Distractor effect on the latent inhibition of conditioned flavor aversion in rats

Using a conditioned flavor aversion procedure with rats as subjects, the effect of the addition of a distractor stimulus on the magnitude of the latent inhibition effect was examined. Experiment 1 showed that latent inhibition to vinegar was attenuated by the addition of sucrose during preexposure. On the other hand, sucrose added during conditioning to vinegar did not attenuate latent inhibition. It was also found that the degree of latent inhibition to the vinegar-sucrose compound solution was less when vinegar alone was preexposed (i.e., when sucrose was added only during conditioning) than when the compound solution was preexposed (i.e., when sucrose was added both during preexposure and during conditioning). Experiment 2 gave similar results but with sucrose assigned as the target flavor and vinegar as the distractor. These findings are in full agreement with the generalization decrement account of latent inhibition.     

Reinstatement of fear to an extinguished conditioned stimulus: two roles for context

The authors studied the role of context in reinstatement. Freezing was reinstated when the conditioned stimulus (CS) was extinguished in 1 context and rats moved to another context for reexposure to the shock unconditioned stimulus (US) and test. It was also reinstated (rather than renewed) when rats were shocked in the extinction context and moved to another context for test. This reinstatement was CS specific and reduced by nonreinforced exposures to the extinction context. Rats shocked in the context in which a stimulus had been preexposed froze when tested in another context. These findings suggest 2 roles for context in reinstatement: conditioning of the test context (M. E. Bouton, 1993) and mediated conditioning by the extinction context (P. C. Holland, 1990).     

Enhanced Latent Inhibition Following Compound Pre-exposure

The influence of non-reinforced exposure to compound stimuli on subsequent appetitive classical conditioning was examined in five experiments with rats as the subjects. Non-reinforced exposure to a visual stimulus retarded subsequent acquisition of conditioned responding relative to a non-pre-exposed condition (latent inhibition). If the target stimulus was pre-exposed in compound with a second (non-target) stimulus, then latent inhibition was abolished. Exposure to the non-target stimulus prior to compound exposure had varying effects on subsequent conditioning to the target: 40 exposures to the non-target stimulus resulted in latent inhibition to the target stimulus that was comparable in magnitude to that observed when the target stimulus was exposed in isolation; 120 exposures to the non-target stimulus enhanced latent inhibition to the target stimulus.     

Latent inhibition and stimulus generalization of the classically conditioned nictitating membrane response in rabbits (Oryctolagus cuniculus) following dorsal hippocampal ablation.

Rabbits received 0 to 450 exposures of a tone conditioned stimulus (CS) prior to classical defensive conditioning of the nicitating membrane response based on an infraorbital eye shock unconditioned stimulus. Tone preexposure resulted in retarded conditioning in normal rabbits. This latent inhibition effect was not present in animals with bilateral dorsal hippocampectomy produced by aspiration. Control animals with bilateral neocortical and callosal aspiration lesions demonstrated a latent inhibition effect similar to that shown by normal nonoperated animals. The failure of CS preexposure to retard conditioning in hippocampal rabbits was not due to differences in threshold of the conditioned response to the CS or to differences in response mechanisms as determined by tests of habituation and dishabituation of the unconditioned response. A subsequent experiment employed combined-cue summation tests to confirm the fact that preexposure did not endow the tone with conditioned as well as latent inhibitiory properties. Finally, tests of stimulus generalization along the auditory frequency dimension indicated flatter relative gradients for hippocampals than for nonoperated controls, with cortical controls in between. These findings were discussed in terms of Douglas' model of hippocampal function.     

Attenuation of latent inhibition after compound conditioning

Using a conditioned suppression procedure with rats, three experiments examined the effects of compound conditioning on the degree of latent inhibition. Experiment 1 suggested that latent inhibition of the preexposed target was not enhanced but rather attenuated when a second stimulus was presented in compound conditioning. Experiment 2 showed that a similar result was obtained when, in subjects given only compound conditioning, the salience of the target was reduced to the level where it was overshadowed by the second stimulus. Experiment 3 proved that addition of the second stimulus only during preexposure, or during both preexposure and conditioning, did not attenuate the latent inhibition to the target. These results are difficult to explain by any model of Pavlovian conditioning which assumes that both latent inhibition and overshadowing effects are a consequence of acquisition deà cit; other possible accounts are discussed.     

The Importance of Context in the US Preexposure Effect in CTA: Novel versus Latently Inhibited Contextual Stimuli

Three experiments were conducted to examine the role of novel contextual stimuli in producing the unconditioned stimulus (US) preexposure effect. Experiment 1 demonstrated that novel contextual stimuli produce a significantly stronger US preexposure effect than familiar or “latently inhibited” contextual stimuli. Moreover, subjects preexposed in the presence of latently inhibited contextual cues failed to show a significant US preexposure effect. Experiments 2 and 3 attempted to provide evidence that the addition of a single novel stimulus to the latently inhibited context would result in a significantly stronger US preexposure effect than when no such novel cue was present. Experiment 3 was able to demonstrate this effect. Results are consistent with the Rescorla–Wagner (1972) model of conditioning.     

Potentiation of latent inhibition

Rats were given exposure either to an odor (almond) or a compound of odor plus taste (almond plus saline), prior to training in which the odor served as the conditioned stimulus. It was found, for both appetitive and aversive procedures, that conditioning was retarded by preexposure (a latent inhibition effect), and the extent of the retardation was greater in rats preexposed to the compound (i.e., latent inhibition to the odor was potentiated by the presence of the taste). In contrast, the presence of the taste during conditioning itself overshadowed learning about the odor. We argue that the presence of the salient taste in compound with the odor enhances the rate of associative learning, producing a rapid loss in the associability of the odor. This loss of associability will generate both overshadowing and the potentiation of latent inhibition that is observed after preexposure to the compound.     

Extinction of the context and latent inhibition

The hypothesis that latent inhibition could be reduced by extinguishing the experimental context, that is, exposing the rats to the context between exposure to the conditional stimulus (CS) and conditioning, was tested. All experiments used the conditioned emotional response procedure. In Experiment 1, extinction was not effective when the animals were exposed to the clicker 40 times off the baseline of responding for food and when the clicker CS was partially reinforced with shocks during the test phase. In Experiments 2 and 3, latent inhibition could be reduced by extinction if the animals were exposed to the CS 24 or 16 times on-baseline, and if continuous reinforcement was used during the test. In Experiments 4, 5, and 6, we attempted to determine which variable was responsible for the discrepant results. In Experiment 4, extinction was effective with 20 or 40 on-baseline exposures to the CS, using continuous reinforcement during the test. In Experiment 5, extinction was not effective with exposure on- or off-baseline, using 24 exposures and partial reinforcement. Finally, in Experiment 6, extinction reduced latent inhibition using continuous, but not partial, reinforcement with 40 exposures off-baseline. From these results, we concluded that Wagner's model of habituation was not sufficient to account for latent inhibition and that a hybrid model, using both associative and cognitive representational processes, was necessary.     

Further evidence for the summation of latent inhibition and overshadowing in rats’ conditioned taste aversion

Repeated exposures to a target taste (X) attenuated subsequent development of rats’ conditioned aversion to X (latent inhibition effect). Presentation of another taste (A) after X in conditioning (serial X-A compound conditioning) also attenuated conditioned X aversion compared with conditioning without A (overshadowing). Furthermore, the latent inhibition and overshadowing effects summed to show the least conditioned aversion in the rats given both the target preexposures and the serial X-A compound conditioning treatment. These results question the validity of the comparator hypothesis as an explanation for Pavlovian conditioning of rats’ conditioned taste aversion.     

The number of trials of compound conditioning: its effect on latent inhibition in rats.

Two experiments interspersed nonreinforced tests for the strength of the preexposed and nonpreexposed elements of a compound CS throughout conditioning. The nonreinforced exposures and the conditioning trials were superimposed on an appetitive operant baseline, and conditioned suppression of the rats' bar pressing was the dependent variable. In both experiments, the preexposed element failed to suppress responding and failed to gain strength over trials. The nonpreexposed element at first suppressed responding completely but gradually lost control of responding, even while the compound continued to suppress responding asymptotically. The results are discussed in terms of 'loss of salience' and 'discrimination' hypotheses.     

Mechanisms underlying retarded emergence of conditioned responding following inhibitory training: evidence for the comparator hypothesis

The comparator hypothesis posits that conditioned responding is determined by a comparison at the time of testing between the associative strengths of the conditioned stimulus (CS) and stimuli proximal to the CS at the time of conditioning. The hypothesis treats all associations as being excitatory and treats conditioned inhibition as the behavioral consequence of a CS that is less excitatory than its comparator stimuli. Conditioned lick suppression by rats was used to differentiate four possible sources of retarded responding to an inhibitory CS. These include habituation to the unconditioned stimulus (US), latent inhibition to the CS, blocking of the CS-US association by the conditioning context, and enhanced excitatory associations to the comparator stimuli. Prior research has demonstrated the first three phenomena. Therefore, we employed parameters expected to highlight the fourth one--the comparator process. In Experiment 1, our negative contingency training was shown to produce a conditioned inhibitor that passed inhibitory summation and retardation tests. In Experiment 2 we found transfer of retardation from an inhibitory CS to a novel stimulus when the location where retardation-test training occurred was excitatory, which is indicative of contextual blocking and/or comparator effects. In Experiment 3, extinction of the conditioning context was found to attenuate retardation regardless of whether extinction occurred before or after the CS-US pairings of the retardation test. This indicates that much of the present retardation was due to the comparator process rather than to contextual blocking. Experiment 4 demonstrated that habituation to the US did not contribute to retardation in the present case. Collectively, these studies suggest that retardation following inhibitory training can be explained without recourse to any of the traditional mechanisms of conditioned inhibition.     

Latent inhibition and contextual associations

Attenuated responding to a conditioned stimulus (CS) after preexposure to that CS (latent inhibition) has traditionally been attributed to reductions in CS associability. Alternatively, CS-context associations formed during CS preexposure later interfere with the acquisition or expression of CS-outcome associations. Three lick suppression experiments with rats contrasted these accounts. Presumably, exposure to the context attenuates the CS-context association without altering CS associability. With a fixed amount of CS preexposure, latent inhibition decreased with increasing context exposure during or after (but not before) CS preexposure. When the ratio of context preexposure duration to CS preexposure was fixed, latent inhibition increased with CS preexposure. These results suggest that latent inhibition is a direct function of the strength of CS-context associations formed during preexposure.     

Latent inhibition experiments with goldfish (Carassius auratus)

Evidence of latent inhibition was sought in a series of experiments with goldfish. In Experiment 1, goldfish were given nonreinforced preexposure to a color that subsequently predicted shock in an activity conditioning situation; their performance did not differ from that of control animals preexposed to a markedly different color. In Experiment 2, a group of goldfish given nonreinforced preexposure to a tone and an unstimulated control group were trained in an appetitive situation, with the tone serving either as a conditioned excitor or as a conditioned inhibitor. Preexposure had no significant effect in the conditioned excitation training, but it reduced the level of responding both to the positive stimulus and to the negative compound in the conditioned inhibition training. In Experiments 3 and 4, classical aversive conditioning was studied in the shuttle box. In Experiment 3, excitatory conditioning to a color was found to be impaired (relative to the performance of nonpreexposed control animals) as much by nonreinforced preexposure to the training color as by nonreinforced preexposure to a markedly different color; substantial variation in amount of preexposure was without significant effect. In the conditioned inhibition training of Experiment 4, animals with nonreinforced preexposure responded less than did unstimulated control animals both to the positive stimulus and to the negative compound. The results for goldfish can be understood on the assumption that the effect of preexposure in these animals is simply to reduce general responsiveness or level of arousal.     

Reinstatement of latent inhibition following a reminder treatment in a conditioned taste aversion paradigm.

Reminder treatments have been shown to facilitate the retrieval of a variety of conditioned responses. Whether or not similar results would occur with an experimental paradigm which involves primarily memory for a stimulus, i.e., where no particular response is specified, is unclear. Accordingly, using Sprague-Dawley rats, we employed a latent inhibition paradigm with a long (10 days) retention interval between sucrose (CS) preexposure and sucrose-illness pairing (training). The results demonstrated a loss of latent inhibition following the 10-day retention interval suggesting "forgetting" of the CS preexposure. However, placing a single reminder exposure to the CS within the preexposure-to-training interval reinstated the preexposure effect. Controls indicated that in the absence of the initial preexposure the reminder per se did not produce latent inhibition. Thus, a reminder can reinstate a stimulus attribute (flavor representation) and explicit conditioned responses.     

Inhibition as a "slave" process: deactivation of conditioned inhibition through extinction of conditioned excitation

Rats were used in a conditioned-suppression paradigm to investigate why a conditioned inhibition (CS-) does not extinguish when presented alone. Experiment 1 assessed the role of blocking by excitatory contextual cues and/or an evoked representation of the conditioned excitor (CS+), which had been nonreinforced in compound with the CS-. When the CS+ and context were extinguished prior to presentations of the CS- alone, the CS- showed a retardation effect, evidently reflecting latent inhibition, because no inhibition was detected in controls for which presentations of the CS- alone had been omitted. Experiment 2 showed that the loss of conditioned inhibition (CI) was due to excitatory extinction and not to time since conditioning. Furthermore, when excitation was reconditioned to the extinguished CS+ (Experiment 1), or to a novel CS in the same context (Experiment 2), CI was restored. Two other experiments evaluated whether the maintenance of CI depended upon excitation that was generic in form or associatively tied to the training context. They showed no loss of CI when groups received CS+ extinction in that context, along with concomitant presentations in a different context of the US by itself, for a novel CS, or correlated either positively or negatively with the original CS+. Collectively, the findings argue that CI is a "slave" to excitation, for when excitation is extinguished, CI is deactivated; and yet when excitation is reconditioned to the original or a new CS+ in the same or a different context, CI is restored.     

Facilitation and Retardation of Flavour Aversion Conditioning Following prior Exposure to the CS

Two sets of experiments on flavour aversion conditioning in rats examined the effects of the following factors: the nature of the flavour serving as the conditioned stimulus (CS), which was either a compound of monsodium glutamate, sucrose and quinine, or an elemental flavour, either sucrose or dilute acid; the amount of the CS rats consumed on a single conditioning trial-4 ml or 1 ml; and whether rats had been pre-exposed to the CS before this conditioning trial. There were three main findings. First, conditioning to an elemental CS was unaffected by the amount consumed on the conditioning trial, but conditioning to 1 ml of the compound CS was poorer than to 4 ml. Second, pre-exposure produced a latent inhibition effect if the CS was an elemental flavour, or if rats were allowed to drink 4 ml of the compound solution on their conditioning trial. But, third, pre-exposure facilitated conditioning in rats given only 1 ml of the compound solution as their CS.     

Conditioned Inhibition Produced by Extinction of a Conditioned Stimulus

Four experiments used a conditioned taste aversion procedure to examine the potential for CS-alone extinction treatment to produce a conditioned stimulus that possesses inhibitory properties. In Experiment 1, saccharin was paired with LiCl, and then saccharin was presented alone for several trials to produce extensive behavioral extinction. Animals receiving this treatment were retarded in reacquiring conditioned responding to saccharin relative to control subjects receiving conditioning to the flavor for the first time. In Experiment 2, the extinguished saccharin stimulus was shown to decrease conditioned responding to a known excitor when the two stimuli were presented in compound as a summation test. Experiments 3A and 3B replicated the findings of Experiments 1 and 2 while providing evidence that the effects were not due to the differential effects of neophobia during testing. These three experiments revealed that an extinguished conditioned excitor passes retardation and summation tests for conditioned inhibition. Experiment 4 found that extinction of a known excitor was slowed when the excitor was extinguished in compound with a previously extinguished conditioned stimulus. That is, an extinguished CS provided protection from extinction to another CS, a finding also consistent with the view that extinction produces conditioned inhibition.     

Interactions between conditioned stimuli for food and water in the rat

In two experiments rats were initially trained with one conditioned stimulus (CS) signalling food and another CS signalling water. The main purpose of the experiments was to examine the effects of presenting these stimuli in compound. Experiment 1 demonstrated that the level of conditioned responding to this compound was equal to that of a group for which the compound was composed of elements that had separately signalled the same reinforcer. Moreover, the level of this responding was greater than for a group that had received only compound conditioning. In the second experiment it was found that after the initial training, repeatedly pairing the compound with a single reinforcer resulted in an equivalent loss of conditioned responding to both elements. These findings suggest that the interaction between conditioned stimuli for food and water is facilitatory rather than inhibitory.     

Reminder-induced recovery from blocking as a function of the number of compound trials

Conditioned lick suppression in rats was employed to examine changes in the associative status of a blocked stimulus as a function of the number of compound stimulus conditioning trials. In each of two experiments, prior tone-footshock pairings produced similar blocking of conditioned responding to the light element of a tone-light compound when either two or six compound trials were used. In experiment 1, two exposures to the light alone in a dissimilar context during the retention interval, i.e., a reminder treatment, resulted in a restoration of responding to the light which was complete and similar for animals receiving two or six compound trials. This indicated that latent acquisition with respect to the blocked stimulus was largely complete after two compound trials. Because of a potential ceiling effect for reminder-induced recovery from blocking, Experiment 2 employed an attenuated reminder (one light exposure). This reminder reversed the blocking when six, but not when two, compound trials were used. These results suggest that, after latent acquisition to the blocked stimulus is complete, the blocked stimulus continues to be processed during additional compound stimulus conditioning trials, with a consequent latent facilitation of retrieval of associations to the blocked stimulus.     

Interaction between preexposure and overshadowing: Further analysis of the extended comparator hypothesis

Three experiments with rats used conditioned suppression of barpress to test predictions of the extended comparator hypothesis, which assumes that the effectiveness of (first-order) comparator stimuli in modulating responding to a target conditioned stimulus (CS) is itself modulated by other (second-order) comparator stimuli. Experiment 1 demonstrated that both pretraining exposure to the target CS alone (i.e., CS-preexposure effect, also known as latent inhibition) and pretraining exposure to a compound of the target CS and nontarget CS (i.e., compound-CS-preexposure effect) counteract overshadowing, and that posttraining deflation (i.e., extinction) of the overshadowing stimulus attenuates responding to the target CS when overshadowing is preceded by a CS-preexposure treatment (i.e., yields a CS-preexposure effect), but not when overshadowing is preceded by a compound-CS-preexposure treatment. Experiment 2 examined the consequences of posttraining associative inflation of the overshadowing stimulus or the preexposure companion stimulus following conjoint compound-CS-preexposure and overshadowing treatment. Experiment 3 examined the consequences of posttraining inflation of the overshadowing stimulus or the context following conjoint CS-alone preexposure and overshadowing treatment. The results support the expression-focused comparator view in contrast to recent acquisition-focused models of retrospective revaluation.