Noninvasive mapping of human motor cortex

Human motor cortex was stimulated using brief, high-voltage electrical stimulation. Constant-voltage stimuli were delivered through a bipolar surface stimulator with the anode placed at multiple positions on the scalp and the cathode situated 2.5 cm anterior to the anode. Recordings were bilateral from the abductor pollicis brevis, tibialis anterior, and risorius. We averaged the amplitudes of three muscle responses obtained from stimulation of each scalp position and assigned the resultant value to that position. The findings in eight normal volunteers were similar and reproducible. The maximal responses of the right hand were obtained when stimulating over C3, of the left hand when stimulating over C4, of the right and left legs when stimulating over Cz, and of the right side of the mouth when stimulating over T3.     

Reliability of motor cortex transcranial magnetic stimulation in four muscle representations.

OBJECTIVE: Motor cortex plasticity may underlie motor recovery after stroke. Numerous studies have used transcranial magnetic stimulation (TMS) to investigate motor system plasticity. However, research on the reliability of TMS measures of motor cortex organization and excitability is limited. We sought to test the reliability of these TMS measurements. METHODS: Twenty healthy volunteers were tested twice over a two-week period using TMS to determine motor threshold, map topography, and stimulus-response curves for first dorsal interosseous (FDI), abductor pollicis brevis (APB), extensor digitorum communis (EDC), and flexor carpi radialis (FCR) muscles. RESULTS: We found moderate to good test-retest reliability TMS measurements of motor threshold (ICC=0.90-0.97), map area (ICC=0.63-0.86) and location (ICC=0.69-0.86), and stimulus-response curves (ICC=0.60-0.83). CONCLUSIONS: TMS assessments of motor representation size, location, and excitability are generally reliable measures, although their reliability may vary according to the muscle under investigation. SIGNIFICANCE: These results suggest that TMS measurements of motor cortex function are reliable enough to be potentially useful in investigation of motor system plasticity.     

Functional somatotopy of finger representations in human primary motor cortex

To assess the degree of fine-scale somatotopy within the hand area of the human primary motor cortex (M1), functional mapping of individual movements of all fingers was performed in healthy young subjects (n = 7) using MRI at 0.8 x 0.8 mm2 resolution and 4 mm section thickness. The experimental design comprised both a direct paradigm contrasting single digit movements vs. motor rest and multiple differential paradigms contrasting single digit movements vs. the movement of another digit. Direct mapping resulted in largely overlapping activations. A somatotopic arrangement was only recognizable when considering the mean center-of-mass coordinates of individual digit representations averaged across subjects. In contrast, differential paradigms revealed more segregated and somatotopically ordered activations in single subjects. The use of center-of-mass coordinates yielded inter-digit distances ranging from 2.0 to 16.8 mm, which reached statistical significance for pairs of more distant digits. For the middle fingers, the functional somatotopy obtained by differential mapping was dependent on the choice of the digit used for control. These results confirm previous concepts that finger somatotopy in the human M1 hand area emerges as a functional predominance of individual digit representations sharing common areas in a distributed though ordered network.     

The comparable size and overlapping nature of upper limb distal and proximal muscle representations in the human motor cortex

The purpose of this study was to determine the relative size and location of proximal and distal upper limb muscle representations in the human motor cortex. Motor-evoked potentials (MEPs) evoked by transcranial magnetic stimulation were recorded in the proximal muscle anterior deltoid (AD) and in the distal muscles extensor carpi radialis (ECR) and first dorsal interosseus (1DI). The coil was moved in steps of 1 cm along a grid drawn on a tight-fitting polyester cap placed on the subject's head. At each location, four stimuli were delivered at 1.2 times the active motor threshold (AMT), and MEPs averaged in real-time. The peak-to-peak amplitude of each muscle's mean MEP was measured at each stimulation site. The area of a muscle's representation was measured by a pixel-counting algorithm. The optimal point of each muscle's areal representation, which corresponds to the locus near which the largest MEPs are obtained, was determined by fitting a 3D Lorentzian function to the data points. The optimal point of distal muscles tended to be situated more laterally along the motor strip than that of proximal muscles. However, there was no statistically significant difference between the size of the areal representations and they overlapped considerably. Additionally, in another five subjects, using a small 45-mm coil placed in a hyper-focal orientation, maps were obtained at a stimulus intensity of 1.1-1.15 times the AMT of the muscle with the lowest threshold, usually the 1DI. Even in this very stringent condition, the mapped representations of the AD, ECR and 1DI overlapped, notwithstanding that sharp demarcations between borders were also apparent. These observations demonstrate that stimulus spread alone does not explain the overlap of muscle representations. These results show that commonly used proximal and distal upper-limb muscles, taken individually, are controlled by motor cortical territories of approximately equal size that significantly overlap despite differences in the location of their optimal points.     

Noninvasive extraction of microsecond‐scale dynamics from human motor cortex

State‐of‐the‐art noninvasive electromagnetic recording techniques allow observing neuronal dynamics down to the millisecond scale. Direct measurement of faster events has been limited to in vitro or invasive recordings. To overcome this limitation, we introduce a new paradigm for transcranial magnetic stimulation. We adjusted the stimulation waveform on the microsecond scale, by varying the duration between the positive and negative phase of the induced electric field, and studied corresponding changes in the elicited motor responses. The magnitude of the electric field needed for given motor‐evoked potential amplitude decreased exponentially as a function of this duration with a time constant of 17 µs. Our indirect noninvasive measurement paradigm allows studying neuronal kinetics on the microsecond scale in vivo.     

Interhemispheric differences of hand muscle representation in human motor cortex

Focal magnetic transcranial stimulation (TCS) is employed for mapping of the motor cortical output to abductor digiti minimi (ADM) muscle. The aim of this study was to evaluate the interhemispheric asymmetries in normals. Motor maps were obtained through motor evoked potentials (MEPs) recordings from ADM muscle in 20 healthy subjects in right and left hemispheres TCS. Measurement of several indexes such as excitability threshold, MEPs amplitude, MEPs latency, and silent period duration did not show differences between the hemispheres. Moreover, no interhemispheric asymmetries were found when the amplitude ratio values were analyzed. The hand motor cortical area, as represented by the number of responsive sites (3.6 vs. 3.5) and the “hot spot” site localization presented a fairly symmetrical organization. Absolute values displayed a relatively wide intersubject variability, while their interhemispheric differences were extremely restricted. This observation can offer a new tool in diagnosing and following up neurological disorders affecting the central motor system, mainly for those concerning monohemispheric lesions. © 1997 John Wiley & Sons, Inc. Muscle Nerve, 20, 535–542, 1997.     

Rapid mapping of finger representations in human primary somatosensory cortex applying neuromagnetic steady-state responses

We analyzed somatosensory evoked steady-state fields in order to localize finger representations in the hand area of the primary somatosensory cortex (S1). Using a 122-channel whole-head neuromagnetometer we recorded in six healthy subjects neuromagnetic responses to high frequency electrical stimuli delivered simultaneously to digit I, II, III and V at 22, 24, 27 and 30 Hz, respectively, and to transient stimulation of each single digit with a frequency of 3 Hz. Responses were averaged separately for each digit and were modeled by single equivalent current dipoles. Both conditions yielded the typical somatotopic finger representations within S1 hand area. Dipole locations did not differ significantly between the transient and the steady-state stimulation. Therefore, simultaneous high-frequency stimulation of the digits seems to be a reliable method for rapid and detailed mapping of the S1 hand area. This procedure has potential advantages over recording of transient responses. With simultaneous steady-state stimulation the measurement times are reduced to 2 min for mapping the whole hand area. Because of this our method probably increases spatial accuracy and permits repeated short interval recordings, e.g. in experiments studying short term plasticity.     

Training-induced changes of motor cortex representations in stroke patients

OBJECTIVE: To study changes in motor cortex representations after a single session of physiotherapy in stroke patients. METHODS: TMS mapping was used to evaluate the motor output map of the abductor pollicis brevis (APB) in both hemispheres. Stroke patients (4-8 weeks after the infarction) were studied prior to a training session aimed at improving dexterity and 1 h and 1 day after the training. RESULTS: Prior to the training, the APB representation area in the affected hemisphere was significantly smaller than on the non-affected side. After therapy, the cortical motor output to the paretic APB was significantly enlarged, and motor function was improved. One day later, these effects were partially reversed. Motor thresholds remained significantly increased in the affected hemisphere before and after the therapy. CONCLUSION: A single session of physiotherapy produces a use-dependent enlargement of motor cortex representations paralleled by an improvement of motor function in stroke patients.     

Analogous corticocortical inhibition and facilitation in ipsilateral and contralateral human motor cortex representations of the tongue.

How the human brain controls activation of the ipsilateral part of midline muscles is unknown. We studied corticospinal and corticocortical network excitability of both ipsilateral and contralateral motor representations of the tongue to determine whether they are under analogous or disparate inhibitory and facilitatory corticocortical control. Motor evoked potentials (MEPs) to unilateral focal transcranial magnetic stimulation (TMS) of the tongue primary motor cortex were recorded simultaneously from the ipsilateral and contralateral lingual muscles. Single-pulse TMS was used to assess motor threshold (MT) and MEP recruitment. Paired-pulse TMS was used to study intracortical inhibition (ICI) and intracortical facilitation (ICF) at various interstimulus intervals (ISIs) between the conditioning stimulus (CS) and the test stimulus (TS), and at different CS and TS intensities, respectively. Focal TMS invariably produced MEPs in both ipsilateral and contralateral lingual muscles. MT was lower and MEP recruitment was steeper when recorded from the contralateral muscle group. ICI and ICF were identical in the ipsilateral and contralateral representations, with inhibition occurring at short ISIs (2 and 3 ms) and facilitation occurring at longer ISIs (10 and 15 ms). Moreover, changing one stimulus parameter regularly produced analogous changes in MEP size bilaterally, revealing strong linear correlations between ipsilateral and contralateral ICI and ICF (P < 0.0001). These findings indicate that the ipsilateral and contralateral representations of the tongue are under analogous inhibitory and facilitatory control, possibly by a common intracortical network.     

Methodology for non-invasive mapping of human motor cortex with electrical stimulation

Different techniques for mapping motor evoked potentials recorded from hand, upper arm, leg and mouth were analyzed. The best results were obtained when: (1) delivering constant voltage stimuli through a bipolar surface stimulator, (2) positioning the anode over the desired scalp location and the cathode 2.5 cm anterior to the anode, (3) maintaining low impedances, and (4) increasing the stimulus intensity over the theoretical motor representation area until a 500-1000 microV muscle response is achieved and then delivering the same stimulus over different scalp locations. This technique allows bilateral mapping of the different body part representations in 1-2 h. Areas so identified are small, clearly separate from each other and from the corresponding somatosensory areas.     

皮质脑电图小波分析定位运动区的探索性研究

目的研究术中皮质脑电图(ECoG)小波分析对运动区皮质定位的可行性。方法利用小波变换,对ECoG信号进行多层分解和重构,提取4个主要频带(δ、θ、μ和β)重构信号的运动前后能量比(ERD)为特征量,并构造特定阈值进行分类,然后与相应手指弯曲运动数据进行对照比较,分析检测的正确率。结果d6子频带(μ频带)的运动前后ERD变化最明显;以40%为阈值进行分类,其定位运动区的正确率达到93%。结论通过小波分析对ECoG的特征进行提取和分类,可有效定位运动区皮质。     

Overlapping representations of the neck and whiskers in the rat motor cortex revealed by mapping at different anaesthetic depths

The primary motor cortex of mammals has an orderly representation of different body parts. Within the representation of each body part the organization is more complex, with groups of neurons representing movements of a muscle or a group of muscles. In rats, uncertainties continue to exist regarding organization of the primary motor cortex in the whisker and the neck region. Using intracortical microstimulation (ICMS) we show that movements evoked in the whisker and the neck region of the rat motor cortex are highly sensitive to the depth of anaesthesia. At light anaesthetic depth, whisker movements are readily evoked from a large medial region of the motor cortex. Lateral to this is a small region where movements of the neck are evoked. However, in animals under deep anaesthesia whisker movements cannot be evoked. Instead, neck movements are evoked from this region. The neck movement region thus becomes greatly expanded. An analysis of the threshold currents required to evoke movements at different anaesthetic depths reveals that the caudal portion of the whisker region has dual representation, of both the whisker and the neck movements. The results also underline the importance of carefully controlling the depth of anaesthesia during ICMS experiments.     

Task-free electrocorticography frequency mapping of the motor cortex

ObjectiveElectrocortical stimulation mapping (ESM) is the current gold standard for functional mapping of the eloquent cortex prior to epilepsy surgery. The procedure is, however, time-consuming and quite demanding for patients. Electrocorticography frequency mapping (ECoG mapping) has been suggested as an adjunct method. Here, we investigated whether it is possible to perform mapping of motor regions using ECoG data of spontaneous movements.MethodsUsing the video registration of seven epilepsy patients who underwent electrocorticography and ESM, we selected periods of spontaneous hand and arm movements and periods of rest. Frequency analysis was performed, and electrodes showing a significant change in power (4–7, 8–14, 15–25, 26–45 or 65–95 Hz) were compared with those being identified as relevant for hand and/or arm movement by ESM.ResultsAll frequency bands showed a high specificity (>0.80), and the 65–95 Hz frequency band additionally had a high sensitivity (0.82) for identifying ESM positive electrodes.ConclusionsOur data show a good match between ECoG mapping of spontaneous movements and ESM data.SignificanceThe accurate match suggests that ECoG mapping of the motor cortex using spontaneous movements may be a valuable complement to ESM, especially when other options requiring patient cooperation fail.     

Stereotactic functional mapping of the cat motor cortex

Epidural motor cortex stimulation is an increasingly used method to control refractory neuropathic pain although its mechanisms of action remain poorly understood. Animal models are currently developed that allow reproducing the conditions of this neurosurgical approach and clarifying its mechanisms. In this study we validate a new stereotactic functional map of the cat motor cortex carried out in epidural conditions, thus allowing future experimentations that closely mimic the technique used in humans.     

Topographic mapping of the human motor cortex with magnetic stimulation: factors affecting accuracy and reproducibility.

We recorded motor evoked potentials (MEPs) from deltoid, biceps brachii, abductor pollicis brevis and flexor carpi radialis muscles of 5 normal volunteers during transcranial magnetic stimulation. With the subjects at rest, an 8-shaped magnetic coil was used to deliver 30 stimuli to different scalp positions 0.5 or 1.0 cm apart. The variability in amplitude and latency of MEPs was studied as a function of the scalp position stimulated, the number of stimuli at each position, and the percentage of maximal peripheral M responses (%M) elicited. The results were used to estimate the optimal number of stimuli at each position and the optimal spacing of scalp positions for topographic mapping of the human motor cortex. The amplitude and latency variability of MEPs were higher when suboptimal scalp positions were stimulated. Consequently, a larger number of stimuli were required to determine representative MEP amplitudes at suboptimal positions. In addition, there was an inverse relationship between %M recruited by transcranial magnetic stimuli in different subjects and the variability in MEP amplitude and latency. Latency variability was less pronounced than amplitude variability. Optimal sampling conditions are required to produce the best topographic maps, particularly to show subtle reorganization patterns in the human motor cortex.     

Electroencephalographic measurement of motor cortex control of muscle activity in humans.

OBJECTIVE: To detect and measure correlation between cortical and muscle activities, coherence analysis was used. METHODS: The electroencephalogram (EEG) and electromyogram (EMG) were recorded in 9 normal volunteers during tonic contraction of upper and lower limb muscles on the right side. Coherence between EEG and EMG was computed to analyze their linear association. RESULTS: EEG over the contralateral sensorimotor area was coherent with EMG, with peak coherence at 11-36 Hz (mean, 22 Hz). For the abductor pollicis brevis (APB) muscle, peak coherence, as determined by functional brain mapping with focal transcranial magnetic stimulation (TMS), was over or slightly posterior to the hand area on the primary motor cortex determined by focal transcranial magnetic stimulation (TMS). Peak coherence over the scalp was somatotopically organized. The temporal relation between EEG and EMG was analyzed with a new model for interpreting the phase shift ('constant phase shift plus constant time lag' model). For the APB muscle, the phase relation between cortical and muscular oscillations differed in the frequency ranges of 3-13 Hz and 14-50 Hz, respectively, suggesting that different coupling mechanisms operate in different bands. Only the phase shift between cortical and motoneuronal firing at 14-50 Hz was reliably estimated by a linear model. At 14-50 Hz, motoneuronal firing was led by surface-negative cortical activity with a constant time lag that depended on the cortical-muscular distance. For the APB muscle, the time lag was slightly shorter than the cortical-muscular conduction time determined by TMS. Vibratory stimulation (100 Hz) of a muscle tendon during tonic contraction had no significant effect on cortical-muscular coherence, indicating that cortical oscillation reflected motor rather than sensory activity. CONCLUSIONS: The present findings suggest temporal coding of the oscillatory motor control system (3-13 Hz vs. 14-50 Hz), and confirm the functional importance of cortical beta and gamma rhythms in the motor efferent command. Cortical-muscular synchronization is most likely mediated by the direct corticospinal pathway within the frequency range of 14-50 Hz.     

Functional metabolic mapping during forelimb movement in rat. I. Stimulation of motor cortex

The quantitative 14C-deoxyglucose (DG) autoradiographic technique has been used to study changes in cerebral metabolism during forelimb movements induced by graded stimulation of motor cortex. Experiments were directed at studying basic physiologic and anatomic aspects of the metabolic changes. Single shocks caused movement without metabolic change, whereas low-frequency trains caused seizures. Repetitive high-frequency train stimuli of short duration (500 Hz for 20 msec) caused jerk movements coupled with DG uptake in pathways. With stimulation of the forelimb motor zone at frequencies of 15-30/min there was prominent activation of cortical columns and strips in ipsilateral SI, SII, and MII, and contralateral MI and SI. Higher frequencies (120/min) were required to cause significant changes in DG in subcortical circuits. The most prominent changes occurred within a longitudinal corridor in dorsal thalamus and a ventral corridor in second-order sites in basal ganglia. Metabolic activation also occurred in contralateral cerebellum, the cuneate nucleus, and dorsal horn of the cervical spinal cord. Changes in these latter two sites were largely eliminated by removing feedback sensory activity. Stimulation of the forelimb sensory zone activated different sites in caudatoputamen and thalamus but similar zones in midbrain and cerebellum. The magnitude of the metabolic response in distant sites depended on the frequency of cortical stimulation. Different frequency-response relationships in different sites seemed to reflect the nature of the cortical input as well as differential effects of anesthesia. The pattern of the metabolic response was studied by comparing sites of activation with sites of the anatomic projections from motor and sensory cortical zones. 3H- and 14C-labeled amino acids were used to map the site and relative strength of pathways. Results revealed good correlation between the site of anatomic projection and the site of DG uptake but no consistent relationship between the relative strength of a projection and the magnitude of metabolic change within its field. Changes in glucose utilization with metabolic mapping experiments depend on the nature, strength, and frequency of stimulation; the site and nature of anatomic projection; the effects of anesthesia; and the strength of sensory feedback associated with the induced behavior.     

The human stretch reflex and the motor cortex.

The spinal stretch reflex, exemplified by the tendon jerk, appears to be less important in humans than a delayed 'long-latency' response. This is easily observed when muscles of the hand are stretched while they are already contracting voluntarily. On limited evidence, many have long held that the delayed response is a transcortical reflex and have tended to neglect alternative possibilities, particularly that it might be a spinal reflex dependent upon slow afferents. New experiments have now eliminated the alternatives, leaving the transcortical hypothesis in command of the field.