Priming and trial spacing in extinction: Effects on extinction performance,spontaneous recovery,and reinstatement in appetitive conditioning

Previous research in this laboratory suggests that priming of the conditional stimulus (CS) in short-term memory may play a role in the trial-spacing effects in appetitive conditioning. For example, a nonreinforced presentation of a CS 60 s before a reinforced trial with the same CS produced slower acquisition than a CS presentation that occurred 240 s before the reinforced trial. The results were consistent with the self-generated priming mechanism proposed by Wagner (e.g., Wagner 1978, 1981). The present experiments extended the earlier work by examining the effects of trial spacing in extinction rather than acquisition. After conditioning with a mixture of intertrial intervals (ITIs), rats received extinction with ITIs of 60 or 240 s, longer or shorter values, or different ways of “chunking” extinction trials in time. Although trial spacing produced effects on extinction performance that were consistent with our previous research on acquisition, there were few long-term differences in spontaneous recovery or in reinstatement. Short ITIs in extinction appear to affect extinction performance more than they affect extinction learning. Mechanisms of trial spacing in conditioning and extinction are discussed.     

The extinction of an aversion: Role of the interval between non-reinforced presentations of the averted stimulus

A toxiphobia conditioning paradigm was used to examine the relation between the intertrial interval (ITI) and the extinction of an aversion. The design employed was based upon that developed by Davis (1970) to study the relation between the ITI and the habituation of a response. After conditioning, the conditioned stimulus (CS) was presented on two occasions at times T1 and T2, and this interval (the ITI) was varied across groups. However, the interval from the CS exposure at T2 to the extinction test was common for all groups. On the test, ITIs of 6 to 24 hr were found to have promoted more extinction than ITIs of 0.5 hr with odour (Experiments 1a and b) and flavour (Experiment 1c) CSs. Further, this facilitation of extinction by the long ITIs compared to the short ones was not due to differences in the intervals between the initial CS exposure at T1 and the test (Experiment 2). The final experiment examined the relation between the ITI and extinction when different CSs were presented at T1 and T2. A short interval between the presentation of CS1 at T1 and the presentation of CS2 at T2 was found to have facilitated the extinction of the aversion to CS2 compared either to a long interval between these presentations or to the presentation of CS2 in the absence of a prior CS1 presentation. The results were discussed in terms of the model developed by Wagner (1981).     

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.     

CS Intensity effects on rabbit nictitating membrane conditioning,extinction and generalization

One purpose of the present experiments was to determine if the Grice and Hunter (1964) observation of augmented within-versus between-Ss CS intensity effects in human eyelid conditioning would be obtained in conditioning of the rabbit’s nictitating membrane response under two (Experiment 1) and four (Experiment 2) CS intensity values. In addition, a determination was made of the effects of CS intensity upon extinction and stimulus intensity generalization gradients. The studies revealed that: (a) while acquisition performance was positively related to CS intensity, the effect was independent of the between and within-S manipulation of CS intensity; (b) rate of response decrement in extinction was an inverse function of CS intensity; and (c) a positively sloped intensity generalization gradient was obtained when the training stimulus was the low-intensity one. Overall, these results are most consistent with Hull’s (1949) stimulus intensity dynamism account of CS intensity effects in conditioning.     

Adrenergic transmission facilitates extinction of conditional fear in mice

Extinction of classically conditioned fear, like its acquisition, is active learning, but little is known about its molecular mechanisms. We recently reported that temporal massing of conditional stimulus (CS) presentations improves extinction memory acquisition, and suggested that temporal spacing was less effective because individual CS exposures trigger two opposing processes: (1) fear extinction, which is favored by CS massing, and (2) fear incubation (increase), which is favored by spacing. We here report the effects of manipulating the adrenergic system during massed or spaced CS presentations in fear-conditioned mice. We administered yohimbine (5 mg/kg), an alpha(2)-receptor antagonist, or propranolol (10 mg/kg), a beta-receptor antagonist, systemically prior to CS presentation sessions and recorded both short- and long-term changes in conditional freezing. Yohimbine treatment facilitated extinction of both cue and context fear with massed protocols. When given before spaced CS presentations, propranolol led to a persistent incubation of cue fear, whereas yohimbine led to persistent extinction, compared with vehicle-treated animals, which showed no change in fear. These results suggest that norepinephrine positively modulates the formation of fear extinction memories in mice. They also provide clear evidence that spaced CS presentations trigger both fear-reducing (extinction) and fear-increasing (incubation) mechanisms.     

A partial reinforcement extinction effect despite equal rates of reinforcement during Pavlovian conditioning

In 4 experiments rats received appetitive Pavlovian conditioning followed by extinction. Food accompanied every trial with the conditioned stimulus (CS) for the continuously reinforced groups and only half of the trials for the partially reinforced groups. In contrast to previous experiments that have compared the effects of partial and continuous reinforcement, the rate at which food was delivered during the CS was the same for both groups. The strength of the conditioned response during extinction weakened more rapidly in the continuously than in the partially reinforced groups. The results demonstrate that the partial reinforcement extinction effect is a consequence of the nonreinforced trials with the CS, rather than the rate at which the unconditioned stimulus is delivered during the CS.     

Resistance to continuous delay of reinforcement or extinction following partial delay or partial reinforcement in acquisition: A direct comparison

Rats received either partial reward (PR) or partial delay (PD) in acquisition with one, two, or three delay or nonreward trials followed by an immediately rewarded trial or one delay or nonreward trial followed by an immediately rewarded trial. These four groups were then split in half and given either continuous delay or continuous nonreward (extinction) in a “response persistence” phase. In addition, two continuously reinforced groups, one experiencing continuous delay, and the other experiencing extinction were included. The results showed that response persistence was greater when PD groups were given continuous delay rather than extinction, but the opposite was true for PR groups. The “length” (1, 2, or 3 versus 1 nonreward or delay trial) also transferred to the response persistence phase with the length 1, 2, and 3 conditions being more persistent than the 1 length conditions. The results were discussed with respect to theoretical notions of response persistence.     

Fear conditioning and extinction in anxiety- and depression-prone persons

Anxiety and depression frequently co-occur and may share similar deficits in the processing of emotional stimuli. High anxiety is associated with a failure in the acquisition and extinction of fear conditioning. Despite the supposed common deficits, no research has been conducted on fear acquisition and extinction in depression. The main aim of the present study was to investigate and compare fear acquisition and extinction in anxiety- and depression-prone participants. Non-clinical anxious, depressive, anxious-depressive and control participants performed a fear discrimination task. During acquisition, the CS+ predicted an aversive event (unconditioned stimulus, US) and the CS? safety (no US). During extinction, the CS+ was no longer followed by the US, rendering it (temporarily) into a safety signal. On each CS participants rated their US expectancy; skin conductance responses (SCRs) were measured throughout. The expectancy scores indicated that high anxiety resulted in less safety learning during acquisition and extinction; no effect of depression was observed. SCRs showed that high-anxiety persons displayed less discrimination learning (CS+ minus CS?) during acquisition than low-anxiety persons. During extinction, high-depression persons demonstrated more discriminative SCR than low-depression persons. The observed discrepancies in response patterns of high-anxiety and -depression persons seem to indicate distinctive information processing of emotional stimuli.     

Revisiting the learning-without-awareness question in human pavlovian autonomic conditioning: Focus on extinction in a dichotic listening paradigm

Numerous studies have indicated that, consistent with current “cognitive” accounts of information processing, human Pavlovian autonomic discrimination acquisition cannot occur without awareness of the CS-US relationship. However, extinction studies have suggested that awareness is not necessary, findings that, in information-processing terms, have been explained by assuming that the processing by the extinction stage is parallel (automatic) rather than serial (controlled). This explanation was tested in an 80-subject study. The first, acquisition phase was a standard semantic differential conditioning arrangement with a 96-db white noise as US, and a “long” CS-US interval of 8 s, with ten trials each of CS+ (paired with US) and CS− (unpaired) trials. In extinction (USs omitted), in order to obtain non-autonomic indices of processing and thereby test the information-processing account of “unaware” autonomic conditioning during extinction, a dichotic listening task was implemented, with the CSs presented in the unattended channel (ear), while the subject had to perform a semantic differential reaction task in an attended-to channel (other ear). In early extinction, the electrodermal response occurring at an interval of 9–15 s after CS onset (i.e., following placement of the US during acquisition) and the finger-pulse-volume response occurring at an interval of 4–11 s after CS onset both showed reliable conditioning, but reaction-time and subjective-report data for the recognized critical words indicated serial rather than parallel processing of the CSs during extinction.     

An analysis of sequential variables in Pavlovian conditioning employing extended and limited acquisition training

Sequential theory’s memory model of learning has been successfully applied in response contingent instrumental conditioning experiments (Capaldi, 1966, Capaldi, 1967, Capaldi, 1994 and Capaldi and Miller, 2003). However, it has not been systematically tested in nonresponse contingent Pavlovian conditioning experiments. The present experiments attempted to determine if several sequential variables affect responding in Pavlovian situations as they do in instrumental ones. Of primary concern here were the effects on extinction of number of NR transitions (the number of times a nonreinforced trial is followed by a reinforced trial), N-length (the number of successive nonreinforced trials that precede a reinforced trial), and percentage of reinforcement (50 versus 100%) following either extended acquisition training (Experiment 1, 720 trials) or limited acquisition training (Experiment 3, 24 trials). In agreement with a sequential analysis, N-length increased resistance to extinction more than number of NR transitions following extensive training with the opposite occurring following limited training. In Experiment 1, greater resistance to extinction was associated with 50% than with 100% reinforcement, a partial reinforcement extinction effect (PREE). Experiment 2 examined an anomalous finding obtained in Experiment 1. A major theoretical difference between instrumental and Pavlovian conditioning has been held to be the greater ease of producing a PREE in instrumental than in Pavlovian conditioning (Kimble, 1961 and Mackintosh, 1974). However, the findings obtained here suggest that the probability of obtaining a PREE and other Pavlovian extinction effects, as in instrumental conditioning, increases along with the effectiveness of the sequential variables employed.     

Duration of extinction trials as a determinant of instrumental extinction in terrestrial toads (<Emphasis Type="Italic">Rhinella arenarum</Emphasis>)

Instrumental learning guides behavior toward resources. When such resources are no longer available, approach to previously reinforced locations is reduced, a process called extinction. The present experiments are concerned with factors affecting the extinction of acquired behaviors in toads. In previous experiments, total reward magnitude in acquisition and duration of extinction trials were confounded. The present experiments were designed to test the effects of these factors in factorial designs. Experiment 1 varied reward magnitude (900, 300, or 100 s of water access per trial) and amount of acquisition training (5 or 15 daily trials). With total amount of water access equated in acquisition, extinction with large rewards was faster (longer latencies in 900/5 than 300/15), but with total amount of training equated, extinction with small rewards was faster (longer latencies in 100/15 than 300/15). Experiment 2 varied reward magnitude (1200 or 120 s of water access per trial) while holding constant the number of acquisition trials (5 daily trials) and the duration of extinction trials (300 s). Extinction performance was lower with small, rather than large reward magnitude (longer latencies in 120/300 than in 1200/300). Thus, instrumental extinction depends upon the amount of time toads are exposed to the empty goal compartment during extinction trials.     

Facilitation of extinction by an increase or a decrease in trial duration

Five experiments examined the effects of altering the duration of a conditioned stimulus (CS) for extinction. For the first 3 experiments, rats received conditioning with a 10-s CS before different groups received extinction with a CS that was either the same duration or longer than that used for conditioning. For the remaining 2 experiments, conditioning was conducted with a 60-s CS before different groups received extinction with a CS of either the same duration or a shorter duration than that used for conditioning. In all experiments, extinction progressed more readily when the CS duration was different for the 2 stages than when it was constant. The results are discussed in terms of rate expectancy theory and associative learning theory.     

Revisiting the learning-without-awareness question in human Pavlovian autonomic conditioning: focus on extinction in a dichotic listening paradigm.

Numerous studies have indicated that, consistent with current "cognitive" accounts of information processing, human Pavlovian autonomic discrimination acquisition cannot occur without awareness of the CS-US relationship. However, extinction studies have suggested that awareness is not necessary, findings that, in information-processing terms, have been explained by assuming that the processing by the extinction stage is parallel (automatic) rather than serial (controlled). This explanation was tested in an 80-subject study. The first, acquisition phase was a standard semantic differential conditioning arrangement with a 96-db white noise as US, and a "long" CS-US interval of 8 s, with ten trials each of CS+ (paired with US) and CS- (unpaired) trials. In extinction (USs omitted), in order to obtain non-autonomic indices of processing and thereby test the information-processing account of "unaware" autonomic conditioning during extinction, a dichotic listening task was implemented, with the CSs presented in the unattended channel (ear), while the subject had to perform a semantic differential reaction task in an attended-to channel (other ear). In early extinction, the electrodermal response occurring at an interval of 9-15 s after CS onset (i.e., following placement of the US during acquisition) and the finger-pulse-volume response occurring at an interval of 4-11 s after CS onset both showed reliable conditioning, but reaction-time and subjective-report data for the recognized critical words indicated serial rather than parallel processing of the CSs during extinction.     

Evidence of the origin of specific spontaneous head turns during intertrial intervals

Direction and the frequency of spontaneous head movements during the ITIs following forward and backward paired trials were compared to an acquisition of a conditioned orienting (alpha) response directed to the side of the tone source. The head movements were analyzed from video recordings using classification of head turns to preferred and to nonpreferred directions. The results showed a significant increase in the alpha responses during the forward paired conditioning to the preferred direction and rapid extinction during the subsequent backward conditioning sessions. Spontaneous head movements during the ITIs increased to the same preferred direction as the conditioned alpha responses. The results of this experiment suggest that the response initially elicited by the CS can later appear as “spontaneous,” instrumental behavior, the form and the nature of which is determined by the characteristics of the conditioned alpha response developing as a result of classical conditioning.     

Reinstatement of fear in humans: autonomic and experiential responses in a differential conditioning paradigm

The present study investigated reinstatement of fear in humans using an aversive differential conditioning paradigm. Two neutral human face pictures were presented during habituation, acquisition, extinction, and postreinstatement phases. One picture served as a conditioned stimulus (CS) reinforced by an unconditioned stimulus (US) in the form of electrical stimulation (CS+) and the second picture as a control stimulus that was never reinforced (CS-). The prediction that in a reinstatement manipulation a previously extinguished fear response in humans can be reinstated in a reinstatement group by the mere presentation of three unpredicted electrical stimulations (USs) was tested. Participants in the control group were not exposed to unpredicted USs and no reinstatement effect was expected. Outcome measures included subjective US expectancy ratings and skin conductance responses. Results showed non-selective return of the fear response due to fear recovery associated with both CSs (CS+/CS-) in the reinstatement group. Unexpected fear recovery was observed for both CSs (CS+/CS-) in control participants. Results are discussed with respect to context conditioning, fear generalisation, and anxiety-related cognitive mechanisms underlying fear recovery after extinction.     

Instructed fear learning,extinction, and recall: additive effects of cognitive information on emotional learning of fear

The effects of instruction on learning of fear and safety are rarely studied. We aimed to examine the effects of cognitive information and experience on fear learning. Fourty healthy participants, randomly assigned to three groups, went through fear conditioning, extinction learning, and extinction recall with two conditioned stimuli (CS+). Information was presented about the presence or absence of conditioned stimulus–unconditioned stimulus (CS–US) contingency at different stages of the experiment. Information about the CS–US contingency prior to fear conditioning enhanced fear response and reduced extinction recall. Information about the absence of CS–US contingency promoted extinction learning and recall, while omission of this information prior to recall resulted in fear renewal. These findings indicate that contingency information can facilitate fear expression during fear learning, and can facilitate extinction learning and recall. Information seems to function as an element of the larger context in which conditioning occurs.     

Fear conditioning in panic disorder: Enhanced resistance to extinction

Enhanced conditionability has been proposed as a crucial factor in the etiology and maintenance of panic disorder (PD). To test this assumption, the authors of the current study examined the acquisition and extinction of conditioned responses to aversive stimuli in PD. Thirty-nine PD patients and 33 healthy control participants took part in a differential aversive conditioning experiment. A highly annoying but not painful electrical stimulus served as the unconditioned stimulus (US), and two neutral pictures were used as either the paired conditioned stimulus (CS+) or the unpaired conditioned stimulus (CS-). Results indicate that PD patients do not show larger conditioned responses during acquisition than control participants. However, in contrast to control participants, PD patients exhibited larger skin conductance responses to CS+ stimuli during extinction and maintained a more negative evaluation of them, as indicated by valence ratings obtained several times throughout the experiment. This suggests that PD patients show enhanced conditionability with respect to extinction.     

Contextual control of the extinction of conditioned fear

Rats received 15 pairings of a CS and shock in one context, and then a series of CS-alone trials in a second context. Even though this extinction procedure produced a complete loss of conditioned suppression, when the animals were returned to the site of original conditioning, suppression was renewed to a level comparable to that of animals that had not undergone extinction. Controls indicated that the renewed suppression was not due solely to pseudoconditioning, suggesting that the CS-US association had survived extinction. Renewed suppression was also demonstrated in a third context that was never associated with shock. Loss of suppression did not necessarily depend upon inhibitory conditioning of the extinction context. The data suggest that extinction of conditioned fear is specific to the context in which it occurs. They also suggest the possibility that animals might discriminate episodes in which a CS is reinforced and nonreinforced independently of the excitatory or inhibitory status of cues, like contextual stimuli, that are coincidentally present during those episodes.     

Pavlovian Disgust Conditioning and Generalization: Specificity and Associations With Individual Differences

Although studies have identified differences between fear and disgust conditioning, much less is known about the generalization of conditioned disgust. This is an important gap in the literature given that overgeneralization of conditioned disgust to neutral stimuli may have clinical implications. To address this knowledge gap, female participants (n = 80) completed a Pavlovian conditioning procedure in which one neutral food item (conditioned stimulus; CS+) was followed by disgusting videos of individuals vomiting (unconditioned stimulus; US) and another neutral food item (CS–) was not reinforced with the disgusting video. Following this acquisition phase, there was an extinction phase in which both CSs were presented unreinforced. Importantly, participants also evaluated generalization stimuli (GS+, GS?) that resembled, but were distinct from, the CS after each conditioning phase. As predicted, the CS+ was rated as significantly more disgusting and fear inducing than the CS? after acquisition and this pattern persisted after extinction. However, disgust ratings of the CS+ after acquisition were significantly larger than fear ratings. Participants also rated the GS+ as significantly more disgusting, but not fear inducing, than the GS? after acquisition. However, this effect was not observed after extinction. Disgust proneness did predict a greater increase in disgust and fear ratings of the CS+ relative to the CS? after acquisition and extinction. In contrast, trait anxiety predicted only higher fear ratings to the CS+ relative to the CS? after acquisition and extinction. Disgust proneness nor trait anxiety predicted the greater increase in disgust to the GS+ relative to the GS? after acquisition. These findings suggest that while conditioned disgust can generalize, individual difference variables that predict generalization remain unclear. The implications of these findings for disorders of disgust are discussed.     

Partial reinforcement effects in instrumental escape conditioning

In Experiment I acquisition and extinction of instrumental escape conditioning with rats (N = 64) were studied as a function of reinforcement magnitude under conditions of partial and continuous reinforcement. In Experiment II the effects of partial and continuous reinforcement were studied in rats (N = 96) during acquisition followed by small, medium, and large reductions in reinforcement magnitude. A water-tank escape apparatus was used with temperature as the relevant variable. It was found that (1) with large reinforcement magnitude a continuously reinforced group was superior in acquisition to one that was partially reinforced; there were no differences with small reinforcement; (2) disruptive effects of a nonreinforced trial (a) appear early in learning, (b) are quite strong after each nonreinforced trial, and (c) persist through several succeeding reinforced trials; (3) major competing behaviors persist throughout acquisition for small reinforcement magnitude regardless of schedule, decline with large reinforcement (more so with continuous than with partial), and return to a high level in extinction for all conditions; (4) the partial reinforcement extinction effect occurs after large reinforcement but not after small, and it appears only with large reductions in reinforcement magnitude which approach extinction conditions. Only the first part of the last finding appears to be consistent with the appetitive conditioning literature.