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     

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.     

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.     

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.     

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...     

An infant chimpanzee (Pan troglodytes) follows human gaze

The ability of non-human primates to follow the gaze of other individuals has recently received much attention in comparative cognition. The aim of the present study was to investigate the emergence of this ability in a chimpanzee infant. The infant was trained to look at one of two objects, which an experimenter indicated by one of four different cue conditions: (1) tapping on the target object with a finger; (2) pointing to the target object with a finger; (3) gazing at the target object with head orientation; or (4) glancing at the target object without head orientation. The subject was given food rewards independently of its responses under the first three conditions, so that its responses to the objects were not influenced by the rewards. The glancing condition was tested occasionally, without any reinforcement. By the age of 13 months, the subject showed reliable following responses to the object that was indicated by the various cues, including glancing alone. Furthermore, additional tests clearly showed that the subject's performance was controlled by the "social" properties of the experimenter-given cues but not by the non-social, local-enhancing peripheral properties. Electronic Publication     

Development of combinatory manipulation in chimpanzee infants (Pan troglodytes)

I made systematic observations of three infant chimpanzees aged 2–4 years, who participated in a series of diagnostic tests of combinatory manipulation. The tasks were stacking blocks, seriating nesting cups, and inserting an object into the corresponding hole in a plate or a box. These tasks were originally devised for developmental diagnosis of human infants. The chimpanzee infants displayed combinatory manipulation comparable to that of 1-year-old human infants. Common motor characteristics were observed across the tasks, namely "repetition" of actions, "adjustment" of actions, "reversal" of actions, and "shifts" of attention. Humans and chimpanzees share these actions when manipulating multiple objects to complete a task. Repetition, adjustment, and reversal of actions and shifts of attention underlie higher levels of cognition common to both species. Accepted after revision: 5 May 2001 Electronic Publication     

Chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) quantify split solid objects

Recent research suggests that gorillas’ and orangutans’ object representations survive cohesion violations (e.g., a split of a solid object into two halves), but that their processing of quantities may be affected by them. We assessed chimpanzees’ (Pan troglodytes) and bonobos’ (Pan paniscus) reactions to various fission events in the same series of action tasks modelled after infant studies previously run on gorillas and orangutans (Cacchione and Call in Cognition 116:193–203, 2010b). Results showed that all four non-human great ape species managed to quantify split objects but that their performance varied as a function of the non-cohesiveness produced in the splitting event. Spatial ambiguity and shape invariance had the greatest impact on apes’ ability to represent and quantify objects. Further, we observed species differences with gorillas performing lower than other species. Finally, we detected a substantial age effect, with ape infants below 6 years of age being outperformed by both juvenile/adolescent and adult apes.     

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.     

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.     

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.     

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.     

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.     

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     

Sequential responding to arabic numerals with wild cards by the chimpanzee (Pan troglodytes)

One adult female chimpanzee (Pan troglodytes) was trained to respond serially to three arabic numerals between 1 and 9, presented on a cathode-ray-tube (CRT) screen. To examine the factors affecting her sequential responding behavior, wild-card items were added to the three-item sequences. When this wild-card item remained until the subject responded to the last numeral (i.e., the terminator condition), her response to the terminator at each point of the sequence was controlled by the ordinal distance between numerals. Thus, the number of responses to the terminator increased as the ordinal distance between numerals increased. When the wild-card item was eliminated by the subject’s response (wild-card conditions), the probability of responses to the wild card before the first numeral increased as a function of the serial position of the first numeral. These results were consistent with previous studies of response time and suggest both serial position and symbolic distance effects. It is suggested that the subject might form the integrated 9-item linear representations by training of possible subsets of three-item sequences. Knowledge concerning the ordinal position of each numeral was established through this training. Received: 27 October 1999 / Accepted: 22 November 1999     

Spatial memory and monitoring of hidden items through spatial displacements by chimpanzees (Pan troglodytes)

This study examined chimpanzee (Pan troglodytes) short-term memory for food location in near space. In Experiments 1 and 2, either 1 or 2 items (chocolate pieces) were hidden in an array of 3 or 5 containers that either remained stationary or were rotated 180 degrees or 360 degrees. When the array remained stationary, the chimpanzees remembered both item locations. When arrays were rotated, however, chimpanzees found only 1 item. In Experiment 3, 2 items were hidden in an array of 7 cups. Both items were found at levels significantly better than chance. Ninety percent of errors were made after the 1st item was found, and errors reflected memory failure rather than a failure of inhibitory control.     

Chimpanzees (Pan troglodytes) remember the location of a hidden food item after altering their orientation to a spatial array

Two chimpanzees (Pan troglodytes) had a direct view of an experimenter placing a food item beneath one of several cups within a horizontal spatial array. The chimpanzees then were required to move around the spatial array, shifting their orientation to the array by 180 degrees . Both chimpanzees remembered the location of the food item. In the next experiment, a visual barrier was placed between the chimpanzees and the spatial array after the food item had been hidden to prevent visual tracking of the location of the object during the chimpanzees' movement. One chimpanzee remembered the location of the hidden item in this variation. These results demonstrate another capacity for spatial memory in this species that complements data indicating chimpanzee spatial memory for invisible displacements, array rotations, and array transpositions.     

Referential communication by chimpanzees (Pan troglodytes)

Two experiments were conducted to assess the referential function of chimpanzee (Pan troglodytes) gestures to obtain food. The chimpanzees received 1 trial per condition. In Experiment 1 (N = 101), in full view of the chimpanzee, a banana was placed on top of 1 of 2 inverted buckets or was hidden underneath 1 of the buckets. In Experiment 2 (N = 35), 4 conditions were presented in constant order: (a) no food, no observer; (b) no food, observer present; (c) food present, no observer; and (d) food present, observer present. Gestures and visual orienting were used socially and referentially. The capacity for nonverbal reference may predate the Hominidae-Pongidae split, and the development of nonverbal reference may be independent of human species-specific adaptations for speech.     

Scale-model comprehension by chimpanzees (Pan troglodytes)

The ability of chimpanzees (Pan troglodytes) to recognize the correspondence between a scale model and its real-world referent was examined. In Experiments 1 and 2, an adult female and a young adult male watched as an experimenter hid a miniature model food in 1 of 4 sites in a scale model. Then, the chimpanzees were given the opportunity to find the real food item that had been hidden in the analogous location in the real room. The female performed significantly above chance, whereas the male performed at chance level. Experiments 3 and 4 tested 5 adult and 2 adolescent chimpanzees in a similar paradigm, using a scale model of the chimpanzees' outdoor area. Results indicate that some adult chimpanzees were able to reliably demonstrate an understanding of the relationship between a scale model and the larger space it represented, whereas other subjects were constrained by inefficient and unsuccessful search patterns.