Memory for sequences of stimuli and responses

Two experiments sought to determine if pigeons could discriminate and remember recent sequences of stimuli and responses. A variant of Konorski's short-term memory procedure involving successive presentation of sample and test stimuli was used. The samples were stimulus-response pairs of the form, (S-R)₁–(S-R)₂. Differential test responding disclosed memory of the two-item samples, with birds showing earlier and greater control by the second item than the first (Experiment 1). When the retention interval separating the second item of the sample sequence from the test stimulus was lenghtened from .5 to 2.0 or 4.0 sec, a systematic loss of stimulus control resulted; however, when varied over the same temporal range, the interval between the two items of the sample sequence had a much smaller effect, or none at all (Experiment 2). These results support an account of response-sequence differentiation that stresses short-term memory of organized behavior patterns.     

Memory for two stimulus-response items in pigeons

Six pigeons served in a discrete-trial experiment on short-term memory. Combinations of three key positions and two ordinal positions, yielding six possible sequences of stimulus-response pairs, were used as lists of items. A retention interval separated list presentation from the test phase in which two (for Group 1) or three (for Group 2) keys were illuminated with either red or green light. A reinforcer was delivered if a subject pecked the key of the first item on a red trial and the key of the second item on a green trial. When the retention interval was lengthened from one to five or nine seconds, a systematic loss of stimulus control resulted. Lengthening the interval between items from one to eight seconds had a much smaller effect for the birds in Group 1, whereas a systematic loss of stimulus control was found in red trials for the birds in Group 2. The functional relations between choice accuracy and delay provided an empirical basis for analysis of what relations among temporal events can become discriminative stimuli.     

Short-term remembering of discriminative stimuli in pigeons.

Pigeons learned to peck the left or right of two white keys depending on whether a red or a green stimulus was displayed on a third key. The opportunity to peck the white keys was then dealyed for zero to six seconds after the red or green (to-be-remembered) stimulus. On half the trials, the feeder operated during the delay to interrupt behavior that might mediate discriminated responding. No events were scheduled on the remaining trials. In a later condition, the pigeons had the opportunity to peck the white keys during the delay. In general, accuracy decreased as delay increased in all conditions, but performance was least accurate following feeder operations and most accurate when pecking was allowed during the delay. The procedures may be analogous to varying the opportunity for rehearsal in studies of human short-term memory.     

Delay-interval illumination changes interfere with pigeon short-term memory.

Pigeons acquired a successive depayed matching-to-sample task at delay intervals ranging from 2.5 to 7 seconds. Test sessions were conducted during which delay-interval illumination conditions were changed from those illumination conditions that prevailed during the baselines. Compared to baseline delayed matching performance, changing delay-interval illumination disrupted matching. This disruption occurred whether the change in delay-interval illumination represented an increase or a decrease, relative to the baseline, and whether there was or was not a change in illumination during the test session. It was concluded that illumination per se introduced during delay intervals of delayed matching tasks does not interfere with pigeon short-term memory. Rather, a change in delay-interval illumination, relative to the baseline, appears to retroactively interfere in pigeon short-term memory.     

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.     

Conditioning of two-response patterns of key pecking in pigeons

On discrete trials, two response keys were made available to hungry pigeons and food reinforcement depended on the order in which the required two key pecks occurred. In different phases, only one of the four possible two-peck sequences (left-left, left-right, right-left, and right-right) produced food reinforcement. In each case, the pigeons learned to perform the correct two-peck sequence more often than the incorrect sequences. Furthermore, the course of differentiation mastery indicated that both reinforcement history and response-reinforcer contiguity influenced performance. These results reveal that response patterns comprising two instances of the same response left-left and right-right) or instances of two different responses (left-right and right-left) may function as operants, thereby extending the generality of conditioning principles from discrete responses to structured sequences of behavior. These and other results are discussed in terms of contiguity-based and memory-based models of learning.     

Delayed choice responding by pigeons when the correct response is not predictable from the sample stimulus.

Food-deprived pigeons were presented with a row of four response keys situated above a grain hopper aperture. At the start of a trial, three of four keys were randomly selected and illuminated white for six seconds. After a variable blackout period, one of the three previously white keys and the previously dark key were illuminated green, and the remaining white keys were reilluminated as before. A response to the green key that was previously white was reinforced with three-second access to gain, a response to any other key resulted in a three-second blackout and the start of a new trial. Five of six subjects responded to the correct green key more often than chance at an interstimulus interval of 1.5 seconds, and they displayed maximal performance at different intertrial interval values ranging from 15 to 60 seconds. Choice accuracy decreased for all but one subject as the interstimulus interval was increased. For the range of interstimulus interval durations employed, decrements in choice accuracy were qualitatively similar to, but lower than those typically obtained from, delayed-matching-to-sample or delayed-pair comparison procedures.     

Some factors that influence the acquisition of complex, stereotyped, response sequences in pigeons

Two pigeons were required to peck six to nine illuminated response keys. A response on any one of the keys darkened that key. When each key had been darkened, a reinforcer was delivered. No specific sequence of key pecking was ever required. The keys were presented in various matrices: three by two, three by three, horizontal rows, and vertical columns. The keys either presented the same stimulus, white light; or each key presented a different stimulus, a color or form. The results indicated that although there were 720 to 362,880 different sequences that would produce reinforcement, each bird developed a particular, stereotyped sequence that dominated its behavior. Variability among the birds across phases yielded less than 60 sequences, .0001 to 6 percent of the possible sequences. The data suggest that a reinforcement contingency that includes “free choice” of response sequence will produce stereotypical response sequences that function as complex “units” of behavior.     

Context specificity of operant discrimination performance in pigeons

In an experiment designed to investigate odors as potential retrieval cues in pigeons' memory (i.e., as conditional stimuli), 8 pigeons first learned to peck a red keylight (S+, reinforced) and not a blue one (S−, extinguished) in the presence of either a eucalyptus oil or isoamyl acetate odor. They were repeatedly switched between two chambers with the same odor to habituate any reaction to the switching that would be required for eventual testing for conditional control by the odors. Next, the birds learned the reversal (blue S+, red S−) in the presence of the alternative odor in one of these chambers. When the birds were then switched to the alternative chamber for additional training, although the odor was still appropriate to the reversal problem, behavior appropriate to the original training condition recurred. Testing indicated that reversal performance was specific to the one chamber in which it had been trained.     

Short-term memory in the pigeon: delayed-pair-comparison procedures and some results.

A discrete-trials, delayed-pair-comparison procedure was developed to study visual short-term memory for tilted lines. In four experiments, pigeons' responses on left or right keys were reinforced with food depending on whether a comparison stimulus was or was not the same as a standard stimulus presented earlier in the same trial. In Experimental I, recall was an increasing function of the exposure time of the to-be-remembered stimulus and was a decreasing function of the retention interval. In Experiment II, retroactive interference was investigated: recall was poorer after a retention interval during which was presented either a tilted line or contextual stimuli in the form of the illuminated experimental chamber. In Experiment III, a subject was required to engage, throughout the retention interval, in one or the other of two different behaviors, depending on which of two stimuli a subject was to remember. This mnemonic strategy vastly improved recall after 15- and 20-second retention intervals. In Experiment IV, the opposite end of the performance continuum was studied: by combining the effects of a larger stimulus set and the effects of what presumably was an increased memory load, performance was reduced to approximately chance levels after retention intervals shorter than 1 second.     

The development of emergent differential sample behavior in pigeons

Three experiments attempted to replicate Manabe, Kawashima, and Staddon's (1995) finding of emergent differential sample behavior in budgerigars that has been interpreted as evidence of functional equivalence class formation. In Experiments 1 and 2, pigeons initially learned two-sample/ two-alternative matching to sample in which comparison presentation was contingent on pecking one sample on a differential-reinforcement-of-low-rate (DRL) schedule and the other on a fixed-ratio (FR) schedule. Later, two new samples were added to the task. Comparison presentation on these trials occurred after the first sample peck following a predetermined interval (Experiment 1) or after completion of either the DRL or FR requirement, whichever occurred first (Experiment 2). Experiment 1 found no evidence for emergent spaced versus rapid responding to the new samples as they established conditional control over the familiar choices. By contrast, differential responding did emerge for some pigeons in Experiment 2, with responding to each new sample coinciding with the pattern explicitly conditioned to the original sample occasioning the same comparison choice. This emergent effect, however, disappeared for most pigeons with continued training. Experiment 3 systematically replicated Experiment 2 using differential peck location as the sample behavior. Differential location pecking emerged to the new samples for most pigeons and remained intact throughout training. Our findings demonstrate a viable pigeon analogue to the budgerigar emergent calling paradigm and are discussed in terms of equivalence- and non-equivalence-based processes.     

Quantification of the effects of chlorpromazine on performance under delayed matching to sample in pigeons.

The effects of four doses of chlorpromazine (dose range 0.5 to 12.5 mg/kg) on performance under a delayed matching-to-sample procedure in pigeons was investigated, using the exponential model of memory (White, 1985). Performance was measured using a bias-free measure of discriminability, log d (Davison & Tustin, 1978), and negative exponential functions were fitted to individual-subject and group data at each dose level. A decrease in matching accuracy was found to be caused by an increase in the rate of forgetting, b, and a decrease in the initial discriminability, log d0. Changes in rate of forgetting and discriminability occurred at doses that had no statistically significant effect on response latency. The exponential model of memory accounted well for the data and provided a useful way of quantifying the effects of chlorpromazine on the processes involved in delayed matching-to-sample performance.     

Factors affecting conditional discrimination learning by pigeons

In Experiment 1 (within subjects) and Experiment 2 (between subjects) it was shown that the sequential training of pigeons on a color discrimination and then on its reversal, each in a different floor-tilt/texture context, failed to produce conditional control of discriminative performance by those contexts. Daily alternation between the two problems (with correlated contexts) was successful, however. In each of these experiments conditional control was better reflected in generalization test performance in extinction than during sessions of training with reinforcement.     

Hill-climbing by pigeons

Pigeons were exposed to two types of concurrent operant-reinforcement schedules in order to determine what choice rules determine behavior on these schedules. In the first set of experiments, concurrent variable-interval, variable-interval schedules, key-peck responses to either of two alternative schedules produced food reinforcement after a random time interval. The frequency of food-reinforcement availability for the two schedules was varied over different ranges for different birds. In the second series of experiments, concurrent variable-ratio, variable-interval schedules, key-peck responses to one schedule produced food reinforcement after a random time interval, whereas food reinforcement occurred for an alternative schedule only after a random number of responses. Results from both experiments showed that pigeons consistently follow a behavioral strategy in which the alternative schedule chosen at any time is the one which offers the highest momentary reinforcement probability (momentary maximizing). The quality of momentary maximizing was somewhat higher and more consistent when both alternative reinforcement schedules were time-based than when one schedule was time-based and the alternative response-count based. Previous attempts to provide evidence for the existence of momentary maximizing were shown to be based upon faulty assumptions about the behavior implied by momentary maximizing and resultant inappropriate measures of behavior.     

Effects of reinforcement amount on attack induced under a fixed-interval schedule in pigeons

Key pecking by pigeons was maintained on a chained fixed-interval 4-min (12-min for 1 subject) fixed-ratio 1 schedule of food presentation. Attacks toward a restrained and protected conspecific were recorded. In the first experiment, the amount of food presented per interval was manipulated across phases by varying the number of fixed ratios required in the terminal link of the chain. Measures of attack for all pigeons during the fixed-interval component increased monotonically as a function of food amount. In the second experiment, two different food amounts alternated within each experimental session under a multiple schedule. For both pigeons in this experiment, measures of attack were higher during the component that delivered the larger food amount per interval. The differences in levels of attack induced by the two food amounts in Experiment 2, however, were not as great as in Experiment 1; apparently this was because attack during the first interval of each component was controlled in part (P-5626) or entirely (P-7848) by the reinforcement amount delivered at the end of the previous component. Attack was also a function of the location of the interfood interval within the session. For both pigeons, attack tended to decrease throughout the session. The results of both experiments suggest that attack is an increasing function of reinforcement amount under fixed-interval schedules, but that this function may be influenced by the manner in which reinforcement amount is manipulated, by the duration of the interfood interval, and by the location of the interfood interval within the experimental session. In general, these results are compatible with theories of induced attack and other schedule-induced behavior that emphasize aversive after-effects of reinforcement presentation.     

Transfer of oddity-from-sample performance in pigeons

Four pigeons were trained on a modified three-key oddity-from-sample task in which an observing response to the sample (center-key) stimulus lighted a single comparison (side-key) stimulus. If the comparison stimulus was different from the sample stimulus, a single peck to the lighted comparison was reinforced. If the comparison and sample stimuli were identical, the pigeons had to refrain from pecking the comparison for 4.6 seconds to terminate the matching comparison and to produce immediately a nonmatching comparison on the remaining side key. Each peck to the matching comparison reset the 4.6-second delay interval. Three hues were used during acquisition. During tests for transfer of the oddity performance, two novel hues were substituted either individually or together for one or two of the original training hues. For three birds, latencies to novel nonmatching hues were identical to baseline nonmatching latencies. Latencies to novel matching hues were shorter than baseline matching latencies but were consistently longer than novel nonmatching latencies. These transfer data demonstrate that the pigeons learned the oddity concept.     

Stimulus stringing by pigeons

Pigeons were trained to peck one, two, three, and then four colors in a predetermined sequence from a five-key array where, over trials, each color appeared equally often in each position of the array. Incorrect pecks resulted in a buzzer and trial termination, with the same array presented for the next trial. Correct pecks produced feedback and correct strings could produce food. All subjects performed at a high level of accuracy with no difference at asymptote between a continuous and a mixed spectral sequence as the required order. Transfer to a new set of arrays had little effect on accuracy. Errors forward in the sequence had the highest probability, followed by repeat errors, backward errors, and dark-key errors. Some arrays had a higher level of accuracy than others but a corresponding systematic variable could not be identified.     

Nonstable concurrent choice in pigeons

Six pigeons were trained on concurrent variable-interval schedules in which the arranged reinforcer ratios changed from session to session according to a 31-step pseudorandom binary sequence. This procedure allows a quantitative analysis of the degree to which performance in an experimental session is affected by conditions in previous sessions. Two experiments were carried out. In each, the size of the reinforcer ratios arranged between the two concurrent schedules was varied between 31-step conditions. In Experiment 1, the concurrent schedules were arranged independently, and in Experiment 2 they were arranged nonindependently. An extended form of the generalized matching law described the relative contribution of past and present events to present-session behavior. Total performance in sessions was mostly determined by the reinforcer ratio in that session and partially by reinforcers that had been obtained in previous sessions. However, the initial exposure to the random sequence produced a lower sensitivity to current-session reinforcers but no difference in overall sensitivity to reinforcement. There was no evidence that the size of the reinforcer ratios available on the concurrent schedules affected either overall sensitivity to reinforcement or the sensitivity to reinforcement in the current session. There was also no evidence of any different performance between independent and nonindependent scheduling. Because of these invariances, this experiment validates the use of the pseudorandom sequence for the fast determination of sensitivity to reinforcement.     

Most directed forgetting in pigeons can be attributed to the absence of reinforcement on forget trials during training or to other procedural artifacts.

In research on directed forgetting in pigeons using delayed matching procedures, remember cues, presented in the delay interval between sample and comparisons, have been followed by comparisons (i.e., a memory test), whereas forget cues have been followed by one of a number of different sample-independent events. The source of directed forgetting in delayed matching to sample in pigeons was examined in a 2 x 2 design by independently manipulating whether or not forget-cue trials in training ended with reinforcement and whether or not forget-cue trials in training included a simultaneous discrimination (involving stimuli other than those used in the matching task). Results were consistent with the hypothesis that reinforced responding following forget cues is sufficient to eliminate performance deficits on forget-cue probe trials. Only when reinforcement was omitted on forget-cue trials in training (whether a discrimination was required or not) was there a decrement in accuracy on forget-cue probe trials. When reinforcement is present, however, the pattern of responding established during and following a forget cue in training may also play a role in the directed forgetting effect. These findings support the view that much of the evidence for directed forgetting using matching procedures may result from motivational and behavioral artifacts rather than the loss of memory.     

Stimulus control of delayed matching in pigeons: Directed forgetting

Pigeons were trained in delayed matching-to-sample with two postsample stimuli. A postsample R-cue signaled that a matching choice phase would follow. A postsample F-cue signaled that a matching choice phase would not follow. Previous research found reduced matching accuracy on F-cued probe trials when comparison stimuli were presented in the choice phase. The present four experiments systematically varied the events following an F-cue to determine the conditions under which the F-cue reduces delayed-matching accuracy. When F-cues and R-cues controlled different behavior, matching on probe trials was poor. When both cues controlled the same behavior, matching on probe trials was good. This result is best explained by the theory that comparison stimuli retrieve the sample representation, but only in the behavioral context established by the R-cue. The present research supports the view that response-produced stimuli serve a contextual role in animal short-term memory.