Behavioral and pharmacological influences on phencyclidine discrimination in the pigeon

Discriminative stimuli produced by drugs control behavior in the same way as exteroceptive discriminative stimuli control behavior, despite the difficulty in controlling the intensity of the discriminative stimulus properties of drugs throughout test sessions. In recent years, many investigators have correlated the potency of drugs as discriminative stimuli, with their affinity for specific pharmacological receptors. High correlations have been interpreted as evidence that the discriminative stimulus properties of drugs are mediated by these specific pharmacological receptors. However, the relationship between discriminative stimulus potency and receptor affinity can be confounded by other pharmacological effects of drugs, such as their ability to produce position responding, and by behavioral variables, such as the schedule of reinforcement under which the drug discrimination is established and measured.     

Behavioral and associative effects of differential outcomes in discrimination learning

The role of the reinforcer in instrumental discriminations has often been viewed as that of facilitating associative learning between a reinforced response and the discriminative stimulus that occasions it. The differential-outcome paradigm introduced by Trapold (1970), however, has provided compelling evidence that reinforcers are also part of what is learned in discrimination tasks. Specifically, when the availability of different reinforcing outcomes is signaled by different discriminative stimuli, the conditioned anticipation of those outcomes can provide another source of stimulus control over responding. This article reviews how such control develops and how it can be revealed, its impact on behavior, and different possible mechanisms that could mediate the behavioral effects. The main conclusion is that differential-outcome effects are almost entirely explicable in terms of the cue properties of outcome expectancies—namely, that conditioned expectancies acquire discriminative control just like any other discriminative or conditional stimulus in instrumental learning.     

Training reinforcement rates,resistance to extinction,and the role of context in reinstatement

Behavior reduced as a consequence of extinction or intervention can relapse. According to behavioral momentum theory, the extent to which behavior persists and relapses once it has been eliminated depends on the relative training reinforcement rate among discriminative stimuli. In addition, studies of context renewal reveal that relapse depends on the similarity between the training stimulus context and the test stimulus context following disruption by extinction. In the present experiments with pigeons, we arranged different reinforcement rates in the presence of distinct discriminative stimuli across components of a multiple schedule. Following extinction, we attempted to reinstate responding in the presence of those target components with response-independent food presentations. Importantly, we arranged the reinstating food presentations either within the target components or in separate components, either paired with extinction (Experiment 1) or reinforcement (Experiment 2) during baseline. Reinstatement increased with greater training reinforcement rates when the reinstating food presentations were arranged in the target components and the separate components paired with reinforcement during training. Reinstatement was smaller and was not systematically related to training reinforcement rates in the target components when reinstating food presentation occurred in separate components paired with extinction. These findings suggest that relapse depends on the history of reinforcement associated with the discriminative stimuli in which the relapse-inducing event occurs.     

Overshadowing by food of stimulus control by a keylight in a two-element serial compound

Pigeons were given successive discrimination training in which pecking during a choice period when the key was white was either reinforced or not, depending upon the prior presence or absence of a discriminative stimulus, which was a two-element serial compound. The compound consisted of a keylight and food, with food presented second or first in a forward or backward pairing for different groups of pigeons. In Experiment 1, the sequence was an S+ indicating reinforced trials, while in Experiment 2, the sequence was an S? indicating nonreinforced trials. Following acquisition of discriminated operant behavior, a sequence generalization test was administered during which all possible orders of the two stimuli were presented on test trials prior to the onset of the choice period. The results showed that food overshadowed stimulus control by the color of the light on the key on the sequence-generalization test, independently of whether food was presented first or second during training and independently of whether food was associated with reinforcement or nonreinforcement. The similarity of results for the two experiments suggests that overshadowing occurs independently of whether the compound is a discriminative stimulus for reinforcement or nonreinforcement. Simultaneous presentation of elements of a compound stimulus is not necessary for overshadowing because the phenomenon was captured with sequentially presented stimuli.     

Higher order occasion setting

Higher order occasion setting with serially presented stimuli was investigated in an appetitively motivated, discrete-trial operant study with rats. Reinforcement of barpressing during an occasion-setting light (a discriminative stimulus) was contingent on immediately preceding second-order occasion setters (i.e., a click train or a buzzer served as a conditional discriminative stimulus). Moreover, the meanings of the clicks and buzzer were themselves indicated by a third-order occasion setter that preceded them (i.e., a white noise acted as a second-order conditional discriminative stimulus). Subjects responded more frequently and had shorter latencies to the first response in the presence of the light on trials during which barpressing was reinforced than on trials during which barpressing was not reinforced. The likelihood that the subjects solved the problem by responding to unique compound stimuli was minimized by the insertion of a 5-sec gap between the different controlling stimuli presented on each trial. Thus, these subjects appear to have mastered a second-order conditional discrimination, which is equivalent to third-order occasion setting if the discriminative stimulus (light) is viewed as a first-order occasion setter. Although the subjects learned to respond appropriately to each of the compound stimuli, differences in responding to specific stimuli were consistent with a higher order feature-positive effect. Some implications of higher order occasion setting are discussed, including the issue of independence between the different levels of occasion setting signaled by a single stimulus.     

The effect of the controllability of auditory discriminative stimuli in the performance of go/no-go discriminations by pigeons

Compared with their performance with localized (on-key) visual stimuli, pigeons are notoriously poor at performing go/no-go discriminations when keypecking for food in the presence of auditory discriminative stimuli. The difference might reflect the fact that an aversive visual onkey stimulus signaling nonreward can be escaped by looking away and not pecking, which contributes to the measure of good discriminative performance, while an auditory stimulus cannot be escaped. In Experiment 1, discriminative performance was significantly improved by providing pigeons with a response incompatible with keypecking by which they could escape a tone S+ and a tone S?. However, the pattern, frequency, and duration of escape responses were found to be insufficient to explain the improvement. In Experiment 2, it was found that the capacity to escape only S+ or only S? enhanced discriminative performance as much as the capacity to escape both. It is theorized that the Pavlovian relationship between the absence of the discriminative stimuli and the nonoccurrence of food might transfer to the instrumental relationships learned in a go/no-go discrimination. The possibility that intermittent stimuli command more attention than continuous stimuli is also considered.     

Cross-modal transfer of stimulus control in the albino rat: A stimulus delay procedure

The present experiment demonstrated in a simultaneous discrete trial discrimination that the stimulus control of a rat’s leverpress response can be errorlessly transferred across stimulus modalities, i.e., from light to click location and from click to light location. Subsequent to acquisition of the original discrimination, the original and new discriminative stimuli were simultaneously presented for several sessions. Then the new discriminative stimulus was presented 3 sec prior to the onset of the original discriminative stimulus. Within the direction of transfer, e.g., from light to click location, the delay group emitted fewer trial and intertriai errors than the control group. As the new discriminative stimuli acquired control over responding, the response latency distributions were differentially affected. The results suggest that the transfer of control from the original to the new discriminative stimuli is mediated by the temporal aspects of the delay interval.     

Behavioral momentum and relapse of extinguished operant responding

Previous experiments on behavioral momentum have shown that relative resistance to extinction of operant behavior in the presence of a stimulus depends on the rate of reinforcement associated with that stimulus, even if some of those reinforcers occur independently of the behavior. We present three experiments examining whether the rate of reinforcement in the presence of a stimulus similarly modulates the relative relapse of operant behavior produced by reinstatement, resurgence, and renewal paradigms. During baseline conditions, pigeons responded for food reinforcement on variable-interval 120-sec schedules in alternating periods of exposure to two stimuli arranged by a multiple schedule. Additional response-independent food presentations were also delivered in the presence of one of the multiple-schedule stimuli. Consistent with previous research, baseline response rates were lower in the presence of the stimulus with the added response-independent reinforcement, and relative resistance to extinction was greater in the presence of that stimulus. In addition, following extinction, the relative relapse of responding produced by reinstatement, resurgence, and renewal paradigms was greater in the presence of the stimulus associated with the higher rate of reinforcement. We suggest that a model of extinction from behavioral momentum theory may be useful for understanding these results.     

Some tests of the additivity (autoshaping) theory of behavioral contrast

Two experiments were performed to determine whether the location of the discriminative stimuli affects the amount of positive behavioral contrast exhibited during discrimination learning by pigeons. When signals for reinforcement and nonreinforcement of keypecking were situated directly on the response key (different line tilts), pecking rates during the positive stimulus were higher than when the source of the signals was located elsewhere (changes in chamber illumination or auditory click frequency). These results are in general agreement with the additivity theory of behavioral contrast, which attributes contrast to the combined effects of stimulus-reinforcer and response-reinforcer correlations on behavior directed at signals of reinforcement. Some shortcomings of the theory were discussed, and the notion that behavioral contrast is a basic, unitary phenomenon was criticized.     

Multiple-schedule interactions and discrimination

Pigeons received variable-interval (VI) reinforcement for keypecking during randomized presentations of seven line-orientation stimuli forming a continuum ranging from horizontal (0 deg) to vertical (90 deg). Each line presentation lasted for 30 sec and was preceded and followed by 30-sec time-outs. After responding stabilized, only responding in the two extreme stimuli (0 and 90 deg) was reinforced. As discrimination training proceeded, strong behavioral contrast and dimensional contrast effects appeared. However, only marginal local effects (local contrast and local dimensional effects), exerted by one line-orientation component upon a second, appeared, indicating that behavioral and dimensional contrast may be independent of parallel local effects. An attempt was made to apply Blough’s (1975) quantitative model of operant generalization and discrimination to the present discrimination procedure. However, this model did not predict the generalization gradient shape that was experimentally obtained. This experiment also yielded two serendipitous findings: (1) Positive behavioral contrast appeared in an extinction-related stimulus (time-out) when other stimuli were switched from reinforcement to extinction (hitherto, positive behavioral contrast had been observed only in responding to a reinforcementrelated stimulus when other stimuli were switched from reinforcement to extinction), and (2) final responding was higher in the presence of an extinction stimulus that had always been an extinction stimulus than it was in the presence of other extinction stimuli that had previously been paired with VI reinforcement.     

Stimulus functions within a fixed-interval clock schedule: Reinforcement,punishment, and discriminative stimulus control

Pigeons received food for responding on a fixed-interval 32-sec schedule divided into three equal parts, each correlated with a distinctive, response-independent, visual stimulus. Response rate was very low during the first two thirds of the interval but high during the terminal third. When a response-dependent brief stimulus correlated with the terminal third was arranged for each response in the presence of the stimuli correlated with the first two thirds, response rate was enhanced, especially in the middle third. However, response rate was suppressed when each response in the presence of the stimulus correlated with the final third produced a brief stimulus correlated with the initial third. A similar suppressive effect occurred when each response produced a brief stimulus correlated with the middle third. Response suppression decreased over successive response-dependent brief-initial-stimulus manipulations. The results were interpreted in terms of reinforcement, punishment, and discriminative stimulus control by visual stimuli correlated with parts of a fixed-interval schedule.     

Instrumental responses become associated with reinforcers that differ in one feature

In three experiments, thirsty rats were trained to make several instrumental responses whose outcomes differed in which of two relatively inconsequential flavor features they contained. In Experiment 1, one of the features was subsequently devalued by pairing it with lithium chloride; in Experiment 2, it was enhanced in value by pairing it with sucrose. In both experiments, differences in the value of the features resulted in parallel differences in the likelihood of the responses during a subsequent extinction test. In Experiment 3, the animals chose between these responses in the presence of discriminative stimuli that had signaled the occurrence of these different features following another response. The stimuli selectively augmented the likelihood of the response with which they shared training by the same-flavored consequence. These results indicate that rats can separately encode features that differ along one dimension, both in the association between an instrumental response and its outcome, and in the association between a discriminative stimulus and that outcome.     

Differential inhibition and stimulus generalization cannot account for value transfer in simultaneous discrimination learning by pigeons: Reply to Aitken

Aitken (1999) argues that, in a simultaneous discrimination, reports of the transfer of value from the positive to the negative stimulus can be more readily explained in terms of an artifact produced by the procedure in which differential inhibition accrues to the negative test stimuli during training, together with stimulus generalization (similarity between the positive and negative stimuli). We argue that (1) there is little evidence for differential inhibition, and it often occurs in the wrong direction; (2) value transfer can be demonstrated when differential value is established to the positive stimuli afterdiscrimination training, when differential inhibition is not likely to be a factor; and (3) on both logical and empirical grounds, stimulus similarity does not provide an adequate account of the transfer of value from the positive to the negative stimulus (i.e., the strongest evidence for value transfer occurs when there is least stimulus similarity). We propose that value transfer occurs whenever there is relatively little experience with the negative stimuli. However, when there is extended experience with the negative stimuli, contrast will be found.     

Failure to block control by a relevant stimulus

Pigeons were trained to depress a treadle in the presence of a discriminative stimulus, either a tone or illumination of red houselights, in order to obtain access to grain or avoid electric shock. In avoidance training, the auditory discriminative stimulus yielded faster acquisition than did the visual one. In appetitive training, the visual discriminative stimulus yielded faster acquisition than the auditory one. Experiments 2 and 3 used these stimuli in Kamin’s (1969) blocking design. In Experiment 2, when the pigeons were trained to depress a treadle in the presence of tone to obtain grain and then red light was added as the redundant stimulus, the light acquired stimulus control over treadlepressing; blocking was not observed. In Experiment 3, when the pigeons were trained to depress a treadle in the presence of red light to avoid electric shock and then tone was added as the redundant stimulus, the tone acquired stimulus control over treadle-pressing. Again, blocking was not observed. The implications of these results for several models of stimulus control are discussed.     

Sequential processes in the generalization and transfer of stimulus control

Prior research indicated that a training sequence consisting of a negative stimulus followed by a positive stimulus constitutes the minimal condition for the production of postdiscrimination phenomena typically observed after training with random sequences of the discriminanda. The present experiments, employing multiple schedules with pigeon subjects, confirmed the earlier findings but indicated that they are restricted to procedures in which the reinforcing stimulus may acquire a discriminative function that competes with the control exerted by the nominal discriminanda. The sequences in which the discriminanda were presented did not differentially affect subsequent measures of generalization and transfer if the discriminative function of reinforcement were degraded either by introducing some reinforcers during the negative stimulus (Experiment 1) or by omitting some reinforcers during the positive stimulus (Experiment 2). It was concluded that the sequence in which the discriminanda are presented during discrimination training does not contribute fundamentally to the processes responsible for discrimination formation with random training sequences.     

Time-course of control by specific stimulus features and relational cues during same-different discrimination training

We trained 7 pigeons to discriminate visual displays of 16same items from displays of 16different items. The specific stimulus features of the items and the relations among the items could serve as discriminative stimuli. Unlike in most studies of same-different discrimination behavior, we gave a small number of probe tests during each session of acquisition to measure the time-course of control by the learning of specific stimulus features and relational cues. Both the specific stimulus features and relational cues exerted reliable stimulus control, with the specific stimulus features exerting more control during the final three fourths of same-different learning. These findings replicate research suggesting that pigeons encode both the specific stimulus features and relational cues, and for the first time document the time-course of control by each kind of cue.     

Signaling the omission of a response-contingent outcome reduces discriminative control

The effect of training a positive discriminative stimulus (S+ ) as a signal for the nonreinforcement of an instrumental response (S?) on the ability of that stimulus to evoke its original instrumental response was examined in three experiments using rats. In all three experiments, two different stimuli were established as S+s for different response-outcome relations. In Experiment 1, an S+ was less effective in controlling its original response after it had undergone training as an S? for a new response that earned the same outcome than it was after training as an S? for a response that earned a different outcome. Experiment 2 established that this effect was not mediated by Pavlovian inhibitory conditioning produced by the negative correlation between the S+ and the outcome during S? training. Simply arranging a negative correlation between S+ and the outcome whose occurrence it had previously signaled did not impair the ability of that S+ to elicit its original response. In Experiment 3, the response-evoking properties of an S+ were found to be undermined by using the S+ as a signal for the simple extinction of a new response trained with the same outcome, but not with a different outcome. These results suggest that positive discriminative stimuli use their associations with the outcomes earned in their presence to control the responses that earned those outcomes.     

The learned function of food-deprivation cues: A role for conditioned modulation

Rats trained in one context to use stimuli arising from food deprivation as discriminative signals for shock were tested in other contexts to assess the basis of conditioned responding (i.e., freezing or behavioral immobility). In Experiment 1, discriminative control by 24-h food-deprivation cues failed to promote transfer responding in a test context that had no association with shock. This indicated that food deprivation cues had little direct excitatory power. However, transfer of behavioral control by 24-h food-deprivation cues was obtained in a context paired with shock only when the rats were 19 h water deprived. This finding agrees with the idea that food-deprivation cues become conditioned modulators of the capacity of external stimuli to activate their association with an unconditioned stimulus. In Experiment 2, rats trained to use 24-h food-deprivation cues as signals for shock exhibited significantly greater transfer performance when the transfer context had undergone partial extinction relative to when the transfer context had undergone only simple excitatory training. This finding with deprivation cues and transfer contexts (1) paralleled earlier results obtained with discrete (auditory and visual) conditioned modulators and transfer targets, and (2) posed difficulties for associative summation and generalization interpretations of transfer performance.     

Response rate is not an effective mediator of learned stimulus equivalence in pigeons

We explored response rate as a possible mediator of learned stimulus equivalence. Five pigeons were trained to discriminate four clip art pictures presented during a 10-sec discrete-trial fixed interval (FI) schedule: two paired with a one-pellet reinforcer, which supported a low rate of responding, and two paired with a nine-pellet reinforcer, which supported a high rate of responding. After subjects associated one stimulus from each of these pairs with a discriminative choice response, researchers presented two new clip art stimuli during a 10-sec FI: one trained with a differential reinforcement of low rate schedule (DRL) after the FI and the other trained with a differential reinforcement of high rate schedule (DRH) after the FI. Each of the stimuli that were withheld during choice training was later shown to see if the choice responses would transfer to these stimuli. The results suggest that response rate alone does not mediate learned stimulus equivalence.     

Blocking and enhancement of fear conditioning by appetitive CSs

Prior research on Pavlovian-to-instrumental transfer has shown that when a CS previously associated with shock (AvCS+) is presented contingent upon a choice response to a discriminative stimulus for food reinforcement, it facilitates discrimination learning. Conversely, a response-contingent CS previously associated with the absence of shock (AvCS?) retards discrimination learning. To evaluate whether these findings reflect across-reinforcement blocking and enhancement effects, two experiments investigated the effects of appetitively conditioned stimuli on fear conditioning to a novel stimulus that was serially compounded with the appetitive CS during conditioned-emotional-response (CER) training. Although there were no differential effects of the appetitive CSs in CER acquisition, Experiment 1, using a relatively weak shock US, showed that a CS previously associated with food (ApCS+) retarded CER extinction to the novel stimulus, in evidence of enhanced fear conditioning to that stimulus. In addition, Experiment 2, using a stronger shock US, showed that a CS previously associated with the absence of food (ApCS?) facilitated CER extinction to the novel stimulus, in evidence of weaker fear conditioning to that stimulus. These results parallel traditional blocking effects and indicate not only that an ApCS+ and an ApCS? are functionally similar to AvCSs of opposite sign, but that their functional similarity is mediated by common central emotional states.