The extrastriate body area (EBA) is a body‐selective focal region located in the lateral occipito‐temporal cortex that responds strongly to images of human bodies and body parts in comparison with ...other classes of stimuli. Whether EBA contributes also to the body recognition of self versus others remains in debate. We investigated whether EBA contributes to self‐other distinction and whether there might be a hemispheric‐side specificity to that contribution using double‐pulse transcranial magnetic stimulation (TMS) in right‐handed participants. Prior to the TMS experiment, all participants underwent an fMRI localizer task to determine individual EBA location. TMS was then applied over either right EBA, left EBA or vertex, while participants performed an identification task in which images of self or others' right, or left hands were presented. TMS over both EBAs slowed responses, with no identity‐specific effect. However, TMS applied over right EBA induced significantly more errors on other's hands than noTMS, TMS over left EBA or over the Vertex, when applied at 100–110 ms after image onset. The last three conditions did not differ, nor was there any difference for self‐hands. These findings suggest that EBA participates in self/other discrimination.
The present study provides evidence that EBA in the right hemisphere participates in identity processing. TMS over the right EBA might disrupt the early processing of visual signals that would be important in an explicit recognition task, leading to decreased response speed and accuracy.
Some language processing theories propose that, just as for other somatic actions, self-monitoring of language production is achieved through internal modeling. The cerebellum is the proposed center ...of such internal modeling in motor control, and the right cerebellum has been linked to an increasing number of language functions, including predictive processing during comprehension. Relating these findings, we tested whether the right posterior cerebellum has a causal role for self-monitoring of speech errors. Participants received 1 Hz repetitive transcranial magnetic stimulation during 15 min to lobules Crus I and II in the right hemisphere, and, in counterbalanced orders, to the contralateral area in the left cerebellar hemisphere (control) in order to induce a temporary inactivation of one of these zones. Immediately afterwards, they engaged in a speech production task priming the production of speech errors. Language production was impaired after right compared to left hemisphere stimulation, a finding that provides evidence for a causal role of the cerebellum during language production. We interpreted this role in terms of internal modeling of upcoming speech through a verbal working memory process used to prevent errors.
The purpose of the present study was to investigate whether corticospinal projections from human supplementary motor area (SMA) are functional during precise force control with the precision grip ...(thumb-index opposition). Since beta band corticomuscular coherence (CMC) is well-accepted to reflect efferent corticospinal transmission, we analyzed the beta band CMC obtained with simultaneous recording of electroencephalographic (EEG) and electromyographic (EMG) signals. Subjects performed a bimanual precise visuomotor force tracking task by applying isometric low grip forces with their right hand precision grip on a custom device with strain gauges. Concurrently, they held the device with their left hand precision grip, producing similar grip forces but without any precision constraints, to relieve the right hand. Some subjects also participated in a unimanual control condition in which they performed the task with only the right hand precision grip while the device was held by a mechanical grip. We analyzed whole scalp topographies of beta band CMC between 64 EEG channels and 4 EMG intrinsic hand muscles, 2 for each hand. To compare the different topographies, we performed non-parametric statistical tests based on spatio-spectral clustering. For the right hand, we obtained significant beta band CMC over the contralateral M1 region as well as over the SMA region during static force contraction periods. For the left hand, however, beta band CMC was only found over the contralateral M1. By comparing unimanual and bimanual conditions for right hand muscles, no significant difference was found on beta band CMC over M1 and SMA. We conclude that the beta band CMC found over SMA for right hand muscles results from the precision constraints and not from the bimanual aspect of the task. The result of the present study strongly suggests that the corticospinal projections from human SMA become functional when high precision force control is required.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
The corticospinal (CS) system plays an important role in fine motor control, especially in precision grip tasks. Although the primary motor cortex (M1) is the main source of the CS projections, other ...projections have been found, especially from the supplementary motor area proper (SMAp). To study the characteristics of these CS projections from SMAp, we compared muscle responses of an intrinsic hand muscle (FDI) evoked by stimulation of human M1 and SMAp during an isometric static low‐force control task. Subjects were instructed to maintain a small cursor on a target force curve by applying a pressure with their right precision grip on a force sensor. Neuronavigated transcranial magnetic stimulation was used to stimulate either left M1 or left SMAp with equal induced electric field values at the defined cortical targets. The results show that the SMAp stimulation evokes reproducible muscle responses with similar latencies and amplitudes as M1 stimulation, and with a clear and significant shorter silent period. These results suggest that (i) CS projections from human SMAp are as rapid and efficient as those from M1, (ii) CS projections from SMAp are directly involved in control of the excitability of spinal motoneurons and (iii) SMAp has a different intracortical inhibitory circuitry. We conclude that human SMAp and M1 both have direct influence on force production during fine manual motor tasks.
We studied the muscle responses evoked by TMS of either SMAp or M1 during a precision grip task. TMS of SMAp evoked muscle responses with similar latencies and amplitudes as TMS of M1, but with a shorter silent period. This suggests that SMAp has a different intracortical inhibitory circuitry and that corticospinal projections from SMAp are directly involved in the control of excitability of spinal motoneurons. We conclude that SMAp and M1 both have direct influence on precise force production.
Charles Capaday ,
Brigitte A. Lavoie ,
Hugues Barbeau ,
Cyril Schneider , and
Mireille Bonnard
Centre de Recherche Université Laval-Robert Giffard, Department of Anatomy and Physiology, Université ...Laval, Quebec City, Quebec H3G 1Y5, Canada
Capaday, Charles, Brigitte A. Lavoie, Hugues Barbeau, Cyril Schneider, and Mireille Bonnard. Studies on the corticospinal control of human walking. I. Responses to focal transcranial magnetic stimulation of the motor cortex. J. Neurophysiol. 81: 129-139, 1999. Experiments were done to determine the extent to which the corticospinal tract is linked with the segmental motor circuits controlling ankle flexors and extensors during human walking compared with voluntary motor tasks requiring attention to the level of motor activity. The motor cortex was activated transcranially using a focal magnetic stimulation coil. For each subject, the entire input-output (I-O) curve i.e., the integral of the motor evoked-potential (MEP) versus stimulus strength was measured during a prescribed tonic voluntary contraction of either the tibialis anterior (TA) or the soleus. Similarly, I-O curves were measured in the early part of the swing phase, or in the early part of the stance phase of walking. The I-O data points were fitted by the Boltzmann sigmoidal function, which accounted for 80% of total data variance. There was no statistically significant difference between the I-O curves of the TA measured during voluntary ankle dorsiflexion or during the swing phase of walking, at matched levels of background electromyographic (EMG) activity. Additionally, there was no significant difference in the relation between the coefficient of variation and the amplitude of the MEPs measured in each task, respectively. In comparison, during the stance phase of walking the soleus MEPs were reduced on average by 26% compared with their size during voluntary ankle plantarflexion. Furthermore, during stance the MEPs in the inactive TA were enhanced relative to their size during voluntary ankle plantarflexion and in four of six subjects the TA MEPs were larger than those of the soleus. Finally, stimulation of the motor cortex at various phases of the step cycle did not reset the cycle. The time of the next step occurred at the expected moment, as determined from the phase-resetting curve. One interpretation of this result is that the motor cortex may not be part of the central neural system involved in timing the motor bursts during the step cycle. We suggest that during walking the corticospinal tract is more closely linked with the segmental motor circuits controlling the flexor, TA, than it is with those controlling the extensor, soleus. However, during voluntary tasks requiring attention to the level of motor activity, it is equally linked with the segmental motor circuits of ankle flexors or extensors.
While part of the left ventral occipito-temporal cortex (left-vOT), known as the Visual Word Form Area, plays a central role in reading, the area also responds to speech. This cross-modal activation ...has been explained by three competing hypotheses. Firstly, speech is converted to orthographic representations that activate, in a top-down manner, written language coding neurons in the left-vOT. Secondly, the area contains multimodal neurons that respond to both language modalities. Thirdly, the area comprises functionally segregated neuronal populations that selectively encode different language modalities. A transcranial magnetic stimulation (TMS)-adaptation protocol was used to disentangle these hypotheses. During adaptation, participants were exposed to spoken or written words in order to tune the initial state of left-vOT neurons to one of the language modalities. After adaptation, they performed lexical decisions on spoken and written targets with TMS applied to the left-vOT. TMS showed selective facilitatory effects. It accelerated lexical decisions only when the adaptors and the targets shared the same modality, i.e., when left-vOT neurons had initially been adapted to the modality of the target stimuli. Since this within-modal adaptation was observed for both input modalities and no evidence for cross-modal adaptation was found, our findings suggest that the left-vOT contains neurons that selectively encode written and spoken language rather than purely written language coding neurons or multimodal neurons encoding language regardless of modality.
•fMRI data showed that the Visual Word Form Area also responds to spoken input.•We explored the underlying neuronal mechanisms of this cross-modal activation.•TMS adaptation protocol was used to examine the properties of neurons in this brain area.•We found that VWFA contained functionally segregated neuronal populations.•These populations selectively encode either written or spoken language input.
As an interface between the visual and language system, the left ventral occipitotemporal cortex (left-vOT) plays a key role in reading. This functional role is supported by anatomical and functional ...connections between the area and other brain regions within and outside the language network. Nevertheless, only a few studies have investigated how the functional state of this area, which is dependent upon the nature of the task demand and the stimulus being processed, could influence the activity of the connected brain regions. In the present combined TMS-EEG study, we studied the left-vOT effective connectivity by adopting a direct, causal intervention approach. Using TMS, we probed left-vOT activation in different processing contexts and measured the neural propagation of activity from this area to other brain regions. A comparison of neural propagation measured during low-level visual detection of language versus non-language stimuli showed that processing language stimuli reduced neural propagation from the left-vOT to the right occipital cortex. Additionally, compared to the low-level visual detection of language stimuli, performing semantic judgments on the same stimuli further reduced neural propagation to the bilateral posterior cingulate, right superior parietal lobule and the right anterior temporal lobe. This reduction of cross-hemispheric neural propagation was accompanied by an increase in the collaboration between areas within the lefthemisphere language network. Together, this first evidence from a direct causal intervention approach suggests that processing language stimuli and performing a high-level language task reduce effective connectivity from the left-vOT to the right hemisphere, and may contribute to the left-hemisphere lateralization typically observed during language processing.
As an interface between the visual and language system, the left ventral occipito-temporal cortex (left-vOT) plays a key role in reading. This functional role is supported by anatomical and ...functional connections between the area and other brain regions within and outside the language network. Nevertheless, only a few studies have investigated how the functional state of this area, which is dependent upon the nature of the task demand and the stimulus being processed, could influence the activity of the connected brain regions. In the present combined TMS-EEG study, we studied the left-vOT effective connectivity by adopting a direct, causal intervention approach. Using TMS, we probed left-vOT activation in different processing contexts and measured the neural propagation of activity from this area to other brain regions. A comparison of neural propagation measured during low-level visual detection of language versus non-language stimuli showed that processing language stimuli reduced neural propagation from the left-vOT to the right occipital cortex. Additionally, compared to the low-level visual detection of language stimuli, performing semantic judgments on the same stimuli further reduced neural propagation to the posterior part of the corpus callosum, right superior parietal lobule and the right anterior temporal lobe. This reduction of cross-hemispheric neural propagation was accompanied by an increase in the collaboration between areas within the left-hemisphere language network. Together, this first evidence from a direct causal intervention approach suggests that processing language stimuli and performing a high-level language task reduce effective connectivity from the left-vOT to the right hemisphere, and may contribute to the left-hemisphere lateralization typically observed during language processing.
This experiment investigates the interaction of different sensory cues in the control of propulsive forces in human gait which in turn allow the body's forward progression to be regulated. The aim of ...this work was to determine how optic flow and leg-somatosensory feedback interact in this control. We therefore determined whether the responses to sinusoidal perturbations of optic flow were accentuated when leg-somatosensory feedback was modified by varying the support resistance. Subjects walked on a treadmill which was driven by their own locomotor activity (1) with a sinusoidal variation of optic flow velocity, (2) with a sinusoidal variation of support resistance which modified leg-somatosensory information and (3) with both visual and leg-somatosensory modification at different frequencies. The response of the subject was measured as changes in speed and propulsive power. The response to sinusoidal perturbations of optic flow was found to be increased and time delayed when visual perturbations are coupled with support perturbations in comparison with the response observed with visual perturbations only. This result shows the influence of leg-somatosensory feedback on the weighting of optic flow. Inversely, it was also found that the motor response to support perturbation was different when the flow was congruent (i.e., corresponding to the subject's virtual speed) and when it was not. This latter result shows the influence of optic flow on the weighting of leg-somatosensory feedback. The interaction between optic flow and leg-somatosensory feedback argues in favor of a multimodal sensory control of propulsive forces. This multimodal sensory control would be based on all the sensory feedback and all their mutual sensorial interaction. Therefore, the modification of one sensory input modifies not only this input but also the integration of the other inputs.
Voluntary movement is often perturbed by the external forces in the environment. Because corticospinal (CS) control of wrist muscles during preparation of voluntary movement has been extensively ...studied without variation in the external forces, very little is known about the way CS control adapts when subjects expect motor perturbations. Here, we studied the CS control of wrist muscles during expectation of an imposed wrist extension. Subjects were instructed either to compensate (COMP) the perturbation (applied at variable delays) or not to intervene (NINT). In a quarter of all trials at random, in the time window when perturbation might occur, TMS was applied over contralateral M1. Motor evoked potentials (MEPs) were measured in the FCR (flexor carpi radialis) and ECR (extensor carpi radialis) muscles, as well as the silent period (SP) in the FCR. Following the perturbation, we found a larger long-latency stretch reflex in COMP than in NINT. During the expectation of the perturbation, MEP amplitudes did not differ across conditions in FCR. However, those evoked in ECR were greater in COMP than in NINT condition. Moreover in the FCR, the silent period lasted longer in NINT. Thus, we showed a selective effect of the prepared reaction on the anticipatory tuning of CS excitability and cortical inhibition in the agonist/antagonist muscles. This tuning clearly differed from the tuning during voluntary movement preparation without variation in the external forces. This shows that the tuning of the CS system during motor preparation depends on the dynamical context of movement production.
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