The repeated acquisition of behavioral chains

Monkeys were trained with food reinforcement in a chamber containing four groups of three levers. For each session the monkey's task was to learn a new four-response chain by pressing the correct lever in each group. A stable pattern of learning resulted, and the number of errors reached a steady state from session to session. The technique was then used to determine how various durations of timeouts, following errors, affected the acquisition of new chains. With no timeout, the monkeys made a great many errors, due in large part to superstitious responses within the reinforced chain. Timeout durations ranging from 1 sec to 4 min reduced the number of errors substantially. A second experiment investigated the effects upon acquisition errors of presenting a single light (an “instruction” stimulus) over the correct lever. When this light did not influence the monkeys' responses to the three alternatives, the chains were learned as without it. When the light did control responding, the monkey pressed the appropriate sequence of levers but did not learn the sequence. Thus, when the light was removed, the monkey performed as if learning that sequence for the first time.     

Navigating two-dimensional mazes: chimpanzees (Pan troglodytes) and capuchins (Cebus apella sp.) profit from experience differently

We examined whether navigation is impacted by experience in two species of nonhuman primates. Five chimpanzees (Pan troglodytes) and seven capuchin monkeys (Cebus apella) navigated a cursor, using a joystick, through two-dimensional mazes presented on a computer monitor. Subjects completed 192 mazes, each one time. Each maze contained one to five choices, and in up to three of these choices, the correct path required moving the cursor away from the Euclidean direction toward the goal. Some subjects completed these mazes in a random order (Random group); others in a fixed order by ascending number of choices and ascending number of turns away from goal (Ordered group). Chimpanzees in both groups performed equivalently, demonstrated fewer errors and a higher rate of self-correcting errors with increasing experience at solving the mazes, and made significantly fewer errors than capuchin monkeys. Capuchins were more sensitive to the mode of presentation than chimpanzees; monkeys in the Ordered group made fewer errors than monkeys in the Random group. However, capuchins’ performance across testing changed little, and they remained particularly susceptible to making errors when the correct path required moving away from the goal. Thus, these two species responded differently to the same spatial challenges and same learning contexts. The findings indicate that chimpanzees have a strong advantage in this task compared to capuchins, no matter how the task is presented. We suggest that differences between the species in the dynamic organization of attention and motor processes contribute to their differences in performance on this task, and predict similar differences in other tasks requiring, as this one does, sustained attention to a dynamic visual display and self-produced movements variably towards and away from a goal.     

Effects of d-amphetamine, cocaine, and phencyclidine on the acquisition of response sequences with and without stimulus fading.

In each of three components of a multiple schedule, monkeys were required to emit a different sequence of four responses in a predetermined order on four levers. Sequence completions produced food on a fixed-ratio schedule. Errors produced a brief timeout. One component of the multiple schedule was a repeated-acquisition task where the four-response sequence changed each session (learning). The second component of the multiple schedule was also a repeated-acquisition task, but acquisition was supported through the use of a stimulus-fading procedure (faded learning). In a third component of the multiple schedule, the sequence of responses remained the same from session to session (performance). At higher doses, d-amphetamine, cocaine, and phencyclidine decreased the overall rate of responding and increased the percent errors in all three components. At lower doses, however, the three drugs produced selective effects on errors. Errors were increased in the learning component at lower doses than those required to disrupt the behavior in the faded-learning component. The performance component tended to be the least sensitive to disruptive drug effects. The data are consistent with the view that stimulus fading can modulate the effects of drugs on acquisition.     

Two-year-olds' search strategies and visual tracking in a hidden displacement task

Children younger than 3 years have difficulty with search tasks that involve hidden displacement. Partial visual information was provided about a ball's path as it moved toward a hiding place. Children (2.0 and 2.5 years old) saw a ball rolling down a ramp placed behind a transparent screen with 4 opaque doors. A wall, placed on the ramp and directly behind 1 of the doors, protruded above the screen and stopped the ball. Children were asked to find the ball. The transparency of the screen permitted visual tracking of the ball between the doors, but its final resting place was obscured. Both age groups were equally proficient at tracking the ball as it rolled behind the screen, but the 2.5-year-olds were more likely to reach to the correct door. Looking behavior was related to errors in the younger group in that tracking that stopped short or continued past the correct door was associated with incorrect choices.     

An experimental analysis of the effects of d-amphetamine and cocaine on the acquisition and performance of response chains in monkeys.

In one component of a multiple schedule of food presentation, monkeys acquired a different four-response chain each session by responding sequentially on three keys in the presence of four geometric forms (learning). In the other component, the four-response chain was the same each session (performance). Both d-amphetamine and cocaine, at the higher doses, disrupted the behavior in the learning component; the overall response rate decreased, the overall accuracy was impaired (i.e., percent errors increased), and there was less within-session error reduction. The performance component was generally less sensitive than the learning component to the disruptive effects of both drugs on rate and accuracy. After pre-feeding or during an extended session, the response rate decreased in both components, but accuracy was generally unaffected. When the four discriminative stimuli in both components were removed, the behavior was disrupted to a greater extent in the performance component. The disruptive effects of both drugs on behavior in the learning component were attenuated when the drugs were administered during the session after the response chain had been acquired. It was concluded that the greater sensitivity of the learning component to disruptive drug effects is related to the relatively weak stimulus control and/or the lower rate of reinforcement associated with that component.     

Timeout as a reinforcer for errors in a serial position task

After learning to press keys in a predetermined serial position sequence, with timeouts scheduled as a consequence of errors, monkeys developed stereotyped errors. As soon as a new trial started, the animals would make an error. On trial after trial, they pressed the same incorrect key at the first member of the sequence, even though they had previously learned the sequence. First-member errors occurred even when sequences of fully bright keys marking correct choices were presented. When timeout was eliminated as a consequence of one first-member error, subjects switched to an error that did produce the timeout. When all first-member errors failed to produce timeout the monkeys ceased responding. Both prefeeding and reduction in reinforcement density resulted in stereotyped errors occurring earlier in the session.     

Perseverative errors in object discrimination learning by aged Japanese monkeys (Macaca fuscata).

To examine the nature of age-dependent cognitive decline, performance in terms of concurrent object discriminations was assessed in aged and nonaged Japanese monkeys (Macaca fuscata). Aged monkeys required more sessions and committed more errors than nonaged ones in the discriminations, even in simple object discriminations. Analyses of errors suggest that aged monkeys repeated the same errors and committed more errors when they chose a negative object at the 1st trial. A hypothesis analysis of behavior suggests that their incorrect choices were mainly due to object preference. Therefore, the impairment was probably caused by a failure to inhibit inappropriate responses. Together with previous neuropsychological findings, deficits of aged monkeys in the performance of object discriminations can be explained by dysfunction of the frontal cortex.     

Long-term proactive interference and novelty enhancement effect in monkey list memory

Serial-probe-recognition (SPR) performance by 2 monkeys deteriorated over several months of training. Three hundred and twenty different items were presented without repetition within a session (trial unique) but were repeated between sessions. The cause of the deterioration was identified as proactive interference (PI) due to repetitive use of items from day to day. Introduction of novel stimuli across days improved performance from 63% to 82% correct (Experiment 1). Tests with only probe items and no list items (Experiment 2) revealed that the monkeys were using a familiar/novel response strategy in combination with a relational strategy (relating the probe item to the list items) to further improve their SPR performance. Intermixing familiar baseline trials and novel transfer trials within a session (Experiment 3) encouraged the subjects to use a relational strategy, and it improved performance on baseline trials as well as on transfer trials. Possible qualitative similarity between the relational strategy and the familiar/novel response strategy is discussed along with theoretical implications of these findings for experiments which have used small number of repeating stimuli within a session.     

Facilitation of a Distributed Shuttle-Box Conditioning with Posttraining Intracranial Self-Stimulation in Old Rats

Old Wistar rats (16–17 months) were trained in a two-way active avoidance task for 5 consecutive days (10 trials/day). Immediately after each training session a lateral hypothalamic intracranial self-stimulation session (ICSS group) or a sham-treatment session (Control group) was given to the animals. Long-term retention was tested 7 days after the last acquisition session. ICSS treatment led to a significant improvement in acquisition. In the long-term retention session the level of avoidance in both groups was similar to that achieved in the last acquisition session, although differences among groups failed to reach statistical significance. These results are compared with those obtained in previous experiments with young adult rats. While ICSS facilitated the process of acquisition in both young and old rats (however, it was much more powerful in young animals), further experiments are needed to elucidate whether this effect is long-lasting in old rats, as occurs in young adult subjects.     

Interference effects by spatial proximity and age-related declines in spatial memory by Japanese monkeys (Macaca fuscata): deficits in the combined use of multiple spatial cues

Spatial information processing was assessed in 3 young (4-10 years old) and 4 aged (24-25 years old) Japanese monkeys (Macaca fuscata) on 3 delayed nonmatching-to-position (DNMP) tests with relatively short delays of 5 s. Each test had 3 conditions of different horizontal distances between sample and to-be-nonmatched positions. Experiment 1 demonstrated that the performance on the DNMP test in both age groups was impaired when 2 stimulus positions were located next to each other; however, it was fairly accurate when they were located farther apart, suggesting that interference is introduced by spatial proximity. Experiment 2 revealed age-related differences in the situation in which an additional spatial cue, depth information, was available by extending the stimulus array of the DNMP test to a 4 x 2 matrix. In this test, young monkeys performed accurately irrespective of position distance between stimuli, whereas the aged monkeys' performance remained the same as before. Experiment 3 confirmed that the recognition ability in aged monkeys was well preserved on DNMP tests with different objects. These patterns of results indicate that the ability to use information from multiple spatial cues is not accessible to the aged monkeys.     

Contrasts in visual responsiveness and emotional arousal between rhesus monkeys raised with living and those raised with inanimate substitute mothers.

Rhesus monkeys were raised with dogs or inanimate surrogates in outdoor cages which provided them with complex, highly varied visual surroundings. Visual responsiveness to a variety of colored transparencies was investigated in three experiments, completed when the monkeys were between 18 and 30 mo old. Results indicated that the frequency and duration of looking at slides was significantly higher for dog-raised than for inanimate-surrogate-raised monkeys and that dog-raised monkeys were much more responsive to the novelty, complexity, ansal were obtained during the final experiment. Heart rate, vocalization, and changes in plasma cortisol were higher for monkeys raised with dogs. The frequency of most self-directed behaviors, however, was higher for monkeys raised with inanimate surrogates. Differences between rearing groups can only be the result of contrasts in attributes of the substitute mothers.     

Dissociation of data-based and expectancy-based memory following hippocampal lesions in rats

Sham-operated and nonoperated animals or animals with hippocampal lesions were presented with sets of trials to test both expectancy-based and data-based memory within the same task. During the study phase of each trial the animals were presented with a constant sequence of five arms on an eight-arm radial maze followed by a test phase in which a recognition test requiring a win-stay rule was used. Expectancy-based memory was measured during the study phase of the trials as a pattern of correct or incorrect orienting responses in anticipation of the ensuing doors in the constant sequence. Both groups of animals acquired correct orienting responses at the same rate, emitted the same pattern of correct orienting responses, and made the same number and pattern of intralist and extralist intrusion errors. Data-based memory was measured during the test phase of the trial as correct recognition test performance. During the test phase the animals with hippocampal lesions were impaired relative to controls on both immediate and 24-h recognition tests. These results suggest that the hippocampus might mediate only data-based, but not expectancy-based, memory and imply a possible dissociation between expectancy-based and data-based memory systems.     

Predicting memory for pictures from rankings of interestingness, pleasingness, complexity, figure-ground and clarity.

Pictures used in the test phase of a picture-recognition task were ranked by 10 university students for interestingness, pleasingness, complexity, figure-ground, and clarity. The scale values obtained from these rankings were then correlated with the errors made by other subjects in the test phase of the picture-recognition task. Figure-ground and clarity were found, for the most part, to be reliable predictors of the errors made to both the "old" and "new" subsets of pictures in two experiments. Complexity, while not as consistent a predictor of either figure-ground or clarity, was a reasonably good predictor. Interestingness and pleasingness were not reliable predictors of errors.     

Perseverative interference with object-in-place scene learning in rhesus monkeys with bilateral ablation of ventrolateral prefrontal cortex

Surgical disconnection of the frontal cortex and inferotemporal cortex severely impairs many aspects of visual learning and memory, including learning of new object-in-place scene memory problems, a monkey model of episodic memory. As part of a study of specialization within prefrontal cortex in visual learning and memory, we tested monkeys with bilateral ablations of ventrolateral prefrontal cortex in object-in-place scene learning. These monkeys were mildly impaired in scene learning relative to their own preoperative performance, similar in severity to that of monkeys with bilateral ablation of orbital prefrontal cortex. An analysis of response types showed that the monkeys with lesions were specifically impaired in responding to negative feedback during learning: The post-operative increase in errors was limited to trials in which the first response to each new problem, made on the basis of trial and error, was incorrect. This perseverative pattern of deficit was not observed in the same analysis of response types in monkeys with bilateral ablations of the orbital prefrontal cortex, who were equally impaired on trials with correct and incorrect first responses. This may represent a specific signature of ventrolateral prefrontal involvement in episodic learning and memory.     

Within-session reversal learning in rhesus macaques (<Emphasis Type="Italic">Macaca mulatta</Emphasis>)

In a midsession reversal (MSR) task, animals are typically presented with a simple, simultaneous discrimination (S1+, S2?) where contingencies are reversed (S1?, S2+) half-way through each session. This paradigm creates multiple, relevant cues that can aid in maximizing overall reinforcement. Recent research has shown that pigeons show systematic anticipatory and perseverative errors across the session, which increase as a function of proximity to the reversal trial. This behavior has been theorized to indicate primary control by temporal cues across the session, instead of the cues provided by recent reinforcement history that appear to control behavior shown by humans. Rats, however, appear to be guided by recent reinforcement history when tested in an operant context, thereby demonstrating behavior that parallels that seen in humans, but they appear to be guided by temporal cues when tested in an open-field apparatus, showing behavior more akin to that seen in pigeons. We tested rhesus macaques (Macaca mulatta) on the MSR with a computerized simultaneous visual discrimination to assess whether they would show errors indicative of control by time or by recent reinforcement history. When a single reversal point occurred midsession, rhesus macaques showed no anticipation of the reversal and a similar level of perseveration to rats tested in an operant setting. Nearly identical results also were observed when the monkeys were trained with a single, variable reversal point or with multiple, variable reversal points within a session. These results indicate that temporal cues are not guiding response flexibility in rhesus macaque visual discrimination.     

Autobiographical remembering and hypermnesia: a comparison of older and younger adults

This study examined age differences in autobiographical memory and extended findings concerning hypermnesia in laboratory tasks to a real world event, the announcement of the verdict in the O. J. Simpson murder trial. Older and younger adults repeatedly recalled the event in a single session. Interviews were coded for amount and type of accurate information and for errors. The age groups did not differ in ability to recall the gist of the event or in the number of errors made. Younger adults were better at remembering when the event had occurred. Both age groups showed hypermnesia. The results are discussed in terms of the importance of autobiographical memory across the life span and the phenomenon of hypermnesia in everyday memory.     

Conditioned suppression by a stimulus associated with nalorphine in morphine-dependent monkeys

Three rhesus monkeys, physically dependent on morphine, were trained to press a lever for food on a fixed ratio of 10 responses. A tone, initially a neutral stimulus, was aperiodically presented every third or fourth session, 5 min before and after the intravenous injection of nalorphine, a morphine antagonist which produces an immediate withdrawal syndrome in morphine-dependent monkeys. After several sessions, conditioned suppression of food-lever response rate was observed. Conditioned bradycardia, emesis, and excessive salivation also occurred. In 40 to 45 sessions the conditioned suppression of food-lever response rate and the conditioned autonomic changes were extinguished by presenting pairings of a tone and saline injection. The monkeys were then reconditioned by presenting the tone aperiodically, every third or fourth session, 5 min before and after the intravenous injection of nalorphine. Results were similar to the initial conditioning sessions. Two rhesus monkeys not dependent on morphine were stabilized on a food schedule similar to that used for the first three monkeys. These monkeys showed no change in food-lever response rate during or after nalorphine injections.     

Response coding in recall of colours by monkeys

Two monkeys learned a recall task with colours as the samples and spatial discriminanda at the retention test. The relationship between the sample colours and the retention test discriminanda was such that it was possible to distinguish between errors that were related to the visual similarity of the sample colours and those that were related to the similarity of the recall responses. When errors were induced by lengthening the delay between sample and retention test, they were related to the similarity of the recall responses. But in a comparable recognition memory task with colours, performance was related to the visual similarity of the colours. These results imply that the monkeys recognised the colours directly from a visual trace but recalled them indirectly from a response-coded trace.     

Post-reinforcement pauses and response rate of monkeys on a two-hand fixed-ratio schedule

Fixed-ratio behavior of monkeys was analyzed separately for two hands. While one hand responded on the fixed-ratio schedule the other performed a holding response and the function of the hands changed in alternate ratio runs. After performance was stable on the fixed ratio (70 responses, two monkeys; 100 responses, two monkeys, 120 responses, two monkeys) 90 sessions of further training equalized post-reinforcement pauses and the mean interresponse time of the two hands. Hand preference in reaching for food remained unchanged. Then, the fixed-ratio requirement was changed (a) in small sequential steps, (b) in two large steps, and, (c) within sessions alternating two runs at a high ratio with two runs at a low ratio. The mean duration of post-reinforcement pauses was correlated with a fixed ratio maintained throughout a session but single pauses were neither controlled by the immediately preceding nor by the following ratio run when a cue to its length was available. The mean interresponse time was insensitive to changes in fixed ratio. The fixed-ratio performance was generally similar to that of pigeons and rats.     

Repeated acquisition of response sequences: stimulus control and drugs.

Pigeons obtained food by making four responses on three keys in a specified sequence, e.g., left, right, center, right. Under the "tandem-learning" condition, all three keys were the same color throughout the response sequence, and the sequence was changed from session to session. After total errors per session (overall accuracy) and within-session error reduction (learning) had stabilized, the effects of varying doses phenobarbital and chlordiazepoxide were assessed. For comparison, the drug tests were also conducted under a "tandem-performance" condition, in which the response sequence was the same from session to session, and under corresponding "chain-learning" and "chain-performance" conditions, where different colored keylights were associated with the response sequence. Under all four baseline conditions, the largest dose of each drug impaired overall accuracy. Under the two learning conditions, the error rate decreased across trials within each session, but the degree of negative acceleration was less in the drug sessions than in the control sessions. In contrast, under the two performance conditions, the error rate was relatively constant across trials, but was higher in the drug sessions than in the control sessions. Of the four baselines, the chain-learning condition was the most sensitive to the drug effects.