Pigeons' spatial memory: III. Effect of distractors on delayed matching of key location.

The effect of distractors on pigeons' delayed matching of key location was investigated. Baseline trials began with a "ready" stimulus (brief operation of the grain feeder). Then one (randomly chosen) key from a three-by-three matrix was lit briefly as the sample. After a short delay (retention interval) the sample key was lit again along with one of the other eight keys. A peck at the key that had served as the sample (correct comparison) produced grain reinforcement, whereas a peck to the other key (incorrect comparison) produced only the intertrial interval. In Experiment 1, a houselight distractor, presented during either the sample, retention interval, or choice phases of the trial, had little if any effect on accuracy of matching key location. In Experiment 2, one of three types of spatial stimuli was interpolated during the retention interval, or the interval was blank as during baseline trials. The three stimuli were: the sample (correct comparison) location for that trial, the incorrect comparison location for that trial, or one of the seven unused locations for that trial. Relative to blank trials, accuracy improved slightly on sample-interpolated trials, decreased slightly on unused location-interpolated trials, and decreased considerably on incorrect comparison-interpolated trials. In Experiment 3, retention intervals were blank or had one of six types of interpolation: the sample, the incorrect comparison, two presentations of the sample, two presentations of the incorrect comparison, the sample followed by the incorrect comparison, or the incorrect comparison followed by the sample.(ABSTRACT TRUNCATED AT 250 WORDS)     

Discrimination training facilitates pigeons' performance on one-trial-per-day delayed matching of key location

Six pigeons were tested on a one-trial-per-day variant of delayed matching of key location. In one condition, a trial began with the illumination of a pair of quasi-randomly selected pecking keys in a large 10-key test box. Pigeons' pecks to one key (the sample) were reinforced with 8-second access to grain on a variable-interval 30-second schedule, whereas pecks to the other key (the distractor) had no scheduled consequences. In the second condition, the nonreinforced distractor was not presented. In both conditions, subjects were removed from the apparatus after 15 minutes and placed in a holding cage. Subjects were subsequently replaced in the box after a delay (retention interval) of 30 seconds and were reexposed to the illuminated sample and distractor keys for 1 minute. If a pigeon made more pecks to the sample during this interval, the distractor was extinguished and subsequent pecks to the sample were reinforced on the previous schedule for an additional 15 minutes. If, however, a pigeon made more pecks to the distractor, both keys were extinguished and the subject was returned to its home cage. For all subjects, matching-to-sample accuracy was higher in the first condition. In a second experiment, the retention interval was increased to 5, 15, and 30 minutes, and then to 1, 2, 4, 8, 12, and 24 hours. Most subjects remembered the correct key location for up to 4 hours, and in one case, up to 24 hours, demonstrating a spatial-memory proficiency far better than previously reported in this species on delayed matching tasks. The results are discussed in terms of the commonly held distinction between working and reference memory.     

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.     

Effect of delay-interval stimuli on delayed symbolic matching to sample in the pigeon

In Experiment 1, food-deprived pigeons received delayed symbolic matching to sample training in a darkened Skinner box. Trials began with the illumination of the grain feeder lamp (no food sample), or illumination of this lamp, accompanied by the raising of the feeder tray (food sample). After a delay of a few seconds, the two side response keys were illuminated, one with red and one with green light, with positions counterbalanced over trials. Pecking the red (green) comparison produced grain reinforcement if the trial had started with food (no food); pecking red after a no-food sample or green after a food sample was not reinforced. Once matching performance was stable, four stimuli were presented during the delay interval, and their effects on matching accuracy were evaluated. Both illumination of the houselight and the center key with white geometric forms decreased matching accuracy, whereas presentation of a tone and vibration of the test chamber did not. In Experiment 2, pecking the red center key was reinforced with food according to a variable interval schedule. The effects of occasional brief presentations of the four stimuli used in the first experiment on ongoing pecking were assessed. The houselight and form disturbed key pecking, but the tone and vibration did not. Thus, stimuli that interfered with delayed matching also interfered with simple operant behavior. Implications of these results for theories of remembering are discussed.     

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.     

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.     

Characteristics of forgetting functions in delayed matching to sample

Performance of pigeons in delayed matching-to-sample procedures was measured in terms of an index of discriminability derived from the difference between logarithms of ratios of choice responses to comparison stimuli following the different sample stimuli. Forgetting functions that plotted discriminability as a function of delay-interval duration were well described by a simple negative exponential function with two parameters, one describing initial discriminability of sample stimuli at zero delay (log d₀) and the other describing rate of decrement in discriminability with increasing delay-interval duration (b). With the difference between wavelength values of the comparison stimuli held constant, a large difference between wavelengths of the sample stimuli resulted in a higher log d₀ value than that for a small difference between sample stimuli, without changing the rate of decrement in discriminability, b. An increase in the fixed-ratio requirement for sample-key responding produced an increase in log d₀ without affecting b, and interpolation of ambient illumination in the delay interval increased b without influencing log d₀. Both parameters changed when intertrial-interval duration was varied. The result of variation in the point of interpolation of ambient illumination in the delay interval indicated that levels of discriminability at longer delays were independent of discriminability levels at earlier delays, consistent with the properties of the exponential function. Functions relating performance to delay-interval duration were suggested to have two characteristics: discriminability of the sample stimuli in the absence of a delay between the stimuli and the behavior they occasion, and rate of attenuation in discriminability with increasing delay-interval duration.     

Sample-duration effects on pigeons' delayed matching as a function of predictability of duration

Three experiments assessed the impact of sample duration on pigeons' delayed matching as a function of whether or not the samples themselves signaled how long they would remain on. When duration was uncorrelated with the sample appearing on each matching trial, the typical effect of duration was observed: Choice accuracy was higher with long (15-s) than with short (5-s) durations. By contrast, this difference either disappeared or reversed when the 5- and 15-s durations were correlated with the sample stimuli. Sample duration itself cued comparison choice by some birds in the latter (predictable) condition when duration was also correlated with the reinforced choice alternatives. However, even when duration could not provide a cue for choice, pigeons matched predictably short-duration samples as accurately as, or more accurately than, predictably long-duration samples. Moreover, this result was observed independently of whether the contextual conditions of the retention interval were the same as, or different from, those of the intertrial interval. These results strongly support the view that conditional stimulus control by the samples is partly a function of their conditioned reinforcing properties, as determined by the relative reduction in overall delay to reinforcement that they signal.     

Transfer of pigeons' matching to sample to novel sample locations

This study examined the conditions under which conditional stimulus control by the sample stimuli in three-key matching-to-sample paradigms would generalize across the different possible sample locations. In Experiments 1 and 2, the samples appeared on the left and right side keys during initial training and then on the center key during testing. Transfer of pigeons' matching performances to the center-key samples was evident after both identity and symbolic matching training. In Experiment 3, pigeons trained on symbolic matching with two side-key samples or with a side-key and a center-key sample generally transferred their learned matching performances to those samples when they subsequently appeared in the remaining (novel) location. These results indicate that, when two-choice conditional discriminations are learned with more than one sample location, the visual characteristics of the sample per se predominantly come to control the pigeons' comparison choices. This finding encourages the use of the multiple-location training procedure as a way of reducing control by location, thus providing a more discriminating test of symmetry in animals.     

Short-term memory for responses: the "choose-small" effect

Pigeons' short-term memory for fixed-ratio requirements was assessed using a delayed symbolic matching-to-sample procedure. Different choices were reinforced after fixed-ratio 10 and fixed-ratio 40 requirements, and delays of 0, 5, or 20 s were sometimes placed between sample ratios and choice. All birds made disproportionate numbers of responses to the small-ratio choice alternative when delays were interposed between ratios and choice, and this bias increased as a function of delay. Preference for the small fixed-ratio alternative was also observed on "no-sample" trials, during which the choice alternatives were presented without a prior sample ratio. This "choose-small" bias is analogous to results obtained by Spetch and Wilkie (1983) with event duration as the discriminative stimulus. The choose-small bias was attenuated when the houselight was turned on during delays, but overall accuracy was not influenced systematically by the houselight manipulation.     

On the development and mechanics of delayed matching-to-sample performance

Despite its frequent use to assess effects of environmental and pharmacological variables on short-term memory, little is known about the development of delayed matching-to-sample (DMTS) performance. This study was designed to examine the dimensions and dynamics of DMTS performance development over a long period of exposure to provide a more secure foundation for assessing stability in future research. Six pigeons were exposed to a DMTS task with variable delays for 300 sessions (i.e., 18,000 total trials; 3,600 trials per retention interval). Percent-correct and log-d measures used to quantify the development of conditional stimulus control under the procedure generally and at each of five retention intervals (0, 2, 4, 8 and 16-s) individually revealed that high levels of accuracy developed relatively quickly under the shorter retention intervals, but increases in accuracy under the longer retention intervals sometimes were not observed until 100-150 sessions had passed, with some still increasing at Session 300. Analyses of errors suggested that retention intervals induced biases by shifting control from the sample stimulus to control by position, something that was predicted by observed response biases during initial training. These results suggest that although it may require a great deal of exposure to DMTS prior to obtaining asymptotic steady state, quantification of model parameters may help predict trends when extended exposure is not feasible.     

Pigeons' preference for free choice: number of keys versus key area.

In concurrent-chains schedules, pigeons prefer terminal links that provide two keys correlated with reinforcers (free choice) over those that provide only one key (forced choice), terminal-link reinforcement rates being equal. With same-size keys, free choice provides a larger area available for pecking. Preferences were examined using terminal links that differed in key number only (one or two) or key size only (small and medium or medium and large), or that equated the area of the two free-choice keys with that of the forced-choice key. Medium (standard) keys were typically preferred to small keys, but indifference was typically obtained between medium and large keys. The size preference usually overrode free-choice preference with one medium key pitted against two small keys, but free-choice preference was reliably observed with one large key pitted against two medium keys. In other words, preferences were a joint function of key number and key area, implying that free-choice preference is not reducible to preference for larger key areas. Free-choice preference requires separate keys rather than larger areas; the relevant behavioral units are the discriminated operants correlated with each terminal-link key rather than classes defined by topographical features such as area or perimeter.     

DOES EFFORT PLAY A ROLE IN THE EFFECT OF RESPONSE REQUIREMENTS ON DELAYED MATCHING TO SAMPLE?

The possible role of "effort" in the accuracy of pigeons' performance on a delayed matching-to-sample procedure was investigated by examining the effects of response requirements that accompanied a trial-initiating stimulus and that accompanied a sample stimulus. In the first experiment, the effect of varying the size of a fixed-ratio requirement for responses during an initiating stimulus was compared to that of varying a similar requirement for responses during the sample stimulus. Accuracy increased reliably with increases in the ratio scheduled during the sample stimulus, but was not significantly affected by increases in the ratio scheduled on the key during the initiating stimulus. In another phase of Experiment 1, sample duration was held constant while the ratio requirement was varied during the initiating stimulus. Again, accuracy of matching to sample was not significantly affected by the size of the ratio scheduled during the initiating stimulus. Experiment 2 provided a systematic replication of these results in another group of pigeons and included a more detailed analysis of responding. These results support the view that increases in sample-response requirement facilitate accuracy of delayed matching by increasing the durations of exposure to the sample stimuli, and do not support a role of effort in the sample-response effect. In Experiment 3, the facilitative effect of responses on the sample but not of those on the initiating stimulus was replicated using a simultaneous matching-to-sample procedure. This finding provides further evidence against an interpretation of response-requirement effects that appeals to effort; the finding also suggests that sample exposure might affect initial discrimination of the sample rather than remembering the sample.     

Detecting a nonevent: Delayed presence-versus-absence discrimination in pigeons

Eight pigeons were trained on a delayed presence-versus-absence discrimination paradigm in which a sample stimulus was presented on some trials but not on others. If a sample was presented, then a response to one choice key produced food. If no sample was presented, a response to the other choice key produced food. The basic finding was that performance remained constant and well above 50% correct on no-sample trials as the retention interval increased, whereas performance dropped precipitously (to below 50% correct) on sample trials. In the second phase of the experiment, all of the trials were no-sample trials, and reinforcers were delivered probabilistically for one group of pigeons and according to time-based schedules for the other group. The exact reinforcement probabilities used in Phase 2 were those calculated to be in effect on no-sample trials in Phase 1 (according to a discrete-state model of performance). Subjects did not show exclusive preference for the richer alternative on no-sample trials in the first phase, but those in the probabilistic group developed near-exclusive preference for the richer alternative during the second phase. These data are inconsistent with the predictions of the discrete-state model, but are easily accommodated by an account based on signal detection theory, which also can be applied effectively to discrimination of event duration and the “subjective shortening” effect.     

RELATIONS AMONG ACUTE AND CHRONIC NICOTINE ADMINISTRATION,SHORT-TERM MEMORY,AND TACTICS OF DATA ANALYSIS

Emerging evidence suggests that nicotine may enhance short‐term memory. Some of this evidence comes from nonhuman primate research using a procedure called delayed matching‐to‐sample, wherein the monkey is trained to select a comparison stimulus that matches some physical property of a previously presented sample stimulus. Delays between sample stimulus offset and comparison stimuli onset are manipulated and accuracy is measured. The present research attempted to systematically replicate these enhancement effects with pigeons. In addition, the effects of nicotine were assessed under another, more dynamic, memory task called titrating‐delay matching‐to‐sample. In this procedure, the delay between sample offset and comparison onset adjusts as a function of the subject's performance. Correct matches increase the delay, mismatches decrease the delay, and titrated delay values serve as the primary dependent measure. Both studies examined nicotine's effects under acute and chronic administration. Neither provided clear or compelling evidence of memory enhancement following nicotine administration despite reliable and systematic dose‐related changes in response latency measures. A modest dose‐related effect on accuracy was found, but the magnitude of the effect appears to be directly related to tactics of data analysis involving best‐dose analyses of a very circumscribed subset of trial types.     

Use of an ambiguous-sample procedure to establish a cue to forget in pigeons

Pigeons were trained on a variation of the matching-to-sample task in which on double-sample trials two samples, one associated with each of the comparison stimuli, were presented successively. Responding to the comparison associated with the first sample was reinforced on half the double-sample trials, and responding to the comparison associated with the second sample was reinforced on the remaining half. One of two postsample stimuli was presented following the termination of each colored sample. A vertical line was presented after a correct or target sample, and a horizontal line was presented after an incorrect or interfering sample. With extended training, each bird demonstrated above-chance accuracy on double-sample trials, providing prima facie evidence that one or both of the postsample stimuli exerted control over matching behavior. Experiment 2 provided evidence that the horizontal line functioned as a cue to forget the code activated by the preceding sample stimulus. It was concluded that a condition sufficient to establish a postsample stimulus as a cue to forget is that the postsample immediately follow presentation of a sample that, if it were to control test responding, would lead to nonreinforcement.     

Formation of transitivity in conditional matching to sample by pigeons

Four homing pigeons were trained over 5 months in a zero-delay, “arbitrary” matching-to-sample procedure with sample and comparison stimuli presented on any of three response keys. Birds were also required to complete a fixed-ratio 10 requirement on both sample and comparison stimuli to terminate their presentation. The procedure resulted in the establishment of relations that were not specifically trained and that can be characterized by the property of transitivity in a stimulus equivalence context. This result was in contrast with the findings obtained from most previous research with nonhuman subjects.     

Pigeons' discounting of probabilistic and delayed reinforcers

Pigeons' discounting of probabilistic and delayed food reinforcers was studied using adjusting-amount procedures. In the probability discounting conditions, pigeons chose between an adjusting number of food pellets contingent on a single key peck and a larger, fixed number of pellets contingent on completion of a variable-ratio schedule. In the delay discounting conditions, pigeons chose between an adjusting number of pellets delivered immediately and a larger, fixed number of pellets delivered after a delay. Probability discounting (i.e., subjective value as a function of the odds against reinforcement) was as well described by a hyperboloid function as delay discounting was (i.e., subjective value as a function of the time until reinforcement). As in humans, the exponents of the hyperboloid function when it was fitted to the probability discounting data were lower than the exponents of the hyperboloid function when it was fitted to the delay discounting data. The subjective values of probabilistic reinforcers were strongly correlated with predictions based on simply substituting the average delay to their receipt in each probabilistic reinforcement condition into the hyperboloid discounting function. However, the subjective values were systematically underestimated using this approach. Using the discounting function proposed by Mazur (1989), which takes into account the variability in the delay to the probabilistic reinforcers, the accuracy with which their subjective values could be predicted was increased. Taken together, the present findings are consistent with Rachlin's (Rachlin, 1990; Rachlin, Logue, Gibbon, & Frankel, 1986) hypothesis that choice involving repeated gambles may be interpreted in terms of the delays to the probabilistic reinforcers.     

Sample-stimulus discriminability and sensitivity to reinforcement in delayed matching to sample

Five pigeons were trained in a delayed matching-to-sample task with red and green stimuli. The retention interval between sample-stimulus presentation and the availability of the choice stimuli was varied between 0.01 s and 12 s within each session. The probability of food produced by correct-red and correct-green responses was varied across conditions. Sample-stimulus discriminability and response bias were measured at four different retention intervals. The results of these analyses showed an interaction between the discriminability of the sample stimuli and the control exerted by differential reinforcement. At longer retention intervals, sample discriminability decreased and sensitivity of choice behavior to changes in the red/green reinforcer ratio increased. An analogous relation has been reported in conditional discriminations in which the physical disparity of stimuli has been varied. This correspondence suggests that increasing the delay between presentation of one of two stimuli and an opportunity to respond discriminatively to it may be functionally similar to increasing the physical similarity of the two stimuli.     

Control by sample location in pigeons'' matching to sample.

Three experiments assessed the impact of sample location in pigeons' matching to sample. Experiments 1 and 2 demonstrated that after line or hue identity matching was acquired to high levels of accuracy with center-key samples, varying sample location across the three keys disrupted performances. The drop in accuracy occurred following both zero-delay and simultaneous training and was mostly confined to trials in which the sample appeared on a side key. Experiment 3 attempted to diminish control by location by training birds to match samples that could appear in any location prior to center-key sample training and moving-sample testing with another set of stimuli. In testing, all birds performed accurately on center-sample trials and on side-key sample trials in which the matching choice appeared on the center key. Accuracy was below chance, however, on side-key sample trials in which the matching choice appeared on the other side key. One implication of the persistent control by sample location in the three-key paradigm is that it precludes the possibility of symmetry because symmetry tests require a change in the locations at which samples and comparisons appear.