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.

Top–down control is critical to select goal-directed actions in changeable environments, particularly when several conflicting options compete for selection. In humans, this control system is thought to involve an inhibitory mechanism that suppresses the motor representation of unwanted responses to favor selection of the most appropriate action. Here, we aimed to evaluate the role of a region of the medial frontal cortex, the pre-SMA, in this form of inhibition by using a double coil TMS protocol combining repetitive TMS (rTMS) over the pre-SMA and a single-pulse TMS over the primary motor cortex (M1) during a visuomotor task that required participants to choose between a left or right button press according to an imperative cue. M1 stimulation allowed us to assess changes in motor excitability related to selected and nonselected (unwanted) actions, and rTMS was used to produce transient disruption of pre-SMA functioning. We found that when rTMS was applied over pre-SMA, inhibition of the nonselected movement representation was reduced. Importantly, this effect was only observed when the imperative cue produced a substantial amount of competition between the response alternatives. These results are consistent with previous studies pointing to a role of pre-SMA in competition resolution. In addition, our findings indicate that this function of pre-SMA involves the control of inhibitory influences directed at unwanted action representations.

Complex actions can be regarded as a concatenation of simple motor acts, arranged according to specific rules. Because the caudal part of the Broca's region (left Brodmann's area 44, BA 44) is involved in processing hierarchically organized behaviors, we aimed to test the hypothesis that this area may also play a role in learning structured motor sequences. To address this issue, we investigated the inhibitory effects of a continuous theta-burst TMS (cTBS) applied over left BA 44 in healthy subjects, just before they performed a serial RT task (SRTT). SRTT has been widely used to study motor skill learning and is also of interest because, for complex structured sequences, subjects spontaneously organize them into smaller subsequences, referred to as chunks. As a control, cTBS was applied over the vertex in another group, which underwent the same experiment. Control subjects showed both a general practice learning effect, evidenced by a progressive decrease in RT across blocks and a sequence-specific learning effect, demonstrated by a significant RT increase in a pseudorandom sequence. In contrast, when cTBS was applied over left BA 44, subjects lacked both the general practice and sequence-specific learning effects. However, surprisingly, their chunking pattern was preserved and remained indistinguishable from controls. The present study indicates that left BA 44 plays a role in motor sequence learning, but without being involved in elementary chunking. This dissociation between chunking and sequence learning could be explained if we postulate that left BA 44 intervenes in high hierarchical level processing, possibly to integrate elementary chunks together.

Some objects in the visual field are more likely to attract attention because they are either intrinsically eye catching or relevant in the context of a particular task. These two factors, known as stimulus-driven and goal-directed factors, respectively, are thought to be integrated into a unique salience map, possibly located in the frontal or the parietal cortex. However, the distinct contribution of these two regions to salience representation is difficult to establish experimentally and remains debated. In an attempt to address this issue, we designed several dual tasks composed of a letter reporting task and a visual search task, allowing us to quantify the salience of each visual item by measuring its probability to be selected by attention. In Experiment 1, the salience of the visual search items depended on a combination of conspicuity and relevance factors, whereas in Experiment 2, stimulus-driven and goal-directed factors were tested separately. Then, we used transcranial magnetic stimulation to interfere transiently with the function of the right angular gyrus (ANG) or right FEFs in healthy subjects performing these dual tasks. We found that interfering with the ANG and the FEF function specifically altered the influence of salience on the letter report rate without affecting the overall letter reporting rate, suggesting that these areas are involved in salience representation. In particular, the present study suggests that ANG is involved in goal-directed salience representation, whereas FEF would rather house a global salience map integrating both goal-directed and stimulus-driven factors.

One remarkable aspect of the human motor repertoire is the multitude of bimanual actions it contains. Still, the neural correlates of coordinated movements, in which the two hands share a common goal, remain debated. To address this issue, we designed two bimanual circling tasks that differed only in terms of goal conceptualization: a “coordination” task that required movements of both hands to adapt to each other to reach a common goal and an “independent” task that imposed a separate goal to each hand. fMRI allowed us to pinpoint three areas located in the right hemisphere that were more strongly activated in the coordination condition: the superior temporal gyrus (STG), the SMA, and the primary motor cortex (M1). We then used transcranial magnetic stimulation (TMS) to disrupt transiently the function of those three regions to determine their causal role in bimanual coordination. Right STG virtual lesions impaired bimanual coordination, whereas TMS to right M1 enhanced hand independence. TMS over SMA, left STG, or left M1 had no effect. The present study provides direct insight into the neural correlates of coordinated bimanual movements and highlights the role of right STG in such bimanual movements.

It has been suggested that both the posterior parietal cortex (PPC) and the extrastriate occipital cortex (OC) participate in the spatial processing of sounds. However, the precise time-course of their contribution remains unknown, which is of particular interest, considering that it could give new insights into the mechanisms underlying auditory space perception. To address this issue, we have used event-related transcranial magnetic stimulation (TMS) to induce virtual lesions of either the right PPC or right OC at different delays in subjects performing a sound lateralization task. Our results confirmed that these two areas participate in the spatial processing of sounds. More precisely, we found that TMS applied over the right OC 50 msec after the stimulus onset significantly impaired the localization of sounds presented either to the right or to the left side. Moreover, right PPC virtual lesions induced 100 and 150 msec after sound presentation led to a rightward bias for stimuli delivered on the center and on the left side, reproducing transiently the deficits commonly observed in hemineglect patients. The finding that the right OC is involved in sound processing before the right PPC suggests that the OC exerts a feedforward influence on the PPC during auditory spatial processing.

The finding that number processing activates a cortical network partly overlapping that recruited for hand movements has renewed interest in the relationship between number and finger representations. Further evidence about a possible link between fingers and numbers comes from developmental studies showing that finger movements play a crucial role in learning counting. However, increased activity in hand motor circuits during counting may unveil unspecific processes, such as shifting attention, reciting number names, or matching items with a number name. To address this issue, we used transcranial magnetic stimulation to measure changes in corticospinal (CS) excitability during a counting task performed silently and using either numbers or letters of the alphabet to enumerate items. We found an increased CS excitability of hand muscles during the counting task, irrespective of the use of numbers or letters, whereas it was unchanged in arm and foot muscles. Control tasks allowed us to rule out a possible influence of attention allocation or covert speech on CS excitability increase of hand muscles during counting. The present results support a specific involvement of hand motor circuits in counting because no CS changes were found in arm and foot muscles during the same task. However, the contribution of hand motor areas is not exclusively related to number processing because an increase in CS excitability was also found when letters were used to enumerate items. This finding suggests that hand motor circuits are involved whenever items have to be put in correspondence with the elements of any ordered series.

Interhemispheric inhibition (IHI) between motor cortical areas is thought to play a critical role in motor control and could influence manual dexterity. The purpose of this study was to investigate IHI preceding movements of the dominant and nondominant hands of healthy volunteers. Movement-related IHI was studied by means of a double-pulse transcranial magnetic stimulation protocol in right-handed individuals in a simple reaction time paradigm. IHI targeting the motor cortex contralateral (IHI c ) and ipsilateral (IHI i ) to each moving finger was determined. IHI c was comparable after the go signal, a long time preceding movement onset, in both hands. Closer to movement onset, IHI c reversed into facilitation for the right dominant hand but remained inhibitory for left nondominant hand movements. IHI i displayed a nearly constant inhibition with a trough early in the premovement period in both hands. In conclusion, our results unveil a more important modulation of interhemispheric interactions during generation of dominant than nondominant hand movements. This modulation essentially consisted of a shift from a balanced IHI at rest to an IHI predominantly directed toward the ipsilateral primary motor cortex at movement onset. Such a mechanism might release muscles from inhibition in the contralateral primary motor cortex while preventing the occurrence of the mirror activity in ipsilateral primary motor cortex and could therefore contribute to intermanual differences in dexterity.