Stephan Salenius

Involvement of Primary Motor Cortex in Motor Imagery: A Neuromagnetic Study

Functional brain imaging studies have indicated that several cortical and subcortical areas active during actual motor performance are also active during imagination or mental rehearsal of movements. Recent evidence shows that the primary motor cortex may also be involved in motor imagery. Using whole-scalp magnetoencephalography, we monitored spontaneous and evoked activity of the somatomotor cortex after right median nerve stimuli in seven healthy right-handed subjects while they kinesthetically imagined or actually executed continuous finger movements. Manipulatory finger movements abolished the poststimulus 20-Hz activity of the motor cortex and markedly affected the somatosensory evoked response. Imagination of manipulatory finger movements attenuated the 20-Hz activity by 27% with respect to the rest level but had no effect on the somatosensory response. Slight constant stretching of the fingers suppressed the 20-Hz activity less than motor imagery. The smallest possible, kinesthetically just perceivable finger movements resulted in slightly stronger attenuation of 20-Hz activity than motor imagery did. The effects were observed in both hemispheres but predominantly contralateral to the performing hand. The attempt to execute manipulatory finger movements under experimentally induced ischemia causing paralysis of the hand also strongly suppressed 20-Hz activity but did not affect the somatosensory evoked response. The results indicate that the primary motor cortex is involved in motor imagery. Both imaginative and executive motor tasks appear to utilize the cortical circuitry generating the somatomotor 20-Hz signal.

Cortico‐muscular synchronization during isometric muscle contraction in humans as revealed by magnetoencephalography

1 Magnetoencephalographic (MEG) and electromyographic (EMG) signals were recorded from six subjects during isometric contraction of four different muscles. 2 Cortical sources were located from the MEG signal which was averaged timelocked to the onset of motor unit potentials. A spatial filtering algorithm was used to estimate the source activity. Sources were found in the primary motor cortex (M1) contralateral to the contracted muscle. Significant coherence between rectified EMG and M1 activity was seen in the 20 Hz frequency range in all subjects. 3 Interactions between the motor cortex and spinal motoneuron pool were investigated by separately studying the non‐stationary phase and amplitude dynamics of M1 and EMG signals. 4 Delays between M1 and EMG signals, computed from their phase difference, were found to be in agreement with conduction times from the primary motor cortex to the respective muscle. The time‐dependent cortico‐muscular phase synchronization was found to be correlated with the time course of both M1 and EMG signals. 5 The findings demonstrate that the coupling between the primary motor cortex and motoneuron pool is at least partly due to phase synchronization of 20 Hz oscillations which varies over time. Furthermore, the consistent phase lag between M1 and EMG signals, compatible with conduction time between M1 and the respective muscle with the M1 activity preceding EMG activity, supports the conjecture that the motor cortex drives the motoneuron pool.

Task-dependent modulation of 15-30 Hz coherence between rectified EMGs from human hand and forearm muscles

1 Recent reports have shown task‐related changes in oscillatory activity in the 15‐30 Hz range in the sensorimotor cortex of human subjects and monkeys during skilled hand movements. In the monkey these oscillations have been shown to be coherent with oscillatory activity in the electromyographic activity of hand and forearm muscles. 2 In this study we investigated the modulation of oscillations in the electromyogram (EMG) of human volunteers during tasks requiring precision grip of two spring‐loaded levers. 3 Two tasks were investigated: in the ‘hold’ task, subjects were required to maintain a steady grip force (ca 2·1 N or 2·6 N) for 8 s. In the ‘ramp’ task, there was an initial hold period for 3 s (force ca 2·1 N) followed by a linear increase in grip force over a 2 s period. The task ended with a further steady hold for 3 s at the higher force level (ca 2·6 N). 4 Surface EMGs were recorded from five hand and forearm muscles in 12 subjects. The coherence of oscillatory activity was calculated between each muscle pair. Frequencies between 1 and 100 Hz were analysed. 5 Each subject showed a peak in the coherence spectra in the 15‐30 Hz bandwidth during the hold task. This coherence was absent during the initial movement of the levers. During the ramp task the coherence in the 15‐30 Hz range was also significantly reduced during the movement phase, and significantly increased during the second hold period, relative to the initial hold. 6 There was coherence between the simultaneously recorded magnetoencephalogram (MEG) and EMG during steady grip in the hold task; this coherence disappeared during the initial lever movement. Using a single equivalent current dipole source model, the coherent cortical activity was localized to the hand region of the contralateral motor cortex. This suggests that the EMG‐EMG coherence was, therefore, at least in part, of cortical origin. 7 The results are discussed in terms of a possible role for synchrony in the efficient recruitment of motor units during maintained grip.

Modulation of human cortical rolandic rhythms during natural sensorimotor tasks

We studied modulation of cortical neuromagnetic rhythms in association with left and right median nerve stimulation, during rest, finger movements, and passive tactile hand stimulation, in seven healthy, right-handed adults. In the rest condition, the amplitude of the rhythmic sensorimotor activity decreased immediately after the median nerve stimuli and increased above the prestimulus level within 0.4 s afterward, especially in the 7- to 25-Hz band. The rebound occurred 100-300 ms earlier for 20 (7-15)-than for 10 (15-25)-Hz activity. Suppressions and rebounds were strongest in the contralateral sensorimotor hand area for the 20-Hz, but not for the 10-Hz, activity. The maximum rebound was on average 22-34% stronger in the left than in the right hemisphere. Active exploration of objects abolished rebounds of both 10- and 20-Hz signals in the contralateral hemisphere and markedly diminished them ipsilaterally. Finger movements without touching an object and passive tactile stimulation produced a weaker effect. The sensorimotor rhythms thus show a characteristic suppression and subsequent rebound after electrical median nerve stimulation. The rebound is left-hemisphere dominant in right-handed subjects and its suppression reveals bilateral cortical activation during both motor tasks and passive tactile stimulation, especially for explorative finger movements.

Synchronous cortical oscillatory activity during motor action.

Oscillations of the motor cortex interact with similar activity of the spinal motoneuron pool in the 15-30 Hertz frequency range. Recent observations have demonstrated how this interaction affects the firing of single corticospinal neurons. The interaction, reflected as corticomuscular coherence, occurs for both distal and proximal muscles and it constitutes one connection in a larger web of oscillatory interactions, including several other motor areas in the cortex, thalamus, and cerebellum. New results cast light on the possible functional significance of this interaction. The rhythmic interaction may reveal interesting information in several motor disorders, including essential tremor, Parkinsons disease, myoclonus epilepsy, and mirror movements.

Human cortical 40 Hz rhythm is closely related to EMG rhythmicity

We recorded cortical neuromagnetic rhythms during self-paced index-finger movements from a subject previously reported to show prominent 40 Hz electroencephalographic activity during motor behavior. The 10 and 20 Hz components of the rolandic mu rhythm were bilaterally suppressed, whereas the contralateral 40 Hz (35-41 Hz) activity was slightly enhanced before both fast and slow movements and strongly enhanced during slow movements. The 40 Hz rhythm originated mainly in the hand motor cortex and was clearly correlated with the rhythmicity of the electromyogram from the extensor muscles, with a systematic time lag. In this subject motor preparation, and especially control of finger movements, may thus be associated with enhanced cortical rhythms near 40 Hz. The coherence of these rhythms with muscular firing patterns likely reflects communication between the sensorimotor cortex and the motor units.

Tactile information from the human hand reaches the ipsilateral primary somatosensory cortex

Neuromagnetic responses to median nerve stimulation were studied in six healthy right-handed subjects. In the rest condition, only the right and left median nerves were alternately stimulated at the wrists. In two other conditions, continuous superficial tactile stimulation was concurrently applied to either the left or right hand. Tactile stimulation of palm and fingers of one hand enhanced, in the ipsilateral primary somatosensory cortex (SI), responses to median nerve stimulation of the other hand. This effect was stronger in the left than the right SI. Our data provide evidence in humans for the access of cutaneous information from the hands to ipsilateral SI, probably via excitatory transcallosal pathways. This interhemispheric information transfer may represent a neurophysiological substrate of somatosensory fusion between the hands.

Task-dependent modulations of cortical oscillatory activity in human subjects during a bimanual precision grip task

Oscillations are a widespread feature of normal brain activity and have been reported at a variety of different frequencies in different neuronal systems. The demonstration that oscillatory activity is present in motor command signals has prompted renewed interest in the possible functions of synchronous oscillatory activity within the primate sensorimotor system. In the current study, we investigated task-dependent modulations in coupling between sensorimotor cortical oscillators during a bimanual precision grip task. The task required a hold-ramp-hold pattern of grip force to be exerted on a compliant object with the dominant right hand, while maintaining a steady grip with the nondominant hand. We found significant task-related modulation of 15- to 30-Hz coherence between magnetoencephalographic (MEG) activity recorded from the left sensorimotor cortex and electromyographic (EMG) activity in hand muscles on the right side. This coherence was maximal during steady hold, but disappeared during the ramp movements. Interestingly coherence between the right sensorimotor MEG and left-hand EMG showed a similar, although less deeply modulated, task-related pattern, even though this hand was maintaining a simple steady grip. No significant ipsilateral MEG-EMG coherence was observed in the 15- to 30-Hz passband for either hand. These results suggest that the cortical oscillators in the two sensorimotor cortices are independent to some degree but that they may share a common mechanism that attenuates the cortical power in both hemispheres in the 15- to 30-Hz range during movements of one hand. The results are consistent with the hypothesis that oscillatory activity in the motor system is important in resetting the descending motor commands needed for changes in motor state, such as those that occur in the transition from movement to steady grip.

Three‐dimensional integration of brain anatomy and function to facilitate intraoperative navigation around the sensorimotor strip

We studied 12 patients with brain tumors in the vicinity of the sensorimotor region to provide a preoperative three‐dimensional visualization of the functional anatomy of the rolandic cortex. We also evaluated the role of cortex‐muscle coherence analysis and anatomical landmarks in identifying the sensorimotor cortex. The functional landmarks were based on neuromagnetic recordings with a whole‐scalp magnetometer, coregistred with magnetic resonance images. Evoked fields to median and tibial nerve and lip stimuli were recorded to identify hand, foot and face representations in the somatosensory cortex. Oscillatory cortical activity, coherent with surface electromyogram during isometric muscle contraction, was analyzed to reveal the hand and foot representations in the precentral motor cortex. The central sulcus was identified also by available anatomical landmarks. The source locations, calculated from the neuromagnetic data, were displayed on 3‐D surface reconstructions of the individual brains, including the veins. The preoperative data were verified during awake craniotomy by cortical stimulation in 7 patients and by cortical somatosensory evoked potentials in 5 patients. Sources of somatosensory evoked fields identified correctly the postcentral gyrus in all patients. Useful corroborative information was obtained from anatomical landmarks in 11 patients and from cortex‐muscle correlograms in 8 patients. The preoperative visualization of the functional anatomy of the sensorimotor strip assisted in designing the operational strategy, facilitated orientation of the neurosurgeon during the operation, and speeded up the selection of sites for intraoperative stimulation or mapping, thereby helping to prevent damage of eloquent brain areas during surgery. Hum. Brain Mapping 12:180–192, 2001.

Reactivity of magnetic parieto-occipital alpha rhythm during visual imagery ☆

Spontaneous MEG signals were recorded during visual imagery from 13 healthy adults with a whole-scalp neuromagnetometer. The parieto-occipital 7-14 Hz alpha activity was suppressed strongly while subjects visualized and evaluated letters. The act of forming a visual image caused a smaller suppression than did inspection of the imaged pattern for a named property. The maximum suppression depended on the baseline alpha level and, for the majority of the subjects, occurred close to the area with the strongest alpha, showing no systematic hemispheric asymmetry. Sources for the alpha activity, modeled with equivalent current dipoles, clustered in the parietal and occipital lobes. The strongest suppression of the activity occurred near the parieto-occipital sulcus.