Dependence of the transcranially induced silent period on the 'instruction set' and the individual reaction time

OBJECTIVES AND METHODS: We looked for influences of the experimental condition on the silent period (SP) from transcranial motor cortex stimulation and analyzed how the instruction given to the subject, as well as the individual reaction time, might affect the duration of the SP in the biceps brachii muscle. RESULTS: The duration of the SP was found to critically depend on the subject's voluntary reaction of the target muscle immediately after the stimulus. With low stimulus intensity and low background force, the duration of the silent period was significantly longer in 10 of 13 subjects (P = 0.002) when they were instructed to relax quickly after the stimulus rather than to maintain the the force at a constant level. A significant shortening of the SP (P = 0.02) was observed when the subjects were instructed to perform a rapid contraction of the target muscle in reaction to the cortical stimulus. With low stimulus intensity and high background force, the same influence of the instruction set was found in 6 of 13 subjects. When the subjects were left without precise instruction, the SP duration was unpredictable. In 10 subjects, the SP corresponded to that obtained with the instruction to maintain the force at a constant level. However, in 3 subjects it was prolonged to the value observed in the 'relax' instruction. With greater stimulus intensities, the effect of the instruction set on the SP duration was generally smaller. A significant prolongation was nevertheless found at low background forces with rapid relaxation (P < 0.001), and a significant shortening was found at high background forces with rapid contraction (P < 0.001) after the stimulus. The SP duration observed with 20% of maximal voluntary contraction (MVC) significantly correlated with the individual reaction time. No such correlation was found for the SP obtained with 80% MVC. The SP was slightly longer at 20% MVC, as compared to 80% MVC within each instruction group. This effect was significant (P < 0.05) at low stimulus intensities. CONCLUSIONS: Therefore, when assessing the SP duration for diagnostic purposes, not only the stimulus intensity but also the background force and the voluntary reaction must be standardized. Furthermore, great stimulus intensities and high background forces should be used to minimise the effects of instruction set and individual reaction time.     

Muscle inactivation: assessment of interpolated twitch technique

The validity, reliability, and protocol for the interpolated twitch technique (ITT) were investigated with isometric plantar flexor and leg extension contractions. Estimates of muscle inactivation were attempted by comparing a variety of superimposed with potentiated evoked torques with submaximal and maximal voluntary contraction (MVC) torques or forces. The use of nerve and surface stimulation to elicit ITT was reliable, except for problems in maintaining maximal stimulation with nerve stimulation at 20 degrees plantar flexion and during leg extension. The interpolated twitch ratio-force relationship was best described by a shallow hyperbolic curve resulting in insignificant MVC prediction errors with second-order polynomials (1.1-6.9%). The prediction error under 40% MVC was approximately double that over 60% MVC, contributing to poor estimations of MVC in non-weight-bearing postimmobilized ankle fracture patients. There was no significant difference in the ITT sensitivity when twitches, doublets, or quintuplets were used. The ITT was valid and reliable when high-intensity contractions were analyzed with a second-order polynomial.     

Task-dependent facilitation of motor evoked potentials during dynamic and steady muscle contractions

Task-dependent differences in the facilitation of motor evoked potentials (MEPs) following cortex stimulation were studied in a proximal (deltoid) and a distal muscle (abductor digiti minimi; ADM) in 23 healthy subjects during both dynamic and steady contractions of the target muscle under isometric and under nonisometric conditions. In the deltoid, MEP amplitudes were significantly greater if stimulation was performed during dynamic contractions than during steady contractions, despite equal background electromyographic levels just prior to the stimulus. The same task-specific extra facilitation of deltoid MEP amplitudes was also found with magnetic stimulation of the brain stem instead of the cortex in 3 subjects. In the ADM, no such task-dependent extra facilitation of MEPs during dynamic contractions was found. It is concluded that in the deltoid, during dynamic contractions, a greater proportion of the spinal motoneurons is close to depolarization threshold (greater "subliminal fringe") whereas the number of firing motoneurons is similar to that during steady contraction. The lack of task-dependent extra facilitation of MEPs in the ADM is explained by the predominant recruitment principle for force gradation in small hand muscles, which is in contrast to the predominant frequency principle used in proximal muscles.     

Crossed and direct effects of digital nerves stimulation on motor evoked potential: a study with magnetic brain stimulation

We studied the influence of contralateral and ipsilateral cutaneous digital nerve stimulation on motor evoked potentials (MEPs) elicited in hand muscles by transcranial magnetic stimulation (TMS). We tested the effect of different magnetic stimulus intensities on MEPs recorded from the thenar eminence (TE) muscles of the right hand while an electrical conditioning stimulus was delivered to the second finger of the same hand with an intensity four times above the sensory threshold. Amplitude decrement of conditioned MEPs as a function of magnetic stimulus intensity was observed. The lowest TMS stimulus intensity produced the largest decrease in conditioned MEPs. Moreover, we investigated the effects of ipsilateral and contralateral electrical digital stimulation on MEPs elicited in the right TE and biceps muscle using an intensity 10% above the threshold. Marked MEP inhibition in TE muscles following both ipsilateral and contralateral digital stimulation is the main finding of this study. The decrease in conditioned MEP amplitude to ipsilateral stimulation reached a level of 50% of unconditioned MEP amplitude with the circular coil and 30% with the focal coil. The amplitude of conditioned MEPs to contralateral digital stimulation showed a decrease of 60% with the circular coil and more than 50% with the focal coil. The onset of the inhibitory effect of contralateral stimulation using the focal coil occurred at a mean of 15 ms later than that of ipsilateral stimulation. No MEP inhibition was observed when recording from proximal muscles. Ipsilateral and contralateral digital stimulation had no effect on F wave at appropriate interstimulus intervals, where the main MEP suppression was noted. We stress the importance of selecting an appropriate test stimulus intensity to evaluate MEP inhibition by digital nerves stimulation. Spinal and cortical sites of sensorimotor integration are adduced to explain the direct and crossed MEP inhibition following digital nerves stimulation.     

Essential dynamics from NMR clusters: dynamic properties of the Myb DNA-binding domain and a hinge-bending enhancing variant

We investigated factors affecting maximal voluntary torque and the assessment of the level of voluntary drive in the elbow flexor muscles. First, the effective compliance of the system was tested by using single, paired, and trains of four stimuli to measure voluntary activation. At high voluntary torques the responses to all these stimuli were identical, suggesting that single stimuli are adequate for estimating voluntary drive. Second, the contribution of torque from synergist elbow flexor muscles was assessed. In attempted maximal voluntary contractions (MVCs), the voluntary activation of brachioradialis (median 91.5%, range 68.9-100%) was lower than for biceps brachii (median 99.1%, range 78.5-100%; P < 0.01). This suggests extra torque may be generated by brachioradialis during elbow flexion, beyond the torque where biceps brachii is maximally activated. Finally, lengthening of the elbow flexors occurred during MVCs, due to slight shoulder movements. This would allow force to increase independently of an increase in voluntary drive.     

Mechanical properties and behaviour of motor units in the tibialis anterior during voluntary contractions

The present work was carried out to analyse the properties and behaviour of Tibialis anterior motor units (MUs) during voluntary contractions in humans. A total of 528 single MU mechanical properties was recorded in 10 subjects by means of the spike-triggered averaging (STA) technique. MU recruitment thresholds and discharge frequencies were recorded during linearly increasing maximal voluntary contraction (MVC). The results indicate a mean (+/- SD) MU torque of 25.5 +/- 21.5 mN.m. and a mean time-to-peak of 45.6 +/- 13.6 ms. A comparison of the average MU twitch torque with that of the muscle allowed an estimate of about 300 MUs in the Tibialis anterior. A positive linear relationship was recorded between the MU twitch torque and the recruitment threshold. The mean minimal and maximal discharge frequencies of MUs were 8.4 +/- 3.0 Hz and 33.2 +/- 14.7 Hz, respectively. The results of the present work indicate that MU behaviour during voluntary contractions is different in the tibialis anterior and in the adductor pollicis.     

Motor unit recruitment during prolonged isometric contractions

Motor unit recruitment patterns were studied during prolonged isometric contraction using fine wire electrodes. Single motor unit potentials were recorded from the brachial biceps muscle of eight male subjects, during isometric endurance experiments conducted at relative workloads corresponding to 10% and 40% of maximal voluntary contraction (MVC), respectively. The recordings from the 10% MVC experiment demonstrated a characteristic time-dependent recruitment. As the contraction progressed both the mean number of motor unit spikes counted and the mean amplitude of the spikes increased significantly (P < 0.01). This progressive increase in spike activity was the result of a discontinuous process with periods of increasing and decreasing activity. The phenomenon in which newly recruited motor units replace previously active units is termed "motor unit rotation" and appeared to be an important characteristic of motor control during a prolonged low level contraction. In contrast to the 10% MVC experiment, there was no indication of de novo recruitment in the 40% MVC experiment. Near the point of exhaustion a marked change in action potential shape and duration dominated the recordings. These findings demonstrate a conspicuous difference in the patterns of motor unit recruitment during a 10% and a 40% MVC sustained contraction. It is suggested that there is a close relationship between intrinsic muscle properties and central nervous system recruitment strategies which is entirely different in fatiguing high and low level isometric contractions.     

EMG power spectrum of respiratory and skeletal muscles during static contraction in healthy man

Changes in EMG power spectrum during isometric voluntary contraction maintained until exhaustion in the range of 20-80% MVC were studied in three skeletal muscles (adductor pollicis or AP, vastus lateralis, and medialis) and two respiratory muscles (diaphragm and rectus abdominis). Quantitative EMG analysis consisted of computation of the median frequency (MF) of power spectra and also the continuous measurement of EMG power in two bands of high (EH) and low (EL) frequencies using bandpass filters. This allowed the calculation of the H/L ratio and its time constant of decay rate (TC delta H/L) throughout the sustained static contraction. The main results were: (1) highly significant, positive correlations between TC delta H/L and the maximal MF changes and also the endurance time to fatigue; (2) EMG changes were determined early, within the first 10-20 s of contraction; and (3) EL always increased throughout the fatiguing isometric contraction, but EH changes markedly varied within the five muscle groups studied. These observations are discussed in terms of the differences in muscle fiber composition and also the variations in motor unit recruitment.     

Ultrasound-guided Tru-cut biopsy of the breast

BACKGROUND: Although H reflexes cannot be reliably recorded from resting human hand intrinsic muscles, a short latency R1 response, thought to be similar to the H reflex, is readily obtained from upper extremity muscles during voluntary contraction. METHODS: The right and left median nerves of 20 normal subjects were repetitively stimulated at 3 Hz at stimulus intensities corresponding to threshold and 20%, 40% and 60% of maximal M response, recording from the abductor pollicis brevis muscle. Studies were done during both minimal and moderate voluntary contraction. RESULTS: The R1 response was present in all subjects at the 40% stimulation intensity level during moderate contraction. The mean latency was 27 ms (SD 1.77 ms) with a good correlation to arm length. The mean amplitude was 1.17 mV (SD 0.79 mV). CONCLUSIONS: Abductor pollicis muscle R1 response can be reliably measured, although latency showed much less intersubject and side to side variability than amplitude. This technique may be useful for the assessment of demyelinative lesions of the inferior segments of the brachial plexus and C8-T1 roots.     

Mechanisms underlying spinal motor neuron excitability during the cutaneous silent period in humans

The transient suppression of muscle contraction during the cutaneous silent period (CSP) could be produced either through postsynaptic inhibition of motoneurons or through presynaptic inhibition of the excitatory inputs to motoneurons that sustain voluntary contraction. We sought to delineate the mechanisms underlying the CSP in hand muscles by measuring changes in H-reflexes and motor-evoked potentials (MEPs) produced by transcranial magnetic stimulation (TMS) during the CSP in 10 healthy volunteers. H-reflexes and MEPs both measure the excitability of the motoneuron pool and activate similar subpopulations of motoneurons through different pathways. Inhibition of H-reflexes and MEPs of similar size was maximal at the midpoint of the CSP and gradually returned to baseline. The similar time course of recovery suggests that the H-reflex and MEP are affected by inhibition at a common site, most likely postsynaptic inhibition of the motoneurons.     

Cortical excitability in patients with essential tremor

We used transcranial magnetic stimulation in 10 patients with essential tremor and 8 matched healthy subjects. A round stimulating coil was placed over the vertex and electromyographic activity was recorded from the first dorsal interosseous muscle. Paired transcranial stimuli were delivered at interstimulus intervals of 3, 5, 20, 100, 150, and 200 ms. The intensity of the conditioning stimulus was 80% of motor threshold at short and 150% at long interstimulus intervals (ISIs). We also measured the silent period obtained after a single magnetic pulse delivered at 150% of motor threshold during a submaximal muscle contraction. Patients and controls had similar motor threshold and similar latencies. Paired magnetic stimuli given at short and long ISIs at rest, and during a voluntary muscle contraction, elicited similar responses in both groups. The silent period evoked by transcranial magnetic stimulation had a similar duration in patients with ET and controls. In conclusion, these findings suggest that patients with essential tremor have normal cortical motor area excitability.     

Task-dependent modulation of inhibitory actions within the primary motor cortex

In 11 healthy subjects motor-evoked potentials (MEPs) and silent periods (SPs) were measured in the right first dorsal interosseus (FDI) and abductor pollicis brevis muscles (APB): (1) when transcranial magnetic cortex stimulation (TMS) was applied at tonic isometric contraction of 20% of maximum force, (2) when TMS was applied during tactile exploration of a small object in the hand, (3) when TMS was applied during visually guided goal-directed isometric ramp and hold finger flexion movements, and (4) when at tonic isometric contraction peripheral electrical stimulation (PES) of the median nerve was delivered at various intervals between PES and TMS. Of the natural motor tasks, duration of SPs of small hand muscles was longest during tactile exploration (APB 205+/-42 ms; FDI 213+/-47 ms). SP duration at tonic isometric contraction amounted to 172+/-35 ms in APB and 178+/-31 ms in FDI, respectively. SP duration in FDI was shortest when elicited during visually guided isometric finger movements (159+/-15 ms). At tonic isometric contraction, SP was shortened when PES was applied at latencies -30 to +70 ms in conjunction with TMS. The latter effect was most pronounced when PES was applied 20 ms before TMS. PES-induced effects increased with increasing stimulation strength up to a saturation level which appeared at the transition to painful stimulation strengths. Both isolated stimulation of muscle afferents and of low-threshold cutaneous afferents shortened SP duration. However, PES of the contralateral median nerve had no effect on SPs. Amplitudes of MEPs did not change significantly in any condition. Inhibitory control of motor output circuitries seems to be distinctly modulated by peripheral somatosensory and visual afferent information. We conclude that somatosensory information has privileged access to inhibitory interneuronal circuits within the primary motor cortex.     

Intracortical inhibition of lower limb motor-evoked potentials after paired transcranial magnetic stimulation

The aim of the present study was to determine the characteristics of intracortical inhibition in the motor cortex areas representing lower limb muscles using paired transcranial magnetic (TMS) and transcranial electrical stimulation (TES) in healthy subjects. In the first paradigm (n=8), paired magnetic stimuli were delivered through a double cone coil and motor evoked potentials (MEPs) were recorded from quadriceps (Q) and tibialis anterior (TA) muscles during relaxation. The conditioning stimulus strength was 5% of the maximum stimulator output below the threshold MEP evoked during weak voluntary contraction of TA (33+/-5%). The test stimulus (67+/-2%) was 10% of the stimulator output above the MEP threshold in the relaxed TA. Interstimulus intervals (ISIs) from 1-15 ms were examined. Conditioned TA MEPs were significantly suppressed (P<0.01) at ISIs of less than 5 ms (relative amplitude from 20-50% of the control). TA MEPs tended to be only slightly facilitated at 9-ms and 10-ms ISIs. The degree of MEP suppression was not different between right and left TA muscles despite the significant difference in size of the control responses (P<0.001). Also, conditioned MEPs were not significantly different between Q and TA. The time course of TA MEP suppression, using electrical test stimuli, was similar to that found using TMS. In the second paradigm (n=2), the suppression of TA MEPs at 2, 3, and 4 ms ISIs was examined at three conditioning intensities with the test stimulation kept constant. For the pooled 2- to 4-ms ISI data, relative amplitudes were 34+/-6%, 61+/-5%, and 98+/-9% for conditioning intensities of 0.95, 0.90, and 0.85x active threshold, respectively (P<0.01). In conclusion, the suppression of lower limb MEPs following paired TMS showed similar characteristics to the intracortical inhibition previously described for the hand motor area.     

Effect of radial shortening on muscle length and moment arms of the wrist flexors and extensors

Manual muscle test scores (MMTS) and maximum voluntary contraction (MVC) force measurements were made from triceps brachii muscles of 70 individuals with chronic cervical spinal cord injury (SCI). Both MMTS and strength assessments showed that asymmetrical motor deficits were common. Muscles with MMTS of 3 generated an average of nine percent MVC force produced by control muscles. In this SCI population, little residual voluntary force is apparently needed for triceps brachii to work against gravity. Only 24 percent of muscles tested had this strength, however, indicating the need to develop strategies to alleviate this muscle weakness. MMTS and force were related positively but each MMTS was not associated with a unique force range. MVC force generating capacity is therefore only one factor that determines whether or not a muscle can work with or against gravity and against resistance.     

A case of cortical reflex positive-negative myoclonus--electrophysiological study

An 11-year-old girl who had the positive-negative myoclonus and the history of the generalized tonic clonic seizure was electrophysiologically studied. She had no siblings with either myoclonus or epilepsy, and her intellectual level was normal. She had no other neurological deficits including ataxia, pyramidal and extrapyramidal signs. Surface EMG showed a brief increase in the EMG activity followed by the silent period associated with positive and negative myoclonus during sustained wrist extension. Giant SEP and C reflex (38.6 ms) following electric stimulation of the median nerve at the wrist were obtained in the resting condition and the silent period (about 180 ms) following C reflex was obtained during voluntary contraction. Jerk-locked back averaging of the EEG time-locked to the onset of the myoclonic discharge recorded from the right biceps muscle showed a cortical spike at the left central region preceding the myoclonus onset by 12.6 ms. The latency of C reflex in this case was very short compared with that of previously reported cortical reflex myoclonus. The estimated cortical delay between the arrival of the somatosensory volley and the motor cortex discharge responsible for the C reflex was -1.0 ms and this value was shorter than that in patients with typical cortical reflex myoclonus (mean 3.7 +/- 1.1 ms). Conditioning stimuli (C) of the right median nerve at the wrist started to facilitate the amplitude of the motor evoked potential recorded from the right abductor pollicis brevis muscle after magnetic test stimuli (T) of the left motor cortex at 20 ms of the C-T interval. This C-T interval was shorter than that (24.6 +/- 1.6 ms) in patients with the typical cortical myoclonus. These electrophysiological findings suggested the shorter reflex pathway of the cortical reflex myoclonus in this case than in typical cortical reflex myoclonus. We speculated that the myoclonus was based upon the direct sensory projection from the thalamus to the motor cortex in this case.     

Acute leg ischaemia--a case for the junior surgeon?

To study the regulation of microvascular blood flow in a compartment muscle, laser-Doppler measurements of muscle microcirculation were recorded in the supraspinatus muscle in eight volunteers during and following submaximal isometric muscle contractions. The subjects performed isometric shoulder abductions at five contraction levels from 5% to 50% maximal voluntary contraction for 1 min each and a sustained 30 degrees shoulder abduction for 20 min. The subjects' perceived exertion increased from "no perceived exertion" to "near maximal exertion" during the 20-min period with 30 degrees shoulder abduction. Microcirculation increased during all 1-min contractions. Following the contractions at 20%, 30% and 50% MVC post-exercise reactive hyperaemia was seen for a period of at least 1 min. The reactive hyperaemia increased in magnitude in response to increasing contraction level. The results showed the same time-history of the blood flow at microvascular level as previously seen in larger peripheral vessels in response to muscle contractions. During the 20-min contraction microcirculation increased in line with the findings during the brief contractions. However, in contrast to the brief contractions no postexercise reactive hyperaemia occurred following the prolonged contraction. Lack of postexercise reactive hyperaemia following the prolonged shoulder abduction would suggest insufficient regulation of the vascular resistance. Alternatively, lack of hyperaemia could be taken as an indication of sufficient microcirculation during the preceding contraction. From previous studies on intramuscular pressure and metabolism the latter alternative would seem unlikely.     

Sexually dimorphic effects of anti-NGF treatment in neonatal rats

To investigate the sensitivity to changes in excitability of motoneuron pool dependent on voluntary motor commands, we recorded motor evoked potentials (MEPs) and H-reflexes from the right flexor carpi radialis (FCR) muscle of normal human subjects. Amplitudes of MEPs were always larger than those of the H-reflex in both tonic and phasic muscle contractions. Furthermore, amplitudes of MEP and H-reflex were larger in phasic than in tonic muscle contraction. These results indicate that there are differences in the sensitivity to changes in motoneuronal excitability related to the production of excitatory postsynaptic potentials for H-reflex and MEP responses, respectively.     

Motor evoked potentials: appropriate positioning of recording electrodes for diagnosis of spinal disorders

The interpretation of normal and pathological findings of motor evoked potential obtained by the use of transcranial magnetic stimulation depends on adequate examination technique, including the appropriate positioning of the recording electrodes over the muscle. On the basis of knowing the location of the motor end plate zones in muscles, magnetic stimulation of the motor cortex of 30 healthy adults was performed in order to explore the influence of the position of the surface recording electrodes on potential parameters and to establish the standard location of the recording electrodes over the biceps brachii, medial vastus, anterior tibial and abductor hallucis muscles for diagnostic use in spine disorders. The cortical latencies and peak-to-peak amplitudes of the evoked potentials were analysed by varying the location of the recording electrodes and the stimulus intensities. The latencies were significantly shorter when the different electrode lay more proximally over the muscle belly. Reproducible potentials with sharp negative onset and maximum amplitude were recorded with a separation of 5-7.5 cm between the different electrode, located over the motor end plate area, and the different electrode, located over the distal myotendinous junction. This implies that the parameters of evoked potentials depend on the position and separation distance of the recording electrodes over the muscles and that it is possible to record the potentials using a lower stimulus intensity and, above all, on relaxed muscles, which may prove to be applicable for intraoperative monitoring of the spinal cord using magnetic stimulation.(ABSTRACT TRUNCATED AT 250 WORDS)     

Motor-unit behavior in humans during fatiguing arm movements

1. The activity of 40 triceps brachii motor units was recorded from the dominant arms of 9 healthy adult volunteers (age 27.8 +/- 4.4 yr, mean +/- SD) during a fatigue task that included both isometric and anisometric contractions. The fatigue task lasted 8.3 min and consisted of 50 extension and 50 flexion movements of the elbow. Each movement (40 degrees in 0.8s) was separated by an isometric contraction. A constant load resisting extension of 17.7 +/- 3.0% of maximal voluntary contractions (MVC) was applied throughout the task. This paradigm enabled the direct contrast of motor-unit discharge behavior during the different types of fatiguing contractions. 2. Motor-unit behavior was examined to determine the relative contribution of two mechanisms for optimizing force production under fatiguing conditions: recruitment of motor units and modulation of motor-unit discharge following recruitment. Threshold torques for motor-unit recruitment thresholds were determined by ramp-and-hold isometric contractions. Motor-unit discharge was evaluated during the fatigue task by contrasting the number of motor-unit potentials (spikes) per contraction for concentric eccentric, and isometric contractions. 3. The fatigue task resulted in a 30 +/- 12% decline in the mean MVC of elbow extension. Recruitment of nine new motor units (23%) was evident during the fatiguing extension movements, often within five to seven movements (i.e., within 25-35 s). Each newly recruited motor unit had the largest recruitment threshold torque in that experiment. 4. Analysis of the motor units that were active from the beginning of the fatigue task revealed that the mean number of motor-unit spikes per contraction increased, or remained constant as fatigue ensued, yet for the majority of motor units it increased or remained constant. None of the newly recruited motor units demonstrated decreased number of mean spikes per contraction after recruitment. Further, concurrently active motor units displayed different discharge behavior in two-thirds of the subjects. It is proposed that if the neural drive to the muscle is distributed uniformly upon the motoneuron pool, peripheral feedback from the exercising muscle may modulate specific motoneuron discharge levels during fatigue.     

Studies on the corticospinal control of human walking. I. Responses to focal transcranial magnetic stimulation of the motor cortex

Experiments were done to determine the extent to which the corticospinal tract is linked with the segmental motor circuits controlling ankle flexors and extensors during human walking compared with voluntary motor tasks requiring attention to the level of motor activity. The motor cortex was activated transcranially using a focal magnetic stimulation coil. For each subject, the entire input-output (I-O) curve [i.e., the integral of the motor evoked-potential (MEP) versus stimulus strength] was measured during a prescribed tonic voluntary contraction of either the tibialis anterior (TA) or the soleus. Similarly, I-O curves were measured in the early part of the swing phase, or in the early part of the stance phase of walking. The I-O data points were fitted by the Boltzmann sigmoidal function, which accounted for >/=80% of total data variance. There was no statistically significant difference between the I-O curves of the TA measured during voluntary ankle dorsiflexion or during the swing phase of walking, at matched levels of background electromyographic (EMG) activity. Additionally, there was no significant difference in the relation between the coefficient of variation and the amplitude of the MEPs measured in each task, respectively. In comparison, during the stance phase of walking the soleus MEPs were reduced on average by 26% compared with their size during voluntary ankle plantarflexion. Furthermore, during stance the MEPs in the inactive TA were enhanced relative to their size during voluntary ankle plantarflexion and in four of six subjects the TA MEPs were larger than those of the soleus. Finally, stimulation of the motor cortex at various phases of the step cycle did not reset the cycle. The time of the next step occurred at the expected moment, as determined from the phase-resetting curve. One interpretation of this result is that the motor cortex may not be part of the central neural system involved in timing the motor bursts during the step cycle. We suggest that during walking the corticospinal tract is more closely linked with the segmental motor circuits controlling the flexor, TA, than it is with those controlling the extensor, soleus. However, during voluntary tasks requiring attention to the level of motor activity, it is equally linked with the segmental motor circuits of ankle flexors or extensors.