Up right, not right up: Primacy of verticality in both language and movement.

When describing motion along both the horizontal and vertical axes, languages from different families express the elements encoding verticality before those coding for horizontality (e.g., going up right instead of right up). In light of the motor grounding of language, the present study investigated whether the prevalence of verticality in Path expression also governs the trajectory of arm biological movements. Using a 3D virtual-reality setting, we tracked the kinematics of hand pointing movements in five spatial directions, two of which implied the vertical and horizontal vectors equally (i.e., up right +45° and bottom right -45°). Movement onset could be prompted by visual or auditory verbal cues, the latter being canonical in French ("en haut à droite"/up right) or not ("à droite en haut"/right up). In two experiments, analyses of the index finger kinematics revealed a significant effect of gravity, with earlier acceleration, velocity, and deceleration peaks for upward (+45°) than downward (-45°) movements, irrespective of the instructions. Remarkably, confirming the linguistic observations, we found that vertical kinematic parameters occurred earlier than horizontal ones for upward movements, both for visual and congruent verbal cues. Non-canonical verbal instructions significantly affected this temporal dynamic: for upward movements, the horizontal and vertical components temporally aligned, while they reversed for downward movements where the kinematics of the vertical axis was delayed with respect to that of the horizontal one. This temporal dynamic is so deeply anchored that non-canonical verbal instructions allowed for horizontality to precede verticality only for movements that do not fight against gravity. Altogether, our findings provide new insights into the embodiment of language by revealing that linguistic path may reflect the organization of biological movements, giving priority to the vertical axis.

Tool use and language share syntactic processes and neural patterns in the basal ganglia.

Does tool use share syntactic processes with language? Acting with a tool is thought to add a hierarchical level into the motor plan. In the linguistic domain, syntax is the cognitive function handling interdependent elements. Using functional magnetic resonance imaging, we detected common neurofunctional substrates in the basal ganglia subserving both tool use and syntax in language. The two abilities elicited similar patterns of neural activity, indicating the existence of shared functional resources. Manual actions and verbal working memory did not contribute to this common network. Consistent with the existence of shared neural resources, we observed bidirectional behavioral enhancement of tool use and syntactic skills in language so that training one function improves performance in the other. This reveals supramodal syntactic processes for tool use and language.

The toolish hand illusion: embodiment of a tool based on similarity with the hand.

A tool can function as a body part yet not feel like one: Putting down a fork after dinner does not feel like losing a hand. However, studies show fake body-parts are embodied and experienced as parts of oneself. Typically, embodiment illusions have only been reported when the fake body-part visually resembles the real one. Here we reveal that participants can experience an illusion that a mechanical grabber, which looks scarcely like a hand, is part of their body. We found changes in three signatures of embodiment: the real hand's perceived location, the feeling that the grabber belonged to the body, and autonomic responses to visible threats to the grabber. These findings show that artificial objects can become embodied even though they bear little visual resemblance to the hand.

Elementary visuospatial perception deficit in children with neurodevelopmental disorders.

AIM: To assess the prevalence of elementary visuospatial perception (EVSP) deficit in children with neurodevelopmental disorders. METHOD: Using a screening test designed and validated to measure dorsal EVSP ability, 168 children (122 males, 46 females; mean age 10y [SD 1y 10mo], range 4y 8mo-16y 4mo) diagnosed with developmental coordination disorder (DCD), specific learning disorder (SLD), attention-deficit/hyperactivity disorder (ADHD), and/or oral language disorder were compared with a group of 184 typically developing children. We also tested 14 children with binocular vision dysfunction and no neurodevelopmental disorder. RESULTS: Children with SLD scored below the interquartile range of typically developing children as frequently (59%) as children with DCD, but only 5% were severely impaired (i.e. scored as outliers). Children with DCD were the most severely impaired (22% of outliers), even more so when they exhibited a co-occuring disorder. Children with language disorder and those with binocular vision dysfunction scored similarly to the group of typically developing children. INTERPRETATION: These results confirm the importance of assessing EVSP in the clinical evaluation of children with neurodevelopmental disorders, in particular those presenting with DCD or SLD. What this paper adds More than half of children with developmental coordination disorder (DCD) scored below the normal interquartile range on the elementary visuospatial perception (EVSP) test. More than half of children with specific learning disorder (SLD) scored below the normal interquartile range on the EVSP test. Twenty-two percent of children with DCD performed as outliers on the EVSP test. Children with language disorder and those with binocular vision dysfunction scored similarly to typically developing children.

Online proprioception feeds plasticity of arm representation following tool-use in healthy aging.

Following tool-use, the kinematics of free-hand movements are altered. This modified kinematic pattern has been taken as a behavioral hallmark of the modification induced by tool-use on the effector representation. Proprioceptive inputs appear central in updating the estimated effector state. Here we questioned whether online proprioceptive modality that is accessed in real time, or offline, memory-based, proprioception is responsible for this update. Since normal aging affects offline proprioception only, we examined a group of 60 year-old adults for proprioceptive acuity and movement's kinematics when grasping an object before and after tool-use. As a control, participants performed the same movements with a weight-equivalent to the tool-weight-attached to their wrist. Despite hampered offline proprioceptive acuity, 60 year-old participants exhibited the typical kinematic signature of tool incorporation: Namely, the latency of transport components peaks was longer and their amplitude reduced after tool-use. Instead, we observed no kinematic modifications in the control condition. In addition, online proprioception acuity correlated with tool incorporation, as indexed by the amount of kinematics changes observed after tool-use. Altogether, these findings point to the prominent role played by online proprioception in updating the body estimate for the motor control of tools.

Validation of a simple screening test for elementary visuo-spatial perception deficit.

OBJECTIVE: The reliability and validity of a screening test for a deficit in elementary visuo-spatial perception (EVSP) were evaluated. METHOD: This prospective study collected performance from 210 typically developing individuals and evaluated the internal consistency of the EVSP screening test. Test-retest reliability was examined with 25 individuals. Validity also involved retrospective clinical data collected from 223 non-typically developing children coming to the hospital for outpatient consultation. Since EVSP matures through childhood, we standardized the EVSP screening test scores by age category and performed Pearson correlations with standardized clinical tests scores. RESULTS: Test-retest reliability (intraclass correlation coefficient=0.76) and internal consistency (Cronbach's alpha=0.76) were satisfactory. Construct validity included correlation with the subtests of the Wechsler Intelligence Scale IV (WISC-IV) involving visuo-spatial analysis (Matrix Reasoning and Block Design, P<0.01; Symbol Search and Coding, P<0.05) and was reinforced by the expected non-correlation between the Verbal Comprehension Index and EVSP scoring class. The EVSP scoring class was correlated with Manual dexterity of the M-ABC (P<0.05) and the Working Memory Index (P<0.05) of the WISC-IV including the subtest Arithmetic (P<0.01). CONCLUSION: This screening test is reliable and valid to evaluate EVSP before more complex cognitive or motor assessment.

Language as a Tool: Motor Proficiency Using a Tool Predicts Individual Linguistic Abilities.

Different disciplines converge to trace language evolution from motor skills. The human ability to use tools has been advocated as a fundamental step toward the emergence of linguistic processes in the brain. Neuropsychological and neuroimaging research has established that linguistic functions and tool-use are mediated by partially overlapping brain networks. Yet, scholars still theoretically debate whether the relationship between tool-use and language is contingent or functionally relevant, since empirical evidence is critically missing. Here, we measured both linguistic production and tool-use abilities in the same participants, as well as manual and linguistic motor skills. A path analysis ruling out unspecific contributions from manual or linguistic motor skills, showed that motor proficiency using a tool lawfully predicts differences in individual linguistic production. In addition, more complex tool-use reveals stronger association between linguistic production and tool mastery. These findings establish the existence of shared cognitive processes between tool-use and language.

Tracking the acquisition of anticipatory postural adjustments during a bimanual load-lifting task: A MEG study.

During bimanual coordination, that is, manipulating with the dominant hand an object held by the postural hand, anticipatory postural adjustments are required to cancel the perturbations and ensure postural stabilization. Using magnetoencephalography (MEG), we investigated changes mediating the acquisition of anticipatory postural adjustments during a bimanual load-lifting task. Participants lifted a load with their right hand, hence triggering the fall of a second load fixed to their left (postural) forearm. During Acquisition, the onset of load-lifting and the fall of the second load were experimentally delayed after few trials. During Control, load-lifting triggered the fall of the second load without delay. Upward elbow rotation decreased with trial repetition during Acquisition, hence attesting the ongoing acquisition of anticipatory postural adjustments. Bilateral event-related desynchronisation (ERD) of the alpha rhythm (8-12 Hz) was recorded. Generators of the mu rhythm were found within central and associative motor regions. Their spatial distribution within the hemisphere contralateral to the load-lifting arm was less refined and circumscribed during Acquisition compared to Control. Regression analyses emphasized the specific involvement of the precuneus in the right hemisphere contralateral to the postural forearm, and a medial prefrontal region in the left hemisphere. Analyses of the time course power showed that an increase in preunloading activation within the precuneus and a decrease in postunloading inhibition within the medial prefrontal region were associated with the acquisition of anticipatory postural adjustments. The study provides original insights into cortical activations mediating the progressive tuning of anticipatory postural adjustments during the acquisition stage of motor learning.

Proprioception Is Necessary for Body Schema Plasticity: Evidence from a Deafferented Patient.

The ability of using a large variety of tools is important in our daily life. Behind human tool-use abilities lays the brain capacity to incorporate tools into the body representation for action (Body Schema, BS), thought to rely mainly on proprioceptive information. Here, we tested whether tool incorporation is possible in absence of proprioception by studying a patient with right upper-limb deafferentation. We adopted a paradigm sensitive to changes of the BS and analyzed the kinematics of free-hand movements before and after tool-use, in three sessions over a period of 2 years. In the first session, before tool-use, the kinematics of the deafferented hand was disrupted. Similarly, the first movements with the tool (a mechanical grabber elongating the arm by ~40 cm) showed an abnormal profile that tended to normalize at the end of the session. Subsequent free-hand movements were also normalized. At session 2, 6 months later, the patient exhibited normal free-hand kinematic profiles, additionally showing changes in grasping kinematics after tool-use, but no sign of tool incorporation. A follow-up 2 years later, further confirmed the normalized kinematic profile but the absence of tool incorporation. This first description of tool-use in absence of proprioception shows the fundamental role of proprioception in the update of the BS. These results provide an important further step in understanding human motor control and have implications for future development of rehabilitation programs for patients with sensory deficits.

Changing ideas about others' intentions: updating prior expectations tunes activity in the human motor system.

Predicting intentions from observing another agent's behaviours is often thought to depend on motor resonance - i.e., the motor system's response to a perceived movement by the activation of its stored motor counterpart, but observers might also rely on prior expectations, especially when actions take place in perceptually uncertain situations. Here we assessed motor resonance during an action prediction task using transcranial magnetic stimulation to probe corticospinal excitability (CSE) and report that experimentally-induced updates in observers' prior expectations modulate CSE when predictions are made under situations of perceptual uncertainty. We show that prior expectations are updated on the basis of both biomechanical and probabilistic prior information and that the magnitude of the CSE modulation observed across participants is explained by the magnitude of change in their prior expectations. These findings provide the first evidence that when observers predict others' intentions, motor resonance mechanisms adapt to changes in their prior expectations. We propose that this adaptive adjustment might reflect a regulatory control mechanism that shares some similarities with that observed during action selection. Such a mechanism could help arbitrate the competition between biomechanical and probabilistic prior information when appropriate for prediction.

Tool-use: An open window into body representation and its plasticity.

Over the last decades, scientists have questioned the origin of the exquisite human mastery of tools. Seminal studies in monkeys, healthy participants and brain-damaged patients have primarily focused on the plastic changes that tool-use induces on spatial representations. More recently, we focused on the modifications tool-use must exert on the sensorimotor system and highlighted plastic changes at the level of the body representation used by the brain to control our movements, i.e., the Body Schema. Evidence is emerging for tool-use to affect also more visually and conceptually based representations of the body, such as the Body Image. Here we offer a critical review of the way different tool-use paradigms have been, and should be, used to try disentangling the critical features that are responsible for tool incorporation into different body representations. We will conclude that tool-use may offer a very valuable means to investigate high-order body representations and their plasticity.

Tool use imagery triggers tool incorporation in the body schema.

Tool-use has been shown to modify the way the brain represents the metrical characteristics of the effector controlling the tool. For example, the use of tools that elongate the physical length of the arm induces kinematic changes affecting selectively the transport component of subsequent free-hand movements. Although mental simulation of an action is known to involve -to a large extent- the same processes as those at play in overt motor execution, whether tool-use imagery can yield similar effects on the body representation remains unknown. Mentally simulated actions indeed elicit autonomic physiological responses and follow motor execution rules that are comparable to those associated with the correspondent overt performance. Therefore, here we investigated the effects of the mental simulation of actions performed with a tool on the body representation by studying subsequent free-hand movements. Subjects executed reach to grasp movements with their hand before and after an imagery task performed with either a tool elongating their arm length or, as a control, with their hand alone. Two main results were found: First, in agreement with previous studies, durations of imagined movements performed with the tool and the hand were similarly affected by task difficulty. Second, kinematics of free-hand movements was affected after tool-use imagery, but not hand-use imagery, in a way similar to that previously documented after actual tool-use. These findings constitute the first evidence that tool-use imagery is sufficient to affect the representation of the user's arm.

Syntax at hand: common syntactic structures for actions and language.

Evidence that the motor and the linguistic systems share common syntactic representations would open new perspectives on language evolution. Here, crossing disciplinary boundaries, we explore potential parallels between the structure of simple actions and that of sentences. First, examining Typically Developing (TD) children displacing a bottle with or without knowledge of its weight prior to movement onset, we provide kinematic evidence that the sub-phases of this displacing action (reaching + moving the bottle) manifest a structure akin to linguistic embedded dependencies. Then, using the same motor task, we reveal that children suffering from specific language impairment (SLI), whose core deficit affects syntactic embedding and dependencies, manifest specific structural motor anomalies parallel to their linguistic deficits. In contrast to TD children, SLI children performed the displacing-action as if its sub-phases were juxtaposed rather than embedded. The specificity of SLI's structural motor deficit was confirmed by testing an additional control group: Fragile-X Syndrome patients, whose language capacity, though delayed, comparatively spares embedded dependencies, displayed slower but structurally normal motor performances. By identifying the presence of structural representations and dependency computations in the motor system and by showing their selective deficit in SLI patients, these findings point to a potential motor origin for language syntax.

Grab an object with a tool and change your body: tool-use-dependent changes of body representation for action.

Along the evolutionary history, humans have reached a high level of sophistication in the way they interact with the environment. One important step in this process has been the introduction of tools, enabling humans to go beyond the boundaries of their physical possibilities. Here, we focus on some "low level" aspects of sensorimotor processing that highlight how tool-use plays a causal role in shaping body representations, an essential plastic feature for efficient motor control during development and skilful tool-use in the adult life. We assess the evidence supporting the hypothesis that tools are incorporated in body representation for action, which is the body schema, by critically reviewing some previous findings and providing new data from on-going work in our laboratory. In particular, we discuss several experiments that reveal the effects of tool-use both on the kinematics of hand movements and the localization of somatosensory stimuli on the body surface, as well as the conditions that are necessary for these effects to be manifested. We suggest that overall these findings speak in favour of genuine tool-use-dependent plasticity of the body representation for the control of action.

A neuropsychological perspective on the link between language and praxis in modern humans.

Hypotheses about the emergence of human cognitive abilities postulate strong evolutionary links between language and praxis, including the possibility that language was originally gestural. The present review considers functional and neuroanatomical links between language and praxis in brain-damaged patients with aphasia and/or apraxia. The neural systems supporting these functions are predominantly located in the left hemisphere. There are many parallels between action and language for recognition, imitation and gestural communication suggesting that they rely partially on large, common networks, differentially recruited depending on the nature of the task. However, this relationship is not unequivocal and the production and understanding of gestural communication are dependent on the context in apraxic patients and remains to be clarified in aphasic patients. The phonological, semantic and syntactic levels of language seem to share some common cognitive resources with the praxic system. In conclusion, neuropsychological observations do not allow support or rejection of the hypothesis that gestural communication may have constituted an evolutionary link between tool use and language. Rather they suggest that the complexity of human behaviour is based on large interconnected networks and on the evolution of specific properties within strategic areas of the left cerebral hemisphere.

The agent is right: when motor embodied cognition is space-dependent.

The role of embodied mechanisms in processing sentences endowed with a first person perspective is now widely accepted. However, whether embodied sentence processing within a third person perspective would also have motor behavioral significance remains unknown. Here, we developed a novel version of the Action-sentence Compatibility Effect (ACE) in which participants were asked to perform a movement compatible or not with the direction embedded in a sentence having a first person (Experiment 1: You gave a pizza to Louis) or third person perspective (Experiment 2: Lea gave a pizza to Louis). Results indicate that shifting perspective from first to third person was sufficient to prevent motor embodied mechanisms, abolishing the ACE. Critically, ACE was restored in Experiment 3 by adding a virtual "body" that allowed participants to know "where" to put themselves in space when taking the third person perspective, thus demonstrating that motor embodied processes are space-dependent. A fourth, control experiment, by dissociating motor response from the transfer verb's direction, supported the conclusion that perspective-taking may induce significant ACE only when coupled with the adequate sentence-response mapping.

Encoding of human action in Broca's area.

Broca's area has been considered, for over a century, as the brain centre responsible for speech production. Modern neuroimaging and neuropsychological evidence have suggested a wider functional role is played by this area. In addition to the evidence that it is involved in syntactical analysis, mathematical calculation and music processing, it has recently been shown that Broca's area may play some role in language comprehension and, more generally, in understanding actions of other individuals. As shown by functional magnetic resonance imaging, Broca's area is one of the cortical areas activated by hand/mouth action observation and it has been proposed that it may form a crucial node of a human mirror-neuron system. If, on the one hand, neuroimaging studies use a correlational approach which cannot offer a final proof for such claims, available neuropsychological data fail to offer a conclusive demonstration for two main reasons: (i) they use tasks taxing both language and action systems; and (ii) they rarely consider the possibility that Broca's aphasics may also be affected by some form of apraxia. We administered a novel action comprehension test--with almost no linguistic requirements--on selected frontal aphasic patients lacking apraxic symptoms. Patients, as well as matched controls, were shown short movies of human actions or of physical events. Their task consisted of ordering, in a temporal sequence, four pictures taken from each movie and randomly presented on the computer screen. Patient's performance showed a specific dissociation in their ability to re-order pictures of human actions (impaired) with respect to physical events (spared). Our study provides a demonstration that frontal aphasics, not affected by apraxia, are specifically impaired in their capability to correctly encode observed human actions.

Subliminal display of action words interferes with motor planning: a combined EEG and kinematic study.

Recent evidence has shown that processing action-related language and motor action share common neural representations to a point that the two processes can interfere when performed concurrently. To support the assumption that language-induced motor activity contributes to action word understanding, the present study aimed at ruling out that this activity results from mental imagery of the movements depicted by the words. For this purpose, we examined cross-talk between action word processing and an arm reaching movement, using words that were presented too fast to be consciously perceived (subliminally). Encephalogram (EEG) and movement kinematics were recorded. EEG recordings of the "Readiness potential" ("RP", indicator of motor preparation) revealed that subliminal displays of action verbs during movement preparation reduced the RP and affected the subsequent reaching movement. The finding that motor processes were modulated by language processes despite the fact that words were not consciously perceived, suggests that cortical structures that serve the preparation and execution of motor actions are indeed part of the (action) language processing network.

Phonological and lexical motor facilitation during speech listening: a transcranial magnetic stimulation study.

In the present study, we used transcranial magnetic stimulation (TMS) to investigate the influence of phonological and lexical properties of verbal items on the excitability of the tongue's cortical motor representation during passive listening. In particular, we aimed to clarify if the difference in tongue motor excitability found during listening to words and pseudo-words [Fadiga, L., Craighero, L., Buccino, G., Rizzolatti, G., 2002. Speech listening specifically modulates the excitability of tongue muscles: a TMS study. European Journal of Neuroscience 15, 399-402] is due to lexical frequency or to the presence of a meaning per se. In order to do this, we investigated the time-course of tongue motor-evoked potentials (MEPs) during listening to frequent words, rare words, and pseudo-words embedded with a double consonant requiring relevant tongue movements for its pronunciation. Results showed that at the later stimulation intervals (200 and 300 ms from the double consonant) listening to rare words evoked much larger MEPs than listening to frequent words. Moreover, by comparing pseudo-words embedded with a double consonant requiring or not tongue movements, we found that a pure phonological motor resonance was present only 100 ms after the double consonant. Thus, while the phonological motor resonance appears very early, the lexical-dependent motor facilitation takes more time to appear and depends on the frequency of the stimuli. The present results indicate that the motor system responsible for phonoarticulatory movements during speech production is also involved during speech listening in a strictly specific way. This motor facilitation reflects both the difference in the phonoarticulatory characteristics and the difference in the frequency of occurrence of the verbal material.