Food delivery as a conditional stimulus: Feature-learning and memory in pigeons

Three experiments investigated the learning and memory of discriminations based on presence versus absence of a pre-trial food delivery. In Experiment 1 half the illuminations of a response key were followed by food regardless of the subject's behavior. In one group an extra food delivery preceded only reinforced trials (feature-positive condition), whereas in a second group it preceded only nonreinforced trials (feature-negative condition). Key pecks and approaches revealed more rapid and superior discrimination learning in the first group. Experiment 2 replicated the results of Experiment 1 but yielded no evidence that greater “unexpectedness” of pretrial food conditions facilitates discriminative performance. In Experiment 3, individual pigeons trained on a conditional discrimination exhibited a within-subject feature-positive superiority. Delay between pretrial and trial stimuli interacted with feature-positive versus feature-negative training in both the between-group (Experiment 2) and within-subject (Experiment 3) procedures: performance was decremented at both short and long delays in the feature-positive condition but was decremented only at longer delays in the feature-negative condition. The feature-positive superiority obtained here is incompatible with explanations based on either the general concept of “perceptual organization” or on the conditional nature of feature-negative discriminations.     

Blocking of occasion setting in feature-positive discriminations

In two experiments rats received feature-positive discrimination training in which brief conditioned stimuli (CSs) were paired with food during presentations of an extended feature stimulus, and non-reinforced in its absence. In Experiment 1 a novel feature was trained in compound with a second, pretrained feature. Acquisition of control over responding to the CS by the novel feature was blocked if the pretrained feature had also been trained in a feature-positive discrimination, compared to a group for whom the pretrained feature had been accompanied by uncorrelated presentations of CSs and food. Experiment 2 employed a within-subjects design. It demonstrated that the feature from a feature-positive discrimination with a particular CS, x, blocked acquisition of control by an added, novel feature over responding to x, compared to the control acquired by the same novel feature over a novel, CS y.     

Effect of proximity of elements on the feature-positive effect

Eight groups of pigeons were trained to discriminate between two stimulus displays that could be differentiated only by a single distinctive feature on one of the displays. For half of the pigeons, responses to displays containing the distinctive feature were reinforced (feature-positive), and for the remaining pigeons responses to displays without the distinctive feature were reinforced (feature-negative). The pigeons were further grouped so that half were presented displays in which the distinctive feature was in close proximity to other features (compact displays) and half were presented displays in which the features were not close together (distributed displays). Pigeons in the feature-positive groups localized responses on the distinctive feature of the displays and seldom responded to displays without the distinctive feature. Pigeons in the distributed feature-negative groups localized responses on features common to the two displays and did not learn the discrimination. Compacting the displays facilitated discrimination performance for the subjects in the feature-negative condition. Tests carried out in extinction indicated that responding in the compact feature-negative group was largely controlled by pattern rather than by the individual elements on the display.     

Feature-positive effect in adults and attention to portion of stimulus array

In three experiments, adult humans were tested in a feature-positive or feature-negative simultaneous symbol task. In Experiment 1, some persons focused on the correct side of the stimulus cards, whereas other persons focused on the not-correct side of the stimulus cards. The feature-positive group learned faster than the feature-negative group did in the correct side condition; the feature-negative group learned faster than the feature-positive group did in the not-correct side condition. In Experiments 2 and 3, all persons focused on both the correct and not-correct sides of the stimulus cards. Under these circumstances, feature-positive and feature-negative performances were comparable. These results indicated that the usual superiority of feature-positive over feature-negative learning results from a tendency to attend to only a portion of the stimulus array.     

Occasion-setting training renders stimuli more similar: Acquired equivalence between the targets of feature-positive discriminations

In two experiments rats received training on two concurrent appetitive feature-positive discriminations. A preliminary test in Experiment 1 confirmed previous demonstrations of the transfer of occasion-setting properties—the feature from one of these discriminations was better able to facilitate responding to the occasion-set target CS from the second discrimination than to a control stimulus that had not been the subject of occasion-setting. The source of this transfer was investigated in a second phase of training, and in Experiment 2. In both experiments one of the occasion-set CSs was paired with food, and generalization of appetitive conditioned responding from this stimulus to the second occasion-set CS, and to a control cue, was examined. There was more generalization from the first occasion-set CS to the second CS that had also been occasion-set than to the control cue. This is taken as evidence that occasion-set CSs are rendered more similar as a result of their common training history. The implications of these findings for explaining transfer of occasion setting are discussed.     

Transfer across unconditioned stimuli in serial feature discrimination training

The transfer of conditioned modulation across conditioned stimuli (CS) and unconditioned stimuli (US) was examined in 3 experiments that used Pavlovian appetitive training procedures with rats. In Experiment 1, after training in a positive patterning discrimination (X-->A+/X-/A-), X increased conditioned responding elicited by another trained-then-extinguished CS as long as that CS had been trained with the same US as was used in discrimination training. In Experiment 2, after training with a feature-negative discrimination (X-->A-/A+), X inhibited conditioned responding elicited by another trained-then-extinguished CS as long as that CS had been trained with the same US. Experiments 1 and 2 used a between-groups design, whereas Experiment 3 used a more powerful within-groups design. In Experiment 3, rats were trained in a feature-positive discrimination (X-->A+/A-). In transfer tests, X increased conditioned responding elicited by another CS trained then extinguished with the same US from training. This increase was greater than the X increased conditioned responding elicited by another CS trained then extinguished with a different US from training. The results supported the suggestion that features trained in serial discrimination tasks influence behavior indirectly by transiently raising or lowering the threshold for activation of the US representations by its target stimuli and by any other stimuli that may be associated with that US. Other interpretations of the findings were also considered.     

Intertrial interval as a contextual stimulus: further analysis of a novel asymmetry in temporal discrimination learning

Four experiments investigated discrimination learning when the duration of the intertrial interval (ITI) signaled whether or not the next conditional stimulus (CS) would be paired with food pellets. Rats received presentations of a 10-s CS separated half the time by long ITIs and half the time by short ITIs. When the long ITI signaled that the CS would be reinforced and the short interval signaled that it would not be (Long+/Short-), rats learned the discrimination readily. However, when the short ITI signaled that the CS would be reinforced and the long interval signaled that it would not (Short+/Long-), discrimination learning was much slower. Experiment 1 compared Long+/Short- and Short+/Long- discrimination learning with 16-min/4-min or 4-min/1-min ITI combinations. Experiment 2 found no evidence that Short+/Long- learning is inferior because the temporal cue corresponding to the short interval is ambiguous. Experiment 3 found no evidence that Short+/Long- learning is poor because the end of a long ITI signals a substantial reduction in delay to the next reinforcer. Long+/Short- learning may be faster than Short+/Long-because elapsing time involves exposure to a sequence of hypothetical stimulus elements (e.g., A then B), and feature-positive discriminations (AB+/A-) are learned quicker than feature-negative discriminations (A+/AB-). Consistent with this view, Experiment 4 found a robust feature-positive effect when sequentially presented CSs played the role of elements A and B.     

Selective transfer of responding in conditional discriminations

In each of three experiments rats received discrimination training in which whether or not a 10-sec target stimulus was followed by food was signalled by a 2-min background stimulus. In the first experiment the target was paired with food in the presence but not the absence of the background stimulus. Subsequent tests revealed that the background elevated responding to a target that had taken part in a similar discrimination. However, it had no influence on the responses elicited by a partially reinforced conditioned stimulus. In the remaining experiments the target was paired with food in the absence but not the presence of the background. Test trials then revealed that although the background had an inhibitory influence on the responses elicited by a target from a similar discrimination, it had no influence on the responses elicited by either a partially or a continuously reinforced conditioned stimulus. Various explanations for this selective influence of a background stimulus are considered.     

Blockade of GABAA receptors in the interpositus nucleus modulates expression of conditioned excitation but not conditioned inhibition of the eyeblink response

The cerebellum and related brainstem structures are essential for excitatory eyeblink conditioning. Recent evidence indicates that the cerebellar interpositus and lateral pontine nuclei may also play critical roles in conditioned inhibition (CI) of the eyeblink response. The current study examined the role of GABAergic inhibition of the interpositus nucleus in retention of CI. Male Long-Evans rats were implanted with a cannula positioned just above or in the anterior interpositus nucleus before training. The rats were trained with two different tones and a light as conditioned stimuli, and a periorbital shock as the unconditioned stimulus. CI training consisted of four phases: 1) excitatory conditioning (8 kHz tone paired with shock); 2) feature-negative discrimination (2 kHz tone paired with shock or 2 kHz tone concurrent with light); 3) summation test (8 kHz tone or 8 kHz tone concurrent with light); and 4) retardation test (light paired with shock) After reaching a criterion level of performance on the feature-negative discrimination (40% discrimination), 0.5 μl picrotoxin (a GABA_A receptor antagonist) was infused at one of four concentrations, each concentration infused during separte test sessions. Picrotoxin transiently impaired conditioned responses during trials with the excitatory stimulus (tone) in a dose-dependent manner, but did not significantly impact responding to the inhibitory compound stimulus (tone-light). The results suggest that expression of conditioned inhibition of the eyeblink conditioned response does not require GABAergic inhibition of neurons in the anterior interpositus nucleus.     

Perirhinal cortex lesions impair feature-negative discrimination

The role of the perirhinal cortex in inhibitory eyeblink conditioning was examined. In Experiment 1, rats were given lesions of the perirhinal cortex or control surgery and subsequently trained with a feature-negative discrimination procedure followed by summation and retardation tests for conditioned inhibition. Perirhinal cortex lesions impaired, but did not prevent acquisition of feature-negative discrimination. Results from the summation test showed that rats with perirhinal cortex lesions could not generalize feature-negative discrimination to a new stimulus. There were no group differences during the retardation test. Experiment 2 showed that lesions of the perirhinal cortex did not impair simple excitatory conditioning. Experiment 3 showed that perirhinal cortex lesions had no effect on acquisition of a simple tone-light discrimination. The results suggest that the perirhinal cortex plays a role in eyeblink conditioning when using discrimination procedures involving overlapping stimuli.     

Stimulus control of Pavlovian facilitation

Two experiments were conducted using an autoshaping procedure with pigeons to examine whether dimensional stimulus control by a Pavlovian facilitator parallels the control established following operant discrimination training. Facilitation training consisted of the presentation of a black vertical line on a white background as the B stimulus in a feature-positive discrimination in which the A stimulus (white keylight) was followed by grain presentation only if preceded by B. In this way, B facilitates or sets the occasion for pecking at A. Subsequent testing for generalization along the line-orientation dimension produced decremental gradients when the facilitation paradigm incorporated an explicit feature-negative stimulus (B−). These results parallel the decremental control obtained following operant discrimination training and suggest that Pavlovian facilitators and instrumental discriminative stimuli are functionally equivalent.     

Summation: Further assessment of a configural theory

Rats received Pavlovian conditioning in which food was signalled by a visual stimulus, A+, an auditory stimulus, B+, and a compound composed of different visual and auditory stimuli, CD+. Test trials were then given with the compound AB. Experiments 1 and 2A revealed stronger responding during AB than during CD. In Experiment 2B, there was no evidence of a summation of responding during AB when A+ B+ training was conducted in the absence of CD+ trials. A further failure to observe abnormally strong responding during ABwas found in Experiment 3 for which the training trials with A+ B+ CD+ were accompanied by trials in which C and D were separately paired with food. The results are explained in terms of a configural theory of conditioning, which assumes that responding during a compound is determined by generalization from its components, as well as from other compounds to which it is similar.     

Reminder-induced recovery from blocking as a function of the number of compound trials

Conditioned lick suppression in rats was employed to examine changes in the associative status of a blocked stimulus as a function of the number of compound stimulus conditioning trials. In each of two experiments, prior tone-footshock pairings produced similar blocking of conditioned responding to the light element of a tone-light compound when either two or six compound trials were used. In experiment 1, two exposures to the light alone in a dissimilar context during the retention interval, i.e., a reminder treatment, resulted in a restoration of responding to the light which was complete and similar for animals receiving two or six compound trials. This indicated that latent acquisition with respect to the blocked stimulus was largely complete after two compound trials. Because of a potential ceiling effect for reminder-induced recovery from blocking, Experiment 2 employed an attenuated reminder (one light exposure). This reminder reversed the blocking when six, but not when two, compound trials were used. These results suggest that, after latent acquisition to the blocked stimulus is complete, the blocked stimulus continues to be processed during additional compound stimulus conditioning trials, with a consequent latent facilitation of retrieval of associations to the blocked stimulus.     

Choice as a dependent measure in autoshaping: sensitivity to frequency and duration of food presentation.

Previous investigations have shown that rate, latency, and percentage of trials with at least one response are somewhat insensitive measures of the strength of autoshaped responding. In the present studies, these measures were contrasted with the allocation of responding during simultaneous choice tests, a measure of response strength frequently used in operant paradigms. In two experiments, nine pigeons were exposed to a forward pairing autoshaping procedure. Training sessions consisted of the successive presentation of three stimuli, each followed by food on either 100%, 50%, or 0% of the trials. Choice testing involved the simultaneous presentation of the three stimuli. In Experiment I, all pigeons consistently directed their initial choice responses and the majority of subsequent responses to the stimulus always followed by food, despite the fact that during training sessions the response rates of most birds were highest in the presence of the stimulus followed by food on 50% of the trials. In Experiment II, rate, latency, and percentage of trials with at least one response did not change appreciably as a function of duration of feeder presentations. However, choice responding was lawfully affected by duration of feeder presentations. These data suggest that choice is perhaps a more sensitive measure of the strength of autoshaped responding than other, more commonly employed, indices.     

Discrimination and differentiation of response number in stimulus directed pecking of pigeons.

In Experiment 1, autoshaping trials terminated with food only if pigeons emitted more than a target number of responses during a trial in one condition and fewer than a target number in another. The median number of responses per trial shifted in accordance wtih the requirements. The responding of yoked-control birds that received response-independent reinforcers did not vary with the response requirements. In Experiment 2, the number of responses in autoshaping trial became the discriminative stimulus for reinforcement in the second component of a chained schedule. In one condition, responding was reinforced only if the number of responses in the first component was above a target value; in the other condition, responding was reinforced only if the number was below the target value. The distribution of the first-component response numbers did not shift systematically between discrimination conditions, but response rates in the second component indicated that the number of responses in the autoshaping trial was a discriminable property behavior.     

Extinction reveals stimulus control: latent learning of feature-negative discriminations in pigeons

Various discriminations based on the presence versus absence of a single feature are supposedly learned much better when the feature appears on reinforced rather than nonreinforced trials. However, failures to show discriminative acquisition with the feature on negative trials could reflect a deficiency in control of performance rather than a lack of learning. Five experiments supported this alternative possibility. Pigeons that had yielded little or no evidence of learning (with distinguishing features like a small white square on the response key or a tone located some distance away) revealed clear differences between keypecking to the formerly positive and negative stimuli when all food was removed from the situation. Besides extinction, several other procedures for decreasing the positive predictiveness of the most informative stimulus element also unmasked feature-negative learning, whereas general and specific contextual changes did not. Incompletely mastered feature-positive discriminations improved during extinction, too. The findings of better discrimination performance in extinction were related to analogous effects in previous generalization and discrimination research employing other tasks and arrangements. A sign-tracking analysis could not completely account for the present results.     

Blockade of GABAA Receptors in the Interpositus Nucleus Modulates Expression of Conditioned Excitation but not Conditioned Inhibition of the Eyeblink Response

The cerebellum and related brainstem structures are essential for excitatory eyeblink conditioning. Recent evidence indicates that the cerebellar interpositus and lateral pontine nuclei may also play critical roles in conditioned inhibition (CI) of the eyeblink response. The current study examined the role of GABAergic inhibition of the interpositus nucleus in retention of CI. Male Long-Evans rats were implanted with a cannula positioned just above or in the anterior interpositus nucleus before training. The rats were trained with two different tones and a light as conditioned stimuli, and a periorbital shock as the unconditioned stimulus. CI training consisted of four phases: 1) excitatory conditioning (8 kHz tone paired with shock); 2) feature-negative discrimination (2 kHz tone paired with shock or 2 kHz tone concurrent with light); 3) summation test (8 kHz tone or 8 kHz tone concurrent with light); and 4) retardation test (light paired with shock). After reaching a criterion level of performance on the feature-negative discrimination (40% discrimination), 0.5 microl picrotoxin (a GABAA receptor antagonist) was infused at one of four concentrations, each concentration infused during separate test sessions. Picrotoxin transiently impaired conditioned responses during trials with the excitatory stimulus (tone) in a dose-dependent manner, but did not significantly impact responding to the inhibitory compound stimulus (tone-light). The results suggest that expression of conditioned inhibition of the eyeblink conditioned response does not require GABAergic inhibition of neurons in the anterior interpositus nucleus.     

Short-Term Retention of “Surprising” Events By Pigeons

Terry and Wagner (1975) have suggested that the short-term retention of information about an event is enhanced if the occurrence of the event is made unexpected or surprising. Three experiments tested this idea using delayed conditional position (Experiment I) and colour discriminations (Experiments II and III). The subjects were pigeons and the presentation of food was the target event to be remembered. Choice of one of two simultaneously presented stimuli was reinforced if the retention interval had been initiated by presentation of target food. Contrary to Terry and Wagner's claim, retention was superior on probe test trials in Experiments I and II if the presentation of the target food was preceded by a previously established signal for food (CS+) rather than by a stimulus which had not been paired with food (CS-). Experiment III systematically manipulated the signalling conditions used during initial discrimination training. Retention was better following a CS+ presentation if the target food had been signalled during initial discrimination training but worse if the food presentation had been unsignalled. These results do not favour the idea that retention of an event is directly affected by whether or not the target event is surprising.     

Reversibility of single-incentive selective associations.

Rats were trained to press a lever in the presence of a tone-light compound stimulus and not to press in its absence. In each of two experiments, schedules were designed to make the compound a conditioned punisher for one group and a conditioned reinforcer for the other. In Experiment 1, one group's responding produced food in the presence of the compound but not in its absence. The other group's responding terminated the compound stimulus, and food was presented only in its absence. When tone and light were later presented separately, light controlled more responding than did tone in the former group, but tone gained substantial control in the latter. The same effects were also observed within subjects when the training schedules were switched over groups. In Experiment 2, two groups avoided shock in the presence of the compound stimulus. In the absence of the compound, one group was not shocked, and the other received both response-independent and response-produced shock. When tone and light were presented separately, the former group's responding was mainly controlled by tone, but the latter group's responding was almost exclusively controlled by light. These effects were also observed within subjects when the training schedules were switched over groups. Thus, these single-incentive selective association effects (appetitive in Experiment 1 and aversive in Experiment 2) were completely reversible. The schedules in which the compound should have been a conditioned reinforcer consistently produced visual control, and auditory control increased when the compound should have become a conditioned punisher. Currently accepted accounts of selective associations based on affinities between shock and auditory stimuli and between food and visual stimuli (i.e., stimulus-reinforcer interactions) do not adequately address these results. The contingencies of reinforcement most recently associated with the compound and with its absence, rather than the nature of the reinforcer, determined whether auditory or visual stimulus control developed.     

Derived relational responding as generalized operant behavior

The major aim of the present study was to demonstrate that derived relational responding may be viewed as a form of generalized operant behavior. In Experiment 1, 4 subjects were divided into two conditions (2 in each condition). Using a two-comparison matching-to-sample procedure, all subjects were trained and tested for the formation of two combinatorially entailed relations. Subjects were trained and tested across multiple stimulus sets. Each set was composed of novel stimuli. Both Conditions 1 and 2 involved explicit performance-contingent feedback presented at the end of each block of test trials (i.e., delayed feedback). In Condition 1, feedback was accurate (consistent with the experimenter-designated relations) following exposure to the initial stimulus sets. When subjects' responding reached a predefined mastery criterion, the feedback then switched to inaccurate (not consistent with the experimenter-designated relations) until responding once again reached a predefined criterion. Condition 2 was similar to Condition 1, except that exposure to the initial stimulus sets was followed by inaccurate feedback and once the criterion was reached feedback switched to accurate. Once relational responding emerged and stabilized, response patterns on novel stimulus sets were controlled by the feedback delivered for previous stimulus sets. Experiment 2 replicated Experiment 1, except that during Conditions 3 and 4 four comparison stimuli were employed during training and testing. Experiment 3 was similar to Condition 1 of Experiment 1, except that after the mastery criterion was reached for class-consistent responding, feedback alternated from accurate to inaccurate across each successive stimulus set. Experiment 4 involved two types of feedback, one type following tests for mutual entailment and the other type following tests for combinatorial entailment. Results from this experiment demonstrated that mutual and combinatorial entailment may be controlled independently by accurate and inaccurate feedback. Overall, the data support the suggestion, made by relational frame theory, that derived relational responding is a form of generalized operant behavior.