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.     

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     

Chimpanzee (Pan troglodytes) intentional communication is not contingent upon food

Studies of great apes have revealed that they use manual gestures and other signals to communicate about distal objects. There is also evidence that chimpanzees modify the types of communicative signals they use depending on the attentional state of a human communicative partner. The majority of previous studies have involved chimpanzees requesting food items from a human experimenter. Here, these same communicative behaviors are reported in chimpanzees requesting a tool from a human observer. In this study, captive chimpanzees were found to gesture, vocalize, and display more often when the experimenter had a tool than when she did not. It was also found that chimpanzees responded differentially based on the attentional state of a human experimenter, and when given the wrong tool persisted in their communicative efforts. Implications for the referential and intentional nature of chimpanzee communicative signaling are discussed.     

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     

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.     

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.     

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.     

Use of numerical symbols by the chimpanzee (Pan troglodytes): Cardinals, ordinals, and the introduction of zero

An adult female chimpanzee with previous training in the use of Arabic numerals 1–9 was introduced to the meaning of "zero" in the context of three different numerical tasks. The first two were cardinal tasks where the subject was required either to select numerals corresponding to the number of items presented on a computer screen (productive use of numerals) or to match sets of the appropriate size to numerals presented as samples (receptive use). The third task addressed the ordinal meaning of the same symbols where the subject was required to respond to numerals sequentially, arranging them into an ascending series. The subject mastered the recognition of the meaning of zero in all three tasks. However, details of her usage of the symbol revealed that transfer of the meaning between different kinds of tasks was incomplete, suggesting that the level of abstraction characteristic of human numerical ability was not attained in the chimpanzee. Over the course of acquisition leading to the high levels of accuracy eventually observed, the newly introduced zero appeared to shift along the length of a continuous numerical scale toward the lower end, while confusions with 1 remained the most frequently encountered mistakes. Such patterns of error thus suggest that Ai's understanding of the meaning of zero in relation to the rest of the number symbols was not consistent with an "absence of items versus presence of items" scheme. 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...     

Unprompted recall and reporting of hidden objects by a chimpanzee (Pan troglodytes) after extended delays

The ability to recall features of environments not present to the senses is important in human thinking, planning, and communication, but to date there are almost no data on recall capabilities in nonverbal animals. In this study, the author used symbol knowledge as a tool to study chimpanzee memory. An 11-year-old female chimpanzee (Pan troglodytes) that had already learned a large number of arbitrarily designated geometric forms (lexigrams) watched as an experimenter hid an object in the woods outside her outdoor enclosure. The type and location of the object varied across trials. After an imposed delay of up to 16 h, the chimpanzee could interact indoors with a person who did not know that an object had been hidden, let alone the type or location of the object. A keyboard in the indoor cage displayed 256 lexigrams. From Trial 1, the chimpanzee seemed to do whatever it took to catch the person's attention and then touched the lexigram corresponding to the type of object hidden, pointed outdoors, went outdoors (if followed), and continued to point manually toward the object and vocalize until the person found the object. The subject indicated nonfood objects as well as more than 20 food types.     

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     

Addendum to "Summation in the chimpanzee (Pan troglodytes)"

Rumbaugh, Savage-Rumbaugh, and Hegel (1987) reported that two chimpanzees (Pan troglodytes) could select, with better than 90% accuracy, the greater of two paired quantities of chocolate chips. In that study, no one quantity of chocolates (from 0 through 5) was used in both pairs on a given trial. We investigated the effect of having one quantity in common (CQ) in both pairs. Whether the other quantities (OQs) of chocolates were both less than or greater than the CQ, summation still occurred. Accuracy was primarily a function of the ratios of sums to be differentiated. This finding substantiated the earlier conclusion that summation was based on both quantities of each pair and not on some simpler process such as the avoidance of the tray with the smallest single amount or selection of the tray with the single largest amount.     

Cognitive capacities for behavioural flexibility in wild chimpanzees (Pan troglodytes): the effect of snare injury on complex manual food processing

In chimpanzees, it is only in the restricted context of tool use that manual and cognitive skills have been described, comparable to those that gorillas and orang-utans display in obtaining plant foods. We report the complex food preparation skills used to eat, without tools, the leaves of the tree Broussonettia papyrifera in the Sonso community of chimpanzees at Budongo Forest, Uganda. Able-bodied individuals used multi-stage techniques that required bimanual role differentiation at several stages, and were hierarchical in organisation. A total repertoire of 14 techniques was found, with strong preference in all individuals for either of two of these; 6 additional techniques were found when flowers and leaves were eaten together. However, in this community over 20% of individuals suffer from some form of upper- or lower-limb injury as a result of snares. We investigated the manner of compensation for upper-limb injury. Only the most severely injured showed reduced feeding efficiency. Injured individuals were found to use the same repertoire of techniques as able-bodied chimpanzees. We found no evidence to suggest that injured individuals were able to develop wholly novel techniques optimal for their specific injuries, although shifts in preference for particular techniques did occur. Rather, injured individuals used novel ways of achieving particular steps in the process; by "working around" their impairments; in this way, they managed to use the same techniques as the able-bodied. Since snare injuries generally befall young animals, these results suggest that chimpanzees learn techniques partly through observational learning (of, necessarily, able-bodied individuals). Accepted after revision: 8 February 2001 Electronic Publication     

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.     

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.     

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.     

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.     

Skill mastery inhibits adoption of observed alternative solutions among chimpanzees (Pan troglodytes)

Geographic variation in socially transmitted skills and signals, similar to human culture, has been well documented for great apes. The rules governing the adoption of novel behaviours, however, are still largely unknown. We conducted an innovation-and-transmission experiment with two groups of chimpanzees living at hopE Primate Sanctuary Gänserndorf, Austria, presenting a board on which food had to be manoeuvred around obstacles to be acquired. Most chimpanzees used sticks to acquire the food, but five adults independently invented a novel technique, rattling, which was subsequently tested by almost all group members. However, individuals who had become proficient with sticks were reluctant to switch to rattling, despite it being more efficient. Similarly, after rattling was prevented, rattle specialists kept trying to rattle and made no attempt to use the stick technique, despite their knowledge about its existence. We conclude that innovators stimulate others to experiment with the solutions they display, but that chimpanzees are nevertheless conservative; mastery of a skill inhibits further exploration, and hence adoption of alternative techniques even if these are more efficient. Consequently, conformity among group members should not be expected in great apes when individuals develop proficiency at different techniques. Conservatism thus joins conformity as a mechanism to bring about cultural uniformity and stability.     

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.