Scale-invariant changes in corticospinal excitability reflect multiplexed oscillations in the motor output.

In the absence of disease, humans produce smooth and accurate movement trajectories. Despite such 'macroscopic' aspect, the 'microscopic' structure of movements reveals recurrent (quasi-rhythmic) discontinuities. To date, it is unclear how the sensorimotor system contributes to the macroscopic and microscopic architecture of movement. Here, we investigated how corticospinal excitability changes in relation to microscopic fluctuations that are naturally embedded within larger macroscopic variations in motor output. Participants performed a visuomotor tracking task. In addition to the 0.25 Hz modulation that is required for task fulfilment (macroscopic scale), the motor output shows tiny but systematic fluctuations at ∼2 and 8 Hz (microscopic scales). We show that motor-evoked potentials (MEPs) elicited by transcranial magnetic stimulation (TMS) during task performance are consistently modulated at all (time) scales. Surprisingly, MEP modulation covers a similar range at both micro- and macroscopic scales, even though the motor output differs by several orders of magnitude. Thus, corticospinal excitability finely maps the multiscale temporal patterning of the motor output, but it does so according to a principle of scale invariance. These results suggest that corticospinal excitability indexes a relatively abstract level of movement encoding that may reflect the hierarchical organisation of sensorimotor processes. KEY POINTS: Motor behaviour is organised on multiple (time)scales. Small but systematic ('microscopic') fluctuations are engrained in larger and slower ('macroscopic') variations in motor output, which are instrumental in deploying the desired motor plan. Corticospinal excitability is modulated in relation to motor fluctuations on both macroscopic and microscopic (time)scales. Corticospinal excitability obeys a principle of scale invariance, that is, it is modulated similarly at all (time)scales, possibly reflecting hierarchical mechanisms that optimise motor encoding.

Speech perception difficulty modulates theta-band encoding of articulatory synergies.

The human brain tracks available speech acoustics and extrapolates missing information such as the speaker's articulatory patterns. However, the extent to which articulatory reconstruction supports speech perception remains unclear. This study explores the relationship between articulatory reconstruction and task difficulty. Participants listened to sentences and performed a speech-rhyming task. Real kinematic data of the speaker's vocal tract were recorded via electromagnetic articulography (EMA) and aligned to corresponding acoustic outputs. We extracted articulatory synergies from the EMA data with principal component analysis (PCA) and employed partial information decomposition (PID) to separate the electroencephalographic (EEG) encoding of acoustic and articulatory features into unique, redundant, and synergistic atoms of information. We median-split sentences into easy (ES) and hard (HS) based on participants' performance and found that greater task difficulty involved greater encoding of unique articulatory information in the theta band. We conclude that fine-grained articulatory reconstruction plays a complementary role in the encoding of speech acoustics, lending further support to the claim that motor processes support speech perception.NEW & NOTEWORTHY Top-down processes originating from the motor system contribute to speech perception through the reconstruction of the speaker's articulatory movement. This study investigates the role of such articulatory simulation under variable task difficulty. We show that more challenging listening tasks lead to increased encoding of articulatory kinematics in the theta band and suggest that, in such situations, fine-grained articulatory reconstruction complements acoustic encoding.

Excitatory/inhibitory motor balance reflects individual differences during joint action coordination.

Joint action (JA) is a continuous process of motor co-regulation based on the integration of contextual (top-down) and kinematic (bottom-up) cues from partners. The fine equilibrium between excitation and inhibition in sensorimotor circuits is, thus, central to such a dynamic process of action selection and execution. In a bimanual task adapted to become a unimanual JA task, the participant held a bottle (JA), while a confederate had to reach and unscrew either that bottle or another stabilized by a mechanical clamp (No_JA). Prior knowledge was manipulated in each trial such that the participant knew (K) or not (No_K) the target bottle in advance. Online transcranial magnetic stimulation (TMS) was administered at action-relevant landmarks to explore corticospinal excitability (CSE) and inhibition (cortical silent period [cSP]). CSE was modulated early on before the action started if prior information was available. In contrast, cSP modulation emerged later during the reaching action, regardless of prior information. These two indexes could thus reflect the concurrent elaboration of contextual priors (top-down) and the online sampling of partner's kinematic cues (bottom-up). Furthermore, participants selected either one of two possible behavioural strategies, preferring early or late force exertion on the bottle. One translates into a reduced risk of motor coordination failure and the other into reduced metabolic expenditure. Each strategy was characterised by a specific excitatory/inhibitory profile. In conclusion, the study of excitatory/inhibitory balance paves the way for the neurophysiological determination of individual differences in the combination of top-down and bottom-up processing during JA coordination.

Cortico-cortical paired associative stimulation conditioning superficial ventral premotor cortex-primary motor cortex connectivity influences motor cortical activity during precision grip.

The ventral premotor cortex (PMv) and primary motor cortex (M1) represent critical nodes of a parietofrontal network involved in grasping actions, such as power and precision grip. Here, we investigated how the functional PMv-M1 connectivity drives the dissociation between these two actions. We applied a PMv-M1 cortico-cortical paired associative stimulation (cc-PAS) protocol, stimulating M1 in both postero-anterior (PA) and antero-posterior (AP) directions, in order to induce long-term changes in the activity of different neuronal populations within M1. We evaluated the motor-evoked potential (MEP) amplitude, MEP latency and cortical silent period, in both PA and AP, during the isometric execution of precision and power grip, before and after the PMv-M1 cc-PAS. The repeated activation of the PMv-M1 cortico-cortical network with PA orientation over M1 did not change MEP amplitude or cortical silent period duration during both actions. In contrast, the PMv-M1 cc-PAS stimulation of M1 with an AP direction led to a specific modulation of precision grip motor drive. In particular, MEPs tested with AP stimulation showed a selective increase of corticospinal excitability during precision grip. These findings suggest that the more superficial M1 neuronal populations recruited by the PMv input are involved preferentially in the execution of precision grip actions. KEY POINTS: Ventral premotor cortex (PMv)-primary motor cortex (M1) cortico-cortical paired associative stimulation (cc-PAS) with different coil orientation targets dissociable neural populations. PMv-M1 cc-PAS with M1 antero-posterior coil orientation specifically modulates corticospinal excitability during precision grip. Superficial M1 populations are involved preferentially in the execution of precision grip. A plasticity induction protocol targeting the specific PMv-M1 subpopulation might have important translational value for the rehabilitation of hand function.

Mechanisms of Hebbian-like plasticity in the ventral premotor - primary motor network.

The functional connection between ventral premotor cortex (PMv) and primary motor cortex (M1) is critical for the organization of goal-directed actions. Repeated activation of this connection by means of cortico-cortical paired associative stimulation (cc-PAS), a transcranial magnetic stimulation (TMS) protocol, may induce Hebbian-like plasticity. However, the physiological modifications produced by Hebbian-like plasticity in the PMv-M1 network are poorly understood. To fill this gap, we investigated the effects of cc-PAS on PMv-M1 circuits. We hypothesized that specific interactions would occur with I2 -wave interneurons as measured by the short intracortical facilitation protocol (SICF). We used different paired-pulse TMS protocols to examine the effects of PMv-M1 cc-PAS on SICF, on GABAergic circuits as measured by short (SICI) and long (LICI) intracortical inhibition protocols, and varied the current direction in M1 to target different M1 neuronal populations. Finally, we examined the effects of cc-PAS on PMv-M1 connectivity using a dual coil approach. We found that PMv-M1 cc-PAS induces both a long-term potentiation (LTP)- or long-term depression (LTD)-like after-effect in M1 neuronal activity that is strongly associated with a bidirectional-specific change in I2 -wave activity (SICF = 2.5 ms ISI). Moreover, cc-PAS induces a specific modulation of the LICI circuit and separately modulates PMv-M1 connectivity. We suggest that plasticity within the PMv-M1 circuit is mediated by a selective mechanism exerted by PMv on M1 by targeting I2 -wave interneurons. These results provide new mechanistic insights into how PMv modulates M1 activity that are relevant for the design of brain stimulation protocols in health and disease. KEY POINTS: The I2 -wave is specifically modulated by the induction of ventral premotor cortex - primary motor cortex (PMv-M1) plasticity. After PMv-M1 cortico-cortical paired associative stimulation (cc-PAS), corticospinal excitability correlates negatively with I2 -wave amplitude. Different cc-PAS coil orientations can lead to a long-term potentiation- or long-term depression-like after-effect in M1.

Modulatory Effects of Communicative Gaze on Attentional Orienting Are Driven by Dorsomedial Prefrontal Cortex but Not Right Temporoparietal Junction.

Communicative gaze (e.g., mutual or averted) has been shown to affect attentional orienting. However, no study to date has clearly separated the neural basis of the pure social component that modulates attentional orienting in response to communicative gaze from other processes that might be a combination of attentional and social effects. We used TMS to isolate the purely social effects of communicative gaze on attentional orienting. Participants completed a gaze-cueing task with a humanoid robot who engaged either in mutual or in averted gaze before shifting its gaze. Before the task, participants received either sham stimulation (baseline), stimulation of right TPJ (rTPJ), or dorsomedial prefrontal cortex (dmPFC). Results showed, as expected, that communicative gaze affected attentional orienting in baseline condition. This effect was not evident for rTPJ stimulation. Interestingly, stimulation to rTPJ also canceled out attentional orienting altogether. On the other hand, dmPFC stimulation eliminated the socially driven difference in attention orienting between the two gaze conditions while maintaining the basic general attentional orienting effect. Thus, our results allowed for separation of the pure social effect of communicative gaze on attentional orienting from other processes that are a combination of social and generic attentional components.

The microstructure of intra- and interpersonal coordination.

Movements are naturally composed of submovements, i.e. recurrent speed pulses (2-3 Hz), possibly reflecting intermittent feedback-based motor adjustments. In visuomotor (unimanual) synchronization tasks, partners alternate submovements over time, indicating mutual coregulation. However, it is unclear whether submovement coordination is organized differently between and within individuals. Indeed, different types of information may be variably exploited for intrapersonal and interpersonal coordination. Participants performed a series of bimanual tasks alone or in pairs, with or without visual feedback (solo task only). We analysed the relative timing of submovements between their own hands or between their own hands and those of their partner. Distinct coordinative structures emerged at the submovement level depending on the relevance of visual feedback. Specifically, the relative timing of submovements (between partners/effectors) shifts from alternation to simultaneity and a mixture of both when coordination is achieved using vision (interpersonal), proprioception/efference-copy only (intrapersonal, without vision) or all information sources (intrapersonal, with vision), respectively. These results suggest that submovement coordination represents a behavioural proxy for the adaptive weighting of different sources of information within action-perception loops. In sum, the microstructure of movement reveals common principles governing the dynamics of sensorimotor control to achieve both intra- and interpersonal coordination.

Visual detection is locked to the internal dynamics of cortico-motor control.

Movements overtly sample sensory information, making sensory analysis an active-sensing process. In this study, we show that visual information sampling is not just locked to the (overt) movement dynamics but to the internal (covert) dynamics of cortico-motor control. We asked human participants to perform continuous isometric contraction while detecting unrelated and unpredictable near-threshold visual stimuli. The motor output (force) shows zero-lag coherence with brain activity (recorded via electroencephalography) in the beta-band, as previously reported. In contrast, cortical rhythms in the alpha-band systematically forerun the motor output by 200 milliseconds. Importantly, visual detection is facilitated when cortico-motor alpha (not beta) synchronization is enhanced immediately before stimulus onset, namely, at the optimal phase relationship for sensorimotor communication. These findings demonstrate an ongoing coupling between visual sampling and motor control, suggesting the operation of an internal and alpha-cycling visuomotor loop.

Movement kinematics drive chain selection toward intention detection.

The ability to understand intentions based on another's movements is crucial for human interaction. This ability has been ascribed to the so-called motor chaining mechanism: anytime a motor chain is activated (e.g., grasp-to-drink), the observer attributes to the agent the corresponding intention (i.e., to drink) from the first motor act (i.e., the grasp). However, the mechanisms by which a specific chain is selected in the observer remain poorly understood. In the current study, we investigate the possibility that in the absence of discriminative contextual cues, slight kinematic variations in the observed grasp inform mapping to the most probable chain. Chaining of motor acts predicts that, in a sequential grasping task (e.g., grasp-to-drink), electromyographic (EMG) components that are required for the final act [e.g., the mouth-opening mylohyoid (MH) muscle] show anticipatory activation. To test this prediction, we used MH EMG, transcranial magnetic stimulation (TMS; MH motor-evoked potentials), and predictive models of movement kinematics to measure the level and timing of MH activation during the execution (Experiment 1) and the observation (Experiment 2) of reach-to-grasp actions. We found that MH-related corticobulbar excitability during grasping observation varied as a function of the goal (to drink or to pour) and the kinematics of the observed grasp. These results show that subtle changes in movement kinematics drive the selection of the most probable motor chain, allowing the observer to link an observed act to the agent's intention.

Motor cortical inhibition during concurrent action execution and action observation.

Action Execution (AE) and Action Observation (AO) share an extended cortical network of activated areas. During coordinative action these processes also overlap in time, potentially giving rise to behavioral interference effects. The neurophysiological mechanisms subtending the interaction between concurrent AE and AO are substantially unknown. To assess the effect of AO on observer's corticomotor drive, we run one electromyography (EMG) and three Transcranial Magnetic Stimulation (TMS) studies. Participants were requested to maintain a steady hand opening or closing posture while observing the same or a different action (hand opening and closing in the main TMS study). By measuring Cortical Silent Periods (CSP), an index of GABAB-mediated corticospinal inhibitory strength, we show a selective reduction of inhibitory motor drive for mismatching AE-AO pairs. The last two TMS experiments, show that this mismatch is computed according to a muscle-level agonist-antagonist representation. Combined, our results suggest that corticospinal inhibition may be the central neurophysiological mechanism by which one's own motor execution is adapted to the contextual visual cues provided by other's actions.

The neural oscillatory markers of phonetic convergence during verbal interaction.

During a conversation, the neural processes supporting speech production and perception overlap in time and, based on context, expectations and the dynamics of interaction, they are also continuously modulated in real time. Recently, the growing interest in the neural dynamics underlying interactive tasks, in particular in the language domain, has mainly tackled the temporal aspects of turn-taking in dialogs. Besides temporal coordination, an under-investigated phenomenon is the implicit convergence of the speakers toward a shared phonetic space. Here, we used dual electroencephalography (dual-EEG) to record brain signals from subjects involved in a relatively constrained interactive task where they were asked to take turns in chaining words according to a phonetic rhyming rule. We quantified participants' initial phonetic fingerprints and tracked their phonetic convergence during the interaction via a robust and automatic speaker verification technique. Results show that phonetic convergence is associated to left frontal alpha/low-beta desynchronization during speech preparation and by high-beta suppression before and during listening to speech in right centro-parietal and left frontal sectors, respectively. By this work, we provide evidence that mutual adaptation of speech phonetic targets, correlates with specific alpha and beta oscillatory dynamics. Alpha and beta oscillatory dynamics may index the coordination of the "when" as well as the "how" speech interaction takes place, reinforcing the suggestion that perception and production processes are highly interdependent and co-constructed during a conversation.

Passive sensorimotor stimulation triggers long lasting alpha-band fluctuations in visual perception.

Movement planning and execution rely on the anticipation and online control of the incoming sensory input. Evidence suggests that sensorimotor processes may synchronize visual rhythmic activity in preparation of action performance. Indeed, we recently reported periodic fluctuations of visual contrast sensitivity that are time-locked to the onset of an intended movement of the arm. However, the origin of the observed visual modulations has so far remained unclear because of the endogenous (and thus temporally undetermined) activation of the sensorimotor system that is associated with voluntary movement initiation. In this study, we activated the sensorimotor circuitry involved in the hand control in an exogenous and controlled way by means of peripheral stimulation of the median nerve and characterized the spectrotemporal dynamics of the ensuing visual perception. The stimulation of the median nerve triggers robust and long-lasting (∼1 s) alpha-band oscillations in visual perception, whose strength is temporally modulated in a way that is consistent with the changes in alpha power described at the neurophysiological level after sensorimotor stimulation. These findings provide evidence in support of a causal role of the sensorimotor system in modulating oscillatory activity in visual areas with consequences for visual perception. NEW & NOTEWORTHY This study shows that the peripheral activation of the somatomotor hand system triggers long-lasting alpha periodicity in visual perception. This demonstrates that not only the endogenous sensorimotor processes involved in movement preparation but also the passive stimulation of the sensorimotor system can synchronize visual activity. The present work suggests that oscillation-based mechanisms may subserve core (task independent) sensorimotor integration functions.

Disruption of Broca's Area Alters Higher-order Chunking Processing during Perceptual Sequence Learning.

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.

Listener-speaker perceived distance predicts the degree of motor contribution to speech perception.

Listening speech sounds activates motor and premotor areas in addition to temporal and parietal brain regions. These activations are somatotopically localized according to the effectors recruited in the production of particular phonemes. Previous work demonstrated that transcranial magnetic stimulation (TMS) of speech motor centers somatotopically altered speech perception, suggesting a role for the motor system. However, these effects seemed to occur only under adverse listening conditions, suggesting that degraded speech may stimulate listeners to adopt unnatural neural strategies relying on motor centers. Here, we investigated whether naturally occurring interspeaker variability, which did not affect task difficulty, made a speech discrimination task sensitive to TMS interference. In this paradigm, TMS over tongue and lips motor representations somatotopically altered the discrimination time of speech. Furthermore, the TMS-induced effect correlated with listeners' similarity judgments between listeners' and speakers' speech productions. Thus, the degree of motor recruitment depends on the perceived distance between listener and speaker. This result supports the claim that discriminating others' speech pattern requires the contribution of the listener's own motor repertoire. We conclude that motor recruitment in speech perception can be a natural product of discriminating speech in a normally variable and unpredictable environment, not merely related to task difficulty.

A theory for how sensorimotor skills are learned and retained in noisy and nonstationary neural circuits.

During the process of skill learning, synaptic connections in our brains are modified to form motor memories of learned sensorimotor acts. The more plastic the adult brain is, the easier it is to learn new skills or adapt to neurological injury. However, if the brain is too plastic and the pattern of synaptic connectivity is constantly changing, new memories will overwrite old memories, and learning becomes unstable. This trade-off is known as the stability-plasticity dilemma. Here a theory of sensorimotor learning and memory is developed whereby synaptic strengths are perpetually fluctuating without causing instability in motor memory recall, as long as the underlying neural networks are sufficiently noisy and massively redundant. The theory implies two distinct stages of learning--preasymptotic and postasymptotic--because once the error drops to a level comparable to that of the noise-induced error, further error reduction requires altered network dynamics. A key behavioral prediction derived from this analysis is tested in a visuomotor adaptation experiment, and the resultant learning curves are modeled with a nonstationary neural network. Next, the theory is used to model two-photon microscopy data that show, in animals, high rates of dendritic spine turnover, even in the absence of overt behavioral learning. Finally, the theory predicts enhanced task selectivity in the responses of individual motor cortical neurons as the level of task expertise increases. From these considerations, a unique interpretation of sensorimotor memory is proposed--memories are defined not by fixed patterns of synaptic weights but, rather, by nonstationary synaptic patterns that fluctuate coherently.

Generalization of motor resonance during the observation of hand, mouth, and eye movements.

Transcranial magnetic stimulation (TMS) of the motor cortex shows that hand action observation (AO) modulates corticospinal excitability (CSE). CSE modulation alternatively maps low-level kinematic characteristics or higher-level features, like object-directed action goals. However, action execution is achieved through the control of muscle synergies, consisting of coordinated patterns of muscular activity during natural movements, rather than single muscles or object-directed goals. This synergistic organization of action execution also underlies the ability to produce the same functional output (i.e., grasping an object) using different effectors. We hypothesize that motor system activation during AO may rely on similar principles. To investigate this issue, we recorded both hand CSE and TMS-evoked finger movements which provide a much more complete description of coordinated patterns of muscular activity. Subjects passively watched hand, mouth and eyelid opening or closing, which are performing non-object-directed (intransitive) actions. Hand and mouth share the same potential to grasp objects, whereas eyelid does not allow object-directed (transitive) actions. Hand CSE modulation generalized to all effectors, while TMS evoked finger movements only to mouth AO. Such dissociation suggests that the two techniques may have different sensitivities to fine motor modulations induced by AO. Differently from evoked movements, which are sensitive to the possibility to achieve object-directed action, CSE is generically modulated by "opening" vs. "closing" movements, independently of which effector was observed. We propose that motor activities during AO might exploit the same synergistic mechanisms shown for the neural control of movement and organized around a limited set of motor primitives.

Domain-specific and domain-general processes in social perception--A complementary approach.

In this brief discussion, we explicate and evaluate Heyes and colleagues' deflationary approach to interpreting apparent evidence of domain-specific processes for social perception. We argue that the deflationary approach sheds important light on how functionally specific processes in social perception can be subserved at least in part by domain-general processes. On the other hand, we also argue that the fruitfulness of this approach has been unnecessarily hampered by a contrastive conception of the relationship between domain-general and domain-specific processes. As an alternative, we propose a complementary conception: the identification of domain-general processes that are engaged in instances of social perception can play a positive, structuring role by adding additional constraints to be accounted for in modelling the domain-specific processes that are also involved in such instances.

Automatic imitation of the arm kinematic profile in interacting partners.

Cognitive neuroscience, traditionally focused on individual brains, is just beginning to investigate social cognition through realistic interpersonal interaction. However, quantitative investigation of the dynamical sensorimotor communication among interacting individuals in goal-directed ecological tasks is particularly challenging. Here, we recorded upper-body motion capture of 23 dyads, alternating their leader/follower role, in a tower-building task. Either a strategy of joining efforts or a strategy of independent action could in principle be used. We found that arm reach velocity profiles of participants tended to converge across trials. Automatic imitation of low-level motor control parameters demonstrates that the task is achieved through continuous action coordination as opposed to independent action planning. Moreover, the leader produced more consistent and predictable velocity profiles, suggesting an implicit strategy of signaling to the follower. This study serves as a validation of our joint goal-directed non-verbal task for future applications. In fact, the quantification of human-to-human continuous sensorimotor interaction, in a way that can be predicted and controlled, is probably one of the greatest challenges for the future of human-robot interaction.

Submovement interpersonal coupling is associated to audio-motor coordination performance.

Acting in concert with others, a key aspect of our social life, requires behavioral coordination between persons on multiple timescales. When zooming in on the kinematic properties of movements, it appears that small speed fluctuations, called submovements, are embedded within otherwise smooth end-point trajectories. Submovements, by occurring at a faster timescale than that of movements, offer a novel window upon the functional relationship between distinct motor timescales. In this regard, it has previously been shown that when partners visually synchronize their movements, they also coordinate the timing of their submovement by following an alternated pattern. However, it remains unclear whether the mechanisms behind submovement coordination are domain-general or specific to the visual modality, and whether they have relevance for interpersonal coordination also at the scale of whole movements. In a series of solo and dyadic tasks, we show that submovements are also present and coordinated across partners when sensorimotor interactions are mediated by auditory feedback only. Importantly, the accuracy of task-instructed interpersonal coordination at the movement level correlates with the strength of submovement coordination. These results demonstrate that submovement coordination is a potentially fundamental mechanism that participates in interpersonal motor coordination regardless of the sensory domain mediating the interaction.

The role of dorsal premotor cortex in joint action inhibition.

Behavioral interpersonal coordination requires smooth negotiation of actions in time and space (joint action-JA). Inhibitory control may play a role in fine-tuning appropriate coordinative responses. To date, little research has been conducted on motor inhibition during JA and on the modulatory influence that premotor areas might exert on inhibitory control. Here, we used an interactive task in which subjects were required to reach and open a bottle using one hand. The bottle was held and stabilized by a co-actor (JA) or by a mechanical holder (vice clamp, no-JA). We recorded two TMS-based indices of inhibition (short-interval intracortical inhibition-sICI; cortical silent period-cSP) during the reaching phase of the task. These reflect fast intracortical (GABAa-mediated) and slow corticospinal (GABAb-mediated) inhibition. Offline continuous theta burst stimulation (cTBS) was used to interfere with dorsal premotor cortex (PMd), ventral premotor cortex (PMv), and control site (vertex) before the execution of the task. Our results confirm a dissociation between fast and slow inhibition during JA coordination and provide evidence that premotor areas drive only slow inhibitory mechanisms, which in turn may reflect behavioral co-adaptation between trials. Exploratory analyses further suggest that PMd, more than PMv, is the key source of modulatory drive sculpting movements, according to the socio-interactive context.