Mangabeys (<Emphasis Type="Italic">Cercocebus torquatus lunulatus</Emphasis>) solve the reverse contingency task without a modified procedure

Problem solving often relies on generating new responses while inhibiting others, particularly prepotent ones. A paradigm to study inhibitory abilities is the reverse contingency task (Boysen and Berntson in J Exp Psychol Anim Behav Process 21:82–86, 1995), in which two different quantities of food are offered to an individual who receives the array he did not choose. Therefore, mastery of the task demands selecting the smaller quantity to obtain the larger one. Several non-human primates have been tested in the reverse contingency task. To date, only great apes and rhesus monkeys (Macaca mulatta) have succeeded in the original task, with no need of procedural modifications as the large-or-none contingency, correction trials or symbolic stimuli substituting for actual food quantities. Here, four mangabeys were presented with two stimulus arrays of one and four raisins in the context of the reverse contingency task. Three of them learned to perform the task well above chance without a modified procedure. They also reached above-chance performance when presented with two stimulus arrays of zero and four raisins, despite the initial difficulty of choosing a null quantity. After a period of 7–10 months, in which the animals were not tested on any task, all three subjects continued to perform well, even when presented with novel quantity pairs.     

The whole-hand point: the structure and function of pointing from a comparative perspective

Pointing by monkeys, apes, and human infants is reviewed and compared. Pointing with the index finger is a species-typical human gesture, although human infants exhibit more whole-hand pointing than is commonly appreciated. Captive monkeys and feral apes have been reported to only rarely "spontaneously" point, although apes in captivity frequently acquire pointing, both with the index finger and with the whole hand, without explicit training. Captive apes exhibit relatively more gaze alternation while pointing than do human infants about 1 year old. Human infants are relatively more vocal while pointing than are captive apes, consistent with paralinguistic use of pointing.     

Capuchin monkeys’ use of human and conspecific cues to solve a hidden object-choice task

Learning by watching others can provide valuable information with adaptive consequences, such as identifying the presence of a predator or locating a food source. The extent to which nonhuman animals can gain information by reading the cues of others is often tested by evaluating responses to human gestures, such as a point, and less often evaluated by examining responses to conspecific cues. We tested whether ten brown capuchin monkeys (Cebus [Sapajus] apella) were able to use cues from monkeys and a pointing cue from a human to obtain hidden rewards. A monkey could gain access to a reward hidden in one of two locations by reading a cue from a conspecific (e.g., reaching) or a human pointing. We then tested whether they could transfer this skill from monkeys to humans, from humans to monkeys, and from one conspecific to another conspecific. One group of monkeys was trained and tested using a conspecific as the cue-giver and was then tested with a human cue-giver. The second group of monkeys was trained and tested with a human cue-giver and was then tested with a monkey cue-giver. Monkeys that were successful with a conspecific cue-giver were also tested with a novel conspecific cue-giver. Monkeys learned to use a human point and conspecific cues to obtain rewards. Monkeys that had learned to use the cues of a conspecific to obtain rewards performed significantly better than expected by chance when they were transferred to the cues of a novel conspecific. Monkeys that learned to use a human point to obtain rewards performed significantly better than expected by chance when tested while observing conspecific cues. Some evidence suggested that transferring between conspecific cue-givers occurred with more facility than transferring across species. Results may be explained by simple rules of association learning and stimulus generalization; however, spontaneous flexible use of gestures across conspecifics and between different species may indicate capuchins can generalize learned social cues within and partially across species.     

Olive baboons communicate intentionally by pointing

A pointing gesture creates a referential triangle that incorporates distant objects into the relationship between the signaller and the gesture’s recipient. Pointing was long assumed to be specific to our species. However, recent reports have shown that pointing emerges spontaneously in captive chimpanzees and can be learned by monkeys. Studies have demonstrated that both human children and great apes use manual gestures (e.g. pointing), and visual and vocal signals, to communicate intentionally about out-of-reach objects. Our study looked at how monkeys understand and use their learned pointing behaviour, asking whether it is a conditioned, reinforcement-dependent response or whether monkeys understand it to be a mechanism for manipulating the attention of a partner (e.g. a human). We tested nine baboons that had been trained to exhibit pointing, using operant conditioning. More specifically, we investigated their ability to communicate intentionally about the location of an unreachable food reward in three contexts that differed according to the human partner’s attentional state. In each context, we quantified the frequency of communicative behaviour (auditory and visual signals), including gestures and gaze alternations between the distal food and the human partner. We found that the baboons were able to modulate their manual and visual communicative signals as a function of the experimenter’s attentional state. These findings indicate that monkeys can intentionally produce pointing gestures and understand that a human recipient must be looking at the pointing gesture for them to perform their attention-directing actions. The referential and intentional nature of baboons’ communicative signalling is discussed.     

Gaze alternation during “pointing” by squirrel monkeys (<Emphasis Type="Italic">Saimiri sciureus</Emphasis>)?

Gaze alternation (GA) is considered a hallmark of pointing in human infants, a sign of intentionality underlying the gesture. GA has occasionally been observed in great apes, and reported only anecdotally in a few monkeys. Three squirrel monkeys that had previously learned to reach toward out-of-reach food in the presence of a human partner were videotaped while the latter visually attended to the food, a distractor object, or the ceiling. Frame-by-frame video analysis revealed that, especially when reaching toward the food, the monkeys rapidly and repeatedly switched between looking at the partner’s face and the food. This type of GA suggests that the monkeys were communicating with the partner. However, the monkeys’ behavior was not influenced by changes in the partner’s focus of attention.     

An experimental social relation between two monkeys

A technique was developed for studying the reinforcement of one organism by another. Two pairs of monkeys served as subjects in adjoining but separate lever-pressing chambers. However, they were in visual, aural, and tactile contact with each other. After both pairs were trained to tolerate delays of reinforcement and one pair was trained under stimulus control to exchange reinforcements, monkey A of each pair pressed a lever to feed monkey B, and monkey B pressed to feed monkey A. The experiment sought to determine if this social interaction could be maintained. With a free responding procedure where the monkeys could work at any time in any order, the social relation proved unstable. After several oscillations in which one monkey did most of the responding and the other monkey did most of the eating, the reinforcement frequency for both pairs of animals decreased to very low levels. The final outcome would have been starvation had the experimenter not intervened.     

An assessment of memory awareness in tufted capuchin monkeys (Cebus apella)

Humans, apes, and rhesus monkeys demonstrate memory awareness by collecting information when ignorant and acting immediately when informed. In this study, five capuchin monkeys searched for food after either watching the experimenter bait one of four opaque tubes (seen trials), or not watching (unseen trials). Monkeys with memory awareness should look into the tubes before making a selection only on unseen trials because on seen trials they already know the location of the food. In Experiment 1, one of the five capuchins looked significantly more often on unseen trials. In Experiment 2, we ensured that the monkeys attended to the baiting by interleaving training and test sessions. Three of the five monkeys looked more often on unseen trials. Because monkeys looked more often than not on both trial types, potentially creating a ceiling effect, we increased the effort required to look in Experiment 3, and predicted a larger difference in the probability of looking between seen and unseen trials. None of the five monkeys looked more often on unseen trials. These findings provide equivocal evidence for memory awareness in capuchin monkeys using tests that have yielded clear evidence in humans, apes, and rhesus monkeys.     

Schedule control of the vocal behavior of Cebus monkeys

The vocal behavior of three Cebus monkeys was maintained by fixed-ratio schedules of response dependent reinforcement at values between fixed-ratio 1 and fixed-ratio 15. In one monkey that was exposed to variable-interval, fixed-interval, and conjunctive fixed-ratio fixed-interval schedules of reinforcement, vocal responding occurred at a low rate, but schedule-appropriate patterns were maintained. The rates and patterns of responding engendered indicated that the vocal operant can be brought under schedule control in the monkey by the use of response-dependent reinforcement.     

How the great apes (Pan troglodytes, Pongo pygmaeus, Pan paniscus, and Gorilla gorilla) perform on the reversed contingency task: the effects of food quantity and food visibility

S. T. Boysen and G. G. Berntson (1995) found that chimpanzees performed poorly on a reversed contingency task in which they had to point to the smaller of 2 food quantities to acquire the larger quantity. The authors compared the performance of 4 great ape species (Pan troglodytes, Pongo pygmaeus, Pan paniscus, and Gorilla gorilla) on the reversed contingency task while manipulating food quantity (0-4 or 1-4) and food visibility (visible pairs or covered pairs). Results showed no systematic species differences but large individual differences. Some individuals of each species were able to solve the reversed contingency task. Both quantity and visibility of the food items had a significant effect on performance. Subjects performed better when the disparity between quantities was smaller and the quantities were not directly visible.     

Multiple perceptual strategies used by macaque monkeys for face recognition

Successful integration of individuals in macaque societies suggests that monkeys use fast and efficient perceptual mechanisms to discriminate between conspecifics. Humans and great apes use primarily holistic and configural, but also feature-based, processing for face recognition. The relative contribution of these processes to face recognition in monkeys is not known. We measured face recognition in three monkeys performing a visual paired comparison task. Monkey and humans faces were (1) axially rotated, (2) inverted, (3) high-pass filtered, and (4) low-pass filtered to isolate different face processing strategies. The amount of time spent looking at the eyes, mouth, and other facial features was compared across monkey and human faces for each type of stimulus manipulation. For all monkeys, face recognition, expressed as novelty preference, was intact for monkey faces that were axially rotated or spatially filtered and was supported in general by preferential looking at the eyes, but was impaired for inverted faces in two of the three monkeys. Axially rotated, upright human faces with a full range of spatial frequencies were also recognized, however, the distribution of time spent exploring each facial feature was significantly different compared to monkey faces. No novelty preference, and hence no inferred recognition, was observed for inverted or low-pass filtered human faces. High-pass filtered human faces were recognized, however, the looking pattern on facial features deviated from the pattern observed for monkey faces. Taken together these results indicate large differences in recognition success and in perceptual strategies used by monkeys to recognize humans versus conspecifics. Monkeys use both second-order configural and feature-based processing to recognize the faces of conspecifics, but they use primarily feature-based strategies to recognize human faces.     

Trials per problem and age as factors in learning set formation of children

Discrimination learning set performance was examined in preschool children as a function of age and number of trials per problem. Subjects 3, 4, 5, and 6 years old were trained over three 72-trial sessions. Half of the children at each age were given problems three trials in length, and half received 12-trial problems. As predicted, younger children (3 and 4-year-olds) performed significantly better for 12 than for three trials per problem, whereas no differences in performance were evident for older children. Response pattern analyses revealed that younger children, especially those receiving less exposure within problems, produced a higher proportion of stimulus alternation and position determined response sequences than did older children. These results contradict the commonly held assumption that learning set acquisition is simply a function of the total number of trials presented and indicate that the amount of exposure to individual problems is a factor in problem solution for young human subjects.     

Treatment of food selectivity: An evaluation of video modeling of contingencies

Many children with disabilities have feeding problems including, but not limited to, food selectivity and/or food refusal. The purpose of this study was to evaluate the effectiveness of video modeling of contingencies alone and combined with direct exposure to the contingencies in the treatment of food selectivity. Treatment procedures included sequentially introducing videos in which models consumed nonpreferred foods or were exposed to differential reinforcement or differential reinforcement plus escape extinction. In addition, during feeding sessions, participants were exposed to differential reinforcement. Results indicated that video modeling of differential reinforcement plus direct exposure to differential reinforcement may be effective at increasing consumption of some nonpreferred foods, but the results were not replicated across all foods. For one participant, consumption of one food increased with video modeling alone.     

Learning by observation in rhesus monkeys

Habit memory provides us with a vast repertoire of learned rules, including stimulus-reward associations, that ensures fast and adapted decision making in daily life. Because we share this ability with monkeys, lesion and recording studies in rhesus macaques have played a key role in understanding the neural bases of individual trial-and-error habit learning. Humans, however, can learn new rules at a lower cost via observation of conspecifics. The neural properties underlying this more ecological form of habit learning remain unexplored, and it is unclear whether the rhesus macaque can be a useful model in this endeavor. We addressed this issue by testing four monkeys from the same social group in their usual semi-natural habitat using a well-established marker of habit memory, concurrent discrimination learning. Each monkey learned 24 lists of 10 object-reward associations each. For one list out of two, monkeys could observe the testing session of another member of the group prior to being tested with the same list themselves. Learning was faster for these lists than for those learned solely by trial-and-error. Errors to criterion (9/10 correct responses) were reduced by 39%, and faultless performance could be achieved for up to 5 of the 10 pairs. These data demonstrate that rhesus macaques spontaneously observe a conspecific learning new stimulus-reward associations, and substantially benefit from this observation. They ascertain that the neural underpinnings of socially-mediated forms of habit learning can be explored using the powerful tools of monkey research, including neurophysiological recordings.     

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.     

Oral self-administration of pentobarbital by rhesus monkeys: relative reinforcing effects under concurrent fixed-ratio schedules.

During daily 3-hr sessions, orally delivered pentobarbital solutions and water, or two separate pentobarbital solutions, were concurrently available to rhesus monkeys according to fixed-ratio schedules of mouth contacts with a spout. First water, and then each of four "comparison-concentration" pentobarbital solutions (0.0625, 0.25, 1, and 4 mg/mL), was successively available from one spout for a block of sessions under a fixed-ratio-64 (three monkeys) or fixed-ratio-16 (one monkey) schedule. Under an identically sized fixed-ratio schedule, deliveries of a "standard-concentration" pentobarbital solution were concurrently available from a second spout. The concentration of the standard solution remained unchanged throughout testing of the series of comparison solutions. Each of three pentobarbital concentrations (4, 1, and 0.25 mg/mL) in turn served as the standard concentration. Within each pair of concurrently available solutions, the higher drug concentration maintained more behavior than the lower concentration. Thus when monkeys were provided with concurrent access to different pentobarbital concentrations, relative reinforcing effects were directly related to drug concentration. Further, the amount of behavior maintained by a particular drug concentration was dependent on the concentration of the concurrently available drug solution. Thus, the relative effectiveness of a reinforcer in maintaining behavior is a function of both the reinforcer's magnitude and the availability of alternative reinforcers in the environment.     

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.     

Ordinal-list integration for symbolic, arbitrary, and analog stimuli by rhesus macaques (Macaca mulatta)

Two numeral-trained monkeys learned to produce 3 5-item lists of Arabic numerals, colors, and arbitrary signs in the correct sequence. The monkeys then responded at above-chance levels when the authors tested them with nonrewarded pair-wise comparisons of items from different lists, indicating their use of ordinal-position information. The authors also tested the monkeys with nonrewarded pair-wise comparisons of an analog quantity and an item from 1 of the 3 learned lists. Although the monkeys were not trained to serially order analog quantities, 1 monkey correctly integrated the analog quantities with the lists of numerals, colors, and signs. The consistent use of an ordinal rule, despite different types of training and varying degrees of experience with the 4 types of stimuli, suggested that the monkey had a robust concept of ordinality.     

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.     

Squirrel monkeys (Saimiri sciureus) choose smaller food arrays: long-term retention, choice with nonpreferred food, and transposition

Squirrel monkeys (Saimiri sciureus) that had learned to reach toward 1 piece of food instead of 4 in a reverse-reward contingency were tested after an 8-month delay with no intervening relevant experiences. All monkeys except 1 continued to show inhibitory control by reliably reaching toward the smaller quantity, most of them doing so within 2 sessions. Performance was maintained when a low-preference food replaced prized foods as arrays and rewards. When the quantities 1 and 4 were replaced with different ones, there was strong evidence of transposition at group level, although individual differences in bias toward the smaller quantities became apparent. Individual differences in mastering the original task more than 8 months previously were quite stable, suggesting robustness in the operations required for this form of self-control.     

What meaning means for same and different: Analogical reasoning in humans (Homo sapiens), chimpanzees (Pan troglodytes), and rhesus monkeys (Macaca mulatta)

Thus far, language- and token-trained apes (e.g., D. Premack, 1976; R. K. R. Thompson, D. L. Oden, & S. T. Boysen, 1997) have provided the best evidence that nonhuman animals can solve, complete, and construct analogies, thus implicating symbolic representation as the mechanism enabling the phenomenon. In this study, the authors examined the role of stimulus meaning in the analogical reasoning abilities of three different primate species. Humans (Homo sapiens), chimpanzees (Pan troglodytes), and rhesus monkeys (Macaca mulatta) completed the same relational matching-to-sample (RMTS) tasks with both meaningful and nonmeaningful stimuli. This discrimination of relations-between-relations serves as the basis for analogical reasoning. Meaningfulness facilitated the acquisition of analogical matching for human participants, whereas individual differences among the chimpanzees suggest that meaning can either enable or hinder their ability to complete analogies. Rhesus monkeys did not succeed in the RMTS task regardless of stimulus meaning, suggesting that their ability to reason analogically, if present at all, may be dependent on a dimension other than the representational value of stimuli.