Event-related desynchronization and synchronization. Reactivity of electrocortical rhythms in relation to the planning and execution of voluntary movement]

Cortical electroencephalographic rhythms reactivity may be quantified using event-related desynchronization (ERD) and synchronization (ERS) methods. We therefore studied cortical activation occurring during programming and performance of voluntary movement in healthy subjects. EEG power evolution within the reactive frequency bands (mu and beta central rhythms) was averaged before, during and after a minimum of 50 self-paced flexions of the thumb. Recordings in 18 normal adults showed that ERD (decrease in power) of mu rhythm started 2,000 ms before movement onset, while ERD of beta rhythm started 1,500 ms before movement onset. Early ERD of mu and beta rhythms were located over the contralateral central region covering primary motor cortex. They were followed by bilateral ERD occurring over ipsilateral and contralateral central regions during performance of the movement. At the end of the movement, an ERS (increase in power) of beta rhythm occurred. These results suggest that programming of voluntary movement induces early activation in contralateral sensorimotor areas, while performance of the movement induces bilateral activation in sensorimotor areas. ERS of beta rhythm occurring at the end of the movement could correspond to inactivation of motor areas activated by movement. Based on EEG activity, ERD and ERS prove to be useful methods to analyze cortical activation during programming and performance of voluntary movements with good spatial and temporal resolution.     

Roles of primate spinal interneurons in preparation and execution of voluntary hand movement

To study the contribution of primate cervical interneurons (INs) to preparation and execution of normal voluntary hand movement we investigated their activity and correlational linkages to muscles in monkeys performing tracking tasks. During ramp-and-hold flexion–extension torques about the wrist most task-related spinal INs exhibited some activity during both flexion and extension, in unexpected contrast to the strictly unidirectional activity of corticomotoneuronal (CM) cells and motoneurons. Most INs increased their activity more in one of these two directions; response patterns in their preferred direction were typically tonic or phasic-tonic. Spike-triggered averages of EMG detected significant features in muscle activity for many task-related INs. Premotor INs (PreM-INs) were identified by post-spike facilitation or suppression with appropriate onset latencies after the trigger spike. Muscle fields of PreM-INs were smaller than those of supraspinal PreM cells in cortex and red nucleus, and rarely involved reciprocal effects on antagonist muscles. To investigate the relation of spinal INs to a repertoire of different muscle synergies, activity of INs was recorded from a macaque performing a multidirectional wrist task. The monkey generated isometric torques in flexion/extension, radial/ulnar deviation, pronation/supination, and executed a power grip that co-contracted wrist flexor and extensor muscles. Many INs showing task-modulated activity had preferred directions in this multidirectional space, typically with broadly tuned activation. The role of spinal INs in preparation for voluntary movement was revealed in monkeys performing instructed delay tasks. During the delay between a transient visual cue and a go signal a third of the tested INs showed significant delay modulation (SDM) of firing rate relative to the pre-cue rate. The SDM responses often differed from the INs’ responses during the subsequent active torque period. In a monkey instructed by either visual or proprioceptive cues the delay period activity for many INs was similar in visual and perturbation trials, although other INs exhibited different SDM for visually and proprioceptively cued trials. These results suggest that spinal INs are involved, with cortex, in the earliest stages of movement preparation. The sensory input to INs could be identified in transient responses to the torque pulse, which showed two predominant patterns, consistent with inputs from cutaneous or proprioceptive receptors. We also investigated the task-dependent modulation of neural responses to peripheral input in a monkey performing wrist flexion/extension movements in a visually cued instructed delay task. Monosynaptic responses evoked by electrical stimulation of the superficial radial nerve through a cuff electrode were suppressed or abolished during the dynamic movement phase. Since task-related activity of the INs increased at the same time, the suppression was mediated by presynaptic rather than postsynaptic inhibition. These observations indicate that under normal behavioral conditions many spinal INs have response properties comparable to those previously documented for cortical neurons in behaving animals.     

The mechanisms of movement preparation: a precuing study

It is well known that precues about the possible locations of upcoming targets reduce the manual reaction time. The present study investigates at which stage of the sensorimotor system such precues act. Several light dots were displayed as a precue; one of them became the target, and subjects had to produce a manual response to the target as fast as possible. Exp. A varied independently the number of precues and the area of space they occupy; we found that the reaction time of pointing movements depended on spatial extent, but not on the number of choices. The outcome was similar in Exp. B, where subjects produced stereotyped 'tapping' movements irrespective of target position. Taken together, both findings support the view that precues act mainly at a stage concerned with the internal representation of space, rather than with response selection or movement preparation. The effects of precues were preserved when subjects fixated throughout the precuing period (Exp. C), but not when precue and target positions were uncorrelated (Exp. D). These findings do not support the alternative interpretations, that precues act by guiding the eyes into the vicinity of targets, or by elevating the subjects' arousal level.     

Gating of sensory input at spinal and cortical levels during preparation and execution of voluntary movement

All bodily movements stimulate peripheral receptors that activate neurons in the brain and spinal cord through afferent feedback. How these reafferent signals are processed within the CNS during movement is a key question in motor control. We investigated cutaneous sensory-evoked potentials in the spinal cord, primary somatosensory and motor cortex, and premotor cortex in monkeys performing an instructed delay task. Afferent inputs from cutaneous receptors were suppressed at several levels in a task-dependent manner. We found two types of suppression. First, suppression during active limb movement was observed in the spinal cord and all three cortical areas. This suppression was induced by both bottom-up and top-down gating mechanisms. Second, during preparation for upcoming movement, evoked responses were suppressed exclusively in the motor cortical areas and the magnitude of suppression was correlated with the reaction time of the subsequent movement. This suppression could be induced by a top-down gating mechanism to facilitate the preparation and execution of upcoming movement.     

The stability of synergy in agonists during the execution of a simple voluntary movement.

The characteristics of muscular synergy between the main elbow flexors (biceps brachii, brachialis, brachoradialis were considered. The activities of these different muscles were recorded simultaneously with surface and wire electrodes. A quantitative analysis of the activity of each of these muscles and of their excitation levels was carried out during movements performed at various velocities and against different inertias. It was shown that: (1) the onset as well as the cessation of activity in the different muscles occur practically simultaneously and independently of the velocity and inertia of the movement; (2) the well-known linear relation between the integrated EMG of biceps brachii and the work can be extended to the other main flexors. This implies that the relation between the activities of the main flexors remains constant whatever the velocity and inertia may be. These results confirm the notion of 'Flexor Equivalent'. They also demonstrate a stability of the synergy between agonist muscles which must be distinguished particularly from the synergy between agonists and antagonists.     

Functional dissociation of lower and upper frequency mu rhythms in relation to voluntary limb movement.

OBJECTIVE: The goal of this study is to investigate the reactivity of central rhythms in the alpha band during self-paced voluntary finger and foot movement and to give an answer to the question, whether different types of mu rhythms exist. METHODS: The effect of self-paced, voluntary finger and foot movement was studied in a group of 12 right-handed healthy volunteers. The EEG was recorded from a grid of 34 electrodes placed over sensorimotor areas with inter-electrode distances of approximately 2.5 cm. The event-related desynchronization (ERD) was quantified in the 8-10 and 10-12 Hz bands. RESULTS: Both frequency components are blocked prior to and during movement and therefore, they have to be considered as mu rhythms. The lower frequency component results in a widespread movement-type non-specific ERD pattern, whereas the upper frequency component shows a more focused and movement-type specific pattern, clearly different with finger and foot movement. CONCLUSIONS: The distinct reactivity patterns provide evidence for the existence of two types of mu rhythms, a somatotopically non-specific lower frequency mu rhythm and a somatotopically specific mu rhythm characteristically found in the upper alpha frequency band.     

Activities of the primary and supplementary motor areas increase in preparation and execution of voluntary muscle relaxation: an event-related fMRI study.

Brain activity associated with voluntary muscle relaxation was examined by applying event-related functional magnetic resonance imaging (fMRI) technique, which enables us to observe change of fMRI signals associated with a single motor trial. The subject voluntarily relaxed or contracted the right upper limb muscles. Each motor mode had two conditions; one required joint movement, and the other did not. Five axial images covering the primary motor area (M1) and supplementary motor area (SMA) were obtained once every second, using an echoplanar 1.5 tesla MRI scanner. One session consisted of 60 dynamic scans (i.e., 60 sec). The subject performed a single motor trial (i.e., relaxation or contraction) during one session in his own time. Ten sessions were done for each task. During fMRI scanning, electromyogram (EMG) was monitored from the right forearm muscles to identify the motor onset. We calculated the correlation between the obtained fMRI signal and the expected hemodynamic response. The muscle relaxation showed transient signal increase time-locked to the EMG offset in the M1 contralateral to the movement and bilateral SMAs, where activation was observed also in the muscle contraction. Activated volume in both the rostral and caudal parts of SMA was significantly larger for the muscle relaxation than for the muscle contraction (p < 0.05). The results suggest that voluntary muscle relaxation occurs as a consequence of excitation of corticospinal projection neurons or intracortical inhibitory interneurons, or both, in the M1 and SMA, and both pre-SMA and SMA proper play an important role in motor inhibition.     

Neural mechanisms of voluntary and involuntary recall: a PET study

Neuropsychological and neuroimaging studies on episodic memory retrieval have primarily focused on volitional memory tasks. However, some conscious memories arise involuntarily, i.e. without a strategic retrieval attempt, yet little is known about the neural network underlying involuntary episodic memory. The aim of this study was to determine whether voluntary and involuntary recall are mediated by separate cortical networks. We used positron emission tomography (PET) to measure changes in regional cerebral blood flow (rCBF) in 12 healthy subjects during voluntary and involuntary cued recall of pictures and a control condition with no episodic memory requirements. Involuntary recall was elicited by using an incidental memory task. Compared to the control condition, voluntary and involuntary recall were both associated with significant regional cerebral blood flow (rCBF) increases in posterior cingulate gyrus (PCG; BA 23), left precuneus (BA 7), and right parahippocampal gyrus (BA 35/36). In addition, rCBF in right dorsolateral prefrontal cortex (PFC; BA 8/9) and left precuneus (BA 7) was significantly larger during voluntary compared to involuntary recall, while rCBF was enhanced in left dorsolateral PFC (BA 9) during involuntary recall. The findings corroborate an association of the right PFC with a strategic component of episodic memory retrieval. Moreover, they show for the first time that it is possible to activate the medial temporal lobe, the PCG, and the precuneus, regions normally associated with retrieval success, without this strategic element. The relatively higher activity in precuneus during voluntary compared to involuntary recall suggests that activity in this region co-varies not only with retrieval success but also with retrieval intentionality.     

Interactions between imagined movement and the initiation of voluntary movement: A TMS study

ObjectiveThe purpose was to examine motor imagery-induced enhancement in corticospinal excitability during a reaction time (RT) task.MethodsNine young and healthy subjects performed an isometric finger flexion tasks in response to a visual imperative cue. In the pre-cue period, they were instructed to: (1) rest; (2) imagine flexing their fingers isometrically (ImFlex); or (3) imagine extending their fingers isometrically (ImExt). Surface EMGs from the finger flexors and extensors were monitored to ensure EMG silence before movement onset. Transcranial magnetic stimulation (TMS) was used to evaluate changes in motor-evoked potentials (MEP) in the finger flexor and extensor muscles during the response phase. TMS was delivered either with the imperative cue, or 120 ms before and after the imperative cue.ResultsRT was slower when they were imagining finger extension prior to the visual imperative cue. MEPs were significantly increased for the finger flexors during imagined finger flexion and for the finger extensors during imagined finger extension at both TMS delivery time points, reflecting movement specific enhancement in corticospinal excitability during motor imagery. When TMS was delivered 120 ms after the cue, finger flexor MEPs were further facilitated under the Rest and ImFlex conditions, but not under the ImExt condition, suggesting additive interactions between imagery-induced enhancement and early rise in corticospinal excitability during the initiation of a reaction time response.ConclusionsOur results provide neurophysiological evidence mediating dynamic interactions between imagined movement and the initiation of voluntary movement.SignificanceMotor imagery can be integrated into a rehabilitation protocol to facilitate motor recovery.     

Gilles de la Tourette syndrome and voluntary movement: a functional MRI study

Tourette syndrome (TS) is hypothesised to be caused by an abnormal organization of movement control. The aim of this study was to use functional magnetic resonance imaging to study motor cortex activation in a TS patient. Usual and unusual self-paced voluntary movements were performed. The TS patient displayed supplementary motor area (SMA) activation during both tasks. This activation reflects a continuous use of the SMA to perform the voluntary motor movements required in both tasks. Moreover, the absence of tics during the execution of these voluntary motor tasks suggests that tic activity may be suppressed by additional mental effort.     

Pyramidal tract neurons in somatosensory cortex: central and peripheral inputs during voluntary movement

Recordings from pyramidal tract neurons (PTNs) in the primary somatosensory cortex of the monkey show that these neurons have 3 properties in common with PTNs of primary motor cortex: (1) they exhibit discharge prior to the onset of voluntary movement, (2) their discharge frequency varies as a function of strength of muscular contraction, and (3) they show reflex responses to afferent stimuli that occur during movement. These findings support the view that in addition to its widely recognized role in somesthetic perception, somatosensory cortex has a direct role in the control of movement.     

Cortical potentials preceding voluntary movement: evidence for three periods of preparation in man

We have recorded movement-related cortical potentials (MRCPs) to voluntary middle finger extension from 10 young and 10 old subjects free of neurological disease using the method of detecting EMG onset associated with each movement described by Barrett et al. (1985). The slow potential shifts preceding movement were measured by fitting a linear regression line to the wave forms to obtain a measure of their slope. Three separate potential shifts were identified. The first had a scalp distribution and onset latency similar to the Bereitschaftspotential (BP) first reported by Kornhuber and Deecke (1964, 1965). The potential shift immediately preceding movement corresponded with the NS' of Shibasaki et al. (1980). We identified, for the first time, a third shift intervening between BP and NS' and named it the intermediate shift (IS). The onset of BP occurred about 1.6 sec before EMG onset and was followed by IS which began about 875 msec before movement. The onset of NS' occurred 300 msec before EMG onset and terminated about 90 msec before this event. The slope of BP preceding right finger movement was steeper than that preceding left hand movement in all our right-handed subjects. The distribution of BP was symmetric about the midline. The IS potential shift had a slope which was steeper on the average preceding left finger movement than right. The distribution of IS was symmetric about the midline preceding left finger movement but had a contralateral tendency preceding right hand movement. NS' had a maximum slope at contralateral electrodes over the hand motor area and parietal areas. It was suggested that the BP potential shift originates in the supplementary motor area on the medial surface of the cerebral cortex. The differing distribution of the IS shift for the two hands suggests that this potential may be generated bilaterally preceding left finger movement but from the contralateral hemisphere only preceding movement of the right finger. The most likely origin of this potential was thought to be superior premotor cortex. NS' was considered to originate in primary motor cortex with possible contributions from other cortical areas associated with movement.     

Inhibitory effect of voluntary movement preparation on cutaneous heat pain and laser-evoked potentials

In our study, preparation of voluntary movement was used to physiologically activate the motor cortex areas and the effect of this activation on CO(2) laser-evoked potentials (LEPs) was explored. LEPs were recorded from 31 scalp electrodes in 10 healthy subjects after painful stimulation of the right C6-C7 skin dermatomes. LEP stimuli were delivered in the time interval between a visual warning stimulus followed after 1 s. by an imperative stimulus. The imperative stimulus triggered: (i) no task in the baseline condition (Pain); (ii) flexion-extension movements of the second finger of the right hand in the movement condition (Pain + Movement); (iii) cognitive task (mathematic computation) in the distraction condition (Pain + Cognition). The experimental conditions were also repeated during application of laser stimuli on the left C6-C7 skin dermatomes. Compared with the baseline condition (no task required), during preparation of right-hand voluntary movement there was a significant reduction in LEP amplitude and subjective pain rating after right- but not after left-hand stimulation, which suggests that the observed effect cannot be attributed to a nonspecific reduction in attention toward painful stimulus. During preparation of a cognitive task, LEP amplitude was reduced compared to baseline. Our results represent the first neurophysiological suggestion that physiological activation of the motor cortex, occurring during movement preparation, inhibits cortical pain processing by a centrifugal mechanism.     

Bilateral neuromagnetic activation of human primary sensorimotor cortex in preparation and execution of unilateral voluntary finger movements

Extracranial magnetoencephalographic activity was separately recorded (25 channels) from bilateral primary sensorimotor cortex (M1-S1) of normal right-handers during unilateral finger movements. Standard dipole analysis indicated only a contralateral M1-S1 source for first movement-evoked field (MEF1) peaking at about 115 ms after electromyographic onset. However, the subtraction of the magnetic field generated by this source from the recorded magnetic field disclosed a low-amplitude ipsilateral central-parietal MEF1 that was explained by an ipsilateral M1-S1 source.     

Studies of parkinsonian movement: 1. Programming and execution of eye movements

Rapid voluntary eye movements in bradykinetic parkinsonian patients and normal subjects were recorded when the movement was executed with visual feedback (closed-loop mode) and in darkness without visual feedback (open-loop mode). The patients had a tendency to generate abnormal saccades consisting of multiple small steps (multiple step saccade), both in the closed-loop and open-loop mode. They were, however, also capable of generating large amplitude saccades. The amplitude-velocity relation of both the small step saccades and the large saccades was normal. The presence of multiple step saccades in the open loop mode suggests that the patients used internal rather than external (visual) feedback to compare the actual eye position with the desired (programmed) eye position and the program for rapid movement is normal but its execution is defective.
Horizontal eye movements were also recorded when the head was stationary with a target moving sinusoidally, and when the target was stationary with the head rotated sinusoidally. In both cases the amplitude of the eye movement relative to the head was about 50°. The patients were observed to generate irregular, saccadic eye movements in pursuit of a slowly moving target when the head was stationary, but their eyes could follow a stationary target smoothly when their head was moved sinusoidally. These findings suggest that the neuronal circuitry in the paramedian pontine reticular formation, responsible for the final integration of different types of eye movements, is physiologically normal in Parkinsonism.     

Extension of mental preparation positively affects motor imagery as compared to motor execution: a functional near-infrared spectroscopy study

Motor imagery (MI) is widely used to study cognitive action control. Although, the neural simulation theory assumes that MI and motor execution (ME) share many common features, the extent of similarity and whether it spreads into the preparation phase is still under investigation. Here we asked, whether an extension of physiological mental preparation has a comparable effect on MI and ME.Data were recorded using wireless functional near-infrared spectroscopy (fNIRS) in a two-stage task design where subjects were cued with or without preparatory stimuli to either execute or imagine complex sequential thumb-finger tasks.The main finding is that the extended mental preparation has a significant positive effect on oxy-hemoglobin (?[O₂Hb]) in response to MI, which is proportionally larger as that found in response to ME. Furthermore, fNIRS was capable to discriminate within each task whether it was preceded by preparatory stimuli or not. Transition from mental preparation to actual performance (ME or MI) was reflected by a dip of the fNIRS signal presumably related to underlying cortical processes changing between preparation and task performance. Statistically significant main effects of ‘Preparation’ and ‘Task’ showed that ?[O₂Hb] during preparation was preparation-specific, i.e., positively affected by the presence of preparatory stimuli, whereas during task performance ?[O₂Hb] was both preparation- and task-specific, i.e., additionally affected by the task mode.These results are particularly appealing from a practical point of view for making use of MI in neuroscientific applications. Especially neurorehabilitation and neural interfaces may benefit from utilizing positive interactions between mental preparation and MI performance.     

Studies of parkinsonian movement: 2. Initiation of fast voluntary eye movement during postural disturbance

Delay in initiation of rapid voluntary eye movements (saccades) in brady-kinetic parkinsonian patients and normal subjects was recorded with and without postural disturbances (rotation of the body and head).
Parkinsonian patients as a group exhibited longer delays in the initiation of saccades. The delay increased during postural disturbance in both the patients and the normal subjects.
The study failed to substantiate the hypothesis that postural reflexes interfere with the initiation of voluntary movement in Parkinsonism.