Asymmetrical effects of Pavlovian excitatory and inhibitory aversive transfer on Pavlovian appetitive responding and acquisition

Predictions of a theory of Pavlovian motivational transfer, which incorporates principles of both the theory of reciprocal inhibition and the Rescorla-Wagner model, were tested in several Pavlovian aversive to Pavlovian appetitive transfer tasks. As predicted, the presence of a signal for an aversive event, conditioned stimulus (AV CS+), reliably suppressed performance of appetitive conditioned responses (CRs) whether imposed during acquisition or on independently established responding. Acquisition of appetitive responding to a novel CS reinforced in compound with an AV CS+, however, was enhanced (“superconditioning”). This observation suggests that the effects of a discrepancy between expectation and actual outcome on a conditioning trial are influenced by the affective value of both the expectation and the reinforcer. These transfer effects were not symmetrical for an inhibitory aversive stimulus (AV CS?). An AV CS? did not enhance appetitive responding compared to a random control condition, nor did the AV CS? reduce (i.e., block) appetitive conditioning to a novel CS when appetitive reinforcement occurred in the presence of the AV CS?. Comparison of the two shock-exposed conditions with a naive control condition suggests that previous results that were apparently consistent with inhibitory aversive enhancement and blocking of appetitive conditioning may have been due to aversive context conditioning.     

Emotional eating and Pavlovian learning: evidence for conditioned appetitive responding to negative emotional states

Appetitive learning has been demonstrated several times using neutral cues or contexts as a predictor of food intake and it has been shown that humans easily learn cued desires for foods. It has, however, never been studied whether internal cues are also capable of appetitive conditioning. In this study, we tested whether humans can learn cued eating desires to negative moods as conditioned stimuli (CS), thereby offering a potential explanation of emotional eating (EE). Female participants were randomly presented with 10 different stimuli eliciting either negative or neutral emotional states, with one of these states paired with eating chocolate. Expectancy to eat, desire to eat, salivation, and unpleasantness of experiencing negative emotions were assessed. After conditioning, participants were brought into a negative emotional state and were asked to choose between money and chocolate. Data showed differential conditioned responding on the expectancy and desire measures, but not on salivation. Specific conditioned effects were obtained for participants with a higher BMI (body mass index) on the choice task, and for participants high on EE on the unpleasantness ratings. These findings provide the first experimental evidence for the idea that negative emotions can act as conditioned stimuli, and might suggest that classical conditioning is involved in EE.     

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.     

The roles of event contingencies and reinforcement in human autoshaping and omission responding

Four experiments were conducted to test those features deemed to play an important role in the acquisition and/or maintenance of autoshaping and omission responding with human subjects, viz., event contingencies, reinforcement, stimulus generalization, the type of CS employed, conditioned reinforcement, and response topography. One important aspect of these studies is that the boundary conditions were made to resemble those pertaining to the majority of nonhuman animal studies, i.e., magazine training followed by paired CS-US presentations. The results demonstrated the importance of CS-UR or, alternatively, CS-US contiguity together with generalization and a response-reinforcer contingency for autoshaped response initiation and maintenance, respectively. The conditioned reinforcing effect of CS offset appears to play a part in generating and maintaining some responding during omission training, especially during a response-terminated CS procedure. Similar response topography was observed when the form of the UR was compared to that of the CR or autoshaped response. Taken as a whole the results of these experiments with humans are directly comparable to results previously reported using a wide variety of nonhuman animals.     

Enhancement of food-rewarded instrumental responding by an appetitive conditioned stimulus

Four experiments are reported in which a stimulus (with a minimum duration of 60 s) signalling the delivery of “free” food was presented to rats lever-pressing for food available on a variable interval schedule. It was found that responding was enhanced in the presence of the stimulus when the baseline schedule of reinforcement was lean (Experiment I) and that the enhancement was dependent upon the pairing of the stimulus with free food (Experiments II and III). Experiment IV showed that an enhancement could be found after initial training in which stimulus-food pairings were given to subjects that were not concurrently lever pressing for food. It is argued that these results are consistent with the suggestion that an appetitive conditioned stimulus can energise appetitive instrumental behaviour.     

Differential effects of two ways of devaluing the unconditioned stimulus after Pavlovian appetitive conditioning

Three experiments with rat subjects investigated the effects of two methods of devaluing a food unconditioned stimulus (US) after pairings of an auditory conditioned stimulus (CS) with that US. Experiment 1 found no effect of postconditioning pairings of the food US with lithium chloride (LiCl) on general activity to a tone CS, even though those pairings substantially reduced food consumption. Experiments 2 and 3 compared the effects on conditioned responding of postconditioning pairings of food with LiCl and with high-speed rotation. In these experiments the general activity measure was supplemented by a detailed visual analysis of the rats' behavior. Experiment 2 found that food-rotation pairings had larger effects than food-LiCl pairings on general activity responding and on two detailed behavioral measures but that food-LiCl pairings had larger effects on food consumption and on one behavioral measure. Experiment 3 replicated the findings of Experiment 2 and found that the ability of the CS to serve as a reinforcer for second-order conditioning after US devaluation was reduced more by food-LiCl pairings.     

Temporal distributions of responding during discrete-trial omission training in rats

Within-session temporal distributions of responding were investigated in three experiments using rats pressing a lever in a discrete-trial omission procedure. This schedule entailed 60, one-minute trials, and a sucrose solution was made available at the end of each trial in which no lever press occurred. In Experiment I, nonnaive rats acquired and maintained responding during this training. Moreover, the probability of a response during any session showed a strong and reliable tendency to increase from the beginning to the end of the session. These results were replicated in Experiment II, using naive animals. In Experiment III, alterations were made in the training procedure, including elimination of response-contingent and noncontingent stimulus changes. Results indicate that stimulus change may be sufficient to maintain low levels of responding whether or not this change is contingent on responding.     

The effects of aversive conditioned stimuli on the timing of unsignalled avoidance responding in rats

Rats received unsignalled avoidance training in a shuttlebox. After acquisition, classical conditioning sessions were administered. The Excitatory Group received signals paired with shock, the Inhibitory Group received signals explicitly unpaired with shock, while the Random Control Group received signals and shocks randomly in time. When subsequently superimposed on the avoidance baseline, the excitatory signal increased response rate, the inhibitory signal decreased rate and the random signal had no effect compared to signal-only control data. Unsignalled avoidance generates temporal gradients of responding; the major purpose of this study was to determine how these rate changes were reflected in the baseline temporal behavior. Very clear and simple summation rules emerged: In the presence of the excitatory signal, rats acted as if they were ahead of their actual location in the response-shock interval regardless of where the signal occurred; the inhibitory signal had the opposite effect while the random signal did not change the temporal behavior relative to signal-only control data. These results were interpreted within a motivational framework.     

Naloxone decreases responding for conditioned reinforcement in rats

That opioids can mediate unconditioned reinforcement is well established, but there is little evidence indicating whether they modify conditioned reinforcement. Here, a tone which initially served as a discriminative stimulus for the availability of water reinforcement was established as a conditioned stimulus. When later given a choice between pressing a lever producing the tone (but not water) or one which produced no effect, rats chose the tone-producing lever 66% of the time. Naloxone (3.0 mg/kg) reduced overall responding and completely eliminated the preference for the tone-producing lever. Results are briefly discussed in terms of the importance of understanding mechanisms serving conditioned reinforcement.     

Drug effects on responding maintained by stimulus-reinforcer and response-reinforcer contingencies.

The effects of pentobarbital and d-amphetamine were assessed on key pecking by pigeons under conventional single-key multiple schedules and under two-key multiple schedules in which discriminative stimuli appeared on one key (stimulus key) while pecks on a second key (constant key) produced food. Pecks on the stimulus key had no scheduled consequences. A 60-second variable-interval schedule operated in one component of each multiple schedule: either extinction or a 60-second variable-time schedule operated in the alternate component. When the alternate-component schedule was extinction, a high rate of responding was maintained in the variable-interval component of the single-key schedule; responding on both keys was maintained in the variable-interval component of the two-key schedule. Pentobarbital increased responding in the variable-interval component of the single-key schedule and increased stimulus-key, but not constant-key responding in that component of the two-key schedule. When the alternate-component schedule was changed to variable time, responding declined in the variable-interval component of the single-key schedule; stimulus-key responding was no longer maintained under the two-key schedule. Pentobarbital decreased responding in the variable-interval component of both schedules. With an exception, d-amphetamine only decreased responding in the variable-interval component of the single- and two-key schedules both when the alternate-component schedule was extinction and when it was variable time. The results suggest that the effects of pentobarbital, but not d-amphetamine, depend on the nature of the contingency (stimulus-reinforcer, response-reinforcer) that maintains responding.     

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.     

Remote effects of aversive contingencies: Disruption of appetitive behavior by adjacent avoidance sessions

Disruption of ongoing appetitive behavior before and after daily avoidance sessions was examined. After baselines of appetitive responding were established under a fixed-interval 180-s schedule of food presentation, 4 rats were exposed to 40-min sessions of the appetitive schedule just prior to 100-min sessions of electric shock postponement, while another 4 rats received the 40-min appetitive sessions just following daily sessions of shock postponement. In all 8 subjects, fixed-interval response rates decreased relative to baseline levels, the effect being somewhat more pronounced when the avoidance sessions immediately followed. The disruption of fixed-interval responding was only partially reversed when avoidance sessions were discontinued. During the initial exposure to the avoidance sessions, patterns of responding under the fixed-interval schedule were differentially sensitive to disruption, with high baseline response rates generally more disturbed than low rates. These disruptions were not systematically related to changes in reinforcement frequency, which remained fairly high and invariant across all conditions of the experiment; they were also not systematically related to the response rates or to the shock rates of the adjacent avoidance sessions. The results, while qualitatively resembling patterns of conditioned suppression as typically studied, occurred on a greatly expanded time scale. As disruption of behavior extending over time, the present data suggest that some forms of conditioned suppression are perhaps best viewed within a larger temporal context.     

Intertrial pellets influence the acquisition and expression of timed appetitive responding in rats

Two experiments examined temporally based changes in the conditioned magazine-entries of rats when a target food pellet arrived at a fixed time before the termination of a conditioned stimulus. Both experiments found that increasing the rate of intertrial pellets systematically interfered with the rate of acquisition. When intertrial pellets were featured in acquisition, their omission on test trials also resulted in a very poorly expressed conditioned response. On test trials preceded by intertrial pellets, however, peak times in groups trained with intertrial pellets were shifted in a rightward direction under a long but not short interstimulus interval. This rightward shift was accompanied by a smaller standard deviation. Our key findings are best accommodated by real-time associative models.     

Stimulus- and response-reinforcer contingencies in autoshaping, operant, classical, and omission training procedures in rats

Separate groups of rats received 500 trials of lever-press training under autoshaping (food delivery followed 10-second lever presentations, or occurred immediately following a response); operant conditioning (responding was necessary for food delivery); and classical conditioning (food followed lever presentations regardless of responding). Each group then received 500 trials on an omission procedure in which food was omitted on trials with a response. Another group received 1000 trials on the omission procedure, and a fifth group, random control, received 1000 uncorrelated presentations of lever and food. The autoshaping, operant, and classical groups reached high response levels by the end of initial training. Acquisition was fastest in the autoshaping group. Responding remained consistently low in the control group. The omission group responded at a level between the control group and the other three groups. During omission training, responding in these three groups declined to the omission-group level. During omission training, the rats continued contacting the lever frequently after lever pressing had declined. Response maintenance under omission training seems not to require topographic similarity between the response and reinforcer-elicited consummatory behaviors.     

Analysis of the learned irrelevance effect in appetitive Pavlovian conditioning

In Experiments 1 and 2 rats received uncorrelated presentations of a light conditioned stimulus (CS) and a food unconditioned stimulus (US) on each day of a preexposure phase. Control subjects received the same number of USs during the first half of preexposure and the same number of CSs during the second. Uncorrelated preexposure retarded inhibitory conditioning. Experiment 3 showed, however, that the different patterns of US preexposure experienced by the two groups could in itself influence the course of subsequent inhibitory conditioning. When this factor was equated by restricting the uncorrelated treatment to the first half of the pre-exposure phase (Experiment 2) or by extending the control treatment throughout the phase (Experiment 4) it was found that uncorrelated preexposure retarded excitatory conditioning, but facilitated inhibitory conditioning. This outcome challenges an interpretation in terms of the concept of learned irrelevance, which predicts that uncorrelated preexposure should retard both forms of conditioning.     

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.     

Aftereffects of the surprising presentation and omission of appetitive reinforcers on key-pecking performance in pigeons

The reinforcement-omission effect (ROE), also known as frustration effect, refers to greater response strength immediately after nonreinforcement (N) than reinforcement (R). The ROE was traditionally interpreted as transient invigoration after N induced by primary frustration. Pigeons demonstrate similar ROEs whether outcomes are surprising (partial R) or expected (discrimination training) in runway (Experiment 1) and Skinner box situations (Experiments 2-3). Variations in the interval between N and the opportunity to respond indicate that the ROE results from an aftereffect of food consumption (Experiment 4). Increasing reinforcer magnitude increased the after-R effect, without modifying the after-N function (Experiment 5). These results are reviewed in the context of comparative research on spaced-trial successive negative contrast and related phenomena that have failed to appear in experiments involving nonmammalian vertebrates.