Effects of tendon vibration on the spatiotemporal characteristics of human locomotion

The present study addressed the involvement of proprioceptive input of the muscle spindles in the spatiotemporal control of human locomotion. Blindfolded subjects walked along a walkway while tendon vibration, a powerful stimulus of Ia afferents, was applied to various muscles of the lower limb. The effects of tendon vibration were measured on joint kinematics and on intralimb and interlimb coordination. Tendon vibration of the tibialis anterior during locomotion led to a decreased plantar flexion at toe-off, whereas vibration of the triceps surae led to a decreased dorsiflexion during swing. Vibration of the quadriceps femoris at the knee led to a decreased knee flexion during swing. These local effects of vibration can be explained in the light of a lengthening illusion of the vibrated muscle in that phase of the gait cycle where the muscle is lengthened. Tendon vibration did not affect the qualitative features of intralimb coordination. With respect to interlimb coordination, only vibration of the biceps femoris showed a significant increase in phase lead of the vibrated limb. The present results suggest the involvement of Ia afferent input in the online control of joint rotations. Additionally it is hypothesized that the proprioceptive input of biceps femoris might be involved in the control of coordination between the limbs, whereas the coordination between the segments of one limb appears to be unaffected by disturbance of muscle spindle input of one muscle.     

Increases in muscle activity produced by vibration of the thigh muscles during locomotion in chronic human spinal cord injury

The purpose of this study was to determine whether the muscle vibration applied to the quadriceps has potential for augmenting muscle activity during gait in spinal cord injured (SCI) individuals. The effects of muscle vibration on muscle activity during robotic-assisted walking were measured in 11 subjects with spinal cord injury (SCI) that could tolerate weight-supported walking, along with five neurologically intact individuals. Electromyographic (EMG) recordings were made from the tibialis anterior (TA), medial gastrocnemius (MG), rectus femoris (RF), vastus lateralis (VL), and medial hamstrings (MH) during gait. Vibration was applied to the anterior mid-thigh using a custom vibrator oscillating at 80 Hz. Five vibratory conditions were tested per session including vibration applied during: (1) swing phase, (2) stance phase, (3) stance-swing transitions, (4) swing-stance transitions, and (5) throughout the entire gait cycle. During all vibration conditions, a significant increase in EMG activity was observed across both SCI and control groups in the RF, VL, and MH of the ipsilateral leg. In the SCI subjects, the VL demonstrated a shift toward more appropriate muscle timing when vibration was applied during stance phase and transition to stance of the gait cycle. These observations suggest that the sensory feedback from quadriceps vibration caused increased muscle excitation that resulted in phase-dependent changes in the timing of muscle activation during gait.     

Contribution of afferent feedback to the soleus muscle activity during human locomotion

During the stance phase of the human step cycle, the ankle undergoes a natural dorsiflexion that stretches the soleus muscle. The afferent feedback resulting from this stretch enhances the locomotor drive. In this study a robotic actuator was used to slightly enhance or reduce the natural ankle dorsiflexion, in essence, mimicking the small variations in the ankle dorsiflexion movement that take place during the stance phase of the step cycle. The soleus (SOL) and tibialis anterior EMG were analyzed in response to the ankle trajectory modifications. The dorsiflexion enhancements and reductions generated gradual increments and decrements, respectively, in the ongoing SOL EMG. We exercised care to ensure that the imposed ankle movements were too slow to elicit distinct burst-like stretch reflex responses that have been investigated previously. The increased SOL EMG after the dorsiflexion enhancements was reduced when the group Ia afferents were blocked with peripheral ischemia at the thigh, and during high-frequency Achilles tendon vibration. However, neither ischemia nor tendon vibration affected the decrements in the SOL EMG during the dorsiflexion reductions. These findings give evidence of the contribution of afferent feedback to the SOL activity in an ongoing basis during the stance phase. The results suggest that mainly feedback from the group Ia pathways is responsible for the increments in the SOL EMG during the dorsiflexion enhancements. However, the decrements in the SOL activity might be mediated by different afferent mechanisms.     

Amplitude modulation of the quadriceps H-reflex in the human during the early stance phase of gait

Summary Amplitude modulation of the quadriceps H reflex was investigated during the early part of the stance phase of gait in normal human subjects. Stability of the M wave was used to ensure constancy of the effective stimulus strength. In all subjects there was a progressive decrease in the reflex amplitude throughout the early knee flexion (yield of the knee), whereas the quadriceps EMG activity remained constant or even increased. At an equal stimulus strength and EMG level, the reflex was often larger at the onset of the stance phase of gait than during voluntary contraction, whereas it was always smaller during the knee extension following the yield of the knee. It is argued that changes in presynaptic inhibition of quadriceps Ia terminals could account for this amplitude modulation of the monosynaptic reflex during gait. The possible role of changes in the gain of the quadriceps stretch reflex during bipedal gait is discussed.     

Fast voluntary trunk flexion movements in standing: motor patterns

The electromyographical (EMG) activity was studied during voluntary flexion movements of the trunk in erect standing man. The movements were performed at maximal velocity with successively increasing amplitude to cover the whole range of motion. The EMG activity was recorded from agonist-antagonist pairs of muscles at the ankle, knee, hip and trunk. The angular displacements at the corresponding joints were recorded using a Selspot optoelectronic system. The duration of initiating activity in prime movers (rectus abdominis and rectus femoris) as well as time to onset of activity in muscles braking the primary movement (erector spinae, gluteus maximus and hamstrings) were highly correlated with amplitude, duration, peak velocity and time to peak velocity of the movement (r = 0.59-0.91). The corresponding correlations for peak acceleration and deceleration of the movement were low (r = 0.03-0.38), indicating that acceleration and deceleration of a movement was not coded in the temporal aspects of the EMG. Onset of activity in rectus abdominis and rectus femoris as well as an early appearing burst of activity in vastus lateralis were invariant in relation to start of movement over the whole movement range. In the initial phase of a fast trunk flexion, activity in tibialis anterior appeared successively earlier with increasing movement amplitude. This resulted in a changed order of activation for the muscles from proximal to distal (rectus abdominis first) to distal to proximal (tibialis anterior first). Two different forms of associated postural adjustments are present during a fast trunk flexion, one early fast knee flexion and a later slower angle extension. Prior to knee flexion, no activity was recorded from muscles flexing at the knee implying that some other force must create a flexing torque around the knee. It is suggested that activity in rectus abdominis initiating the primary movement also initiates knee flexion through the upward pulling of pelvis. This would be possible since rectus femoris stabilizes the pelvis in relation to the leg, allowing the force in rectus abdominis to be transmitted below the hip joint and act extending around the ankle joint. However, when tibialis anterior is activated it stabilizes the shank which in turn will cause a knee flexion controlled by a lengthening contraction in vastus lateralis. During the subsequent ankle extension activity appears in lateral gastrocnemius and soleus causing the associated postural adjustment at the ankle. It can be concluded that activation of postural muscles prior to prime mover muscles is not always necessary.(ABSTRACT TRUNCATED AT 400 WORDS)     

Adaptive control for backward quadrupedal walking. II. Hindlimb muscle synergies

1. To compare the basic hindlimb synergies for backward (BWD) and forward (FWD) walking, electromyograms (EMG) were recorded from selected flexor and extensor muscles of the hip, knee, and ankle joints from four cats trained to perform both forms of walking at a moderate walking speed (0.6 m/s). For each muscle, EMG measurements included burst duration, burst latencies referenced to the time of paw contact or paw off, and integrated burst amplitudes. To relate patterns of muscle activity to various phases of the step cycle, EMG records were synchronized with kinematic data obtained by digitizing high-speed ciné film. 2. Hindlimb EMG data indicate that BWD walking in the cat was characterized by reciprocal flexor and extensor synergies similar to those for FWD walking, with flexors active during swing and extensors active during stance. Although the underlying synergies were similar, temporal parameters (burst latencies and durations) and amplitude levels for specific muscles were different for BWD and FWD walking. 3. For both directions, iliopsoas (IP) and semitendinosus (ST) were active as the hip and knee joints flexed at the onset of swing. For BWD walking, IP activity decreased early, and ST activity continued as the hip extended and the knee flexed. For FWD walking, in contrast, ST activity ceased early, and IP activity continued as the hip flexed and the knee extended. For both directions, tibialis anterior (TA) was active throughout swing as the ankle flexed and then extended. A second ST burst occurred at the end of swing for FWD walking as hip flexion and knee extension slowed for paw contact. 4. For both directions, knee extensor (vastus lateralis, VL) activity began at paw contact. Ankle extensor (lateral gastrocnemius, LG) activity began during midswing for BWD walking but just before paw contact for FWD walking. At the ankle joint, flexion during the E2 phase (yield) of stance was minimal or absent for BWD walking, and ankle extension during BWD stance was accompanied by a ramp increase in LG-EMG activity. At the knee joint, the yield was also small (or absent) for BWD walking, and increased VL-EMG amplitudes were associated with the increased range of knee extension for BWD stance. 5. Although the uniarticular hip extensor (anterior biceps femoris, ABF) was active during stance for both directions, the hip flexed during BWD stance and extended during FWD stance.(ABSTRACT TRUNCATED AT 400 WORDS)     

The normal peak of electromyographic activity of the quadriceps femoris muscle in the stair cycle

The quadriceps femoris muscles of 18 subjects with no history of knee joint pathology were analysed climbing stairs. Temporal data was obtained from bilateral contact closing footswitches. Knee joint data was measured using a specially constructed flexible linkage-bar electrogonimeter. Electromyographic activity was obtained from bipolar Beckman surface electrodes placed on four components of the quadriceps femoris, vastus medialis oblique, vastus medialis longus, vastus lateralis and rectus femoris. Results showed that within the stair cycle, stance occupied 60% and swing 40%. Cadence values were greater during descending than ascending stairs. Joint angle data demonstrated 2 changes in direction of the angular motion of the knee joint in both ascending and descending. Electromyographic analysis identified a peak of EMG activity for each component of the quadriceps femoris in both ascending and descending stairs. Results identified the location of peak EMG activity at specific knee joint angles. The quadriceps components also demonstrated a regular sequence of recruitment. EMG amplitude levels obtained were higher in ascending than descending stairs. The results have clinical implications in the design of lower extremity prostheses and in the application of functional electrical stimulation.     

Modulation of heteronymous reflexes from ankle dorsiflexors to hamstring muscles during human walking

In 16 human subjects, stimulation of the common peroneal nerve (CPN) was applied during walking and standing. The effect of the stimulation was evaluated from the rectified and averaged biceps femoris (BF) electromyographic (EMG) activity. In the swing phase of walking, the CPN stimulation evoked a suppression in the BF EMG in 12 of the subjects. In the early stance phase, the suppression was replaced by facilitation at a similar latency in 9 of the subjects. Of the other 3 subjects, in whom a suppression was observed during swing, a decrease in the suppression was observed in the stance phase in two of them. During a voluntary co-contraction of BF and tibialis anterior while standing, a suppression similar to that observed in the swing phase was observed. The thresholds of the suppression and facilitation were identical, suggesting that afferents of similar diameter were responsible. Cutaneous stimuli, which mimicked the sensation evoked by the CPN stimulation, but without activation of muscle afferents, did not produce similar effects in the BF EMG activity. It is suggested that the observed response and reflex reversal may reflect opening of an excitatory group I pathway in the early stance phase of walking with a concomitant shut-down of heteronymous group I inhibition.     

Early corrective reactions of the leg to perturbations at the torso during walking in humans

The contribution of afferent feedback to the regulation of locomotion in humans is not well understood. Animal experiments have suggested that loading of the leg during the stance phase may enhance the magnitude of extensor burst activity and delay the onset of swing phase. The aim of the present study was to determine whether transient loading of the leg at the end of stance would enhance extensor-muscle activity and delay the onset of swing in walking humans. To test this hypothesis, we applied loads to the hips of subjects so that the load was applied along the long axis of the leg at the end of stance (down-back unsupported, DBU). This resulted in an unexpectedly complex reaction characterised by rapid co-contraction of antagonist pairs of muscles around the ankle and knee and a prolongation of the stance phase. We speculated that the complexity of the reaction was, in part, due to a disturbance in equilibrium. To address this possibility, two additional perturbation paradigms were tested: (1) subjects held a rail during the loading paradigm (down-back supported, DBS), or (2) subjects received only a posteriorly directed perturbation of the hips, which added no additional load to the leg (backward unsupported, BU). As predicted, the DBS perturbation resulted in an enhancement of the ongoing soleus-muscle activity, and the unexpected tibialis anterior burst that was observed during the DBU paradigm was absent. Allowing the subjects to hold a rail substantially reduced the change in the timing of the step cycle observed in the DBU paradigm. The BU perturbation prolonged the stance phase duration and, as expected, resulted in a burst of activity in tibialis activity. This was usually accompanied by a reduction in the ongoing soleus activity. Two important conclusions are drawn from the present study. First, loading of the leg at the end of stance phase enhances the ongoing extensor-muscle activity. We suggest that afferent feedback responding to the increase load supported by the leg leads to rapid enhancement of the active extensor muscles to compensate for the increased load and prevent collapse of the leg. Interestingly, the duration of the stance phase was only marginally increased when loading was applied without a postural disturbance (DBS). Second, posterior perturbation of the centre of mass at the end of stance phase evokes an "automatic postural response" in tibialis anterior. Of particular interest, this evoked postural response can occur simultaneously with an enhanced activation of soleus. This indicates that the DBU perturbation employed in this study elicited two responses, one to prevent the collapse of the leg and the other to stabilise the centre of mass.     

Effect of knee joint laxity on long-loop postural reflexes: evidence for a human capsular-hamstring reflex

Summary The onset latency and discharge amplitude of preprogrammed postural responses were evaluated in order to determine if the structure of synergistic activation could be altered by ligamentous laxity at the knee joint. Twelve subjects with unilateral and one subject with bilateral anterior cruciate ligament (ACL) insufficiency were tested while standing on a moveable platform. External balance perturbations (6 cm anterior or posterior horizontal displacements of the platform) were presented at velocities ranging from 15 to 35 cm/s. Perturbations were presented under the following experimental conditions: unilateral and bilateral stance, knees fully straight or flexed, and with ankle motion restricted or free. These stance, knee position, and ankle motion conditions were introduced to alter the stress transmitted to the knee joint during movement of the support surface. The automatic postural response was recorded from the tibialis anterior (T), quadriceps (Q), and medial hamstrings muscles (H) bilaterally. The normal response to an externally induced backward sway involved the automatic activation of T and Q at latencies of 80 ms and 90 ms respectively. Activation of the hamstrings in the non-injured extremity was not coupled with the postural response. Hamstrings are not typically involved in the correction posterior sway because H activation would tend to pull the center of mass further backwards. However, when the response in the ACL-deficient extremity was compared to the non-injured limb: (1) the automatic postural response in the ACL-deficient extremity was restructured to include hamstrings activation (100 ms latency), (2) H activation time was faster and less variable in the ACL-deficient limb, and (3) the ratio of H/Q discharge amplitude integrated over 100 ms and 200 ms from the onset of EMG activation showed a dominance of hamstring activity during unilateral stance on the lax limb. In addition, H/Q ratios integrated over 200 ms showed dominant hamstring activity in the ACL-deficient limb during bilateral stance. (4) Crosslimb comparisons showed greater normalized IEMG amplitudes for T, H, and Q during unilateral stance on the lax limb. These results suggest that a capsular-hamstring reflex is integrated into the existing structure of a preprogrammed postural synergy in order to compensate for ligamentous laxity. Furthermore, the generalized increase of response gain observed during perturbations of unilateral stance on the lax limb indicates that joint afference can modulate central programming to control localized joint hypermobility. A concept of postural control is discussed with respect to the capsular reflex, joint loading and displacement of the center of gravity.     

Characterisation of the quadriceps stretch reflex during the transition from swing to stance phase of human walking

The main objective of this study was to characterize the stretch reflex response of the human thigh muscles to an unexpected knee flexion at the transition from stance to swing during walking. Eleven healthy subjects walked on a treadmill at their preferred speed. Reliable and constant knee flexions (6–12° amplitude, 230–350°/s velocity, 220 ms duration) were applied during the late swing and early stance phase of human walking by rotating the knee joint with a specifically designed portable stretch apparatus affixed to the left knee. Responses from rectus femoris (RF), vastus lateralis (VL), vastus medialis (VM), biceps femoris (BF), medial hamstrings (MH) and medial gastrocnemius (GM) were recorded via bipolar surface electromyograms (EMG). The onset of the response in the RF, VL and VM, remained stable and independent of the time in the step cycle when the stretch was applied. Across all subjects the response onset (mean ± SD) occurred at 23±1, 24±1 and 23±1 ms for RF, VL and VM, respectively. The duration of the initial response was 90–110 ms, at which time the EMG signal returned towards baseline levels. Three reflex response windows, labelled the short latency reflex (SLR), the medium latency reflex (MLR) and the late latency reflex response (LLR), were analysed. The medium and late reflex responses of all knee extensors increased significantly (p=0.008) as the gait cycle progressed from swing to stance. This was not related to the background EMG activity. In contrast, during standing at extensor EMG levels similar to those attained during walking the reflex responses were dependent on background EMG. During walking, LLR amplitudes expressed as a function of the background activity were on average two to three times greater than SLR and MLR reflex amplitudes. Distinct differences in SLR and LLR amplitude were observed for RF, VL and VM but not in the MLR amplitude. This may be related to the different pathways mediating the SLR, MLR and LLR components of the stretch response. As for the knee extensor antagonists, they exhibited a response to the stretch of the quadriceps at latencies short enough to be monosynaptic. This is in agreement with the suggestion by Eccles and Lundberg (1958) that there may be functional excitatory connections between the knee extensors and flexors in mammals.     

Head stabilization on a continuously oscillating platform: the effect of a proprioceptive disturbance on the balancing strategy

When standing and balancing on a continuously and predictably moving platform, body equilibrium relies on both anticipatory control and proprioceptive feedback. We have vibrated different postural muscles of the body to assess any effect of confounding the proprioceptive input on balance during such unstable conditions. Low and high platform oscillation frequencies were used, because different strategies are used to withstand the two perturbations. Eyes open (EO) and closed (EC) conditions were also tested, to assess whether the stabilizing effect of vision is independent from the proprioceptive disturbance. Subjects (n=14) performed two series of trials, EO and EC: (1) quiet erect stance, (2) stance on the platform translating at 0.2 or 0.6 Hz sinusoidally in the anteroposterior (A-P) direction (dynamic conditions). Continuous bilateral vibration (90 Hz) was produced by two vibrators fixed to the following homonymous muscles: dorsal neck, quadriceps, biceps femoris, tibialis anterior, and triceps surae. Acquisition of body segments displacement began 10 s after the start of platform translation. From markers fixed to head, hip, and malleolus, we computed the A-P oscillation of head and hip, body orientation in space, and cross-correlation (CC) and time-delay between malleolus and head trajectories. The results were (a) the head A-P oscillation was smaller with EO than EC, under both quiet stance and dynamic conditions; (b) vibration of tibialis and triceps surae, but not of other muscles, slightly increased head and body A-P oscillation with EC under dynamic conditions; (c) at 0.2 Hz but not at 0.6 Hz, for all visual and vibration conditions, there was a significant association between head and feet; (d) at 0.2 Hz, EC, neck muscle vibration increased this association, whereas vibration of the other muscles induced a major time delay in the oscillation of head compared with feet; (e) vibration of either neck or tibialis induced forward body leaning, while vibration of either triceps surae or biceps femoris induced backward leaning, with both EO and EC, under both static and dynamic conditions; (f) the head A-P oscillation, however, under dynamic conditions was not dependent on body leaning. The relatively scarce effects of proprioceptive disturbance on head stabilization and multijoint coordination (in spite of effects on body orientation similar to those observed during stance) speak for a major role of anticipatory control in the dynamic equilibrium task. However, the significant vibration-induced time delay in segments coordination at low translation frequency, EC, suggests that the normally patterned Ia input promotes continuous adjustments of the feed-forward control mode.     

Treadmill locomotion in normal mice—Step related multi-channel EMG profiles of thigh muscles

Mouse models are increasingly used in current research on motor disorders. In mice, the myoelectrical activation of thigh muscles during locomotion has not yet, however, been investigated in depth. Especially intramuscular coordination has hardly been clarified. Therefore, the aims of this study were to characterize myoelectrical activity in the vastus lateralis (VL) and the biceps femoris (BF) muscle of the healthy mouse for reference purposes. The VL and the BF muscles of 12 healthy mice performing a total of 1985 steps during treadmill locomotion were investigated with two subcutaneous arrays each incorporating four electrodes. Eight-channel EMG was recorded simultaneously with high-speed videography. The EMG curves of each step were rectified and smoothed by calculating root mean square (RMS) profiles and then time-normalized for comparisons within and between animals. The EMG-activity of both muscles increased during late swing phase. The VL activity rose steeply and peaked during mid-stance phase, while the biceps activity reached a plateau during early stance phase. With increasing gait velocity, stance time decreased. The increase in gait velocity was also associated with greater EMG amplitudes. The results suggest that the BF lifts the lower hind leg during swing phase and stabilizes the leg during stance, while the VL bears the weight of the body during the stance phase.     

Functionally complex muscles of the cat hindlimb

The biceps femoris (BF) muscle is divided into three neuromuscular compartments defined by the innervation patterns of the main nerve branches (English and Weeks 1987). The goals of this study were i) to determine how different regions of the biceps femoris muscle are activated in the intact cat during a broad range of limb movements evoked by perturbations of stance posture, and ii) to determine the relationship between the anatomical compartments of biceps femoris and the functional units as defined in this task. Cats were trained to stand on a moveable platform with each paw on a triaxial force plate. The animal's stance was perturbed by linear translation of the platform in each of sixteen different directions in the horizontal plane. EMG activity was recorded from eight sites across the width of the left biceps femoris muscle. During quiet stance only the anterior compartment was tonically active, presumably contributing to hip extensor torque in the maintenance of stance. During platform translation, evoked EMG activity was recorded from each electrode pair for a wide range of directions of perturbation; as direction changed progressively, the amplitude of evoked activity from any electrode pair increased to a maximum and then decreased. When the EMG amplitude was plotted in polar coordinates as a function of translation direction, the region of response formed a petal shaped area in the horizontal plane, termed the EMG tuning curve. The compartments of the BF muscle were not activated homogeneously. The tuning curve of the anterior BF compartment was similar to that of other hip extensors, and coincided with the region of postero-lateral force production by the hindlimb against the support. The tuning curve of the middle BF compartment was shifted in a counterclockwise direction from that of the anterior compartment, but overlapped extensively with it; the middle BF tuning curve was similar to that of anterior gracilis. The tuning curve of the posterior biceps compartment was rotated further counterclockwise and overlapped very little with that of the middle BF compartment. The posterior BF was activated in a pattern similar to that of other knee flexors. The functional units of BF activation were not identical with the neuromuscular compartments defined by the main nerve branches. As direction of the perturbation changed, the region of BF that was activated moved progressively across the muscle. This progression of the active region was continuous across BFa and BFm, whereas there was a jump, or discontinuity at the border between BFm and BFp.(ABSTRACT TRUNCATED AT 400 WORDS)     

Pattern of monosynaptic heteronymous Ia connections in the human lower limb

Changes in the firing probability of single motor units in response to electrical stimulation of muscle nerves were used to derive the projections of muscle spindle Ia afferents to the motoneurones of various leg and thigh muscles. Discharges of units in soleus, gastrocnemius medialis, peroneus brevis, tibialis anterior, quadriceps, biceps femoris and semitendinosus were investigated after stimulation of inferior soleus, gastrocnemius medialis, superficial peroneal, deep peroneal and femoral nerves. Homonymous facilitation, occurring at the same latency as the H reflex and therefore attributed to monosynaptic Ia EPSPs, was found in virtually all the sampled units. In many motor nuclei an early facilitation was also evoked by heteronymous low-threshold afferents. The heteronymous facilitation was considered to be mediated through a monosynaptic pathway when the difference between the central latencies of heteronymous and homonymous peaks was not more than 0.2 ms. The heteronymous Ia connections were widely distributed. In particular, monosynaptic coupling between muscles operating at different joints appears to be the rule in humans, though it is rare between ankle and knee muscles in the cat and the baboon.     

The amplitude modulation of the Quadriceps H-reflex in relation to the knee joint action during walking

Previously the modulation of the quadriceps H-reflex has only been investigated in the initial part of the gait cycle, and it was suggested that the quadriceps H-reflex modulates with relative high reflex gain at heel contact and decreases during the subsequent part of stance (Dietz et al. 1990b). The objectives of the present study was to elaborate on the previous results by increasing the measurement resolution around heel contact and include additional measures in order to relate the H-reflex modulation to the mechanical function of the knee extensors throughout the gait cycle. EMG profiles were measured in quadriceps and the antagonistic hamstring muscles simultaneously with the knee joint kinematics in ten subjects during treadmill walking at preferred speed. H-reflex excitability was measured in vastus lateralis (VL) and rectus femoris (RF) at 11 selected positions during the gait cycle. The resulting excitability curves showed a significant modulation of the quadriceps H-reflex during the gait cycle. The H-reflex amplitude increases shortly after heel contact and reflex inhibition is present in the remaining part of stance and most of the swing phase. The modulation of the quadriceps H-reflex during walking does not follow the classical pattern of reciprocal inhibition between antagonistic muscles. It is suggested that at least during the stance phase the modulation of the quadriceps H-reflex is controlled by presynaptic inhibition. The present results confirm the idea that the excitability of the quadriceps H-reflex is controlled to comply with the different mechanical demands on the muscle during the gait cycle in humans.     

Modulation of the startle response during human gait

While many studies have shown that there is a phase-dependent modulation of proprioceptive and exteroceptive reflexes during gait, little is known about such modulation for auditory reflexes. To examine how startle reactions are incorporated in an ongoing gait pattern, unexpected auditory stimuli were presented to eight healthy subjects in six phases of the step cycle during walking on a treadmill at 4 km/h. For both legs, electromyographic activity (EMG) was recorded in the biceps femoris (BF), the rectus femoris (RF), the tibialis anterior (TA), and the soleus (SO). In addition, stance and swing phases of both legs, along with knee angles of both legs and the left ankle angle, were measured. All subjects showed various response peaks. Responses with latencies of approximately 60 ms (F1), approximately 85 ms (F2), and approximately 145 ms (F3) were found. The amplitude of the reflex responses was dependent on the timing of the startle stimulus in the step cycle. Although the startle response habituated rapidly, the phase-dependent modulation pattern generally remained the same. The phase-dependent amplitude modulations were not strictly correlated with the modulation of the background activity. The TA even showed a transition from facilitatory F2 responses during stance to suppressive responses during midswing. Responses were observed in both flexors and extensors, often in coactivation, especially during stance. Furthermore the gait characteristics showed a shortening of the subsequent step cycle and a small decrease in the range of motion of ankle and knees. These results suggest that the responses are adapted to achieve extra stability dependent on the phase of the step cycle. However, even in the first trials, the changes in kinematics were small allowing a smooth progression of gait.     

Influence of leg muscle vibration on human walking

We studied the effect of vibratory stimulation of different leg muscles [bilateral quadriceps (Q), hamstring (HS) muscles, triceps surae (TS), and tibialis anterior (TA)] in seven normal subjects during 1) quiet standing, 2) stepping in place movements, and 3) walking on the treadmill. The experiments were performed in a dimly illuminated room, and the subjects were given the instruction not to resist the applied perturbation. In one condition the velocity of the treadmill was controlled by a feedback from the subject's current position. In normal standing, TA vibration elicited a prominent forward body tilt, whereas HS and TS vibration elicited backward trunk or whole body inclination, respectively. Q vibration had little effect. During stepping in place, continuous HS vibration produced an involuntary forward stepping at about 0.3 m s(-1) without modifying the stepping frequency. When the subjects (with eyes closed) kept a hand contact with an external still object, they did not move forward but perceived an illusory forward leg flexion relative to the trunk. Q, TS, and TA vibration did not cause any systematic body translation nor illusory changes in body configuration. In treadmill locomotion, HS vibration produced an involuntary steplike increase of walking speed (by 0.1-0.6 m.s(-1)). Continuous vibration elicited larger speed increments than phasic stimulation during swing or stance phase. For phasic stimulation, HS vibration tended to be more effective when applied during swing than during stance phase. Q, TA, and TS vibration had little if any effect. Vibration of thigh muscles altered the walking speed depending on the direction of progression. During backward locomotion, the walking speed tended to decrease after HS vibration, whereas it significantly increased after Q vibration. Thus the influence of leg muscle vibration on stepping in place and locomotion differed significantly from that on normal posture. We suggest that the proprioceptive input from thigh muscles may convey information about the velocity of the foot movement relative to the trunk.     

The role of cutaneous afferents in controlling locomotion evoked by epidural stimulation of the spinal cord in decerebrate cats

The effects of the cutaneous input on the formation of the locomotor pattern in conditions of epidural stimulation of the spinal cord in decerebrate cats were studied. Locomotor activity was induced by rhythmic stimulation of the dorsal surface of spinal cord segments L4-L5 at a frequency of 3-5 Hz. Electromyograms (EMG) recorded from the antagonist muscles quadriceps, semitendinosus, tibialis anterior, and gastrocnemius lateralis were recorded, along with the kinematics of stepping movements during locomotion on a moving treadmill and reflex responses to single stimuli. Changes in the pattern of reactions observed before and after exclusion of cutaneous receptors (infiltration of lidocaine solution at the base of the paw or irrigation of the paw pads with chlorothane solution) were assessed. This treatment led to impairment of the locomotor cycle: the paw was placed with the rear surface downward and was dragged along in the swing phase, and the duration of the stance phase decreased. Exclusion of cutaneous afferents suppressed the polysynaptic activity of the extensor muscles and the distal flexor muscle of the ipsilateral hindlimb during locomotion evoked by epidural stimulation of the spinal cord. The effects of exclusion of cutaneous afferents on the monosynaptic component of the EMG response were insignificant.     

On the origin of the soleus H-reflex modulation pattern during human walking and its task-dependent differences

Recently, Brooke and colleagues have suggested "that the strong inhibition arising from passive movement about the knee and hip joints, lays down the base for the soleus H-reflex gain modulation seen during human gait." In particular stretch-evoked afferent activity from the quadriceps muscle was emphasized as the most important source of movement-induced inhibition of the H-reflex. To test this hypothesis we examined the kinematics and electromyographic (EMG) activity of the leg during human walking and correlated these with the modulation pattern of the soleus H-reflex. To further test the possible contribution of stretch-evoked quadriceps afferent activity to the soleus H-reflex modulation pattern during walking different walking gaits were studied. In one condition subjects were asked to walk with their knee locked in full extension by a rigid knee brace. In a second condition subjects were asked to walk backwards. During normal walking, the soleus H-reflex modulation pattern is strongly correlated with the EMG events of the soleus and tibialis anterior (TA), but not with hip, knee, or ankle angular displacement or velocity. When subjects walked with the knee locked in full extension, the amplitude of the H-reflex, its modulation pattern, and the task-dependent changes of its amplitude were the same as during normal walking. During backward walking, the H-reflex increases in late swing before activity of the soleus has begun and while the knee is flexing, an observation that highlights central control of the H-reflex amplitude. The effects of imposed flexion of the knee in passive subjects were also reexamined. The knee flexion imposed by the experimenter followed the same trajectory as that which occurred during the swing phase of the subject's step cycle. It was found that imposed knee flexions elicited a burst of TA EMG activity with an average latency of 81.6 ms (SD = 21 ms) in six out of eight subjects. Inhibition of the H-reflex, when it occurred, was associated with the occurrence of this burst. When subjects voluntarily flexed their right knee from an initial quiet standing posture, the inhibition of the soleus H-reflex began before flexion of the knee or that of any other leg segment. Once again the onset of inhibition was closely associated with the onset of activity in the TA. In the discussion section the present observations are examined in light of the predictions made by the movement-induced inhibition hypothesis of Brooke et al. It will be concluded that none of the predictions of this hypothesis were corroborated by present tests done during human walking. In consequence, we suggest that the modulation pattern of the H-reflex observed during normal human walking is centrally determined, as are the task-dependent differences of its amplitude (e.g., standing versus the stance phase of human walking).