Of the importance to reconsider individual variability in infant studies: Family traits do impact turn-taking perception in 6-month-olds.

Turn-taking is a universal pattern of human conversations characterized by a fast exchange of turns between speakers and an avoidance of overlaps. Language is embedded in this conversational skill acquired well before it during infancy, through everyday interactions with caregivers. The earliness of this skill and its link with language allows us to test whether social environment shapes early language development. We therefore study turn-taking perception of 6-month-old infants by measuring their gazes during video presentation of three different conversational situations where the turn is explicitly given, normally taken or taken with an overlap. We studied 51 infants to cover several family and infant characteristics: infants' sex, presence of siblings, and family socioeconomic status (SES). We found that infants looked more at the second speaker when she overlapped the first speaker than in the other situations, but not all infants were equally sensitive. Indeed, infants from high-SES families reacted differently to the three situations, while infants from the two lower SES categories did not. Also, only singletons reacted differently by looking more at the second speaker after the overlapping and turn-giving situations, and not after the turn-taking situation. Our results emphasize the importance of early social experiences on language development.

Human laterality for manipulation and gestural communication: A study of beach-volleyball players during the Olympic Games.

Comparative studies can help understand better brain functional lateralization for manipulation and language. This study investigated and compared, for the first time, human adults' laterality for manipulation and gestures in a non-experimental social context. We analysed the manual laterality of 48 beach volleyball athletes for four frequently expressed behaviours: a complex throwing action (jump serve) and three gestures (CLAP HAND, PUMP FIST and SLAP HAND-TO-HAND). We evaluated population-level laterality bias for each of the four behaviours separately, compared manual laterality between behaviours and investigated factors influencing gestural laterality. We furthered our between-gestures comparison by taking into account three categories of factors simultaneously: gesture characteristics (sensory modality), interactional context components (positions of interactants and emotional valence), and individual demographic characteristics (age, sex and country). Our study showed that (1) each behaviour considered presented a population-level right-hand bias, (2) differences of laterality between behaviours were probably related to gesture sensory modality and (3) signaller's laterality was modulated differently in relation to positions of interactants, emotional valence, age and sex. Our results support the literature suggesting that left-hemisphere specialization for manipulation and language (speech and gestures) may have evolved from complex manual activities such as throwing and from gestural communication.

The origins of gestures and language: history, current advances and proposed theories.

Investigating in depth the mechanisms underlying human and non-human primate intentional communication systems (involving gestures, vocalisations, facial expressions and eye behaviours) can shed light on the evolutionary roots of language. Reports on non-human primates, particularly great apes, suggest that gestural communication would have been a crucial prerequisite for the emergence of language, mainly based on the evidence of large communication repertoires and their associated multifaceted nature of intentionality that are key properties of language. Such research fuels important debates on the origins of gestures and language. We review here three non-mutually exclusive processes that can explain mainly great apes' gestural acquisition and development: phylogenetic ritualisation, ontogenetic ritualisation, and learning via social negotiation. We hypothesise the following scenario for the evolutionary origins of gestures: gestures would have appeared gradually through evolution via signal ritualisation following the principle of derived activities, with the key involvement of emotional expression and processing. The increasing level of complexity of socioecological lifestyles and associated daily manipulative activities might then have enabled the acquisition and development of different interactional strategies throughout the life cycle. Many studies support a multimodal origin of language. However, we stress that the origins of language are not only multimodal, but more broadly multicausal. We propose a multicausal theory of language origins which better explains current findings. It postulates that primates' communicative signalling is a complex trait continually shaped by a cost-benefit trade-off of signal production and processing of interactants in relation to four closely interlinked categories of evolutionary and life cycle factors: species, individual and context-related characteristics as well as behaviour and its characteristics. We conclude by suggesting directions for future research to improve our understanding of the evolutionary roots of gestures and language.

Captive chimpanzees' manual laterality in tool use context: Influence of communication and of sociodemographic factors.

Understanding variations of apes' laterality between activities is a central issue when investigating the evolutionary origins of human hemispheric specialization of manual functions and language. We assessed laterality of 39 chimpanzees in a non-communication action similar to termite fishing that we compared with data on five frequent conspecific-directed gestures involving a tool previously exploited in the same subjects. We evaluated, first, population-level manual laterality for tool-use in non-communication actions; second, the influence of sociodemographic factors (age, sex, group, and hierarchy) on manual laterality in both non-communication actions and gestures. No significant right-hand bias at the population level was found for non-communication tool use, contrary to our previous findings for gestures involving a tool. A multifactorial analysis revealed that hierarchy and age particularly modulated manual laterality. Dominants and immatures were more right-handed when using a tool in gestures than in non-communication actions. On the contrary, subordinates, adolescents, young and mature adults as well as males were more right-handed when using a tool in non-communication actions than in gestures. Our findings support the hypothesis that some primate species may have a specific left-hemisphere processing gestures distinct from the cerebral system processing non-communication manual actions and to partly support the tool use hypothesis.

A multifactorial investigation of captive gorillas' intraspecific gestural laterality.

Multifactorial investigations of intraspecific laterality of primates' gestural communication aim to shed light on factors that underlie the evolutionary origins of human handedness and language. This study assesses gorillas' intraspecific gestural laterality considering the effect of various factors related to gestural characteristics, interactional context and sociodemographic characteristics of signaller and recipient. Our question was: which factors influence gorillas' gestural laterality? We studied laterality in three captive groups of gorillas (N = 35) focusing on their most frequent gesture types (N = 16). We show that signallers used predominantly their hand ipsilateral to the recipient for tactile and visual gestures, whatever the emotional context, gesture duration, recipient's sex or the kin relationship between both interactants, and whether or not a communication tool was used. Signallers' contralateral hand was not preferentially used in any situation. Signallers' right-hand use was more pronounced in negative contexts, in short gestures, when signallers were females and its use increased with age. Our findings showed that gorillas' gestural laterality could be influenced by different types of social pressures thus supporting the theory of the evolution of laterality at the population level. Our study also evidenced that some particular gesture categories are better markers than others of the left-hemisphere language specialization.

Human laterality for manipulation and gestural communication related to 60 everyday activities: Impact of multiple individual-related factors.

Literature on laterality emphasises the importance of implementing a comprehensive investigation of humans' and non-humans' laterality for both non-communication and communication functions. Adopting a global approach should enhance our understanding of the mechanistic drivers of human brain functional lateralisation and help to explore further the nature of the left-hemispheric systems for both functions. This study investigated human laterality for both functions by taking, for the first time, numerous behaviours and multiple potential influential factors into consideration. We analysed replies to the Rennes Laterality Questionnaire that takes into account simultaneously: participants' behavioural, demographic and social characteristics as well as their genetic/social learning components and collateral factors such as health problems. We collected and analysed a large data set including 450,220 item responses (317,594 items related to behavioural laterality and 132,626 items related to personal information) by 5904 participants. The majority of participants were right-lateralised for nine behavioural categories. The laterality of right-lateralised individuals for the behaviours considered varied very little between behavioural categories, contrary to the laterality of ambiguously- and left-lateralised individuals. This is the first evidence of the stability of right-lateralised individuals and the relative flexibility of ambiguously- and left-lateralised individuals with regards to behavioural laterality related to 60 everyday activities. Moreover, the laterality patterns of our study population were linked in particular to the following individual-related factors: demographic characteristics (current continent of residence), social characteristics (socio-professional group and study level), and collateral factors (writing hand, forced right-hand use, guiding eye, and health problems related to ear and back). We discuss our findings in relation to the evolutionary roots of human brain lateralisation for non-communication and communication functions.

Captive gorillas' manual laterality: The impact of gestures, manipulators and interaction specificity.

Relationships between humans' manual laterality in non-communicative and communicative functions are still poorly understood. Recently, studies showed that chimpanzees' manual laterality is influenced by functional, interactional and individual factors and their mutual intertwinement. However, what about manual laterality in species living in stable social groups? We tackled this question by studying three groups of captive gorillas (N=35) and analysed their most frequent manual signals: three manipulators and 16 gesture types. Our multifactorial investigation showed that conspecific-directed gestures were overall more right-lateralized than conspecific-directed manipulators. Furthermore, it revealed a difference between conspecific- and human-directed gestural laterality for signallers living in one of the study groups. Our results support the hypothesis that gestural laterality is a relevant marker of language left-brain specialisation. We suggest that components of communication and of manipulation (not only of an object but also of a conspecific) do not share the same lateralised cerebral system in some primate species.

Assessment and analysis of human laterality for manipulation and communication using the Rennes Laterality Questionnaire.

Despite significant scientific advances, the nature of the left-hemispheric systems involved in language (speech and gesture) and manual actions is still unclear. To date, investigations of human laterality focused mainly on non-communication functions. Although gestural laterality data have been published for infants and children, relatively little is known about laterality of human gestural communication. This study investigated human laterality in depth considering non-communication manipulation actions and various gesture types involving hands, feet, face and ears. We constructed an online laterality questionnaire including 60 items related to daily activities. We collected 317 594 item responses by 5904 randomly selected participants. The highest percentages of strong left-lateralized (6.76%) and strong right-lateralized participants (75.19%) were for manipulation actions. The highest percentages of mixed left-lateralized (12.30%) and ambidextrous (50.23%) participants were found for head-related gestures. The highest percentage of mixed right-lateralized participants (55.33%) was found for auditory gestures. Every behavioural category showed a significant population-level right-side bias. More precisely, participants were predominantly right-lateralized for non-communication manual actions, for visual iconic, visual symbolic, visual deictic (with and without speech), tactile and auditory manual gestures as well as for podial and head-related gestures. Our findings support previous studies reporting that humans have left-brain predominance for gestures and complex motor activities such as tool-use. Our study shows that the Rennes Laterality Questionnaire is a useful research instrument to assess and analyse human laterality for both manipulation and communication functions.

Intraspecific gestural laterality in chimpanzees and gorillas and the impact of social propensities.

A relevant approach to address the mechanisms underlying the emergence of the right-handedness/left-hemisphere language specialization of humans is to investigate both proximal and distal causes of language lateralization through the study of non-human primates' gestural laterality. We carried out the first systematic, quantitative comparison of within-subjects' and between-species' laterality by focusing on the laterality of intraspecific gestures of chimpanzees (Pan troglodytes) and gorillas (Gorilla gorilla) living in six different captive groups. We addressed the following two questions: (1) Do chimpanzees and gorillas exhibit stable direction of laterality when producing different types of gestures at the individual level? If yes, is it related to the strength of laterality? (2) Is there a species difference in gestural laterality at the population level? If yes, which factors could explain this difference? During 1356 observation hours, we recorded 42335 cases of dyadic gesture use in the six groups totalling 39 chimpanzees and 35 gorillas. Results showed that both species could exhibit either stability or flexibility in their direction of gestural laterality. These results suggest that both stability and flexibility may have differently modulated the strength of laterality depending on the species social structure and dynamics. Furthermore, a multifactorial analysis indicates that these particular social components may have specifically impacted gestural laterality through the influence of gesture sensory modality and the position of the recipient in the signaller's visual field during interaction. Our findings provide further support to the social theory of laterality origins proposing that social pressures may have shaped laterality through natural selection.

Correction: Can Population-Level Laterality Stem from Social Pressures? Evidence from Cheek Kissing in Humans.

[This corrects the article DOI: 10.1371/journal.pone.0124477.].

Sex Differences in Language Across Early Childhood: Family Socioeconomic Status does not Impact Boys and Girls Equally.

Child sex and family socioeconomic status (SES) have been repeatedly identified as a source of inter-individual variation in language development; yet their interactions have rarely been explored. While sex differences are the focus of a renewed interest concerning emerging language skills, data remain scarce and are not consistent across preschool years. The questions of whether family SES impacts boys and girls equally, as well as of the consistency of these differences throughout early childhood, remain open. We evaluated consistency of sex differences across SES and age by focusing on how children (N = 262), from 2;6 to 6;4 years old, from two contrasting social backgrounds, acquire a frequent phonological alternation in French - the liaison. By using a picture naming task eliciting the production of obligatory liaisons, we found evidence of sex differences over the preschool years in low-SES children, but not between high-SES boys and girls whose performances were very similar. Low-SES boys' performances were the poorest whereas low-SES girls' performances were intermediate, that is, lower than those of high-SES children of both sexes but higher than those of low-SES boys. Although all children's mastery of obligatory liaisons progressed with age, our findings showed a significant impeding effect of low-SES, especially for boys.

The maintenance of regional dialects: a matter of gender? Boys, but not girls, use local varieties in relation to their friends' nativeness and local identity.

The linguistic diversity enduring beyond institutional pressures and social prejudices against non-standard dialects questions the social forces influencing language maintenance across generations and how children contribute to this process. Children encounter multi-dialectal interactions in their early environment, and increasing evidence shows that the acquisition of sociolinguistic variation is not a side issue but an inherent part of the general acquisition process. Despite these recent advances in sociolinguistic acquisition, children's sociolinguistic uses remain under-studied in relation to peer social networks and the ability to use dialect for identity purposes. Our study focused on a grammatical sociolinguistic variable consisting of the alternation between a regional and a standard variant of the third person object pronoun in French. The regional variant is a remnant of the Francoprovençal language and its usage by adults is strongly associated with local identity in the French Alps. We described, using questionnaires, the social networks of 117 10-11 year-old girls and boys living in the same restricted rural area. Thirteen native target children (7 girls and 6 boys) were selected from the sample, as well as 39 same-sex friends chosen according to their place of birth (native vs. non-native) and the duration of their friendship with the targets (number of years they have known each other). The target children were recorded during spontaneous dyadic conversations during free play at school with each category of friends. Target boys, but not girls, used the regional variant significantly more frequently with their long-term native friends than with their non-native friends. This adjustment mirrored their partners' uses. Moreover, with long-term native friends, boys used the regional variant twice as frequently as girls. Boys appeared thus as key actors in the maintenance and the diffusion of regional cues in local social networks.

Age- and sex-dependent contact call usage in Japanese macaques.

The question of the flexibility of nonhuman primate vocal communication remains open today, especially due to early evidence of innately guided vocal production. However, socially determined flexibility can be found when the debate is moved from vocal structure to vocal usage. While increasing evidence shows that the audience quality influences the vocal behaviour of nonhuman primates, the impact of the caller's characteristics has been far less studied. Here, we tested the influence of an individual's sex and age on the usage style of contact calls. We recorded contact calls of male and female Japanese macaques and compared the vocal usage styles of approximately 1-year-old juveniles with those of adults at various ages. We found, first, important differences in call usage style between juveniles and adults, the latter forming temporally ruled vocal exchanges respecting an interindividual turntaking principle. Moreover, sex differences were substantial in adults but nonexistent in juveniles. Finally, age continued to influence female vocal behaviour during adulthood, whereas dominance rank explained differences between adult males. Two nonexclusive mechanisms can explain this phenomenon, that is, a socially guided development of the appropriate form of calling versus an emotional maturation to control call emission, opening new lines of research on nonhuman primate vocal development of appropriate usages.

Boys and girls on the playground: sex differences in social development are not stable across early childhood.

Sex differences in human social behaviors and abilities have long been a question of public and scientific interest. Females are usually assumed to be more socially oriented and skillful than males. However, despite an extensive literature, the very existence of sex differences remains a matter of discussion while some studies found no sex differences whereas others reported differences that were either congruent or not with gender stereotypes. Moreover, the magnitude, consistency and stability across time of the differences remain an open question, especially during childhood. As play provides an excellent window into children's social development, we investigated whether and how sex differences change in social play across early childhood. Following a cross-sectional design, 164 children aged from 2 to 6 years old, divided into four age groups, were observed during outdoor free play at nursery school. We showed that sex differences are not stable over time evidencing a developmental gap between girls and boys. Social and structured forms of play emerge systematically earlier in girls than in boys leading to subsequent sex differences in favor of girls at some ages, successively in associative play at 3-4 years, cooperative play at 4-5 years, and social interactions with peers at 5-6 years. Preschool boys also display more solitary play than preschool girls, especially when young. Nevertheless, while boys catch up and girls move on towards more complex play, sex differences in social play patterns are reversed in favor of boys at the following ages, such as in associative play at 4-5 years and cooperative play at 5-6 years. This developmental perspective contributes to resolve apparent discrepancies between single-snapshot studies. A better understanding of the dynamics of sex differences in typical social development should also provide insights into atypical social developments which exhibit sex differences in prevalence, such as autism.

The mere perception of eye contact increases arousal during a word-spelling task.

Eye contact is a highly salient and fundamentally social signal. This entails that the mere perception of direct gaze may trigger differentiated neurobehavioral responses as compared to other gaze directions. We investigated this issue using a visual word-spelling task where faces under different gaze directions and head orientations were displayed on-screen concomitantly with the words. We show evidence for automatic increase of skin conductance response (SCR), indicative of arousal, associated with the perception of direct gaze as compared to both averted gaze and closed eyes. Moreover, the perception of averted gaze was associated with an increase of electromyographic (EMG) corrugator activity. These effects were observed in two demanding word-spelling tasks, but not in a simple letter decision task. We propose to interpret these findings in terms of the social value of direct and averted gaze and conclude that some circumstances such as the task at hand may be essential for uncovering the neurobehavioral responses associated with the perception of others' gaze.

Stability and flexibility in preschoolers' social networks: a dynamic analysis of socially directed behavior allocation.

The author studied preschoolers' social networks by investigating the allocation of children's social investment within and across time in a classroom of a French nursery school during an academic year. Observations of children's social exchanges during free play revealed that social behaviors were directed toward particular group members. After an important turnover in the peer group at the beginning of the school year, the social network became more structured. Children's strong associations were mostly same sex and small sized. Even if the stability of children's connections remained low, it increased over time. High-frequency partners as well as same-sex partners were more likely to be maintained over time. These findings as well as conceptual and methodological issues are discussed from a developmental perspective.