The repertoire and intentionality of gestural communication in wild chimpanzees

A growing body of evidence suggests that human language may have emerged primarily in the gestural rather than vocal domain, and that studying gestural communication in great apes is crucial to understanding language evolution. Although manual and bodily gestures are considered distinct at a neural level, there has been very limited consideration of potential differences at a behavioural level. In this study, we conducted naturalistic observations of adult wild East African chimpanzees (Pan troglodytes schweinfurthii) in order to establish a repertoire of gestures, and examine intentionality of gesture production, use and comprehension, comparing across manual and bodily gestures. At the population level, 120 distinct gesture types were identified, consisting of 65 manual gestures and 55 bodily gestures. Both bodily and manual gestures were used intentionally and effectively to attain specific goals, by signallers who were sensitive to recipient attention. However, manual gestures differed from bodily gestures in terms of communicative persistence, indicating a qualitatively different form of behavioural flexibility in achieving goals. Both repertoire size and frequency of manual gesturing were more affiliative than bodily gestures, while bodily gestures were more antagonistic. These results indicate that manual gestures may have played a significant role in the emergence of increased flexibility in great ape communication and social bonding.     

How cross-fostered chimpanzees (Pan troglodytes) initiate and maintain conversations

This study systematically sampled typical attention-getting sounds and sign language conversations between each of 4 originally cross-fostered chimpanzees (Pan troglodytes), still living freely, but now in a laboratory setting, and a familiar human interlocutor. Videotape records showed that when they encountered a human interlocutor sitting alone at his desk with his back turned to them, the cross-fosterlings either left the scene or made attention-getting sounds. The only signs they made to the interlocutor's back were noisy signs. When the human turned and faced them, the chimpanzees promptly signed to him (98% of the time) and rarely made any sounds during the ensuing signed conversations. Under systematic experimental conditions, the signed responses of the chimpanzees were appropriate to the conversational styles of the human interlocutor, confirming daily field observations.     

Interactive use of sign language by cross-fostered chimpanzees (Pan troglodytes)

Cross-fostered as infants in Reno, Nevada, chimpanzees (Pan troglodytes) Washoe, Moja, Tatu, and Dar freely converse in signs of American Sign Language with each other as well as with humans in Ellensburg, Washington. In this experiment, a human interlocutor waited for a chimpanzee to initiate conversations with her and then responded with 1 of 4 types of probes: general requests for more information, on-topic questions, off-topic questions, or negative statements. The responses of the chimpanzees to the probes depended on the type of probe and the particular signs in the probes. They reiterated, adjusted, and shifted the signs in their utterances in conversationally appropriate rejoinders. Their reactions to and interactions with a conversational partner resembled patterns of conversation found in similar studies of human children.     

Different social motives in the gestural communication of chimpanzees and human children

Both chimpanzees and human infants use the pointing gesture with human adults, but it is not clear if they are doing so for the same social motives. In two studies, we presented chimpanzees and human 25-month-olds with the opportunity to point for a hidden tool (in the presence of a non-functional distractor). In one condition it was clear that the tool would be used to retrieve a reward for the pointing subject (so the pointing was selfish or 'for-me'), whereas in the other condition it was clear that the tool would be used to retrieve the reward for the experimenter (so the pointing was helpful or 'for-you'). The chimpanzees pointed reliably only when they themselves benefited, whereas the human children pointed reliably no matter who benefited. These results are interpreted as evidence for the especially cooperative nature of human communication.     

Communicative intentions in wild chimpanzees: persistence and elaboration in gestural signalling

We examine evidence for communicative intent during conspecific interactions in wild chimpanzees (Budongo Forest, Uganda), focusing on persistence in gestural communication. Previous research indicates that great apes have large gestural repertoires and produce gestural communication in a flexible and intentional manner, including the production of gesture sequences. Although there is a lack of consensus on the form and function of sequences, there is some evidence that sequences are produced when signallers fail to receive any response from a recipient. Here, we provide first systematic evidence for communicative persistence in wild chimpanzees. Rather than examining only the presence or absence of a response, we used the most commonly observed response to assign meanings to gestures and examined sequence production in relation to response congruency. Chimpanzees ceased communication if successful, but persevered when unsuccessful. Chimpanzees repeated gestures when a response partially matched their goal but substituted the original gesture when a response was incongruent. Persistence was also mediated by recipient intent to respond, with more sequences produced within competitive than affiliative contexts. Gestures within sequences were homogenous in semantic meaning and signallers continued until the response matched the assigned meaning of the initial gesture. Gestural sequence production was not primarily affective; gesture intensity (in terms of modality) did not increase within sequences. Chimpanzee gestural sequences emerged to achieve specific outcomes; given variability in recipient behaviour following initial gestures, signallers were flexible in their persistence towards these goals.     

Vocal learning of a communicative signal in captive chimpanzees, Pan troglodytes

We hypothesized that chimpanzees could learn to produce attention-getting (AG) sounds via positive reinforcement. We conducted a vocal assessment in 76 captive chimpanzees for their use of AG sounds to acquire the attention of an otherwise inattentive human. Fourteen individuals that did not produce AG sounds during the vocal assessment were evaluated for their ability to acquire the use of an AG sound through operant conditioning and to employ these sounds in an attention-getting context. Nine of the 14 chimpanzees were successfully shaped using positive reinforcement to produce an AG sound. In a post-training vocal assessment, eight of the nine individuals that were successfully trained to produce AG sounds generalized the use of these newly acquired signals to communicatively relevant situations. Chimpanzees possess the ability to acquire the use of a communicative signal via operant conditioning and can generalize the use of this newly acquired signal to appropriate communicative contexts.     

Differential use of vocal and gestural communication by chimpanzees (Pan troglodytes) in response to the attentional status of a human (Homo sapiens)

This study examined the communicative behavior of 49 captive chimpanzees (Pan troglodytes), particularly their use of vocalizations, manual gestures, and other auditory- or tactile-based behaviors as a means of gaining an inattentive audience's attention. A human (Homo sapiens) experimenter held a banana while oriented either toward or away from the chimpanzee. The chimpanzees' behavior was recorded for 60 s. Chimpanzees emitted vocalizations faster and were more likely to produce vocalizations as their 1st communicative behavior when a human was oriented away from them. Chimpanzees used manual gestures more frequently and faster when the human was oriented toward them. These results replicate the findings of earlier studies on chimpanzee gestural communication and provide new information about the intentional and functional use of their vocalizations.     

A prelinguistic gestural universal of human communication

Several cognitive accounts of human communication argue for a language-independent, prelinguistic basis of human communication and language. The current study provides evidence for the universality of a prelinguistic gestural basis for human communication. We used a standardized, semi-natural elicitation procedure in seven very different cultures around the world to test for the existence of preverbal pointing in infants and their caregivers. Results were that by 10-14 months of age, infants and their caregivers pointed in all cultures in the same basic situation with similar frequencies and the same proto-typical morphology of the extended index finger. Infants' pointing was best predicted by age and caregiver pointing, but not by cultural group. Further analyses revealed a strong relation between the temporal unfolding of caregivers' and infants' pointing events, uncovering a structure of early prelinguistic gestural conversation. Findings support the existence of a gestural, language-independent universal of human communication that forms a culturally shared, prelinguistic basis for diversified linguistic communication.     

The gestural repertoire of the wild chimpanzee

Great ape gestural communication is known to be intentional, elaborate and flexible; yet there is controversy over the best interpretation of the system and how gestures are acquired, perhaps because most studies have been made in restricted, captive settings. Here, we report the first systematic analysis of gesture in a population of wild chimpanzees. Over 266 days of observation, we recorded 4,397 cases of intentional gesture use in the Sonso community, Budongo, Uganda. We describe 66 distinct gesture types: this estimate appears close to asymptote, and the Sonso repertoire includes most gestures described informally at other sites. Differences in repertoire were noted between individuals and age classes, but in both cases, the measured repertoire size was predicted by the time subjects were observed gesturing. No idiosyncratic usages were found, i.e. no gesture type was used only by one individual. No support was found for the idea that gestures are acquired by ‘ontogenetic ritualization’ from originally effective actions; moreover, in detailed analyses of two gestures, action elements composing the gestures did not closely match those of the presumed original actions. Rather, chimpanzee gestures are species-typical; indeed, many are ‘family-typical’, because gesture types recorded in gorillas, orangutans and chimpanzee overlap extensively, with 24 gestures recorded in all three genera. Nevertheless, chimpanzee gestures are used flexibly across a range of contexts and show clear adjustment to audience (e.g. silent gestures for attentive targets, contact gestures for inattentive ones). Such highly intentional use of a species-typical repertoire raises intriguing questions for the evolution of advanced communication.     

Urinary cortisol responses to unusual events in captive chimpanzees (Pan troglodytes)

This study investigated the urinary cortisol stress response to one known stressor (anaesthesia) and three unusual events hypothesized to result in increases in cortisol (confinement to one half of an enclosure for several days due to a hurricane, an enrichment exercise, and a change in group composition) in young chimpanzees (Pan troglodytes). Although a cortisol stress response to a variety of laboratory experiences has been documented in captive animals, it is unclear whether other types of atypical events are stressful, including those that are not necessarily negative. Cortisol was measured in 519 urine samples collected from 20 awake, unrestrained chimpanzees; individuals were compared against their own baseline values. A significant increase in urinary cortisol concentration was found as a result of the stress of anaesthesia, but no significant change in urinary cortisol resulted from the three other potential stressors. A lack of a urinary cortisol response to these events may indicate that the events were not actually stressful for the chimpanzees, but may have resulted from the limited temporal resolution of measuring cortisol excretion as an indicator of integrated secretion, or from changes in rates of agonistic behaviors.     

Evocation of functional and volumetric gestural knowledge by objects and words

We distinguish between grasping gestures associated with using an object for its intended purpose (functional) and those used to pick up an object (volumetric) and we develop a novel experimental framework to show that both kinds of knowledge are automatically evoked by objects and by words denoting those objects. Cued gestures were carried out in the context of depicted objects or visual words. On incongruent trials, the cued gesture was not compatible with gestures typically associated with the contextual item. On congruent trials, the gesture was compatible with the item's functional or volumetric gesture. For both gesture types, response latency was longer for incongruent trials indicating that objects and words elicited both functional and volumetric manipulation knowledge. Additional evidence, however, clearly supports a distinction between these two kinds of gestural knowledge. Under certain task conditions, functional gestures can be evoked without the associated activation of volumetric gestures. We discuss the implication of these results for theories of action evoked by objects and words, and for interpretation of functional imaging results.     

Language-based access to gestural components of conceptual knowledge

We report two experiments in which production of articulated hand gestures was used to reveal the nature of gestural knowledge evoked by sentences referring to manipulable objects. Two gesture types were examined: functional gestures (executed when using an object for its intended purpose) and volumetric gestures (used when picking up an object simply to move it). Participants read aloud a sentence that referred to an object but did not mention any form of manual interaction (e.g., Jane forgot the calculator) and were cued after a delay of 300 or 750 ms to produce the functional or volumetric gesture associated with the object, or a gesture that was unrelated to the object. At both cue delays, functional gestures were primed relative to unrelated gestures, but no significant priming was found for volumetric gestures. Our findings elucidate the types of motor representations that are directly linked to the meaning of words referring to manipulable objects in sentences.     

Gestural impairment and gestural ability in aphasia: A review

Studies of gestural ability in aphasic subjects have found impairment of gestural expression and comprehension, with a close relationship between severity of aphasia and degree of gestural impairment. A few investigators have correlated gestural ability with specific language functions or subcategories of aphasia. Reading appears to correlate better with gestural recognition than does auditory comprehension. Despite the general findings of gestural impairment in aphasia, successes have been reported with gestural training including artificial language techniques, pantomime, Amerind, and American Sign Language. Future studies of gestural therapy in aphasia should examine specific language deficits in order to identify those aphasic individuals who can benefit from gestural training.     

The gestural repertoire and its evolution over the second year

The entire repertoire of communicative gestures was documented in a longitudinal, observational study of 10 infants, whose combined ages covered the range from 9 to 22 months. Early in the second year, giving as a request to do something with the object increased, while emotive gestures decreased. Later in the second year, pointing gestures increased, while protest gestures tended to decrease. Combining gestures with vocalization tended to increase only for protest gestures later in the second year. Eye contact showed a small but continuous increase in coordination with gestures over the second year, particularly with comment, request, and emotive gestures. These findings indicate an increasing use of the parent as an agent and of the informative function in non-verbal communication during this period of transition to verbal communication.     

Triggering social interactions: chimpanzees respond to imitation by a humanoid robot and request responses from it

Even the most rudimentary social cues may evoke affiliative responses in humans and promote social communication and cohesion. The present work tested whether such cues of an agent may also promote communicative interactions in a nonhuman primate species, by examining interaction-promoting behaviours in chimpanzees. Here, chimpanzees were tested during interactions with an interactive humanoid robot, which showed simple bodily movements and sent out calls. The results revealed that chimpanzees exhibited two types of interaction-promoting behaviours during relaxed or playful contexts. First, the chimpanzees showed prolonged active interest when they were imitated by the robot. Second, the subjects requested ‘social’ responses from the robot, i.e. by showing play invitations and offering toys or other objects. This study thus provides evidence that even rudimentary cues of a robotic agent may promote social interactions in chimpanzees, like in humans. Such simple and frequent social interactions most likely provided a foundation for sophisticated forms of affiliative communication to emerge.