Time course of corticospinal excitability in reaction time and self-paced movements

We used transcranial magnetic stimulation (TMS) to study the time course of corticospinal excitability before and after brisk thumb abduction movements, either in a simple reaction time (RT) paradigm or self-paced. Premovement increase in corticospinal excitability began about 20 msec earlier for self-paced compared with simple RT movements. For both simple RT and self-paced movements after electromyographic (EMG) offset, there was a first period of increased excitability from 0 to 100 msec, followed by a second period from 100 to 160 msec. Corticospinal excitability was decreased from about 500 to 1,000 msec after EMG offset for both types of movements. Our results show that motor preparation that begins 1.5 to 2 seconds before self-paced movement is not associated with increased corticospinal excitability. The first phase of increased corticospinal excitability after EMG offset may be due to activity of motor cortex neuron subthreshold for activating spinal motor neurons, and the second phase may reflect a subthreshold second agonist burst. The period of decreased corticospinal excitability after movement corresponds to the onset of event-related synchronization (ERS) of electroencephalographic signals in the 20-Hz band, and supports the hypothesis that ERS may be related to an inactive, idling state of the motor cortex.     

Event-related desynchronization in reaction time paradigms: a comparison with event-related potentials and corticospinal excitability.

OBJECTIVES: To study cortical activity in different motor tasks, we compared event-related desynchronization (ERD) and event-related potentials (ERPs) in different reaction time (RT) paradigms with the time course of corticospinal excitability. METHODS: Nine right-handed, normal subjects performed right or left thumb extensions in simple, choice and go/no go auditory RT paradigms. Eight subjects had participated in a previous study evaluating changes in corticospinal excitability during the same paradigms. Twenty-nine EEG channels with electrooculogram and bilateral EMG monitoring were collected. ERPs and ERD of 10 and 18-22 Hz bands were obtained with respect to tone administration and EMG onset. RESULTS: Trials with movement showed lateralized ERP components, corresponding to the motor potential (MP), both in the averages on the tone and on EMG. The MP corresponded well in time and location to the rise in corticospinal excitability on the moving side observed in the previous study. Sensorimotor ERD, followed by event-related synchronization (ERS), was present for trials with movements and for the no go. ERD was present contralaterally during movement preparation and in no go trials, while it was bilateral during motor execution. No go ERD was followed more rapidly by ERS than in trials with movement. This finding suggests that in no go trials, there is a brief active process in the sensorimotor areas. ERD and ERS do not correspond, respectively, in time and location to increases and decreases in corticospinal excitability. In fact, ERD is bilateral during movement execution, when corticospinal inhibition of the side at rest is observed. Contralateral no go ERS occurs later than corticospinal inhibition, which is bilateral. CONCLUSIONS: These findings may suggest that ERD is compatible with both corticospinal activation and inhibition, ERS indicating the removal of either, resulting in cortical idling.     

Event-related desynchronization and excitability of the ipsilateral motor cortex during simple self-paced finger movements.

OBJECTIVE: To study the time course of oscillatory EEG activity and corticospinal excitability of the ipsilateral primary motor cortex (iM1) during self-paced phasic extension movements of fingers II-V. METHODS: We designed an experiment in which cortical activation, measured by spectral-power analysis of 28-channel EEG, and cortical excitability, measured by transcranial magnetic stimulation (TMS), were assessed during phasic self-paced extensions of the right fingers II-V in 28 right-handed subjects. TMS was delivered to iM1 0-1500 ms after movement onset. RESULTS: Ipsilateral event-related desynchronization (ERD) during finger movement was paralleled by increased cortical excitability of iM1 from 0-200 ms after movement onset and by increased intracortical facilitation (ICF) without changes in intracortical inhibition (ICI) or peripheral measures (F waves). TMS during periods of post-movement event-related synchronization (ERS) revealed no significant changes in cortical excitability in iM1. CONCLUSIONS: Our findings indicate that motor cortical ERD ipsilateral to the movement is associated with increased corticospinal excitability, while ERS is coupled with its removal. These data are compatible with the concept that iM1 contributes actively to motor control. No evidence for inhibitory modulation of iM1 was detected in association with self-paced phasic finger movements. SIGNIFICANCE: Understanding the physiological role of iM1 in motor control.     

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.     

Impairment of motor cortex activation and deactivation in Parkinson's disease.

OBJECTIVES: To examine the time course of corticospinal excitability before and after voluntary movement in Parkinson's disease (PD). METHODS: We studied 9 mild PD patients at least 12 h off medications and 11 healthy volunteers in a simple reaction time (RT) paradigm. Suprathreshold transcranial magnetic stimulation was delivered to the left motor cortex at intervals covering the periods before and after movement. RESULTS: PD patients (284+/-90 ms) and normal subjects (282+/-56 ms) had similar median RT. The time courses of both the premovement increase and the postmovement decrease in corticospinal excitability were significantly different between PD patients and normal subjects. The increase in motor-evoked potential (MEP) amplitudes began earlier for PD patients (200 ms before electromyographic (EMG) onset) than for normal subjects (150 ms before EMG onset), but the rate of increase was slower in PD patients than controls. After EMG offset, MEP amplitudes were increased for about 150 ms in normal subjects, but in PD patients this period was prolonged to about 350 ms. CONCLUSIONS: Impairment of motor cortex activation and deactivation is an early feature of PD and may be a physiological correlate of bradykinesia. The normal RT in our patients may be related to the earlier occurrence of the premovement increase in corticospinal excitability compensating for the slower rate of rise.     

Altered pattern of motor cortical activation–inhibition during voluntary movements in Tourette syndrome

In patients with Gilles de la Tourette syndrome (GTS) alterations of motor cortex (M1) excitability at rest have been evidenced. In contrast, there has so far been little research into changes of motor cortical reactivity during the time course of voluntary movements in GTS patients. The present study investigates neuromagnetic event‐related desynchronization (ERD) and event‐related synchronization (ERS) of bilateral M1 in 11 GTS patients and 11 healthy control subjects. ERD represents motor cortical activation, whereas ERS most likely indicates its inhibition. Subjects performed a self‐paced finger movement task while magnetoencephalography was used to record neuromagnetic activity. In GTS patients, ERD at beta frequency was significantly increased in the contralateral hemisphere before and during movements, whereas ERS following movement termination was increased in M1 ipsilateral. Ipsilateral ERS was inversely correlated with tic severity as determined by the Yale Global Tic Severity Rating Scale. The data of the present study support the hypothesis that during voluntary movements, motor cortical reactivity is pathologically altered in GTS patients. The observed pattern of increased activation (ERD) prior to and during movement execution followed by increased inhibition (ERS) after movement termination at beta frequency suggests abnormally increased motor cortical activation, possibly driving stronger inhibition. The stronger this inhibition is, the better symptoms appear to be controlled. © 2010 Movement Disorder Society     

Early non‐specific modulation of corticospinal excitability during action observation

Activity of the primary motor cortex (M1) during action observation is thought to reflect motor resonance. Here, we conducted three studies using transcranial magnetic stimulation (TMS)‐induced motor‐evoked potentials (MEPs) of the first dorsal interosseus muscle (FDI) during action observation to determine: (i) the time course of M1 corticospinal excitability during the observation of a simple finger movement; (ii) the specificity of M1 modulation in terms of type of movement and muscle; and (iii) the relationship between M1 activity and measures of empathy and autistic traits. In a first study, we administered single‐pulse TMS at 30‐ms intervals during the observation of simple finger movements. Results showed enhanced corticospinal excitability occurring between 60 and 90 ms after movement onset. In a second experiment, TMS‐induced MEPs were recorded from the FDI and abductor digiti minimi muscles while pulses were delivered 90 ms after movement onset during observation of simple finger movement and dot movement. Increased corticospinal excitability was restricted to finger movement and was present in both muscles. Finally, in an exploratory experiment, single‐pulse TMS was administered at 30, 90 and 150 ms after movement onset, and participants were asked to complete the Empathy Quotient (EQ) and the Autism Spectrum Quotient (AQ). Correlational analysis revealed a significant link between motor facilitation at 90 ms and the EQ and AQ scores. These results suggest that corticospinal excitability modulation seen at M1 during action observation is the result of a rapid and crude automatic process, which may be related to social functioning.     

Inhibition in the human motor cortex is reduced just before a voluntary contraction.

OBJECTIVE: To examine inhibition in the human motor cortex before and during voluntary movements. METHODS: The balance between the excitation and inhibition of corticospinal neurons in the human motor cortex was tested by conditioning the motor evoked potentials (MEP) evoked in forearm muscles by transcranial magnetic stimulation with a preceding subthreshold stimulus delivered through the same coil. RESULTS: When normal individuals (n = 9) made a tonic wrist extension, inhibition of the forearm extensor MEP decreased, whereas that of the forearm flexors was unchanged. When these individuals made a tonic wrist flexion, inhibition of the forearm flexor MEP diminished, whereas that of the forearm extensors was unchanged. When normal individuals (n = 10) made a phasic wrist extension in response to an auditory signal, inhibition of the extensor MEP began to decline about 95 msec before the onset of the agonist EMG activity. CONCLUSIONS: The changes in balance of excitation and inhibition of corticospinal neurons associated with a voluntary movement precede the movement and are directed at the corticospinal neurons projecting to the agonists. These changes may help to select the population of cortical neurons responsible for the movement.     

Motor cortex excitability during ballistic forearm and finger movements

In a ballistic forearm flexion movement, a centrally programmed triphasic pattern of electromyogram (EMG) is seen with two bursts in biceps and a single burst in triceps. Rapid abduction of the index finger, in contrast, is achieved with a single agonist burst. Transcranial magnetic and electrical stimuli, triggered at the onset of the EMG burst, have been used to probe cortical and spinal cord excitability during and after self-paced ballistic finger and forearm movements. In both, the motor cortex has two phases of increased excitability. The first phase is coincident with the initial agonist burst. The second phase in biceps is associated with the second agonist burst, but in the finger movement, the raised motor cortical excitability is not associated with any EMG. It is argued that the motor program for the two movements may be similar, despite there being large differences in the EMG pattern generated. © 1996 John Wiley & Sons, Inc.     

Corticospinal excitability is specifically modulated by motor imagery: a magnetic stimulation study

Transcranial magnetic stimulation (TMS) was used to investigate whether the excitability of the corticospinal system is selectively affected by motor imagery. To this purpose, we performed two experiments. In the first one we recorded motor evoked potentials from right hand and arm muscles during mental simulation of flexion/extension movements of both distal and proximal joints. In the second experiment we applied magnetic stimulation to the right and the left motor cortex of subjects while they were imagining opening or closing their right or their left hand. Motor evoked potentials (MEPs) were recorded from a hand muscle contralateral to the stimulated cortex. The results demonstrated that the excitability pattern during motor imagery dynamically mimics that occurring during movement execution. In addition, while magnetic stimulation of the left motor cortex revealed increased corticospinal excitability when subjects imagined ipsilateral as well as contralateral hand movements, the stimulation of the right motor cortex revealed a facilitatory effect induced by imagery of contralateral hand movements only. In conclusion, motor imagery is a high level process, which, however, manifests itself in the activation of those same cortical circuits that are normally involved in movement execution.     

Functional relationship between human rolandic oscillations and motor cortical excitability: an MEG study

Synchronization and desynchronization of the neural rhythm in the brain play an important role in the orchestration of perception, motor action and conscious experience. Based on the results of electrocorticographic and magnetoencephalographic (MEG) recordings, it has been considered that human rolandic oscillations originate in the anterior bank of the central sulcus (20-Hz rhythm) and the postcentral cortex (10-Hz rhythm): the 20-Hz oscillation is closely related to motor function, while the 10-Hz rhythm is attributed mainly to sensory function. To test whether the rolandic oscillations are functionally relevant to the motor cortical excitability, we examined effects of 1-Hz repetitive transcranial magnetic stimulation (rTMS) of the left primary motor cortex (M1) on movement-related changes of the rolandic oscillations in 12 normal subjects. MEG data recorded during brisk extension of the right index finger in two different sessions (with and without rTMS conditioning) were compared. Motor-evoked potential (MEP) of the right hand muscle was also measured before and after rTMS to assess the motor cortical excitability. We found that 1-Hz rTMS over M1 significantly reduced the movement-related rebound of the 20-Hz oscillation in association with decreased motor cortical excitability. In particular, movement-related rebound of the 20-Hz rhythm was closely tied with motor cortical excitability. These findings further strengthen the notion of functional relevance of 20-Hz cortical oscillation to motor cortical excitability. In the framework of previous studies, the decrease in movement-related rebound may be regarded as a compensatory reaction to the inhibited cortical activity.     

Primary motor cortex and movement prevention: where Stop meets Go

Processes that engage frontal cortex and the basal ganglia are responsible for the prevention of planned movements. Here, we review the role of primary motor cortex (M1) in this function. M1 receives and integrates input from a range of cortical and subcortical sites. It is also the final cortical processing site for voluntary motor commands, before they descend to the spinal cord. Inhibitory networks within M1 may be an important mechanism for the prevention or suppression of movement. Transcranial magnetic stimulation (TMS) has been used to evaluate corticospinal excitability and intracortical inhibition in humans, during the performance of a range of movement selection and prevention tasks. This review explores how M1 intracortical inhibition is selectively reduced to initiate desired voluntary movements, while movement prevention is associated with rapid, non-selective recruitment of inhibition within M1. The relationship between deficient intracortical inhibition and behavioural inhibition is also explored. Examples of neuropathology are reviewed, including focal dystonia, attention deficit hyperactivity disorder and Tourette syndrome. The strengths and limitations of TMS in the study of movement prevention are also discussed. While the precise functional links between M1 neuronal populations and the fronto-basal-ganglia network activated by movement prevention have yet to be elucidated, it is clear that M1 plays a critical role in the final processing stage of response inhibition.     

Sensory and movement-related cortical potentials in nociceptive and auditory reaction time tasks

The processing of a sensory stimulus leading to a simple motor command was studied with scalp-recorded long latency cortical potentials in humans. Two sensory modalities were tested in their ability to activate descending motor pathways: auditory stimuli and painful cutaneous stimuli produced by a CO2 laser. Subjects were asked to react to stimuli with voluntary index finger movements. The stimulus-related and movement-related cortical potentials were recorded simultaneously with five midline electrodes on the scalp. The auditory reaction time, measured from the stimulus to the onset of electromyogram (EMG), was faster (150 ms) than the laser reaction time (350 ms). The onset of EMG of finger movements occurred only after the first negative components following auditory or laser stimuli but before the positive components. The latency from the auditory negativity to the onset of EMG was about 50 ms and the latency from the laser negativity to the onset of EMG was about 110 ms. This finding indicates that not only the peripheral afferent conduction but also central processing takes longer in a pain-related somatosensory task than in an auditory task. The frontal peak of Motor Potential (fpMP), a cortical potential related to the sensory feedback from movement, occurred with a constant latency after the onset of EMG (100 ms) and was unaffected by the task.     

Pathophysiology of tics and Tourette syndrome

Tics are involuntary movements that can affect one or more muscles producing simple or complex movements. Blink reflex and startle reflex studies disclose an increased excitability of brainstem interneurons. Analysis of voluntary movement shows that when advance visual information is reduced, patients with tics and Tourette syndrome become progressively slower in completing motor sequences. Sensorimotor integration is abnormally processed. Studies of the contingent negative variation demonstrate abnormalities of movement preparation and the investigation of premotor potentials shows that in some patients tics are not preceded by a normal premotor potential. Magnetic stimulation studies demonstrate an increased excitability of cortical motor cortex. Functional MRI, PET and SPECT studies show abnormal activation of cortical and subcortical areas. Dysfunction of basal ganglia-thalamo-cortical projections affects sensorimotor, language and limbic cortical circuits, and may explain why patients with Tourette syndrome have difficulty in inhibiting unwanted behaviors and impulses.     

Independent control of voluntary movements and associated anticipatory postural responses in a bimanual task.

OBJECTIVE: When one hand loads the other arm, EMG responses in the stationary arm anticipate the load. This study used transcranial magnetic stimulation over each hemisphere to clarify the relationship between a voluntary movement on one side and the anticipatory postural response on the other. METHODS: Subjects (n = 7) performed elbow flexion movements of one arm as a reaction-time task. Because subjects' arms were linked, flexion about one elbow resulted in extension force about the other, and an anticipatory response occurred in those elbow flexor muscles. After the 'go' signal and before the predicted onset of EMG, transcranial magnetic stimuli were delivered over one or other motor cortex. RESULTS: Stimulation contralateral to the reaction-time movement delayed the onset of voluntary EMG (46 ms in right biceps, 77 ms left) but did not alter the onset of EMG in the postural arm. Stimulation contralateral to the anticipatory postural response delayed only the postural EMG (left 96 ms, right 52 ms). CONCLUSIONS: Thus, the associated voluntary and postural responses were delayed independently by stimuli over their respective contralateral motor cortex. SIGNIFICANCE: This suggests that, although timing of responses may be linked by an initial signal, the response from each motor cortex develops independently.     

Modulation of electroencephalographic responses to transcranial magnetic stimulation: evidence for changes in cortical excitability related to movement

Transcranial magnetic stimulation (TMS) and multichannel electroencephalography (EEG) were used for the investigation of cortical excitability preceding voluntary movement in human subjects. The study showed the practical value of the combined TMS-EEG approach in differentiating between cortical and spinal-cord mechanisms, which is difficult with conventional electromyographic measures alone. TMS induced a pronounced negativity (N100) lasting for 150-200 ms, with the amplitude maximum in the stimulated hemisphere. When TMS was applied just before the onset of the visually triggered movement, N100 was markedly attenuated, although motor evoked potentials (MEPs) became larger. We suggest that the N100 component represents an inhibitory response following TMS. This interpretation is in agreement with intracellular recordings in animals, paired-pulse TMS studies and experiments showing increased premovement excitability on the basis of MEPs. N100 was not affected only by the subsequent movement, but also by the switching from rest to the motor-task condition, which caused a slight attenuation of the N100 component; no changes, however, were found in the amplitude of MEPs, suggesting that modified excitability did not affect the output of the corticospinal pyramidal cells. By contrast to MEPs, N100 was modulated also by the presentation of the visual stimulus alone, i.e. when no movement was required. This attenuation suggests that even in a rest condition visual stimuli have an access to the sensorimotor regions of the cortex, most probably through ascending arousal brain systems.     

Corticospinal excitability accompanying ballistic wrist movements in primary dystonia.

Current models of basal ganglia dysfunction in primary dystonia propose that the excessive muscle activity results from an increase in the excitability of the primary motor cortex. Neurophysiological and neuroimaging studies, however, have shown consistently reduced movement-related sensorimotor cortical activity. To explore this paradox, we used transcranial magnetic stimulation (TMS) to examine changes in corticospinal excitability preceding and during ballistic movements of the wrist in 9 patients with primary dystonia affecting the arm and 9 matched control subjects. The onset time, rate of rise, and duration of changes in the excitability of corticospinal projections to the agonist muscle were normal in the patients with dystonia. Increases in excitability were selective to the initial agonist muscle, suggesting that the spatial recruitment of corticospinal neurons was normal. Nonetheless, movements were slower in the patients by an average of 26%. The onset of the first agonist muscle burst was normal in magnitude and timing but the activity in this muscle subsequently became attenuated as movement progressed. Muscle activity in antagonist and proximal muscles of the upper arm was reduced significantly in the dystonia patients. These findings support the view that movement preparation and initiation at the level of the primary motor cortex is normal in patients with dystonia. Bradykinesia could not be attributed to co-contraction or overflow of activity and was associated with reduced rather than excessive muscle activity.     

Effects of force–load on cortical activity preceding voluntary finger movement: Whole-scalp magnetoencephalography of the Bereitschaftsfeld

Neural activity preceding force-loaded voluntary finger movement (the Bereitschaftsfeld) was recorded using 143-channel whole-scalp magnetoencephalography (MEG) in order to determine how the level of force produced during voluntary finger movement is represented in activity over different premovement time intervals localized to different cortical areas. Eighteen healthy subjects performed voluntary right index-finger extension movements against an inertial load of either 0, 100, or 200 g. Results showed that the earliest component of premovement activity, beginning between 1.5 and 1.0 s prior to movement and localized to the central midline around the region of supplementary/cingulate motor areas, was not modulated by the level of force required for movement. However, later premovement activity, occurring between 500 and 200 ms prior to movement onset, was significantly greater for the highest force movements compared with both intermediate (p < 0.05) and no weight-load conditions (p < 0.01). This component was localized to primary sensorimotor cortical areas, with greater source strength on the left side contralateral to movement. Results indicate that, although early premovement activity of the supplementary/cingulate motor areas does not appear to encode movement force, later premovement activity of the primary motor cortex is significantly greater for movements made with more force, not only during movement execution but also up to 500 ms prior in readiness for intended movements of greater force.     

Motor control in simple bimanual movements: a transcranial magnetic stimulation and reaction time study.

OBJECTIVE: Simple reaction time (RT) can be influenced by transcranial magnetic stimulation (TMS) to the motor cortex. Since TMS differentially affects RT of ipsilateral and contralateral muscles a combined RT and TMS investigation sheds light on cortical motor control of bimanual movements. METHODS: Ten normal subjects and one subject with congenital mirror movements (MM) were investigated with a RT paradigm in which they had to move one or both hands in response to a visual go-signal. Suprathreshold TMS was applied to the motor cortex ipsilateral or contralateral to the moving hand at various interstimulus intervals (ISIs) after presentation of the go-signal. EMG recordings from the thenar muscles of both hands were used to determine the RT. RESULTS: TMS applied to the ipsilateral motor cortex shortened RT when TMS was delivered simultaneously with the go-signal. With increasing ISI between TMS and go-signal the RT was progressively delayed. This delay was more pronounced if TMS was applied contralateral to the moving hand. When normal subjects performed bimanual movements the TMS-induced changes in RT were essentially the same as if they had used the hand in an unimanual task. In the subject with MM, TMS given at the time of the go-signal facilitated both the voluntary and the MM. With increasing ISI, however, RT for voluntary movements and MM increased in parallel. CONCLUSIONS: Ipsilateral TMS affects the timing of hand movements to the same extent regardless of whether the hand is engaged in an unimanual or a bimanual movement. It can be concluded, therefore, that in normal subjects simple bimanual movements are controlled by each motor cortex independently. The results obtained in the subject with MM are consistent with the hypothesis that mirror movements originate from uncrossed corticospinal fibres. The alternative hypothesis that a deficit in transcallosal inhibition leads to MM in the contralateral motor cortex is not compatible with the presented data, because TMS applied to the motor cortex ipsilateral to a voluntary moved hand affected voluntary movements and MM to the same extent.     

Phasic stabilization of motor output after auditory and visual distractors

To maintain steady motor output, distracting sensory stimuli need to be blocked. To study the effects of brief auditory and visual distractors on the human primary motor (M1) cortex, we monitored magnetoencephalographic (MEG) cortical rhythms, electromyogram (EMG) of finger flexors, and corticomuscular coherence (CMC) during right‐hand pinch (force 5–7% of maximum) while 1‐kHz tones and checkerboard patterns were presented for 100 ms once every 3.5–5 s. Twenty‐one subjects (out of twenty‐two) showed statistically significant ～20‐Hz CMC. Both distractors elicited a covert startle‐like response evident in changes of force and EMG (～50% of the background variation) but without any visible movement, followed by ～1‐s enhancement of CMC (auditory on average by 75%, P < 0.001; visual by 33%, P < 0.05) and rolandic ～20‐Hz rhythm (auditory by 14%, P < 0.05; visual by 11%, P < 0.01). Directional coupling of coherence from muscle to the M1 cortex (EMG→MEG) increased for ～0.5 s at the onset of the CMC enhancement, but only after auditory distractor (by 105%; P < 0.05), likely reflecting startle‐related proprioceptive afference. The 20‐Hz enhancements occurred in the left M1 cortex and were for the auditory stimuli preceded by an early suppression (by 7%, P < 0.05). Task‐unrelated distractors modulated corticospinal coupling at ～20 Hz. We propose that the distractors triggered covert startle‐like responses, resulting in proprioceptive afference to the cortex, and that they also transiently disengaged the subject's attention from the fine‐motor task. As a result, the corticospinal output was readjusted to keep the contraction force stable. Hum Brain Mapp 36:5168–5182, 2015. © 2015 Wiley Periodicals, Inc.