CS-preexposure Effect Research Articles

Flavor Aversion Learning Based on Running: A Review

Wheel running establishes aversion in rats to a flavored solution consumed shortly before the running. Many studies have shown that this is a case of Pavlovian conditioning, in which the flavor and running respectively act as the conditioned stimulus (CS) and the unconditioned stimulus (US). The present article introduces some procedural variables of this running-based flavor aversion learning (FAL), including subjects, CS agents, US agents, and drive operations. This article also summarize various behavioral features of Pavlovian conditioning demonstrated in running-based FAL including the law of contiguity despite long-delay learning, extinction and spontaneous recovery, CS-preexposure effect, remote and proximal US-preexposure effects, degraded contingency effect, inhibitory learning by backward conditioning, stimulus overshadowing, associative blocking, and higher-order contextual control. Also reviewed are several hypotheses proposed for the underlying psychophysiological causes of running-based FAL (activation of mesolimbic dopamine system, gastrointestinal discomfort, motion sickness, energy expenditure, general stress, and anticipatory contrast). At the end of the article, we visit the question of most general interest about running-based FAL: why pleasurable activity of voluntary running yields aversive learning in rats.

Effect of CS preexposure on the conditioned ejaculatory preference of the male rat: behavioral analyses and neural correlates.

Early experiences with sexual reward play a pivotal role in the formation of sexual behavior and partner preference. Associations of salient partner cues, or even neutral cues on a partner, with sexual reward states are a product of Pavlovian learning. However, the extent to which first experiences that associate a neutral stimulus with no immediate consequence, and how that association may affect subsequent associability after being paired with a sexual reward state after copulation to ejaculation, remains unclear. To address this question, sexually naïve males were preexposed over one or five trials to almond scented gauze pads prior to training during which half of the males were trained 10 times with scented receptive females, and the other half with unscented receptive females. A final test of partner preference was conducted in a large open field containing two sexually receptive females, one scented and the other unscented. Males developed a conditioned ejaculatory preference for the type of female they were trained with, except when they were preexposed five times to the odor and then trained with females bearing the same odor, indicating a significant CS preexposure effect. One CS preexposure was not sufficient to inhibit subsequent conditioning. Exposure to the scent before perfusion for inmunohistochemistry, revealed different patterns of brain activation in brain areas previously associated with the development of partner preference, like the medial preoptic area, ventral tegmental area, nucleus accumbens, basolateral amygdala, among others, depending on group membership. Thus, CS preexposure results in a subsequent impairment of the association that links the odor cue to sexual reward and preference. This highlights the impact of the first sexual experiences in future partner preference.

Characteristics of retrograde amnesia for CS preexposure.

Two experiments using rats evaluated the susceptibility of CS preexposure to retrograde amnesia induced by the protein synthesis inhibitor cycloheximide and tested whether amnesia for CS preexposure shares similar characteristics with amnesia for other memories. In Experiment 1, rats received cycloheximide either immediately, 60minutes, or 120minutes after preexposure. Following preexposure, rats received fear conditioning. When later tested, the subjects that received the amnestic treatment shortly after preexposure showed no CS preexposure effect (i.e., no reduction of fear). The amnesia for CS preexposure was attenuated with longer post-preexposure delays, showing a temporal gradient. In Experiment 2, following the replication of amnesia for CS preexposure, the amnestic treatment was readministered to the rats prior to testing. It was demonstrated that the amnestic-preexposure memory could be recovered (i.e., readministration of the drug alleviated the amnesia for CS preexposure). These two experiments show that memories for CS preexposure are susceptible to retrograde amnesia and share similar characteristics with memories for original acquisition and extinction. The results are explained using a retrieval hypothesis of retrograde amnesia.

Context-dependent latent inhibition in preweanling rats.

Preexposure to a conditioned stimulus (CS) usually weakens conditioning, an effect known as latent inhibition. Similar to other learning interference effects, latent inhibition has been characterized as context-dependent, which means that the magnitude of this effect can be attenuated by changing the context between the different phases of the procedure (e.g., preexposure and conditioning). Latent inhibition has been found with a variety of procedures in infant rats, but the few studies that examined the context-dependency of this phenomenon during this ontogenetic period found no context-change effect. The present study explored the context-dependency of latent inhibition during infancy using a conditioned taste aversion preparation and employing contexts enriched with distinctive odors to increase the possible efficacy of the context manipulation. Experiment 1 showed that three preexposures to the CS (saccharin) were sufficient to retard conditioning to the same CS, although this effect was also observed in a control group preexposed to an alternative taste stimulus (saline), in comparison with a non-preexposed control group. In Experiment 2a, the CS-preexposure effect was found to be specific to the preexposed CS when the number of preexposures was increased. This effect was revealed as context-dependent in Experiment 2b, since it was attenuated by changing the context between preexposure and conditioning. The present result is consistent with recent studies showing the context-dependency of extinction in preweanling rats, thus demonstrating these animals' capacity to learn about context early on in their development.

An assessment of context-specificity of the CS-preexposure effect in Pavlovian excitatory and inhibitory conditioning

Non-reinforced preexposure to a to-be-conditioned stimulus (CS) results in retarded development of conditioned excitation and inhibition. In a magazine-approach preparation in rats, we explored the role of background context on this CS-preexposure effect by changing contexts after the preexposure treatment. Experiment 1 demonstrated with a typical three-group design that changing background contexts attenuated the CS-preexposure effect in conditioned excitation. Experiment 2 employed the identical design except that conditioned inhibition was the target of study. Preexposure to stimulus X retarded subsequent differentiation of responding to reinforced A trials and non-reinforced AX trials, suggesting that CS-preexposure retarded development of inhibitory conditioning. However, changing contexts did not attenuate the preexposure effect. We discuss these results in the framework of the extended comparator hypothesis.

Interaction of retention interval with CS-preexposure and extinction treatments: Symmetry with respect to primacy

Imposition of a retention interval between cue-outcome pairings and testing can alleviate the retardation of conditioned responding induced by pretraining exposure to the cue (i.e., the CS-preexposure effect). However, recent studies have reported an enhanced effect of CS-preexposure treatment with longer retention intervals (De la Casa & Lubow, 2000, 2002; Lubow & De la Casa, 2002). In a series of conditioned barpress suppression studies with rats, we examined the effects of imposing a retention interval just prior to testing following either CS-preexposure (cue alone before cue-outcome pairings) or extinction (cue alone after cue-outcome pairings) treatments. Experiment 1 replicated in a different preparation recent reports of CS-preexposure treatment effects increasing with longer retention intervals. Experiment 2 showed that spontaneous recovery of stimulus control of behavior after extinction can be obtained with the same parameters as those used to observe the augmented effect of CS-preexposure treatment. In Experiment 3, both the augmented effect of CS-preexposure treatment and spontaneous recovery from extinction were found when we used, in place of a retention interval, an associative priming manipulation.

The CS-Preexposure Effect in Conditioned Taste-Aversion Learning in Golden Hamsters

In Experiment 1, golden hamsters were injected with either 0.9% saline or the nausea-inducing agent, lithium chloride (LiCL), immediately after consuming a flavored diet that was either novel or familiar. The LiCl-induced aversion was strong in hamsters for which the flavored diet was novel, but no significant aversion was observed in hamsters that were familiar with the flavored diet. In Experiment 2, the strength of the LiCl-induced aversion was related inversely to the amount of conditioned-stimulus (CS) preexposure and directly to the duration of the preexposure-conditioning interval. Thus, although some previous researchers have suggested that hamsters may not demonstrate the CS-preexposure effect in a conditioned taste-aversion paradigm, they clearly did so under the conditions of the present experiments, and moreover, the characteristics of the CS-preexposure effect in hamsters were generally similar to those observed in rats.

CSs and USs: what's the difference?

Differences in processing representations of conditioned and unconditioned stimuli (CSs and USs) may result from either their temporal order in training (i.e., CSs precede USs) or the greater biological significance of USs. The CS- and US-preexposure effects were used to probe this question. These effects are similar except that context extinction between preexposure and training more readily attenuates the US- than the CS-preexposure effect. In Experiments 1, 2, and 5, context extinction following preexposure to the stimulus that later served as Event 1 in Event 1-->Event 2 pairings alleviated the response deficit due to Event 1 preexposure if Event 1 was biologically significant. In Experiments 3 and 4, context extinction alleviated the response deficit due to Event 2 preexposure if Event 2 was biologically significant. Thus, biological significance and not temporal order determines how a representation will be processed.

The effect of US preexposure on conditioned taste aversion: Lack of postconditioning recovery of the aversion

Although the CS preexposure effect in CTA was once viewed exclusively as an acquisition failure, recent studies have suggested that the latent inhibition phenomenon is the result of retrieval impairment. This interpretive challenge is based on the unexpected finding that recovery of the aversion occurs over a long retention interval following conditioning (Kraemer, Lariviere, & Spear, Animal Learning and Behavior, 16, 185-190, 1988; Bakner, Strohen, Nordeen, & Riccio, Physiology & Behavior, 50, 1269-1272, 1991). This study examined whether a similar recovery occurs after US preexposure. Following preexposure to the US (LiCl), rats received a sucrose-illness pairing and were subsequently tested after either short or long training-to-test intervals. In contrast to the findings with the CS preexposure effect, US-preexposed subjects did not show a spontaneous increase in CTA following the long retention interval.

Attenuation of the CS-preexposure effect after a retention interval in preweanling rats

The effects of flavor preexposure and retention interval were assessed in 6- and 12-day-old rats. Conditioned aversions to a flavor appeared at both ages. The conditioning of the younger pups was unaffected by conditioned stimulus (CS) preexposure and was not evident after a 10-day retention interval. For the 12-day-old rats, preexposure to either the flavor CS or a different flavor attenuated aversion strength when the rats were tested soon after conditioning. Other 12-day-old rats that were tested 10 days after conditioning also expressed substantial aversions, but with a retention interval of this length, the aversions were equivalent for animals preexposed to the CS and those not preexposed before conditioning. This loss of the CS-preexposure effect over a long interval, which has also been observed in adult rats, identifies the locus of this effect as postacquisition and perhaps at the stage of memory retrieval.

The effect of CS-preexposure on conditioned taste aversion in young and adult rats

Young and adult rats were given nonreinforced exposures to the flavor-CS prior to a taste aversion conditioning session. CS-preexposure prevented the conditioning of a taste aversion in young rats but only attenuated conditioning in adults. These results suggest that an enhanced CS-preexposure effect may account for the previously reported weaker conditioned taste aversion observed in young rats, as compared to adults, when there is a protracted interstimulus interval.