"Constructive" enumeration by chimpanzees (Pan troglodytes) on a computerized task

Two chimpanzees used a joystick to collect dots, one at a time, on a computer monitor (see video-clip in the electronic supplementary material), and then ended a trial when the number of dots collected was equal to the Arabic numeral presented for the trial. Both chimpanzees performed substantially and reliably above chance in collecting a quantity of dots equal to the target numeral, one chimpanzee for the numerals 1–7, and the second chimpanzee for the numerals 1–6. Errors that were made were seldom discrepant from the target by more than one dot quantity, and the perceptual process subitization was ruled out as an explanation for the performance. Additionally, analyses of trial duration data indicated that the chimpanzees were responding based on the numerosity of the constructed set rather than on the basis of temporal cues. The chimpanzees' decreasing performance with successively larger target numerals, however, appeared to be based on a continuous representation of magnitude rather than a discrete representation of number. Therefore, chimpanzee counting in this type of experimental task may be a process that represents magnitudes with scalar variability in that the memory for magnitudes associated with each numeral is imperfect and the variability of responses increases as a function of the numeral's value. Accepted after revision: 11 June 2001 Electronic Publication     

A reversed-reward contingency task reveals causal knowledge in chimpanzees (Pan troglodytes)

In the reversed-reward contingency task, subjects are required to choose the less preferred of two options in order to obtain the more preferred one. Usually, this task is used to measure inhibitory skills, but it could also be used to measure how strong the subjects’ preferences are. We presented chimpanzees with support tasks where only one of two paper strips could physically bring food into reach. Subjects were rewarded for choosing the non-functional strip. In Experiment 1, subjects failed to pick the non-baited strip. In Experiment 2, subjects failed to pick the broken strip. Chimpanzees performed worse in these tasks than in other similar tasks where instead of paper strips, there were similar shapes painted on a platform. The fact that subjects found the reversed-reward contingency task based on causality more difficult to solve than a perceptually similar task with no causality involved (i.e., arbitrary) suggests that they did not treat real strips as an arbitrary task. Instead, they must have had some causal knowledge of the support problem that made them prefer functional over non-functional strips despite the contrary reward regime.     

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     

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) do not develop contingent reciprocity in an experimental task

Chimpanzees provide help to unrelated individuals in a broad range of situations. The pattern of helping within pairs suggests that contingent reciprocity may have been an important mechanism in the evolution of altruism in chimpanzees. However, correlational analyses of the cumulative pattern of interactions over time do not demonstrate that helping is contingent upon previous acts of altruism, as required by the theory of reciprocal altruism. Experimental studies provide a controlled approach to examine the importance of contingency in helping interactions. In this study, we evaluated whether chimpanzees would be more likely to provide food to a social partner from their home group if their partner had previously provided food for them. The chimpanzees manipulated a barpull apparatus in which actors could deliver rewards either to themselves and their partners or only to themselves. Our findings indicate that the chimpanzees’ responses were not consistently influenced by the behavior of their partners in previous rounds. Only one of the 11 dyads that we tested demonstrated positive reciprocity. We conclude that contingent reciprocity does not spontaneously arise in experimental settings, despite the fact that patterns of behavior in the field indicate that individuals cooperate preferentially with reciprocating partners.     

Chimpanzees (Pan troglodytes) transfer tokens repeatedly with a partner to accumulate rewards in a self-control task

There has been extensive research investigating self-control in humans and nonhuman animals, yet we know surprisingly little about how one’s social environment influences self-control. The present study examined the self-control of chimpanzees in a task that required active engagement with conspecifics. The task consisted of transferring a token back and forth with a partner animal in order to accumulate food rewards, one item per token transfer. Self-control was required because at any point in the trial, either chimpanzee could obtain their accumulated rewards, but doing so discontinued the food accumulation and ended the trial for both individuals. Chimpanzees readily engaged the task and accumulated the majority of available rewards before ending each trial, and they did so across a number of conditions that varied the identity of the partner, the presence/absence of the experimenter, and the means by which they could obtain rewards. A second experiment examined chimpanzees’ self-control when given the choice between immediately available food items and a potentially larger amount of rewards that could be obtained by engaging the token transfer task with a partner. Chimpanzees were flexible in their decision-making in this test, typically choosing the option representing the largest amount of food, even if it involved delayed accumulation of the rewards via the token transfer task. These results demonstrate that chimpanzees can exhibit self-control in situations involving social interactions, and they encourage further research into this important aspect of the self-control scenario.     

Facial perception of conspecifics: chimpanzees (Pan troglodytes) preferentially attend to proper orientation and open eyes

This paper reports on the use of an eye-tracking technique to examine how chimpanzees look at facial photographs of conspecifics. Six chimpanzees viewed a sequence of pictures presented on a monitor while their eye movements were measured by an eye tracker. The pictures presented conspecific faces with open or closed eyes in an upright or inverted orientation in a frame. The results demonstrated that chimpanzees looked at the eyes, nose, and mouth more frequently than would be expected on the basis of random scanning of faces. More specifically, they looked at the eyes longer than they looked at the nose and mouth when photographs of upright faces with open eyes were presented, suggesting that particular attention to the eyes represents a spontaneous face-scanning strategy shared among monkeys, apes, and humans. In contrast to the results obtained for upright faces with open eyes, the viewing times for the eyes, nose, and mouth of inverted faces with open eyes did not differ from one another. The viewing times for the eyes, nose, and mouth of faces with closed eyes did not differ when faces with closed eyes were presented in either an upright or inverted orientation. These results suggest the possibility that open eyes play an important role in the configural processing of faces and that chimpanzees perceive and process open and closed eyes differently.     

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.     

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.     

Imitation in neonatal chimpanzees (Pan troglodytes)

This paper provides evidence for imitative abilities in neonatal chimpanzees (Pan troglodytes), our closest relatives. Two chimpanzees were reared from birth by their biological mothers. At less than 7 days of age the chimpanzees could discriminate between, and imitate, human facial gestures (tongue protrusion and mouth opening). By the time they were 2 months old, however, the chimpanzees no longer imitated the gestures. They began to perform mouth opening frequently in response to any of the three facial gestures presented to them. These findings suggest that neonatal facial imitation is most likely an innate ability, developed through natural selection in humans and in chimpanzees. The relationship between the disappearance of neonatal imitation and the development of social communicative behavior is discussed from an evolutionary perspective.     

Gravity bias in young and adult chimpanzees (Pan troglodytes): tests with a modified opaque-tubes task

Young human children at around 2 years of age fail to predict the correct location of an object when it is dropped from the top of an S-shape opaque tube. They search in the location just below the releasing point (Hood, 1995). This type of error, called a 'gravity bias', has recently been reported in dogs and monkeys. In the present study, we investigated whether young and adult chimpanzees also show such a gravity bias in a modified version of the original opaque-tube task. The original task by Hood and colleagues required the subject to search in a location after the object had fallen, while in the task reported here, subjects were required to predict the location before the object was dropped. Thus the present procedure does not involve explicit invisible displacement operations, one of the important components of the original procedure. In Experiment 1 both young (1.5-2.5-year-old) and adult chimpanzees predicted the location of falling food items below the releasing point even when crossed tubes were used. These gravity errors remained after the extensive experience of using the tubes themselves. Experiment 2 further tested adult and 4-year-old chimpanzees under the set-up in which the straight and crossed tubes were simultaneously presented. The results were the same as those in the previous test, suggesting that developmental changes and learning effect do not affect the gravity bias in chimpanzees.     

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...     

The effect of response contingencies on scale model task performance by chimpanzees (Pan troglodytes)

The effects of modified procedures on chimpanzees' (Pan troglodytes) performance in a scale model comprehension task were examined. Seven chimpanzees that previously participated in a task in which they searched an enclosure for a hidden item after watching an experimenter hide a miniature item in the analogous location in a scale model were retested under procedures incorporating response costs. In Experiment 1, chimpanzees were trained under procedures that rewarded only item retrievals occurring on the 1st search attempt. During test trials, 6 chimpanzees performed above chance, including 4 that were previously unsuccessful under the original procedures (V. A. Kuhlmeier, S. T. Boysen, & K. L. Mukobi, 1999). Experiment 2 compared performance under the new and original procedures. Results indicated that for some chimpanzees, performance depended on procedures that decreased the use of competing search strategies and encouraged strategies based on information from the scale model.     

Dominance assertion in male chimpanzees (Pan troglodytes)

The social relations of three adult males in a captive chimpanzee group were evaluated for 12 months. The observations encompassed a period of transition in the dominance hierarchy providing an opportunity to assess behavioral patterns involved in the initiation of status changes. This study shows that dominance in the chimpanzee is expressed primarily through ritualized and non-contact behavior rather than by overt aggression. In particular, display behavior is important for indicating dominance rank, in communicating aggressive intent, and as a prelude to status struggles. The functional relationships between status, displays and other socio-sexual behavior patterns are presented.     

Lateral bias in infant chimpanzees (Pan troglodytes)

This study documents the presence, strength, and direction of lateralization in chimpanzees (Pan troglodytes) over the first 3 months of life. Nursery-reared chimpanzees (7 males and 5 females) were repeatedly assessed on a behavioral scale. Lateral bias was measured for 4 behaviors: hand-to-mouth, hand-to-hand, defensive grasp, and first step. Hand-to-mouth was significantly lateralized for the sample. Eight of the 10 chimpanzees that showed hand-to-mouth used the right hand. Lateral bias for defensive grasp was positively related to lateral bias both of first step and of hand-to-mouth. Lateral bias in hand-to-mouth was inversely related to lateral bias in hand-to-hand. Strength of lateralization increased as chimpanzees matured. These laterality effects in infant chimpanzees were expressed under conditions of emotional arousal. Moreover, degree of laterality may be a predictor of responsivity to stress.     

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.     

Recognition of Line-Drawing Representations by a Chimpanzee (Pan troglodytes)

The ability of a chimpanzee to recognize individuals portrayed in line drawings was evaluated. A 12-year-old female chimpanzee with extensive prior experience in the use of visual symbols matched the line drawings of chimpanzees, humans, and an orangutan with a specific letter of the alphabet. When a line drawing of a familiar individual was presented on the computer screen, the chimpanzee responded by punching a key with the letter of the alphabet that corresponded to the individual's name. Results indicate that the chimpanzee is able to categorize individuals from novel line-drawing representations.     

Numerical judgments by chimpanzees (Pan troglodytes) in a token economy

We presented four chimpanzees with a series of tasks that involved comparing two token sets or comparing a token set to a quantity of food. Selected tokens could be exchanged for food items on a one-to-one basis. Chimpanzees successfully selected the larger numerical set for comparisons of 1 to 5 items when both sets were visible and when sets were presented through one-by-one addition of tokens into two opaque containers. Two of four chimpanzees used the number of tokens and food items to guide responding in all conditions, rather than relying on token color, size, total amount, or duration of set presentation. These results demonstrate that judgments of simultaneous and sequential sets of stimuli are made by some chimpanzees on the basis of the numerousness of sets rather than other non-numerical dimensions. The tokens were treated as equivalent to food items on the basis of their numerousness, and the chimpanzees maximized reward by choosing the larger number of items in all situations.