Because Broca's area is known to be involved in many cognitive functions, including language, music, and action processing, several attempts have been made to propose a unifying theory of its role that emphasizes a possible contribution to syntactic processing. Recently, we have postulated that Broca's area might be involved in higher-order chunk processing during implicit learning of a motor sequence. Chunking is an information-processing mechanism that consists of grouping consecutive items in a sequence and is likely to be involved in all of the aforementioned cognitive processes. Demonstrating a contribution of Broca's area to chunking during the learning of a nonmotor sequence that does not involve language could shed new light on its function. To address this issue, we used offline MRI-guided TMS in healthy volunteers to disrupt the activity of either the posterior part of Broca's area (left Brodmann's area [BA] 44) or a control site just before participants learned a perceptual sequence structured in distinct hierarchical levels. We found that disruption of the left BA 44 increased the processing time of stimuli representing the boundaries of higher-order chunks and modified the chunking strategy. The current results highlight the possible role of the left BA 44 in building up effector-independent representations of higher-order events in structured sequences. This might clarify the contribution of Broca's area in processing hierarchical structures, a key mechanism in many cognitive functions, such as language and composite actions.

Human genetic and phenotypic diversity declines with distance from Africa, as predicted by a serial founder effect in which successive population bottlenecks during range expansion progressively reduce diversity, underpinning support for an African origin of modern humans. Recent work suggests that a similar founder effect may operate on human culture and language. Here I show that the number of phonemes used in a global sample of 504 languages is also clinal and fits a serial founder-effect model of expansion from an inferred origin in Africa. This result, which is not explained by more recent demographic history, local language diversity, or statistical non-independence within language families, points to parallel mechanisms shaping genetic and linguistic diversity and supports an African origin of modern human languages.

Mirror neurons (MNs) are a fascinating class of cells originally discovered in the ventral premotor cortex (PMv) and, subsequently, in the inferior parietal lobule (IPL) of the macaque, which become active during both the execution and observation of actions. In this review, I will first highlight the mounting evidence indicating that mirroring others' actions engages a broad system of reciprocally connected cortical areas, which extends well beyond the classical IPL-PMv circuit and might even include subcortical regions such as the basal ganglia. Then, I will present the most recent findings supporting the idea that the observation of one's own actions, which might play a role in the ontogenetic origin and tuning of MNs, retains a particular relevance within the adult MN system. Finally, I will propose that both cortical and subcortical mechanisms do exist to decouple MN activity from the motor output, in order to render it exploitable for high-order perceptual, cognitive, and even social functions. The findings reviewed here provide an original framework for envisaging the main challenges and experimental directions of future neurophysiological and neuroanatomical studies of the monkey MN system.

Research has shown a close relationship between gestures and language development. In this study, we investigate the cross-lagged relationships between different types of gestures and two lexicon dimensions: number of words produced and comprehended. Information about gestures and lexical development was collected from 48 typically developing infants when these were aged 0;9, 1;0 and 1;3. The European Portuguese version of the MacArthur-Bates Communicative Development Inventory: Words and Gestures (PT CDI:WG) was used. The results indicated that the total number of actions and gestures and the number of early gestures produced at 0;9 and at 1;0 year predicted the number of words comprehended three months later. Actions and gestures' predictive power of the number of words produced was limited to the 0;9-1;0 year interval. The opposite relationship was not found: word comprehension and production did not predict action and gestures three months later. These results highlight the importance of non-verbal communicative behavior in language development.

The widespread existence of cooperation is difficult to explain because individuals face strong incentives to exploit the cooperative tendencies of others. In the research reported here, we examined how the spread of reputational information through gossip promotes cooperation in mixed-motive settings. Results showed that individuals readily communicated reputational information about others, and recipients used this information to selectively interact with cooperative individuals and ostracize those who had behaved selfishly, which enabled group members to contribute to the public good with reduced threat of exploitation. Additionally, ostracized individuals responded to exclusion by subsequently cooperating at levels comparable to those who were not ostracized. These results suggest that the spread of reputational information through gossip can mitigate egoistic behavior by facilitating partner selection, thereby helping to solve the problem of cooperation even in noniterated interactions.

How does sign language compare with gesture, on the one hand, and spoken language on the other? Sign was once viewed as nothing more than a system of pictorial gestures without linguistic structure. More recently, researchers have argued that sign is no different from spoken language, with all of the same linguistic structures. The pendulum is currently swinging back toward the view that sign is gestural, or at least has gestural components. The goal of this review is to elucidate the relationships among sign language, gesture, and spoken language. We do so by taking a close look not only at how sign has been studied over the past 50 years, but also at how the spontaneous gestures that accompany speech have been studied. We conclude that signers gesture just as speakers do. Both produce imagistic gestures along with more categorical signs or words. Because at present it is difficult to tell where sign stops and gesture begins, we suggest that sign should not be compared with speech alone but should be compared with speech-plus-gesture. Although it might be easier (and, in some cases, preferable) to blur the distinction between sign and gesture, we argue that distinguishing between sign (or speech) and gesture is essential to predict certain types of learning and allows us to understand the conditions under which gesture takes on properties of sign, and speech takes on properties of gesture. We end by calling for new technology that may help us better calibrate the borders between sign and gesture.

How does gesturing help children learn? Gesturing might encourage children to extract meaning implicit in their hand movements. If so, children should be sensitive to the particular movements they produce and learn accordingly. Alternatively, all that may matter is that children move their hands. If so, they should learn regardless of which movements they produce. To investigate these alternatives, we manipulated gesturing during a math lesson. We found that children required to produce correct gestures learned more than children required to produce partially correct gestures, who learned more than children required to produce no gestures. This effect was mediated by whether children took information conveyed solely in their gestures and added it to their speech. The findings suggest that body movements are involved not only in processing old ideas, but also in creating new ones. We may be able to lay foundations for new knowledge simply by telling learners how to move their hands.

Human evolutionary scholars have long supposed that the earliest stone tools were made by the genus Homo and that this technological development was directly linked to climate change and the spread of savannah grasslands. New fieldwork in West Turkana, Kenya, has identified evidence of much earlier hominin technological behaviour. We report the discovery of Lomekwi 3, a 3.3-million-year-old archaeological site where in situ stone artefacts occur in spatiotemporal association with Pliocene hominin fossils in a wooded palaeoenvironment. The Lomekwi 3 knappers, with a developing understanding of stone's fracture properties, combined core reduction with battering activities. Given the implications of the Lomekwi 3 assemblage for models aiming to converge environmental change, hominin evolution and technological origins, we propose for it the name 'Lomekwian', which predates the Oldowan by 700,000 years and marks a new beginning to the known archaeological record.

Human ancestors first modified stones into tools 2.6 million years ago, initiating a cascading increase in technological complexity that continues today. A parallel trend of brain expansion during the Paleolithic has motivated over 100 years of theorizing linking stone toolmaking and human brain evolution, but empirical support remains limited. Our study provides the first direct experimental evidence identifying likely neuroanatomical targets of natural selection acting on toolmaking ability. Subjects received MRI and DTI scans before, during, and after a 2-year Paleolithic toolmaking training program. White matter fractional anisotropy (FA) showed changes in branches of the superior longitudinal fasciculus leading into left supramarginal gyrus, bilateral ventral precentral gyri, and right inferior frontal gyrus pars triangularis. FA increased from Scan 1-2, a period of intense training, and decreased from Scan 2-3, a period of reduced training. Voxel-based morphometry found a similar trend toward gray matter expansion in the left supramarginal gyrus from Scan 1-2 and a reversal of this effect from Scan 2-3. FA changes correlated with training hours and with motor performance, and probabilistic tractography confirmed that white matter changes projected to gray matter changes and to regions that activate during Paleolithic toolmaking. These results show that acquisition of Paleolithic toolmaking skills elicits structural remodeling of recently evolved brain regions supporting human tool use, providing a mechanistic link between stone toolmaking and human brain evolution. These regions participate not only in toolmaking, but also in other complex functions including action planning and language, in keeping with the hypothesized co-evolution of these functions.

The main lines of evidence taken as support for the

The present study attempted to reconstruct 3D brain shape of Neanderthals and early Homo sapiens based on computational neuroanatomy. We found that early Homo sapiens had relatively larger cerebellar hemispheres but a smaller occipital region in the cerebrum than Neanderthals long before the time that Neanderthals disappeared. Further, using behavioural and structural imaging data of living humans, the abilities such as cognitive flexibility, attention, the language processing, episodic and working memory capacity were positively correlated with size-adjusted cerebellar volume. As the cerebellar hemispheres are structured as a large array of uniform neural modules, a larger cerebellum may possess a larger capacity for cognitive information processing. Such a neuroanatomical difference in the cerebellum may have caused important differences in cognitive and social abilities between the two species and might have contributed to the replacement of Neanderthals by early Homo sapiens.

The prefrontal cortex subserves executive control, i.e., the organization of action or thought in relation to internal goals. This brain region hosts a system of executive processes extending from premotor to the most anterior prefrontal regions that governs the temporal organization of behavior. Little is known, however, about the prefrontal executive system involved in the hierarchical organization of behavior. Here, we show using magnetic resonance imaging in humans that the posterior portion of the prefrontal cortex, including Broca's area and its homolog in the right hemisphere, contains a system of executive processes that control start and end states and the nesting of functional segments that combine in hierarchically organized action plans. Our results indicate that Broca's area and its right homolog process hierarchically structured behaviors regardless of their temporal organization, suggesting a fundamental segregation between prefrontal executive systems involved in the hierarchical and temporal organization of goal-directed behaviors.

We introduce a set of biologically and computationally motivated design choices for modeling the learning of language, or of other types of sequential, hierarchically structured experience and behavior, and describe an implemented system that conforms to these choices and is capable of unsupervised learning from raw natural-language corpora. Given a stream of linguistic input, our model incrementally learns a grammar that captures its statistical patterns, which can then be used to parse or generate new data. The grammar constructed in this manner takes the form of a directed weighted graph, whose nodes are recursively (hierarchically) defined patterns over the elements of the input stream. We evaluated the model in seventeen experiments, grouped into five studies, which examined, respectively, (a) the generative ability of grammar learned from a corpus of natural language, (b) the characteristics of the learned representation, (c) sequence segmentation and chunking, (d) artificial grammar learning, and (e) certain types of structure dependence. The model's performance largely vindicates our design choices, suggesting that progress in modeling language acquisition can be made on a broad front-ranging from issues of generativity to the replication of human experimental findings-by bringing biological and computational considerations, as well as lessons from prior efforts, to bear on the modeling approach.

Developmental psychologists have long recognized the extraordinary influence of action on learning (Held & Hein, 1963; Piaget, 1952). Action experiences begin to shape our perception of the world during infancy (e.g., as infants gain an understanding of others' goal-directed actions; Woodward, 2009) and these effects persist into adulthood (e.g., as adults learn about complex concepts in the physical sciences; Kontra, Lyons, Fischer, & Beilock, 2012). Theories of embodied cognition provide a structure within which we can investigate the mechanisms underlying action's impact on thinking and reasoning. We argue that theories of embodiment can shed light on the role of action experience in early learning contexts, and further that these theories hold promise for using action to scaffold learning in more formal educational settings later in development.

Three laboratory experiments involving students' behavior and brain imaging and one randomized field experiment in a college physics class explored the importance of physical experience in science learning. We reasoned that students' understanding of science concepts such as torque and angular momentum is aided by activation of sensorimotor brain systems that add kinetic detail and meaning to students' thinking. We tested whether physical experience with angular momentum increases involvement of sensorimotor brain systems during students' subsequent reasoning and whether this involvement aids their understanding. The physical experience, a brief exposure to forces associated with angular momentum, significantly improved quiz scores. Moreover, improved performance was explained by activation of sensorimotor brain regions when students later reasoned about angular momentum. This finding specifies a mechanism underlying the value of physical experience in science education and leads the way for classroom practices in which experience with the physical world is an integral part of learning.

Using an epidemiological sample (N = 1,117) and a prospective longitudinal design, this study tested the direct and indirect effects of preverbal and verbal communication (15 months to 3 years) on executive function (EF) at age 4 years. Results indicated that whereas gestures (15 months), as well as language (2 and 3 years), were correlated with later EF (phis >/= .44), the effect was entirely mediated through later language. In contrast, language had significant direct and indirect effects on later EF. Exploratory analyses indicated that the pattern of results was comparable for low- and not-low-income families. The results were consistent with theoretical accounts of language as a precursor of EF ability, and highlighted gesture as an early indicator of EF.

I introduce seven criteria for determining the validity of competing theories for the original function of language. I go on to present a novel explanation that meets all the criteria: language originally evolved to teach kin. I suggest that the use of symbols subsequently generated evolutionary feedback at two levels, in the form of self-modified selection pressures that favored structures in the mind that functioned to manipulate and use symbols with efficiency, and cultural selection on languages for learnability.

In the current study we investigated whether 12-month-old infants gesture appropriately for knowledgeable versus ignorant partners, in order to provide them with needed information. In two experiments we found that in response to a searching adult, 12-month-olds pointed more often to an object whose location the adult did not know and thus needed information to find (she had not seen it fall down just previously) than to an object whose location she knew and thus did not need information to find (she had watched it fall down just previously). These results demonstrate that, in contrast to classic views of infant communication, infants' early pointing at 12 months is already premised on an understanding of others' knowledge and ignorance, along with a prosocial motive to help others by providing needed information.

Purpose: This longitudinal study compared the development of hand and index-finger pointing in children with typical language development (TD) and children with language delay (LD). First, we examined whether the number and the form of pointing gestures during the second year of life are potential indicators of later LD. Second, we analyzed the influence of caregivers' gestural and verbal input on children's communicative development. Method: Thirty children with TD and 10 children with LD were observed together with their primary caregivers in a seminatural setting in 5 sessions between the ages of 12 and 21 months. Language skills were assessed at 24 months. Results: Compared with children with TD, children with LD used fewer index-finger points at 12 and 14 months but more pointing gestures in total at 21 months. There were no significant differences in verbal or gestural input between caregivers of children with or without LD. Conclusions: Using more index-finger points at the beginning of the second year of life is associated with TD, whereas using more pointing gestures at the end of the second year of life is associated with delayed acquisition. Neither the verbal nor gestural input of caregivers accounted for differences in children's skills.

Co-speech gestures constitute a unique form of multimodal communication because here the hand movements are temporally synchronized and semantically integrated with speech. Recent neuroimaging studies indicate that the perception of co-speech gestures might engage a core set of frontal, temporal, and parietal areas. However, no study has compared the neural processes during perception of different types of co-speech gestures, such as beat, deictic, iconic, and metaphoric co-speech gestures. The purpose of this study was to review the existing literature on the neural correlates of co-speech gesture perception and to test whether different types of co-speech gestures elicit a common pattern of brain activity in the listener. To this purpose, we conducted a meta-analysis of neuroimaging studies, which used different types of co-speech gestures to investigate the perception of multimodal (co-speech gestures) in contrast to unimodal (speech or gestures) stimuli. The results show that co-speech gesture perception consistently engages temporal regions related to auditory and movement perception as well as frontal-parietal regions associated with action understanding. The results of this study suggest that brain regions involved in multisensory processing and action understanding constitute the general core of co-speech gesture perception. Crown Copyright (C) 2014 Published by Elsevier Ltd.

Humans differ from other animals in many aspects of anatomy, physiology, and behaviour; however, the genotypic basis of most human-specific traits remains unknown. Recent whole-genome comparisons have made it possible to identify genes with elevated rates of amino acid change or divergent expression in humans, and non-coding sequences with accelerated base pair changes. Regulatory alterations may be particularly likely to produce phenotypic effects while preserving viability, and are known to underlie interesting evolutionary differences in other species. Here we identify molecular events particularly likely to produce significant regulatory changes in humans: complete deletion of sequences otherwise highly conserved between chimpanzees and other mammals. We confirm 510 such deletions in humans, which fall almost exclusively in non-coding regions and are enriched near genes involved in steroid hormone signalling and neural function. One deletion removes a sensory vibrissae and penile spine enhancer from the human androgen receptor (AR) gene, a molecular change correlated with anatomical loss of androgen-dependent sensory vibrissae and penile spines in the human lineage. Another deletion removes a forebrain subventricular zone enhancer near the tumour suppressor gene growth arrest and DNA-damage-inducible, gamma (GADD45G), a loss correlated with expansion of specific brain regions in humans. Deletions of tissue-specific enhancers may thus accompany both loss and gain traits in the human lineage, and provide specific examples of the kinds of regulatory alterations and inactivation events long proposed to have an important role in human evolutionary divergence.

The gene ASPM (abnormal spindle-like microcephaly associated) is a specific regulator of brain size, and its evolution in the lineage leading to Homo sapiens was driven by strong positive selection. Here, we show that one genetic variant of ASPM in humans arose merely about 5800 years ago and has since swept to high frequency under strong positive selection. These findings, especially the remarkably young age of the positively selected variant, suggest that the human brain is still undergoing rapid adaptive evolution.

Incongruencies between auditory and visual signals negatively affect human performance and cause selective activation in neuroimaging studies; therefore, they are increasingly used to probe audiovisual integration mechanisms. An open question is whether the increased BOLD response reflects computational demands in integrating mismatching low-level signals or reflects simultaneous unimodal conceptual representations of the competing signals. To address this question, we explore the effect of semantic congruency within and across three signal categories (speech, body actions, and unfamiliar patterns) for signals with matched low-level statistics. In a localizer experiment, unimodal (auditory and visual) and bimodal stimuli were used to identify ROIs. All three semantic categories cause overlapping activation patterns. We find no evidence for areas that show greater BOLD response to bimodal stimuli than predicted by the sum of the two unimodal responses. Conjunction analysis of the unimodal responses in each category identifies a network including posterior temporal, inferior frontal, and premotor areas. Semantic congruency effects are measured in the main experiment. We find that incongruent combinations of two meaningful stimuli (speech and body actions) but not combinations of meaningful with meaningless stimuli lead to increased BOLD response in the posterior STS (pSTS) bilaterally, the left SMA, the inferior frontal gyrus, the inferior parietal lobule, and the anterior insula. These interactions are not seen in premotor areas. Our findings are consistent with the hypothesis that pSTS and frontal areas form a recognition network that combines sensory categorical representations (in pSTS) with action hypothesis generation in inferior frontal gyrus/premotor areas. We argue that the same neural networks process speech and body actions.

Children often find it difficult to map verbs to specific referents within complex scenes, often believing that additional features are part of the referents. This study investigated whether 3-year-olds could use iconic gestures to map novel verbs to specific referents. One hundred and twenty children were taught verbs that could be interpreted as change-of-state or manner verbs while presented with manner, end-state, or no iconic gestures. Children were then presented with a choice that forced them to generalize either on the basis of manner or end state. Results showed that children who saw manner gestures showed a stronger manner bias compared to the other groups. Thus, the specific feature of an event encoded in gestures guides children's interpretations of novel words.

Previous research has shown that children benefit from gesturing during math instruction. We asked whether gesturing promotes learning because it is itself a physical action, or because it uses physical action to represent abstract ideas. To address this question, we taught third-grade children a strategy for solving mathematical-equivalence problems that was instantiated in one of three ways: (a) in a physical action children performed on objects, (b) in a concrete gesture miming that action, or (c) in an abstract gesture. All three types of hand movements helped children learn how to solve the problems on which they were trained. However, only gesture led to success on problems that required generalizing the knowledge gained. The results provide the first evidence that gesture promotes transfer of knowledge better than direct action on objects and suggest that the beneficial effects gesture has on learning may reside in the features that differentiate it from action.

A great deal of attention has recently been paid to gesture and its effects on thinking and learning. It is well established that the hand movements that accompany speech are an integral part of communication, ubiquitous across cultures, and a unique feature of human behavior. In an attempt to understand this intriguing phenomenon, researchers have focused on pinpointing the mechanisms that underlie gesture production. One proposal--that gesture arises from simulated action (Hostetter & Alibali Psychonomic Bulletin & Review, 15, 495-514, 2008)--has opened up discussions about action, gesture, and the relation between the two. However, there is another side to understanding a phenomenon and that is to understand its function. A phenomenon's function is its purpose rather than its precipitating cause--the why rather than the how. This paper sets forth a theoretical framework for exploring why gesture serves the functions that it does, and reviews where the current literature fits, and fails to fit, this proposal. Our framework proposes that whether or not gesture is simulated action in terms of its mechanism--it is clearly not reducible to action in terms of its function. Most notably, because gestures are abstracted representations and are not actions tied to particular events and objects, they can play a powerful role in thinking and learning beyond the particular, specifically, in supporting generalization and transfer of knowledge.

Vocal learning, in which animals modify their vocalizations to imitate those of others, has evolved independently in scattered lineages of birds and mammals. Comparative evidence supports two hypotheses for the selective advantages leading to the origin of vocal learning. The sexual selection hypothesis proposes that vocal learning evolves to allow expansion of vocal repertoires in response to mating preferences for more complex vocalizations. The information-sharing hypothesis also proposes that vocal learning evolves to allow expansion of vocal repertoires, but in this case in response to kin selection favoring sharing of information among relatives.

Much debate has been stimulated by the recent hypothesis that human language consists of a faculty that is shared with non-human animals (faculty of language in a broad sense; FLB) and a faculty that is specific to human language (faculty of language in a narrow sense; FLN). This hypothesis has encouraged a tendency to emphasize one component of FLN: the cognitive operation of recursion. In consequence, non-syntactical, yet unique, aspects of human language have been neglected. One of these properties consists of vocal learning that enables an abundance of learned syllables. I suggest that FLN is not an independent faculty, but an ‘emergent’ property, arising from interactions between several other non-syntactical subfaculties of FLB, including vocal learning ability.

Inferring the intentions and beliefs of another is an ability that is fundamental for social and affiliative interactions. A substantial amount of empirical evidence suggests that making sense of another's intentional and belief states (i.e. theory of mind) relies on exteroceptive (e.g. visual and auditory) and proprioceptive (i.e. motor) signals. Yet, despite its pivotal role in the guidance of behaviour, the role of the observer's interoceptive (visceral) processing in understanding another's internal states remains unexplored. Predicting and keeping track of interoceptive bodily states - which inform intentions and beliefs that guide behaviour - is one of the fundamental purposes of the human brain. In this paper, we will focus on the role of interoceptive predictions, prescribed by the free energy principle, in making sense of internal states that cause another's behaviour. We will discuss how multimodal expectations induced at deep (high) hierarchical levels - that necessarily entail interoceptive predictions - contribute to inference about others that is at the heart of theory of mind.

Typically developing (TD) children refer to objects uniquely in gesture (e.g., point at cat) before they produce verbal labels for these objects (

Previous brain imaging studies investigating motor sequence complexity have mainly examined the effect of increasing the length of pre-learned sequences. The novel contribution of this research is that we varied the structure of complex visuo-motor sequences along two different dimensions using mxn paradigm. The complexity of sequences is increased from 12 movements (organized as a 2 x 6 task) to 24 movements (organized as 4 x 6 and 2 x 12 tasks). Behavioral results indicate that although the success rate attained was similar across the two complex tasks (2 x 12 and 4 x 6), a greater decrease in response times was observed for the 2 x 12 compared to the 4 x 6 condition at an intermediate learning stage. This decrease is possibly related to successful chunking across sets in the 2 x 12 task. In line with this, we observed a selective activation of the fronto-parietal network. Shifts of activation were observed from the ventral to dorsal prefrontal, lateral to medial premotor and inferior to superior parietal cortex from the early to intermediate learning stage concomitant with an increase in hyperset length. We suggest that these selective activations and shifts in activity during complex sequence learning are possibly related to chunking of motor sequences. (C) 2011 Elsevier Inc.

The extent to which nonhuman animals can learn actual human language is a controversial question, but many nonhuman species have acquired elements of a two-way communication system that is, and was, sophisticated enough to enable its use in evaluating cognitive capacities. This article is a personal view of the history of these animal language studies.

For humans, the ability to communicate and use language is instantiated not only in the vocal modality but also in the visual modality. The main examples of this are sign languages and (co-speech) gestures. Sign languages, the natural languages of Deaf communities, use systematic and conventionalized movements of the hands, face, and body for linguistic expression. Co-speech gestures, though non-linguistic, are produced in tight semantic and temporal integration with speech and constitute an integral part of language together with speech. The articles in this issue explore and document how gestures and sign languages are similar or different and how communicative expression in the visual modality can change from being gestural to grammatical in nature through processes of conventionalization. As such, this issue contributes to our understanding of how the visual modality shapes language and the emergence of linguistic structure in newly developing systems. Studying the relationship between signs and gestures provides a new window onto the human ability to recruit multiple levels of representation (e.g., categorical, gradient, iconic, abstract) in the service of using or creating conventionalized communicative systems.

A set of left frontal, temporal, and parietal brain regions respond robustly during language comprehension and production (e.g., Fedorenko E, Hsieh PJ, Nieto-Castanon A, Whitfield-Gabrieli S, Kanwisher N. J Neurophysiol 104: 1177-1194, 2010; Menenti L, Gierhan SM, Segaert K, Hagoort P. Psychol Sci 22: 1173-1182, 2011). These regions have been further shown to be selective for language relative to other cognitive processes, including arithmetic, aspects of executive function, and music perception (e.g., Fedorenko E, Behr MK, Kanwisher N. Proc Natl Acad Sci USA 108: 16428-16433, 2011; Monti MM, Osherson DN. Brain Res 1428: 33-42, 2012). However, one claim about overlap between language and nonlinguistic cognition remains prominent. In particular, some have argued that language processing shares computational demands with action observation and/or execution (e.g., Rizzolatti G, Arbib MA. Trends Neurosci 21: 188-194, 1998; Koechlin E, Jubault T. Neuron 50: 963-974, 2006; Tettamanti M, Weniger D. Cortex 42: 491-494, 2006). However, the evidence for these claims is indirect, based on observing activation for language and action tasks within the same broad anatomical areas (e.g., on the lateral surface of the left frontal lobe). To test whether language indeed shares machinery with action observation/execution, we examined the responses of language brain regions, defined functionally in each individual participant (Fedorenko E, Hsieh PJ, Nieto-Castanon A, Whitfield-Gabrieli S, Kanwisher N. J Neurophysiol 104: 1177-1194, 2010) to action observation ( experiments 1, 2, and 3a) and action imitation ( experiment 3b). With the exception of the language region in the angular gyrus, all language regions, including those in the inferior frontal gyrus (within

Detection of communicative signals is thought to facilitate knowledge acquisition early in life, but less is known about the role these signals play in adult learning or about the brain systems supporting sensitivity to communicative intent. The current study examined how ostensive gaze cues and communicative actions affect adult recognition memory and modulate neural activity as measured by fMRI. For both the behavioral and fMRI experiments, participants viewed a series of videos of an actress acting on one of two objects in front of her. Communicative context in the videos was manipulated in a 2 x 2 design in which the actress either had direct gaze (Gaze) or wore a visor (NoGaze) and either pointed at (Point) or reached for (Reach) one of the objects (target) in front of her. Participants then completed a recognition memory task with old (target and nontarget) objects and novel objects. Recognition memory for target objects in the Gaze conditions was greater than NoGaze, but no effects of gesture type were seen. Similarly, the fMRI video-viewing task revealed a significant effect of Gaze within right posterior STS (pSTS), but no significant effects of Gesture. Furthermore, pSTS sensitivity to Gaze conditions was related to greater memory for objects viewed in Gaze, as compared with NoGaze, conditions. Taken together, these results demonstrate that the ostensive, communicative signal of direct gaze preceding an object-directed action enhances recognition memory for attended items and modulates the pSTS response to object-directed actions. Thus, establishment of a communicative context through ostensive signals remains an important component of learning and memory into adulthood, and the pSTS may play a role in facilitating this type of social learning.

Children from low-socioeconomic status (SES) families, on average, arrive at school with smaller vocabularies than children from high-SES families. In an effort to identify precursors to, and possible remedies for, this inequality, we videotaped 50 children from families with a range of different SES interacting with parents at 14 months and assessed their vocabulary skills at 54 months. We found that children from high-SES families frequently used gesture to communicate at 14 months, a relation that was explained by parent gesture use (with speech controlled). In turn, the fact that children from high-SES families have large vocabularies at 54 months was explained by children's gesture use at 14 months. Thus, differences in early gesture help to explain the disparities in vocabulary that children bring with them to school.

The gestures children produce predict the early stages of spoken language development. Here we ask whether gesture is a global predictor of language learning, or whether particular gestures predict particular language outcomes. We observed 52 children interacting with their caregivers at home, and found that gesture use at 18 months selectively predicted lexical versus syntactic skills at 42 months, even with early child speech controlled. Specifically, number of different meanings conveyed in gesture at 18 months predicted vocabulary at 42 months, but number of gesture+speech combinations did not. In contrast, number of gesture+speech combinations, particularly those conveying sentence-like ideas, produced at 18 months predicted sentence complexity at 42 months, but meanings conveyed in gesture did not. We can thus predict particular milestones in vocabulary and sentence complexity at age by watching how children move their hands two years earlier.

There is growing evidence that activation of the motor system during observation of actions, a phenomenon first observed in non-human primates, underlies action understanding and even communication. This review (a) examines the evidence on motor system activity as an underlying neural correlate of action understanding; (b) reviews the theoretical and empirical work linking action understanding and the development of communication, with a specific focus on the role that gestures play as an intermediary; and (c) discusses the research on and existing opportunities for understanding the link between the motor system and communication in both humans and non-human primates, through the lens of action perception. Bringing together findings and perspectives from developmental social cognition in both humans and non-human primates and applying recent neuroscientific perspectives will help to elucidate the processes underlying the ability to understand and communicate with others.

In infancy, use of gesture and the ability to engage in joint attention with others both predict later language development. Conceptually, gesture and joint attention abilities may reflect a similar underlying social communicative skill. However, these abilities are often studied separately. Despite the fact that gesture is often used in episodes of joint attention, little is known about the degree to which measures of gesture use and joint attention ability are associated with one another or how they similarly, or differentially, predict children's language abilities. Participants in the current study were 53 infants. At 12-months, multiple measures of infants' gesture use were gleaned from a free-play interaction with a parent. Infants' responding to and initiating joint attention were measured via the Early Social-Communicative Scales (ESCS, Mundy et al., 2003). Infants' expressive and receptive language was measured at 24-months with the Mullen Scales of Early Learning (Mullen, 1995). A factor analysis including gesture and joint attention measures indicated that at 12-months joint attention, particularly responding to joint attention, reflects a similar underlying construct with infant gesture use, yet they uniquely predict later language ability.

It has long been postulated that language is not purely learned, but arises from an interaction between environmental exposure and innate abilities. The innate component becomes more evident in rare situations in which the environment is markedly impoverished. The present study investigated the language production of a generation of deaf Nicaraguans who had not been exposed to a developed language. We examined the changing use of early linguistic structures (specifically, spatial modulations) in a sign language that has emerged since the Nicaraguan group first came together: In tinder two decades, sequential cohorts of learners systematized the grammar of this new sign language. We examined whether the systematicity being added to the language stems from children or adults: our results indicate that such changes originate in children aged 10 and younger Thus, sequential cohorts of interacting young children collectively: possess the capacity not only to learn, but also to create, language.

Humans are an unusually prosocial species-we vote, give blood, recycle, give tithes and punish violators of social norms. Experimental evidence indicates that people willingly incur costs to help strangers in anonymous one-shot interactions, and that altruistic behaviour is motivated, at least in part, by empathy and concern for the welfare of others (hereafter referred to as other-regarding preferences). In contrast, cooperative behaviour in non-human primates is mainly limited to kin and reciprocating partners, and is virtually never extended to unfamiliar individuals. Here we present experimental tests of the existence of other-regarding preferences in non-human primates, and show that chimpanzees (Pan troglodytes) do not take advantage of opportunities to deliver benefits to familiar individuals at no material cost to themselves, suggesting that chimpanzee behaviour is not motivated by other-regarding preferences. Chimpanzees are among the primates most likely to demonstrate prosocial behaviours. They participate in a variety of collective activities, including territorial patrols, coalitionary aggression, cooperative hunting, food sharing and joint mate guarding. Consolation of victims of aggression and anecdotal accounts of solicitous treatment of injured individuals suggest that chimpanzees may feel empathy. Chimpanzees sometimes reject exchanges in which they receive less valuable rewards than others, which may be one element of a 'sense of fairness', but there is no evidence that they are averse to interactions in which they benefit more than others.

Paleolithic stone tools provide concrete evidence of major developments in human behavioural and cognitive evolution. Of particular interest are evolving cognitive mechanisms implied by the cultural transmission of increasingly complex prehistoric technologies, hypothetically including motor resonance, causal reasoning and mentalizing. To test the relevance of these mechanisms to specific Paleolithic technologies, we conducted a functional magnetic resonance imaging study of Naive, Trained and Expert subjects observing two toolmaking methods of differing complexity and antiquity: the simple 'Oldowan' method documented by the earliest tools 2.5 million years ago; and the more complex 'Acheulean' method used to produce refined tools 0.5 million years ago. Subjects observed 20-s video clips of an expert demonstrator, followed by behavioural tasks designed to maintain attention. Results show that observational understanding of Acheulean toolmaking involves increased demands for the recognition of abstract technological intentions. Across subject groups, Acheulean compared with Oldowan toolmaking was associated with activation of left anterior intraparietal and inferior frontal sulci, indicating the relevance of resonance mechanisms. Between groups, Naive subjects relied on bottom-up kinematic simulation in the premotor cortex to reconstruct unfamiliar intentions, and Experts employed a combination of familiarity-based sensorimotor matching in the posterior parietal cortex and top-down mentalizing involving the medial prefrontal cortex. While no specific differences between toolmaking technologies were found for Trained subjects, both produced frontal activation relative to Control, suggesting focused engagement with toolmaking stimuli. These findings support motor resonance hypotheses for the evolutionary origins of human social cognition and cumulative culture, directly linking these hypotheses with archaeologically observable behaviours in prehistory.

The recent blossoming of evolutionary linguistics has resulted in a variety of theories that attempt to provide a selective scenario for the evolution of early language. However, their overabundance makes many researchers sceptical of such theorising. Here, we suggest that a more rigorous approach is needed towards their construction although, despite justified scepticism, there is no agreement as to the criteria that should be used to determine the validity of the various competing theories. We attempt to fill this gap by providing criteria upon which the various historical narratives can be judged. Although individually none of these criteria are highly constraining, taken together they could provide a useful evolutionary framework for thinking about the evolution of human language.

Understanding the factors which affect the age of acquisition (AoA) of words and concepts is fundamental to understanding cognitive development more broadly. Traditionally, studies of AoA have taken two approaches, either exploring the effect of linguistic variables such as input frequency (e.g., Naigles and Hoff-Ginsberg, 1998) or the semantics of the underlying concept, such as concreteness or imageability (e.g., Bird et al., 2001). Embodied theories of cognition, meanwhile, assume that concepts, even relatively abstract ones, can be grounded in the embodied experience. While the focus of such discussions has been mainly on grounding in external modalities, more recently some have argued for the importance of interoceptive features, or grounding in complex modalities such as social interaction. In this paper, we argue for the integration and extension of these two strands of research. We demonstrate that the psycholinguistic factors traditionally considered to determine AoA are far from sufficient to account for the variability observed in AoA data. Given this gap, we propose groundability as a new conceptual tool that can measure the degree to which concepts are grounded both in external and, critically, internal modalities. We then present a mechanistic theory of conceptual representation that can account for groundability in addition to the existing variables argued to influence concept acquisition in both the developmental and embodied cognition literatures, and discuss its implications for future work in concept and cognitive development.

AbstractThe cortex constituting Broca's area does not exist in isolation. Rather, like other cortical regions, Broca's area is connected to other brain structures, which likely play closely related functional roles. This paper focuses on the basal ganglia, a set of subcortical structures that project through topographically organized “channels” via the thalamus to different frontal regions. It is hypothesized that the basal ganglia project to Broca's area. This circuitry is further posited to encompass at least two channels. One channel can be characterized as subserving procedural memory, while the other underlies the retrieval of knowledge from declarative memory. These hypotheses are supported by both anatomical and functional evidence. Implications and issues for further investigation are discussed.]]>

BACKGROUND: The popular theory that complex tool-making and language co-evolved in the human lineage rests on the hypothesis that both skills share underlying brain processes and systems. However, language and stone tool-making have so far only been studied separately using a range of neuroimaging techniques and diverse paradigms. METHODOLOGY/PRINCIPAL FINDINGS: We present the first-ever study of brain activation that directly compares active Acheulean tool-making and language. Using functional transcranial Doppler ultrasonography (fTCD), we measured brain blood flow lateralization patterns (hemodynamics) in subjects who performed two tasks designed to isolate the planning component of Acheulean stone tool-making and cued word generation as a language task. We show highly correlated hemodynamics in the initial 10 seconds of task execution. CONCLUSIONS/SIGNIFICANCE: Stone tool-making and cued word generation cause common cerebral blood flow lateralization signatures in our participants. This is consistent with a shared neural substrate for prehistoric stone tool-making and language, and is compatible with language evolution theories that posit a co-evolution of language and manual praxis. In turn, our results support the hypothesis that aspects of language might have emerged as early as 1.75 million years ago, with the start of Acheulean technology.

Teaching a new concept through gestures-hand movements that accompany speech-facilitates learning above-and-beyond instruction through speech alone (e.g., Singer & Goldin-Meadow, ). However, the mechanisms underlying this phenomenon are still under investigation. Here, we use eye tracking to explore one often proposed mechanism-gesture's ability to direct visual attention. Behaviorally, we replicate previous findings: Children perform significantly better on a posttest after learning through Speech+Gesture instruction than through Speech Alone instruction. Using eye tracking measures, we show that children who watch a math lesson with gesture do allocate their visual attention differently from children who watch a math lesson without gesture-they look more to the problem being explained, less to the instructor, and are more likely to synchronize their visual attention with information presented in the instructor's speech (i.e., follow along with speech) than children who watch the no-gesture lesson. The striking finding is that, even though these looking patterns positively predict learning outcomes, the patterns do not mediate the effects of training condition (Speech Alone vs. Speech+Gesture) on posttest success. We find instead a complex relation between gesture and visual attention in which gesture moderates the impact of visual looking patterns on learning-following along with speech predicts learning for children in the Speech+Gesture condition, but not for children in the Speech Alone condition. Gesture's beneficial effects on learning thus come not merely from its ability to guide visual attention, but also from its ability to synchronize with speech and affect what learners glean from that speech.

Verb learning is difficult for children (Gentner, ), partially because children have a bias to associate a novel verb not only with the action it represents, but also with the object on which it is learned (Kersten & Smith, ). Here we investigate how well 4- and 5-year-old children (N = 48) generalize novel verbs for actions on objects after doing or seeing the action (e.g., twisting a knob on an object) or after doing or seeing a gesture for the action (e.g., twisting in the air near an object). We find not only that children generalize more effectively through gesture experience, but also that this ability to generalize persists after a 24-hour delay.

Human beings routinely help others to achieve their goals, even when the helper receives no immediate benefit and the person helped is a stranger. Such altruistic behaviors (toward non-kin) are extremely rare evolutionarily, with some theorists even proposing that they are uniquely human. Here we show that human children as young as 18 months of age (prelinguistic or just-linguistic) quite readily help others to achieve their goals in a variety of different situations. This requires both an understanding of others' goals and an altruistic motivation to help. In addition, we demonstrate similar though less robust skills and motivations in three young chimpanzees.

It is debated how language and praxis are co-represented in the left hemisphere (LH). As voxel-based lesion-symptom mapping in LH stroke patients with aphasia and/or apraxia may contribute to this debate, we here investigated the relationship between language and praxis deficits at the behavioral and lesion levels in 50 sub-acute stroke patients. We hypothesized that language and (meaningful) action are linked via semantic processing in Broca's region. Behaviorally, half of the patients suffered from co-morbid aphasia and apraxia. While 24% (n = 12) of all patients exhibited aphasia without apraxia, apraxia without aphasia was rare (n = 2, 4%). Left inferior frontal, insular, inferior parietal, and superior temporal lesions were specifically associated with deficits in naming, reading, writing, or auditory comprehension. In contrast, lesions affecting the left inferior frontal gyrus, premotor cortex, and the central region as well as the inferior parietal lobe were associated with apraxic deficits (i.e., pantomime, imitation of meaningful and meaningless gestures). Thus, contrary to the predictions of the embodied cognition theory, lesions to sensorimotor and premotor areas were associated with the severity of praxis but not language deficits. Lesions of Brodmann area (BA) 44 led to combined apraxic and aphasic deficits. Data suggest that BA 44 acts as an interface between language and (meaningful) action thereby supporting parcellation schemes (based on connectivity and receptor mapping) which revealed a BA 44 sub-area involved in semantic processing.

Gestures play an important role in face-to-face communication and have been increasingly studied via functional magnetic resonance imaging. Although a large amount of data has been provided to describe the neural substrates of gesture comprehension, these findings have never been quantitatively summarized and the conclusion is still unclear. This activation likelihood estimation meta-analysis investigated the brain networks underpinning gesture comprehension while considering the impact of gesture type (co-speech gestures vs. speech-independent gestures) and task demand (implicit vs. explicit) on the brain activation of gesture comprehension. The meta-analysis of 31 papers showed that as hand actions, gestures involve a perceptual-motor network important for action recognition. As meaningful symbols, gestures involve a semantic network for conceptual processing. Finally, during face-to-face interactions, gestures involve a network for social emotive processes. Our finding also indicated that gesture type and task demand influence the involvement of the brain networks during gesture comprehension. The results highlight the complexity of gesture comprehension, and suggest that future research is necessary to clarify the dynamic interactions among these networks.

This study examined whether poor pointing gestures and imitative actions at 18months of age uniquely predicted late language production at 36months, beyond the role of poor language at 18months of age. Data from the Norwegian Mother and Child Cohort Study were utilized. Maternal reports of the children's nonverbal skills and language were gathered for 42,517 children aged 18months and for 28,107 of the same children at 36months. Panel analysis of latent variables revealed that imitative actions, language comprehension, and language production uniquely contributed to predicting late development of language production, while pointing gestures did not. It is suggested that the results can be explained by underlying symbolic representational skills at 18months.