Pigeons’ memory for event duration: Intertrial interval and delay effects

The effects of within-session variations in the intertriai interval (ITI) and delay on pigeons’ memory for event duration were studied in delayed symbolic matching-to-sample tasks. Pigeons were trained to peck one color following a long (8 sec) sample and another color following a short (2 sec) sample. In the first three experiments, the baseline conditions included a 10-sec delay (retention interval) and a 45-sec ITI. During testing, the delay was varied from 0 to 20 sec, and the ITI that preceded the trial was varied from 5 to 90 sec. When the ITI and delay were manipulated separately (Experiments 1 and 2), the pigeons displayed a choose-short tendency when the delay was longer than 10 sec or when the ITI was longer than 45 sec, and a choose-long tendency when either the delay or the ITI was shorter than these baseline values. These effects occurred whether the sample was food access or light. When the ITI and delay were manipulated together, the pigeons showed a large choose-long error tendency when the short delay was tested together with a short ITI, and no systematic error tendency when the short delay was tested together with a longer ITI. A very large choose-short error tendency emerged on trials with a long delay and a long ITI; a reduced choose-short tendency was present when the long delay was presented together with a short ITI. In Experiment 4, the baseline conditions were a 0-sec delay and a 45-sec ITI. In this case variations in the ITI had a smaller and unidirectional effect: the pigeons showed a choose-long error tendency when the ITI was decreased, but no effect of ITI increases. Two hypotheses were proposed and discussed: (1) that pigeons judge sample durations relative to a background time composed of the ITI and delay, and (2) that the delay and ITI effects might arise from a combination of subjective shortening and proactive effects of samples from previous trials.     

Pigeons’ memory for empty time intervals marked by visual or auditory stimuli

In Experiment 1, pigeons were trained to discriminate the duration (2 or 8 sec) of an empty interval separated by two 1325-Hz tone markers by responding to red and green comparison stimuli. During delay testing, a choose-short bias occurred at 1 sec, but a robust choose-long bias occurred at 9 sec. Responding in the absence of tone markers indicated that the pigeons were attending to the markers and not simply timing the total trial duration. The birds were then trained to match short (2-sec) or long (8-sec) empty intervals marked by light to blue/yellow comparisons. For both visual and auditory markers, delay testing produced a choose-short bias at 1 sec and a choose-long bias at 9 sec. In Experiment 2, the pigeons were shifted from a fixed to variable intertrial intervals (ITI) within sessions. On trials with tone markers, the duration of both the empty interval and the preceding ITI affected choice responding. On trials with light markers, only the duration of the empty interval influenced choice responding. Subsequent delay testing in the context of variable ITIs replicated the memory biases previously obtained. In Experiment 3, performance was assessed at various delay intervals on trials in which either the first or the second marker was omitted. The data from these omission tests indicated that the first marker initiated timing but that the second marker sometimes initiated the timing of a new interval. Explanations of these effects in terms of the internal clock model of timing are discussed, and a simple quantitative model of the delay interval data is tested.     

A comparison of the short-term memory performances of pigeons and jackdaws

Two experiments employed a delayed conditional discrimination procedure in which half the trials began with the presentation of food and half with no food; following a retention interval, subjects were presented with a choice between red and green keys, a response to one of which was reinforced according to whether the trial had started with food or no food. In Experiment 1, after 38 training sessions during which the retention interval was gradually increased, pigeons performed at a moderate level with intervals of 5 to 7.5 sec. A final test produced a steep forgetting function for food trials, but not for no-food trials; performance was unaffected by the duration of the intertriai interval (10 or 40 sec). Experiment 2 used the delayed conditional discrimination procedure to compare short-term memory in jackdaws (Corvus monedulus) with that in pigeons. Although the performance of the jackdaws was below that of the pigeons at the start of training, they showed more rapid learning over long delays, and, in the final test, a shallower forgetting function for food trials than that shown by pigeons. The results suggested superior short-term memory in jackdaws, which may help to explain the better performance of corvids in general when compared with that of pigeons in certain complex learning tasks.     

Reinforcement expectancy and trial spacing effects in delayed matching-to-sample by pigeons

Four experiments assessed the role of reinforcement expectancies in the trial spacing effect obtained in delayed matching-to-sample by pigeons. In Experiment 1, a differential outcome (DO) group received reinforcement with a probability of 1.0 for correct comparison responses following one sample stimulus and a probability of 0.2 for correct comparison responses following the other sample stimulus. The nondifferential outcome (NDO) group received reinforcement with a probability of 0.6 for correct responses to either stimulus. While matching accuracy was higher for the DO group than for the NDO group, both groups showed an equivalent decline in accuracy as the intertriai interval (ITI) duration was decreased. However, within the DO group, ITI duration affected performance on low-probability-of-reinforcement trials but not on high-probability-of-reinforcement trials. In Experiment 2, delay interval (DI) duration was 5, 10, or 15 sec and accuracy was higher for the DO group than for the NDO group at all DI durations. In addition, accuracy decreased similarly on high- and low-probability-of-reinforcement trials for the DO group as DI was increased. In Experiment 3, all birds were studied under DO conditions and ITI duration was manipulated along with DI duration. At the short DI duration, decreasing ITI duration had a detrimental effect on low-probability-of-reinforcement trials but no effect on high-probability-of-reinforcement trials. At the long DI duration, decreasing ITI duration had detrimental effects on both types of trials. In Experiment 4, unsignaled ITI reinforcers disrupted accuracy when the DI was long and when the ITI was short. The applicability of scalar expectancy theory to these data is discussed.     

Event-duration discrimination by pigeons: The choose-short effect may result from retention-test novelty

Pigeons trained on a conditional event-duration discrimination typically “choose short” when retention intervals are inserted between samples and comparisons. In two experiments, we tested the hypothesis that this effect results from ambiguity produced by the similarity of the novel retention intervals and the familiar intertrial interval by training pigeons with retention intervals from the outset and, for one group, in addition, making retention intervals distinctive from the intertrial intervals. In Experiment 1, when the retention intervals (0–4 sec) were not distinctive from the intertrial intervals, the pigeons did not show a clear choose-short effect even when extended retention intervals (8 sec) were introduced. When the retention intervals were distinctive, the pigeons showed a choose-long effect (they appeared to time through the retention interval), but it was relatively weak until the retention intervals were extended to 8 sec. In Experiment 2, when pigeons were discouraged from timing through the retention intervals by making the intertrial intervals and retention intervals salient distinct events and using long (up to 16-sec) retention intervals in training, parallel retention functions were found. It appears that when ambiguity is removed, forgetting by pigeons does not occur by the process of subjective shortening. These experiments suggest that the accurate interpretation of results of animal memory research using differential-duration samples must consider the novelty of the retention intervals on test trials as well as their similarity to other trial events.     

Unraveling sources of stimulus control in a temporal discrimination task

In temporal discriminations tasks, more than one stimulus may function as a time marker. We studied two of them in a matching-to-sample task, the sample keylight and the houselight that signaled the intertrial interval (ITI). One group of pigeons learned a symmetrical matching-to-sample task with two samples (2 s or 18 s of a center keylight) and two comparisons (red and green side keys), whereas another group of pigeons learned an asymmetrical matching-to-sample task with three samples (2 s, 6 s, and 18 s) and two comparisons (red and green). In the asymmetrical task, 6-s and 18-s samples shared the same comparison. In a subsequent retention test, both groups showed a preference for the comparison associated with the longer samples, a result consistent with the hypothesis that pigeons based their choices on the duration elapsed since the offset of the houselight (i.e., sample duration + retention interval). Results from two no-sample tests further corroborated the importance of the ITI illumination as a time marker: When the ITI was illuminated, the proportion of choices correlated positively with the retention interval; when the ITI was darkened, choices fell to random levels. However, the absolute value of choice proportions suggested that the sample stimulus was also a time marker. How multiple stimuli acquire control over behavior and how they combine remains to be worked out.     

Pigeons’ spatial memory: V. Proactive interference in the delayed matching of key location paradigm occurs only under restricted conditions

In the delayed matching of key location procedure, pigeons must remember the location of the sample key in order to choose correctly between two comparison keys. The deleterious effect of short intertrial intervals on key location matching found in previous studies suggested that pigeons’ short-term spatial memory is affected by proactive interference. However, because a reward expectancy mechanism may account for the intertriai interval effect, additional research aimed at demonstrating proactive interference was warranted. In Experiment 1, matching accuracy did not decline from early to late trials within a session, a finding inconsistent with a proactive interference effect. In Experiment 2, evidence suggestive of proactive interference was found: Matching was more accurate when the locations that served as distractors and as samples were chosen from different sets. However, this effect could have been due to differences in task difficulty, and the results of the two subsequent experiments provided no evidence of proactive interference. In Experiment 3, the distractor on Trialn was either the location that had served as the sample on Trialn ? 1 or one that had been a sample on earlier trials. Matching accuracy was not inferior on the former type of trial. In Experiment 4, the stimuli that served as samples and distractors were taken from sets containing 2, 3, 5, or 9 locations. Matching accuracy was no worse, actually slightly better, with smaller memory set sizes. Overall, these findings suggested that pigeons’ memory for spatial location may be immune to proactive interference. However, when, in Experiment 5, an intratrial manipulation was used, clear evidence of proactive interference was found: Matching accuracy was considerably lower when the sample was preceded by the distractor for that trial than when it was preceded by the sample or by nothing. Possible reasons why interference was produced by intratrial but not intertrial manipulations are discussed, as are implications of these data for models of pigeons’ short-term spatial memory.     

Pigeons’ memory for event duration: Differences between visual and auditory signals

In Experiment 1, pigeons were trained to discriminate short (2 sec) and long (8 sec) durations of tone by responding to red and green comparison stimuli. During delay testing, a systematic response bias to the comparison stimulus correct for the long duration occurred. Tests of responding without the tone reduced accuracy on long-sample trials but not on short-sample trials suggesting that the pigeons were attending to the tone and not simply timing the total trial duration. The pigeons were then trained to match short (2 sec) and long (8 sec) durations of light to blue/yellow comparisons. During delay testing, “choose-long errors” occurred following tone durations, but “choose-short errors” occurred following light durations. In Experiment 2, accuracy was assessed on test trials in which the tone and the light signals were simultaneously presented for the same duration or for different durations. Pigeons responded accurately to durations of light, but were unable to accurately respond to durations of tone simultaneously presented with the light. The data from Experiment 1 suggest that there are important differences between light and tone signals with respect to the events that control the termination of timing. The data from Experiment 2 indicate that pigeons cannot simultaneously time visual and auditory signals independently and without interference. Consequently, they are inconsistent with the idea that there is a single internal clock that times both tone and light durations.     

Directed forgetting in monkeys

Based on several recent demonstrations of a directed forgetting effect in pigeons, three experiments were carried out in an attempt to demonstrate directed forgetting in three squirrel monkeys. During initial training with a delayed matching-to-sample procedure, retention tests were always given for sample stimuli followed by remember cues (R-cues) and were always omitted for sample stimuli followed by forget cues (F-cues). Retention of F-cued items was tested on probe trials after initial training. The first two experiments examined the effects of R- and F-cues on memory for slide-projected pictures, with different pictures used on each trial of a session. In Experiment 1, a complex design was used in which one or two sample pictures were presented on each trial; when two pictures were presented, both could be R-cued or F-cued, or one could be R-cued and the other F-cued. A simpler design was used in Experiment 2, with only single pictures presented as sample stimuli and half the trials within a session R-cued and the other half F-cued. In both of these experiments, no differential retention of R- and F-cued stimuli was found, even at a retention interval as long as 16 sec. In Experiment 3, a series of studies was performed to test for directed forgetting when only two sample stimuli were used repeatedly throughout training and testing. With two pictures as sample stimuli, clear evidence of directed forgetting was found in Experiment 3b. It is suggested that the directed forgetting effect may arise only when a small set of sample stimuli is used.     

Effects of sample duration and spaced repetition upon delayed matching-to-sample in monkeys (Macaca arctoides andSaimiri sciureus)

Delayed matching-to-sample was used to study the effects of sample presentation time and spaced repetition upon delayed matching accuracy in one stumptail monkey and three squirrel monkeys. It was found in Experiment 1 that presenting the sample stimulus for 0.5 sec led to lower matching accuracy than was the case with longer presentation times of 2.5, 5.0, and 10.0 sec. Experiments 2 and 3 investigated the effects of temporally spacing the presentations of the sample stimulus. It was found that spaced repetition led to a deterioration of performance relative to massed repetition. These results are similar to the findings of experiments with pigeons and are contradictory to several previous experiments with monkeys or apes which found no effect of presentation time and a facilitative effect of spaced repetition. It is suggested that the use of monkeys inexperienced in short sample duration matching and tested in operant chambers using a limited set of noncomplex stimuli may be responsible for the discrepancies between these results and those of other experiments with primates.     

Pigeons’ short-term memory for temporal and visual stimuli in delayed matching-to-sample

In the present experiment, we compared directly pigeons’ short-term memory of temporal and visual stimuli in a delayed matching-to-sample task. The sample stimuli consisted of red and green lights presented for 5 and 30 sec, followed by a retention interval and blue and yellow comparisons. For subjects in the visual group, duration was irrelevant and the color of the sample was the conditional cue. For animals in the temporal group, color was irrelevant and duration of the sample was the conditional stimulus. The results showed that acquisition of the matching task was faster and accuracy was higher in the visual than in the temporal group. More importantly, memory of either sample generally declined at a similar rate when the duration of the retention interval was increased and when the intertrial interval was reduced. Taken together, the results indicate that with 1–8-sec retention intervals, short-term memory for temporal stimuli is similar to that found with color-visual samples. The findings are discussed in terms of retrospective and prospective processing.     

Absence of immediate transfer of training of duration symbolic-matching-to-sample in pigeons

Two experiments examined the performance of pigeons on symbolic-matching-to sample in which the relevant sample dimension consisted of duration. Each pigeon was trained on two problems that had the same two sample durations, 2 and 10 sec, but were different with respect to other physical properties of the samples. Durations of light and tone were used in Experiment 1; durations of two different color-location compounds were used in Experiment 2. In each experiment, a unique choice stimulus was associated with each of the four possible combinations of duration and signal type. Test sessions contained probe trials in which the choice stimuli were these appropriate for a long and a short duration of the signal type opposite to that actually presented. Pigeons in both experiments displayed asymmetrical performance deficits. Accuracy on long durations dropped to chance or below, whereas accuracy on short durations remained high. This pattern is similar to the choose-short effect that is obtained when animals are tested with long retention intervals. The implications of these results for duration memory, coding, and transfer of training are discussed.     

Testing the translational-symmetry hypothesis of abstract-concept learning in pigeons

The translational-symmetry hypothesis of abstract-concept learning was tested in a same/different (S/D) task with pairs of pictures. The translational-symmetry hypothesis proposes that subjects discriminate same trials by the simultaneous repetition of features in the two pictures (and different trials by the lack of feature repetition). Pigeons that had learned a simultaneous S/D task were tested with delays between the two pictures to remove emergent perceptual cues. In Experiment 1, we tested delays of 0 and 1 sec. The results did not show the accuracy decrease expected according to the translational-symmetry hypothesis. In Experiment 2, we expanded the delays to 2 and 6 sec. Even at the longest delay, there was no evidence of the precipitous performance decline or default strategy that would be predicted by translational symmetry. The results provide evidence against translational symmetry (or other perceptually emergent features) that might control these pigeons’ performance in our two-item S/D task.     

The effect of scopolamine on reference and working memory in pigeons

A comparison of the effects of scopolamine hydrobromide on working memory and reference memory in White Carneaux pigeons was undertaken by means of a modified delayed matching-to-sample procedure. Performance on working-memory trials was disrupted by decreases in sample duration and intertriai interval and by increases in delay interval. Performance on reference memory trials was not disrupted by any of these parametric manipulations. In Experiment 1, the pigeons received injections of scopolamine hydrobromide (0.01, 0.05, or 0.1 mg/kg), scopolamine methyl bromide (0.1 mg/kg), or saline prior to test sessions. In Experiment 2, the pigeons received injections of scopolamine hydrobromide (0.01 or 0.03 mg/kg), scopolamine methyl bromide (0.03 mg/kg), or saline. In both experiments, scopolamine hydrobromide had greater disruptive effects on working-memory trials than on reference-memory trials. The centrally active form of scopolamine disrupted working-memory trial accuracy more than the peripherally active form. However, no drug dose × delay interval interaction was obtained. Thus, the interference on working-memory-trial accuracy produced by central cholinergic blockade would not appear to be due to alterations in the active maintenance of information during the delay interval.     

Memory for number of light flashes in the pigeon

Two groups of pigeons were trained to perform symbolic delayed matching-to-sample at a 0-sec delay with sample stimuli that consisted of sequences of light flashes. The sequences varied in number but not time for one group (number group) and in time but not number for the other group (time group). When retention was tested at delays up to 10 sec in Experiment 1, a choose-small effect was found in the number group, and a choose-long effect was found in the time group. Transfer tests between number and time samples in Experiment 2 supported the hypothesis that pigeons were discriminating between the number of light flashes at the end of sample sequences in Experiment 1. It was concluded that pigeons in both the number and the time groups were discriminating between number of flashes and that the apparent choose-long effect was actually a choose-small effect. The implications of these findings for the mode-control model of counting and timing (Meck & Church, 1983) were discussed.     

Delayed matching-to-successive-samples in pigeons: Short-term memory for item and order information

Short-term memory for order information in pigeons was explored by using a delayed matching-to-successive-samples task (DMTSS). Experiment 1 indicated that pigeons can accurately report the order of two successively presented samples. Experiments 2, 3 and 4 specifically addressed the representation of order information in short-term memory. Experiment 2 showed that when the duration of the first sample (S1) was very long, or when the duration of the second sample (S2) was very short, order errors increased relative to baseline (S1 and S2 of equal duration), suggesting that memory strength plays an important role in the discrimination of order. The possibility that strength information is necessary for accurate DMTSS performance was tested in Experiments 3 and 4. Pigeons continued to match accurately when memory strength and order were uncorrelated.     

Flexible coding of temporal information by pigeons: Event durations as remember and forget cues for temporal samples

The ability of pigeons to use event durations as remember (R) and forget (F) cues for temporal samples was examined. Pigeons were required to indicate whether a houselight sample stimulus was short (2 sec) or long (6 sec) by pecking a red or a green comparison stimulus. After training with a constant 10-sec delay interval, temporal cues (illumination of the center key) were presented 2 sec after the offset of the temporal samples. For one group, a short (2-sec) temporal cue served as the R cue and a long (6-3ec) temporal cue served as the F cue. This was reversed for a second group of birds. During training, comparison stimuli were always presented following the temporal R cue, but never following the temporal F cue. Tests for the effectiveness of the temporal R and F cues showed that F cues were equally effective in reducing matching accuracy in both groups of birds. It was concluded that pigeons used the duration of the cue to determine whether or not to rehearse the memory code for the temporal sample.     

Sample duration and type of stimuli in delayed matching-to-sample in rhesus monkeys

Two experiments were performed to determine the effect of sample duration (0.1, 2, and 4 sec), delay interval (.03, 4, 8, 16, and 32 sec), and type of stimulus (color and shape) on the matching performance of rhesus monkeys. In Experiment 1, the 15 possible delay-duration combinations were randomly presented in blocks of 15 trials. In Experiment 2, each duration was held constant and the five delays randomly presented. Then each delay interval was held constant with the three durations randomly varied. Matching performance increased as sample duration increased (ps < .01 and .005), while length of delay did not significantly affect performance. The type of stimuli paired in the matching test significantly affected performance (ps < .05 and .10) with the shape/shape choices leading to the poorest performance. Stimulus discriminability and amount of training with brief sample durations were implicated as significant determinants of matching performance.     

More evidence of robust spatial associative memory in the pigeon,Columba livia

Willson and Wilkie (1993) developed a novel procedure for assessing pigeons’ memory for the spatial location of food. Only one of four locations (consisting of an illuminated pecking key and grain feeder) provided food each day. Over days, different locations provided food. The pigeons’ tendency to revisit the location that was profitable on the previous day demonstrated memory for food-spatiallocation associations over a period of 24 h, retention longer than previously reported for this species. This basic finding was replicated and extended in three experiments. Experiment 1 demonstrated that location-food discriminations were also remembered well when established with successive rather than concurrent procedures. Experiment 2 demonstrated that pigeons can remember two location-food associations over 24 h. Experiment 3 showed that the discrimination training inherent in this paradigm is important for retention; retention was impaired when only the rewarded location was presented. Overall, this research suggests that cross-species differences in spatial memory performance may be due to quantitative rather than qualitative differences in the memory system underlying performance.     

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.