Motor Cortical Representation Research Articles

P 74. Practice and stimulation induce shifts in TMS-evoked 2D movements involving multiple joints of the upper limb

Goal-directed reaching is important for activities of daily living. The population of neurons in the primary motor cortex (M1) is known to represent kinematic characteristics of reaching movements. Previous research in our laboratory used transcranial magnetic stimulation (TMS) to reveal maps of reaching within M1 that varied considerably by subject and stimulus location, but remained consistent over time, providing a substrate to assess practice-related changes in motor representation (Jones-Lush et al., 2010). We investigated how repetitive practice of goal-directed reaching movements induces use-dependent plasticity of kinematic characteristics of reaching movements encoded in M1. Healthy individuals (N = 22) sat with their arm in a robotic manipulandum while TMS was applied over M1. Movements were evoked with 120% movement threshold TMS intensity applied over M1. Two baseline measurements were obtained to establish the stability of TMS-evoked movements. Participants then practiced 3 blocks of 160 goal-directed reaching movements in a direction opposite to the baseline direction (14 cm reach 180° from baseline direction) against a 75 N m spring field. Changes in TMS-evoked arm movements were assessed after each practice block and after 5 min following the end of practice. In a second study, subthreshold M1 stimulation coordinated with voluntary reaching movement was added on alternate practice trials, with stimulation either triggered by EMG or preceding EMG by linking to the go cue. Kinematic characteristics of TMS-evoked arm movements were consistent for direction (p = 0.339) and amplitude (p = 0.881) across the two baseline measurements. Direction and the position of the point of peak velocity of TMS-evoked movements were significantly altered following training (p = 0.001 for direction, and p = 0.003 for position), and after 5-min interval following training (p = 0.0316 for direction and p = 0.01 for position). This was accompanied by significant changes in the motor evoked potentials (MEPs) of the shoulder (p = 0.04) and elbow (p = 0.02) agonist muscles indicating an association between the motor cortical physiology and kinematics of TMS-evoked movements. When stimulation was added to practice, preliminary results suggest an increased magnitude and rapidity of shift in kinematics. These findings demonstrate that repetitive practice of functional multi-joint movements modulates the motor cortical representation within M1 in a way that supports the performance of practiced reaching movements, although there are inter-individual differences that may have implications for rehabilitation. Stimulation to M1 has the potential to enhance this plasticity (Bütefisch et al., 2004), and we are currently determining the best timing paradigm to take advantage of this form of associational plasticity. Support: USPHS NIH R01 HD061462.

Differentiation of Motor Cortical Representation of Hand Muscles by Navigated Mapping of Optimal TMS Current Directions in Healthy Subjects

The precision of navigated transcranial magnetic stimulation (TMS) to map the human primary motor cortex may be effected by the direction of TMS-induced current in the brain as determined by the orientation of the stimulation coil. In this study, the authors investigated the effect of current directionality on motor output mapping using navigated brain stimulation. The goal of this study was to determine the optimal coil orientation (and, thus, induced brain current) to activate hand musculature representations relative to each subject's unique neuroanatomical landmarks. The authors studied motor output maps for the first dorsal interosseous, abductor pollicis brevis, and abductor digiti minimi muscles in 10 normal volunteers. Monopolar current pulses were delivered through a figure-of-eight-shaped TMS coil, and motor evoked potentials were recorded using electromyography. At each targeted brain region, the authors systematically rotated the TMS coil to determine the direction of induced current in the brain for induction of the largest motor evoked potentials. These optimal current directions were expressed as an angle relative to each subject's central sulcus. Consistency of the optimal current direction was assessed by repeating the entire mapping procedure on two different occasions across subjects. The authors demonstrate that systematic optimization of current direction as guided by MRI-based neuronavigation improves the resolution of cortical output motor mapping with TMS.

Long-term TENS treatment decreases cortical motor representation in multiple sclerosis

This study investigated the effects of a long-term transcutaneous electrical nerve stimulation (TENS) treatment on cortical motor representation in patients with multiple sclerosis (MS). In this double-blind crossover design, patients received either TENS or sham stimulation for 3weeks (1h per day) on the median nerve region of the most impaired hand, followed by the other stimulation condition after a washout period of 6months. Cortical motor representation was mapped using transcranial magnetic stimulation (TMS) at the baseline and after the 3-week stimulation protocol. Our results revealed that 3weeks of daily stimulation with TENS significantly decreased the cortical motor representation of the stimulated muscle in MS patients. Although the mechanisms underlying this decrease remain unclear, our findings indicate that TENS has the ability to induce long-term reorganization in the motor cortex of MS patients.

AB0836-HPR Functional reeducation program associated with back school improve functional disability and pain in workers with chronic low back pain: a pilot study

BackgroundChronic Low Back Pain (CLBP) may present functional disability that compromises the performance of functional activities (1). Treatment of CLBP patients has been relying on generic exercises such as stretching...

Reliability in the location of hindlimb motor representations in Fischer-344 rats

The purpose of the present study was to determine the feasibility of using a common laboratory rat strain for reliably locating cortical motor representations of the hindlimb. Intracortical microstimulation techniques were used to derive detailed maps of the hindlimb motor representations in 6 adult Fischer-344 rats. The organization of the hindlimb movement representation, while variable across individual rats in topographic detail, displayed several commonalities. The hindlimb representation was positioned posterior to the forelimb motor representation and posterolateral to the motor trunk representation. The areal extent of the hindlimb representation across the cortical surface averaged 2.00 ± 0.50 mm(2). Superimposing individual maps revealed an overlapping area measuring 0.35 mm(2), indicating that the location of the hindlimb representation can be predicted reliably based on stereotactic coordinates. Across the sample of rats, the hindlimb representation was found 1.25-3.75 mm posterior to the bregma, with an average center location approximately 2.6 mm posterior to the bregma. Likewise, the hindlimb representation was found 1-3.25 mm lateral to the midline, with an average center location approximately 2 mm lateral to the midline. The location of the cortical hindlimb motor representation in Fischer-344 rats can be reliably located based on its stereotactic position posterior to the bregma and lateral to the longitudinal skull suture at midline. The ability to accurately predict the cortical localization of functional hindlimb territories in a rodent model is important, as such animal models are being increasingly used in the development of brain-computer interfaces for restoration of function after spinal cord injury.

H‐coil repetitive transcranial magnetic stimulation for pain relief in patients with diabetic neuropathy

Painful neuropathy is associated with plasticity changes in the nervous system. Standard repetitive transcranial magnetic stimulation (rTMS) is a non-invasive technique used to study changes in cortical excitability and to inhibit pain perception. Deep rTMS is a newer development that allows direct activation of deeper neuronal populations, by a unique coil design termed the H-coil. This study was designed to assess whether deep rTMS applied over the motor cortical lower-limb representation relieves pain in patients with diabetic neuropathy. Patients were randomly assigned to receive daily real or sham H-coil rTMS for 5 consecutive days. After a 5-week washout period, they crossed over to the alternative treatment for additional 5 days (according to a crossover study design). Outcome measures were changes in the visual analogue scale (VAS) for pain and in area and threshold of RIII nociceptive flexion reflex (RIII reflex). Of the 25 patients randomized, 23 completed the study. After real rTMS, the VAS scores decreased significantly (p=0.01), and so did RIII reflex area (p<0.01), while no significant effects in these variables were induced by the sham rTMS treatment. The rTMS-induced changes in the outcome measures disappeared about 3 weeks after stimulation. All patients tolerated stimulation well. Deep H-coil rTMS provides pain relief in patients with diabetic neuropathy. This innovative technique can induce a therapeutic effect on brain areas that otherwise remain difficult to target. rTMS may produce its analgesic effects, inducing motor cortex plasticity and activating descending inhibitory pain control systems.

Functional Topography of Cortical Thumb Movement Representations in Human Primary Motor Cortex

Intracortical microstimulation and single cell recordings in non-human primates showed that both, muscles and movements are represented in primary motor cortex (M1). This was also suggested in humans using electrophysiological and neuroimaging techniques. Transcranial magnetic stimulation (TMS) thus far was used to study motor cortical muscle representations, but data on movement representations in man are scarce. Therefore, we used TMS of M1 to evoke directional (flexion, extension) thumb movements and generated topographic movement maps (TMS-mov-map) in fourteen healthy individuals. TMS-mov-maps were related to the individual voluntary motor output (Vol-mov, directional thumb acceleration). Conventional TMS-motor evoked potentials (TMS-MEP) were simultaneously recorded from the prime movers (flexor and extensor pollicis brevis) and also compared to the voluntary motor output. Remarkably stable topographic maps were generated with both, thumb flexion and extension being multiply represented, overlapped and interspersed. Thumb Flexion was usually more robust than extension. TMS-mov-maps rather than TMS-MEP-maps seemed to better reflect the individual voluntary motor output. The findings emphasize existing models of multiple, overlapping finger movement representations in human M1, and indicate that this model also adapts to antagonistic thumb movements. The results suggest that investigating topographic movement maps may be an interesting adjunct in studying human motor cortical topography and its relevance for motor control.

Simultaneous rTMS and Piano Playing Improve Hand Dexterity and Cortical Motor Representation in a Professional Pianist with Multiple Sclerosis: A Case Report (P07.189)

Simultaneous rTMS and Piano Playing Improve Hand Dexterity and Cortical Motor Representation in a Professional Pianist with Multiple Sclerosis: A Case Report (P07.189)

Simultaneous rTMS and Piano Playing Improve Hand Dexterity and Cortical Motor Representation in a Professional Pianist with Multiple Sclerosis: A Case Report (IN11-1.010)

Simultaneous rTMS and Piano Playing Improve Hand Dexterity and Cortical Motor Representation in a Professional Pianist with Multiple Sclerosis: A Case Report (IN11-1.010)

Effect of transcranial direct current stimulation on elbow flexor maximal voluntary isometric strength and endurance

The effects of transcranial direct current stimulation (tDCS) on maximal voluntary isometric contraction (MVC) strength and the time to failure (TTF) in an isometric (30% MVC) muscle endurance test of the elbow flexors were investigated. Fifteen men (mean age, 27.7 ± 8.4 years) were tested for MVC strength and TTF 2 times, separated by a 60-min rest. During the last 10 min of the rest period, 1 of 2 tDCS treatments or 1 sham intervention session was administered, in a randomized order, with 1 week between sessions. In the tDCS intervention, a 2 mA direct current was delivered for 10 min through an anode placed on the scalp, overlying the right motor cortical representation of the left arm; a cathode was secured over the right shoulder. In the sham intervention, the current was delivered for the first 30 s only. No significant differences between the first and second tDCS sessions were evident for MVC strength or TTF. For MVC strength (baseline, 66.0 ± 11.4 Nm), postintervention measures decreased by 5.9% ± 4.2% (p < 0.05), but no significant difference in the changes was evident between tDCS and sham sessions. TTF did not change significantly from preintervention (309.2 ± 91.6 s) to postintervention (327.2 ± 128.5 s), and there was no significant difference between interventions. It was concluded that the tDCS intervention did not affect muscle function, perhaps because of ceiling effects, in which the intervention does not enhance muscle function further when muscle function is already maximal.

Applicability of nTMS in locating the motor cortical representation areas in patients with epilepsy

Transcranial magnetic stimulation (TMS) is increasingly used for non-invasive functional mapping in preoperative evaluation for brain surgery, and the reliability of navigated TMS (nTMS) motor representation maps has been studied in the healthy population and in brain tumor patients. The lesions behind intractable epilepsy differ from typical brain tumors, ranging from developmental cortical malformations to injuries early in development, and may influence the functional organization of the cortical areas. Moreover, the interictal cortical epileptic activity and antiepileptic medication may affect the nTMS motor threshold. The reliability of the nTMS motor representation localization in epilepsy patients has not been addressed. We compared the nTMS motor cortical representation maps of hand and arm muscles with the results of invasive electrical cortical stimulation (ECS) in 13 patients with focal epilepsy. The nTMS maps were projected to the cortical surface segmented from preoperative magnetic resonance images (MRI), and the positions of the subdural electrodes were extracted from the postoperative low-dose computed tomography (CT) images registered with preoperative MRI. The 3D distance between the average nTMS site and average ECS electrode location was 11 ± 4 mm for the hand and 16 ± 7 mm for arm muscle representation areas. In all patients the representation areas defined with nTMS and ECS were located on the same gyrus, also in patients with abundant interictal epileptic activity on the motor gyrus. nTMS can reliably locate the hand motor cortical representation area with the accuracy needed for pre-surgical evaluation in patients with epilepsy.

Cortical motor activity and reorganization following upper-limb amputation and subsequent targeted reinnervation

Previous studies have postulated that the amount of brain reorganization following peripheral injuries may be correlated with negative symptoms or consequences. However, it is unknown whether restoring effective limb function may then be associated with further changes in the expression of this reorganization. Recently, targeted reinnervation (TR), a surgical technique that restores a direct neural connection from amputated sensorimotor nerves to new peripheral targets such as muscle, has been successfully applied to upper-limb amputees. It has been shown to be effective in restoring both peripheral motor and sensory functions via the reinnervated nerves as soon as a few months after the surgery. However, it was unclear whether TR could also restore normal cortical motor representations for control of the missing limb. To answer this question, we used high-density electroencephalography (EEG) to localize cortical activity related to cued motor tasks generated by the intact and missing limb. Using a case study of 3 upper-limb amputees, 2 of whom went through pre and post-TR experiments, we present unique quantitative evidence for the re-mapping of motor representations for the missing limb closer to their original locations following TR. This provides evidence that an effective restoration of peripheral function from TR can be linked to the return of more normal cortical expression for the missing limb. Therefore, cortical mapping may be used as a potential guide for monitoring rehabilitation following peripheral injuries.

Posture-related modulations in motor cortical excitability of the proximal and distal arm muscles

The effect of postural orientation on the motor corticospinal excitability (MCE) of proximal and distal upper extremity (UE) muscles was investigated. In a crossover design, recruitment curves (RCs), short interval cortical inhibition (SICI) and intracortical facilitation (ICF) of resting anterior deltoid (AD) and first dorsal interosseus (FDI) was assessed in two postures: sitting and standing. Six healthy adults without contraindications to transcranial magnetic stimulation (TMS) participated in the study. TMS was applied over the motor cortical representation of FDI and AD at intensities ranging from 90% to 200% of resting motor threshold (RMT) in increments of 10%. SICI and ICF were assessed for each muscle using a conditioning stimulus (80% RMT) preceding a test stimulus (120% RMT) with an interstimulus interval of 2ms and 15ms, respectively. For AD, but not FDI, there was a significant and consistent increase in RC slope during standing compared to sitting. For FDI, there was no difference in ICF and SICI between sitting and standing. However, for AD, while there was no difference in ICF between the two postures, there was a clear trend for SICI to decrease (p=0.06) in standing compared to sitting. These results indicate that postural change from sitting to standing, affects the MCE of proximal but not distal muscles. While this indicates the role of proximal UE muscles in postural control, it also implies that rehabilitation protocols for enhancing proximal arm motor function may be advantaged if administered in a standing posture.

Hand Dominance and Age Have Interactive Effects on Motor Cortical Representations

Older adults exhibit more bilateral motor cortical activity during unimanual task performance than young adults. Interestingly, a similar pattern is seen in young adults with reduced hand dominance. However, older adults report stronger hand dominance than young adults, making it unclear how handedness is manifested in the aging motor cortex. Here, we investigated age differences in the relationships between handedness, motor cortical organization, and interhemispheric communication speed. We hypothesized that relationships between these variables would differ for young and older adults, consistent with our recent proposal of an age-related shift in interhemispheric interactions. We mapped motor cortical representations of the right and left first dorsal interosseous muscles using transcranial magnetic stimulation (TMS) in young and older adults recruited to represent a broad range of the handedness spectrum. We also measured interhemispheric communication speed and bimanual coordination. We observed that more strongly handed older adults exhibited more ipsilateral motor activity in response to TMS; this effect was not present in young adults. Furthermore, we found opposing relationships between interhemispheric communication speed and bimanual performance in the two age groups. Thus, handedness manifests itself differently in the motor cortices of young and older adults and has interactive effects with age.

Skill learning induced plasticity of motor cortical representations is time and age-dependent

Movement representations in the motor cortex can reorganize to support motor skill learning during young adulthood. However, little is known about how motor representations change during aging or whether their change is influenced by continued practice of a skill after it is learned. We used intracortical microstimulation to characterize the organization of the forelimb motor cortex in young and aged C57/BL6 mice after short (2–4weeks) or long (8weeks) durations of training on a skilled reaching task or control procedures. In young mice, a short duration of reach training increased the area of proximal forelimb movement representations at the expense of distal representations. Following a longer training duration, ratios of proximal to distal movements returned to baseline, even with ongoing practice and skill maintenance. However, lingering changes were evident in thresholds for eliciting distal forelimb movements, which declined over the longer training period. In aged mice, movement representations and movement thresholds failed to change after either duration of training. Furthermore, there was an age-related loss of digit representations and performance decrements on other sensorimotor tests. Nevertheless, in quantitative measures of reaching success, aged mice learned and performed the skilled reaching task at least as well as younger mice. These results indicate that experience-driven topographical reorganization of motor cortex varies with age, as well as time, and is partially dissociable from behavioral performance. They also support an enduring capacity to learn new manual skills during aging, even as more youthful forms of cortical plasticity and sensorimotor function are lost.

Motor cortex evaluation by nTMS after surgery of central region tumors: a feasibility study

Largely discussed during the past decade, motor cortex reorganization in brain tumor surgery has been investigated only by few studies. We therefore aimed to investigate cortical motor representation after resection of perirolandic WHO grade II and III gliomas using navigated transcranial magnetic stimulation (nTMS). Five patients were examined before neurosurgery and after a follow-up period of 17.7 ± 6.8months. As a control, five healthy age-matched subjects were equally studied by nTMS in two sessions spaced 12.6 (range 2-35) days apart. Resting motor thresholds (RMT), hotspots and centers of gravity (CoG) were identified for the first dorsal interosseous (FDI), abductor pollicis brevis (APB), extensor digitorum (EXT), tibialis anterior (TA) and abductor hallucis (AH) muscles. Euclidian distances, coefficients of variance and intraclass correlation coefficients (ICC) were calculated. Healthy subjects showed moderate to excellent reliability measurement of RMT (ICC = 0.69-0.94). Average displacement of CoGs across sessions was 0.68 ± 0.34cm in the dominant and 0.76 ± 0.38cm in the non-dominant hemisphere; hotspots moved 0.87 ± 0.51cm and 0.83 ± 0.45cm, respectively. In one patient these parameters differed significantly from the control group (p < 0.05 for both CoGs and hotspots). Overall, all patients' CoGs moved 1.12 ± 0.93cm, and hotspots were 1.06 ± 0.7cm apart. In both patients and healthy subjects, movement of assessed parameters was more important along the X- than the Y-axis. nTMS allows evaluating cortical reorganization after brain tumor surgery. It may contribute to the understanding of neurofunctional dynamics, thus influencing therapeutic strategy.

Functional Plasticity of the Motor Cortical Structures Demonstrated by Navigated TMS in Two Patients with Epilepsy

BackgroundRecently, navigated transcranial magnetic stimulation (nTMS) has been suggested to be useful in preoperative functional localization of motor cortex in patients having tumors close to the somatomotor cortex. Resection of tumors in anatomically predicted eloquent areas without adverse effects have emphasized functional plasticity elicited by intracranial pathology. ObjectiveTo describe functional plasticity of motor cortex indicated by nTMS in two patients with epilepsy. MethodsnTMS, functional MRI (fMRI), diffusion-tensor (DT)-tractography and magnetoencephalography (MEG) were utilized to preoperatively localize motor cortical areas in the workup for epilepsy surgery. The localizations were compared with each other, with the cortical anatomical landmarks, and in one patient with invasive electrical cortical stimulation (ECS). ResultsIn two out of 19 studied patients, nTMS identified motor cortical sites that differed from those indicated by anatomical landmarks. In one patient, nTMS activated preferentially premotor cortex rather than pathways originating from the precentral gyrus. MEG and fMRI localizations conformed with nTMS whereas ECS localized finger motor function into the precentral gyrus. Resection of the area producing motor responses in biphasic nTMS did not produce a motor deficit. In the other patient, nTMS indicated abnormal ipsilateral hand motor cortex localization and confirmed the functionality of aberrant motor cortical representations of the left foot also indicated by fMRI and DT-tractography. ConclusionnTMS may reveal the functional plasticity and shifts of motor cortical function. Epileptic foci may modify cortical inhibition and the nTMS results. Therefore, in some patients with epilepsy, the nTMS results need to be interpreted with caution with regard to surgical planning.

Representation of cricothyroid muscles at the primary motor cortex (M1) in healthy subjects, mapped by navigated transcranial magnetic stimulation (nTMS)

ObjectivesTo establish a methodology for mapping of primary motor cortex (M1) for cricothyroid (CTHY) muscles in a group of healthy subjects using three-dimensional (3D) magnetic resonance imaging (MRI) navigated transcranial magnetic stimulation (nTMS). MethodsTwo independent measurements were performed. Twelve right-handed healthy subjects were included in the study. In the first measurement, mapping of the abductor pollicis brevis (APB) muscle was followed by mapping of the M1 for CTHY. This was performed in 11 subjects. Second, to avoid bias concerning using a hand knob as a landmark, mapping of M1 for CTHY muscle was followed by mapping of M1 for APB. This was performed in three healthy subjects.The nTMS was used, with selective recordings of motor evoked potentials (MEPs) from APB muscle and corticobulbar motor evoked potentials (CoMEPs) from the CTHY muscle. For recording the responses from the CTHY muscle two hook wire electrodes (the size of 76 μm of diametre passing through 27 gauge needle) were inserted in the muscle. For the recording of MEPs from APB muscle, surface electrodes were used. ResultsFirst measurement: Stimulation over the left M1 for APB muscles elicits MEPs in the contralateral APB muscle with a mean latency of 22.8±1.69ms. Stimulation over the left M1 for the CTHY muscle elicits CoMEPs in the contralateral CTHY muscle with a mean latency of 11.89±1.26ms. The distance between the cortical representation for APB and CTHY was 25.19±6.52mm, with CTHY muscle representation lateral to the APB muscle.Second measurement: The results of second measurement of the distance between M1 for CTHY and M1 for APB and their cortical localisation were comparable to the results of the first measurement. ConclusionThis is the first study with the aim to determine the exact cortical localisation of CTHY muscle with nTMS. Mapping of M1 for CTHY and APB muscles by nTMS was successfully performed in all healthy subjects. The exact location of the stimulating points over M1 muscles eliciting responses in CTHY and APB muscles was determined and superimposed over 3D MRI images. The data show that M1 for CTHY muscle is about 25mm more lateral with regard to M1 for the APB muscle. SignificanceMapping of M1 for CTHY muscle might represent an important neurophysiologic marker for facilitating preoperative mapping of motor speech-related cortical areas due to the proximity of motor cortical representation for laryngeal muscles and opercular part of the Broca area.

Distinct EEG Amplitude Suppression to Facial Gestures as Evidence for a Mirror Mechanism in Newborn Monkeys

At birth, human infants and newborns of other primate species demonstrate the capacity to attend and to respond to facial stimuli provided by a caregiver. Newborn infants are also capable of exhibiting a range of facial expressions. Identification of the neural underpinnings of these capacities represents a formidable challenge in understanding social development. One possible neuronal substrate is the mirror-neuron system assumed to activate shared motor cortical representations for both observation and production of actions. We tested this hypothesis by recording scalp EEG from 1- to 7-day-old newborn rhesus macaques who were observing and producing facial gestures. We found that 5-6 Hz EEG activity was suppressed both when the infants produced facial gestures and while they were observing facial gestures of a human experimenter, but not when they were observing nonbiological stimuli. These findings demonstrate the presence of neural reactivity for biological, communicatively relevant stimuli, which may be a likely signature of neuronal mirroring. The basic elements of the mirror-neuron system appear to operate from the very first days of life and contribute to the encoding of socially relevant stimuli.

Controlled movement processing: Evidence for a common inhibitory control of finger, wrist, and arm movements

We used the behavioral task and theoretical construct of the countermanding paradigm to test whether there is any difference between the inhibitory control of the finger, wrist, and arm. Participants were instructed (primary task) to respond to a directional go signal presented at the fovea by pressing a button with either their index or middle fingers, moving a joystick with their wrists, or reaching to a stimulus on a touch screen with their arms. They were also instructed (secondary task) to withhold their responses when a stop signal was presented on 25% of trials. The participants’ ability to inhibit each of the commanded movements was captured by their inhibition probability function, which describes how withholding is increasingly difficult as the delay between the go and stop signals increased. By modeling each participant’s inhibition function, we estimated that the time needed to inhibit a commanded movement was about 240ms, a variable that did not differ significantly between the three limb segments. Moreover, we found that the best-fit model of each segment’s inhibition function could fit equally well the inhibition functions obtained with the other two segments. These results provide evidence that the upper limb segments share a common inhibitory control, which may facilitate the regulation of neuronal activity within the distributed motor cortical representations and thus simplify the voluntary control of multi-segmental movements.