Neuromagnetic Cerebellar Activity Entrains to the Kinematics of Executed Finger Movements.

This magnetoencephalography (MEG) study aims at characterizing the coupling between cerebellar activity and the kinematics of repetitive self-paced finger movements. Neuromagnetic signals were recorded in 11 right-handed healthy adults while they performed repetitive flexion-extensions of right-hand fingers at three different movement rates: slow (~ 1 Hz), medium (~ 2 Hz), and fast (~ 3 Hz). Right index finger acceleration was monitored with an accelerometer. Coherence analysis was used to index the coupling between right index finger acceleration and neuromagnetic signals. Dynamic imaging of coherent sources was used to locate coherent sources. Coupling directionality between primary sensorimotor (SM1), cerebellar, and accelerometer signals was assessed with renormalized partial directed coherence. Permutation-based statistics coupled with maximum statistic over the entire brain volume or restricted to the cerebellum were used. At all movement rates, maximum coherence peaked at SM1 cortex contralateral to finger movements at movement frequency (F0) and its first harmonic (F1). Significant (statistics restricted to the cerebellum) coherence consistently peaked at the right posterior lobe of the cerebellum at F0 with no influence of movement rate. Coupling between Acc and cerebellar signals was significantly stronger in the afferent than in the efferent direction with no effective contribution of cortico-cerebellar or cerebello-cortical pathways. This study demonstrates the existence of significant coupling between finger movement kinematics and neuromagnetic activity at the posterior cerebellar lobe ipsilateral to finger movement at F0. This coupling is mainly driven by spinocerebellar, presumably proprioceptive, afferences.

Movement Kinematics Dynamically Modulates the Rolandic ~ 20-Hz Rhythm During Goal-Directed Executed and Observed Hand Actions.

This study investigates whether movement kinematics modulates similarly the rolandic α and ß rhythm amplitude during executed and observed goal-directed hand movements. It also assesses if this modulation relates to the corticokinematic coherence (CKC), which is the coupling observed between cortical activity and movement kinematics during such motor actions. Magnetoencephalography (MEG) signals were recorded from 11 right-handed healthy subjects while they performed or observed an actor performing the same repetitive hand pinching action. Subjects' and actor's forefinger movements were monitored with an accelerometer. Coherence was computed between acceleration signals and the amplitude of α (8-12 Hz) or ß (15-25 Hz) oscillations. The coherence was also evaluated between source-projected MEG signals and their ß amplitude. Coherence was mainly observed between acceleration and the amplitude of ß oscillations at movement frequency within bilateral primary sensorimotor (SM1) cortex with no difference between executed and observed movements. Cross-correlation between the amplitude of ß oscillations at the SM1 cortex and movement acceleration was maximal when acceleration was delayed by ~ 100 ms, both during movement execution and observation. Coherence between source-projected MEG signals and their ß amplitude during movement observation and execution was not significantly different from that during rest. This study shows that observing others' actions engages in the viewer's brain similar dynamic modulations of SM1 cortex ß rhythm as during action execution. Results support the view that different neural mechanisms might account for this modulation and CKC. These two kinematic-related phenomena might help humans to understand how observed motor actions are actually performed.

Effect of movement rate on corticokinematic coherence.

AIMS OF THE STUDY: This study investigates the effect of movement rate on the coupling between cortical magnetoencephalographic (MEG) signals and the kinematics of repetitive active finger movements, i.e., the corticokinematic coherence (CKC). MATERIAL AND METHODS: CKC was evaluated in ten right-handed healthy adults performing repetitive flexion-extension of the right-hand fingers in three different movement rate conditions: slow (∼1 Hz, duration: 11 min), medium (∼2 Hz, duration: 5 min) and fast (∼3 Hz, duration: 3 min). Neuromagnetic signals were recorded with a whole-scalp-covering MEG (Elekta Oy) and index acceleration was monitored with a 3-axis accelerometer. Coherent sources were estimated on the time-course of the cross-correlogram using equivalent current dipole (ECD) modeling. RESULTS: Significant coherence was found at movement frequency or its first harmonics in all subjects and movement conditions. ECDs clustered at the primary sensorimotor cortex contralateral to hand movements. Movement rate had no effect on the coherence levels and the location of coherent sources. CONCLUSIONS: This study demonstrates that the movement rate does not affect coherence levels and CKC source location during active finger movements. This finding has direct implications for CKC functional mapping applications and studies investigating the pathophysiology of central nervous disorders affecting proprioceptive pathways.

Corticokinematic coherence during active and passive finger movements.

Corticokinematic coherence (CKC) refers to coupling between magnetoencephalographic (MEG) brain activity and hand kinematics. For voluntary hand movements, CKC originates mainly from the primary sensorimotor (SM1) cortex. To learn about the relative motor and sensory contributions to CKC, we recorded CKC from 15 healthy subjects during active and passive right index-finger movements. The fingertip was either touching or not touching table, resulting in active-touch, active-no-touch, passive-touch, and passive-no-touch conditions. The kinematics of the index-finger was measured with a 3-axis accelerometer. Beamformer analysis was used to locate brain activations for the movements; somatosensory-evoked fields (SEFs) elicited by pneumatic tactile stimulation of the index finger served as a functional landmark for cutaneous input. All active and passive movements resulted in statistically significant CKC at the movement frequency (F0) and its first harmonic (F1). The main CKC sources at F0 and F1 were in the contralateral SM1 cortex with no spatial differences between conditions, and distinct from the SEF sources. At F1, the coherence was by two thirds stronger for passive than active movements, with no difference between touch vs. no-touch conditions. Our results suggest that the CKC occurring during repetitive finger movements is mainly driven by somatosensory, primarily proprioceptive, afferent input to the SM1 cortex, with negligible effect of cutaneous input.

A brush stimulator for functional brain imaging.

OBJECTIVE: To describe a novel non-magnetic hand-held device to stimulate various parts of the skin and to evaluate its performance in magnetoencephalographic (MEG) recordings. METHODS: The hand-held part of the device consists of an optic fiber bundle that forms a small brush. Half of the fibers emit modulated red light and the other half detect the reflected light from the skin so that the brush-to-skin contact is detected by means of reflectance. RESULTS: Light tapping of the back of the hand at the innervation area of the radial nerve elicited clear responses in all 10 subjects studied, with the main deflections peaking 40-70 ms after the stimulus. The earliest responses, obtained with a higher number of averaged trials, peaked 27-28 ms after the tap to the left hand dorsum. Source analysis of the MEG signals indicated neuronal sources at the primary somatosensory (SI) cortex, with a clear somatotopical order for face vs. hand. CONCLUSIONS: The device seems feasible for both MEG and functional magnetic resonance imaging experiments to address functional anatomy of the human somatosensory system with a real-life like stimulation. SIGNIFICANCE: Non-magnetic and artefact-free tactile stimulator with a selective stimulus offers new possibilities for experimental designs to study the human mechanoreceptor system.

Dorsal penile nerve stimulation elicits left-hemisphere dominant activation in the second somatosensory cortex.

Activation of peripheral mixed and cutaneous nerves activates a distributed cortical network including the second somatosensory cortex (SII) in the parietal operculum. SII activation has not been previously reported in the stimulation of the dorsal penile nerve (DPN). We recorded somatosensory evoked fields (SEFs) to DPN stimulation from 7 healthy adults with a 122-channel whole-scalp neuromagnetometer. Electrical pulses were applied once every 0.5 or 1.5 sec to the left and right DPN. For comparison, left and right median and tibial nerves were stimulated alternatingly at 1.5-sec intervals. DPN stimuli elicited weak, early responses in the vicinity of responses to tibial nerve stimulation in the primary somatosensory cortex. Strong later responses, peaking at 107-126 msec were evoked in the SII cortices of both hemispheres, with left-hemisphere dominance. In addition to tactile processing, SII could also contribute to mediating emotional effects of DPN stimuli.

Cortical activation associated with passive movements of the human index finger: an MEG study.

We recorded somatosensory evoked fields to passive extensions of the left and right index fingers in eight healthy adults. A new nonmagnetic device was designed to produce calibrated extensions of 19 degrees, with a mean angular velocity of 630 degrees/s. The responses, recorded with a 306-channel neuromagnetometer, were modeled with current dipoles. The earliest activation was in the primary somatosensory cortex, with peaks at 36-58 and 30-82 ms for left and right index finger extensions, respectively. Later signals were observed in the left second somatosensory (SII) cortex in six of eight subjects at 75-175 and 75-155 ms for left- and right-sided extensions, respectively; three subjects showed bilateral SII activation in at least one condition. Our results suggest a predominant role for the human left SII cortex in proprioceptive processing.

Functional overlap of finger representations in human SI and SII cortices.

We aimed to find out to what extent functional representations of different fingers of the two hands overlap at the human primary and secondary somatosensory cortices SI and SII. Somatosensory evoked fields (SEFs) were recorded with a 306-channel neuromagnetometer from 8 subjects. Tactile stimuli, produced by diaphragms driven by compressed air, were delivered to the fingertips in three different conditions. First, the right index finger was stimulated once every 2 s. Then two other stimuli were interspersed, in different sessions, to right- or left-hand fingers (thumb, middle finger, or ring finger) between the successive right index finger stimuli. Strengths of the responses to right index finger stimuli were evaluated in each condition. Responses to right index finger stimuli were modeled by three current dipoles, located at the contralateral SI and the SII cortices of both hemispheres. The earliest SI responses, peaking around 65 ms, were suppressed by 18% (P < 0.05) when the intervening stimuli were presented to the same hand; intervening stimuli to the other hand had no effect. The SII responses were bilaterally suppressed by intervening stimuli presented to either hand: in the left SII, the suppression was 39 and 42% (P < 0.01) and in the right SII 67 and 72% (P < 0.001) during left- and right-sided intervening stimuli, respectively. Left- and right-sided intervening stimuli affected similarly the SII responses and had no effect on the response latencies. The results indicate a strong and symmetric overlap of finger representations for both hands in the human SII cortices, and a weaker functional overlap for fingers of the same hand in the SI cortex.

Evidence for a 7- to 9-Hz "sigma" rhythm in the human SII cortex.

Electrical activity of the human brain features several rhythmical components which can be readily studied with whole-scalp neuromagnetometers. We describe a new 7- to 9-Hz "sigma" rhythm in the human second somatosensory cortex, distinct from both the mu rhythm of the primary sensorimotor cortex and the tau rhythm of the supratemporal auditory cortex. Sigma shows rate-selective responsiveness to rhythmical median nerve stimulation and is enhanced by stimulation at the rhythm's dominant frequency. Single stimuli may trigger several periods of the rhythm. The functional significance of the sigma rhythm remains to be investigated.

Extraction of event-related signals from multichannel bioelectrical measurements.

Independent component analysis (ICA) is a powerful tool for separating signals from their mixtures. In this field, many algorithms were proposed, but they poorly use a priori information in order to find the desired signal. Here, we propose a fixed point algorithm which uses a priori information to find the signal of interest out of a number of sensors. We particularly applied the algorithm to cancel cardiac artifacts from a magnetoencephalogram.

Independent component approach to the analysis of EEG and MEG recordings.

Multichannel recordings of the electromagnetic fields emerging from neural currents in the brain generate large amounts of data. Suitable feature extraction methods are, therefore, useful to facilitate the representation and interpretation of the data. Recently developed independent component analysis (ICA) has been shown to be an efficient tool for artifact identification and extraction from electroencephalographic (EEG) and magnetoencephalographic (MEG) recordings. In addition, ICA has been applied to the analysis of brain signals evoked by sensory stimuli. This paper reviews our recent results in this field.

Speaking modifies voice-evoked activity in the human auditory cortex.

The voice we most often hear is our own, and proper interaction between speaking and hearing is essential for both acquisition and performance of spoken language. Disturbed audiovocal interactions have been implicated in aphasia, stuttering, and schizophrenic voice hallucinations, but paradigms for a noninvasive assessment of auditory self-monitoring of speaking and its possible dysfunctions are rare. Using magnetoencephalograpy we show here that self-uttered syllables transiently activate the speaker's auditory cortex around 100 ms after voice onset. These phasic responses were delayed by 11 ms in the speech-dominant left hemisphere relative to the right, whereas during listening to a replay of the same utterances the response latencies were symmetric. Moreover, the auditory cortices did not react to rare vowel changes interspersed randomly within a series of repetitively spoken vowels, in contrast to regular change-related responses evoked 100-200 ms after replayed rare vowels. Thus, speaking primes the human auditory cortex at a millisecond time scale, dampening and delaying reactions to self-produced "expected" sounds, more prominently in the speech-dominant hemisphere. Such motor-to-sensory priming of early auditory cortex responses during voicing constitutes one element of speech self-monitoring that could be compromised in central speech disorders.

Cognitive response profile of the human fusiform face area as determined by MEG.

Activation in or near the fusiform gyrus was estimated to faces and control stimuli. Activation peaked at 165 ms and was strongest to digitized photographs of human faces, regardless of whether they were presented in color or grayscale, suggesting that face- and color-specific areas are functionally separate. Schematic sketche evoked approximately 30% less activation than did face photographs. Scrambling the locations of facial features reduced the response by approximately 25% in either hemisphere, suggesting that configurational versus analytic processing is not lateralized at this latency. Animal faces evoked approximately 50% less activity, and common objects, animal bodies or sensory controls evoked approximately 80% less activity than human faces. The (small) responses evoked by meaningless control images were stronger when they included surfaces and shading, suggesting that the fusiform gyrus may use these features in constructing its face-specific response. Putative fusiform activation was not significantly related to stimulus repetition, gender or emotional expression. A midline occipital source significantly distinguished between faces and control images as early as 110 ms, but was more sensitive to sensory qualities. This source significantly distinguished happy and sad faces from those with neutral expressions. We conclude that the fusiform gyrus may selectively encode faces at 165 ms, transforming sensory input for further processing.

Tracking functions of cortical networks on a millisecond timescale.

The human cerebral cortex, consisting of six layers and billions of neurons and synapses, processes sensory input from numerous sensory receptors. Noninvasive magnetoencephalographic (MEG) recordings provide a view through the skull to electrophysiological signals of the cortex on a millisecond timescale. For example, magnetic somatosensory evoked fields (SEFs) to a given peripheral somatosensory stimuli, reflect sequential activation of an extensive cortical network. Several cortical areas contributing to the SEFs can be evaluated in time and space by using source modeling. This brief review focuses on MEG studies of the human somatosensory networks with a special emphasis on tactile stimulation.

Somatosensory evoked fields to large-area vibrotactile stimuli.

We describe a method to apply large-area vibrotactile stimuli, based on a vibrating balloon, on the palms of both hands during evoked response studies. Magnetoencephalographic (MEG) signals were recorded with a whole-scalp neuromagnetometer from six healthy subjects while they held their hands on a balloon which was made to vibrate by delivering tones to it through a loudspeaker and a tube. The 200 Hz stimuli, presented once every 1 or 2 s in separate sessions, elicited prominent and replicable somatosensory evoked fields (SEFs) and also auditory evoked fields (AEFs) due to the concomitant sound. Source modelling allowed reliable differentiation between bilateral activation of the primary somatosensory (SI) cortices (peaks at 46-61 ms after the stimulus onset) and of the supratemporal auditory cortices (peaks at 104-126 ms). These simple vibrotactile stimuli could be useful for rapid and reliable identification of the somatosensory and auditory cortices, for example in presurgical evaluation of children.

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.

Magnetoencephalography in presurgical evaluation of children with the Landau-Kleffner syndrome.

PURPOSE: Our aim was (a) to localize the primary epileptogenic cortex for possible multiple subpial transsection in four children with the Landau-Kleffner syndrome (LKS), and (b) to evaluate the impact of magnetoencephalography (MEG) in the localizing process. METHODS: We used EEG to detect the overall epileptiform activity and MEG for selective recording of fissural spikes. The cortical generators of MEG spikes were modeled with dipoles, and their activation order was determined. The voltage distribution, consistent with the earliest MEG sources, was then identified during the course of the patient's EEG spikes to determine the relative timing between stereotypic EEG and MEG spikes and to distinguish the earliest (primary) source area among the secondary ones. RESULTS: In all patients, the earliest spike activity originated in the intrasylvian cortex, spreading in one subject to the contralateral sylvian cortex within 20 ms. Secondary spikes occurred within 10-60 ms in ipsilateral perisylvian, temporooccipital, and parietooccipital areas. A single intrasylvian pacemaker initiated all epileptic activity in two patients, whereas the other two had independent left- and right-hemisphere circuits or focal spikes. MEG source dynamics predicted the results of the methohexital suppression test in two patients and was confirmed by surgery outcome in one patient, in whom all epileptic activity ceased after a small transsection of the sylvian pacemaker. CONCLUSIONS: (a) The intrasylvian cortex is a likely pacemaker of epileptic discharges in LKS, and (b) MEG provides useful presurgical information of the cortical spike dynamics in LKS patients.

Effects of stimulus intensity on signals from human somatosensory cortices.

We recorded somatosensory evoked magnetic fields (SEFs) to left median nerve electric stimulation from seven healthy subjects. The stimulus intensity was varied in three sessions: sensory stimuli evoked a clear tactile sensation without any movement, weak motor stimuli exceeded the motor threshold, and strong motor stimuli caused a vigorous movement. Responses were modelled with sources in the contralateral primary somatosensory cortex (SI), the contralateral and ipsilateral secondary somatosensory cortices (SIIs) and the contralateral posterior parietal cortex (PPC). The amplitude of the 20 ms response from the SI cortex and the subjective magnitude estimations followed the stimulus intensity whereas signals from the three other areas saturated already at the level of the motor threshold. The results implicate differential roles for various somatosensory cortices in intensity coding.

Vibration-induced auditory-cortex activation in a congenitally deaf adult.

Considerable changes take place in the number of cerebral neurons, synapses and axons during development, mainly as a result of competition between different neural activities [1-4]. Studies using animals suggest that when input from one sensory modality is deprived early in development, the affected neural structures have the potential to mediate functions for the remaining modalities [5-8]. We now show that similar potential exists in the human auditory system: vibrotactile stimuli, applied on the palm and fingers of a congenitally deaf adult, activated his auditory cortices. The recorded magnetoencephalographic (MEG) signals also indicated that the auditory cortices were able to discriminate between the applied 180 Hz and 250 Hz vibration frequencies. Our findings suggest that human cortical areas, normally subserving hearing, may process vibrotactile information in the congenitally deaf.