Spontaneous logicomathematical constructions by chimpanzees (Pan troglodytes, P. paniscus)

Two experiments investigated the spontaneous construction of precursory logicomathematical operations by human-enculturated and language-reared chimpanzees (Pan troglodytes, Pan paniscus) when they were interacting freely with objects. In experiment 1, three chimpanzees ranging in age from 6 to 18 years were presented with sets of six objects. Chimpanzees constructed equivalence, order and reversibility relations within single sets of objects as well as between two or three contemporaneous sets of objects. The chimpanzees’ logicomathematical operations were more advanced, including infrequent and minimal operations on three sets, than those of some previously investigated younger nonenculturated common chimpanzees. In experiment 2, six chimpanzees ranging in age from 6 to 21 years were presented with sets of 12 objects. Chimpanzees constructed more advanced operations on single sets, but not on contemporaneous sets. The results suggest partial convergence and partial divergence between development of logicomathematical cognition in chimpanzees and humans. Received: 20 January 1999 / Accepted after revision: 9 August 1999     

Spatial construction skills of chimpanzees (Pan troglodytes) and young human children (Homo sapiens sapiens)

Spatial construction tasks are basic tests of visual‐spatial processing. Two studies have assessed spatial construction skills in chimpanzees (Pan troglodytes) and young children (Homo sapiens sapiens) with a block modelling task. Study 1a subjects were three young chimpanzees and five adult chimpanzees. Study 1b subjects were 30 human children belonging to five age groups (24, 30, 36, 42, 48 months). Subjects were given three model constructions to reproduce: Line, Cross‐Stack and Arch, which differed in type and number of spatial relations and dimensions, but required comparable configurational understanding. Subjects’ constructions were rated for accuracy. Our results show that: (1) chimpanzees are relatively advanced in constructing in the vertical dimension; (2) Among chimpanzees only adults make accurate copies of constructions; (3) Chimpanzees do not develop in the direction of constructing in two dimensions as human children do starting from age 30 months. The pattern of development of construction skills in chimpanzees partially diverges from that of human children and indicates that spatial analysis and spatial representation are partially different in the two species.     

Spontaneous spatial constructions by chimpanzees (Pan troglodytes,Pan paniscus)

We investigated chimpanzees’ spontaneous spatial constructions with objects. Two were common chimpanzees ages 6 and 18 years and one was an 11‐year‐old bonobo raised in a very enriched human environment from very early in life. Chimpanzees’ products and procedures were quite advanced when constructing functional spatial relations by placing objects in or on each other, but not when constructing nonfunctional spatial relations by placing objects next to each other. This difference was found in all their constructions, including their spatial correspondences and spatial symmetries. Compared to those of younger nonencultured chimpanzees all their spatial constructions were more advanced. Compared to human infants, their functional spatial constructions were also more advanced but their nonfunctional spatial constructions were less advanced.     

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     

Tool use by chimpanzees (Pan troglodytes) of the Arnhem Zoo community

Abstract: Chimpanzees have a large repertoire of tool-use behaviors. This study reports on the variety and the extent of tool use exhibited by the chimpanzees of the Arnhem Zoo community in The Netherlands, living in an enriched captive setting since 1971. The use of tools by 29 chimpanzees aged from 0 to 37 years was observed. We identified 13 types of tool use comparable to those found in the wild. Some of these types of tool use seem to be specific to this community, and can be explained by the ecological characteristics of this captive setting. Chimpanzees started to use tools from the age of 2 years. Young chimpanzees, from 5 to 9 years old, showed a greater repertoire of tool use than infants and adults. All types of tool use in the community have appeared by the age of 10, the age of puberty for chimpanzees. Multivariate analysis was applied for the 29 individuals by 13 types of tool use in a one-zero matrix. The results show two major categories of tool use, one in a practical or substantial context and the other in a nonpractical or play context. The subjects clustered into groups reflecting developmental stages, although there are great individual differences. In conclusion, this captive community provides a unique opportunity to clarify the details of tool use by chimpanzees.     

Cues that chimpanzees do and do not use to find hidden objects

Chimpanzees follow conspecific and human gaze direction reliably in some situations, but very few chimpanzees reliably use gaze direction or other communicative signals to locate hidden food in the object-choice task. Three studies aimed at exploring factors that affect chimpanzee performance in this task are reported. In the first study, vocalizations and other noises facilitated the performance of some chimpanzees (only a minority). In the second study, various behavioral cues were given in which a human experimenter either touched, approached, or actually lifted and looked under the container where the food was hidden. Each of these cues led to enhanced performance for only a very few individuals. In the third study – a replication with some methodological improvements of a previous experiment – chimpanzees were confronted with two experimenters giving conflicting cues about the location of the hidden food, with one of them (the knower) having witnessed the hiding process and the other (the guesser) not. In the crucial test in which a third experimenter did the hiding, no chimpanzee found the food at above chance levels. Overall, in all three studies, by far the best performers were two individuals who had been raised in infancy by humans. It thus seems that while chimpanzees are very good at “behavior reading” of various sorts, including gaze following, they do not understand the communicative intentions (informative intentions) behind the looking and gesturing of others – with the possible exception of enculturated chimpanzees, who still do not understand the differential significance of looking and gesturing done by people who have different knowledge about states of affairs in the world. Received: 8 November 1999 / Accepted after revision: 24 January 2000     

Cultural innovation and transmission of tool use in wild chimpanzees: evidence from field experiments

Chimpanzees (Pan troglodytes) are the most proficient and versatile users of tools in the wild. How such skills become integrated into the behavioural repertoire of wild chimpanzee communities is investigated here by drawing together evidence from three complementary approaches in a group of oil-palm nut- (Elaeis guineensis) cracking chimpanzees at Bossou, Guinea. First, extensive surveys of communities adjacent to Bossou have shown that population-specific details of tool use, such as the selection of species of nuts as targets for cracking, cannot be explained purely on the basis of ecological differences. Second, a 16-year longitudinal record tracing the development of nut-cracking in individual chimpanzees has highlighted the importance of a critical period for learning (3–5 years of age), while the similar learning contexts experienced by siblings have been found to result in near-perfect (13 out of 14 dyads) inter-sibling correspondence in laterality. Third, novel data from field experiments involving the introduction of unfamiliar species of nuts to the Bossou group illuminates key aspects of both cultural innovation and transmission. We show that responses of individuals toward the novel items differ markedly with age, with juveniles being the most likely to explore. Furthermore, subjects are highly specific in their selection of conspecifics as models for observation, attending to the nut-cracking activities of individuals in the same age group or older, but not younger than themselves. Together with the phenomenon of inter-community migration, these results demonstrate a mechanism for the emergence of culture in wild chimpanzees.     

Spontaneous classification in action by a human-enculturated and language-reared bonobo (Pan paniscus) and common chimpanzees (Pan troglodytes).

The authors investigated the formal features of spontaneous manipulations used by 1 bonobo (Pan paniscus) and 2 common chimpanzees (Pan troglodytes) to classify objects in action. Chimpanzees' manipulations were evenly split between serial, one-at-a-time acts on 1 object and parallel, two-at-a-time acts on 2 or 3 objects. Chimpanzees systematically combined their manipulations into routines to generate class-consistent categories of objects. Their routines featured much reproduction of the same manipulations, planful acts that anticipated follow-up manipulations, and manipulations that were reciprocal to each other to accomplish an end. In all these respects, chimpanzees' manipulations were similar to those of 2-year-old human infants. In others they differed. Chimpanzees' routines were mainly based on a linear integration of manipulations. Classifying in action was only infrequently produced by hierarchically integrated routines.     

The distribution and development of handedness for manual gestures in captive chimpanzees (Pan troglodytes)

This article describes the distribution and development of handedness for manual gestures in captive chimpanzees. Data on handedness for unimanual gestures were collected in a sample of 227 captive chimpanzees. Handedness for these gestures was compared with handedness for three other measures of hand use: tool use, reaching, and coordinated bimanual actions. Chimpanzees were significantly more right-handed for gestures than for all other measures of hand use. Hand use for simple reaching at 3 to 4 years of age predicted hand use for gestures 10 years later. Use of the right hand for gestures was significantly higher when gestures were accompanied by a vocalization than when they were not. The collective results suggest that left-hemisphere specialization for language may have evolved initially from asymmetries in manual gestures in the common ancestor of chimpanzees and humans, rather than from hand use associated with other, non-communicative motor actions, including tool use and coordinated bimanual actions, as has been previously suggested in the literature.     

Chimpanzees (Pan troglodytes) accurately compare poured liquid quantities

Although many studies have shown that nonhuman animals can choose the larger of two discrete quantities of items, less emphasis has been given to discrimination of continuous quantity. These studies are necessary to discern the similarities and differences in discrimination performance as a function of the type of quantities that are compared. Chimpanzees made judgments between continuous quantities (liquids) in a series of three experiments. In the first experiment, chimpanzees first chose between two clear containers holding differing amounts of juice. Next, they watched as two liquid quantities were dispensed from opaque syringes held above opaque containers. In the second experiment, one liquid amount was presented by pouring it into an opaque container from an opaque syringe, whereas the other quantity was visible the entire time in a clear container. In the third experiment, the heights at which the opaque syringes were held above opaque containers differed for each set, so that sometimes sets with smaller amounts of juice were dropped from a greater height, providing a possible visual illusion as to the total amount. Chimpanzees succeeded in all tasks and showed many similarities in their continuous quantity estimation to how they performed previously in similar tasks with discrete quantities (for example, performance was constrained by the ratio between sets). Chimpanzees could compare visible sets to nonvisible sets, and they were not distracted by perceptual illusions created through various presentation styles that were not relevant to the actual amount of juice dispensed. This performance demonstrated a similarity in the quantitative discrimination skills of chimpanzees for continuous quantities that matches that previously shown for discrete quantities.     

Development of self-recognition in chimpanzees (Pan troglodytes).

Chimpanzees (Pan troglodytes) demonstrate the ability to recognize themselves in mirrors, yet investigations of the development of self-recognition in chimpanzees are sparse. Twelve young chimpanzees, grouped by age, were given mirror exposure and tested for self-recognition and contingent movement. All 6 juveniles, 4 and 5 years old, exhibited mirror-guided, mark-directed behavior and clear evidence of self-recognition. In contrast, among the infants, only the oldest group of 2 1/2-year-olds exhibited clear evidence of self-recognition. All chimpanzees exhibited both self-directed behaviors and contingent movements. These results suggest that self-recognition occurs at a slightly older age in chimpanzees than in human infants. In humans, self-recognition is linked with other cognitive abilities. The results conform to the general pattern that great apes exhibit many cognitive skills comparable to those of 2-year-old humans.     

Erratum to: Chimpanzees (Pan troglodytes) accurately compare poured liquid quantities

Although many studies have shown that nonhuman animals can choose the larger of two discrete quantities of items, less emphasis has been given to discrimination of continuous quantity. These studies are necessary to discern the similarities and differences in discrimination performance as a function of the type of quantities that are compared. Chimpanzees made judgments between continuous quantities (liquids) in a series of three experiments. In the first experiment, chimpanzees first chose between two clear containers holding differing amounts of juice. Next, they watched as two liquid quantities were dispensed from opaque syringes held above opaque containers. In the second experiment, one liquid amount was presented by pouring it into an opaque container from an opaque syringe, whereas the other quantity was visible the entire time in a clear container. In the third experiment, the heights at which the opaque syringes were held above opaque containers differed for each set, so that sometimes sets with smaller amounts of juice were dropped from a greater height, providing a possible visual illusion as to the total amount. Chimpanzees succeeded in all tasks and showed many similarities in their continuous quantity estimation to how they performed previously in similar tasks with discrete quantities (for example, performance was constrained by the ratio between sets). Chimpanzees could compare visible sets to nonvisible sets, and they were not distracted by perceptual illusions created through various presentation styles that were not relevant to the actual amount of juice dispensed. This performance demonstrated a similarity in the quantitative discrimination skills of chimpanzees for continuous quantities that matches that previously shown for discrete quantities.     

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.     

Now you see me,now you don't: evidence that chimpanzees understand the role of the eyes in attention

Chimpanzees appear to understand something about the attentional states of others; in the present experiment, we investigated whether they understand that the attentional state of a human is based on eye gaze. In all, 116 adult chimpanzees were offered food by an experimenter who engaged in one of the four experimental manipulations: eyes closed, eyes open, hand over eyes, and hand over mouth. The communicative behavior of the chimpanzees was observed. More visible behaviors were produced when the experimenter's eyes were visible than when the experimenter's eyes were not visible. More vocalizations were produced when the experimenter's eyes were closed than when they were open, but there were no differences in other attention getting behaviors. There was no effect of age or rearing history. The results suggest that chimpanzees use the presence of the eyes as a cue that their visual gestures will be effective.     

Slow development of vocal sequences through ontogeny in wild chimpanzees (Pan troglodytes verus)

The development of the unique, hierarchical, and endless combinatorial capacity in a human language requires neural maturation and learning through childhood. Compared with most non-human primates, where combinatorial capacity seems limited, chimpanzees present a complex vocal system comprising hundreds of vocal sequences. We investigated how such a complex vocal system develops and the processes involved. We recorded 10,929 vocal utterances of 98 wild chimpanzees aged 0–55 years, from Taï National Park, Ivory Coast. We developed customized Generalized non-Linear Models to estimate the ontogenetic trajectory of four structural components of vocal complexity: utterance length, diversity, probability of panting (requiring phonation across inhalation and exhalation), and probability of producing two adjacent panted units. We found chimpanzees need 10 years to reach adult levels of vocal complexity. In three variables, the steepest increase coincided with the age of first non-kin social interactions (2–5 years), and plateaued in sub-adults (8–10 years), as individuals integrate into adult social life. Producing two adjacent panted units may require more neuromuscular coordination of the articulators, as its emergence and steepest increase appear later in development. These results suggest prolonged maturational processes beyond those hitherto thought likely in species that do not learn their vocal repertoire. Our results suggest that multifaceted ontogenetic processes drive increases in vocal structural complexity in chimpanzees, particularly increases in social complexity and neuro-muscular maturation. As humans live in a complex social world, empirical support for the “social complexity hypothesis” may have relevance for theories of language evolution.

Research Highlights

• Chimpanzees need around 10 years to develop the vocal structural complexity present in the adult repertoire, way beyond the age of emergence of every single vocal unit.
• Multifaceted ontogenetic processes may drive increases in vocal structural complexity in chimpanzees, particularly increases in social complexity and neuro-muscular maturation.
• Non-linear increases in vocal complexity coincide with social developmental milestones.
• Vocal sequences requiring rapid articulatory change emerge later than other vocal sequences, suggesting neuro-muscular maturational processes continue through the juvenile years.

     

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.     

"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     

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.     

Long-term retention of the differential values of Arabic numerals by chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

As previously reported (Beran and Rumbaugh, 2001), two chimpanzees used a joystick to collect dots, one-at-a-time, on a computer monitor, and then ended a trial when the number of dots collected was equal to the Arabic numeral presented for the trial. Here, the chimpanzees were presented with the task again after an interval of 6 months and then again after an additional interval of 3.25 years. During each interval, the chimpanzees were not presented with the task, and this allowed an assessment of the extent to which both animals retained the values of each Arabic numeral. Despite lower performance at each retention interval compared to the original study, both chimpanzees performed above chance levels 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. For the 3.25-year retention, errors were more dispersed around each target numeral than in the original study, but the chimpanzees' performances again appeared to be based on a continuous representation of magnitude rather than a discrete representation of number. These data provide an experimental demonstration of long-term retention of the differential values of Arabic numerals by chimpanzees.     

Stream/bounce perception and the effect of depth cues in chimpanzees (Pan troglodytes)

The stream/bounce display represents an ambiguous motion event in which two identical visual objects move toward one another and the objects overlap completely before they pass each another. In our perception, they can be interpreted as either streaming past one another or bouncing off each other. Previous studies have shown that the streaming percept of the display is generic for humans, suggesting the inertial nature of the motion integration process. In this study, chimpanzees took part in behavioral experiments using an object-tracking task to reveal the characteristics of their stream/bounce perception. Chimpanzees did not show a tendency toward a dominant "stream" perception of the stream/bounce stimulus. However, depth cues, such as X-junctions and local motion coherence, did promote the stream percept in chimpanzees. These results suggest both similarities and differences between chimpanzees and humans with respect to motion integration and object individuation processes.