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The origins of non-human primates’ manual gesturesAuthor(s): Katja Liebal and Josep CallSource: Philosophical Transactions: Biological Sciences , 12 January 2012, Vol. 367, No.1585, From action to language: comparative perspectives on primate tool use, gestureand the evolution of human language (12 January 2012), pp. 118-128Published by: Royal SocietyStable URL: https://www.jstor.org/stable/41433520REFERENCESLinked references are available on JSTOR for this article:https://www.jstor.org/stable/41433520?seq=1&cid=pdfreference#references_tab_contentsYou may need to log in to JSTOR to access the linked references.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a widerange of content in a trusted digital archive. We use information technology and tools to increase productivity andfacilitate new forms of scholarship. For more information about JSTOR, please contact [email protected] use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available athttps://about.jstor.org/termsRoyal Society is collaborating with JSTOR to digitize, preserve and extend access toPhilosophical Transactions: Biological SciencesThis content downloaded from150.135.174.98 on Thu, 03 Dec 2020 03:06:04 UTCAll use subject to httpPHILOSOPHICALTRANSACTIONStoerWK PhiL Trans’ R’ Soc’ B (2012) 361 3 118-128SOCIETY doi:10.1098/rstb.20 11.0044ReviewThe origins of non-human primates’manual gesturesKatja Liebal1’2’3’* and Josep Call31 Department of Education and Psychology, Cluster Languages of Emotion, Freie Universität Berlin,Habelschwerdter Allee 45, 14195 Berlin, Germany2 Department of Psychology, University of Portsmouth, King Henry 1st Street, Portsmouth POI 2DY, UK3 Department of Developmental and Comparative Psychology, Max Planck Institute for EvolutionaryAnthropology, Deutscher Platz 6, 04103 Leipzig, GermanyThe increasing body of research into human and non-human primates’ gestural communicationreflects the interest in a comparative approach to human communication, particularly possible scen-arios of language evolution. One of the central challenges of this field of research is to identifyappropriate criteria to differentiate a gesture from other non-communicative actions. After an intro-duction to the criteria currently used to define non-human primates’ gestures and an overview ofongoing research, we discuss different pathways of how manual actions are transformed intomanual gestures in both phylogeny and ontogeny. Currently, the relationship between actionsand gestures is not only investigated on a behavioural, but also on a neural level. Here, we focuson recent evidence concerning the differential laterality of manual actions and gestures in apes inthe framework of a functional asymmetry of the brain for both hand use and language.Keywords: gesture; manual; ontogenetic ritualization; phylogenetic ritualization;laterality; great apes
INTRODUCTIONDolphins are one of the most gracile and elegant crea-tures of the sea. However, before dolphins becamewhat they are today, they underwent a remarkabletransformation. The terrestrial ancestor of dolphinswas a hippopotamus-like creature that walked on allfours and lacked the stylized forms, and presumablythe elegant movements, of its marine descendant.Over the last 50 million years, dolphins have beenevolving into what they are today. This remarkabletransformation teaches us an important lesson.Complex structures such as legs and snouts can betransmuted over time into equally complex and func-tionally equivalent structures such as fins andblowholes, respectively.The relation that exists between gesture and actionis in some ways analogous to the relation that existsbetween fins and legs or between noses and blowholes.A central thesis of this contribution is that many of thegestures displayed by apes began their existence asactions devoid of a communicative function, but overtime they became co-opted and transformed into com-municative devices that accomplished similarfunctions [1]. Moreover, just like fins and legs, thischange took place over evolutionary time, but in thecase of gestures, it can also take place during the
Author for correspondence ([email protected]).One contribution of 13 to a Theme Issue ‘From action to language:comparative perspectives on primate tool use, gesture, and theevolution of human language’.lifetime of one individual. In both cases, however,one can find clues that inform us about their origin.Comparing the communicative repertoires of monkeysand apes with those of humans can play a crucial rolein the quest for the roots of human language, and morespecifically in the role that gestures might have playedin the evolution of language.The focus of this paper is twofold. First, we willexplore the question of how actions are transformedinto gestures both from a phylogenetic and an onto-genetic perspective. To this end, we will present thelatest advances in ape gestural communication, includ-ing some of the controversies in the field. We will beginby defining gestures, briefly presenting some of thefeatures of the apes’ gestural repertoires and discussingthree ways in which individuals can acquire gestures.Second, having established the connection betweenactions and gestures, we will turn our attention tothe role that gestures may have played in languageevolution. First, we will note the increasing interestin gestural communication of our closest living pri-mate relatives in the framework of the proposed closelink between action and language in humans. Thenwe will link recent data on ape laterality in gesturaluse with language hemispheric specialization.
GESTURE ORIGINS (OUT OF ACTIONS)(a) Defining a gestureHuman gestures are usually very broadly referred to asthe ‘manner of carrying the body’ and ‘movements ofthe body or limbs as an expression of feeling’ ([2],118 This journal is © 201 1 The Royal SocietyThis content downloaded from150.135.174.98 on Thu, 03 Dec 2020 03:06:04 UTCAll use subject to httpReview. Gesture origins К. Liebal & J. Call 119p. 476). According to Kendon [3], a gesture is a formof non-verbal communication in which visible bodilyactions communicate particular messages, either inplace of speech or together and in parallel withspoken words. Before children start to speak, theyuse a variety of gestures to communicate with theircarers, such as showing objects and pointing toobjects, events or persons in their environment [4-7] . Even when they acquire their first words, gesturesare not simply replaced but are incorporated in theirverbal communication [8,9].In adults, a substantial body of research addresses thekinds of manual gestures produced by humans whilespeaking [3,10 – 12] . If spoken language is not possible,then manual gestures can convey very specific andcomplex information, even replacing spoken language,thus becoming a form of a highly conventionalizedsign system [13,14]. Thus, in humans, gestures canvary in their degree of conventionalization and thereforethe degree to which they are linked to or even replacespoken language ([15], pp. 37-40). Therefore, researchinto human gestures is a highly diverse field, since itcovers very different kinds of gestural communication,such as speech-accompanying gestures, gestures of pre-linguistic children or even gestures co-occurring withsign language. The question arising here is whethernon-human primate species, which are clearly lackingspoken language but with bodies and particularly fore-limbs sharing many characteristics with human beings,use gestures to communicate with conspecifics. Totackle this question and to enable any comparison withhumans at all, we need to focus on human gestures notused in combination with language (either spoken orsigned) and thus on the gestural communication of pre-linguistic children. By adopting the corresponding cri-teria, a gesture is defined as a behaviour that unlike anaction is motorically ineffective. It requires the active par-ticipation of a partner to fulfil its purpose, it is producedin the presence of an audience and is tailored to the atten-tional state of the audience. Furthermore, it involves gazealternation or visual checking between social partnersand distant objects or events, is characterized by the sen-der’s waiting for the recipient’s response and displayspersistence and elaboration of communicative behaviourwhen communicative attempts fail [16-19].As our previous introduction to the term gesturepointed out, gestures are not restricted to the use ofhands, but often include movements of limbs andalso head and body movements, as well as postures.Some scholars even include facial expressions as ges-tures [20,21]. However, here we only focus onmanual gestures in non-human primates, that is, ges-tures produced with the whole arm or hands. Wealso mainly discuss studies of gestural communicationin great apes; this is not to neglect gibbons and mon-keys, but so far there is still little evidence of handuse for the purpose of communication in non-greatape species ([22], but see [23-25]).One of the biggest challenges in gestural researchlies in determining when an instrumental action hascrossed the threshold and becomes a gesture. Somegestures are easy to distinguish from instrumentalactions, but there are others that are much more diffi-cult to differentiate. For instance, we would include asPhil. Trans. R. Soc. В (2012)gestures the subtle touches and presses that dancersuse to inform their partner about their impendingactions or to direct them in a certain way. In contrast,we would not consider as gestures holding an infantwhen she is beginning to walk because here the mainfunction would be to help the infant to maintain herequilibrium.The problem of deciding between gestures andactions is further compounded when multiple speciesare considered. Although the potential for confusingactions and gestures represents a potential analyticweakness, it can become a strength since it tells ussomething about the origin of gestures. In particular,it suggests that at least some gestures may havebegun their existence as actions before they weretransformed into a communicative function.From a more practical point of view, one approachthat we find useful in distinguishing actions from ges-tures is to consider how many of the criteria outlinedabove are met. Thus, faced with a potential candidateas a gesture, we must ask whether (i) it is motoricallyineffective, (ii) there is response waiting, (iii) gazealternation, and (iv) persistence. The more criteriaare met, the more sure we can be that a given behaviourqualifies as a gesture. One cannot be 100 per centsure but at least this method can help in reducing ouruncertainty.In the next two sections, we present a brief overviewof the gestural communication of the great apes (see[18] for a more detailed treatment) and then discusstheir potential origins.(b) Gestures of non-human primatesUnlike research into human gestures mostly restrictingthe focus on the visual channel [3], researchers inves-tigating primate gestures also consider tactile gesturessuch as push or throw objects and gestures with an audi-tory component such as hand clap and chest beat.Including gestures that transfer information via non-visual channels captures the richness and subtletiesof non-vocal communication. However, it also raisespotential problems when it comes to distinguishinggestures from instrumental actions. For instance, agesture called reach that consists of extending an armin the direction of a conspecific is easier to identifyas gesture than a gesture called touch-side that consistsof touching an individual on her side to make hermove. The reason for this is simple. The lack of phys-ical contact between the two interacting individualsautomatically makes reach motorically ineffective, oneof the first criteria to identify a gesture as such. Afterall, it is conceivable that the touch-side gesture involvedenough force to make the individual move, thusmaking this action motorically effective and automati-cally disqualifying it as a gesture. In sum, researchersinvestigating non-human primates have faced a trade-off between capturing the richness and subtleties ofnon-vocal communication in primates at the expenseof making the distinction between gestures and instru-mental actions less clear-cut than in human research.In a recent summary of a systematic comparison ofthe four great apes, siamangs and Barbary macaques,Call & Tomasello [18] concluded that those speciesThis content downloaded from150.135.174.98 on Thu, 03 Dec 2020 03:06:04 UTCAll use subject to http120 К. Liebal & J. Call Review . Gesture originsdiffered in repertoire size, composition and function oftheir gestures. They reported between 20 and 35different gesture types depending on the species,which meet the above-mentioned criteria of beingmotorically ineffective and are accompanied byresponse waiting and/or gaze alternation, as well aspersistence in case the recipient did not react. Out ofthose reported gestural repertoires, at least 50 per centof each species’ repertoire consisted of manualgestures, with the highest proportion found in gorillas(73%). (It is important to note that those numbersrefer to the total repertoires found across differentgroups of one species, not average proportions.)For example, tactile gestures, which included somekind of physical contact with another individual (e.g.touch , pull or slap) , were used by all great apes, siamangsand Barbary macaques [18]. Auditory gestures oftenincluded the individual’s own body used to producethat noise, such as hand clap in chimpanzees [26], andchest beat and body beat in gorillas [27] . Alternatively,noise can be produced by using objects while perform-ing gestures such as ground slap, push objects or throwobjects , which are gestures particularly reported forchimpanzees [26]. On the other hand, examples forsilent gestures not involving physical contact includegestures like extend arm (reach), arm raise and wavearm [18]. As opposed to bonobos, siamangs andBarbary macaques, chimpanzees and orangutans oftenincorporated objects in their gestural displays (15% ofthe gestures). For example, orangutans offer food toother individuals by extending one arm with food intheir hand to another individual [28] and chimpanzeesuse branches, which they shake vigorously to get theattention of another group member [26]. The highervalues for gestures involving objects for chimpanzeesand orangutans are interesting in light of their higherpropensity to use tools in the wild than the otherspecies and may be indicative of a common neuralsubstrate for tool use and gestural communication.So far, we have mostly presented the results of ourown research project on gestural communication ofnon-human primates that started with the work byTomasello et al. [29]. Of course, there are manymore scholars working on the question of which ges-tures non-human primates use, how they acquirethem and what the underlying socio-cognitive skillsare, both in wild and captive settings. The first pio-neering field studies report several gestures as partsof ethograms for orangutans [30], gorillas [31], chim-panzees [32] and bonobos [33], but also for gibbons[34,35] and monkeys [36]. Lately, there is an increasein more systematic, mostly observational studies inves-tigating gesture use within social groups of great apes[37,38] and monkeys [22,24,39-41]. This increasingbody of research reflects the interest in the role ges-tures might have played for the evolution of humanlanguage [42-45], although studies addressing facialexpressions or vocalizations still outnumber studiesconcerning gestures [46] .However, the reported gestural repertoires for thedifferent species vary considerably between studies.For instance, while Pika et al. [27] described 33 ges-tures for gorillas, Genty et al. [37] reported morethan 100 gestures for this species. Furthermore, veryPhil. Trans. R. Soc. В (2012)different names are often used to categorize the samebehaviour, complicating comparisons across studiesand species. These discrepancies may be attributedin part to the sampling effort and the differences inthe detail of the coding schema across studies [47],but it remains a fact that gestures are, first of all, diffi-cult to differentiate from actions, and second, althoughthe majority of gestures are not gradual signals like inthe case of facial expressions, they are difficult to cat-egorize because of the often different criteria usedacross studies to define a gesture. This is closelyrelated to a third reason, namely that gestures areoften defined based on their function or the contextthey are used in (e.g. food offer ), resulting in a confla-tion of form and function rather than referring toform and meaning as separate variables.However, although the paucity of data in terms ofthe number of species and groups investigated andalso in terms of consistency of definitions used acrossstudies prevents us from concluding that there areany systematic differences between species (yet), atthe very least we can say that hands play an importantrole in gestural communication among primates.Considering the function of gestural communi-cation, monkeys and apes use the majority of theirgestures to request actions like grooming, play ormating. They use their gestures in a dyadic way andusually not to communicate about events or objectsoutside their dyad, but to request certain actions,expecting an immediate response [48]. In case therecipient is not reacting, they will continue to gestureuntil they finally receive the appropriate response oftheir social partner [49-51]. Apes do take intoaccount the visual access of others (see [52] for areview), use visual gestures only if the recipient isattending [18,53], or use other strategies like movinginto someone’s visual field before starting to gesture[50,54]. In other words, they take into account thebehaviour of others and adjust their communicativemeans accordingly. However, there are inconsistentresults as to to what extent apes are actually able toalter their gestures if their first gesture was not success-ful – for chimpanzees and orangutans, it is shown thatmost often the same gesture is repeated [50,55], whilegorillas seem to show more flexibility in alternating thegestures they use to achieve a certain goal [49]. Itshould be considered that for interactions with ahuman experimenter, both chimpanzees and orangu-tans were shown to not only substitute, but alsoelaborate their gestures depending on the behaviourof a human in case their goal was not met [19,56].Another much-debated topic is the question ofpointing in non-human great apes. In captivity, greatapes and also some monkey species point to requestfood, tools or particular actions from humans[57-61]. Pointing in great apes represents a flexible,intentional behaviour, since the use of this gesture isadjusted to the attentional state of the human and itoccurs in combination with other signals such asfacial expressions and vocalizations [19,57,62,63].Pointing is also frequently used by language-trainedapes [60,64,65], where it often resembles the form ofthe pointing gesture of Western cultures with the armand index finger extended [66] .This content downloaded from150.135.174.98 on Thu, 03 Dec 2020 03:06:04 UTCAll use subject to httpReview. Gesture origins К. Liebal & J. Call 121However, unlike human infants that also point toshow objects, to share attention upon things, or eveninform others [67], non-human primates usuallypoint to request things or actions in their interactionswith humans. The vast majority of great apes’ pointinggestures therefore fall within the category of so-calledimperative gestures, which consist of the ape usingthe gesture to obtain something that they want fromthe human (see [68] for a review). Great apes alsoinform a human by pointing to the location of ahidden tool, but with the aim that the human canuse it to retrieve food for the ape [61]. Unlikehumans, however, non-human primates rarely (ifever) use pointing or other gestures aimed at sharingan attitude about the designated referent (expressivedeclaratives sensu Tomasello [69], e.g. [70,71]).Most importantly, pointing for conspecifics andthus the sharing of information is a rather rare event innon-human primates [72]. There is one report aboutone incidence of pointing in wild bonobos [73], andsome studies with language-trained apes mention theuse of pointing gestures in interactions with other apes[65] . However, note that the communicative behaviourof those language-trained apes is largely influencedby their raising history and thus their close proximityto the human culture [66,74]. Therefore, pointing forother conspecifics is extremely rare among wild andcaptive, non-enculturated apes. The flexible and inten-tional use of this gesture has been only systematicallydocumented for interactions with humans. Gómez[75] argues that captive non-human primates arerestricted by cages and therefore use humans as toolsto make them do things for them. Interestingly, anuncaged hand-reared gorilla grabbed the hand of thehuman and took him to the desired object or target ofaction and therefore preferred contact gestures insteadof pointing [70]. Therefore, it seems unlikely that theysimply learn to point by trial and error, but it issuggested that they recruit existing cognitive skills intothis referential form of communication [75]. Formonkeys, the situation seems to be different, sincepointing seems to be ritualized from previously reachingfor the food [75].To summarize, great apes and to some extent alsogibbons and monkeys use a variety of manual ges-tures to communicate with other group members,mostly to request immediate actions of their socialpartner. Thus, they use their gestures mostly in adyadic, imperative way to get others to do somethingfor them. Interestingly, Bard [76,77] referred to ges-tural communication as ‘social tool use’, which is alsoreflected in the use of pointing gestures in inter-actions with humans. Unlike humans, non-humanprimates do not point for conspecifics and their ges-tures are often derived from functional actionsrather than created as arbitrary ones for communica-tive purposes ([15], pp. 37-40; [78]), although thereare single reports about iconic gestures in gorillas andthe use of pantomime in orangutans [79,80]. Onepossible explanation for those observed differencesbetween different groups of great apes may bebased on the different ways in which gestures areacquired. In the next section, we turn our attentionto this issue.Phil. Trans. R. Soe. В (2012)(c) Gesture originsSince the focus of this paper is on hand-based gestures,we begin this section with a brief description of howhands are used for the purpose of communication bygreat apes and monkeys. Hands did not evolve as com-municative devices in the first place. In fact, the hands ofprimates are characterized by an extraordinary degreeof primitiveness [81], since the basic, five-fingeredappearance is shared not only with other mammals,but even other vertebrates. Still, only in primatesdoes the hand serve a variety of functions includinglocomotion, manipulation and communication [82].Moreover, each of these functions is represented by avariety of forms. Thus, locomotion can include thingslike walking, climbing, jumping or brachiating. Manipu-lation can include actions such as touching, holding orgrasping and more complex forms that combine thesebasic forms with other more elaborate actions thatenable primates to engage in a range of fine-grainedactivities such as grooming and tool use.From an evolutionary point of view, Napier ([81],p. 14) noted a . .trend … to emancipate the handsfrom weight-bearing to sensitive and delicate multipur-pose tools’. However, those different functions arenot representative for all primate species but verymuch depend on the differentiation of the hand in thedifferent taxa. While many monkeys and apes haveprehensile hands with nails and in some cases evenindependently movable or opposable thumbs, otherprimates such as marmosets and tamarins lack thosefeatures. Moreover, the gradual shrinkage of thehands’ palmar pads in phylogeny correlates with anincrease in prehensility and tactile sensitivity [81].With the emancipation of forelimbs for manipulatorypurposes, the stage is set for the development of handsas communicative devices. Indeed, it is not hard tofind potential commonalities between manipulativeactivity and communicative displays. For instance,monkeys and apes touch, push or pull other’s furduring communication. Apes beg for food by placing acupped hand under the chin of a potential food donoras if to catch food that may fall out. Even in the caseof locomotor activity, we can find connections betweenlocomotion and communication.An intriguing and contentious issue refers to theorigin of those communicative displays. One possibilityis that they evolved over evolutionary time solely forcommunicative purposes or that they originally evolvedfor one function (e.g. locomotion) and were co-optedand reused for a communicative function. Alternatively,communicative gestures may have become ritualizednot over evolutionary time but in interactions betweenindividuals and thus over a much shorter time span,an individual’s lifetime. Next, we turn our attentionto the possible changes involved in gesture origindepending on whether changes take place overevolutionary time (phylogenesis) or an individual’slifetime (ontogenesis).(d) Phylogenetic originsAnimal communication can be very complex and highlyritualized. Perhaps the most famous example is thebee ‘language’ consisting of different dances to indicateThis content downloaded from150.135.174.98 on Thu, 03 Dec 2020 03:06:04 UTCAll use subject to http122 К. Liebal & J. Call Review . Gesture originsthe position of food resources to other members ofthe hive [83]. Ritualized communication is not onlyfound in invertebrates. There are many examples fromvertebrates, including the complex mating displaydance of the stickleback or some lek breeding birds[84,85]. In some cases, the communicative displaysare composed of a set of discrete actions that follow afixed sequence, while in other cases they are constitutedby single units. Such signals are displayed by allindividuals of the species under a set of predeterminedconditions and, critically, they appear even if individualshad no opportunity to observe or interact with otherindividuals to acquire them.Whereas some communicative displays seem tohave evolved for communicative purposes only, otherdisplays appear to have been ‘borrowed’ from othercontexts and thus from movements that previouslyhad no communicative function via a process calledphylogenetic ritualization [86]. For instance, domi-nance signals such as mounting in monkeys are likelyto have evolved from mating behaviour, while somecourting displays in birds include elements of foragingbehaviour. This principle of derived activities [87] refersto actions that originally served a different function butwere borrowed and modified to some extent to accom-plish a communicative function, in some cases even ina different context from its original one.If we assume that phylogenetic ritualization is themain mechanism underlying gestural communicationin non-human primates, then repertoires of each speciesshould be highly uniform and species-specific gesturesshould be used even if individuals never had contactwith another conspecific. Gestures appear fully formedeven when subjects have not had a chance to interactwith other individuals. Ground-slapping and chest-beating would be examples of these behaviours [88].However, that they are phylogenetically ritualized doesnot mean that they are totally inflexible because, at thevery least, they are deployed in the right circumstancesand the existence of appropriate substrates/elementsdetermines their appearance. A phylogenetic origin ofgestures would mean that all members of a given speciesshould inherit their gestural repertoire, as is the case forvocalizations and, provided with the right conditions, allmembers of the species would display them. However, itis important to consider that some gestures might belimited to certain developmental stages, resulting inspecies-typical gestures that are restricted to particularage classes.(e) Ontogenetic originsAn alternative mechanism for the origin of gesturesentails individuals acquiring them during their inter-actions with conspecifics during their lifetimes ratherthan inheriting them as postulated above. One suchprocess that involves two individuals mutually shapingeach other’s behaviour during the course of repea-ted interactions is called ontogenetic ritualization[29,89] . Initially, individuals use functional behavioursto affect their partner’s behaviour. For instance, whenthey want to embrace their partner, initially theysimply pull their partner towards themselves andwhen they are within reach, they embrace them.Phil. Trans. R. Soc. В (2012)Over repeated interactions, partners begin to antici-pate the individual’s goal and react before theindividual actually has a chance to pull the partner.Next, the individual (anticipating that their partnerwill react appropriately) does not actually pull butgives an even more abbreviated version of the pulland their partner reacts. Once this stage is reached,we can say that the instrumental action of pullinghas become ritualized into a communicative signal.Ontogenetic ritualization as the main mechanismof gesture acquisition would result in a high degreeof variability of individual repertoires and particularlyin the occurrence of idiosyncratic gestures, which areexclusive for single individuals only [27] . Idiosyncraticgestures, which were found in all great ape species (foran overview, see [18]), seem to rule out phylogeneticritualization and thus a genetic determination of anindividual’s gestural repertoire, since those instancesclearly indicate that new gestures can be acquiredduring an individual’s lifetime. Although we stillknow very little about how such an individuallylearned new gesture spreads across other group mem-bers, there is some evidence that such a transmissiontakes place, as was shown for the grooming handclaspin a captive group of chimpanzees [90] .Variability between groups is evident in the occur-rence of group-specific gestures, which are used bythe majority of individuals in one group, but areabsent in another group. Although group-specific ges-tures are infrequent, they are reported for chimpanzees[26], gorillas [27,38] and orangutans [28] in captivesettings, but also in wild populations, like the groominghandclasp of wild chimpanzees [91].Two basic kinds of gestures have been describedin this context: intention movements and attention-getters [92]. Intention movements result from theabbreviations of full-fledged behaviour. For instance,the gesture arm raise has been hypothesized to orig-inate from play hitting, initially a functionalbehaviour that acquires a value as a signal of impend-ing actions. Intention movements typically convey aclear message and are used in a restricted set ofsocial contexts. Moreover, their meaning and origincan be deduced based on use in those contexts. Thesecond kind of gesture is the so-called attention-getter. It is true that the name attention-getter is notvery fortunate because unlike what its name suggests,attention-getters are not just designed to captureattention. In fact, their main function may be to triggerothers into action, not to call their attention. That theyalso serve to capture attention may be a by-product.However, there are inconsistent results in terms ofwhether great apes actually use their gestures to attractthe attention of others. In interactions with conspeci-fics, chimpanzees use either poke at or throw stuff-both heavily tactile gestures – to attract the attentionof the unattending individual [92]. However, thisseems to account only for those particular gestures,since further research found that chimpanzees alsouse auditory gestures more often towards an attentiverecipient and tactile gestures were used regardless ofthe attentional state of the recipient [26,50]. In otherwords, tactile and auditory gestures are not used par-ticularly often if the recipient is not attending.This content downloaded from150.135.174.98 on Thu, 03 Dec 2020 03:06:04 UTCAll use subject to httpReview . Gesture origins К. Liebal & J. Call 123However, in interactions with humans, orangutans,gorillas and chimpanzees do use attention-gettingbehaviours more when they interact with a humanwho is facing away compared with situations when thehuman is facing them [53,93-95]. The different find-ings for interactions between conspecifics on the onehand and interactions with humans on the other handmight also be explained by the constraints of thecaptive setting. When apes encountered a human withher back turned and they were given a choice betweenpositioning themselves in front of the human or usingan auditory gesture to call the human’s attention, allgreat apes species preferred to walk in front of thehuman to gesture [54]. Thus, similar to the use ofpointing gestures, the use of attention-getters mightdepend very much on the restraints of captivity.Compared with intention movements, attention-getters appear to be less context-dependent as theyappear in multiple contexts for multiple purposes.Additionally, unlike intention movements, it is noteasy to envisage a history of ontogenetic ritualizationfrom pre-existing social behaviours as their origin, sothat they are possibly also phylogenetic in origin.There is a second way in which individuals couldacquire gestures during ontogeny without requiringritualization: learning gestures by observation. Onepossibility is that the individual would copy the ges-tures that another individual is directing to her(second-person imitation). Another possibility is thatthe individual could observe two individuals gesturingto each other and acquires those gestures herself with-out directly interacting with others (third-personimitation). Interestingly, gestures learned by obser-vation walk an opposite path from those that areontogenetically ritualized. They are acquired fullyformed, the individual does not transform an existingbehaviour into a streamlined version that becomesthe gesture. The individual copies the streamlined ver-sion. The resulting outcome would be a high degree ofuniformity within the group, paired with substantialdifferences between groups because each group mayhave developed their own idiosyncratic gestures andtransmitted them across generations. This is clearlythe case in humans but it is unclear whether that isalso the case in non-human primates.(f) Phylogenetic versus ontogenetic origins ofgestures?There is currently some debate about what may be themost likely origin of gestures. We have indicated threepotential origins for gestures. Historically, observationallearning had been proposed as a main mechanismfor gesture acquisition. However, there are very littledata supporting the idea that apes learn gestures,especially visual gestures, by imitation [26]. Notethat the variability in gestural use within groups is aslarge as between groups. This is not what one wouldexpect based on imitation and cultural transmissionsince between-group variability should be higher thanwithin-group variability as is the case in humans.Unlike observational learning, ontogenetic rituali-zation can explain this pattern of results becausethe homogeneity within groups would be reduced byPhil. Trans. R. Soc. В (2012)the presence of idiosyncratic gestures developedby some individuals but not others. The reason foridiosyncrasy stems from the fact that certain dyadicinteractions between individuals are unique, forinstance, mothers and infants may follow differentcaregiving routines. In fact, according to Tomaselloet al. [29], the presence of idiosyncratic gestures is akey indicator of ontogenetic ritualization and evidenceagainst a phylogenetic origin of gestures.Genty et al. [37] have recently challenged this ideaand proposed that ape gestures are not ontogeneticallyritualized but appear fully formed in individuals. Thisphylogenetic origin of gestures leaves no room for modi-fication over time. They argue that the differencesbetween groups and the idiosyncrasy that has beendescribed are a consequence of the sampling methodsthat have been used. In particular, not enoughhours have been observed to be able to obtain thewhole repertoire of gestures. Thus, idiosyncrasy resultsfrom a low sampling effort as opposed to individualizedexperiences with other conspecifics. An increase in thesampling effort showed that all individuals used thesame gestures and virtually eliminated idiosyncraticgestures from the sample. This result casts somedoubt not only on ontogenetic ritualization but alsoon observational learning as acquisition mechanismsbecause there were no differences between groups.However, one limitation of this and many otherstudies is that they are not longitudinal and thereforethey cannot detect change either within an individual’slifespan or across generations. So, although all individ-uals use the same gestures, this does not prove thatgestures have not undergone an ontogenetic ritualiza-tion process. What is needed are long-term studiesactually investigating whether the gestures of greatapes (and monkeys) change over time. Additionally,studies that have investigated gestural acquisition ofapes in contact with humans have described the rituali-zation process [96]. One could argue that apes incontact with humans would be different, but this ishard to reconcile with the idea that human-rearedapes were requesting the same things (e.g. go to anotherlocation) as the ones living with their biologicalmothers. Nevertheless, it is true that the case for onto-genetic ritualization may have been overstated becauseas Genty et al. [37] point out, it is difficult to envisagea history of ontogenetic ritualization for some gesturessuch as chest beating or ground slapping, although itis also true that ontogenetic ritualization may still be aviable alternative for other gestures such as gentletouch or arm raise.After discussing the origins of gestures in non-humanprimates and their close link to actions, we will nowbriefly refer to some of the current theories on languageevolution and the role gestures might have played,before we address the question of laterality in gestureuse in non-human primates.
LANGUAGE ORIGINS (OUT OF ACTIONS)(a) Gestural origin of human languageThe origin of human language is a fiercely debatedquestion, with some scholars favouring a vocal origin(e.g. [97]) and others emphasizing gestures asThis content downloaded from150.135.174.98 on Thu, 03 Dec 2020 03:06:04 UTCAll use subject to http124 К. Liebal & J. Call Review. Gesture originsprecursors to human language (e.g. [43]). To ourknowledge, there is no coherent theory currently avail-able that has attempted to reconcile the two opposingsets of theories, which usually see themselves asmutually exclusive [46]. Gestural theories usuallyrefer to the very flexible use of gestures across differentcontexts and the fact that new gestures can be learnedand incorporated into a species repertoire [98]. Thediscovery of a mirror-neuron system for grasping inmonkeys [99] has nourished a variety of evolutionaryscenarios focusing on the role of gestures in thisprocess, since mirror neurons represent the linkbetween manual, practical actions and communication.(b) From action to languageMirror neurons allow macaques to establish a linkbetween performing an action and being able to recog-nize it [99] . Interestingly, these neurons are located in abrain area that is homologous to Broca’s area in thehuman brain responsible for processing language.Mirror neurons therefore might have played an impor-tant role in the evolution of human language, sincethey were already present in our ancestors representingthe neural prerequisite for the development of inter-individual communication and finally of speech [100].Next, we will give a brief overview of theories suggestinga gestural origin, and second, we will turn to the latera-lization of hand use while gesturing and the evidencecurrently available for non-human primates.(c) Gesture and lateralityGestural theories of language evolution often refer tothe link between lateralization of hand use and language[43]. In humans, the motor systems controlling bothmanual and oral movements are usually lateralized tothe left hemisphere [101]. Therefore, the majority ofthe human population is right-handed, with the lefthemisphere controlling movements of the right hand.Furthermore, both language production and compre-hension are located in distinct areas of the left brainhemisphere [102]. The close link between languageand manual actions becomes evident in studies showingthat while speaking, humans gesture significantly morewith their right hand compared with their left hand[103]. This suggests that the functional asymmetryis not specific for one modality only, and that theproduction of speech apparently also activates motorareas in the left hemisphere, resulting in an increaseduse of the right hand [104] .Comparable evidence has been found for non-human primates since they show a preference forusing their right hand for different manual actionsincluding gestures while vocalizing [105,106]. Thesefindings suggest that the lateralization of manual andoral movements represents a trait shared by bothhumans and other primates.However, results for the preference for one handand particularly the right hand are not completely con-sistent [107,108]. Although a hand preference is foundin many monkey and ape species for different manualactions such as carrying, tool use and locomotion[105,109-111], hand preference is often task-specificand often only evident on an individual, but notPhil. Trans. R. Soc. В (2012)species level [112,113]. So far, there is little evidencethat gestures used to communicate with conspeci-fics – and thus not in interactions with humans whenbegging for food – are mainly produced with theright hand. To our knowledge, there is only onestudy showing that baboons use their right handwhile they gesture, but not when they perform non-communicative actions [22] . In interactions withhumans, however, there is clear evidence that chim-panzees use their right hand preferentially whilegesturing [114-116], and they also used their righthand significantly more while producing gestures com-pared with other manual actions. Hopkins et al. [1 17]therefore concluded that the left-hemisphere specializ-ation for language may have evolved initially fromasymmetries in manual gestures in the common ances-tor of chimpanzees and humans, rather than fromhand use associated with other, non-communicativemotor actions such as tool use and bimanual actions.The laterality of hand use in chimpanzees is alsoreflected in neuroanatomical asymmetries, since chim-panzees that preferably gesture with their right handalso have larger inferior frontal gyri in the left hemi-sphere than those apes that do not show consistenthand use while gesturing [118].To summarize, those studies indicate that manualgestures of at least chimpanzees are lateralized, andthis functional asymmetry is also associated with asym-metries in the corresponding neural substrate.Hopkins et al. [117] therefore suggest that the domi-nance of the left hemisphere for language has evolvedfrom a gestural communication system already latera-lized in the left hemisphere in our common ancestor5-7 Ma.
CONCLUSIONIn our contribution, we wanted to highlight that thehands of non-human primates, and particularly thoseof the great apes, are suitable tools to perform a varietyof gestures of different modalities. They are used toachieve a range of different social goals and display ahigh degree of flexibility as indicated by the possibilityof acquiring new gestures often outside of what wouldbe the species-specific repertoire. Still, gestures ofnon-human primates are different from human ges-tures in many aspects, since they are mostly used ina dyadic and imperative way, and they also lack thehigh level of abstraction typical for human gestures.Thus, gestures of non-human primates may emergefrom actions via three potential pathways.The high degree of variability between individualrepertoires, the occurrence of idiosyncratic gesturesand thus the creation of new gestures support theidea that ontogenetic ritualization may be involvedin the origin of some gestures. However, other gesturesappear more or less fully formed even in the absenceof conspecifics, thus indicating a strong genetic predis-position to develop certain gestures. Finally, someform of social learning might also be implicatedeither in the form of facilitating the appearance ofsome gestures or perhaps even the acquisition ofnovel gestures, although this still needs to be sup-ported by empirical evidence.This content downloaded from150.135.174.98 on Thu, 03 Dec 2020 03:06:04 UTCAll use subject to httpReview. Gesture origins К. Liebal & J. Call 125After elaborating on the close link between manualactions and gestural communication, we turned to thequestion of whether gestures of non-human primatesare lateralized as many human manual actions includ-ing certain gesture types are. There is some evidencefor right-handedness at least in captive chimpanzees,and, interestingly, they use their right hand evenmore while vocalizing, thus suggesting a close linkbetween the manual and oral movements. This factis often used to support a gestural origin of humanlanguage, since the functional asymmetry of handuse while gesturing is also present in the neural sub-strate of chimpanzees, suggesting some continuity inour phylogenetic history. However, one must keep inmind that the evidence of laterality in chimpanzeesand other non-human primate species at the popu-lation level is quite mixed. This means that it may betoo early to generalize a right-hand preference for ges-ture use in our closest relatives.There is much to be done in the future to trace theorigins of gestures. Longitudinal studies are especiallyimportant as they can throw light on how gesturesactually emerge in both monkeys and apes. Someresearch effort devoted to non-great ape species wouldbe particularly welcome. Otherwise the field runsthe risk of underestimating what aspects of gesturalcommunication that are common to human and non-human apes are already present in monkeys. Finally,there is much work to be done in terms of unifyingconcepts and criteria across the various disciplinesthat conduct research on gestural communication.We would like to thank Michael Tomasello, Simone Pika andCornelia Mueller for fruitful discussions on this topic, andDaniel Haun as well as Erica Cartmill and an anonymousreviewer for their very helpful comments on the manuscript.REFERENCES1 Gómez, J. C. 1990 The emergence of intentionalcommunication as a problem-solving strategy in gorilla.In 4 Language 9 and intelligence in monkeys and apes (edsS. T. Parker & K. R. Gibson), pp. 333-355.Cambridge, UK: Cambridge University Press.2 Simpson, J. A. & Weiner, E. S. C. (eds) 1998 TheOxford English Dictionary, 2nd edn. 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