The generalization of deferred imitation in enculturated chimpanzees (Pan troglodytes)

Deferred imitation of object-related actions and generalization of imitation to similar but not identical tasks was assessed in three human-reared (enculturated) chimpanzees, ranging in age from 5 to 9 years. Each ape displayed high levels of deferred imitation and only slightly lower levels of generalization of imitation. The youngest two chimpanzees were more apt to generalize the model's actions when they had displayed portions of the target behaviors at baseline, consistent with the idea that learning is more likely to occur when working within the "zone of proximal development." We argue that generalization of imitation is the best evidence to date of imitative learning in chimpanzees. Electronic Publication     

Summation in the chimpanzee (Pan troglodytes)

In this research, we asked whether 2 chimpanzee (Pan troglodytes) subjects could reliably sum across pairs of quantities to select the greater total. Subjects were allowed to choose between two trays of chocolates. Each tray contained two food wells. To select the tray containing the greater number of chocolates, it was necessary to sum the contents of the food wells on each tray. In experiments where food wells contained from zero to four chocolates, the chimpanzees chose the greater value of the summed wells on more than 90% of the trials. In the final experiment, the maximum number of chocolates assigned to a food well was increased to five. Choice of the tray containing the greater sum still remained above 90%. In all experiments, subjects reliably chose the greater sum, even though on many trials a food well on the "incorrect" tray held more chocolates than either single well on the "correct" tray. It was concluded that without any known ability to count, these chimpanzees used some process of summation to combine spatially separated quantities. Speculation regarding the basis for summation includes consideration of perceptual fusion of pairs of quantities and subitization.     

Numerical competence in a chimpanzee (Pan troglodytes)

A chimpanzee (Pan troglodytes), trained to count foods and objects by using Arabic numbers, demonstrated the ability to sum arrays of 0-4 food items placed in 2 of 3 possible sites. To address representational use of numbers, we next baited sites with Arabic numbers as stimuli. In both cases performance was significantly above chance from the first sessions, which suggests that without explicit training in combining arrays, the animal was able to select the correct arithmetic sum for arrays of foods or Arabic numbers under novel test conditions. These findings demonstrate that counting strategies and the representational use of numbers lie within the cognitive domain of the chimpanzee and compare favorably with the spontaneous use of addition algorithms demonstrated in preschool children.     

Acquisition of navigation by chimpanzees (Pan troglodytes) in an automated fingermaze task

These experiments investigated how chimpanzees learn to navigate visual fingermazes presented on a touch monitor. The aim was to determine whether training the subjects to solve several different mazes would establish a generalized map-reading skill such that they would solve new mazes correctly on the first presentation. In experiment 1, two captive adult female chimpanzees were trained to move a visual object (a ball) with a finger over the monitor surface toward a target through a grid of obstacles that formed a maze. The task was fully automated with storage of movement paths on individual trials. Training progressed from very simple mazes with one obstacle to complex mazes with several obstacles. The subjects learned to move the ball to the target in a curved path so as to avoid obstacles and blind alleys. After training on several mazes, both subjects developed a high level of efficiency in moving the ball to the target in a path that closely approached the ideal shortest path. New mazes were then presented to determine whether the subjects had acquired a more generalized maze-solving performance. The subjects solved 65–100% of the new mazes the first time they were presented by moving the ball around obstacles to the target without making detours into blind alleys. In experiment 2, one of the chimpanzees was trained using mazes with two routes to the target. One of the routes was blocked at one of many possible locations. After training to avoid the blind alley in different mazes, new mazes were presented that also had one route blocked. The subject correctly solved 90.7% of the novel mazes. When the mazes had one short and one long open route to the target the subject preferred the shorter route. When the short route was blocked, the subject solved only 53.3% of the mazes because of the preference for the shorter route even when blocked. The overall results suggest that with the training methods used the subjects learned to solve specific mazes with a trial-and-error method. Although both subjects were able to solve many of the novel mazes they did not fully develop a more general "map-reading" skill. Accepted after revision: 30 July 2001 Electronic Publication     

Discrimination and categorization of photographs of natural objects by chimpanzees (Pan troglodytes)

Two experiments assessed the ability of four adult female chimpanzees to categorize natural objects. Chimpanzees were initially trained to match different color photographs of familiar objects from four possible categories. In training, all the comparison stimuli were from the same category in one condition, and from different categories in another condition. For all subjects, training performance was consistently better for the "different category" than for the "same category" trials. Probe trials were shown after training. In probe trials, the sample and positive comparison stimuli were different items from the same category, and the foils were selected from among the three other test categories. Individual performance was above chance in probe trials, suggesting that categorization by chimpanzees may transcend perceptual resemblance. These results were later replicated with novel stimulus items from the same four categories (experiment 2). Altogether, this research demonstrates that chimpanzees grouped perceptually different exemplars within the same category, and further suggests that these animals formed conceptual representations of the categories. Accepted after revision: 10 August 2001 Electronic Publication     

Prospective memory in a language-trained chimpanzee (Pan troglodytes)

Prospective memory involves the encoding, retention, and implementation of an intended future action. Although humans show many forms of prospective memory, less is known about the future oriented processes of nonhuman animals, or their ability to use prospective memory. In this experiment, a chimpanzee named Panzee, who had learned to associate geometric forms called lexigrams with real-world referents, was given a prospective memory test. Panzee selected between two foods the one she wanted to receive more immediately. That food was scattered in an outdoor yard where she could forage for it. Also outdoors were lexigram tokens, one of which represented the food item that remained indoors throughout a 30 min period, and that could be obtained if Panzee brought in the token that matched that food item. After foraging for the selected food item, Panzee consistently remembered to retrieve and return the correct token when food was available indoors, whereas on control trials involving no indoor food she rarely returned a token. This indicated that Panzee encoded information relevant to the future action of token retrieval after extended delays for one type of food, even when a more immediately preferred food was available.     

Imitation recognition in a captive chimpanzee (Pan troglodytes)

This study investigated the ability of a captive chimpanzee (Pan troglodytes) to recognise when he is being imitated. In the experimental condition of test 1a, an experimenter imitated the postures and behaviours of the chimpanzee as they were being displayed. In three control conditions the same experimenter exhibited (1) actions that were contingent on, but different from, the actions of the chimpanzee, (2) actions that were not contingent on, and different from, the actions of the chimpanzee, or (3) no action at all. The chimpanzee showed more "testing" sequences (i.e., systematically varying his actions while oriented to the imitating experimenter) and more repetitive behaviour when he was being imitated, than when he was not. This finding was replicated 4 months later in test 1b. When the experimenter repeated the same actions she displayed in the experimental condition of test 1a back to the chimpanzee in test 2, these actions now did not elicit those same testing sequences or repetitive behaviours. However, a live imitation condition did. Together these results provide the first evidence of imitation recognition in a nonhuman animal.     

The use of tokens as rewards and tools by chimpanzees (Pan troglodytes)

This paper explores the effectiveness of token rewards in maintaining chimpanzees (Pan troglodytes) in working at intellectually costly tasks, and studies the "saving" behavior of the subjects, investigating the factors that can condition it. Two experiments were run. Tokens were introduced as rewards in a matching-to-sample task and used as exchange tools for food by three adult female chimpanzees. Subjects' performances were maintained at constant high levels of accuracy, suggesting that the tokens were almost equivalent to direct food rewards. The results also showed the emergence of saving behavior. The subjects spontaneously saved the tokens during the matching-to-sample task before exchanging them for food. The chimpanzees also learned a new symbolic discrimination task, with tokens as the reward. During this learning process a rarely reported phenomenon emerged: one of the subjects showed symmetry, a form of stimulus equivalence. Accepted after revision: 7 August 2001 Electronic Publication     

Post-event misinformation effects in a language-trained chimpanzee (Pan troglodytes)

Animal Cognition - Errors of source monitoring are widespread human memory challenges, and our memories are subject to distortion upon the presentation of subsequent misinformation. Less is known...     

Perception of shape from shading in chimpanzees (Pan troglodytes) and humans (Homo sapiens)

The perception of shape from shading was tested in two chimpanzees (Pan troglodytes) and five humans (Homo sapiens), using visual search tasks. Subjects were required to select and touch an odd item (target) from among uniform distractors. Humans found the target faster when shading was vertical than when it was horizontal, consistent with results of previous research. Both chimpanzees showed the opposite pattern: they found the target faster when shading was horizontal. The same difference in response was found in texture segregation tasks. This difference between the species could not be explained by head rotation or head shift parallel to the surface of the monitor. Furthermore, when the shaded shape was changed from a circle to a square, or the shading type was changed from gradual to stepwise, the difference in performance between vertical and horizontal shading disappeared in chimpanzees, but persisted in humans. These results suggest that chimpanzees process shading information in a different way from humans. Received: 20 January 1998 / Accepted after revision: 30 March 1998     

Causal knowledge and imitation/emulation switching in chimpanzees (Pan troglodytes) and children (Homo sapiens)

This study explored whether the tendency of chimpanzees and children to use emulation or imitation to solve a tool-using task was a response to the availability of causal information. Young wild-born chimpanzees from an African sanctuary and 3- to 4-year-old children observed a human demonstrator use a tool to retrieve a reward from a puzzle-box. The demonstration involved both causally relevant and irrelevant actions, and the box was presented in each of two conditions: opaque and clear. In the opaque condition, causal information about the effect of the tool inside the box was not available, and hence it was impossible to differentiate between the relevant and irrelevant parts of the demonstration. However, in the clear condition causal information was available, and subjects could potentially determine which actions were necessary. When chimpanzees were presented with the opaque box, they reproduced both the relevant and irrelevant actions, thus imitating the overall structure of the task. When the box was presented in the clear condition they instead ignored the irrelevant actions in favour of a more efficient, emulative technique. These results suggest that emulation is the favoured strategy of chimpanzees when sufficient causal information is available. However, if such information is not available, chimpanzees are prone to employ a more comprehensive copy of an observed action. In contrast to the chimpanzees, children employed imitation to solve the task in both conditions, at the expense of efficiency. We suggest that the difference in performance of chimpanzees and children may be due to a greater susceptibility of children to cultural conventions, perhaps combined with a differential focus on the results, actions and goals of the demonstrator.     

Cardiac correlates of individual recognition in the chimpanzee (Pan troglodytes)

The ability of a chimpanzee to recognize individuals depicted in photographs was evaluated through the use of heart rate measures. Heart rate was recorded before and during photographic projections of human caregivers, familiar individuals, strangers, and blank control slides. In the absence of explicit training or reinforcement, the chimpanzee displayed a differential pattern of heart rate response to the stimulus categories. Although heart rate responses to all stimuli were predominantly deceleratory, the photographs of caregivers yielded consistently larger responses than other stimuli. Results indicate that the chimpanzee is able to recognize individual humans from novel photographic representations and that heart rate can provide an objective measure of such recognition.     

Use of gesture sequences in captive chimpanzee (Pan troglodytes) play

This study examined the use of sensory modalities relative to a partner’s behavior in gesture sequences during captive chimpanzee play at the Chimpanzee and Human Communication Institute. We hypothesized that chimpanzees would use visual gestures toward attentive recipients and auditory/tactile gestures toward inattentive recipients. We also hypothesized that gesture sequences would be more prevalent toward unresponsive rather than responsive recipients. The chimpanzees used significantly more auditory/tactile rather than visual gestures first in sequences with both attentive and inattentive recipients. They rarely used visual gestures toward inattentive recipients. Auditory/tactile gestures were effective with and used with both attentive and inattentive recipients. Recipients responded significantly more to single gestures than to first gestures in sequences. Sequences often indicated that recipients did not respond to initial gestures, whereas effective single gestures made more gestures unnecessary. The chimpanzees thus gestured appropriately relative to a recipient’s behavior and modified their interactions according to contextual social cues.     

Chimpanzees (Pan troglodytes) and the question of cumulative culture: an experimental approach

There is increasing evidence for cultural variations in behaviour among non-human species, but human societies additionally display elaborate cumulative cultural evolution, with successive generations building on earlier achievements. Evidence for cumulative culture in non-human species remains minimal and controversial. Relevant experiments are also lacking. Here we present a first experiment designed to examine chimpanzees' capacity for cumulative social learning. Eleven young chimpanzees were presented with a foraging device, which afforded both a relatively simple and a more complex tool-use technique for extracting honey. The more complex 'probing' technique incorporated the core actions of the simpler 'dipping' one and was also much more productive. In a baseline, exploration condition only two subjects discovered the dipping technique and a solitary instance of probing occurred. Demonstrations of dipping by a familiar human were followed by acquisition of this technique by the five subjects aged three years or above, whilst younger subjects showed a significant increase only in the elements of the dipping technique. By contrast, subsequent demonstrations of the probing task were not followed by acquisition of this more productive technique. Subjects stuck to their habitual dipping method despite an escalating series of demonstrations eventually exceeding 200. Supplementary tests showed this technique is within the capability of chimpanzees of this age. We therefore tentatively conclude that young chimpanzees exhibit a tendency to become 'stuck' on a technique they initially learn, inhibiting cumulative social learning and possibly constraining the species' capacity for cumulative cultural evolution.     

Simian scribbles: a reappraisal of drawing in the chimpanzee (Pan troglodytes)

Three chimpanzees (Pan troglodytes) were provided with 18 different stimulus pages for drawing. The resulting 618 drawings were coded for drawn marks, and results were compared with early reports on ape drawing (Morris, 1962; Schiller, 1951) and with more recent systematic studies (Smith, 1973). The findings of the present study confirm Smith's observations of a tendency for the animals to draw closer to the center and toward the bottom of the page. No evidence for perceptual balancing was observed in drawings produced on pages with lateral figures, nor was any evidence for closure apparent with complex figures. The present results agree with earlier findings that chimpanzees will engage in drawing activities without training or reinforcement, and this behavior may reflect their intrinsic interest in exploratory and manipulative play.     

Correlation between menstrual cycle and cognitive performance in a chimpanzee (Pan troglodytes)

Extensive research on human subjects has tried to investigate whether there is a correlation between cognitive performance and the menstrual cycle. Less is known about the relationship between the menstrual cycle and task performance in other cognitive animals. We test whether the secretion of a sex hormone [luteinizing hormone(LH)] influences the performance of cognitive tasks by a female chimpanzee (Pan troglodytes) who is part of a long-term cognition research program. We focus on two cognitive tasks: an "easy task," which consists of simple numerical ordering, and a "difficult task," which combines numerical ordering with memorizing the numerals' spatial location. Data on the performance of these cognitive tasks, urine samples, and sexual swelling over six menstrual cycles showed that the chimpanzee's performance accuracy decreased and that the intertrial interval was longer during the LH-surge of the menstrual cycle, but only for the performance of the difficult task. These performance attributes seem to reflect a decrease in attention or motivation during ovulation. In summary, the cognitive performance of a chimpanzee was disturbed by hormonal changes despite her long-term experience in the tasks.     

Tactics to obtain a hidden food item in chimpanzee pairs (Pan troglodytes)

Five dyads of chimpanzees were tested in a competitive situation, as a pilot study to examine chimpanzees' understanding of conspecifics' knowledge. A human experimenter baited one of five containers in an outdoor enclosure. Chimpanzee A (witness) could see where the food was hidden, while chimpanzee B (witness-of-witness) could not see the baited place but could observe the chimpanzee A watching the food being hidden. Then the two were released into the enclosure. This procedure was repeated for a certain number of days along with a control condition in which neither could see the baited location. The witness-of-witness developed tactics to forestall the witness in two pairs. The witness misled the witness-of-witness by taking a route to an empty container in several cases. These episodes might represent examples of deception. Tactics and counter-tactics thus developed through the interaction between the witness and the witness-of-witness, illustrating the high social intelligence of chimpanzees. An examination of the changes in tactics suggests a possibility that the witness-of-witness understands the witness's knowledge of the location of hidden food. Accepted after revision: 22 May 2001 Electronic Publication     

Conspecific recognition in the chimpanzee (Pan troglodytes): cardiac responses to significant others

The ability of a chimpanzee (Pan troglodytes) to recognize photographs of conspecifics was evaluated with heart-rate measures. Heart rate was recorded before, during, and after viewing photographs of an aggressive chimpanzee, a friendly companion animal, and an unfamiliar chimpanzee. The subject displayed a differential pattern of heart-rate response to the stimulus animals, without prior experience with the photographs. Responses to the aggressive animal were acceleratory, which suggests a defensive response. Respiratory sinus arrhythmia suggested that this response was associated with sympathetic activation. In contrast, responses to the familiar animal were minimal, whereas cardiac deceleration was observed in response to the strange chimp, which likely reflects an orienting response. Results suggest the chimpanzee is able to recognize photographs of individual conspecifics and that heart-rate change can reflect the nature of established social relationships between chimpanzees.     

Matching visual stimuli on the basis of global and local features by chimpanzees (Pan troglodytes) and rhesus monkeys (Macaca mulatta)

This study was designed to examine whether chimpanzees and monkeys exhibit a global-to-local precedence in the processing of hierarchically organized compound stimuli, as has been reported for humans. Subjects were tested using a sequential matching-to-sample paradigm using stimuli that differed on the basis of their global configuration or local elements, or on both perceptual attributes. Although both species were able to discriminate stimuli on the basis of their global configuration or local elements, the chimpanzees exhibited a global-to-local processing strategy, whereas the rhesus monkeys exhibited a local-to-global processing strategy. The results suggest that perceptual and attentional mechanisms underlying information-processing strategies may account for differences in learning by primates. Accepted after revision: 11 September 2001 Electronic Publication     

Enumeration of briefly presented items by the chimpanzee (Pan troglodytes) and humans (Homo sapiens)

In this study, we compared the performances on an enumeration task (numerical labeling task) of 1 chimpanzee (Pan troglodytes) and 4 humans. In this task, two types of trials, with different exposure durations of the sample that was to be enumerated, were used. In the unlimited-exposure trials, the sample remained on until the subject made a choice. In the brief-exposure trials, the sample was presented for 100 msec and then was masked. The results show clear differences between the different species. The main differences had to do with accuracy during the unlimited trials and response times during the brief trials. Detailed analyses of the pattern of response times for the chimpanzee and of looking-back behavior during the task suggests that the enumeration process underlying the subject's performance was not counting but estimation.