Restoration of the orienting response to a light by a change in its predictive accuracy

In three experiments the orienting response (OR) to a light was reduced by presenting rats with a serial conditioning schedule in which the light was always followed by a tone, and the tone was intermittently paired with food. The original strength of the OR was then restored by changing the nature of the serial conditioning: the light was followed either by the tone and food, or by nothing. Experiments 1 and 2 demonstrated that this change resulted in an increase in the OR relative to control groups, which continued to receive the original training. Experiment 3 revealed a similar effect using a within-subjects design. A further finding from the first two experiments was that the recovery of the OR was accompanied by an enhancement in the conditionability of the light. Overall, the results suggest that the accuracy with which the light signals the events that immediately follow it is an important determinant of the strength of the OR it elicits.     

Constraints on Pavlovian conditioning of the pigeon: Relative conditioned reinforcing effects of red-light and tone CSs paired with food

Recent Pavlovian conditioning experiments presented all possible CS-US combinations of red-light and tone CSs and food and shock USs to separate groups of pigeons. Pigeons receiving shock USs demonstrated conditioned head raising followed by prancing to both CSs, but CRs were acquired more rapidly to tone than to red light. Although pigeons receiving food USs rapidly acquired a conditioned response of pecking to the red-light CS, there was no evidence of conditioned responding in groups receiving tone-food pairings. This outcome left open the possibility that Pavlovian pairings of tone and food may have resulted in association formation that was not revealed in performance. The present series of experiments attempted to reveal that association, using an indirect method of assessment, conditioned reinforcement. Experiment 1 demonstrated that both red light and tone paired with food became positive conditioned reinforcers, suggesting that an association between tone and food was formed in the same number of Pavlovian conditioning trials that previously failed to yield any direct evidence of conditioning. Experiment 2, which presented fewer conditioning trials, revealed that the tone-food association was formed less rapidly than the red light-food association. Experiment 3 demonstrated that the observed outcomes were not attributable to unconditioned, rather than conditioned, reinforcing effects of the Css.     

A developmental study of attention to auditory and visual signals

Three experiments tested for developmental changes in attention to simple auditory and visual signals. Subject pressed a single button in response to the onset (Experiment 1) or offset (Experiment 2) of either a tone or a light. During one block of trials subjects knew which stimulus would come on or go off on each trial (precue condition) whereas during the other block of trials no precue was provided. In both experiments subjects as young as 4 years old responded more rapidly with precues, indicating that they were able to allocate their attention to the indicated modality. Experiment 3 utilized a choice reaction paradigm (in which subjects pressed different buttons in response to the onset of the light and the tone) in order to examine their attention allocation when no precues were provided. It was found that the adults and 7-year-olds tended to allocate their attention to vision rather than audition when no precue was provided. The results with the 4-year-olds were not entirely consistent, but suggested a similar biasing of attention to vision on their part as well.     

Asymmetrical retention functions for sequences of light flashes or tone bursts in the rat

In Experiment 1, rats were trained in a symbolic delayed matching-to-sample task to discriminate sample stimuli that consisted of sequences of magazine light flashes. The intertrial interval was illuminated by the houselight for Group Light, and it was dark for Group Dark. Retention functions exhibited a choose-many response bias when the delay interval was illuminated by the houselight in both groups, and no consistent response bias when the delay interval was dark. In Experiment 2, rats were trained to discriminate sample stimuli that consisted of sequences of tone bursts. During delay testing, a different tone (i.e., different frequency and location than the sample tone) was present or absent during the delay interval. The retention functions exhibited a significant choose-many bias when tone was present during the delay and a choose-few bias when tone was absent. Asymmetrical retention functions for tone burst and light flash sequences are due to the similarity between the stimulus conditions of the delay interval and the modality of the sequential event being discriminated. These results are consistent with an instructional ambiguity explanation of response biases in memory for number.     

Role of the hippocampus in blocking and conditioned inhibition of the rabbit's nictitating membrane response.

Bilateral aspiration of the dorsal hippocampus produced a disrupttion of blocking of the rabbit's nictitating membrane response in Kamin's two-stage paradigm (Experiment 1) but had no effect on the formation of a Pavlovian conditioned inhibitor (Experiment 2). The results of Experiment 1 indicated that normal animals and those with cortical lesions given conditioning to a light-plus-tone conditioned stimulus (CS) gave conditioned responses (CRs) to both the light and the tone during nonreinforced presentations of each (test phase). If, however, compound conditioning was preceded by tone acquisition, only the tone elicited a CR during testing; that is, blocking was observed. In rabbits with hippocampal lesions, however, CRs were given to both the light and the tone during testing whether or not compound conditioning was preceded by tone acquisition. The data from Experiment 2 showed that rabbits with hippocampal lesions could discriminate as well as normal rabbits and those with cortical lesions between a light (CS+) and light plus tone (CS-). In addition, when the inhibitory tone was subsequently paired with the unconditioned stimulus in retardation testing, animals in all three lesion conditions acquired the CR at the same rate. Thus, it appears that hippocampal lesions do not disrupt conditioned inhibition. The results of these experiments were taken as support for the view that the hippocampus is responsible for "tuning out" stimuli that have no adaptive value to the organism.     

The influence of partial reinforcement on serial autoshaping with pigeons

The effect of partial reinforcement on the rate of responding during the first element of a serial compound was investigated using autoshaping in pigeons. Experiment I employed the illumination of a response key by two different colours as the elements of the compound. Responding during the first element was faster when this stimulus was intermittently paired with the second element and the unconditioned stimulus than when a continuous reinforcement schedule was employed. Experiment II demonstrated that this effect of partial reinforcement is unaffected by maniuplating the associative strength of the second element at the outset of compound conditioning. A similar effect of partial reinforcement was also found in Experiment III which used a tone as the first element of the serial compound.     

One-trial overshadowing

Two experiments on conditioned suppression in rats examined the extent to which conditioning to a tone was overshadowed by the presence of a light. When animals received four conditioning trials, a light of moderate intensity significantly overshadowed the tone. On the first trial of conditioning, however, the light had no effect on conditioning to the tone, unless (Experiment I) its intensity was increased, or (Experiment II) it occurred in closer temporal contiguity to reinforcement than the tone. The results establish that overshadowing can occur on the first trial of conditioning, contrary to the predictions of several theories, but suggest that this may be for different reasons from those operating to produce overshadowing over a series of conditioning trials.     

Trait anxiety and perceptual load as determinants of emotion processing in a fear conditioning paradigm

The impact of trait anxiety and perceptual load on selective attention was examined in a fear conditioning paradigm. A fear-conditioned angry face (CS+), an unconditioned angry face (CS-), or an unconditioned face with a neutral or happy expression were used in distractor interference and attentional probe tasks. In Experiments 1 and 2, participants classified centrally presented letters under two conditions of perceptual load. When perceptual load was high, distractors had no effect on selective attention, even with aversive conditioning. However, when perceptual load was low, strong response interference effects for CS+ face distractors were found for low trait-anxious participants. Across both experiments, this enhanced distractor interference reversed to strong facilitation effects for those reporting high trait anxiety. Thus, high trait-anxious participants were faster, rather than slower, when ignoring CS+ distractors. Using an attentional probe task in Experiment 3, it was found that fear conditioning resulted in strong attentional avoidance in a high trait-anxious group, which contrasted with enhanced vigilance in a low trait-anxious group. These results demonstrate that the impact of fear conditioning on attention is modulated by individual variation in trait anxiety when perceptual load is low. Fear conditioning elicits an avoidance of threat-relevant stimuli in high trait-anxious participants.     

Recovery of an overshadowed association achieved by extinction of the overshadowing stimulus

Three experiments are reported in which conditioned lick suppression by water-deprived rats was used as an index of associative strength. In Experiment 1, overshadowing of a light by a tone was observed when the light-tone compound stimulus was paired with foot shock. After initial compound pairings, the tone-shock association was extinguished in one group of subjects. Subsequently, these animals demonstrated significantly higher levels of suppression to the light relative to a control group in which the tone had not been extinguished. Experiment 2 replicated this effect while failing to find evidence to support the possibilities that extinction presentations of the overshadowing tone act as retrieval cues for the light-shock association, or that, via second-order conditioning, the light-shock association is actually formed during extinction of the tone. Experiment 3 determined that the recovery from overshadowing observed in Experiments 1 and 2 was specific to the extinction of the overshadowing stimulus rather than the extinction of any excitatory cue. Collectively, these results suggest that the debilitated response to an overshadowed stimulus does not represent an acquisition failure, but rather the failure of an acquired association to be manifest in behavior.     

Auditory food cue conditioning: Effects of spatial contiguity and taste quality

The effects of spatial location of an auditory stimulus and quality of a potentiating taste on the aversive conditioning of an auditory food cue were investigated. In Experiment 1 rats ate salty food activating a tone from a speaker either in (spatially contiguous with) or displaced from the food and were then made ill. It was found that spatial contiguity during conditioning resulted in avoidance of food with a contiguous or a displaced tone in testing, and spatial displacement during conditioning resulted in avoidance of food only if the tone was also displaced in testing. Experiment 2 was identical, except rats ate salty, bitter, or sweet food with a displaced tone during conditioning and testing. The salty and bitter food groups demonstrated an avoidance of noisy food relative to the sweet food group. These results indicate that spatial contiguity interacts with taste quality in the conditioning of nongustatory food cues.     

Purkinje cell activity in the cerebellar anterior lobe after rabbit eyeblink conditioning

The cerebellar anterior lobe may play a critical role in the execution and proper timing of learned responses. The current study was designed to monitor Purkinje cell activity in the rabbit cerebellar anterior lobe after eyeblink conditioning, and to assess whether Purkinje cells in recording locations may project to the interpositus nucleus. Rabbits were trained in an interstimulus interval discrimination procedure in which one tone signaled a 250-msec conditioned stimulus-unconditioned stimulus (CS-US) interval and a second tone signaled a 750-msec CS-US interval. All rabbits showed conditioned responses to each CS with mean onset and peak latencies that coincided with the CS-US interval. Many anterior lobe Purkinje cells showed significant learning-related activity after eyeblink conditioning to one or both of the CSs. More Purkinje cells responded with inhibition than with excitation to CS presentation. In addition, when the firing patterns of all conditioning-related Purkinje cells were pooled, it appeared that the population showed a pattern of excitation followed by inhibition during the CS-US interval. Using cholera toxin-conjugated horseradish peroxidase, Purkinje cells in recording areas were found to project to the interpositus nucleus. These data support previous studies that have suggested a role for the anterior cerebellar cortex in eyeblink conditioning as well as models of cerebellar-mediated CR timing that postulate that Purkinje cell activity inhibits conditioned response (CR) generation during the early portion of a trial by inhibiting the deep cerebellar nuclei and permits CR generation during the later portion of a trial through disinhibition of the cerebellar nuclei.     

Fear of the warning signal during overtraining of avoidance

The present experiment assessed the Pavlovian properties of a tone used as a warning signal in discrete-trial avoidance training of rats by measuring its ability to block conditioning to a light when the tone-light compound signalled shock in a conditioned suppression procedure. The results, although based on a different measure of Pavlovian conditioning, confirmed those reported by Starr and Mineka (1977). After moderate levels of avoidance training, the tone blocked conditioning to the light; but after extended avoidance training it no longer did so, even though a yoked group exposed to the same Pavlovian relation between tone and shock showed no such decline.     

Controls for and constraints on auto-shaping

Auto-shaping the pigeon's key-peck response was examined as a respondent conditioning procedure with the use of Rescorla's truly-random control procedure. In the first experiment, pigeons received presentations of brief light on the response key and brief presentations of food where the light and the food were independently presented. All birds failed to key peck after many light and food presentations, but explicit pairing of the light and food rapidly conditioned pecking to the light. Experiment 2 showed that even when an independent light/food presentation schedule was reduced to variable-time 30 sec, additional naive birds would not key peck and only one bird pecked when the schedules were variable-time 15 sec. A third experiment examined an explicit-unpairing control procedure, where the light and food were not only presented on independent schedules but were also separated by a minimum time, and found that auto-shaping did not occur. A fourth experiment investigated a number of control procedures and found them ineffective. A fifth experiment investigated the effects of a physical separation of the locus of the response key and the food dispenser, and a sixth experiment investigated using a tone in place of the light. It was concluded that pecking is generated by auto-shaping procedures only when an intermittently presented keylight is regularly paired with food.     

Blocking in Rabbit Eyeblink Conditioning Is Not Due to Learned Inattention: Indirect Support for an Error Correction Mechanism of Blocking

Blocking is a classical conditioning task in which prior training to one cue such as a tone reduces learning about a second cue such as a light, when subsequently trained as a tone-light compound. Blocking has been theorized to come about through a US-modulated error correction mechanism by Rescorla & Wagner (1972) as well as through a mechanism of learned inattention as theorized by Mackintosh (1973). In the case of eyeblink conditioning, an error correction mechanism has been hypothesized to take place in the cerebellum while some form of inattention has been hypothesized to take place in the hippocampal region. The hypothesis we are testing is whether the mechanism of learned inattention is involved in blocking in rabbit eyeblink conditioning. If blocking in eyeblink conditioning is produced by a mechanism of learned inattention, then training to a previously blocked cue should be slower than training to that cue in a na?ve animal. Rabbits that had received tone training followed by tone-light training exhibited blocking. Rabbits that had been previously blocked to the light acquired conditioned responses to the light at the same rate as naive rabbits. This finding failed to support the hypothesis that blocking in rabbit eyeblink conditioning is due to learned inattention, but does support the Rescorla-Wagner mechanism of error correction. The present finding along with previous work on error correction mechanism in the cerebellar-brainstem circuit (Kim et al., 1998) lend support to the theory that blocking, at least in rabbit eyeblink conditioning, seems to be due to an error correction mechanism rather than a learned inattention mechanism.     

Blocking in rabbit eyeblink conditioning is not due to learned inattention: Indirect support for an error correction mechanism of blocking

Blocking is a classical conditioning task in which prior training to one cue such as a tone reduces learning about a second cue such as a light, when subsequently trained as a tone-light compound. Blocking has been theorized to come about through a US-modulated error correction mechanism, by Rescorla &; Wagner (1972) as well as through a mechanism of learned inattention as theorized by Mackintosh (1973). In the case of eyeblink conditioning, an error correction mechanism has been hypothesized to take place in the cerebellum while some form of inattention has been hypothesized to take place in the hippocampal region. The hypothesis we., are testing is whether the mechanism of learned inattention is involved in blocking in rabbit eyeblink conditioning. If blocking in eyeblink conditioning is produced by a mechanism of learned inattention, then training to a previously blocked cue should be slower than training to that cue in a naïve animal. Rabbits that had received tone training followed by tone-light training exhibited blocking. Rabbits that had been previously blocked to the light acquired conditioned responses to the light at the same rate as naïve rabbits. This finding failed to support the hypothesis that blocking in rabbit eyeblink conditioning is due to learned inattention, but does support the Rescorla-Wagner mechanism of error correction. The present finding along with previous work on error correction mechanism in the cerebellar-brainstem circuit (Kim et al., 1998) lend support to the theory that blocking, at least in rabbit eyeblink conditioning, seems to be due to an error correction mechanism rather than a learned inattention mechanism.     

Compounding of discriminative stimuli from the same and different sensory modalities

Rats' responding was maintained by fixed-interval schedules of reinforcement in the presence of a tone or two separate lights. The lights were either of low, moderate, or high intensity. Compounds of these single discriminative stimuli each maintained a greater frequency of response than did the single stimuli, and the compound composed of stimuli from different sensory modalities (light+tone) maintained a greater level of responding than did the compound composed of stimuli from the same sensory modality (light+light). Combining lights of different intensity had no differential effect on responding. However, in the second experiment, a compound composed of a light and a tone, each of greater intensity than the light and tone of another compound, initially maintained a higher frequency of response, demonstrating intensity effects during stimulus compounding when the increase in intensity occurs through the component stimuli. This intensity effect, however, was only transitory.     

SELECTIVE ASSOCIATIONS PRODUCED SOLELY WITH APPETITIVE CONTINGENCIES: THE STIMULUS-REINFORCER INTERACTION REVISITED

In studies reporting stimulus-reinforcer interactions in traditional conditioning paradigms, when a tone-light compound was associated with food the light gained stimulus control, but when the compound was paired with shock avoidance the tone gained control. However, the physical nature of the reinforcerrelated events (food vs. shock) presented in the presence of the tone-light compound was always confounded with the conditioned hedonic value of the compound's presence relative to its absence. When the compound was paired with shock, its presence was negative relative to its absence (which was shock-free). In contrast, when the compound was paired with food, its presence was positive relative to its absence (which was food-free). The present experiment dealt with this confounding effect by conditioning a tone-light compound to be positive or negative, relative to its absence, solely with food reinforcement. One group of rats received food for responding in the presence of the tone-light compound and no food in its absence. The other group also responded in the presence of the compound, but received food only in its absence. These rats were trained on a chained schedule in which responding in the presence of the tone-light compound produced a terminal link signaled by the absence of the compound; responding ceased in the terminal link because it delayed food delivery. In a test session to assess stimulus control by the elements of the compound, tone and light were presented separately under extinction conditions. Rats that had been exposed to a positive correlation between food and the compound emitted almost double the responses in the presence of the light as in the presence of the tone. In comparison, rats that had been exposed to a negative correlation emitted only two thirds as many responses in the presence of the light as in the presence of the tone. Because this selective association was produced using only food, it appears that the contingencies under which a reinforcer is presented, rather than (or as well as) its physical properties, can generate the selective associations previously attributed to “stimulus-reinforcer interactions.” This could mean that regardless of the class of reinforcer that ultimately maintains responding (appetitive or aversive), the contingency-generated hedonic value of the compound stimulus may influence the dominant modality of stimulus control.     

Revisiting the relation between contingency awareness and attention: evaluative conditioning relies on a contingency focus

Although evaluative conditioning has occasionally been demonstrated in the absence of contingency awareness, many recent studies imply that its acquisition depends on the availability of attentional resources during conditioning. In previous experiments attention has typically been manipulated in a general way rather than looking at the particular focus of attention. The present study investigated the role of a focus on the CS-US contingency. Two separate distraction tasks were designed that either diverted attention from the stimuli or directed it to the stimuli while drawing attention away from the contingency between the stimuli. Both types of distraction were shown to eliminate evaluative conditioning. Significant evaluative conditioning was observed in a third group of participants who were required to attend the contingencies. A mediation analysis showed that the observed discrepancy in evaluative conditioning effects between groups was mediated by contingency awareness. The results imply that attention in terms of a stimulus focus is not sufficient for evaluative conditioning to occur. Rather, attention to the contingencies between stimuli appears to be crucial in evaluative conditioning, because it is supposed to foster the acquisition of contingency awareness.     

Revisiting the relation between contingency awareness and attention: Evaluative conditioning relies on a contingency focus

Although evaluative conditioning has occasionally been demonstrated in the absence of contingency awareness, many recent studies imply that its acquisition depends on the availability of attentional resources during conditioning. In previous experiments attention has typically been manipulated in a general way rather than looking at the particular focus of attention. The present study investigated the role of a focus on the CS–US contingency. Two separate distraction tasks were designed that either diverted attention from the stimuli or directed it to the stimuli while drawing attention away from the contingency between the stimuli. Both types of distraction were shown to eliminate evaluative conditioning. Significant evaluative conditioning was observed in a third group of participants who were required to attend the contingencies. A mediation analysis showed that the observed discrepancy in evaluative conditioning effects between groups was mediated by contingency awareness. The results imply that attention in terms of a stimulus focus is not sufficient for evaluative conditioning to occur. Rather, attention to the contingencies between stimuli appears to be crucial in evaluative conditioning, because it is supposed to foster the acquisition of contingency awareness.     

The effect of reinforcement differences on choice and response distribution during stimulus compounding

In Experiments I and II, rats were trained to respond on one lever during light and another during tone. The absence of tone and light controlled response cessation. In the multiple schedule of Experiment I, all reinforcements were received for responding in tone or light; in the chain schedule of Experiment II, all reinforcements were received in no tone + no light for not responding. Experiment I subjects, for which tone and light were associated with response and reinforcement increase, responded significantly more to tone-plus-light than to tone or light alone (additive summation). Experiment II subjects, for which tone and light were associated with response increase and reinforcement decrease, responded comparably to tone, light, and tone + light. Thus, additive summation was observed when stimulus-response and stimulus-reinforcer associations in tone and light were both positive, but not when they were conflicting. All subjects in both experiments responded predominantly on the light-correlated lever during tone + light, even when light intensity was reduced in testing. Furthermore, when a light was presented to a subject engaged in tone-associated responding, all subjects immediately switched the locus of responding to the light-correlated lever. No change in locus occurred when a tone was presented to a subject engaged in light-associated responding, irrespective of the stimulus-reinforcer association conditioned to tone. The light-lever preference in tone + light indicates that the heightened responding observed in Experiment I was not the summation of tone-associated behavior with light-associated behavior. Rather, it appears to be the result of a facilitation of one operant (light-associated responding) by the reinforcement-associated cue for the other.