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.     

Pentobarbital self-administration in rhesus monkeys: drug concentration and fixed-ratio size interactions.

Performances of three rhesus monkeys were reinforced by the oral delivery of pentobarbital and studied as functions of fixed-ratio size and drug concentration. Pentobarbital solutions and water were concurrently available on identical reinforcement schedules from separate liquid-delivery systems during 3-hour sessions. Under a fixed-ratio 16 schedule of drug availability, a descending series of drug concentrations was tested (4, 2, 1, 0.5, 0.25, 0.125, and 0.0625 mg/ml, followed by a retest at 4 mg/ml). Partial concentration series beginning with the highest concentration were repeated with fixed-ratios of 32 and 64, with a fixed-ratio 128 for two monkeys, and with fixed-ratio 256 for one. At each fixed-ratio value, response rate and number of drug deliveries were inverted U-shaped functions of pentobarbital concentration. Drug intake (mg/kg/session) increased directly with drug concentration. As the fixed-ratio size was increased, the number of liquid deliveries decreased. For each drug concentration, when the numbers of drug deliveries at fixed-ratios of 32, 64, and 128 responses were plotted as percentages of those obtained at fixed-ratio 16, the following orderly relationship emerged: the higher the drug concentration, the less that drug deliveries were decreased by increases in fixed-ratio size. This relationship indicates an increase in reinforcing efficacy with increases in pentobarbital concentration.     

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.     

Monkeys (Macaca mulatta and Cebus apella) track, enumerate, and compare multiple sets of moving items

Despite many demonstrations of numerical competence in nonhuman animals, little is known about how well animals enumerate moving stimuli. In this series of experiments, rhesus monkeys (Macaca mulatta) and capuchin monkeys (Cebus apella) performed computerized tasks in which they had to enumerate sets of stimuli. In Experiment 1, rhesus monkeys compared two sets of moving stimuli. Experiment 2 required comparisons of a moving set and a static set. Experiment 3 included human participants and capuchin monkeys to assess all 3 species' performance and to determine whether responding was to the numerical properties of the stimulus sets rather than to some other stimulus property such as cumulative area. Experiment 4 required both monkey species to enumerate subsets of each moving array. In all experiments, monkeys performed above chance levels, and their responses were controlled by the number of items in the arrays as opposed to nonnumerical stimulus dimensions. Rhesus monkeys performed comparably to adult humans when directly compared although capuchin performance was lower.     

The evolutionary origins of natural pedagogy: Rhesus monkeys show sustained attention following nonsocial cues versus social communicative signals

The natural pedagogy hypothesis proposes that human infants preferentially attend to communicative signals from others, facilitating rapid cultural learning. In this view, sensitivity to such signals is a uniquely human adaptation and as such nonhuman animals should not produce or utilize these communicative signals. We test these evolutionary predictions by examining sensitivity to communicative cues in 206 rhesus monkeys (Macaca mulatta) using an expectancy looking time task modeled on prior work with infants. Monkeys observed a human actor who either made eye contact and vocalized to the monkey (social cue), or waved a fruit in front of her face and produced a tapping sound (nonsocial cue). The actor then either looked at an object (referential look) or looked toward empty space (look away). We found that, unlike human infants in analogous situations, rhesus monkeys looked longer at events following nonsocial cues, regardless of the demonstrator's subsequent looking behavior. Moreover younger and older monkeys showed similar patterns of responses across development. These results provide support for the natural pedagogy hypothesis, while also highlighting evolutionary changes in human sensitivity to communicative signals.     

Control of concurrent avoidance and appetitive behaviors by an indicator of shock proximity

Two monkeys were exposed to concurrent schedules in which every fifth response on one lever had the dual consequences of delivering food and stepping a 20-step light counter upward; responses on a second lever stepped the light downward. By responding appropriately on each lever, a ration of food could be obtained without exposure to a brief shock that occurred when the counter reached the twentieth step. Behavioral patterns of light-counter control were analyzed when the monkeys were handicapped by several types of discontinuities in the 20-step counter: (1) different sets of four consecutive lights inoperative, (2) the upper or lower 10 steps inoperative, and (3) all lights inoperative. Generally, the monkeys maintained the light in the upper five steps of the 20-step light counter if the lights immediately preceding the shock position were functional. However, when these counter positions were inoperative, subjects maintained the counter around the highest operative position. With all positions of the light counter inoperative, no systematic pattern of responding on the food and avoidance levers occurred.     

Sequential planning in rhesus monkeys (<Emphasis Type="Italic">Macaca mulatta</Emphasis>)

In the current study, we examined the planning abilities of rhesus monkeys (Macaca mulatta) by training them on a five-item list composed of coloured photographs and then testing them on switch and mask trials. In contrast to previous studies where monkeys made responses using a joystick, in the current study, monkeys made responses directly to a touch screen. On switch trials, after a response to the first list item, the on-screen positions of two list items were exchanged. Performance on trials in which the second and third list items were exchanged was poorer compared to normal (non-switch) trials for all subjects. When the third and fourth items were exchanged, however, only one subject continued to show performance deficits. On mask trials, following a response to the first item, the remaining items were covered by opaque white squares. When two items were masked, all four subjects responded to each masked item at a level significantly above chance. When three items were masked, however, only one subjected was able to respond to all three masked items at a level significantly above chance. The results of the present study indicate that three of our four monkeys planned one response ahead while a single monkey planned two responses ahead. The significance of these findings is discussed in relation to previous studies on planning in chimpanzees and monkeys.     

Formation of the sameness-difference concept by Japanese monkeys from a small number of color stimuli

Japanese monkeys were trained to form the sameness-difference concept. In Experiment 1, four monkeys were trained with two colors to discriminate matching stimulus pairs from nonmatching pairs by reinforcing only lever-pressing responses to matching pairs with a variable-interval schedule. Three monkeys showed successful transfer of this discrimination to two new colors, thus demonstrating that some Japanese monkeys are able to form this relational concept from a minimum number of stimuli. In Experiment 2, two monkeys were trained, in a Yes/No procedure with three colors, to press one lever under matching pairs and another lever under nonmatching pairs. Poor transfer performances to three new colors suggest that simultaneously establishing two different response patterns to matching and nonmatching pairs is ineffective in forming the concept. In Experiment 3, the amount of transfer to three new colors after mastering a standard three-color matching-to-sample task was compared with that of a modified task in which correct responses were reinforced with a within-trial variable-interval schedule. All three monkeys showed greater transfer with the modified procedure. The results suggest that the variable-interval schedule adopted within trials is effective in forming the sameness-difference concept.     

Individual differences in response to a stranger: social impulsivity as a dimension of temperament in vervet monkeys (Cercopithecus aethiops sabaeus)

Social impulsivity in response to a stranger was assessed in male vervet monkeys (Cercopithecus aethiops sabaeus) using the Intruder Challenge Test. Vervets (N = 128, ages 3-18 years) were presented with an unfamiliar adult male at the periphery of the home enclosure. An index of impulsivity reflecting variation in the tendency to rapidly approach, engage, and challenge the intruder was derived from factor analysis of behavioral responses. Scale reliability (alpha = .84) and test-retest consistency (intraclass correlation = .83) were high, indicating that this index reliably and efficiently measures a stable aspect of temperament from impulsive to inhibited. Impulsivity scores peaked at age 4, when vervet males typically emigrate from the natal group. The highest ranking males in each group were more likely to score in the moderate range, whereas lower ranking males were more likely to score in the highest (impulsive) or lowest (inhibited) quartiles.     

Continuity and Change in Emotional Reactivity in Rhesus Monkeys Throughout the Prepubertal Period

We examined continuity and change in emotional reactiviy as assessed by hypothalamic–pituitary–adrenal (HPA) activity in rhesus monkeys. Plasma samples were obtained from mother-peer–reared and surrogate-peer–reared monkeys and assayed for cortisol concentrations at 5 time points in infancy. Both plasma and saliva samples were collected for cortisol assay from the same monkeys as 1-, 2-, and 3-year-old juveniles, respectively. Plasma cortisol concentrations increased during the first 5 months of life and decreased from 1 to 3 years of age. Females consistently had higher plasma cortisol concentrations than males. Plasma cortisol concentrations were lower in surrogate-peer–reared than mother-peer–reared monkeys during the first month of life. Juvenile plasma cortisol concentrations were significantly correlated with infant cortisol levels at all but the earliest time point. Analysis of salivary cortisol from a subset of animals showed a similar age-related decline and the presence of a significant rearing effect, with lower concentrations in surrogate-peer–reared juveniles, but no sex difference. Salivary and plasma cortisol concentrations in these animals were significantly correlated.     

Recognition of heterospecific alarm vocalizations by bonnet macaques (Macaca radiata)

Recognition of heterospecific alarm vocalizations is an essential component of antipredator behavior in several prey species. The authors examined the role of learning in the discrimination of heterospecific vocalizations by wild bonnet macaques (Macaca radiata) in southern India The bonnet macaques' flight and scanning responses to playbacks of their own alarm vocalizations were compared with their responses to playbacks of vocalizations of Nilgiri langurs (Trachypithecus johnii), Hanuman langurs (Semnopithecus entellus), and sambar deer (Cervus unicolor). The study was conducted in 3 regions that differed in the frequency with which bonnet macaques encountered these species and included an urban setting. Call recognition was highest in adults and in regions where individuals were frequently exposed to the calling species; calls were not recognized by urban monkeys. Thus, age and experience are important factors in heterospecific call recognition by bonnet macaques.     

Concurrent access to two concentrations of orally delivered phencyclidine: effects of feeding conditions.

Two experiments addressed the effects of food satiation and deprivation on oral self-administration of two concurrently available phencyclidine concentrations. In the first experiment, 8 rhesus monkeys self-administered either of two concentrations of phencyclidine ("PCP, angel dust") and water under concurrent fixed-ratio 16 schedules. One concentration was always held constant (0.25 mg/mL) while a series of other phencyclidine concentrations, ranging from 0 (water) to 1.0 mg/mL, was presented in a nonsystematic order. Initially the monkeys were tested while food satiated, and the procedure was then repeated during food deprivation. The monkeys usually selected the higher concentration within the first few minutes of the session, indicating that taste and/or other immediate postingestional effects were important factors. Contrary to a number of previous reports, there were no consistent differences across subjects in the mean number of liquid deliveries or mean drug intake (mg/kg) during food satiation and deprivation. However, for all monkeys the within-session time course of responding during food satiation consistently differed from that during deprivation. A second experiment assessed whether the failure to find consistent differences in drug intake during food satiation and deprivation had been due to the history of concurrent access to different phencyclidine concentrations or to the extended experience with phencyclidine under food-satiation conditions. Six additional monkeys (Group 2) were exposed to the phencyclidine self-administration procedure (during food satiation and deprivation) for the same length of time as the monkeys in Experiment 1 (Group 1), except they received only concurrent access to phencyclidine (0.25 mg/mL) and water. Both groups then received concurrent access to phencyclidine and water during five repeated cycles of food deprivation and satiation. There were also marked individual differences in Group 2: During food satiation, 2 of the monkeys' responding increased, 1 showed no change, and 3 decreased. Examination of a number of historical variables indicated that the greater the percentage of total sessions spent during food satiation with phencyclidine available (before these experiments began), the greater the amounts of phencyclidine consumed during food satiation and the smaller the differences in phencyclidine intake when the two feeding conditions were compared.     

Monkeys exhibit prospective memory in a computerized task

Prospective memory (PM) involves forming intentions, retaining those intentions, and later executing those intended responses at the appropriate time. Few studies have investigated this capacity in animals. Monkeys performed a computerized task that assessed their ability to remember to make a particular response if they observed a PM cue embedded within an ongoing learning-set (LS) task. At a break in the LS task, monkeys selected one of two icons indicating that they had or had not encoded the occurrence of the PM cue (the latter icon resumed the LS task). Critically, during this response period, the PM response icon appeared after a delay during which monkeys could self-initiate the PM response prior to receiving any external prompt. Monkeys selected the PM and LS icons when each was the optimal response, illustrating that they could encode, store, and respond appropriately to a stimulus event in the future. Critically, some monkeys self-initiated the PM response prior to that icon's appearance, indicating that they could retrieve the PM and act on their intention to make that response without the aid of a prompt. These monkeys appeared capable of using PM in this task. Thus, this capacity appears not to be limited to humans.     

Control of responding by sound location in monkeys: rapid acquisition in darkness.

Rapid control of responding by sound location is obtained in squirrel monkeys when sound stimuli are presented from one of two loudspeakers, each one adjacent to a response key. With this arrangement of loudspeakers and response keys, squirrel monkeys quickly learn to respond on the key near the source of the sound stimulus, and this pattern is the same whether or not responses near the sound source are differentially reinforcedmthis result may depend on a pre-experimental tendency in squirrel monkeys to orient head and eyes toward a sound, which would lead the animal to look at the response key in front of the loudspeaker producing the sound. The present experiment sought to determine whether visual stimuli are necessary for rapid control of responding by sound location. Two monkeys were trained in darkness in a sound-localization task similar to that described above. Results were similar to those obtained from animals trained in light, indicating that visual stimuli are not required for rapid acquisition of sound-localization behavior in monkeys.     

I know you are not looking at me: capuchin monkeys’ (<Emphasis Type="Italic">Cebus apella</Emphasis>) sensitivity to human attentional states

The present study asked whether capuchin monkeys recognize human attentional states. The monkeys requested food from the experimenter by extending an arm (pointing) toward the baited one of two transparent cups. On regular trials the experimenter gave the food immediately to the monkeys upon pointing but on randomly inserted test trials she ignored the pointing for 5 s during which she displayed different attentional states. The monkeys looked at the experimenter's face longer when she looked at the monkeys than when she looked at the ceiling in Experiment 1, and longer when she oriented her head midway between the two cups with eyes open than when she did so with eyes closed in Experiment 2. However, the monkeys showed no differential pointing in these conditions. These results suggest that capuchins are sensitive to eye direction but this sensitivity does not lead to differential pointing trained in laboratory experiments. Furthermore, to our knowledge, this is the first firm behavioral evidence that non-human primates attend to the subtle states of eyes in a food requesting task.     

Differences in performance of young and old monkeys on a visuospatial memory task

Two groups of stumptailed macaques (Macaca arctoides) were compared in terms of performance on a visuospatial working memory task involving repeated presentations of an array of 15 doors, each of which concealed one food reward at the start of a session. Rewards obtained by the subject were not replaced. Errors were defined as responses to doors previously pushed open during the session. Both groups generally obtained all rewards and showed improved performance across sessions, but old monkeys consistently made more errors than young ones, especially in the latter half of a given session. Further, old monkeys were less diverse in the errors they made, indicating different response strategies by the two groups.     

SOME RELATIONS BETWEEN CLASSICALLY CONDITIONED AGGRESSION AND CONDITIONED SUPPRESSION IN SQUIRREL MONKEYS

During three experiments with squirrel monkeys, stimulus and shock pairings were given in the presence of a bite tube. Experiments 1 and 2 used a conditioned-suppression procedure in which bar pressing was reinforced with food. A transparent shield prevented biting of the bar. When the stimulus was paired with shock, bar pressing decreased (conditioned suppression) and tube biting increased during the stimulus (classically conditioned aggression). When the bite tube was removed on alternate sessions in Experiment 2, there was more suppression when the tube was present, thus suggesting that biting competed with bar pressing. However, this simple competing-response interpretation was complicated by the findings of Experiment 3 where, with naive monkeys, bar pressing was never reinforced with food, yet bar pressing was induced during the stimulus and was highest when the bite tube was absent. The fact that stimulus-induced bar pressing developed indicates that bar pressing in conditioned-suppression procedures, suppressed or not, may be maintained by two types of control—the food reinforcer and induced CS control. The higher rate of induced bar pressing during the stimulus with the bite tube absent confounds a simple competing response interpretation of conditioned suppression. It suggests that shock-induced responses during conditioned suppression could be both contributing to and competing with responding maintained by food, with the net effect depending on specific but ill-defined features of the situation.     

Heart rate and ratings of perceived exertion during treadmill and elliptical exercise training

Elliptical exercise has recently increased in popularity. However, little is known regarding heart rate and perceptual responses during this exercise mode even though such information is important with respect to prescribing and regulating exercise intensity. In the current study, heart rate and perceived exertion were compared between treadmill and elliptical exercise. During treadmill exercise (TM(EST)) participants estimated RPE-Overall as well as RPE-Legs and RPE-Chest. Two elliptical sessions followed: (1) RPE estimation during elliptical exercise (EL(EST)): HR (b/ min.) from TM(EST) was achieved during elliptical exercise, with participants estimating RPE-Overall, RPE-Legs, and RPE-Chest. (2) RPE production during elliptical exercise (EL(PROD)): RPE-Overall from TM(EST) was produced during elliptical exercise. There were no significant differences between modes for RPE-Overall (TM(EST): 11.2 +/- 2.2 vs EL(EST): 11.9 +/- 3.2) or RPE-Chest (TM(EST): 11.0 +/- 2.4 vs EL(EST): 11.7+/- 3.2). Mean RPE-Legs was significantly different (TM(EST): 11.2 +/- 2.4 vs EL(EST): 12.5 +/- 3.1). Heart rate was not significantly different between TM(EST) (163 +/- 16.6) and EL(PROD) (159 +/- 20.0). Analysis indicated elliptical exercise is perceived as more intense with respect to leg (RPE-Legs) exertion. Further, RPE-Overall appears effective for regulating heart rate during elliptical exercise.     

Observational learning in monkeys

Observer monkeys were housed next to demonstrator monkeys that were conditioned to respond on a multiple reinforcement schedule whose components were fixed-ratio 32, variable-interval 3-min, and extinction 5-min followed by an additional 30 sec of extinction during which every response started a new 30-sec interval. After observational periods from 113 to 210 hr long, during which observers could not perform the response and were given no extrinsic reinforcers, their first-response latencies to fixed ratio and variable interval were as short as the demonstrators; and their rates of responding were well above pre-observational baseline levels. About 8 hr later, a temporal pattern of responding appropriate to the multiple schedule emerged, including non-emission of responses during extinction. Controls lacking the chance to observe did not develop typically patterned responding after 60 hr in one case and, in two other cases, after 80 hr during which, on two occasions, every one of 50 responses was reinforced. In a second experiment, the stimulus lights associated with fixed ratio and variable interval were presented simultaneously. Subjects chose one of the schedules by responding to one of the levers beneath the lights. All subjects initially chose fixed ratio. Seeing the demonstrators switch to variable interval, due to increases in the fixed-ratio requirement, had no effect upon observers, which continued to choose fixed ratio.     

CRT信号显示的刺激率和信号率对警觉作业的影响

本实验系统地考察了在不同刺激率、信号率条件下警觉水平的变化 ,并分析了以往警觉理论的适应条件。实验一采用 5次 /分、1 5次 /分和 30次 /分的刺激率条件 ,发现检测正确率在各条件下随时间的变化是由于 d′下降所致 ,而β值基本不变 ,可以认为警觉随时间下降是由于疲劳所致 ;中等刺激率条件下的 d′最高 ,低刺激率次之 ,高刺激率最差并且高刺激率导致被试唤醒水平降低。实验二在 30次 /分的高刺激率条件下变化信号率 (分别为 3次 /分、6次 /分和 1 2次 /分 ) ,发现信号率变化不会改变 d′,但导致β值的变化 ,可以认为高信噪比导致被试的唤醒水平升高