David A. Leavens

Intentional Communication by Chimpanzees: A Cross-Sectional Study of the Use of Referential Gestures

This study describes the use of referential gestures with concomitant gaze orienting behavior to both distal food objects and communicative interactants by 115 chimpanzees, ranging from 3 to 56 years of age. Gaze alternation between a banana and an experimenter was significantly associated with vocal and gestural communication. Pointing was the most common gesture elicited; 47 subjects pointed with the whole hand, whereas 6 subjects pointed with index fingers. Thus, communicative pointing is commonly used by laboratory chimpanzees, without explicit training to point, language training, or home rearing. Juveniles exhibited striking decrements in their propensity to communicate with adult male experimenters compared with older chimpanzees. Significantly fewer mother-reared chimpanzees exhibited gaze alternation compared with nursery-reared chimpanzees.

Understanding the Point of Chimpanzee Pointing: Epigenesis and Ecological Validity

Pointing has long been considered to be a uniquely human, universal, and biologically based gesture. However, pointing emerges spontaneously, without explicit training, in captive chimpanzees. Because pointing is commonplace in captive chimpanzees and virtually absent in wild chimpanzees, and because both captive and wild chimpanzees are sampled from the same gene pool, pointing by captive apes is attributable to environmental influences on communicative development. If pointing by captive chimpanzees is so variably expressed in different rearing environments, this suggests that pointing by humans may also be attributable to situational factors that make pointing effective in certain developmental contexts.

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.

Hand use and gestural communication in chimpanzees (Pan troglodytes).

Hand use in gestural communication was examined in 115 captive chimpanzees (Pan troglodytes). Hand use was measured in subjects while they gestured to food placed out of their reach. The distribution of hand use was examined in relation to sex, age, rearing history, gesture type, and whether the subjects vocalized while gesturing. Overall, significantly more chimpanzees, especially females and adults, gestured with their right than with their left hand. Foods begs were more lateralized to the right hand than pointing, and a greater prevalence of right-hand gesturing was found in subjects who simultaneously vocalized than those who did not. Taken together, these data suggest that referential, intentional communicative behaviors, in the form of gestures, are lateralized to the left hemisphere in chimpanzees.

Multimodal communication by captive chimpanzees (Pan troglodytes)

Many studies have shown that apes and monkeys are adept at cross-modal matching tasks requiring the subject to identify objects in one modality when information regarding those objects has been presented in a different modality. However, much less is known about non-human primates’ production of multimodal signaling in communicative contexts. Here, we present evidence from a study of 110 chimpanzees demonstrating that they select the modality of communication in accordance with variations in the attentional focus of a human interactant, which is consistent with previous research. In each trial, we presented desirable food to one of two chimpanzees, turning mid-way through the trial from facing one chimpanzee to facing the other chimpanzee, and documented their communicative displays, as the experimenter turned towards or away from the subjects. These chimpanzees varied their signals within a context-appropriate modality, displaying a range of different visual signals when a human experimenter was facing them and a range of different auditory or tactile (attention-getting) signals when the human was facing away from them; this finding extends previous research on multimodal signaling in this species. Thus, in the impoverished circumstances characteristic of captivity, complex signaling tactics are nevertheless exhibited by chimpanzees, suggesting continuity in intersubjective psychological processes in humans and apes.

Human handedness: an inherited evolutionary trait.

Our objective was to demonstrate that human population-level, right-handedness, is not species specific, precipitated from language areas in the brain, but rather is context specific and inherited from a behavior common to both humans and great apes. In general, previous methods of assessing human handedness have neglected to consider the context of action, or employ methods suitable for direct comparison across species. We employed a bottom-up, context-sensitive method to quantitatively assess manual actions in right-handed, typically developing children during naturalistic behavior. By classifying the target to which participants directed a manual action, as animate (social partner, self) or inanimate (non-living functional objects), we found that children demonstrated a significant right-hand bias for manual actions directed toward inanimate targets, but not for manual actions directed toward animate targets. This pattern was revealed at both the group and individual levels. We used a focal video sampling, corpus data-mining approach to allow for direct comparisons with captive gorillas (Forrester et al. Animal Cognition 2011;14(6):903-7) and chimpanzees (Forrester et al. Animal Cognition, in press). Comparisons of handedness patters support the view that population-level, human handedness, and its origin in cerebral lateralization is not a new or human-unique characteristic. These data are consistent with the theory that human right-handedness is a trait developed through tool use that was inherited from an ancestor common to both humans and great apes.

Target animacy influences gorilla handedness.

We investigated the unimanual actions of a biological family group of twelve western lowland gorillas (Gorilla gorilla gorilla) using a methodological approach designed to assess behavior within social context from a bottom-up perspective. Measures of both the lateralization of unimanual actions (left, right) and the target of the action (animate, inanimate) were assessed during dual, synchronized video observations of naturalistic behavior. This paper demonstrates a corelationship between handedness and the animate quality of the target object. Analyses demonstrated a significant interaction between lateralized unimanual actions and target animacy and a right-hand bias for actions directed toward inanimate targets. We suggest that lateralized motor preference reflects the different processing capabilities of the left and right hemispheres, as influenced by the emotive (animate) and/or functional (inanimate) characteristics of the target, respectively.

Deictic gesturing in wild chimpanzees (Pan troglodytes)? Some possible cases.

Referential pointing is important in the development of language comprehension in the child and is often considered a uniquely human capacity. Nonhuman great apes do point in captivity, usually for a human audience, but this has been interpreted as an interaction pattern learned from human caretakers, not indicative of natural deictic ability. In contrast, spontaneous pointing for other apes is almost unknown among wild ape populations, supporting doubts as to whether apes naturally have any capacity to point referentially. Here the authors describe and illustrate 4 cases of gestures by juvenile chimpanzees in the Sonso chimpanzee community in Budongo, Uganda, that, at some level, may appear to be deictic and referential. The authors discuss the possible reasons why chimpanzees, if they possess a capacity for referential pointing, do not use it more frequently.

Target animacy influences chimpanzee handedness

We employed a bottom-up, quantitative method to investigate great ape handedness. Our previous investigation of gorillas (Gorilla gorilla gorilla) demonstrated that contextual information influenced an individual’s handedness toward target objects. Specifically, we found a significant right-hand bias for unimanual actions directed toward inanimate target objects but not for actions directed to animate target objects (Forrester et al. in Anim Cogn 14(6):903–907, 2011). Using the identical methodological technique, we investigated the spontaneous hand actions of nine captive chimpanzees (Pan troglodytes) during naturalistic, spontaneous behavior. We assessed both the frequencies and proportions of lateralized hand actions directed toward animate and inanimate targets employing focal follow video sampling. Like the gorillas, the chimpanzees demonstrated a right-handed bias for actions directed toward inanimate targets, but not toward animate targets. This pattern was evident at the group level and for the majority of subjects at the individual level. We postulate that a right-hand bias for only inanimate targets reflects the left hemisphere’s dominant neural processing capabilities for objects that have functional properties (inanimate objects). We further speculate that a population-level right-hand bias is not a human-unique characteristic, but one that was inherited from a common human-ape ancestor.

Emotional Engagements Predict and Enhance Social Cognition in Young Chimpanzees

Social cognition in infancy is evident in coordinated triadic engagements, that is, infants attending jointly with social partners and objects. Current evolutionary theories of primate social cognition tend to highlight species differences in cognition based on human-unique cooperative motives. We consider a developmental model in which engagement experiences produce differential outcomes. We conducted a 10-year-long study in which two groups of laboratory-raised chimpanzee infants were given quantifiably different engagement experiences. Joint attention, cooperativeness, affect, and different levels of cognition were measured in 5- to 12-month-old chimpanzees, and compared to outcomes derived from a normative human database. We found that joint attention skills significantly improved across development for all infants, but by 12 months, the humans significantly surpassed the chimpanzees. We found that cooperativeness was stable in the humans, but by 12 months, the chimpanzee group given enriched engagement experiences significantly surpassed the humans. Past engagement experiences and concurrent affect were significant unique predictors of both joint attention and cooperativeness in 5- to 12-month-old chimpanzees. When engagement experiences and concurrent affect were statistically controlled, joint attention and cooperation were not associated. We explain differential social cognition outcomes in terms of the significant influences of previous engagement experiences and affect, in addition to cognition. Our study highlights developmental processes that underpin the emergence of social cognition in support of evolutionary continuity.