Working Memory for Temporal and Nontemporal Events in Monkeys

This is the first report that introduces appropriate behavioral tasks for monkeys for investigations of working memory for temporal and nontemporal events. Using several behavioral tests, the study also shows how temporal information is coded during retention intervals in the tasks. Each of three monkeys was trained with two working memory tasks: delayed matching-to-sample of stimulus duration (DMS-D) and delayed matching-to-sample of stimulus color (DMS-C). The two tasks employed an identical apparatus and responses and differed only in the temporal and nontemporal attribute of the stimuli to be retained for correct performance. When a retention interval between the sample and comparison stimuli was prolonged, the monkeys made more incorrect responses to short samples in the DMS-C task, suggesting “trace decay” of memory for short stimuli. However, the same monkeys showed no such increase in incorrect responses to short samples in the DMS-D task, suggesting active coding of temporal information, that is, the length of stimulus duration, during the retention interval. When variable lengths of samples were presented with a fixed retention interval, the monkeys made more incorrect responses when length differences between short and long samples were small in the DMS-D task, but not in the DMS-C task. This suggests that the codes of working memory retained in the DMS-D task were not absolute (analogical) but rather were relative (categorical) and related to differences in the duration of the samples.     

Transfer of serial reversal learning in the pigeon

Pigeons were trained on a series of reversals of a simultaneous visual discrimination and were then shifted to a second series of reversals with different visual discriminanda. Pigeons that were given discrimination reversals with one pair of colours (Group Colour) and then shifted to a second pair of colours made fewer errors with the second pair than the first. In contrast, pigeons that were initially given reversals with a pair of orientations (Group Orientation) and then shifted to colours made as many errors during colour reversals as Group Colour had during initial colour training. When birds in Group Colour were subsequently shifted to orientation discrimination reversals, they performed no better than Group Orientation had during initial orientation training. The present results suggest that positive transfer from one series of discrimination reversals to a second, independent series may be constrained by the nature of the stimulus shift.     

The influence of acoustic similarity on serial recall of letter sequences

Acoustic similarity is known to impair short-term memory (STM) for letter sequences. The present series of experiments investigated the effects of acoustic similarity on long-term retention. In the first experiment, subjects were asked to learn one of two lists of 8 letters, the letters being either of high or low acoustic similarity. Lists were visually presented for three trials, with subjects responding after each trial. Then subjects participated in an immediate memory task for digits which lasted for 20 min. Finally, subjects tried to recall the list of letters they had learned previously. Lists having items of high acoustic similarity were more difficult to recall on the first trial, but were better recalled on the delayed retention test. In a second experiment, groups of subjects were again asked to learn one of two lists of 8 letters differing in acoustic similarity, using different orders of the letters used previously. The procedures were identical except that in two groups, a STM task for digits intervened between the presentation and test of the letters. This intervening task minimized the effects of STM and eliminated the differences in retention found previously. In a third experiment, better long-term retention for material having high acoustic similarity was also obtained when subjects used a backward recall procedure. In the last experiment 14 item lists were learned to a criterion of two correct trials, and retention was tested after each trial and at a delay of 20 min. and 23 hr. No effect of acoustic similarity was found and little retention loss occurred. These results suggest that reducing the STM component by introducing a STM control or by lengthening the list caused the effect of acoustic similarity to disappear.     

Dynamic visual noise reduces confidence in short-term memory for visual information

Previous research has shown effects of the visual interference technique, dynamic visual noise (DVN), on visual imagery, but not on visual short-term memory, unless retention of precise visual detail is required. This study tested the prediction that DVN does also affect retention of gross visual information, specifically by reducing confidence. Participants performed a matrix pattern memory task with three retention interval interference conditions (DVN, static visual noise and no interference control) that varied from trial to trial. At recall, participants indicated whether or not they were sure of their responses. As in previous research, DVN did not impair recall accuracy or latency on the task, but it did reduce recall confidence relative to static visual noise and no interference. We conclude that DVN does distort visual representations in short-term memory, but standard coarse-grained recall measures are insensitive to these distortions.     

Metamemory in tufted capuchin monkeys (Cebus apella)

Whereas evidence for metacognition by nonhuman primates has been obtained in great apes and old world monkeys, it is weaker in new world monkeys. For instance, capuchin monkeys may fail to recognize their own knowledge of the location of invisible bait. In the present study, we tested whether tufted capuchin monkeys would flexibly change their behavior in a delayed matching-to-sample (DMTS) test depending upon the strength of their memory trace of the sample. In Experiment 1, two monkeys were tested on a modified 9-alternative DMTS task with various delays on a computerized display. In some trials, the monkeys could choose whether to go for a memory test or for a simple key touch as an escape from the test. In other trials, they were forced to go for the memory test. Both monkeys escaped from the memory test more often when their matching accuracy on forced tests was lower. In one of the monkeys, the matching accuracies on chosen memory tests decreased more slowly as a function of delay length, and were higher after long delays than those on forced memory tests. This suggests that at least one capuchin monkey was able to recognize the strength of his own memory trace. Experiment 2 employed occasional no-sample tests, in which the monkeys faced the task choice without presentation of any sample for the trial. The monkey who was successful in Experiment 1 declined the memory test more often in no-sample trials than regular trials, further indicating metamemory in this individual. In Experiment 3, this successful monkey received a task, in which he was sometimes able to choose between shape MTS or texture MTS tasks. However, his matching accuracies did not differ between chosen tasks and forced tasks. Thus, the metamemory possessed by this new world monkey species may be more like a flag, showing strength of memory trace, than an elaborate representation showing details of the memory trace.     

Visual similarity in short-term recall for where and when

Two experiments examined the effects of visual similarity on short-term recall for where and when in the visual spatial domain. A series of squares of similar or dissimilar colours were serially presented at various locations on the screen. At recall, all coloured squares were simultaneously presented in a random order at the bottom of the screen, and the locations used for presentation were indicated by white squares. Participants were asked to place the colours at their appropriate location in their presentation order. Performance for location (where) and order (when) was assessed separately. Results revealed that similarity severely hinders both memory for what was where and memory for what was when, under quiet and articulatory suppression conditions. These results provide further evidence that similarity has a major impact on processing relational information in memory.     

Visual similarity in short-term recall for where and when

Two experiments examined the effects of visual similarity on short-term recall for where and when in the visual spatial domain. A series of squares of similar or dissimilar colours were serially presented at various locations on the screen. At recall, all coloured squares were simultaneously presented in a random order at the bottom of the screen, and the locations used for presentation were indicated by white squares. Participants were asked to place the colours at their appropriate location in their presentation order. Performance for location (where) and order (when) was assessed separately. Results revealed that similarity severely hinders both memory for what was where and memory for what was when, under quiet and articulatory suppression conditions. These results provide further evidence that similarity has a major impact on processing relational information in memory.     

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.     

Colour versus quantity as cues in reverse-reward-competent squirrel monkeys (Saimiri sciureus)

To assess the relative salience of colour and quantity cues, squirrel monkeys previously trained to reach for the smaller of two quantities of food in a reverse-reward contingency task received colour discrimination training. After initial failure to discriminate between two colours of dots under a differential reinforcement regime, they learned the task when the S- colour was associated with zero reward. The monkeys then showed good retention on the original reverse-reward task of 1 versus 4 with pairs of dots presented in S+ or S- colours. However, on "mismatch" trials of 1S- versus 4S+ , only 2 of 4 monkeys tested showed a preference--1 monkey chose based on quantity, the other based on colour. Individual differences and the possible roles of overshadowing and blocking are discussed.     

Colour versus quantity as cues in reverse-reward-competent squirrel monkeys (Saimiri sciureus)

To assess the relative salience of colour and quantity cues, squirrel monkeys previously trained to reach for the smaller of two quantities of food in a reverse-reward contingency task received colour discrimination training. After initial failure to discriminate between two colours of dots under a differential reinforcement regime, they learned the task when the S? colour was associated with zero reward. The monkeys then showed good retention on the original reverse-reward task of 1 versus 4 with pairs of dots presented in S + or S? colours. However, on “mismatch” trials of 1S? versus 4S + , only 2 of 4 monkeys tested showed a preference—1 monkey chose based on quantity, the other based on colour. Individual differences and the possible roles of overshadowing and blocking are discussed.     

Landmark Discrimination Learning in the Dog

Allocentric spatial memory was studied in dogs of varying ages and sources using a landmark discrimination task. The primary goal of this study was to develop a protocol to test landmark discrimination learning in the dog. Using a modified version of a landmark test developed for use in monkeys, we successfully trained dogs to make a spatial discrimination on the basis of the position of a visual landmark relative to two identical discriminanda. Task performance decreased, however, as the distance between the landmark and the “discriminandum” was increased. A subgroup of these dogs was also tested on a delayed nonmatching to position spatial memory task (DNMP), which relies on egocentric spatial cues. These findings suggest that dogs can acquire both allocentric and egocentric spatial tasks. These data provide a useful tool for evaluating the ability of canines to use allocentric cues in spatial learning.     

Visual discrimination learning in rats with lesions of superior colliculus: Door-push and approach errors in modified jumping stand

It has been suggested that, for some species, lesions of the superior colliculus affect visual discrimination learning, but only in certain conditions: (a) when problems are first learnt only after operation, or (b) when discriminanda require detailed scanning, or (c) when “approach” responses to the discriminanda are measured, rather than the response of actually touching them. These suggestions were examined in rats learning visual discriminations in a modified jumping-stand apparatus, after sustaining large lesions of the superior colliculus (and in some cases also of the pretectum). The lesions produced open-field hyperactivity and reduced exploration, indicating effective tectal damage, but the rats learnt a series of difficult discriminations in a door-push task as fast as normal rats, and they did not make more approach errors. Their main abnormality in the discrimination apparatus was that they looked less often between the stimulus doors before stepping across to one of them from the central platform. It is suggested that in rats, as in other animals, lesions of the superior colliculus disrupt the control of scanning head and eye movements; in rats, however, such disruption need not affect discrimination learning (at least in some kinds of apparatus), possibly because the retina of the rat has a relatively poorly developed area centralis.     

Association learning and visual discrimination

The effect of learned stimulus associations on visual discrimination was measured in four experiments. The stimuli were visual shapes which had been scaled for similarity. Two shapes were selected as discriminanda, and each S was pretested and posttested for discrimination of these briefly presented simultaneous pairs of “same” and “different” shapes. During the training, each discriminandum was paired with another simultaneously presented associated shape on a paired-associate response-learning task. The two associated shapes were very similar, intermediate in similarity, or very dissimilar. There was more improvement in posttest discrimination following training with dissimilar associates. We conclude that learned stimulus associations affect visual discrimination.     

Some characteristics of concurrent discrimination and retention by monkeys

Before and after learning-set training, 12 rhesus monkeys were tested on the acquisition and retention of tasks consisting of eight concurrent object discrimination problems. Training on the concurrent discrimination was administered unitil a fairly stringent acquisition criterion was met. Under these procedures, retention, unlike acquisition, was little influenced by initial object preferences. Excellent retention was observed both before and after learning-set training. In a second experiment, these same monkeys were tested on a series of concurrent tasks which provided different numbers of objects as the sets of correct and incorrect discriminanda. Task solutions depended largely upon acquiring and retaining a list of correct objects despite designation of the large or small sets as the correct one. The animals seemed not to use “exclusion” strategies even when this might have provided an efficient task solution. It was considered that the monkeys' performances were based on stimulus sampling characteristics like those seen in other discrimination testing situations.     

Linguistic coding by deaf children in relation to beginning reading success

The coding of printed letters in a task of consonant recall was examined in relation to the level of success of prelingually and profoundly deaf children (median age 8.75 years) in beginning reading. As determined by recall errors, the deaf children who were classified as good readers appeared to use both speech and fingerspelling (manual) codes in short-term retention of printed letters. In contrast, deaf children classified as poor readers did not show influence of either of these linguistically based codes in recall. Thus, the success of deaf children in beginning reading, like that of hearing children, appears to be related to the ability to establish and make use of linguistically recoded representations of the language. Neither group showed evidence of dependence on visual cues for recall.     

The influence of similarity on visual working memory representations

In verbal memory, similarity between items in memory often leads to interference and impaired memory performance. The present study sought to determine whether analogous interference effects would be observed in visual working memory by varying the similarity of the to-be-remembered objects in a colour change detection task. Instead of leading to interference and impaired performance, increased similarity among the items being held in memory led to improved performance. Moreover, when two similar colours were presented along with one dissimilar colour, memory performance was better for the similar colours than for the dissimilar colour. Similarity produced better performance even when the objects were presented sequentially and even when memory for the first item in the sequence was tested. These findings show that similarity does not lead to interference between representations in visual working memory. Instead, similarity may lead to improved task performance, possibly due to increased stability or precision of the memory representations during maintenance.     

A method for examining short-term retention of haptic cues in monkeys

Tasks used to examine short-term memory (STM) in animals have almost exclusively required retention of visual cues. To determine if haptic information can be retained, three rhesus monkeys were trained to perform, using only haptic cues, a simultaneous (SMS) and a delayed matching-to-sample (DMS) task. On each trial, the monkeys felt and responded to a sample stimulus on a centrally located manipulandum. They were then presented two comparison stimuli located on both sides of the central manipulandum. A response matching the comparison stimulus with the sample stimulus was reinforced. In SMS a mean of 2,725 trials was required to reach a criterion of 90% correct. As in DMS performance for visual cues, in haptic DMS the monkeys were capable of above-chance responding at retention intervals of greater than 1 min. This haptic DMS task should be useful for testing STM models, for examining the physiological basis of STM, and for examining drug effects.     

Preselection and response biasing in short-term motor memory

Two experiments were performed comparing preselected (subject defined) and constrained (experimenter defined) movements. In the first experiment, subjects made reproduction responses immediately or under unfilled and filled 15-sec retention intervals. Results indicated that recall of preselected movements was clearly superior until the interpolation of information processing activity. In addition, preselected movements demonstrated no forgetting over a 15-sec retention interval while constrained movements evidenced spontaneous memory lass, suggesting that preselected movements possess a stronger representation in memory. The second experiment examined this interpretation in a response biasing paradigm. Subjects made criterion responses under preselected or constrained conditions, while the interpolated movement was always in the constrained mode and ± 40 deg from the criterion. The subjects' task was to attend to both movements and recall each when instructed. While preselected recall was clearly superior' to constrained recall, response biasing was clearly evident in both. The failure to find differential biasing effects was discussed in terms of the relative trace strength hypothesis (Stelmach & Welsh, 1972).     

Visual and phonological components of working memory in children

Previous studies have shown that young children's immediate memory for a short series of drawings of objects is mediated by a visual component of working memory, whereas older children rely chiefly upon a phonological component. Three experiments investigated the hypothesis that older children rely also, but to a lesser extent, on visual working memory. Experiment 1 confirmed previous evidence that 11-year-olds' memory is disrupted by phonemic similarity of object names, but is unaffected by visual similarity of the objects themselves. However, when articulatory suppression was used to prevent phonological coding, levels of recall were sensitive to visual rather than phonemic similarity. Experiment 2 compared the effects of interpolating an auditory-verbal or a visual postlist task on memory for drawings viewed either with or without suppression. The visual task had a clear disruptive effect only in the suppression condition, where it interfered selectively with recall of the most recent item. Experiment 3 compared the effects of interpolating an auditory-verbal or a mixed-modality (visual-auditory) postlist task when subjects were not required to suppress. There was greater interference from the mixed-modality task, and this effect was confined to the last item presented. These experiments are taken as confirming the presence of a small but reliable contribution from visual memory in 11-year-old children's recall. As in younger children, visual working memory in 11-year-olds is sensitive to visual similarity and is responsible for a final-item visual recency effect. The results also show that older children's use of visual working memory is usually masked by the more pervasive phonological component of recall. Some implications for the structure of working memory and its development are discussed.