire associative strength, these data suggest that overshadowing may be expressed, or not expressed, as a result of changes in the relative retrievability of learned associations over time.     

US inflation with trace and simultaneous fear conditioning

Two experiments with rat subjects in a conditioned punishment paradigm are reported. These experiments attempted to determine if the events entering into association with the CS following conditioning with informative (forward) and noninformative (simultaneous) CSs were comparable. In Experiment 1, exposure to intense shock alone following trace (ISI = 10 sec) conditioning with moderate shock enhanced the suppressive effects of a 2-sec CS. A similar manipulation following explicitly unpaired CS-US presentations (ISI = 2 min) had no effect. These data were taken as evidence that the CS and US were associated during trace conditioning. Experiment 2 showed that exposure to intense shock following simultaneous conditioning also enhanced suppression to the CS. These results suggested that simultaneous and forward conditioning procedures yield similar forms of associative learning.     

Effects of expected vs. unexpected proximal US preexposure on taste-aversion learning

Animals were first conditioned to expect lithium treatment following exposure to one taste solution (the CS+) and to expect no drug treatment following exposure to another flavor (the CS?). All subjects then received a saccharin taste-aversion conditioning trial. In Experiment 1, this conditioning trial was preceded 0, 1, 2, 4, or 6 h earlier by exposure to the CS+ flavor for independent groups. The CS+ exposure attenuated saccharin aversion learning if it occurred immediately before the saccharin conditioning trial but not if it occurred 1 h or more before conditioning. In Experiment 2, the saccharin conditioning trial was preceded 3 or 4.5 h earlier by a lithium injection. This proximal US preexposure injection was either unannounced (Li) or preceded by exposure to the CS+ (CS+Li) or the CS? (CS?Li) stimuli. The US preexposure attenuated saccharin aversion learning in all cases. However, the interference effect was less when the preexposure injection was expected (CS+Li) than when it was unexpected (CS?Li). This outcome could not be explained in terms of direct effects of the CS+ and CS? stimuli on the saccharin conditioning trial, and shows that the proximal US preexposure effect is a function of not only the drug dosage and preexposure interval, but also the anticipation of the drug pretreatment.     

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.     

Role of exteroceptive background context in taste-aversion conditioning and extinction

In two experiments, saccharin (CS) and lithium chloride (US) were paired in a context consisting of specific visual, auditory, tactual, and olfactory cues. The saccharin aversion was then extinguished in a context free from conditioning-context cues. Later, saccharin preference tests were given in the presence and absence of these cues. The results indicated that the background cues of the conditioning trial controlled the amount of saccharin drunk on extinction trials, and, furthermore, that extinction of the taste aversion was context specific; i.e., groups given extinction trials in a different (from conditioning) context retained their saccharin aversion in the conditioning context only. The results indicate an important role played by the exteroceptive context in taste-aversion conditioning.     

Expression of a taste aversion conditioned with an odor-taste compound: Overshadowing is relatively weak in weanlings and decreases over a retention interval in adults

Adult rats were injected with lithium chloride (LiCl) after consumption of a novel flavor (chocolate milk) that either was or was not presented together with a novel ambient odor (banana) as a compound conditioned stimulus (CS). In Experiment 1, the adults’ consumption of the flavor 24 h after conditioning was compared with that of weanling rats given the same conditioning treatment on Postnatal Day 21. The results confirmed previous indications that the reduction in aversion observed for adults conditioned with the compound CS (overshadowing) was weak or nonexistent in weanlings. After a longer retention interval (21 days), there was no evidence of overshadowing in adults despite maintained retention of the basic conditioned aversion. In Experiment 2 this decrease in overshadowing after a long retention interval was replicated with adult animals and extended to a different method of testing. The form of the effect was the same as in Experiment 1: The decrease in overshadowing occurred over the retention interval without loss in retention of the basic taste aversion; the decrease in overshadowing was a consequence of anincrease in the flavor aversion displayed by animals conditioned with the compound CS. The impaired flavor aversion (i.e., the overshadowing) observed shortly after conditioning apparently was due to factors associated with memory retrieval, rather than to reduced attentional or associative strength.     

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.     

CS–US interval determines the transition from overshadowing to potentiation with flavor compounds

The present series of five flavor aversion experiments with rat subjects examined compound conditioning at varying CS-US intervals. Using a taste-taste design, Experiments 1A and 1B demonstrated overshadowing at a 0-min CS-US interval and potentiation at a 120-min CS-US interval, and these effects occurred with both tastes of the compound. Experiment 2 showed that the aversion to a single element is reduced when the CS-US interval is increased to 120 min, but the aversion for a compound taste is not. Experiments 3A and 3B explored odor + taste compound conditioning; the results demonstrated odor potentiation across the trace interval and a transition from taste overshadowing to taste potentiation. Collectively, the data show that the change from overshadowing to potentiation was not due to changes in the aversions produced by compound conditioning but, instead, was due to a more rapid loss of conditionability across a trace interval prior to the US in single-element conditioning. These experiments suggest that following compound conditioning, the aversion to each element represents generalization decrement from the configured compound, but the designation of overshadowing or potentiation actually depends on the status of conditioning in the single-element control.     

Effects of CS amount on conditioned taste aversion at different CS-US intervals

Previous studies on the effect of CS amount/duration on the conditioning of taste aversion have reported that animals having greater contact with the CS acquire greater aversion. These findings appear to contradict studies of CS preexposure, which show that greater contact with the CS results in less aversion. In the present research, the effect of CS amount was shown to depend on the CS-US interval. Thus, a 10-ml CS (0.15% saccharin) at 3- and 9-h CS-US intervals produced less aversion than a 1-ml CS, but there was no significant effect of CS amount at a 30-min interval. These results suggest a two-process interpretation of the delay gradient in conditioned taste aversion: one process (learned safety) is dominant at relatively long CS-US intervals, and a different process becomes dominant at short intervals.     

Cross-modality transfer effects in conditioning-enhanced neophobia in chicks (Gallus domesticus): Evidence for the separability of novelty from specific stimulus characteristics

In this study, we investigated the separability of novelty from specific stimulus characteristics (e.g., color or taste quality) in the transfer of aversion effects. Ninety-six chicks (Gallus domesticus) received a novel visual (red water) or taste (3.0% vinegar) CS paired with an injection of lithium chloride or saline. The chicks were then tested for aversion to the CS or for conditioning-enhanced neophobia in response to a different novel visual cue (green water) or taste cue (1.0% saline). Aversions to the CSs were reliable and similar to each other. Reliable evidence of conditioning-enhanced neophobia occurred with respect to each test stimulus, irrespective of the type of CS, but conditioning with the vinegar CS produced reliably greater enhancement of neophobia than did conditioning with red water. For each CS, postconditioning neophobia was more persistent in testing with saline than with green water. The results for postconditioning neophobia suggested that novelty is a general stimulus property that is separable from specific stimulus characteristics.     

The influence of the information value provided by prior-cuing treatment on the reactivation of memory in preweanling rats

In two experiments, the influence of exposure to a CS? on the acquisition and retention of a conditioned odor aversion was examined. Preweanling rats were given exposure to the CS? either prior to (CS?/CS+) or following (CS + /CS?) the pairing of a second odor (the CS+) with footshock. The results of Experiment 1 indicated that subjects in both of the treatment conditions acquired aversions of comparable strength to the odor paired with footshock and that retention of the odor aversion was not affected by order of stimulus presentation during conditioning. Experiment 2 indicated, however, that the effectiveness of pretest exposure to various elements of the conditioning episode in reactivation of the memory for conditioningwas dependent on the order of stimulus presentation during conditioning. This differential effectiveness of the various reactivation treatments is discussed in terms of their relationship to the associative “status” of the stimuli present during conditioning and in terms of the information provided to the animal by the reactivation treatment.     

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.     

The effect of morphine preexposure on the acquisition of morphine-induced taste aversions: A nonassociative effect

In Experiment 1, the amount of time rats spent engaged in a range of behaviors was recorded immediately prior to and following the intraperitoneal administration of morphine sulfate (6 mg/kg) or distilled water. No behavioral differences were observed between these groups. In Experiment 2, preexposure to this low dose of morphine attenuated the subsequent acquisition of a morphine-induced taste aversion independent of the similarity of the preexposure and conditioning environments. These results with a dose of morphine that does not produce any behavioral effects, which in turn could potentially mask associative conditioning during preexposure, confirm that the attenuating effects of morphine preexposure on taste aversion learning are nonassociative.     

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.     

A history of morphine-induced taste aversion learning fails to affect morphine-induced place preference conditioning in rats

Drugs of abuse have both rewarding and aversive effects, as indexed by the fact that they support place preferences and taste aversions, respectively. In the present study, we explored whether having a history with the aversive effects of morphine (via taste aversion conditioning) impacted the subsequent rewarding effects of morphine, as measured in the place preference design. In Experiment 1, rats were exposed to a taste aversion procedure in which saccharin was followed by morphine. Place preference conditioning was then initiated in which animals were injected with morphine and placed on one side of a two-chambered apparatus. Animals with a taste aversion history acquired place preferences to the same degree as controls without such a history, suggesting that morphine’s affective properties condition multiple effects, dependent on the specific stimuli present during conditioning. To determine whether these results were a reflection of processes operating in traditional associative conditioning, in a modified blocking procedure, place preference conditioning was attempted in the presence of a taste previously associated with morphine (Exp. 2). Under these conditions, animals still acquired morphine-induced place preferences comparable to those of animals without a morphine or conditioning history. These results are consistent with the position that drugs of abuse have multiple stimulus effects (positive and negative) that are differentially associated with specific stimuli (environmental and taste) that drive different behavioral responses (approach and avoidance).     

Second-order conditioning detects unexpressed morphine-induced salt aversion

Morphine failed to condition a salt taste aversion at a dose (15 mg/kg) sufficient to produce a robust aversion to a saccharin taste. Indeed, three different concentrations of salt (1%, 1.5%, and 2%) paired with the same morphine dose yielded no direct evidence for conditioned aversion. Yet, when a novel saccharin taste was paired in compound with the previously conditioned salt conditioned stimulus, we found evidence for a conditioning to the saccharin cue alone in three separate experiments. Control groups eliminated alternative accounts such as neophobia and differential exposure to morphine. Combined, these findings indicate that morphine conditioned a salt aversion. Although this aversion was not directly expressed, a second-order conditioning procedure was able to provide a more sensitive index of conditioning.     

US preexposures retard excitatory and facilitate inhibitory conditioning of the rabbit’s nictitating membrane response

Two experiments were conducted to examine the effects of US preexposure on differential conditioning of the rabbit’s nictitating membrane response. Both experiments consisted of three phases: a 10-day US preexposure phase, a 7-day differential conditioning phase, and a 3-day retardation of learning test for inhibition. In Experiment 1, US preexposures retarded the development of excitation to CS+ but facilitated the development of inhibition to CS?. In Experiment 2, half of the preexposed subjects received the preexposures in one experimental environment and differential conditioning in a second environment. The remaining preexposed subjects received both phases in a single environment. Retarded excitatory and facilitated inhibitory conditioning were observed only in the preexposed subjects that received both phases in the same environment. Rabbits that received a context shift performed at control levels. The results are discussed in terms of current theories of US preexposure effects, and the best account was provided by a modified associative theory.     

Habituation to illness: Effects of prior experience with the US on the formation of learned taste aversions in rats

Experiment 1 investigated the effects of US habituation on the acquisition and extinction of learned taste aversions in rats. Subjects receiving five noncontingent LiCl intubations prior to conditioning failed to develop a conditioned taste aversion, while control subjects experiencing a single saccharin/LiCl pairing displayed a pronounced taste aversion which weakened during subsequent poisonings. Experiment 2 examined whether habituation, defined as a waning of responses to repeated presentation of an illness stimulus, was a possible mechanism for explaining the results of Experiment 1. Subjects showed a decrease in motor activity following an initial LiCl intubation, but less attenuation of activity with successive intubations.     

Differential inflation with short and long CS-US intervals: Evidence of a nonassociative process in long-delay taste avoidance

Three experiments were performed to investigate the learning process underlying the phenomenon of long-delay taste conditioning. An associative model views taste avoidance as due to a conditioned stimulus-unconditioned stimulus (CS-US) associative structure, despite the long interval interposed between the flavor and illness. A nonassociative account of this avoidance behavior posits that avoidance stems from the interaction of two nonassociative processes: habituation of neophobic avoidance to a novel taste, and the poison-induced dishabituation of this process. A postconditioning inflation manipulation was used to discriminate between these two views. It has been demonstrated that enhanced responding with a US inflation manipulation depends, in part, on a previously conditioned association. Therefore, if long-delay taste avoidance arises from nonassociative processes, an inflation manipulation should not affect conditional responding. Experiment 1 demonstrated a delay of reinforcement effect, enhanced avoidance of saccharin in the immediate/inflation group, and no effect of inflation in the delay group and sham controls. Experiment 2 revealed that this differential effect of inflation is not due to absolute differences in the strength of the avoidance response. In Experiment 3, we investigated a potential associative learning mechanism that could account for the differential inflation effect. Together, the present results support the various predictions of a nonassociative account of long-delay taste avoidance.     

A transfer of control test for contextual associations

A transfer of control experiment measured the associative properties of contextual stimuli from three standard classical conditioning paradigms. After baseline training on a Sidman avoidance schedule, dogs received aversive conditioning using excitatory, inhibitory, or truly random conditioning procedures in the presence of a manipulable background stimulus. As predicted by current theory (Rescorla & Wagner, 1972; Wagner & Rescorla, 1972), the contextual stimulus was excitatory after serving as the background during conditioning of a CS? and was neutral when it had been part of the background for conditioning of a CS+. The background to the truly random procedure was also neutral. This last result contrasts with Rescorla and Wagner’s theory.