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Ape Gestures and Language Evolution

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Ape Gestures and Language Evolution Ape gestures and language evolution Amy S. Pollick and Frans B. M. de Waal* Living Links, Yerkes National Primate Research Center, Emory University, 954 North Gatewood Road, Atlanta, GA 30329 Contributed by Frans B. M. de Waal, March 20, 2007 (sent for review January 28, 2007) The natural communication of apes may hold clues about language This difference in control was dramatically illustrated by the origins, especially because apes frequently gesture with limbs and failure of efforts to teach chimpanzees vocal modifications to hands, a mode of communication thought to have been the ‘‘speak’’ (21, 22), whereas this species had no trouble learning to starting point of human language evolution. The present study use the gestures of American sign language in a referential aimed to contrast brachiomanual gestures with orofacial move- manner (23). Greater control over natural gestures than facial ments and vocalizations in the natural communication of our expressions is also suggested by observations of (i) deception, in closest primate relatives, bonobos (Pan paniscus) and chimpanzees which apes may use a hand to modify a facial expression (24, 25), (Pan troglodytes). We tested whether gesture is the more flexible and (ii) cultural transmission, with gestures being far more form of communication by measuring the strength of association population-specific in both humans and apes than facial expres- between signals and specific behavioral contexts, comparing sions, which tend to be relatively invariant (8, 26–28). groups of both the same and different ape species. Subjects were All of these observations are in line with the gestural hypoth- two captive bonobo groups, a total of 13 individuals, and two esis of human language origins, which is further supported by captive chimpanzee groups, a total of 34 individuals. The study differential growth of the brain and vocal apparatuses, as seen in distinguished 31 manual gestures and 18 facial/vocal signals. It was paleoarchaeological remains (29), the appearance of gestural found that homologous facial/vocal displays were used very sim- communication in human infants before speech (30), and the ilarly by both ape species, yet the same did not apply to gestures. right-hand (hence left-brain) bias of both ape and human Both within and between species gesture usage varied enor- gestures (31–33). The ape homologue of Broca’s area (i.e., mously. Moreover, bonobos showed greater flexibility in this Brodmann’s area 44) is enlarged in the left hemisphere (34). In monkeys, this area is activated during both the production and regard than chimpanzees and were also the only species in which perception of gestures but not vocalizations (35). It has been multimodal communication (i.e., combinations of gestures and speculated, therefore, that the neural structures underlying facial/vocal signals) added to behavioral impact on the recipient. manual movements in the great apes, perhaps also including tool ͉ ͉ ͉ use (36), are homologous with the lateralized language areas in bonobo chimpanzee communication multimodal the human brain (37, 38). Gesture remains very much alive in human communication. he longstanding debate about the evolution of language Recent research demonstrates the universal importance of ges- Ttypically compares human speech with animal vocalizations ture to human cognitive functioning, such as enhanced infor- (1). The vocal modality offers obvious parallels, yet it has been mation transfer (39), lexical retrieval (40), and even the provi- proposed that our ancestors’ first linguistic utterances were not sion of a supplemental cognitive arena for thought (41). Gesture in the vocal but in the gestural domain (2–5). This proposal is all production in humans is so automatic that it is relatively immune the more intriguing because gestural communication is virtually to audience effects: blind subjects gesture at equal rates as limited to the Hominoidea (i.e., humans and apes). sighted subjects to a known blind audience (42). Gestures are narrowly defined here as communication by Gestures are rarely produced in the absence of other com- means of hands, feet, or limbs (Table 1). One reason to set municative signals, such as facial expressions and vocalizations. gestures apart from other bodily communication is that the two Multimodal communication has been appreciated in humans for are neurologically distinct in both production and perception by several decades (43) and is becoming increasingly important in others (6, 7). Whereas all primates regularly communicate by the study of animal communication (44). Multimodal signaling means of vocalizations, orofacial movements, body postures, and occurs across taxa, from snapping shrimps to spiders and birds, locomotion patterns, free brachiomanual gestures (i.e., manual and in all contexts, although those related to courtship and communication without touching another individual or a sub- mating are best documented (A.S.P., unpublished work). This strate) are typical of humans and apes (8). Gestures were first communication strategy can have a variety of functions, includ- described for chimpanzees (9, 10), followed by other anthropoid ing amplification and modulation of signal meaning. Combined apes: bonobos (11, 12), gorillas (Gorilla gorilla; refs. 13 and 14), with the graded facial/vocal signals typical of the apes (46), and orangutans (Pongo pygmaeus; ref. 15). gestural flexibility has the advantage over the more stereotyped The discontinuity between the Hominoidea and all other signaling by monkeys that it permits greater communicative primates regarding gestural modality suggests a relatively recent complexity. shift toward a more flexible and intentional communicative Despite prior work on ape gesture, no research has investi- gated how these signals combine with other forms of natural strategy in our prehominid ancestors (8). One mark of this shift communication and how variations in usage affect behavior. is contextually defined usage; that is, a single gesture may Here, we set out to test how the natural gestural modality communicate entirely different needs or intentions depending combines with and differs from the facial/vocal modality in the on the social context in which it is used. For example, a chimpanzee stretching out an open hand toward a third party during a fight signals a need for support, whereas the same Author contributions: A.S.P. and F.B.M.d.W. designed research, performed research, ana- gesture toward a possessor of food signals a desire for a share lyzed data, and wrote the paper. (refs. 16 and 17; Fig. 1). This is not to say that other forms of The authors declare no conflict of interest. communication cannot derive extra meaning from the context in *To whom correspondence should be addressed. E-mail: [email protected]. which they occur (18), but gestures seem less closely tied to This article contains supporting information online at www.pnas.org/cgi/content/full/ specific emotions, hence they permit greater cortical control 0702624104/DC1. than other forms of communication (19, 20). © 2007 by The National Academy of Sciences of the USA 8184–8189 ͉ PNAS ͉ May 8, 2007 ͉ vol. 104 ͉ no. 19 www.pnas.org͞cgi͞doi͞10.1073͞pnas.0702624104 Downloaded by guest on September 26, 2021 Table 1. Operational definition of gesture (1) Gesture refers to a nonlocomotory movement with communicative value of the forearm, hand, wrist, or fingers (cf. ref. 14). Leg and foot movements are included, because these movements have been observed to be communicative in bonobos (11). Excluded from this definition are any other movements that are sometimes labeled gestures in the literature (e.g., ref. 53), such as body postures, head movements, and locomotion patterns. (2) The movement must be directed at another individual. Directed means that the performer of the gesture must potentially be in the recipient’s view. This applies only to nontactile gestures; for a tactile gesture the recipient simply needs to be within touching proximity. (3) A tactile action is considered a gesture only if it lasts2sand visibly lacks the mechanical force to bring about the reaction shown by the recipient, and also does not include any attempt to grab or extensively hold a body part of the other. For example, the gesture hard touch applied to the leg neither moves the other’s leg in any observable direction nor does it visibly, forcibly change the other’s physical position (if a positional change does result, it must appear ЉvoluntaryЉ). (4) Play movements are repeated and exaggerated, usually leading to tickling, play hitting, or wrestling. Movements during play are not considered gestures in this study, except for those that initiate play. These gestures must occur at least 0.5 s before any play is observed between two individuals. communication of the two ape species genetically closest to humans: bonobos and chimpanzees. Given the above back- ground, we formulated a gestural flexibility hypothesis according to which the usage of gestures will be more flexible than that of facial/vocal displays, both within and between species. In testing this hypothesis, we will use context as a proxy for usage, meaning, PSYCHOLOGY and function; that is, we predict that gestures will be less context-bound than other types of signals. The Pan line, which includes both chimpanzees and bonobos, split off from the line that produced our species Ϸ6 million years ago (47), whereas the two ape species themselves split apart only Fig. 1. Meaning often needs to be extracted from the specific context in Ϸ2.5 million years ago (48). Chimpanzees and bonobos are, which a gesture is being used. (A) A juvenile chimpanzee tries to reclaim food therefore, genetically equidistant to us. Studying similar types of that a dominant has taken away by combining the reach out up begging gesture with a scream vocalization. (B) An adolescent bonobo male making communicative signals in closely related species allows one to sexual advances to a female adds the arm raise gesture. Photographs by Frans determine homologies, i.e., shared evolutionary ancestry (49).
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