Pavlovian appetitive discriminative conditioning inAplysia californica

Three experiments demonstrated Pavlovian appetitive discrimination learning in the marine mollusc,Aplysia californica. In each experiment, subjects were exposed to two conditioned stimuli; one stimulus (CS+) was paired with food presentations and the other stimulus (CS?) was never followed by food. In Experiments 1 and 3 different chemosensory stimuli were used, and in Experiment 2 different tactile stimuli were used. For both types of conditioned stimuli, bite responses occurred significantly more often to the CS+ than to the CS?. Experiment 2 also showed thatAplysia could learn a reversal of this discrimination. Experiment 3 showed that nonreinforced presentations of CS+ resulted in a decline in the frequency of conditioned biting. The implications of these results for neurobiological analyses of learning are discussed.     

Overexpectation in appetitive Pavlovian and instrumental conditioning

In three experiments that used appetitive preparations with rats, we examined the effects of reinforcing a compound consisting of two previously reinforced stimuli on subsequent responding to those stimuli. Experiment 1 showed that a Pavlovian conditioned stimulus given this treatment evoked fewer magazine entries when presented alone than did a reinforced stimulus that did not receive the compound treatment. Experiment 2 examined inhibition of delay and generalization decrement accounts for the results of Experiment 1. Experiment 3 extended this finding to an instrumental learning paradigm.     

Spontaneous recovery of instrumental discriminative responding

In six experiments, rats received discriminative training in which making a response (R) during a stimulus (S) produced a particular outcome (O). In Experiment 1, that outcome was replaced by a second outcome and responding was tested either immediately or after a delay. More substantial responding was observed with the delayed test. In Experiment 2, a test of transfer to new responses suggested that the growth in performance was not attributable to greater use of particular S-O associations. However, in Experiment 3, the growth in responding was found to be specific to particular S-R combinations. Experiment 4 replicated that specificity and demonstrated the importance of using two different outcomes for obtaining the growth in responding with time. Experiments 5 and 6 repeated these observations for the case of extinction, in which O was replaced by nonreinforcement. These results are interpreted as suggesting that an outcome-independent inhibitory S-R process develops, both with extinction and with the use of a second outcome, but dissipates with time.     

Transfer of control between causal predictive judgments and instrumental responding

Four experiments were conducted to explore outcome-specific transfer from causal predictive judgments to instrumental responding. A video game was designed in which participants had to defend Andalusia from navy and air force attacks. First, they learned the relationship between two instrumental responses (two keys on a standard keyboard) and two different outcomes (destruction of the ships or destruction of the planes). Then they learned to predict which of two different stimuli predicted which outcome. Finally, they had the opportunity of making either of the two instrumental responses in the presence of either stimulus. Transfer was shown as a preference for the response that shared an outcome with the current stimulus. The presentation of the stimulus during the test produced a decrease in the overall rate of response. Responding to a neutral stimulus in Experiments 2 and 3 suggested that this overall decrease in responding was due to a combination of the time needed to process the meaning of the stimulus and the activation of the representation of the outcome in the presence of the stimulus during the test. Transfer between predictive judgments and instrumental responding mirrors the outcome-specific Pavlovian instrumental transfer observed in conditioning studies with rats.     

Behavioral variability is controlled by discriminative stimuli

Previous research has demonstrated that behavioral variability can be modified by reinforcers contingent on it, but there has been no convincing evidence of discriminative stimulus control over such variability. We therefore rewarded 20 rats for variable response sequences in the presence of one stimulus and provided equal rewards independently of sequence variability in the presence of a second stimulus. We found that sequence variability was significantly higher during the first stimulus than during the second, with the greatest difference occurring immediately following onset of the stimuli. Removing the discriminative stimuli caused levels of variability to converge. These experiments provide strong evidence that behavioral variability can be controlled by discriminative stimuli, which may be important for general theories of operant behavior and their applications.     

Modulation of trained and extinguished stimuli by facilitators and inhibitors

Pigeon subjects received training of a diffuse stimulus as either a conditioned inhibitor or a conditioned facilitator of a keylight signal for food. The ability of these diffuse stimuli to modulate responding was then assessed with two other keylights that were undergoing discrimination reversal. At a point where responding was equivalent for the two targets, the modulators had a greater impact on the target undergoing extinction than on the target undergoing acquisition. These results have implications for the nature of modulation and extinction.     

Pavlovian aversive to instrumental appetitive transfer: Evidence for across-reinforcement blocking effects

Rats received Pavlovian aversive (shock) conditioning in which white noise was established for different groups as a CS+, CSO, or CS?. Then, in an appetitive T-maze discrimination, the CSs were presented contingent upon a designated correct response for which food reinforcement was factorially varied at 0, 1, 2, or 4 pellets. Although the CS+ suppressed and the CS? facilitated speed of running in the correct arm at the start of discrimination training, these effects extinguished rapidly and did not interact with reward magnitude. Furthermore, choice learning was faciltated by the CS+ and retarded by the CS?, with these effects being comparable for the 1- to 4-pellet reinforcement conditions, but absent for the 0-pellet condition. These findings are difficult to reconcile with a transfer interpretation positing a general signaling property of the CS and are better interpreted as across-reinforcement blocking effects: By predicting a preferred outcome (safety) comparable to the preferred outcome of food reinforcement, the CS? blocks (retards) the association of reinforcement and the S^D; conversely, by predicting a nonpreferred (shock) outcome discrepant from the preferred food outcome, the CS+ “counterblocks” (enhances) the association of reinforcement and the S^D.     

Renewal of extinguished instrumental responses: independence from Pavlovian processes and dependence on outcome value

The source of renewal of instrumental responding in rats was investigated. In Experiment 1, two responses (R1 and R2) were reinforced with one outcome (O1) in contexts A and B (i.e., R1→O1, R2→O1), and then R2 was extinguished in A and R1 was extinguished in B. At test, the rate of R1 was higher than that of R2 in context A, and the reverse was the case in context B: Renewed responding was independent of the Pavlovian context→O1 associations. In Experiment 2, all rats received R1→O1 and R2→O2 trials in A and then were placed in B, where they were sated on O2 and either did (Group Extinction) or did not (Group No Extinction) receive concurrent extinction of R1 and R2. At test, we found more responding in A than in B for Group Extinction, but not for Group No Extinction, and the renewed responding in A was as sensitive to the current value of the outcome as responding that had not been subject to extinction (i.e., the rate was higher for R1 than for R2). That is, the renewed responding was goal-directed. These results identify the removal of contextual inhibion of either the response or the response→outcome associaon as potenal bases for renewal, and the response→outcome associaon as the source of renewed responding.     

Integration of cardiac responses to serial stimuli after Pavlovian conditioning in rats

The expression of cardiac responses to sequences of two sounds was studied in restrained rats following discriminative trace or delay conditioning. Stimuli paired with a tail shock 10 sec later (CS1) elicited conditioned bradycardia. Unpaired or neutral stimuli (CS0) elicited mostly tachycardia. Rats did not learn to suppress responding to nonreinforced sequences with an interval of 6 sec between sounds. Responses to the second stimulus were significantly augmented following a CS1 stimulus, but not following a CS0 stimulus. Real-time summation of simple responses provided a more complete and quantitative prediction of dual responses than did resetting or facilitation. These results extend the time range over which summation may be observed from less than 2 sec to at least 16 sec. They appear to be inconsistent with models involving competition between unitary representations of stimuli in short-term memory and suggest the existence of multiple stimulus traces with independent time courses.     

Transfer testing after serial feature-ambiguous discrimination in Pavlovian keypeck conditioning

Four experiments were performed to explore modulatory transfer after serial feature-ambiguous (F-A) discrimination (X→A+, A?, X→B?, B+) in Pavlovian keypeck conditioning (autoshaping) with pigeons. Diffuse features were used in the first three experiments, and no modulatory transfer was found: (1) There was no modulatory transfer between two concurrently trained F-A tasks; (2) modulatory transfer to conditioned and then extinguished stimuli was not observed; and (3) responding to the targets (A and B) after their contingencies were reversed was not affected by presentation of their original feature stimulus (X). Transfer between two serial F-A tasks was obtained in the last experiment, in which keylights were used as features, but other evidence indicated that this was due to stimulus generalization between the features. Together, the results strongly suggest that specific target-food occasion setting or configural learning is the underlying mechanism of F-A discrimination.     

Negative discriminative stimuli provide information about the identity of omitted response-contingent outcomes

Three experiments using rats examined whether a signal for the nonreinforcement of an instrumental response (S-) provided information about the identity of the omitted outcome. In all three experiments, one stimulus was established as a signal for the nonreinforcement of a response that earned food pellets and another stimulus signaled the nonreinforcement of a response that earned liquid sucrose. Experiment 1 found that each S-suppressed another instrumental response trained with the same outcome significantly more than a response trained with a different outcome. Using a variant of this transfer design, Experiment 2 demonstrated that an S- was slower to develop discriminative control over a new response reinforced in its presence with the same outcome compared with an outcome different from the one whose omission-the S- had previously signaled. Experiment 3 examined transfer of the S- stimuli to a response trained with two outcomes, one of which had subsequently been devalued. Performance of this response significantly increased in the presence of a signal for the omission of the devalued outcome, but decreased in the presence of a signal for the omission of the valued outcome. These results suggest that S-s do provide information about the identity of omitted response-contingent outcomes.     

Transfer of discriminative control during stimulus fading conducted without reinforcement

Using trial-and-error training, eight pigeons did not learn to discriminate between 45° and 135° lines, but did learn to discriminate between red and green colors. Control by line tilt was induced by stimulus fading that did not include reinforcement while fading out the colors. After establishing the red–green discrimination, low-intensity lines were superimposed on colors and were gradually faded in. All of this was done using reinforcement. At the end of the line fade-in, the lines had not acquired control of responding. Finally, color intensity was gradually faded out in the absence of reinforcement, and the lines acquired discriminative control by six of the eight pigeons. Thus, reinforcement during the color fade-out was not necessary for the acquisition of discriminative control by the lines during fading. Acquisition of control by lines was attributed to overshadowing, the reduction of stimulus blocking by generalization, and the evocation of correct responding by the colors while the participants were attending to the lines. This last process was also responsible for the induction of discriminative control during sensory preconditioning, higher order conditioning, and response transfer in equivalence classes. Errors, however, were not correlated with discrimination learning during stimulus fading. Finally, transfer of control occurred very quickly with or without errors.     

Selective reinstatement of instrumental performance depends on the discriminative stimulus properties of the mediating outcome

We conducted three experiments to investigate the associative structure underlying the reinstatement of instrumental performance after extinction. In each experiment, rats were initially rewarded on two responses with different outcomes. At test, both responses were extinguished in order to assess the impact of a single noncontingent outcome delivery on response selection. Experiment 1 found evidence of outcome-selective reinstatement (i.e., more responses were performed on the lever that was trained with the reinstating outcome than on the other lever). Experiment 2 demonstrated that the outcome’s capacity to reinstate performance was not affected by a reduction in its motivational value. Experiment 3 found evidence that the reinstating outcome selectively retrieved the response it signaled rather than the response it followed during training. Together, these findings are consistent with the view that instrumental reinstatement depends on the discriminative stimulus properties of the reinstating outcome.     

Transfer of instrumental control mediated by a devalued outcome

In three instrumental learning experiments, rat subjects were used to explore transfer of an instrumental discriminative stimulus to a new response. The stimulus increased the likelihood of that response to the degree that the stimulus and response had a history of association with the same outcome. Moreover, devaluation of the outcome by pairing with lithium chloride had no detrimental effects on its ability to mediate transfer to a new response. This result helps one choose among various accounts of transfer.     

The overshadowing of background stimuli by an informative CS in aversive Pavlovian conditioning with rats

Two experiments investigated the capacity of a nominal CS to overshadow background stimuli. Rats received CS-shock pairings in one compartment of a double-compartment apparatus. After training, the shock compartment was re-presented but in the absence of both the CS and shock, and overshadowing was assessed by measuring the rats’ latency to enter the compartment. In the first experiment, predictive values of temporal cues provided by CS onset and CS offset were manipulated. Rats readily entered the shock compartment, provided shock onset had previously been predicted by CS onset or CS offset. This indicates that background cues in the shock compartment had acquired little associative strength during training. In contrast, rats avoided the shock compartment under every condition in which shock onset had not previously been predicted by either CS onset or CS offset. In the second experiment, a precise temporal cue provided by CS onset was dissociated from other factors arising from CS-shock contiguity. It was found that rats’ reactions to the shock compartment after training was governed by both temporal and contiguity factors. The results are relevant to theories which state that conditioning is a function of a competitive relationship between stimuli present in an animal’s environment during training.     

Intrinsic reinforcing properties of putatively neutral stimuli in an instrumental two-lever discrimination task

Four experiments examined the influence of a stimulus presented after one response in a two-lever choice task. In Experiment 1, food-deprived rats trained on a concurrent variable-interval extinction schedule responded more often on the extinction lever when such responding periodically produced a visual stimulus than when it did not. In Experiments 2 and 3, a similar signal-induced enhancement effect was found even when food was delivered randomly with respect to responding on both levers or when no food was presented. In Experiment 4, a response-contingent visual stimulus elevated responding to the lever on which it was presented, but an auditory cue suppressed responding. These findings indicate that visual stimuli may possess intrinsically reinforcing properties for rats.     

Transfer of conditioned suppression and conditioned acceleration from instrumental to consummatory baselines

Following training on a variable-interval food reinforcement schedule, rats were exposed to Pavlovian procedures which produced reliable conditioned suppression and conditioned acceleration of the leverpressing (instrumental) baseline. When free food was simultaneously made available in the test cage, all subjects spent the majority of each session “freeloading,” that is, eating food from a dish rather than leverpressing for it. When superimposed upon the freeloading baseline, the conditioned suppression and conditioned acceleration procedures affected the rate of pellet consumption identically in magnitude and direction to their previous effects on leverpressing. These results suggest a motivational mechanism for conditioned suppression and acceleration, rather than one which depends upon spurious punishment of specific response sequences.     

Interactions between retroactive-interference and context-mediated treatments that impair Pavlovian conditioned responding

In Pavlovian fear conditioning, context-mediated decrements in conditioned responding (e.g., the US preexposure effect) can counteract competition between cues trained together (e.g., overshadowing). Two experiments were conducted using rats in a conditioned lick suppression preparation to determine whether context-mediated competition also counteracts competition between cues trained separately (retroactive interference, or RI). In Experiment 1, a combination of degraded contingency and RI treatments produced less of a decrement in conditioned responding than did either of those treatments alone. Experiment 2 showed that RI treatment attenuates the normally deleterious effect of trial massing. The results suggest that empirical similarities are shared by interference between cues trained apart and competition between cues trained together.