Effectiveness of electrical noise in reducing postural sway: a comparison between imperceptible stimulation applied to the anterior and to the posterior leg muscles

Purpose

The aim of this study was to investigate whether subsensory electrical noise stimulation applied over the tibialis anterior (TA) muscles may reduce postural oscillations during quiet stance and to compare such reductions with those obtained by applying the noise stimulation over the triceps surae (TS) muscles. The rationale relies on the use of noise stimulation to enhance the sensitivity of sensory receptors, thereby improving the performance of the postural control system. As recent evidence suggested that the TA muscle might be a better source of proprioceptive information during quiet stance than the TS muscles, the main hypothesis of the present study was that subsensory noise stimulation applied to the TA muscles should be more efficient for postural stabilization than a similar stimulation to the TS.

Methods

Participants had their postural steadiness assessed while subsensory electrical noise was applied bilaterally to their TA or TS muscles and also while the stimulation was applied bilaterally to both TA and TS muscles. No stimulation was delivered in the control condition.

Results

Time-domain and frequency-domain parameters based on center of mass and center of pressure signals were significantly reduced when noise stimulation was applied to the anterior and/or to the posterior leg muscles. No consistent differences in postural sway parameters were observed among the stimulation conditions.

Conclusions

This is the first investigation showing that subsensory stimulation applied over the TA muscles alone is effective in attenuating postural sway, with TS stimulation being equally effective. These findings may have useful applications for the development of medical/rehabilitation devices designed to improve postural steadiness in people with balance impairments.     

Subthreshold electrical stimulation reduces motor unit discharge variability and decreases the force fluctuations of plantar flexion

The purpose of this study was to examine the influence of subthreshold electrical stimulation on the force fluctuations and motor-unit discharge variability during low-level, steady contraction of the plantar flexor muscles. Seven subjects performed a force-matching task of isometric plantar flexion at 5% of maximal voluntary contraction with and without random electrical stimulation applied to the tibial nerve. During the task, the motor unit action potential was continuously recorded with fine-wire electrodes, and the inter-spike intervals of a single motor unit were calculated. The coefficient of variation (CV) of the force fluctuations and the inter-spike intervals of the motor unit discharge were significantly decreased by the intervention of subthreshold electrical stimulation, although there were no changes in the mean values. These results suggest that subthreshold stimulation reduced the motor-unit discharge variability, which in turn, increased the steadiness of the force.     

Effects of 20-day bed rest with and without strength training on postural sway during quiet standing

Aim: To examine the effect of unweighting as a possible contributory factor to a reduced calf muscle volume on postural sway during quiet standing, changes in postural sway following bed rest with or without strength training were investigated. Methods: Twelve young men participated in a 20‐day bed‐rest study. Subjects were divided into a non‐training group (BR‐Con) and a strength training group (BR‐Tr). For the BR‐Tr group, training was comprised of dynamic calf‐raise and leg‐press exercises to maintain the muscle volume of the plantar flexors. Before and after bed rest, subjects maintained quiet standing in a barefoot position on a force platform with their eyes open or closed. During the quiet stance, foot centre‐of‐pressure (CoP) and the mean velocity of CoP was calculated. Muscle volume of the plantar flexors was computed using axial magnetic resonance images of the leg. Results: After the bed‐rest period, the muscle volume decreased in the BR‐Con group but not in the BR‐Tr group. The mean velocity of CoP as an assessment of postural sway, however, increased in both groups. These results indicate that the strength training during bed rest cannot counteract the increase in postural sway. Conclusion: We concluded that postural sway increases following 20 days of bed rest despite maintenance of the muscle volume of plantar flexors as the main working muscles for the human postural standing.     

Evidence for reduced efficacy of the Ia-pathway during shortening plantar flexions with increasing effort

To determine whether the soleus (SOL) H-reflex is modulated during shortening contractions in a manner that has been observed for isometric contractions, SOL H-reflexes and M-waves were elicited via percutaneous electrical stimulation to the tibial nerve at an intensity that evoked an H-reflex at 50% of its maximum in 11 healthy subjects. Paired electrical stimuli were delivered as the ankle angle passed through 90° at an interval of 400 ms while the subject performed shortening contractions at levels of plantar flexion torque ranging between 2 and 30% of that during a maximal voluntary contraction (MVC). H-reflexes were also recorded during the performance of isomeric contractions of plantar flexors at similar levels of plantar flexion torque and at the same joint angle (muscle length) in an additional five healthy subjects. Correlations were examined between the peak-to-peak amplitude of the first H-reflexes, M-waves and plantar flexion torques in both protocols. It was revealed that no significant correlation was found between the SOL H-reflex and increasing plantar flexion torque during shortening contractions (ρ = −0.07, P = 0.15), while a strong positive correlation was observed for the isometric conditions (ρ = 0.99, P < 0.01). No significant change was observed in the SOL M-wave for either contraction type. Furthermore, the H-reflexes elicited via paired stimuli with the same background activity in voluntary shortening contractions showed almost identical amplitudes, suggesting that the level of homosynaptic post-activation depression did not change in response to the varying levels of activation in voluntary shortening contractions. Therefore, the lack of increase in the H-reflex during shortening contractions at increasing intensities is possibly due to a centrally regulated increase in presynaptic inhibition. Such a downward modulation of the reflex suggests that Ia-excitatory input onto the SOL motoneurone pool needs to be reduced during the performance of shortening contractions.     

TENS to the posterior aspect of the legs decreases postural sway during stance

The purpose of this study was to examine the effect of Transcutaneous Electrical Nerve Stimulation (TENS) applied to the posterior aspect of the legs, on postural sway during stance. Thirty healthy subjects were tested while standing on a force platform under four stimulation conditions: no TENS, bilateral TENS, and unilateral left and right TENS. Thirty-second long tests, employing detection threshold amplitudes, were performed in three blocks. In each block, the four conditions were applied both with and without vision in a random order. The results indicate that the application of TENS brought about a decrease in postural sway as expressed by average sway velocity, in addition to a decrease in the absolute values of maximal and minimal medio-lateral and anterior-posterior velocity. Thus, similar to sub-threshold random electrical noise, it appears that the application of low-amplitude TENS to the lower limbs decreases postural sway during stance. Considering the ease of TENS application and the high prevalence of postural disorders, the potential clinical significance of this observation is to be determined by further studies.     

An enhanced level of motor cortical excitability during the control of human standing

Aim: The study examined the role of the motor cortex in the control of human standing. Methods: Subjects (n = 15) stood quietly with or without body support. The supported standing condition enabled subjects to stand with a reduced amount of postural sway. Peripheral electrical stimulation, transcranial magnetic stimulation (TMS) or transcranial electrical stimulation (TES) was applied to elicit a soleus (SOL) H‐reflex, or motor‐evoked potentials (MEPs) in the SOL and the tibialis anterior (TA). Trials were grouped based on the standing condition (i.e. supported vs. normal) as well as sway direction (i.e. forward and backward) while subjects were standing normally. Results: During normal when compared to supported standing, the SOL H‐reflex was depressed (?11 ± 4%), while the TMS‐evoked MEPs from the SOL and TA were facilitated (35 ± 11% for the SOL, 51 ± 15% for the TA). TES‐evoked SOL and TA MEPs were, however, not different between the normal and supported standing conditions. The data based on sway direction indicated that the SOL H‐reflex, as well as the SOL TMS‐ and TES‐evoked MEPs were all greater during forward when compared to backward sway. In contrast, the TMS‐ and TES‐evoked MEPs from the TA were smaller when swaying forward as compared to backward. Conclusions: The results indicated the presence of an enhanced cortical excitability because of the need to control for postural sway during normal standing. The increased cortical excitability was, however, unlikely to be involved in an on‐going control of postural sway, suggesting that postural sway is controlled at the spinal and/or subcortical levels.     

The force–velocity relationship of the human soleus muscle during submaximal voluntary lengthening actions

In experiments on isolated animal muscle, the force produced during active lengthening contractions can be up to twice the isometric force, whereas in human experiments lengthening force shows only modest, if any, increase in force. The presence of synergist and antagonist muscle activation associated with human experiments in situ may partly account for the difference between animal and human studies. Therefore, this study aimed to quantify the force–velocity relationship of the human soleus muscle and assess the likelihood that co-activation of antagonist muscles was responsible for the inhibition of torque during submaximal voluntary plantar flexor efforts. Seven subjects performed submaximal voluntary lengthening, shortening(at angular, velocities of +5, –5, +15, –15 and +30, and –30° s^–1) and isometric plantar flexor efforts against an ankle torque motor. Angle-specific (90°) measures of plantar flexor torque plus surface and intramuscular electromyography from soleus, medial gastrocnemius and tibialis anterior were made. The level of activation (30% of maximal voluntary isometric effort) was maintained by providing direct visual feedback of the soleus electromyogram to the subject. In an attempt to isolate the contribution of soleus to the resultant plantar flexion torque, activation of the synergist and antagonist muscles were minimised by: (1) flexing the knee of the test limb, thereby minimising the activation of gastrocnemius, and (2) applying an anaesthetic block to the common peroneal nerve to eliminate activation of the primary antagonist muscle, tibialis anterior and the synergist muscles, peroneus longus and peroneus brevis. Plantar flexion torque decreased significantly (P<0.05) after blocking the common peroneal nerve which was likely due to abolishing activation of the peroneal muscles which are synergists for plantar flexion. When normalised to the corresponding isometric value, the force–velocity relationship between pre- and post-block conditions was not different. In both conditions, plantar flexion torques during shortening actions were significantly less than the isometric torque and decreased at faster velocities. During lengthening actions, however, plantar flexion torques were not significantly different from isometric regardless of angular velocity. It was concluded that the apparent inhibition of lengthening torques during voluntary activation is not due to co-activation of antagonist muscles. Results are presented as mean (SEM).     

A new paradigm of electrical stimulation to enhance sensory neural function

The ability to improve peripheral neural transmission would have significant therapeutic potential in medicine. A technology of this kind could be used to restore and/or enhance sensory function in individuals with depressed sensory function, such as older adults or patients with peripheral neuropathies. The goal of this study was to investigate if a new paradigm of subsensory electrical noise stimulation enhances somatosensory function. Vibration (50 Hz) was applied with a Neurothesiometer to the plantar aspect of the foot in the presence or absence of subsensory electrical noise (1/f type). The noise was applied at a proximal site, on a defined region of the tibial nerve path above the ankle. Vibration perception thresholds (VPT) of younger adults were measured in control and experimental conditions, in the absence or presence of noise respectively. An improvement of ∼16% in VPT was found in the presence of noise. These are the first data to demonstrate that modulation of axonal transmission with externally applied electrical noise improves perception of tactile stimuli in humans.     

Compensation for intrinsic muscle stiffness by short-latency reflexes in human triceps surae muscles

The incremental torque resisting rotation of the foot about the ankle joint was studied in normal seated subjects. Prior to each rotation, subjects were required to activate triceps surae (TS) muscles and maintain a constant plantar flexion torque (range 6-14 N X m) on a platform whose position was controlled by a torque motor. Subjects were instructed to increase torque as rapidly as possible once rotation commenced. Rotations ranged from 0.5 to 14 degrees amplitude and from 20 to 300 degrees/s maximum velocity. The torque in response to rotations stretching TS muscles and releasing tibialis anterior (TA) muscles increased steeply and then rapidly decreased with stretch velocity. At approximately 60 ms from stretch onset, the torque reduction terminated, torque then increased again until it began to level off at approximately 120 ms. A further large increase in torque occurred at 180 ms. A burst of short-latency (SL) electromyographic (EMG) activity in soleus (SOL) commenced at 40 ms, and was followed by a second burst at approximately 68 ms, provided that stretch deceleration started later than 20 ms after stretch onset. A period of sustained EMG activity in SOL commenced at approximately 130 ms (long-latency (LL) activity). Incremental torque in response to stretch of TA and release of TS muscles initially showed a step decrease followed by a reversal of the torque trajectory back toward base line. This change was arrested at 60 ms and torque then remained approximately constant until a large increase in torque at 180 ms. Ischemia was used to reduce SL EMG reflexes without significantly modifying the background EMG activity. A comparison between torque curves under control and ischemic conditions indicated that SL EMG activity in TS muscles recruited the force responsible for terminating the torque reduction coincident with decreasing stretch velocity. The torque response prior to the onset of force recruited by SL activity was attributed to the intrinsic properties of active muscle fibers. Thereafter, until the onset of LL activity, the torque response was attributed to intrinsic and reflex-recruited force. Torque in these two time periods was compared under a variety of stretch conditions in order to test the hypothesis that force recruited by segmental reflexes compensates for the non-linear stretch properties of active TS muscles. The relationships of SL EMG amplitudes and areas to stretch velocity and acceleration were also examined.(ABSTRACT TRUNCATED AT 400 WORDS)     

Cutaneous mechanisms of isometric ankle force control

The sense of force is critical in the control of movement and posture. Multiple factors influence our perception of exerted force, including inputs from cutaneous afferents, muscle afferents and central commands. Here, we studied the influence of cutaneous feedback on the control of ankle force output. We used repetitive electrical stimulation of the superficial peroneal (foot dorsum) and medial plantar nerves (foot sole) to disrupt cutaneous afferent input in 8 healthy subjects. We measured the effects of repetitive nerve stimulation on (1) tactile thresholds, (2) performance in an ankle force-matching and (3) an ankle position-matching task. Additional force-matching experiments were done to compare the effects of transient versus continuous stimulation in 6 subjects and to determine the effects of foot anesthesia using lidocaine in another 6 subjects. The results showed that stimulation decreased cutaneous sensory function as evidenced by increased touch threshold. Absolute dorsiflexion force error increased without visual feedback during peroneal nerve stimulation. This was not a general effect of stimulation because force error did not increase during plantar nerve stimulation. The effects of transient stimulation on force error were greater when compared to continuous stimulation and lidocaine injection. Position-matching performance was unaffected by peroneal nerve or plantar nerve stimulation. Our results show that cutaneous feedback plays a role in the control of force output at the ankle joint. Understanding how the nervous system normally uses cutaneous feedback in motor control will help us identify which functional aspects are impaired in aging and neurological diseases.     

Experimental knee pain impairs postural stability during quiet stance but not after perturbations

The objective of this study was to examine the effect of experimental knee-related pain on postural control. Twelve healthy subjects stood as quietly as possible on a movable force platform (that measured the centre of pressure and provided fast perturbations) before, during, and after experimental knee-related pain. Lower limb electromyographic (EMG) activity and joint angles were measured. Experimental pain was induced by injecting hypertonic saline into the infrapatellar fat pad (unilateral and bilateral) and isotonic saline was used for control sessions. Compared with the baseline and control sessions, unilateral and bilateral knee-related pain during quiet standing evoked (1) an increased sway displacement in the anterior-posterior direction (P < 0.05), (2) larger knee flexion (P < 0.05), and (3) larger EMG changes. Bilateral pain also induced (1) larger medial-lateral sway displacement and speed (P < 0.05) and (2) larger left hip flexion (P < 0.05). During forward perturbations, subjects leaned forward during both painful conditions when compared with baseline (P < 0.05). The additional impairment by bilateral pain suggests that the non-painful limb in unilateral pain conditions compensates for the impaired postural control. These results show that knee-related pain impairs postural stability during quiet standing, indicating the vulnerability of patients with knee pain to falls. This measure could potentially help clinicians who seek to assess how pain responses may contribute to patient's postural control and stability during quiet standing.     

Contribution of muscle afferents to prolonged flexion withdrawal reflexes in human spinal cord injury

The contribution of force-sensitive muscular afferents to prolonged flexion withdrawal reflexes, or flexor spasms, after human spinal cord injury (SCI) was investigated. In three separate experimental conditions, flexion reflexes were triggered in subjects with SCI using trains of electrocutaneous stimuli delivered at the foot and lower leg and compared with reflexes elicited via intramuscular (i.m.) electrical stimuli. In the first experiment, flexion reflexes were elicited using i.m. stimuli to the tibialis anterior (TA) in the majority of subjects tested. The ratio of peak isometric ankle to hip torques during i.m.-triggered reflexes were proportionally similar to those evoked by electrocutaneous foot or shank stimulation, although the latency to onset and peak flexion torques were significantly longer with i.m. stimulation. In the second experiments, the amplitude and frequency of i.m. TA stimulation were varied to alter the stimulus-induced muscle torque. Peak ankle and hip torques generated during the flexion reflex responses were correlated to a greater extent with stimulus-induced muscle torques as compared with the modulated stimulus parameters. In the third experimental series, i.m. stimuli delivered to the gastrocnemius (GS) elicited flexion reflexes in approximately half of the subjects tested. The combined data indicate a potentially prominent role of the stimulus-induced muscle contraction to the magnitude and latency of flexor reflex behaviors after i.m. TA stimulation. Results after i.m. GS stimulation indicate multi-joint flexion reflexes can also be elicited, although to a lesser extent than i.m. TA stimulation.     

Stance and sensory feedback influence on postural dynamics

This study examined the effects of ice-induced plantar desensitization and the withdrawal of visual feedback on the magnitude and time-dependent structure of postural sway variability. The magnitude of variability was quantified as the area of an ellipse enclosing 95% of the center of pressure (COP) time-series during normal and tandem stances. The same time-series were also analyzed using Approximate Entropy (ApEn) and Cross-Approximate Entropy (CrossApEn) as indices of irregularity and asynchrony between the mediolateral and anteroposterior COP motions. Variability increased during tandem stance and this increase was compounded by both visual feedback withdrawal and cutaneous desensitization. Both ApEn (mediolateral and anteroposterior COP motion) and CrossApEn increased with the withdrawal of visual feedback during the tandem stance, but decreased significantly during normal stance. The results of the study demonstrate that plantar desensitization only affected the magnitude of sway variability but did not alter its time-dependent structure. Contrasting effects on the structure of postural sway variability with visual feedback withdrawal were observed during the different stances, highlighting the role of task demands in postural dynamics.     

Directional postural responses induced by vibrotactile stimulations applied to the torso

It has been shown that torso-based vibrotactile feedback significantly reduces postural sway in balance-compromised adults during quiet standing and in response to perturbations. This study aimed to determine whether vibrotactile stimulations applied to different torso locations induced directional postural responses and whether torso cutaneous information contributes to body representation. Eleven healthy young adults equipped with an inertial measurement unit (IMU) placed on the torso were asked to maintain an upright posture with closed eyes. Six vibrators (tactors) were placed on the torso in contact with the skin over the left and right external oblique, internal oblique, and erector spinae muscles at the L4/L5 level. Each tactor was randomly activated four times per location at a frequency of 250 Hz for a period of 5 s. The IMU results indicated that vibration applied individually over the internal oblique and erector spinae muscles induced a postural shift of about one degree oriented in the direction of the stimulation, while simultaneous activation of all tactors and activation of tactors over external oblique muscles produced insignificant postural effects. The root mean square of the sway signal was significantly higher during vibration than before or after. However, the center of pressure displacement, measured by a force plate, was uninfluenced by any vibration. These results suggest a multi-joint postural response including a torso inclination associated with vibration-induced changes in cutaneous information. The directional aspect of vibration-induced postural shifts suggests that cutaneous information from the stimulated areas contributes to proprioception and upper body spatial representation.     

Control of the triceps surae during the postural sway of quiet standing

AIM: The present study investigated how the triceps surae are controlled at the spinal level during the naturally occurring postural sway of quiet standing. METHODS: Subjects stood on a force platform as electrical stimuli were applied to the posterior tibial nerve when the center of pressure (COP) was either 1.6 standard deviations anterior (COP(ant)) or posterior (COP(post)) to the mean baseline COP signal. Peak-to-peak amplitudes of the H-reflex and M-wave from the soleus (SOL) and medial gastrocnemius (MG) muscles were recorded to assess the efficacy of the Ia pathway. RESULTS: A significant increase in the H(max) : M(max) ratio for both the SOL (12 +/- 6%) and MG (23 +/- 6%) was observed during the COP(ant) as compared to the COP(post) condition. The source of the modulation between COP conditions cannot be determined from this study. However, the observed changes in the synaptic efficacy of the Ia pathway are unlikely to be simply a result of an altered level of background electromyographic activity in the triceps surae. This was indicated by the lack of differences observed in the H(max) : M(max) ratio when subjects stood without postural sway (via the use of a tilt table) at two levels of background activity. CONCLUSIONS: It is suggested that the phase-dependent modulation of the triceps surae H-reflexes during the postural sway of quiet standing functions to maintain upright stance and may explain the results from previous studies, which, until now, had not taken the influence of postural sway on the H-reflex into consideration.     

Fingertip contact influences human postural control

Touch and pressure stimulation of the body surface can strongly influence apparent body orientation, as well as the maintenance of upright posture during quiet stance. In the present study, we investigated the relationship between postural sway and contact forces at the fingertip while subjects touched a rigid metal bar. Subjects were tested in the tandem Romberg stance with eyes open or closed under three conditions of fingertip contact: no contact, touch contact (< 0.98 N of force), and force contact (as much force as desired). Touch contact was as effective as force contact or sight of the surroundings in reducing postural sway when compared to the no contact, eyes closed condition. Body sway and fingertip forces were essentially in phase with force contact, suggesting that fingertip contact forces are physically counteracting body sway. Time delays between body sway and fingertip forces were much larger with light touch contact, suggesting that the fingertip is providing information that allows anticipatory innervation of musculature to reduce body sway. The results are related to observations on precision grip as well as the somatosensory, proprioceptive, and motor mechanisms involved in the reduction of body sway.     

Fingertip contact influences human postural control

Touch and pressure stimulation of the body surface can strongly influence apparent body orientation, as well as the maintenance of upright posture during quiet stance. In the present study, we investigated the relationship between postural sway and contact forces at the fingertip while subjects touched a rigid metal bar. Subjects were tested in the tandem Romberg stance with eyes open or closed under three conditions of fingertip contact: no contact, touch contact (<0.98 N of force), and force contact (as much force as desired). Touch contact was as effective as force contact or sight of the surroundings in reducing postural sway when compared to the no contact, eyes closed condition. Body sway and fingertip forces were essentially in phase with force contact, suggesting that fingertip contact forces are physically counteracting body sway. Time delays between body sway and fingertip forces were much larger with light touch contact, suggesting that the fingertip is providing information that allows anticipatory innervation of musculature to reduce body sway. The results are related to observations on precision grip as well as the somatosensory, proprioceptive, and motor mechanisms involved in the reduction of body sway.     

Effect of prior isometric muscle action on concentric torque output during plantar flexion

The influence of different levels of prior isometric muscle action on the concentric torque output during plantar flexion was examined at two angular velocities (60°·s^–1 and 120°·s^–1) in ten healthy female subjects. The levels of the prior muscle actions were 25%, 50%, 75% and 100% of the maximal voluntary isometric contraction (MVIC). A KINetic-COMmunicator II dynamometer was used to measure torque output during plantar flexion within a range of motion of 78°-120° of the ankle joint. Simultaneous recordings of electromyograms (low-pass filtered and rectified) were obtained from the gastrocnemius medialis muscle and the soleus muscle. Torque-angle curves were made for the plantar flexions using different prior muscle actions. Up to 75% of MVIC, the torque output in the first part of the range of motion increased with the level of the prior isometric muscle action; at higher levels of MVIC the torque did not appear to increase any further. Later in the range of motion, after 24° in the plantar flexion at a velocity of 60°·s^–1 and 31° at 120°·s^–1, the prior muscle actions had no further influence. No increase was found in the electromyograms, with one exception, during the concentric movements when preceded by higher levels of MVIC. It would seem therefore that the increase in torque output early in the range of motion cannot be explained on the basis of differences in electrical muscle activation in this study.     

Postural control strategy during standing is altered in patients with multiple sclerosis

Disturbances in balance are one of the first reported symptoms of Multiple Sclerosis (MS), yet limited research has been performed to classify the postural control deficits in this population. This study investigated the variability present in the sway patterns during quiet standing in patients with MS (PwMS) and healthy controls. Subjects were assessed (eyes open, closed) standing on a force platform. Variability of the sway patterns was quantified using a measure of amount of variability (root mean square; RMS) and two measures of temporal structure of variability (Lyapunov Exponent – LyE; Approximate Entropy – ApEn). RMS results revealed significantly higher amount of variability in the sway patterns of PwMS. PwMS also exhibit increased regularity (decreased ApEn) and decreased divergence (decreased LyE) during standing compared to healthy controls. Removing vision resulted in significantly decreased divergence (decreased LyE) in the MS subject group. These changes in the temporal structure correspond well with the theoretical model of the optimal movement variability hypothesis and the results support using variability measures to understand the mechanisms that underline postural control in PwMS and possibly other neurodegenerative disease pathologies.     

Phase-dependent reversal of reflexly induced movements during human gait

Summary To investigate whether phase-dependent reversals in reflex responses on electromyography (EMG) are accompanied by movement reversals, a series of human volunteers were studied for their behavioural responses to sural nerve stimulation during running or walking on a treadmill. Low-intensity stimulation (< 2.5 x perception threshold, T) of the sural nerve yielded facilitatory responses in the tibialis anterior muscle (TA), correlated with an induced ankle dorsiflexion (mean maximum 4°) in early swing. The same stimuli yielded primarily TA suppression and weak ankle plantar flexion (mean maximum 1°) at end swing. The correlated induced knee angle changes did not precede the ankle changes, and they were relatively small. Mean maximum flexion in early swing was 6.2°, while mean maximum extension was 3.7°. High-intensity stimulation of the sural nerve (> 2.5 x T) always gave rise to suppression of the ongoing activity. This resulted in a second type of movement reversal. During late stance and early swing the responses in TA were suppressive (i.e. below background activity) and related to ankle plantar flexion. In contrast, the responses during early and middle stance consisted of suppression in extensor activity (gastrocnemius medialis and soleus) and ankle dorsiflexion.The data are discussed in terms of a new hypothesis, which states that the responses to electrical stimulation of cutaneous nerves during locomotion do not correspond directly to corrections for stumbling following mechanical perturbations during the step cycle. Instead, the data invite a reinterpretation in terms of the opening and closing of reflex pathways, presumably by a central pattern generator for locomotion.