Use of a single-code/default strategy by pigeons to acquire duration sample discriminations

Past evidence that pigeons may adopt a single-code/default strategy to solve duration sample discriminations may be attributable to the similarity between the intertrial interval (ITI) and the retention interval. The present experiments tested whether pigeons would adopt a single-code/default strategy when possible ITI-retention-interval ambiguity was eliminated and sample salience was increased. Previous studies of duration sample discriminations that have purported to show evidence for the use of a single-code/default coding strategy have used durations of 0, 2, and 10 sec (Zentall, Klein, & Singer, 2004). However, the results of Experiment 1 suggest that the use of a 0-sec sample may produce an artifact resulting in inadvertent present/absent sample matching. In Experiment 2, when pigeons were trained with three nonzero duration samples (2, 8, and 32 sec), clear evidence for the use of a single-code/default strategy was found.     

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.     

The role of trial tracking in rats’ working memory

The experiments reported in the present study tested whether decreasing intertrial intervals (ITIs) intensifies the disruptive effects of increasing retention intervals (RIs) in a delayed conditional discrimination by decreasing the animal’s trial tracking accuracy (Cohen & Armstrong, 1996; Cohen & Roberts, 1996). Rats responded on a fixed ratio (FR) 1 or fixed interval (FI) 10-sec reinforcement schedule at a second light or tone stimulus, S2, when the first light or tone stimulus, S1, had signaled an FI 10-sec or FR 1 schedule, respectively. RIs between S1 and S2 were increased from 3 to 24 sec and never exceeded ITIs that were reduced from 24 to 6 sec. For some rats, the trials were separated from each other by extending the lever at S1 and retracting it at the end of S2 (ITI lever-retracted group). For other, control rats, the lever remained extended throughout the session (lever-extended group, Experiment 1) or was extended and retracted with the onset and offset of each stimulus (RI/ITI lever-retracted group, Experiment 2). The rats under all trial conditions learned to delay leverpressing on the FI 10-sec schedule. Latency to begin leverpressing on the FI 10-sec schedule declined as RIs were increased, but this effect was attenuated in the ITI lever-retracted groups in both experiments, as would be predicted by thetrial tracking hypothesis. Decreasing ITIs from 24 to 6 sec intensified the disruptive effects of increasing RIs from 3 to 6 sec in the RI/ITI lever-retracted group (Experiment 2), as would be predicted by the trial tracking hypothesis.     

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.     

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.     

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.     

Analysis of a trial-spacing effect with relatively long intertrial intervals

In three experiments with rat subjects, we examined the effects of trial spacing in appetitive conditioning. Previous research in this preparation suggests that self-generated priming of the conditional stimulus (CS) and/or unconditional stimulus (US) in short-term memory is a cause of the trial-spacing effect that occurs with intertrial intervals (ITIs) of less than 240 sec. Experiment 1 nonetheless showed that a trial-spacing effect still occurs when ITIs are increased beyond 240 sec, and that the effect of ITI over 60–1,920 sec on conditioned responding is best described as a linear function. In Experiment 2, some subjects were removed from the context during the ITIs, preventing extinction of the context. Removal abolished the advantage of the long ITI, suggesting the importance of exposure to the context during the long ITI. Experiment 3 still produced a trial-spacing effect in a within-subjects design that controlled for the level of context conditioning and reinforcement rate in the absence of the CS. Overall, the results are most consistent with the idea that adding time to the ITI above 240 sec facilitates conditioning by extinguishing context-CS associations—and possibly context-US associations—that otherwise interfere with CS-US learning through retrieval-generated priming (see, e.g., Wagner, 1981).     

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.     

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.     

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.     

Pigeons’ memory for sequences of light flashes when gap duration is an unreliable discriminative cue

In Experiment 1, pigeons were trained with a 1-sec dark and a 1-sec houselight-illuminated delay interval to discriminate between sequences of two and four flashes of light (feeder illumination). The sequences could be discriminated on the basis of the number of flashes, the number of gaps, or the duration of the gap between flashes. A choose-few bias was obtained at extended dark delays, but not at extended illuminated delays. Pigeons appeared to confuse long dark delays with the longer gap between flashes on few-sample trials. In Experiment 2, additional sample sequences were included that made gap duration an unreliable cue for discriminating between the few and many samples. A significant choose-many bias was obtained at extended dark delay intervals, but no biased forgetting was found at extended illuminated delays. The pigeons appeared to discriminate light flash sequences by relying on multiple temporal features of a sequence rather than using an event switch to count flashes. The biased-forgetting effects observed appear to be due to instructional ambiguity that results from the similarity of the delay interval to features of the flash sequences. nt]mis|This research was supported by Grant OGPOOD6378 from the Natural Sciences and Engineering Research Council of Canada to A.S.     

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.     

Memory for empty time intervals in pigeons

Pigeons were trained to discriminate short (2 sec) and long (8 sec) empty intervals that began each trial. In group consistent, onset of an empty interval was marked by a brief presentation of red keylight, and termination of the interval was marked by a brief presentation of green keylight. In group inconsistent, red and green served equally often as the first and second markers across trials. Testing revealed that, in group consistent, (1) birds were sensitive to the relation between marker color and marker type and (2) presentation of the second marker did not initiate timing a new interval. Testing also revealed a robust choose-long effect at delays longer than the training delay and indifference between the comparisons on no-sample trials. Both of the latter findings differ from those typically obtained when filled intervals are employed. It was concluded that pigeons process filled and empty intervals differently.     

The “disinhibition” of extinguished operant behavior in pigeons: Trial-tempo shifts and novel stimulus effects

After conditioning and extinction of keypecking with 25-see intertrial intervals between key illuminations, an immediate change to 5-sec intertrial intervals reinstated pecking during trials. Brief illuminations of the chamber during intertrial intervals also temporarily restored extinguished keypecking. The same manipulations of trial tempo and chamber illumination usually weakened well established, still reinforced keypecking. No recovery of extinguished behavior occurred when the intertrial interval was shifted upward from 5 sec to 25 sec. These and other behavioral findings were examined in relation to (a) Pavlov’s and Skinner’s research and views on “disinhibition” and “external inhibition” and to (b) analogous phenomena in physiological studies of habituation and dishabituation. The reliable evidence of disinhibition obtained in the present experiments suggests the involvement of an inhibitory process in extinction.     

Short-term retention of “surprising” events following different training conditions

Terry and Wagner (1975) have suggested that short-term retention of information about an event is enhanced if the occurrence of the event is surprising. To investigate this idea, we trained two groups of pigeons in a preparatory-releaser procedure in which half the trials started with the presentation of food (the preparatory event). The preparatory food presensation was signaled by an 8-sec white keylight in the signaled, but not in the unsignaled, group. After a retention interval, varying between 2 and 32 sec, the releaser stimulus (CS_R), a red keylight, was presented for 8 sec in the absence of any reinforcement. The remaining trials were initiated by the presentation of CS_R, and the first peck occurring 8 sec after the onset of CS_R was reinforced by food. The preparatory event controlled responding to CS_R at the short retention interval, with the level of control declining systematically with increasing retention intervals. On probe test trials, the presentation of the preparatory food event was preceded by a stimulus that had previously been paired (CS+) or unpaired with food (CS?). Discriminative responding to CSr was better following CS? than following CS+ in the unsignaled, but not the signaled, group. These results suggest that the enhanced retention following surprising preparatory events reflects a generalization decrement induced by changing the signaling conditions between training and testing.     

Pigeons’ memory for time: Assessment of the role of subjective shortening in the duration-comparison procedure

Pigeons were trained in a duration-comparison procedure to peck one key if the comparison duration (c) was 1 sec shorter than a standard duration (s), and another key if c was 1 sec longer than s. During training, the s-c delay was 1 sec, and the total duration of an s-c pair was not predictive of the correct choice. In Experiment 1, during equal-duration pair test trials, pigeons increasingly responded long (i.e., c τ s) as the s-c delay was lengthened. In Experiment 2, we demonstrated that s affected long responding on equal-duration test trials, even at the 8-sec s-c delay. In Experiment 3, long responding increased as the s-c delay was lengthened, even when stimulus conditions during the s-c delay differed from those during the intertrial interval (ITI). Additional analyses indicated that it was unlikely that the increase in long responding was due to the pigeons’ adding the s-c delay to c and comparing the total against the duration of s. The increase in long responding with an increase in s-c delay is more consistent with subjective shortening of s than with confusion between the s-c delay and the ITI.     

Partial reinforcement in autoshaping with pigeons

The acquisition, maintenance, and extinction of autoshaped responding in pigeons were studied under partial and continuous reinforcement. Five values of probability of reinforcement, ranging from .1 to 1.0, were combined factorially with five values of intertrial interval ranging from 15 to 250 sec for different groups. The number of trials required before autoshaped responding emerged varied inversely with the duration of the intertriai interval and probability of reinforcment, but partial reinforcement did not increase the number of reinforcers before acquisition. During maintained training, partial reinforcement increased the overall rate of responding. A temporal gradient of accelerated responding over the trial duration emerged during maintenance training for partial reinforcement groups, and was evident for all groups in extinction. Partial reinforcement groups responded more than continuous reinforcement groups over an equivalent number of trials in extinction. However, this partial-reinforcment extinction effect disappeared when examined in terms of the omission of “expected” reinforcers.     

Order and duration of stimuli are important determinants of reactivation

Prior cuing treatments intended to alleviate the forgetting of a conditioned avexsion-to an odor were tested with 18-day-old rats. Previous experiments had shown that when such pups were conditioned with the use of a CS?/CS+ procedure, pretest presentation of the CS? or US, but not the CS+, alleviated the forgetting otherwise seen after a 3-h retention interval. In Experiment 1, it was determined that the forgetting was not alleviated if the GS? was either preceded or followed by presentation of the CS+, despite the fact that the CS?/CS+ ordering mimicked that of original conditioning. Experiment 2 was an examination of the balance of extinction and reactivation effects caused by presenting the CS+ for varying durations following the 3-h retention interval. The forgetting over this interval was alleviated if the CS+ was presented for 5 or 15 sec, but not 30 sec. With an increase in duration of exposure from 15 to 30 sec, the consequences of the CS+ as a prior cuing treatment apparently shifted from reactivation to extinction. Experiment 3 was a test of the interaction between the consequences of different lengths of CS+ exposure and the effectiveness of adding CS? to the CS+ as a reactivation treatment. The varied effectiveness of reactivation treatments is discussed interms of a change in stimulus conditions from training to reactivation.     

CS-US temporal relations in blocking

In four trace-conditioning experiments with rats, the influence on the blocking of differences between the blocking cue-unconditioned stimulus (US) and the blocked cue-US trace intervals was explored. Experiment 1 demonstrated blocking despite the blocked cue’s having a shorter trace interval than the blocking cue in both elemental (Phase 1) and compound (Phase 2) training. In Experiment 2, blocking was attenuated when the blocked cue had a longer trace interval than did the blocking cue in both elemental and compound training. In Experiments 3 and 4, the trace intervals of the two cues during compound training were matched (i.e., unlike in Experiments 1 and 2, neither had temporal priority). Blocking was attenuated when the blocking cue trace interval in the elemental phase was shorter (Experiment 3) or longer (Experiment 4) than the compound cue trace during compound training. The findings indicate that subjects encode interstimulus intervals, and they further suggest that cue competition is greatest when the competing cues have the same temporal information as the US.     

Partial reinforcement delayed extinction effect after regularly alternating reward training

When extinction is delayed very long, the superior resistance to extinction of the random schedule group relative to the alternating schedule group disappears (partial reinforcement delayed extinction effect, PRDE). Two experiments assessed the effects of reinforcement/nonreinforcement on Trial 1 on the PRDE. Following extended partial reinforcement acquisition training in a runway, rats received extinction training after a short (1-day) or long (23-day) retention interval. The schedules used in Experiment 1 were: a single-alternation (SA) schedule beginning each day with a rewarded (r) trial, for Group r-SA; an SA schedule beginning with a nonrewarded (n) trial, for Group n-SA; and a random (Rd) schedule, for Group Rd. The schedules and group names used in Experiment 2 were r-SA, Rd, and r-Rd. The results were that (1) rats given r-SA schedules yielded considerable resistance under delayed extinction, (2) those given Rd and r-Rd schedules showed a decline in resistance to extinction over a long retention interval, (3) those given the n-SA schedule showed relatively low resistance at both retention intervals, although retention deficit was not greater than in the case of the Rd schedule, and thus, (4) the PRDE was found in both experiments, although only weakly in Experiment 1. The results indicated that a regularly alternating reward pattern was a more important determinant than was type of reward on Trial 1 for the PRDE. The PRDE due to differential retention deficits among schedules is discussed on the basis of dual-process associative sequential mechanisms and cognitive rule-encoding mechanisms.