Facilitation of instrumental behavior by a Pavlovian appetitive conditioned stimulus

Three experiments examined appetitive Pavlovian-instrumental interactions by presenting separately trained conditioned stimuli (CSs) during reinforced instrumental responding in rabbits. Intra-oral reinforcement was used to minimize interference from peripheral responses such as magazine approach. In experiment 1, the rabbits were first trained to perform an instrumental head-raising response for sucrose reward. A conditioned jaw movement response was then established to a 2-sec CS by pairing it with sucrose; a control stimulus was unpaired with sucrose. Instrumental responding maintained by a variable-interval 40-sec schedule was enhanced during 10-sec presentations of the paired, but not the unpaired, CS. Responding on a variable-ratio 15 schedule was unaffected except on trials on which the pre-CS baseline response rate was low; in such cases the paired CS caused a long-lasting acceleration of responding. Noncontingent presentation of the sucrose reinforcer itself briefly suppressed responding but had no long-term effect. In Experiment 2, a CS that had been conditioned at a 10-sec duration produced the same pattern of effects as in the first study, indicating that facilitation resulted from CS presentation rather than from the frustrative effects of non-reinforcement of the CS. In Experiment 3 an inhibitory CS blocked facilitation by the excitatory CS but did not itself affect instrumental responding. These results support the view that Pavlovian processes play a positive role in instrumental performance and suggest that previous findings of suppression by a short-duration CS reflect peripheral interference. The dependence of facilitation on the baseline level of responding is discussed in terms of associative and motivational theories of Pavlovian mediation.     

Counterconditioning of appetitive and defensive CRs in rabbits

Two experiments examined interactions between conditioned appetitive and defensive responses in the rabbit. In Experiment I, a conditioned jaw-movement response was established by following presentations of a clicker CS with intra-oral sucrose delivery on 50% of trials. The jaw-movement response was then maintained on this partial reinforcement schedule during a counterconditioning phase. A group which received para-orbital shock paired with the CS on non-sucrose trials showed acquisition of eyeblink responding and suppression of jaw-movement responding to the CS, in comparison to control groups which received either no shock or unpaired presentations of the CS and shock. Experiment II was identical in design to Experiment I except that the roles of the sucrose and shock reinforcers were reversed. The paired group acquired a conditioned jaw-movement response when sucrose was added in the counterconditioning phase, but in contrast to Experiment I showed a slight enhancement of the previously established eyeblink response. The asymmetry of appetitive and defensive counterconditioning was discussed in relation to opponent-process theories of motivation and reinforcement.     

Marking effects in instrumental performance on DRH schedules

Three experiments investigated the effect of presenting a brief stimulus after a response sequence on the rate of lever-pressing by rats on differential reinforcement of high rate (DRH) schedules. In Experiment 1 enhanced responding was produced by a visual stimulus presented during a 500-msec delay of reinforcement compared to a condition in which no stimulus was presented. In Experiment 2 rats responded on a multiple DRH DRH schedule in which the DRH contingency was reinforced on a 50% schedule in each component. Equivalent levels of responding occurred in the components when reinforcement was signalled in one component and when the signal was presented following the non-reinforced schedules in the other components. A further group of rats received the stimulus presented after non-reinforced schedules in one component but not at all in the other component; responding was enhanced in the former component relative to the latter component. In Experiment 3 brief stimuli presented after the completion of DRH components on a second-order VR (DRH) schedule elevated response rates irrespective of whether the signal was presented paired or unpaired with reinforcement. The present data support the view that a brief signal may serve to mark a response sequence in memory and facilitate instrumental performance.     

The effects of lateral, medial, or complete septal lesions on positive conditioned suppression in rats

This study investigated the effects of lateral, medial, or complete septal lesions in rats on lever pressing and US-approach behaviors during the presentation of a light followed by two free food pellets. Ten days after surgery, groups of rats received 20 sessions of a random interval (RI) 60-s schedule of food reinforcement. The positive conditioned suppression paradigm consisted of 20 sessions of a 10-s light paired with free food while the rats were responding on the RI 60-s schedule. All groups of rats significantly suppressed lever pressing during the 10-s light presentation. During the last four sessions of the 10-s light condition, rats with lateral septal lesions had a significant increase in food-tray entries during the light presentation, while the other groups decreased lever pressing without a change in food-tray entries during the 10-s light presentation. Although rats with septal damage generally have difficulty inhibiting responses in a variety of operant situations, rats with lateral, medial, or complete septal lesions showed no impairment in positive conditioned suppression. This study also suggests that when the spatial arrangement of the CS and the type of US were manipulated to minimize sign-tracking and goal-tracking behaviors, emotional reactions during the presentation of the CS must be considered a factor influencing positive conditioned suppression.     

Interstimulus interval and delivery cues influence timed conditioned responding in rats

Appetitive contextual excitation supported by intertrial unconditioned stimuli was more easily overcome by timed conditioned responding in rats using quiet (Experiment 1) rather than noisy (Experiment 2) food pellet deliveries. Head-entry responding in acquisition peaked above the contextual baseline when pellet delivery occurred 10, 30, 60, or 90 s after the onset of the 120-s white-noise conditioned stimulus (CS). Special tests in extinction revealed CS onset and offset were conditioned by pellet delivery at 0 and 120 s, respectively. Responding was not undermined in Experiment 3 when noisy pellet deliveries replaced quiet pellet deliveries. Our results suggest that micro-stimuli occasioned at different times during the CS are vulnerable to overshadowing, but do not lose strength if they are already predictive.     

Food-paired stimuli as conditioned reinforcers: effects of d-amphetamine.

Seven pigeons were studied in two experiments in which key pecks were reinforced under a second-order schedule wherein satisfaction of variable-interval schedule requirements produced food or a brief stimulus. In the second part of each session, responses produced only the brief stimulus according to a variable-interval schedule (food extinction). For the 4 pigeons in Experiment 1, the response key was red throughout the session. In separate phases, the brief stimulus was either paired with food, not paired with food, or not presented during extinction. d-Amphetamine (0.3 to 10.0 mg/kg) dose-dependently reduced food-maintained responding during the first part of the session and, at intermediate dosages, increased responding during the extinction portion of the session. The magnitude of these increases, however, did not consistently depend on whether the brief stimulus was paired, not paired, or not presented. It was also true that under nondrug conditions, response rates during extinction did not differ reliably depending on pairing operations for the brief stimulus. In Experiment 2, 3 different pigeons responded under a procedure wherein the key was red in the component with food presentations and blue in the extinction component (i.e., multiple schedule). Again, d-amphetamine produced dose-related decreases in responding during the first part of a session and increases in responding in the second part of the session. These increases, however, were related to the pairing operations; larger increases were observed when the brief stimulus was paired with food than when it was not or when it was not presented at all. Under nondrug conditions, the paired brief stimulus controlled higher response rates during extinction than did a nonpaired stimulus or no stimulus. These findings suggest that d-amphetamine can enhance the efficacy of conditioned reinforcers, and that this effect may be more robust if conditioned reinforcers occur in the context of a signaled period of extinction.     

Effects of contextual conditioning and unconditional stimulus presentation on performance in appetitive conditioning

Four experiments with rat subjects examined the effects of contextual conditioning on conditioned appetitive performance. Experiment 1 compared the effects of contextual conditioning on performance to conditioned stimuli (CSs) with different conditioning histories. Contextual conditioning enhanced performance to the CS if the CS had first been conditioned and then extinguished, but had no effect on performance when the CS had been merely paired or unpaired with food. Experiments 2 and 3 then asked whether the effect on the extinguished CS was due to contextual conditioning acting as a cue for conditioning. In Experiment 2, extinction procedures in which extra unconditioned stimuli (USs) were presented during the intertrial intervals were found to reduce the CS's sensitivity to enhancement by contextual conditioning, but had no effect on spontaneous recovery. In Experiment 3, USs added to conditioning or extinction acquired the ability to cue the corresponding performance. Under some conditions, USs added to conditioning could suppress performance (Experiment 4). The results suggest that contextual conditioning has complex effects that can be better understood by recognizing that contextual conditioning, as well as the USs that create it,Mayacquire discriminative control over conditioned responding.     

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.     

Evidence for expectancy as a mediator of avoidance and anxiety in a laboratory model of human avoidance learning

A laboratory model was developed to study human avoidance learning. Participants could avoid an electric shock signalled by a 5-s conditioned stimulus (CS) by pressing one of a set of response buttons. Self-reported shock expectancy and skin conductance were recorded during a subsequent 10-s interval before shock. Shock expectancy declined when the correct avoidance response was learned and returned when the response was unavailable. Learning transferred to another shock CS. Parallel effects were observed on skin conductance once performance anxiety was controlled by requiring responding on all trials. Learning was faster when the Pavlovian contingencies were trained before introduction of the instrumental response. The results support a cognitive model of anxiety in which performance of an avoidance response reduces expectancy of an aversive outcome and thereby reduces anxiety.     

Evidence for expectancy as a mediator of avoidance and anxiety in a laboratory model of human avoidance learning

A laboratory model was developed to study human avoidance learning. Participants could avoid an electric shock signalled by a 5-s conditioned stimulus (CS) by pressing one of a set of response buttons. Self-reported shock expectancy and skin conductance were recorded during a subsequent 10-s interval before shock. Shock expectancy declined when the correct avoidance response was learned and returned when the response was unavailable. Learning transferred to another shock CS. Parallel effects were observed on skin conductance once performance anxiety was controlled by requiring responding on all trials. Learning was faster when the Pavlovian contingencies were trained before introduction of the instrumental response. The results support a cognitive model of anxiety in which performance of an avoidance response reduces expectancy of an aversive outcome and thereby reduces anxiety.     

Posttraining prefrontal lesions impair jaw movement conditioning performance,but have no effect on accompanying heart rate changes

This experiment examined the role of the medial prefrontal cortex (mPFC) in regulating learned autonomic and somatomotor responses in rabbits using appetitive Pavlovian conditioning. Interstimulus interval (ISI) duration [i.e., the time between the onset of the conditioned stimulus (CS) and unconditioned stimulus (US)] was manipulated in order to determine whether ISI duration was related to the heart rate (HR) responses obtained during conditioning. Two groups received either a 1- or a 4-s ISI, with a tone as the CS and an intraoral pulse of water as the US. Another two groups received explicitly unpaired presentations of either the 1- or 4-s tone CS and water US. Few conditioned jaw movement (JM) or HR conditioned responses (CRs) were observed in the unpaired conditions. Significant JM conditioning was, however, elicited by the paired conditions, especially to the 4-s ISI. Consistent CS-evoked HR accelerations were observed in both ISI conditions. After five sessions of training, the mPFC was lesioned in half the animals. A separate group of paired animals received sham lesions. After surgical recovery, all animals received 3 days of postoperative training. During the first postoperative training session, JM CRs significantly declined in both groups with mPFC lesions in comparison to the groups with sham lesions. The mPFC lesions, however, did not affect the CS-evoked cardiac accelerations, which again occurred during postoperative training.     

Examining stimuli paired with alternative reinforcement to mitigate resurgence in children diagnosed with autism spectrum disorder and pigeons

In two laboratory experiments, we examined whether stimuli paired with alternative reinforcers could mitigate resurgence of a previously reinforced target response with pigeons (Experiment 1) and children diagnosed with Autism Spectrum Disorder (Experiment 2). In Phase 1, we arranged food reinforcement according to a variable-ratio schedule for engaging in a target response. In Phase 2, we arranged extinction for target responding and differentially reinforced alternative responding according to a fixed-ratio schedule, with every alternative-reinforcer delivery paired with a change in keylight color (Experiment 1) or automated verbal (praise) statement (Experiment 2). In Phase 3, we assessed resurgence during extinction of target and alternative responding in the presence versus absence of continued presentation of the paired stimulus. Despite variation across sessions, resurgence on average was lower when continuing to present the paired stimuli in all pigeons and children while maintenance of alternative responding did not differ between assessments. These findings indicate that stimuli paired with alternative reinforcement can modestly decrease resurgence, but further examination of their efficacy and a better understanding of the underlying processes are necessary before they can be recommended for clinical use in reducing resurgence of clinically relevant problem behavior.     

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.     

Conditioned reinforcement and discrimination in second-order schedules

Pigeons were exposed to multiple second-order schedules in which responding on the “main key” was reinforced according to either a variable-interval or fixed-interval schedule by production of a brief stimulus on the “brief-stimulus key”. A response was required to the brief stimulus during its fourth (final) presentation to produce food; responses to the earlier brief stimuli indicated the extent to which the final brief stimulus was discriminated from preceding ones. Main-key response rates were higher in early components of paired brief-stimulus schedules, in which each brief stimulus was the same as that paired with reinforcement, than in comparable unpaired brief-stimulus or tandem schedules. Poor discrimination occurred between paired brief stimuli (Experiment I). When chain stimuli on the main key induced a discrimination between the first two and second two brief stimuli, the response-rate enhancement in the paired brief-stimulus schedule persisted (Experiment II). Rate enhancement diminished when the initial link of the chain included the first three components (Experiment IV). Eliminating the contingency between responding and brief-stimulus production also diminished rate enhancement (Experiment III). The results show that the discriminative and conditioned reinforcing effects of food-paired brief stimuli may be selectively manipulated and suggest that the reinforcing effects are modulated by other reinforcers in the situation.     

Acquisition and extinction of signaled avoidance as a function of intermittent reinforcement

Signaled, shuttle-box avoidance responding in female rats of the Fischer₃₄₄ strain was examined as a function of four separate contingencies of intermittent reinforcement. In Experiment 1, when avoidance responses during acquisition were reinforced 25% of the time with prompt CS termination, animals responded equally often during acquisition and significantly more often during extinction than animals who received such reinforcement on a 100% schedule. Similar results were found under a trace procedure in Experiment 2 when avoidance responses were reinforced 25% of the time with informational feedback stimuli. In contrast, during Experiment 3, when animals were shocked on only 25% of the trials on which they failed to respond, the level of avoidance responding during both acquisition and extinction was significantly less than it was when animals were shocked on a 100% schedule. Comparable results were found in Experiment 4 when avoidance responses during acquisition averted shock on only 25% of the trials. Thus, intermittent reinforcement contingencies involving response-contingent feedback stimuli and shock have differential effects on avoidance responding during both acquisition and extinction trials under the signaled avoidance procedure.     

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.     

Extinction of goal tracking also eliminates the conditioned reinforcing effects of an appetitive conditioned stimulus

Previous studies have suggested that the effects of extinction are response-specific. The present study investigated whether an extinction treatment that eliminated goal tracking elicited by an appetitive conditioned stimulus (CS) would also eliminate the conditioned reinforcing effects of that CS. Rats were first trained on a goal-tracking procedure in which an auditory CS was paired with a food unconditioned stimulus. Animals learned to approach the location where the food was delivered. In a subsequent phase, rats in one group received extinction training that eliminated the goal-tracking elicited by the CS. Rats in the other group did not experience extinction of the food-paired CS. Then, both groups received a test for conditioned reinforcement in which leverpresses resulted in the brief presentation of the stimulus previously paired with food. This stimulus did not act as a conditioned reinforcer in the group that had been subjected to extinction training, but did serve as a conditioned reinforcer in the group that did not experience extinction. These results indicate that the effects of extinction generalize from the approach-eliciting to the conditioned reinforcing effects of an appetitive CS.     

Pavlovian second-order conditioned analgesia

Three experiments with rat subjects assessed conditioned analgesia in a Pavlovian second-order conditioning procedure by using inhibition of responding to thermal stimulation as an index of pain sensitivity. In Experiment 1, rats receiving second-order conditioning showed longer response latencies during a test of pain sensitivity in the presence of the second-order conditioned stimulus (CS) than rats receiving appropriate control procedures. Experiment 2 found that extinction of the first-order CS had no effect on established second-order conditioned analgesia. Experiment 3 evaluated the effects of post second-order conditioning pairings of morphine and the shock unconditioned stimulus (US). Rats receiving paired morphine-shock presentations showed significantly shorter response latencies during a hot-plate test of pain sensitivity in the presence of the second-order CS than did groups of rats receiving various control procedures; second-order analgesia was attenuated. These data extend the associative account of conditioned analgesia to second-order conditioning situations and are discussed in terms of the mediation of both first- and second-order analgesia by an association between the CS and a representation or expectancy of the US, which may directly activate endogenous pain inhibition systems.     

Effects of expectancies of different reward magnitudes in transfer from noncontingent pairings to instrumental performance

In two experiments, rats received noncontingent pairings of two stimuli with food reward, one paired with small reward and the other with large reward, and received bar press training with large reward or with small reward. When the noncontingent stimuli (NS) were presented for test during subsequent rewarded bar pressing and during early extinction of bar pressing, responding for each group was faster in the presence of the NS which was paired with the same reward magnitude that group received in bar press training than to the NS which had been paired with a different reward magnitude. As extinction progressed, all groups responded more slowly in the presence of the NS which had been paired with the large reward than in the presence of the NS which had been paired with small reward. These results were interpreted as indicating that responding in the presence of an NS depends on: (i) whether the reward expectancy elicited by the NS has been conditioned to the instrumental response, and (ii) the relationship between the reward expected in the presence of the NS and that received in test.     

Timeout postponement without increased reinforcement frequency

Three experiments were conducted to examine pigeons' postponement of signaled extinction periods (timeouts) from a schedule of food reinforcement when such responding neither decreased overall timeout frequency nor increased the overall frequency of food reinforcement. A discrete-trial procedure was used in which a response during the first 5 s of a trial postponed an otherwise immediate 60-s timeout to a later part of that same trial but had no effect on whether the timeout occurred. During time-in periods, responses on a second key produced food according to a random-interval 20-s schedule. In Experiment 1, the response-timeout interval was 45 s under postponement conditions and 0 s under extinction conditions (responses were ineffective in postponing timeouts). The percentage of trials with a response was consistently high when the timeout-postponement contingency was in effect and decreased to low levels when it was discontinued under extinction conditions. In Experiment 2, the response-timeout interval was also 45 s but postponement responses increased the duration of the timeout, which varied from 60 s to 105 s across conditions. Postponement responding was maintained, generally at high levels, at all timeout durations, despite sometimes large decreases in the overall frequency of food reinforcement. In Experiment 3, timeout duration was held constant at 60 s while the response-timeout interval was varied systematically across conditions from 0 s to 45 s. Postponement responding was maintained under all conditions in which the response-timeout interval exceeded 0 s (the timeout interval in the absence of a response). In some conditions of Experiment 3, which were designed to control for the immediacy of food reinforcement and food-correlated (time-in) stimuli, responding postponed timeout but the timeout was delayed whether a response occurred or not. Responding was maintained for 2 of 3 subjects, suggesting that behavior was negatively reinforced by timeout postponement rather than positively reinforced by the more immediate presentation of food or food-correlated (time-in) stimuli.