Note on delay-interval illumination effects on retention in monkeys (Cebus apella).

Two monkeys experienced with delayed matching to sample were given a 30-day baseline training period during which the delay interval was illuminated. Both subjects showed an increase in matching accuracy when shifted to dark delay intervals, and accuracy declined when the illuminated delay interval was reinstituted. These results, as well as earlier reports of facilitation of delayed matching behavior by dark delay intervals, support the view that the absolute level of delay-interval illumination can importantly affect visual retention in monkeys and may be indicative of significant differences in the retention mechanisms employed by monkeys and birds.     

A comparison of visual and auditory short-term memory in monkeys (Cebus apella)

Using a visual and an acoustic sample set that appeared to favour the auditory modality of the monkey subjects, in Experiment 1 retention gradients generated in closely comparable visual and auditory matching (go/no-go) tasks revealed a more durable short-term memory (STM) for the visual modality. In Experiment 2, potentially interfering visual and acoustic stimuli were introduced during the retention intervals of the auditory matching task. Unlike the case of visual STM, delay-interval visual stimulation did not affect auditory STM. On the other hand, delay-interval music decreased auditory STM, confirming that the monkeys maintained an auditory trace during the retention intervals. Surprisingly, monkey vocalizations injected during the retention intervals caused much less interference than music. This finding, which was confirmed by the results of Experiments 3 and 4, may be due to differential processing of “arbitrary” (the acoustic samples) and species-specific (monkey vocalizations) sounds by the subjects. Although less robust than visual STM, auditory STM was nevertheless substantial, even with retention intervals as long as 32 sec.     

Directed forgetting of elements in compound samples.

Pigeons were trained in a delayed matching-to-sample procedure in which the sample stimuli consisted of a compound of color (red or green) and spatial location (left or right). A postsample cue (houselight on or off) signaled whether color matching or location matching would be required following the delay. In Experiment 1, the reduction in performance on probe trials (in which the houselight condition was reversed relative to that on regular trials) was greater for location matching than for color matching. The birds showed overt mediational behavior during the delays on location-matching trials. On color-matching trials, the birds exhibited behavior during delays that might have interfered with that mediational behavior. In Experiment 2, the houselight condition was changed shortly before presentation of the comparison stimuli on probe trials. Accuracy of location matching was reduced when the cue initially signaled color matching and was then changed to signal location matching, whereas matching accuracy was not reduced by a change in the opposite direction. Accuracy of color matching was reduced by a change in illumination level from dark to light, regardless of type of the relevant dimension signaled by houselight illumination. Discussion of these findings focuses on the variables critical to establishment of an effective cue to forget.     

Effect of delay-interval illumination on matching behavior in the capuchin monkey

Experiment 1 demonstrated that delayed matching-to-sample in the capuchin monkey was superior when the delay interval was spent in darkness rather than in moderate illumination. In contrast with previous studies in which the delayed-matching ability of primates appeared limited to 60 sec or less, in the dark condition all subjects showed above-chance matching at a 120-sec delay interval. Experiment 2 verified that darkness during the delay interval can facilitate delayed matching and provided evidence that the effective variable was the illumination level of the delay interval rather than change in illumination, which in Exp. 1 was confounded with illumination level.     

Cold-induced impairment of delayed matching in rats

Exposure to moderate, nonhypothermic cold temperature has been reported to affect a variety of behavioral and neural functions. To elucidate the effects of mild cold stress on short-term (working) memory, Long-Evans rats were exposed to an ambient temperature of either 2 degrees or 23 degrees C while performing a delayed matching task. At the beginning of each trial, rats were required to respond on one of two levers cued by a light. Following a delay of 2, 8, or 16 s, a response on the lever previously cued produced food reinforcement. Relative to performance at 23 degrees C, exposure to 2 degrees C occasioned no change in matching accuracy at the 2-s delay, a modest decrement at the 8-s delay, and a larger decrement at the 16-s delay. The cold exposure did not decrease colonic temperature. In addition to accuracy decrements, matching response times were consistently shorter during cold exposures. Cold-induced impairments were absent during removal of the memory component from the task, indicating the observed cold effects on memory were not due to impaired attentional, sensory, or motor processes. These data suggest that mild cold stress may impair active maintenance of information in working memory but not processes related to reference memory.     

Key pecking of pigeons under variable-interval schedules of briefly signaled delayed reinforcement: effects of variable-interval value.

Key pecking of 4 pigeons was maintained under a multiple variable-interval 20-s variable-interval 120-s schedule of food reinforcement. When rates of key pecking were stable, a 5-s unsignaled, nonresetting delay to reinforcement separated the first peck after an interval elapsed from reinforcement in both components. Rates of pecking decreased substantially in both components. When rates were stable, the situation was changed such that the peck that began the 5-s delay also changed the color of the keylight for 0.5 s (i.e., the delay was briefly signaled). Rates increased to near-immediate reinforcement levels. In subsequent conditions, delays of 10 and 20 s, still briefly signaled, were tested. Although rates of key pecking during the component with the variable-interval 120-s schedule did not change appreciably across conditions, rates during the variable-interval 20-s component decreased greatly in 1 pigeon at the 10-s delay and decreased in all pigeons at the 20-s delay. In a control condition, the variable-interval 20-s schedule with 20-s delays was changed to a variable-interval 35-s schedule with 5-s delays, thus equating nominal rates of reinforcement. Rates of pecking increased to baseline levels. Rates of pecking, then, depended on the value of the briefly signaled delay relative to the programmed interfood times, rather than on the absolute delay value. These results are discussed in terms of similar findings in the literature on conditioned reinforcement, delayed matching to sample, and classical conditioning.     

Pigeons may not remember the stimuli that reinforced their recent behavior

In two experiments the conditioned reinforcing and delayed discriminative stimulus functions of stimuli that signal delays to reinforcement were studied. Pigeons' pecks to a center key produced delayed-matching-to-sample trials according to a variable-interval 60-s (or 30-s in 1 pigeon) schedule (Experiment 1) or a multiple variable-interval 20-s variable-interval 120-s schedule (Experiment 2). The trials consisted of a 2-s illumination of one of two sample key colors followed by delays ranging across phases from 0.1 to 27.0 s followed in turn by the presentation of matching and nonmatching comparison stimuli on the side keys. Pecks to the key color that matched the sample were reinforced with 4-s access to grain. Under some conditions of Experiment 1, pecks to nonmatching comparison stimuli produced a 4-s blackout and the start of the next interval. Under other conditions of Experiment 1 and each condition of Experiment 2, pecks to nonmatching stimuli had no effect and trials ended only when pigeons pecked the other, matching stimulus and received food. The functions relating pretrial response rates to delays differed markedly from those relating matching-to-sample accuracy to delays. Specifically, response rates remained relatively high until the longest delays (15.0 to 27.0 s) were arranged, at which point they fell to low levels. Matching accuracy was high at short delays, but fell to chance at delays between 3.0 and 9.0 s. In Experiment 2, both matching accuracy and response rates remained high over a wider range of delays in the variable-interval 120-s component relative to the variable-interval 20-s component. The difference in matching accuracy between the components was not due to an increased tendency in the variable-interval 20-s component toward proactive interference following short intervals. Thus, under these experimental conditions the conditioned reinforcing and the delayed discriminative functions of the sample stimulus depended on the same variables (delay and variable-interval value), but were nevertheless dissociated.     

Feature saliency affects delayed matching of an attended feature

We examined how top-down attentional modulation and bottom-up stimulus saliency interact with feature memory. Experiment 1 used a delayed-matching-to-sample (DMS) task to examine the relative saliency between features by observing the relative accuracy of recognition at different stimulus durations. Feature salience decreased according to the following order: colour, form, and texture. In a modified DMS task (Experiments 2 and 3), participants were required to attend to one of three features and ignore the others. After a delay, they were required to choose which of the two test stimuli matched the reference stimulus on the attended feature, disregarding other task-irrelevant features. The target was either identical to the reference stimulus or mismatched the reference stimulus on one of the irrelevant features. The results showed that colour matching was affected neither by a form change nor by a texture change. Form matching was affected by a colour change, and texture matching was affected by a colour or form change. These results are consistent with the relative saliency hypothesis. Even when all features of an attended object are maintained, a relatively more salient task-irrelevant feature can interfere with the delayed recognition of a less salient feature.     

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.     

Sources of visual interference in delayed matching-to-sample with pigeons

In two experiments, independent groups of pigeons were trained on an identity matching task involving line orientations as sample and comparison stimuli. For some birds an overhead houselight was illuminated continuously throughout each training session. For other birds the houselight was never illuminated during training sessions. During subsequent testing, the lighting conditions during the delay were the same as in training on some trials, but on other trials they were opposite those of training during either the entire delay (Experiment 1) or during a portion of the delay (Experiment 2). In birds trained with the houselight off, turning the houselight on during the delay produced a large and enduring disruption in matching accuracy. On the other hand, in birds trained with the houselight on, turning the houselight off during the delay produced only a moderate and temporary disruption in matching accuracy. These findings are inconsistent with the prevailing view that retroactive interference in pigeons is a function of a change in illumination level relative to that which prevailed during training. In pigeons, as in monkeys, sustained retoactive interference effects obtain only when the level of illumination is increase during the delay interval.     

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.     

Delay-dependent involvement of the rat entorhinal cortex in habituation to a novel environment

Evidence has accumulated that the entorhinal cortex (EC) is involved in memory operations underlying formation of a long-term memory. Because entorhinal-lesioned rats are impaired for long delays in delayed matching and non-matching to sample tasks, it has been proposed that EC contributes to the maintenance of information in short-term memory. In the present study, we asked whether such a time-limited role applies also when learning complex spatial information in a novel environment. We therefore examined the effects of EC lesions on habituation in an object exploration task in which a delay of either 4 min or 10 min is imposed between successive sessions. EC-lesioned rats exhibited a deficit in habituation at 10 min but not 4 min delays. Following habituation, reactions to spatial change (object configuration) and non-spatial change (novel object) were also examined. EC-lesioned rats were impaired in detecting the spatial change but were able to detect a non-spatial change, irrespective of the delay. Overall, the results suggest that EC is involved in maintaining a large amount of novel, multidimensional information in short-term memory therefore enabling formation of long-term memory. Switching to a novelty detection mode would then allow the animal to rapidly adapt to environmental changes. In this mode, EC would preferentially process spatial information rather than non-spatial information.     

The avian hippocampus and short-term memory for spatial and non-spatial information

Three experiments investigated the role of the pigeon hippocampal formation (the hippocampus and area-parahippocampalis) in short-term memory for non-spatial and spatial information. The acquisition of delayed matching-to-sample and the short-term retention of non-spatial visual information, using a small set of sample stimuli, were unaffected by aspiration lesions of the hippocampus or the neostriatum (Experiment 1). Similarly, acquisition and short-term retention of non-spatial information using a successive, trial-unique, delayed non-matching-to-sample procedure were unaffected by hippocampal damage; the same birds had, however, displayed a profound autoshaping impairment (Experiment 2). Acquisition of a spatial delayed matching-to-sample task was unimpaired by hippocampal damage. However, lesioned animals were impaired following the introduction of retention intervals on this procedure (Experiment 3). The correspondence between the behavioural effects of hippocampal lesions in birds and mammals on short-term memory is discussed, and the implications of these results for avian hippocampal function are considered.     

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.     

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.     

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.     

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.     

Stimulus control topographies and tests of symmetry in pigeons

Pigeons were tested for symmetry after A-B training under conditions designed to avoid problems that may prevent its emergence, namely the change of stimulus location in testing relative to training and the lack of requisite discrimination training. In Experiment 1, samples appeared in two locations during baseline training to minimize the impact of stimulus location. Experiments 2 and 3 included multiple-location training along with additional identity and symbolic matching training, respectively, to explicitly train all of the simultaneous and successive stimulus discriminations required for testing. Experiment 4 provided reinforcement for symmetrical matching relations with some stimulus sets (with multiple-location training) prior to symmetry testing with different sets. In all experiments, pigeons showed no evidence of symmetry despite the fact that baseline (A-B) matching transferred to novel locations. Additional tests for reflexivity (Experiment 2) yielded similar outcomes. These results indicate that the change in stimulus location is not the sole reason that pigeons do not show symmetry and increase the plausibility of arguments that symmetry and other indexes of stimulus equivalence may be beyond the capabilities of the pigeon.     

Token reinforcement, choice, and self-control in pigeons.

Pigeons were exposed to self-control procedures that involved illumination of light-emitting diodes (LEDs) as a form of token reinforcement. In a discrete-trials arrangement, subjects chose between one and three LEDs; each LED was exchangeable for 2-s access to food during distinct posttrial exchange periods. In Experiment 1, subjects generally preferred the immediate presentation of a single LED over the delayed presentation of three LEDs, but differences in the delay to the exchange period between the two options prevented a clear assessment of the relative influence of LED delay and exchange-period delay as determinants of choice. In Experiment 2, in which delays to the exchange period from either alternative were equal in most conditions, all subjects preferred the delayed three LEDs more often than in Experiment-1. In Experiment 3, subjects preferred the option that resulted in a greater amount of food more often if the choices also produced LEDs than if they did not. In Experiment 4, preference for the delayed three LEDs was obtained when delays to the exchange period were equal, but reversed in favor of an immediate single LED when the latter choice also resulted in quicker access to exchange periods. The overall pattern of results suggests that (a) delay to the exchange period is a more critical determinant of choice than is delay to token presentation; (b) tokens may function as conditioned reinforcers, although their discriminative properties may be responsible for the self-control that occurs under token reinforcer arrangements; and (c) previously reported differences in the self-control choices of humans and pigeons may have resulted at least in part from the procedural conventions of using token reinforcers with human subjects and food reinforcers with pigeon subjects.     

Assessing the associative deficit of older adults in long-term and short-term/working memory

Older adults exhibit a deficit in associative long-term memory relative to younger adults. However, the literature is inconclusive regarding whether this deficit is attenuated in short-term/working memory. To elucidate the issue, three experiments assessed younger and older adults' item and interitem associative memory and the effects of several variables that might potentially contribute to the inconsistent pattern of results in previous studies. In Experiment 1, participants were tested on item and associative recognition memory with both long-term and short-term retention intervals in a single, continuous recognition paradigm. There was an associative deficit for older adults in the short-term and long-term intervals. Using only short-term intervals, Experiment 2 utilized mixed and blocked test designs to examine the effect of test event salience. Blocking the test did not attenuate the age-related associative deficit seen in the mixed test blocks. Finally, an age-related associative deficit was found in Experiment 3, under both sequential and simultaneous presentation conditions. Even while accounting for some methodological issues, the associative deficit of older adults is evident in short-term/working memory. (PsycINFO Database Record (c) 2012 APA, all rights reserved).