Pigeons' memory for number of events: Effects of intertrial interval and delay interval illumination

In Experiment 1, pigeons were trained at a 0-s baseline delay to discriminate sequences of light flashes (illumination of the feeder) that varied in number but not time (2f/4s and 8f/4s). During training, the intertrial interval was illuminated by the houselight for Group Light, but it was dark for Group Dark. Testing conducted with dark delay intervals produced a strong choose-small bias in both groups. All birds then received baseline training with a 5-s dark delay and were subsequently tested at shorter and longer dark delays. A choose-small bias was again observed at delays longer than the training delay, while a choose-large bias occurred at delays shorter than the training delay. Differentiating the ambient chamber illumination during the intertrial interval and the delay interval did not attenuate choose-small or choose-large effects. In Experiment 2, all birds received baseline training with a 5-s illuminated delay and were subsequently tested at shorter and longer illuminated delays. A choose-large bias was observed at delays longer than the training delay, while a choose-small bias occurred at delays shorter than the training delay. In Experiment 3, on intermittent test trials, when the duration of the second flash on small-sample trials was equal to the total flash duration on large-sample trials (i.e., 1600 ms), accuracy fell to approximately chance. These results suggest that pigeons discriminated the sequences of light flashes that varied in number but not in total duration of the sequence by relying on other temporal properties of the sequence rather than by using an event switch to count flashes.     

Performance of pigeons on delayed simple and conditional discriminations under equivalent training procedures

A pair of experiments investigated the short-term memory of pigeons under delayed simple and conditional discriminations. Trial sequences in both discriminations consisted of a color as the sample stimulus, a memory interval, a line orientation as the test stimulus, and a trial outcome, which was either food reinforcement or blackout. Pecking rates during the test stimulus defined discrimination performance. In the simple discrimination, the sample provided the necessary information regarding the subsequent trial outcome. In the conditional discrimination, the sample and test stimuli conjointly provided this information. In Experiment 1, the two procedures were compared with independent groups of pigeons. In Experiment 2, the comparison was made within subjects. The simple discrimination was acquired more quickly and was performed better with a memory requirement. Introduction of long delays disrupted performance even at shorter delays in both discriminations. Postulation of prospective as well as retrospective mediating processes facilitates the interpretation of these results.     

Alan Baron: A pioneer in translational science

Primates take longer to choose between alternatives with smaller differences in value. This effect—a particular instance of the distance effect in symbolic comparisons—has not been replicated in birds. Instead, birds appear to respond independently to each alternative, such that the latency to choose depends primarily on the alternative of highest value. Three experiments tested for the distance effect in pigeons under conditions not previously considered. Experiment 1 presented pigeons with forced‐ and binary free‐choice trials, where each alternative was one of three possible delays to reinforcement (4, 8, and 16 s). Pigeons were exposed to the choice stimuli for different amounts of time and with different sample response requirements prior to the choice response. Experiment 2 added a fourth (0‐s delay) alternative. Experiment 3 substituted the 16‐s delay with a second 4‐s delay. In all experiments, pigeons systematically chose the shortest delay to reinforcement. Latency to choose the 4‐s delay did not vary when choosing against the 8‐s or 16‐s delay, regardless of whether choice stimuli were exposed for the duration of nine pecks (Experiment 1), or whether a 0‐s delay alternative was sometimes present (Experiment 2). Latency to choose the preferred of two identical alternatives (4‐s vs. 4‐s) was shorter than the latency to choose between different alternatives (4‐s vs. 8‐s; Experiment 3); this is the opposite of a distance effect. These results show no evidence of a distance effect in pigeon choice, consistent with the hypothesis that pigeons respond independently to each choice alternative.     

工作记忆的存储时间及目标时距对时间知觉的影响

本研究操作记忆信息与计时开始之间的时间间隔（ISI）和目标时距,探讨工作记忆影响时间判断的灵活性。被试首先记忆一个客体,然后在每个trial的最后判断测试刺激是否与记忆项相同;在延迟阶段,被试完成时间判断任务,即判断相继出现的两个刺激的时距哪个更长（或更短）。时间任务中的一个刺激与记忆内容完全相同,相应的另一个刺激与记忆内容在形状和颜色上都不同。重复条件下,被试忽略第一个刺激,仅完成时间判断任务。结果发现,时间间隔（ISI）短时,记忆匹配条件下的准确率更高,匹配刺激延长了知觉的时间;但随着时间间隔的增加,工作记忆匹配对时间判断的影响降低甚至消失。并且,长或短ISI,记忆任务或重复条件下,目标时距长时,记忆匹配反而缩短了知觉的时间。研究说明工作记忆对时间判断的影响是灵活的,受到注意或工作记忆等高级认知系统的调控。     

The Effect of Nontemporal Information Processing on Time Estimation in Pigeons

Pigeons' ability to time a stimulus while simultaneously engaged in another information-processing task was examined in three experiments using a matching-to-sample procedure. Pigeons were trained to match temporal samples of 2 and 10 s and were tested with durations of 2, 3, 4.5, 6.7, or 10 s while simultaneously processing information from another dimension. Experiment 1 revealed that the psychophysical curve for long judgments taken when pigeons also had to evaluate line orientation was shifted to the right of a control curve taken from trials on which only time had to be judged. In Experiment 2, results from probe duration tests showed that processing the spatial location of a stimulus while attending to duration caused a more general loss of timing ability across the probe durations (shorter durations were judged as longer and longer durations were judged as shorter). In Experiment 3, a distracter light was illuminated on some probe trials to determine how such a perceptual distraction would affect time judgments on the probe trials. Results showed a general loss of timing ability similar to that found in Experiment 2. It is proposed that the data might best be explained by a divided-attention effect rather than by a systematic effect on the timing mechanism.     

Short-term memory and information processing in pigeons

Six pigeons were trained to asymptotic performance on a variable-delay matching-to-sample task in which the samples were sometimes line or color elements and sometimes line-color compounds. On compound-sample trials, the comparison stimuli were sometimes color elements and sometimes line-tilts. Sample type and delay (0, 1.5, and 4.5 sec) were varied within sessions, and sample duration (.4, 1.0, and 3.0 sec) was varied between sessions. Forgetting curves were steeper for line-tilt than for color. As sample duration increased, matching performance improved more for colors than for line-tilts, especially at delays greater than zero. Performance was better with element samples than with compound samples only on the line-tilt dimension at zero delay. Some predictions of a unitary trace growth and decay theory of pigeon short-term memory were not confirmed. A dual-code hypothesis was proposed to account for the data.     

Required pecking alters judgments of the passage of time by pigeons

There is evidence that how humans perceive time is affected by the activity in which they are engaged when they are judging time. In humans, typically, the more demanding the task, the faster time seems to pass. We asked whether a similar effect could be found in pigeons. Pigeons were trained to discriminate between a short-(2-sec) and a long-(10-sec) duration stimulus. Depending on the color of the stimulus (white or blue), the pigeons were required to peck (at least once per second or the trial was aborted) or to refrain from pecking (pecks aborted the trial). Once these tasks had been acquired to a high degree, probe trials involving white and blue stimuli were presented at durations between 2 and 10 sec. On trials in which the pigeons were required to peck, the point of subjective equality (i.e., the point at which pigeons are equally likely to choose the stimulus associated with long stimuli as the stimulus associated with short stimuli) was almost 1 sec longer than on trials in which the pigeons were required to refrain from pecking. In other words, on trials that required pecking, more time passed before the pigeons indicated that the probe duration was at the subjective midpoint between 2 and 10 sec than on trials that did not require pecking. This result suggests that like humans, the pigeons underestimated the passage of time when they were active or when attention to time-related cues had to be shared with attention to satisfying the response rate requirement.     

Short-term memory for event duration: Modality specificity and goal dependency

Time perception is involved in various cognitive functions. This study investigated the characteristics of short-term memory for event duration by examining how the length of the retention period affects inter- and intramodal duration judgment. On each trial, a sample stimulus was followed by a comparison stimulus, after a variable delay period (0.5?C5?s). The sample and comparison stimuli were presented in the visual or auditory modality. The participants determined whether the comparison stimulus was longer or shorter than the sample stimulus. The distortion pattern of subjective duration during the delay period depended on the sensory modality of the comparison stimulus but was not affected by that of the sample stimulus. When the comparison stimulus was visually presented, the retained duration of the sample stimulus was shortened as the delay period increased. Contrarily, when the comparison stimulus was presented in the auditory modality, the delay period had little to no effect on the retained duration. Furthermore, whenever the participants did not know the sensory modality of the comparison stimulus beforehand, the effect of the delay period disappeared. These results suggest that the memory process for event duration is specific to sensory modality and that its performance is determined depending on the sensory modality in which the retained duration will be used subsequently.     

Pigeons group time intervals according to their relative duration

In the present research, we asked whether pigeons tended to judge time intervals not only in terms of their absolute value but also relative to a duration from which they must be discriminated (i.e., longer or shorter). Pigeons were trained on two independent temporal discriminations. In one discrimination, sample durations of 2 and 8 sec were associated with, for example, red and green hue comparisons, respectively, and in the other discrimination, sample durations of 4 and 16 sec were associated with vertical and horizontal line comparisons, respectively. If pigeons are trained on a temporal discrimination and tested with intermediate durations, the subjective midpoint typically occurs close to the geometric mean of the two trained values. The 4- and 8-sec values were selected to be the geometric mean of the two values in the other discrimination. When a 4-sec test sample was presented with the comparisons from the 2- and 8-sec discrimination, the pigeons preferred the comparison associated with the shorter sample. Similarly, when an 8-sec test sample was presented with the comparisons from the 4-and 16-sec discrimination, the pigeons preferred the comparison associated with the longer sample. Thus, a relative grouping effect was found. That is, durations that should have produced indifferent choice were influenced by their relative durations (shorter than or longer than the alternative) during training.     

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.     

Relative judgments affect assessments of stimulus duration

Humans were trained on two independent temporal discriminations, with correct choice dependent on the initial stimulus duration. In Experiment 1, the durations were 1.0 and 4.0 sec, with one set of choice stimuli, and 2.0 and 8.0 sec, with a different set of choice stimuli. The 2.0- and 4.0-sec values were selected to be the geometric mean of the two values in the other discrimination. In Experiment 2, the durations were 2.0 and 5.0 sec for one discrimination and 3.5 and 6.5 sec for the other. The 3.5- and 5.0-sec values were selected to be the arithmetic mean of the two values in the other discrimination. In both experiments, participants showed evidence for relational coding of the duration pairs. That is, the test durations were selected to be at the presumed bisection point (i.e., they should have produced indifferent choice), but instead the shorter test duration from the longer duration pair produced a “short” bias (in both experiments), whereas the longer duration from the shorter duration pair produced a “long” bias (in the second experiment). Results were similar to those from Zentall, Weaver, and Clement (2004) with pigeons.     

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.     

The Effects of Hippocampal and Area Parahippocampalis Lesions in Pigeons: I. Delayed Matching to Sample

Four experiments were conducted to determine the effects of bilateral damage to the hippocampus and area parahippocampalis (Hp-APH) on visual memory in pigeons using the delayed matching-to-sample (DMS) procedure. In Experiment 1, we generated visual retention gradients with delays of 0, 1.5, 3, 6, and 12 sec both preoperatively and postoperatively in three pigeons with considerable preoperative visual DMS experience. Bilateral Hp-APH lesions had no effect whatsoever on visual retention. In Experiment 2, we examined the effects of Hp-APH lesions on both the acquisition of a visual DMS task with a 0-sec delay, and the subsequent retention performance with delays of 0, 3, 6, 12, and 24 sec. There was no difference between unoperated control pigeons and Hp-APH pigeons either in terms of the number of sessions required to learn the visual DMS task or in terms of their subsequent visual retention performance levels. In Experiments 3 and 4, we examined whether Hp-APH pigeons might be more sensitive than control pigeons to the effects of proactive interference (by reducing the duration of the intertrial interval) and retroactive interference (by introducing delay-interval illumination). Although reducing the duration of the intertrial interval and increasing the level of delay-interval illumination both resulted in lower performance levels on the visual DMS task, there was no indication that the Hp-APH pigeons were any more affected by the changes in interference levels than were unoperated control pigeons. These findings support the view that the Hp-APH in pigeons plays little role in the processing and retention of purely visual information.     

Separating The Effects Of Trial-specific And Average Sample-stimulus Duration In Delayed Matching To Sample In Pigeons

Pigeons were studied in two experiments employing delayed matching-to-sample (DMTS) tasks in which the reduction in delay to reinforcement signaled by the onset of the sample stimulus was manipulated by varying sample-stimulus duration. In Experiment 1, the duration of the sample stimulus was either 5 s or 10 s for one sample stimulus and 10 s or 20 s for the other. Subjects matched more frequently when the sample duration was 10 s following the sample associated with the shorter average duration. This finding is analogous to the memory distribution effect found by Honig (1987) in a successive DMTS task that varied retention interval. In Experiment 2, sample duration was either 5 s or 15 s. In Phases 1 and 3 each sample duration was correlated with a particular sample color, and in Phase 2 sample duration and color were uncorrelated. When sample duration was 5 s, subjects matched more frequently when sample duration and color were correlated than when they were uncorrelated. Overall, subjects matched more frequently when sample duration and color were correlated. The data from both experiments support Wixted's (1989) model, which states that one determinant of choice in a DMTS task is the delay-reduction value of the sample stimulus.     

Differences in matching-to-sample performance with element and compound sample stimuli in pigeons

This series of experiments examined zero-delay matching-to-sample performance in pigeons with element and compound sample stimuli. In Experiment 1, compound sample stimuli were consistently followed by compound comparison stimuli and matching accuracy during testing was equivalent to element sample-element comparison trials on the color dimension. In Experiment 2, element comparisons suddenly introduced following compound samples produced a decrement on the line dimension only. Subsequent testing at various sample durations revealed higher matching accuracy following element samples than following compound samples on both the color and line-tilt dimensions. Experiment 3 replicated the results of Experiments 1 and 2 and also demonstrated that the superiority of element over compound matching performance remains constant over the sample durations tested. In Experiment 4 testing at sample durations up to 30 sec produced an overall decrement in matching performance, but again the element vs compound matching difference remained constant. The stimulus-generalization decrement hypothesis provided a better explanation of these results than either the information-overload hypothesis or the rule hypothesis.     

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.     

Visual dominance in the pigeon

In Experiment 1, three pigeons were trained to obtain grain by depressing one foot treadle in the presence of a 746-Hertz tone stimulus and by depressing a second foot treadle in the presence of a red light stimulus. Intertrial stimuli included white light and the absence of tone. The latencies to respond on auditory element trials were as fast, or faster, than on visual element trials, but pigeons always responded on the visual treadle when presented with a compound stimulus composed of the auditory and visual elements. In Experiment 2, pigeons were trained on the auditory-visual discrimination task using as trial stimuli increases in the intensity of auditory or visual intertrial stimuli. Again, pigeons showed visual dominance on subsequent compound stimulus test trials. In Experiment 3, on compound test trials, the onset of the visual stimulus was delayed relative to the onset of the auditory stimulus. Visual treadle responses generally occurred with delay intervals of less than 500 milliseconds, and auditory treadle responses generally occurred with delay intervals of greater than 500 milliseconds. The results are discussed in terms of Posner, Nissen, and Klein's (1976) theory of visual dominance in humans.     

Overshadowing and stimulus duration

In 3 experiments, the authors investigated the effect of stimulus duration on overshadowing. Experiments 1 and 2 examined responding to a target conditioned stimulus (CS1) when it was conditioned in compound with a coterminating overshadowing stimulus (CS2) that was longer, shorter, or of the same duration (the long, short, and matched groups, respectively). Equal overshadowing of conditioning to CS1 was obtained in all 3 conditions relative to a control group conditioned to the light alone. There were, however, differences in responding to CS2 as a function of its absolute duration. Experiment 3 examined the contribution of the food-food interval/CS onset-food interval ratio to these findings. In Experiments 1 and 2, the ratio differed for the overshadowing CS but not for the target CS. In Experiment 3, this arrangement was reversed, but the pattern of results remained the same. The implications of these findings for trial-based and real-time models of conditioning are discussed.     

The effect of scopolamine and physostigmine on working and reference memory in pigeons

A comparison of the effects of scopolamine and physostigmine on working memory and reference memory in White Carneaux pigeons was undertaken. In Experiment 1, the pigeons received injections of scopolamine hydrobromide (0.03 mg/kg), or saline. Scopolamine hydrobromide had greater disruptive effects on working memory trials than on reference memory trials, and the centrally active form of scopolamine disrupted working memory trial accuracy more than the peripherally active form. The differential sensitivity of accuracy on working memory trials to disruption by central cholinergic blockade was obtained even though the discrimination required on reference memory trials was more difficult. In Experiment 2, the pigeons received injections of scopolamine hydrobromide (0.015 mg/kg), physostigmine (0.075 mg/kg) both scopolamine and physostigmine, or saline. Physostigmine given with scopolamine was able to reverse the scopolamine-induced reduction of accuracy on working memory trials. In neither study did scopolamine promote accelerated forgetting as the delay interval was increased. These results indicate that manipulation of central cholinergic neurotransmitter systems influences working memory processes in the pigeon, but these effects occur without alterations in the ability of the birds to actively maintain information during the retention interval.     

Time,change, and motion: The effects of stimulus movement on temporal perception

The effects of stimulus motion on time perception were examined in five experiments. Subjects judged the durations (6–18 sec) of a series of computer-generated visual displays comprised of varying numbers of simple geometrical forms. In Experiment 1, subjects reproduced the duration of displays consisting of stationary or moving (at 20 cm/sec) stimulus figures. In Experiment 2, subjects reproduced the durations of stimuli that were either stationary, moving slowly (at 10 cm/sec), or moving fast (at 30 cm/sec). In Experiment 3, subjects used the production method to generate specified durations for stationary, slow, and fast displays. In Experiments 4 and 5, subjects reproduced the duration of stimuli that moved at speeds ranging from 0 to 45 cm/sec. Each experiment showed that stimulus motion lengthened perceived time. In general, faster speeds lengthened perceived time to a greater degree than slower speeds. Varying the number of stimuli appearing in the displays had only limited effects on time judgments. Other findings indicated that shorter intervals tended to be overestimated and longer intervals underestimated (Vierordt’s law), an effect which applied to both stationary and moving stimuli. The results support a change model of perceived time, which maintains that intervals associated with more changes are perceived to be longer than intervals with fewer changes.