Appetitive differential conditioning of the rabbit’s jaw movement response

Two experiments examined appetitive differential conditioning of the rabbit’s jaw movement response (JMR) in a two-phase procedure. The first phase entailed reinforced training with one conditioned stimulus (CS+), and the second phase involved intermixed presentations of CS+ and an unreinforced stimulus (CS?). In Experiment 1, CS+ was a 600-Hz tone, and CS-was either a 660-,1,000-, or 2,100-Hz tone. In Experiment 2, the magnitude of the water unconditioned stimulus (US) paired with CS+ was either 1, 3, or 9 ml. The experiments revealed that 1) the level of responding to CS-rose for several days and then declined over the remainder of training; 2) the physical similarity between CS+ and CS?directly affected the level of responding to CS?but had no discernable effect on the level of responding to CS+ ; and 3) US magnitude positively affected the level of responding to CS + and, to a lesser extent, CS?. The results are discussed in terms of their implications for Spence’s gradient interaction theory and Pavlov’s external inhibition hypothesis.     

Timing at the Start of Associative Learning

Some theories of associative learning imply that time plays a fundamental role in the acquisition process. Consistent with these theories, this paper presents evidence that the time from the onset of a conditioned stimulus (CS) until presentation of the unconditioned stimulus (US) is learned very rapidly at the start of training. We report two autoshaping studies and a study on aversive conditioning in goldfish in which we examine timing at the start of conditioning. We also review data from a number of other conditioning preparations, including fear-potentiated startle, appetitive conditioning in rats, and eyeblink conditioning in rabbits, that report conditioned response (CR) timing early in training. Acquisition speed and the very first expressions of conditioned responding often show sensitivity to the time of US presentation. In instances where temporal control is slowly expressed, it is likely due to performance factors, not to slow learning about time. In fact, the learning about time may be a necessary condition for associative learning.     

Reinforcement magnitude effects in first- and second-order conditioning of directed action

Three experiments investigated the effects of reinforcement magnitude on conditioned key pecking in pigeons. Experiment 1, which included between-groups and within-subject designs, yielded significant effects of unconditioned stimulus (US) magnitude on the within-conditioned stimulus (CS) distribution of key pecks and on choice behavior, but no effect on the overall rate of key pecking. Experiment 2 employed a larger US-magnitude difference in a within-subject design. This manipulation resulted in differential rates of key pecking as well as a significant choice effect and differential within-CS key-peck distributions. A second-order conditioning procedure was used in Experiment 3, in which diffuse, visual stimuli (S1's) served as Pavlovian reinforcers for two key-light S2's. The S1 previously paired with a large US was more effective in conditioning second-order key-peck behavior to an S2 than was the S1 paired with a small US. The results of these experiments demonstrate that the associative effects of US magnitude can be expressed in the strength of CS-directed motor responding. The distinctive within-CS key-peck distributions in first-order conditioning suggests an interaction between CS- and US-directed responses.     

Effects of US-Alone Presentations on Human Evaluative Conditioning

In the context of evaluative conditioning, the effects of additional presentations of the unconditioned stimulus (US) prior to conditioning (US preexposure) or after conditioning (US postexposure) were examined using between- and within-subjects control conditions. Two experiments that differed with respect to the nationality of the subjects were conducted. In both experiments, US-alone presentations reduced the magnitude of the evaluative response. The US pre- and postexposure effects were observed in subjects classified as aware as well as in subjects classified as unaware of the experimental contingencies. Another finding is that the evaluative conditioning procedure described by Martin and Levey (1978; Levey & Martin, 1975) resulted in reliable conditioning effects also in an American sample, thus extending the scope of that special evaluative conditioning paradigm. The findings are discussed in the context of recent models of classical and evaluative conditioning. Especially, the unexpected US postexposure effect gives rise to speculations concerning the learning process underlying evaluative conditioning.     

Pavlovian conditioning of distinct components of Hermissenda's responses to rotation

In two experiments with the nudibranch mollusk Hermissenda, distinct characteristics of conditioned and unconditioned responses to high-speed orbital rotation were examined. In Experiment 1, two principle unconditioned responses to rotation were identified, namely, reduced rate of locomotion and contraction of the foot. The magnitude of the foot contraction increased throughout a 20-s period of rotation, whereas locomotion was reduced immediately after the onset of the rotation and was maintained at this constant low rate throughout the stimulus presentation. These divergent response patterns suggest that the two responses emerge independently. In Experiment 2, a classical conditioning procedure was employed in which a light (CS) was paired with the rotation (US) employed in Experiment 1. In a subsequent test, it was found that the light had acquired the capability to evoke both foot contraction and decreased locomotion. Although the magnitude of these conditioned responses was reduced relative to the corresponding unconditioned response, the patterns of responding were virtually identical; that is, foot contraction developed gradually whereas locomotion decreased immediately. In contrast, animals that received unpaired presentations of the light and rotation, light alone, or no prior exposure to those stimuli exhibited foot extension in response to the light. These results illustrate a transfer of some of the response-evoking properties of the US to the CS as a result of conditioning, as well as the emergence of two independent conditioned responses. Moreover, these results suggest modulation of at least two distinct motor pathways as a function of learning.     

Motivational control of second-order conditioning

Two experiments examined the motivational specificity of the associations that support 2nd-order conditioning. In the 1st phase of each experiment rats were exposed to 2 visual conditioned stimuli (CSs) paired with either a saline or food pellet unconditioned stimulus (US) prior to exposure to 2nd-order conditioning using 2 auditory CSs, 1 paired with each visual CS. Rats' motivational state was then shifted prior to a test such that if and only if specific motivational features of the 1st-order training US played a role in the 2nd-order associative structure would responding to the 2nd-order cues shift appropriately with the state change. Even when the US was irrelevant to the training motivational state, shifts in state revealed that it was encoded within the associative structure supporting 2nd-order responding.     

Time course of cardiac conditioned responses in restrained rats as a function of the trace CS-US interval

Two experiments aimed at understanding the temporal characteristics of trace-conditioned heart-rate responses to a 0.5-s tone (conditioned stimulus [CS]) in restrained rats. A CS paired with a tail-shock (unconditioned stimulus [US]) elicited lasting bradycardiac responses. The magnitude and extinction rate of conditioned responses (CR) were independent of the CS-US interval (interstimulus interval [ISI], 3 s to 20 s). Unreinforced test trials were analyzed for CR topography. Responding was delayed in groups with longer ISIs, but CR latencies, peak and decay times were not proportional to the ISI. Response peaks tended to cluster either about 6 s after CS onset, or about 10 s with a slow decay, depending on the ISI. The authors postulated 2 components of auditory stimulus traces involved in cardiac conditioning, maximally active 6 s and 10 s respectively after CS onset. The topography of the CR could be constrained to combinations of associative strength and instantaneous activity of these 2 components.     

Overshadowing of subsequent events and recovery thereafter

Four experiments using a conditioned lick suppression preparation with rats were conducted to examine whether overshadowing of subsequent events could be obtained in Pavlovian backward conditioning (i.e. unconditioned stimulus [US] before conditioned stimulus [CS]), and to determine whether such overshadowing could be reversed without further training with the overshadowed CS, as has been reported in overshadowing of antecedent events. In Experiment 1, a backward-conditioned CS overshadowed a second backward-conditioned CS. Two posttraining manipulations, extinction of the overshadowing CS (Experiment 2) and shifting of the temporal relationship of the overshadowing CS to the US (Experiment 3), increased responding to the overshadowed CS. These results constitute the first unambiguous demonstration of stimulus competition between subsequent events using first-order conditioning, and they show that, like overshadowing with forward conditioning, such overshadowing is due, at least in part if not completely, to a failure to express information that had been acquired.     

Interoceptive fear conditioning as a learning model of panic disorder: an experimental evaluation using 20% CO(2)-enriched air in a non-clinical sample

Despite the role afforded interoceptive fear conditioning in etiologic accounts of panic disorder, there are no good experimental demonstrations of such learning in humans. The aim of the present study was to evaluate the interoceptive conditioning account using 20% carbon dioxide (CO(2))-enriched air as an interoceptive conditioned stimulus (CS) (i.e., physiologically inert 5-s exposures) and unconditioned stimulus (US) (i.e., physiologically prepotent 15-s exposures). Healthy participants (N=42) were randomly assigned to one of three conditions: a CS-only, contingent CS-US pairings, or unpaired/non-contingent CS and US presentations. Electrodermal and self-report (e.g., distress, fear) served as indices of conditioned emotional responding. Results showed greater magnitude electrodermal and evaluative fear conditioning in the paired relative to the CS-only condition. The explicitly unpaired condition showed even greater electrodermal and evaluative responding during acquisition, and marked resistance to extinction. The latter results are consistent with the possibility that the unpaired procedure constituted a partial reinforcement procedure in which CO(2) onset was paired with more extended CO(2) exposure on 50% of the trials. Overall, the findings are consistent with contemporary learning theory accounts of panic.     

A role for CS-US contingency in Pavlovian conditioning

Two experiments evaluated the role of conditioned stimulus-unconditioned stimulus (CS-US) contingency in appetitive Pavlovian conditioning in rats. In both experiments, some groups received a positively contingent CS signaling an increased likelihood of the US relative to the absence of the CS. These groups were compared with control treatments in which the likelihood of the US was the same in the presence and absence of the CS. A trial marker served as a trial context. Experiment 1 found contingency sensitivity. There was a reciprocal relationship between responding to the CS and the trial marker. Experiment 2 showed that this result was not stimulus or response specific. These results are consistent with associative explanations and the idea that rats are sensitive to CS-US contingency.     

Occasional reinforced trials during extinction can slow the rate of rapid reacquisition

Two appetitive conditioning experiments with rats examined reacquisition after conditioned responding was eliminated by either extinction or by a partial reinforcement procedure in which reinforced trials were occasionally presented among many nonreinforced trials. In Experiment 1, reacquisition to a conditional stimulus (CS) that had been conditioned and extinguished was more rapid than acquisition in a group that had received no prior conditioning. However, the addition of occasional reinforced trials to extinction slowed this rapid reacquisition effect. Experiment 2 replicated the result and showed that a procedure in which the CS and the unconditional stimulus (US) were unpaired in extinction interfered even further with reacquisition. The results suggest that rapid reacquisition is ordinarily produced when reinforced trials provide a contextual cue that can renew responding by signaling other acquisition trials (Ricker & Bouton, 1996). The effects of partial reinforcement in extinction are surprising from several theoretical perspectives and have useful clinical implications.     

Conditioned stimulus informativeness governs conditioned stimulus-unconditioned stimulus associability

In a conditioning protocol, the onset of the conditioned stimulus ([CS]) provides information about when to expect reinforcement (unconditioned stimulus [US]). There are two sources of information from the CS in a delay conditioning paradigm in which the CS-US interval is fixed. The first depends on the informativeness, the degree to which CS onset reduces the average expected time to onset of the next US. The second depends only on how precisely a subject can represent a fixed-duration interval (the temporal Weber fraction). In three experiments with mice, we tested the differential impact of these two sources of information on rate of acquisition of conditioned responding (CS-US associability). In Experiment 1, we showed that associability (the inverse of trials to acquisition) increased in proportion to informativeness. In Experiment 2, we showed that fixing the duration of the US-US interval or the CS-US interval or both had no effect on associability. In Experiment 3, we equated the increase in information produced by varying the C/T ratio with the increase produced by fixing the duration of the CS-US interval. Associability increased with increased informativeness, but, as in Experiment 2, fixing the CS-US duration had no effect on associability. These results are consistent with the view that CS-US associability depends on the increased rate of reward signaled by CS onset. The results also provide further evidence that conditioned responding is temporally controlled when it emerges.     

Testing response generation rules

Robbins (1988) reported data that he viewed as inconsistent with Miller and Schachtman's (1985a) comparator hypothesis of conditioned response generation. Here we explain why we do not find his experiments a compelling test of the comparator hypothesis. We also briefly review other studies that tested the same predictions of the comparator hypothesis that Robbins examined. We conclude that there is considerable evidence that following excitatory or inhibitory conditioning with a target conditioned stimulus (CS) and unconditioned stimulus (US), extinction of other cues that were present during CS training ordinarily increases excitatory responding and decreases inhibitory responding to the CS. However, consistent with Robbins's conclusion, there is scant evidence that after CS-US training, enhancing the associative value of other cues that were present during CS training influences excitatory or inhibitory responding to the CS. The implications of these conclusions for the comparator hypothesis as an explanation of differences in acquired behavior and as a heuristic tool are considered.     

Parallel acquisition of awareness and differential delay eyeblink conditioning

There is considerable debate about whether differential delay eyeblink conditioning can be acquired without awareness of the stimulus contingencies. Previous investigations of the relationship between differential-delay eyeblink conditioning and awareness of the stimulus contingencies have assessed awareness after the conditioning session was finished using a post-experimental questionnaire. In two experiments, the point at which contingency awareness developed during the conditioning session was estimated from a button-press measure of expectancy of the unconditioned stimulus (US). In both experiments, knowledge of the stimulus contingencies and acquisition of differential delay eyeblink conditioning developed approximately in parallel. In Experiment 1 it was shown that predicting the US facilitated eyeblink conditioning compared with predicting the eyeblink response. In Experiment 2, a masking task was used that slowed down the emergence of awareness, and it was shown that differential conditioning only occurred in participants who were able to predict the US. The current findings challenge the hypothesis that differential delay eyeblink conditioning is entirely mediated by a functionally and neurally distinct nondeclarative learning system.     

Form and characteristics of the cardiovascular conditional response in Rhesus monkeys

Three Rhesus monkeys restrained in primate chairs were exposed to several Pavlovian control procedures, followed by 13 sessions of cardiac conditioning under a delay paradigm. The conditional stimulus (CS) was a vertical line and the unconditional stimulus (US) was electric foot shock. Heart rate (HR) was analyzed in successive fivesecond intervals beginning five seconds before CS onset. The major finding was that the conditional response was consistently biphasic and consisted of an initial acceleration followed by deceleration toward the baseline, but rarely reaching it before onset of US. The subjects differed in magnitude of acceleration and subsequent deceleration as well as the location of the maximum rate in the CS-US interval. A breakdown of trials on the basis of the pre-CS HR revealed that the magnitude of effect was inversely related to the pre-CS rate.     

Non-differential return of fear in humans after a reinstatement procedure

The present experiment investigated reinstatement of fear in humans using a differential fear conditioning preparation. In this experiment, one neutral stimulus (conditioned stimulus; CS+) was paired with an aversive stimulus (unconditioned stimulus; US) during the acquisition phase, while another neutral stimulus was not (CS−). This procedure led to a difference in responding between the CS+ and the CS− (i.e., differential conditioning). After this acquisition phase, an extinction phase followed, during which both CSs were presented without the US, resulting in a decrease in differential conditioned responding. Reinstatement refers to the return of extinguished conditioned responses due to the experience of US-only trials after the extinction phase. This phenomenon was investigated by presenting half of the participants (reinstatement group) with unpredictable USs after the extinction phase. The control group did not receive these USs after the extinction procedure. The results show that return of fear was clearly apparent after the reinstating USs. This return of fear was, however, not limited to the CS+. An increase in ‘conditioned’ responding was also observed for the control stimulus. This interesting observation will be discussed against the background of a number of recent theoretical conceptualizations of reinstatement.     

Conditioning the unconditioned response: modification of the rabbit's (Oryctolagus cuniculus) unconditioned nictitating membrane response

Conditioning-specific reflex modification (CRM) occurs when classical conditioning modifies responding to an unconditioned stimulus (US) in the absence of a conditioned stimulus (CS). Three experiments monitored rabbit nictitating (Oryctolagus cuniculus) membrane unconditioned responses to 5 intensities and 4 durations of periorbital electrical stimulation before and after CS or US manipulation. CRM occurred after 12 days of CS-US pairings but not following unpaired CS/US presentations or restraint. CRM survived CS-alone and CS/US-unpaired extinction of the conditioned response (CR) but not presentations of the US alone, although CRs remained intact. Thus, CRs could be weakened without eliminating CRM and CRM could be weakened without eliminating CRs. Data indicate CRM is a reliable, associative effect that is more than a generalized CR and may not be explained by habituation, stimulus generalization, contextual conditioning, or bidirectional conditioning.     

Blocking and unblocking of conditioned analgesia

Two experiments with rat subjects assessed the blocking phenomenon using inhibition of responding to painful thermal stimulation as an index of conditioned analgesia established through conditioned stimulus (CS)-shock pairings. In Experiment 1, a group of rats receiving the standard blocking procedure ($\text{A+}\text{AB+}$) showed shorter response latencies during a hot plate test of pain sensitivity in the presence of the added CS than groups of rats receiving various blocking control procedures. Experiment 2 replicated the blocking outcome of Experiment 1 and also demonstrated that the addition of a second shock unconditioned stimulus (US) during the compound conditioning phase attenuated the blocking effect; i.e., unblocking was observed. However, the deletion of a second shock US during compound conditioning in a group that had received two shocks on each trial in the first phase of the experiment failed to result in unblocking. These data extend the associative account of conditioned analgesia to situations involving stimulus selection and the implications for the analysis of aversive learning are discussed.     

Stimulus and temporal cues in classical conditioning

In 2 experiments, separate groups of rats were given stimulus conditioning, temporal conditioning, untreated control and (in Experiment 2) learned irrelevance control procedures, followed by a compound with both stimulus and temporal cues. Stimulus conditioning consisted of a random 15-s duration conditioned stimulus (CS) followed by food; temporal conditioning consisted of food-food intervals of fixed 90 s (Experiment 1) or fixed 75 + random 15 s (Experiment 2). The stimulus group abruptly increased responding after CS onset, and the temporal group gradually increased responding over the food-food interval. When the food-food interval was fixed 90 s, the temporal cue exerted stronger control in the compound, whereas when the food-food interval was fixed 75 + random 15 s, the stimulus cue exerted stronger control. The strength of conditioning, temporal gradients of responding, and cue competition effects appear to reflect simultaneous timing of multiple intervals.     

Contextual Control of Latent Inhibition by the Reinforcer

Three experiments examined the contextual control of latent inhibition (LI) by the unconditioned stimulus (US) using a within-subjects conditioned suppression procedure with rats. The effect of reducing the context change produced by the introduction of the shock US was investigated by presenting this US during preexposure to the conditioned stimulus (CS). Although limited CS preexposure in the absence of the US had no impact on subsequent conditioning, preexposure in the presence of the shock retarded both excitatory and inhibitory conditioning. We conclude that the introduction of the US during the conditioning phase of a normal LI experiment can produce a contextual change that reduces the observed magnitude of LI.