The relationship between dynamic balancing ability and posture-related modulation of the soleus H-reflex

Soleus H-reflex reveals down modulation with increased postural difficulty. Role of this posture-related reflex modulation is thought to shift movement control toward higher motor centers in order to facilitate more precise postural control. Present study hypothesized that the ability to modulate H-reflex is related to one’s ability to dynamically balance while in an unstable posture. This study examined the relationship between dynamic balancing ability and soleus H-reflex posture-related modulation. Thirty healthy adults participated. The soleus maximal H-reflex (Hmax), motor response (Mmax), and background EMG activity (bEMG) were obtained during three postural conditions: prone, open-legged standing, and closed-legged standing. Hmax/Mmax ratios were normalized via the corresponding bEMG in order to remove the effects of background muscle activity from the obtained H-reflex. Reflex modulation was calculated as the ratio of the normalized Hmax/Mmax ratios in one postural condition to another posture in a more difficult condition. Dynamic balancing ability was assessed by testing stability while standing on a wobble board. A significant negative correlation was observed between balancing scores and reflex modulation from open-legged standing to closed-legged standing. This suggests that the ability to modulate monosynaptic stretch reflex excitability in response to a changing posture is a significant factor for dynamic balancing.     

Daily muscle vibration amelioration of neural impairments of the soleus muscle during 2 weeks of immobilization

V-wave, F wave and H-reflex responses of soleus were used to determine neural adaptations to 2-week immobilization and whether muscle vibration intervention during immobilization would attenuate the negative adaptations induced by immobilization. Thirty subjects were divided into the ankle immobilization group and the immobilization with muscle vibration group. Mechanical vibrations with constant low amplitude (0.3 mm) were applied (12 × 4 min daily) with a constant frequency of 100 Hz on the soleus muscle of the subjects in vibration group during the ankle immobilization period. Soleus maximal M-wave (Mmax) and H-reflex (Hmax) were evoked at rest. F-wave was recorded by supramaximal stimulation delivered at rest and V-wave during maximum voluntary contraction (MVC). The EMG during MVC was represented by its root-mean-square (RMS) value. Each subject was examined before and after 2 weeks of immobilization. Results showed that following 2 weeks of immobilization, Mmax, Hmax and F wave all did not change with immobilization in either group (P > 0.05). After 2 weeks of immobilization, significant reductions in V/Mmax (of 30.78%) (P < 0.01) and EMG RMS (24.82%) (P < 0.001) were found in the immobilization group. However, no significant changes occurred in the immobilization with muscle vibration group. Such findings suggested that 2 weeks of immobilization resulted in neural impairments as evidenced by the reduction in EMG and V wave, and that such decrease was prevented by the intervention of muscle vibration during the immobilization period.     

Differences in H-reflex between athletes trained for explosive contractions and non-trained subjects

The efficacy of type Ia synapse on alpha-motoneurons of soleus and lateral gastrocnemius muscles has been investigated, using the H-reflex technique, in athletes engaged in sports requiring very rapid and intense contractions (sprinters and volley-ball players) as well as in non-trained subjects. It has been observed, in both muscles, that the ratio between the mean value of the maximal reflex response (Hmax) and the mean value of the maximal direct response (Mmax) elicited upon electrical stimulation of the tibial nerve is significantly smaller in athletes trained for explosive-type movements than in non-trained subjects. This difference in the Hmax: Mmax ratio was dependent on a smaller amplitude of Hmax and not on a greater amplitude of Mmax. No significant differences were observed between sprinters and volley-ball players. In both trained and non-trained subjects, soleus and lateral gastrocnemius muscles displayed significant differences in Hmax: Mmax ratio and Mmax amplitude but not in Hmax amplitude. Since the H-response is considered to be due mainly to activation of the smallest motoneurons in the motoneuronal pools, the difference in Hmax amplitude and Hmax: Mmax ratio between athletes and non-trained subjects could have been dependent on a lower incidence of these motoneurons in the athletes. This is in accord with the mechanical needs of muscles during explosive-type power training. Although this difference may have been wholly determined genetically, the possibility is discussed as to whether the lower incidence in sprinters and volley-ball players of small motoneurons could have been related to a training-induced transformation of small and slow motoneurons into large and fast ones.     

Passive knee movement-induced modulation of the soleus H-reflex and alteration in the fascicle length of the medial gastrocnemius muscle in humans

In humans, an inhibitory via Ia afferent pathway from the medial gastrocnemius (MG) to the soleus (SOL) motoneuron pool has been suggested. Herein, we examined the relation between MG fascicle length changes and the SOL H-reflex modulation during passive knee movement. Twelve subjects performed static and passive (5° s^?1) knee movement tasks with the ankle immobilized using an isokinetic dynamometer in sitting posture. The maximal H- and M-waves were measured at four target angles (20°, 40°, 60°, and 80° flexion from full knee extension). The MG fascicles length and velocity were measured using a B-mode ultrasonic apparatus. Results demonstrated that the SOL Hmax/Mmax; i.e., ratio of the maximal H- to M-waves, was attenuated with increasing MG fascicle length in static tasks. The SOL Hmax/Mmax at 20° was significantly attenuated compared with 60° and 80° with increasing MG fascicle length and lengthening velocity in passive knee extension. However, no significant differences in the SOL Hmax/Mmax were found across the target angles in the passive knee flexion task. In conclusion, as muscle spindles increase their discharge with lengthening fascicle velocity, but keep silent when fascicles shorten, our data suggest that lengthening the MG facilitates an inhibitory Ia pathway from MG to SOL, and modulates SOL motoneuron activity during movements.     

Influence of stimulus intensity on the soleus H-reflex amplitude and modulation during locomotion

Diverging results have been reported regarding the modulation and amplitude of the soleus H-reflex measured during human walking and running. A possible explanation to this could be the use of too high stimulus strength in some studies while not in others. During activities like walking and running it is necessary to use a small M-wave to control the effective stimulus strength during all phases of the movement. This implies that the descending part of the H-reflex recruitment curve is being used, which may lead to an unwanted suppression of the H-reflex due to limitations imbedded within the H-reflex methodology itself.Accordingly, the purpose of the present study was to study the effect on the soleus H-reflex during walking and running using stimulus intensities normally considered too high (up to 45% Mmax).Using M-waves of 25–45% Mmax as opposed to 5–25% Mmax showed a significant suppression of the peak H-reflex during the stance phase of walking, while no changes were observed during running. No differences were observed regarding modulation pattern. So a possible use of too high stimulus intensity cannot explain the differences mentioned. The surprising result in running may be explained by the much higher voluntary muscle activity, which implies the existence of a V-wave influencing the H-reflex amplitude in positive direction.     

The acute effect of whole-body vibration on the hoffmann reflex

The extent to which motoneuron pool excitability, as measured by the Hoffmann reflex (H-reflex), is affected by an acute bout of whole-body vibration (WBV) was recorded in 19 college-aged subjects (8 male and 11 female; mean age 19 +/- 1 years) after tibial nerve stimulation. H/M recruitment curves were mapped for the soleus muscle by increasing stimulus intensity in 0.2- to 1.0-volt increments with 10-second rest intervals between stimuli, until the maximal M-wave and H-reflex were obtained. After determination of Hmax and Mmax, the intensity necessary to generate an H-reflex approximately 30% of Mmax (mean 31.5% +/- 4.1%) was determined and used for all subsequent measurements. Fatigue was then induced by 1 minute of WBV at 40 Hz and low amplitude (2-4 mm). Successive measurements of the H-reflex were recorded at the test intensity every 30 seconds for 30 minutes post fatigue. All subjects displayed a significant suppression of the H-reflex during the first minute post-WBV; however, four distinct recovery patterns were observed among the participants (alpha = 0.50). There were no significant differences between genders across time (P = 0.401). The differences observed in this study cannot be explained by level or type training. One plausible interpretation of these data is that the multiple patterns of recovery may display variation of muscle fiber content among subjects. Future investigation should consider factors such as training specificity and muscle fiber type that might contribute to the differing H-reflex response, and the effect of WBV on specific performance measures should be interpreted with the understanding that there may be considerable variability among individuals. Recovery times and sample size should be adjusted accordingly.     

Modulation of spinal inhibitory reflexes depends on the frequency of transcutaneous electrical nerve stimulation in spastic stroke survivors

Neurophysiological studies in healthy subjects suggest that increased spinal inhibitory reflexes from the tibialis anterior (TA) muscle to the soleus (SOL) muscle might contribute to decreased spasticity. While 50?Hz is an effective frequency for transcutaneous electrical nerve stimulation (TENS) in healthy subjects, in stroke survivors, the effects of TENS on spinal reflex circuits and its appropriate frequency are not well known. We examined the effects of different frequencies of TENS on spinal inhibitory reflexes from the TA to SOL muscle in stroke survivors. Twenty chronic stroke survivors with ankle plantar flexor spasticity received 50-, 100-, or 200-Hz TENS over the deep peroneal nerve (DPN) of the affected lower limb for 30?min. Before and immediately after TENS, reciprocal Ia inhibition (RI) and presynaptic inhibition of the SOL alpha motor neuron (D1 inhibition) were assessed by adjusting the unconditioned H-reflex amplitude. Furthermore, during TENS, the time courses of spinal excitability and spinal inhibitory reflexes were assessed via the H-reflex, RI, and D1 inhibition. None of the TENS protocols affected mean RI, whereas D1 inhibition improved significantly following 200-Hz TENS. In a time-series comparison during TENS, repeated stimulation did not produce significant changes in the H-reflex, RI, or D1 inhibition regardless of frequency. These results suggest that the frequency-dependent effect of TENS on spinal reflexes only becomes apparent when RI and D1 inhibition are measured by adjusting the amplitude of the unconditioned H-reflex. However, 200-Hz TENS led to plasticity of synaptic transmission from the antagonist to spastic muscles in stroke survivors.     

Hypoxia and monosynaptic reflexes in humans

The recruitment curves of the monosynaptic Hoffmann (H) reflex and of the direct motor (M) excitation of alpha-motor fibers of the posterior popliteal nerve were studied in seven human subjects in normoxic and hypoxic conditions at sea level. The amplitude of the H and M responses were determined from the computerized full-wave rectified and integrated surface electromyographic (EMG) signal derived from bipolar surface electrodes placed over the soleus muscle. Hypoxic exposure [end-tidal O2 fraction (FETO2) = 0.066 +/- 0.003 and end-tidal CO2 fraction (FETCO2) = 0.0504 +/- 0.001 (SE)] did not affect the maximal M (Mmax) response but decreased significantly (7%) the maximal H (Hmax) response. The Hmax/Mmax ratio decreased from 0.60 to 0.53. Furthermore, by fitting the rising phase of the recruitment curves of the H and M responses vs. stimulus intensity with linear regressions, hypoxia was found to produce a significant decrease of similar magnitude (6%) in the threshold of both the H and M responses with no change in slope. Using a constant stimulus strength eliciting an H response of half the maximum (H50%) of the control conditions, hypoxia resulted in a 50% increase in the amplitude of the H response within 12 min. These results suggest that the effects of hypoxia on the nervous system consist of a direct depolarizing action on the peripheral alpha-fibers and 1A sensory fibers and of a central effect on supraspinal structures affecting the spinal alpha-motoneurons.     

Post-tetanic potentiation of reciprocal Ia inhibition in human lower limb.

The purpose of this study was to investigate how reciprocal Ia inhibition is changed during muscle fatigue of lower limb muscle, induced with a voluntary contraction or height frequency electrical stimulation. Reciprocal Ia inhibition from ankle flexors to extensors has been investigated in 12 healthy subjects. Hoffmann reflex (H-reflex) in the soleus muscle was used to monitor changes in the amount of reciprocal Ia inhibition from common peroneal nerve as demonstrated during voluntary dorsi or planterflexion and 50 Hz electrical stimulation induced dorsi or planterflexion. The test soleus H-reflex was kept at 20-25% of maximum directly evoked motor response (M response) and the strength of the conditioning common peroneal nerve stimulation was kept at 1.0 x motor threshold. At rest, weak la inhibition was demonstrated in 12 subjects, maximal inhibition from the common peroneal nerve was 28.8%. During voluntary dorsiflexion and 50 Hz electrical stimulation induced dorsiflexion, there absolute amounts of inhibition increased as compared to at rest, and decreased or disappeared during voluntary planterflexion and 50 Hz electrical stimulation induced planterflexion as compared to at rest. During voluntary or electrical stimulation induced agonist muscle fatigue, the inhibition of the soleus H-reflex from the common peroneal nerve was greater during voluntary dorsiflexion (maximal, 11.1%) and 50 Hz (maximal, 6.7%) electrical stimulation induced dorsiflexion than at rest. The inhibition was decreased or disappeared during voluntary planterflexion 50 Hz electrical stimulation induced planterflexion. It was concluded that the results were considered to support the hypothesis that alpha-motoneurones and la inhibitory intemeurones link to antagonist motoneurones in reciprocal inhibition. The diminished reciprocal Ia inhibition of voluntary contraction during muscle fatigue as compared to electrical stimulation, is discussed in relation to its possible contribution to ankle stability.     

Joint kinetic response during unexpectedly reduced plantar flexor torque provided by a robotic ankle exoskeleton during walking

During human walking, plantar flexor activation in late stance helps to generate a stable and economical gait pattern. Because plantar flexor activation is highly mediated by proprioceptive feedback, the nervous system must modulate reflex pathways to meet the mechanical requirements of gait. The purpose of this study was to quantify ankle joint mechanical output of the plantar flexor stretch reflex response during a novel unexpected gait perturbation. We used a robotic ankle exoskeleton to mechanically amplify the ankle torque output resulting from soleus muscle activation. We recorded lower-body kinematics, ground reaction forces, and electromyography during steady-state walking and during randomly perturbed steps when the exoskeleton assistance was unexpectedly turned off. We also measured soleus Hoffmann- (H-) reflexes at late stance during the two conditions. Subjects reacted to the unexpectedly decreased exoskeleton assistance by greatly increasing soleus muscle activity about 60 ms after ankle angle deviated from the control condition (p<0.001). There were large differences in ankle kinematic and electromyography patterns for the perturbed and control steps, but the total ankle moment was almost identical for the two conditions (p=0.13). The ratio of soleus H-reflex amplitude to background electromyography was not significantly different between the two conditions (p=0.4). This is the first study to show that the nervous system chooses reflex responses during human walking such that invariant ankle joint moment patterns are maintained during perturbations. Our findings are particularly useful for the development of neuromusculoskeletal computer simulations of human walking that need to adjust reflex gains appropriately for biomechanical analyses.     

Soleus H-reflex dynamics during fast plantarflexion in humans.

The relationship between the size of the soleus (Sol) Hoffmann (H-) reflex and the level of background (BG) electromyographic (EMG) activity was examined during plantarflexing at different force levels. The experiments were carried out on seven healthy male subjects aged 20-37 years. The subjects were asked to perform fast plantarflexion under a reaction-time condition. The amounts of contraction force were 10, 20, 50 and 80% of maximum voluntary contraction (MVC). Since the maximum size of the M-wave (Mmax) changed systematically during the plantarflexion, we tried to maintain the size of the reference M-wave, an indicator of the efficiency of the electrical stimulation, at a constant value (20% of Mmax) throughout the experiment. The size of the H-reflex was rapidly increased at the very beginning of the movement, and then it tended to decrease in the later phase of the movement. Consequently, even with the same level of BG EMG, the size of the H-reflex was always larger in the early rising phase of the EMG activity than in the later falling phase. The maximum size of the H-reflex was poorly correlated with the force exerted. In contrast, the size of the F-response was proportional to the force exerted. The non-linear relationship between the size of the H-reflex and the BG EMG suggests that the level of the presynaptic inhibition onto Ia terminals was modified depending on the required force level and during the course of the movement.     

The influence of increased muscle spindle sensitivity on Achilles tendon jerk and H-reflex in relaxed human subjects

Whether the fusimotor system contributes to reflex gain changes during reinforcement maneuvers is re-examined in the light of new data. Recently, from direct recordings of spindle afferent activity originating from ankle flexor muscles, we showed that mental computation increased the muscle spindle mechanical sensitivity in completely relaxed human subjects without concomitant alpha-motoneuron activation, providing evidence for selective fusimotor drive activation. In the present study, the effects of mental computation were investigated on monosynaptic reflexes elicited in non-contracting soleus muscle either by direct nerve stimulation (Hoffmann reflex, H) or by tendon tap (Tendinous reflex, T). The aim was to relate the time course of the changes in reflex size to the increase in spindle sensitivity during mental task in order to explore whether fusimotor activation can influence the size of the monosynaptic reflex. The results show changes in reflex amplitude that parallel the increase in muscle spindle sensitivity. When T-reflex is consistently facilitated during mental effort, the H-reflex is either depressed or facilitated, depending on the subjects. These findings suggest that the increased activity in muscle spindle primary endings may account for mental computation-induced changes in both tendon jerk and H-reflex. The facilitation of T-reflex is attributed to the enhanced spindle mechanical sensitivity and the inhibition of H-reflex is attributed to post-activation depression following the increased Ia ongoing discharge. This study supports the view that the fusimotor sensitization of muscle spindles is responsible for changes in both the mechanically and electrically elicited reflexes. It is concluded that the fusimotor drive contributed to adjustment of the size of tendon jerk and H-reflex during mental effort. The possibility that a mental computation task may also operate by reducing the level of presynaptic inhibition is discussed on the basis of H-reflex facilitation.     

On the methods employed to record and measure the human soleus H-reflex

The aim of this study was to investigate if the magnitude of the soleus H-reflex is different depending on the method employed to measure its size (peak-to-peak amplitude vs. area). In this study, 13 healthy human subjects participated, while the soleus H-reflex was induced via conventional methods. In the first experiment, the soleus H-reflex was recorded via two monopolar electrodes and was evoked at least at eight different stimulation intensities in respect to the recovery curve of the H-reflex and at three different inter-stimulus intervals (ISIs) (8, 5, and 2 s). The ISI refers to the time delay between the single pulses delivered to the posterior tibial nerve within a single trial. In the second experiment, the effects of common peroneal nerve (CPN) stimulation at short (2-4 ms) and at long (60-120 ms) conditioning test (C-T) intervals on the soleus H-reflex elicited every 5 s were established. Control and conditioned reflexes were recorded via a single differential bipolar electrode. In both experiments, H-reflexes were quantified by measuring their size as peak-to-peak amplitude and as area under the full-wave rectified waveform. The reflex responses recorded through two monopolar electrodes across stimulation intensities and ISIs measured as peak-to-peak amplitude had larger values than measured as area. In contrast, the magnitude of the reflexes, conditioned by CPN stimulation at either short or long C-T intervals and recorded via a single differential electrode, were not significantly different when measured as peak-to-peak amplitude or as area. Our findings indicate that monopolar recordings yield different reflex sizes depending on the method employed to measure the reflex size, and that the H-reflex measured as area might detect better the homosynaptic reflex depression. The lack of observing such differences with bipolar recordings might be related to changes of the reflex shape at a given stimulus intensity due to inhibitory inputs. The implications of our findings are discussed in respect to human reflex studies.     

On the methods employed to record and measure the human soleus H-reflex

The aim of this study was to investigate if the magnitude of the soleus H-reflex is different depending on the method employed to measure its size (peak-to-peak amplitude vs. area). In this study, 13 healthy human subjects participated, while the soleus H-reflex was induced via conventional methods. In the first experiment, the soleus H-reflex was recorded via two monopolar electrodes and was evoked at least at eight different stimulation intensities in respect to the recovery curve of the H-reflex and at three different inter-stimulus intervals (ISIs) (8, 5, and 2?s). The ISI refers to the time delay between the single pulses delivered to the posterior tibial nerve within a single trial. In the second experiment, the effects of common peroneal nerve (CPN) stimulation at short (2–4?ms) and at long (60–120?ms) conditioning test (C-T) intervals on the soleus H-reflex elicited every 5?s were established. Control and conditioned reflexes were recorded via a single differential bipolar electrode. In both experiments, H-reflexes were quantified by measuring their size as peak-to-peak amplitude and as area under the full-wave rectified waveform. The reflex responses recorded through two monopolar electrodes across stimulation intensities and ISIs measured as peak-to-peak amplitude had larger values than measured as area. In contrast, the magnitude of the reflexes, conditioned by CPN stimulation at either short or long C-T intervals and recorded via a single differential electrode, were not significantly different when measured as peak-to-peak amplitude or as area. Our findings indicate that monopolar recordings yield different reflex sizes depending on the method employed to measure the reflex size, and that the H-reflex measured as area might detect better the homosynaptic reflex depression. The lack of observing such differences with bipolar recordings might be related to changes of the reflex shape at a given stimulus intensity due to inhibitory inputs. The implications of our findings are discussed in respect to human reflex studies.     

Neural Mechanisms Influencing Interlimb Coordination during Locomotion in Humans: Presynaptic Modulation of Forearm H-Reflexes during Leg Cycling

Presynaptic inhibition of transmission between Ia afferent terminals and alpha motoneurons (Ia PSI) is a major control mechanism associated with soleus H-reflex modulation during human locomotion. Rhythmic arm cycling suppresses soleus H-reflex amplitude by increasing segmental Ia PSI. There is a reciprocal organization in the human nervous system such that arm cycling modulates H-reflexes in leg muscles and leg cycling modulates H-reflexes in forearm muscles. However, comparatively little is known about mechanisms subserving the effects from leg to arm. Using a conditioning-test (C-T) stimulation paradigm, the purpose of this study was to test the hypothesis that changes in Ia PSI underlie the modulation of H-reflexes in forearm flexor muscles during leg cycling. Subjects performed leg cycling and static activation while H-reflexes were evoked in forearm flexor muscles. H-reflexes were conditioned with either electrical stimuli to the radial nerve (to increase Ia PSI; C-T interval  = 20 ms) or to the superficial radial (SR) nerve (to reduce Ia PSI; C-T interval  = 37–47 ms). While stationary, H-reflex amplitudes were significantly suppressed by radial nerve conditioning and facilitated by SR nerve conditioning. Leg cycling suppressed H-reflex amplitudes and the amount of this suppression was increased with radial nerve conditioning. SR conditioning stimulation removed the suppression of H-reflex amplitude resulting from leg cycling. Interestingly, these effects and interactions on H-reflex amplitudes were observed with subthreshold conditioning stimulus intensities (radial n., ∼0.6×MT; SR n., ∼ perceptual threshold) that did not have clear post synaptic effects. That is, did not evoke reflexes in the surface EMG of forearm flexor muscles. We conclude that the interaction between leg cycling and somatosensory conditioning of forearm H-reflex amplitudes is mediated by modulation of Ia PSI pathways. Overall our results support a conservation of neural control mechanisms between the arms and legs during locomotor behaviors in humans.     

Enhanced D1 and D2 Inhibitions Induced by Low-Frequency Trains of Conditioning Stimuli: Differential Effects on H- and T-Reflexes and Possible Mechanisms

Mechanically evoked reflexes have been postulated to be less sensitive to presynaptic inhibition (PSI) than the H-reflex. This has implications on investigations of spinal cord neurophysiology that are based on the T-reflex. Preceding studies have shown an enhanced effect of PSI on the H-reflex when a train of ~10 conditioning stimuli at 1 Hz was applied to the nerve of the antagonist muscle. The main questions to be addressed in the present study are if indeed T-reflexes are less sensitive to PSI and whether (and to what extent and by what possible mechanisms) the effect of low frequency conditioning, found previously for the H-reflex, can be reproduced on T-reflexes from the soleus muscle. We explored two different conditioning-to-test (C-T) intervals: 15 and 100 ms (corresponding to D1 and D2 inhibitions, respectively). Test stimuli consisted of either electrical pulses applied to the posterior tibial nerve to elicit H-reflexes or mechanical percussion to the Achilles tendon to elicit T-reflexes. The 1 Hz train of conditioning electrical stimuli delivered to the common peroneal nerve induced a stronger effect of PSI as compared to a single conditioning pulse, for both reflexes (T and H), regardless of C-T-intervals. Moreover, the conditioning train of pulses (with respect to a single conditioning pulse) was proportionally more effective for T-reflexes as compared to H-reflexes (irrespective of the C-T interval), which might be associated with the differential contingent of Ia afferents activated by mechanical and electrical test stimuli. A conceivable explanation for the enhanced PSI effect in response to a train of stimuli is the occurrence of homosynaptic depression at synapses on inhibitory interneurons interposed within the PSI pathway. The present results add to the discussion of the sensitivity of the stretch reflex pathway to PSI and its functional role.     

Static otolithic drive alters presynaptic inhibition in soleus motor pool

The vestibular system has both direct and indirect connections to the soleus motor pool via the vestibulospinal and reticulospinal tracts. The exact nature of how this vestibular information is integrated within the spinal cord is largely unknown. The purpose of this study was to identify whether changes in static otolithic drive altered the amount of presynaptic inhibition in the soleus H-reflex pathway. Changes in static otolithic drive were investigated in sixteen healthy participants using a tilt table. Two presynaptic pathways (common peroneal and femoral) to the soleus H-reflex were tested in three weight conditions (supine, non-weight bearing, and weight bearing). The dependent variable was the peak-to-peak amplitude of the soleus H-reflex. Inhibition to the soleus motor pool through the common peroneal nerve pathway differed significantly during weight conditions and tilt. During tilt and non-weight bearing there was greater inhibition of the soleus H-reflex compared to supine, however, this effect was reversed during tilt and weight bearing. Facilitation from the femoral nerve pathway was reduced by tilt compared to supine, but this reduction was unaffected by weight condition. This supports a role of the vestibular system as providing complex, task-dependent presynaptic input to motoneurons in the lower limbs.     

Spinal reciprocal inhibition in human locomotion.

The purpose of this paper was to study spinal inhibition during several different motor tasks in healthy human subjects. The short-latency, reciprocal inhibitory pathways from the common peroneal (CP) nerve to the soleus muscle and from the tibial nerve to the tibialis anterior muscle were studied as a depression of ongoing voluntary electromyograph (EMG) activity. First, the effect of stimulus intensity on the amount of inhibition was examined to decide an appropriate stimulation to study the task-dependent modulation of inhibition. Then, the inhibition at one level of stimulation (1.5 x motor threshold) was investigated during standing, walking, and running. The change in slope of inhibition vs. EMG level, which approximates the fraction of ongoing activity that is inhibited, decreased with CP stimulation from 0.52 during standing to 0.30 during fast walking (6 km/h) to 0.17 during running at 9 km/h. Similarly, the slope decreased with tibial nerve stimulation from 0.68 (standing) to 0.42 (fast walking) to 0.35 (running at 9 km/h). All differences, except the last one, were highly significant (P < 0.01, Student's t-test). However, the difference between walking (0.42) and running (0.36) at the same speed (6 km/h) was not significant with tibial nerve stimulation and only significant at P < 0.05 with CP nerve stimulation (0.30, 0.20). Also, the difference between standing (0.52) and slow walking (3 km/h; 0.41) with CP stimulation was not significant, but it was significant (P < 0.01) with tibial nerve stimulation (0.68, 0.49). In conclusion, our findings indicate that spinal reciprocal inhibition decreases substantially with increasing speed and only changes to a lesser extent with task.     

Direct corticospinal pathways contribute to neuromuscular control of perturbed stance.

The antigravity soleus muscle (Sol) is crucial for compensation of stance perturbation. A corticospinal contribution to the compensatory response of the Sol is under debate. The present study assessed spinal, corticospinal, and cortical excitability at the peaks of short- (SLR), medium- (MLR), and long-latency responses (LLR) after posterior translation of the feet. Transcranial magnetic stimulation (TMS) and peripheral nerve stimulation were individually adjusted so that the peaks of either motor evoked potential (MEP) or H reflex coincided with peaks of SLR, MLR, and LLR, respectively. The influence of specific, presumably direct, corticospinal pathways was investigated by H-reflex conditioning. When TMS was triggered so that the MEP arrived in the Sol at the same time as the peaks of SLR and MLR, EMG remained unaffected. Enhanced EMG was observed when the MEP coincided with the LLR peak (P < 0.001). Similarly, conditioning of the H reflex by subthreshold TMS facilitated H reflexes only at LLR (P < 0.001). The earliest facilitation after perturbation occurred after 86 ms. The TMS-induced H-reflex facilitation at LLR suggests that increased cortical excitability contributes to the augmentation of the LLR peaks. This provides evidence that the LLR in the Sol muscle is at least partly transcortical, involving direct corticospinal pathways. Additionally, these results demonstrate that approximately 86 ms after perturbation, postural compensatory responses are cortically mediated.     

Invariable H-reflex and sustained facilitation of stretch reflex with heightened sympathetic outflow

Stretch reflex shows sustained (3-min) increase with heightened sympathetic outflow [Hjortskov N, Skotte J, Hye-Knudsen C, Fallentin N. Sympathetic outflow enhances the stretch reflex response in the relaxed soleus muscle in humans. J Appl Physiol 2005;98:1366–70], but it is unknown if it accompanies a sustained increase in H-reflex. The purpose of the study was to test if there is a sustained facilitation in the H-reflex in the human soleus muscle during a variety of sustained tasks that are known to elevate sympathetic outflow. Mean arterial blood pressure, heart rate, and H- and stretch reflexes in the relaxed soleus muscle were obtained in healthy young adults who performed mental arithmetic, static handgrip exercise, post-handgrip ischemia, and cold stimulation. Each task lasted 3 min with a 3-min rest in between tasks. Data were analyzed for the initial 30 s and entire 3 min of each task. There was a heightened cardiovascular response in all tasks for both durations of analysis. An increase in H-reflex amplitude was not observed for either the initial or entire duration of the analysis. The tasks increased stretch reflex amplitude for both durations of analysis. Invariable H-reflex and sustained facilitation of stretch reflex with heightened sympathetic outflow would imply sympathetic modulation of muscle spindle sensitivity.