Hiroki Obata

Differential effects of plantar cutaneous afferent excitation on soleus stretch and H-reflex

Previous studies have demonstrated that plantar cutaneous afferents can adjust motoneuron excitability, which may contribute significantly to the control of human posture and locomotion. However, the role of plantar cutaneous afferents in modulating the excitability of stretch and H‐reflex with respect to the location of their excitation remains unclear. In the present study, it was hypothesized that electrical stimulation delivered to the sole of the foot might be followed by modulation of spinal excitability that depends on: (1) the stimulation location and (2) the reflex studied. In these experiments, conditioned and unconditioned stretch and H‐reflexes were evoked in 16 healthy subjects in a seated position. Both reflexes were conditioned by non‐noxious electrical plantar cutaneous afferent stimulation at two different sites, the heel and metatarsal regions, at four different conditioning–test (CT) intervals. The conditioning stimulation delivered to the heel caused a significant facilitation of the soleus stretch reflex for all CT intervals, whereas the soleus H‐reflex had significant facilitation only at CT interval of 50 ms and significant inhibition at longer CT intervals. Stimulation delivered to the metatarsal region, however, resulted mainly in reduced stretch and H‐reflex sizes. This study extends the reported findings on the contribution of plantar cutaneous afferents within spinal interneuron reflex circuits as a function of their location and the reflex studied. Muscle Nerve, 2008

Facilitation of both stretch reflex and corticospinal pathways of the tibialis anterior muscle during standing in humans

Excitability of both stretch reflex (SR) and motor evoked potential (MEP) elicited in the tibialis anterior (TA) muscle by transcranial magnetic stimulation were tested in standing humans. The results demonstrated significantly greater values for both SR and MEP in the TA while standing than while in the supine posture, although background electromyographic activity was silent in the two conditions. Taken together with previous reports that both pathways are facilitated in the TA at the early stance phase of human walking, our findings suggest that a common neural mechanism underlies both observations, one that might be functionally relevant for securing ankle joint stabilization during upright standing.

Age-related changes of the stretch reflex excitability in human ankle muscles

The purpose of this study was to characterize the effects of aging on the stretch reflex in the ankle muscles, and in particular to compare the effects on the ankle dorsi-flexor (tibialis anterior: TA) and the plantar-flexor (soleus: SOL). Stretch reflex responses were elicited in the TA and SOL at rest and during weak voluntary contractions in 20 elderly and 23 young volunteers. The results indicated that, in the TA muscle, the elderly group had a remarkably larger long-latency reflex (LLR), whereas no aging effect was found in the short latency reflex (SLR). These results were very different from those in the SOL muscle, which showed significant aging effects in the SLR and medium latency reflex (MLR), but not in the LLR. Given the fact that the LLR of the TA stretch reflex includes the cortical pathway, it is probable that the effects of aging on the TA stretch reflex involve alterations not only at the spinal level but also at the cortical level. The present results indicate that the stretch reflexes of each of the ankle antagonistic muscles are affected differently by aging, which might have relevance to the neural properties of each muscle.

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.

Posture-related modulation of cortical excitability in the tibialis anterior muscle in humans

Corticospinal excitability in the lower leg muscles is enhanced during standing as compared to other postures. In the present study, we investigated how the excitability of intracortical circuits that control the tibialis anterior muscle (TA) is modulated during standing. Short-interval intracortical inhibition (SICI) and intracortical facilitation (ICF) were assessed by the paired-pulse transcranial magnetic stimulation technique during standing (STD) and sitting (SIT) with a comparable background activity level in both the soleus and the TA muscle. The results demonstrated that SICI was less effective during STD than during SIT, whereas ICF was more effective during STD than during SIT. These findings suggest that the excitabilities of these cortical neural circuits are modulated depending on posture. A decrease in SICI and an increase in ICF may reflect subliminal enhancement of the cortical excitability in the TA muscle during standing as compared with that during sitting.

The Effects of Temporal and Spatial Predictions on Stretch Reflexes of Ankle Flexor and Extensor Muscles While Standing.

The purpose of the present study was to investigate how stretch reflex (SR) responses in the ankle extensor (soleus: SOL) and flexor (tibialis anterior: TA) muscles would be modulated with temporal and/or spatial predictions of external perturbations and whether their effects are specific to the standing posture. SR responses in the SOL/TA were elicited by imposing quick ankle toes-up/toes-down rotations while standing upright and in the supine position. We designed four experimental conditions based on pre-information about perturbations: no information (No Cue), the timing of the perturbation onset (TIM), the direction of the perturbation (DIR), and both the timing and direction of the perturbation (TIM/DIR). Each condition was separated and its order was counterbalanced. In the SR of TA evoked by toes-down rotation, integrated electromyography activities of the late component were significantly reduced in the TIM and TIM/DIR conditions as compared with those in the No Cue and DIR conditions. The occurrence rate of late SR components that reflects how often the reflex response was observed was also lower in the TIM and TIM/DIR conditions as compared with that in the No Cue and DIR conditions. On the other hand, no significant changes were seen among the four conditions in the early SR component in the TA and both SR components in the SOL. The same results in the occurrence rate were found in the supine position. The present results suggest (1) only temporal predictions have a remarkable effect on the SR excitability of the TA, and (2) this effect is independent of posture.

Mode-dependent control of human walking and running as revealed by split-belt locomotor adaptation

ABSTRACT Here, we investigate the association of neural control between walking and running, and in particular, how these two gait modes at different velocities are controlled by the central nervous system. The subjects were fully adapted by acquiring modified motor patterns to either walk or run on a split-belt treadmill driven in split mode (asymmetry in the velocities of two belts at 1.0 and 2.0 m s−1). Subsequently, we tested how the adaptation affected walking and running at three different velocities in the tied mode (equal belt velocities). At 0.75 m s−1, we found a preference to walk, at 1.50 m s−1, there was a preference to both walk and run, and at a velocity of 2.25 m s−1 there was a preference to run. Both walking and running on the split belt resulted in the emergence of a significant aftereffect (asymmetrical movement) at all of the velocities tested when walking after adapting to walk and running after adapting to run. However, for contrasting modes (i.e. running after adapting to walk and walking after adapting to run), such aftereffects were far less evident at all velocities; thus showing only limited transfer across gait modes. The results demonstrate a clear mode dependency in the neural control of human walking and running. In addition, only for walking, was there a degree of velocity dependency. Summary: Walking and running are not simply dependent on velocity, but are controlled by the CNS as two completely distinct forms of locomotion in humans.

Distinct Motor Strategies Underlying Split- Belt Adaptation in Human Walking and Running

The aim of the present study was to elucidate the adaptive and de-adaptive nature of human running on a split-belt treadmill. The degree of adaptation and de-adaptation was compared with those in walking by calculating the antero-posterior component of the ground reaction force (GRF). Adaptation to walking and running on a split-belt resulted in a prominent asymmetry in the movement pattern upon return to the normal belt condition, while the two components of the GRF showed different behaviors depending on the gaits. The anterior braking component showed prominent adaptive and de-adaptive behaviors in both gaits. The posterior propulsive component, on the other hand, exhibited such behavior only in running, while that in walking showed only short-term aftereffect (lasting less than 10 seconds) accompanied by largely reactive responses. These results demonstrate a possible difference in motor strategies (that is, the use of reactive feedback and adaptive feedforward control) by the central nervous system (CNS) for split-belt locomotor adaptation between walking and running. The present results provide basic knowledge on neural control of human walking and running as well as possible strategies for gait training in athletic and rehabilitation scenes.

Neural effects of muscle stretching on the spinal reflexes in multiple lower-limb muscles

While previous studies have shown that muscle stretching suppresses monosynaptic spinal reflex excitability in stretched muscles, its effects on non-stretched muscles is still largely unknown. The purpose of this study was to examine the effects of muscle stretching on monosynaptic spinal reflex in non-stretched muscles. Ten healthy male subjects participated in this study. Muscle stretching of the right triceps surae muscle was performed using a motor torque device for 1 minute. Three different dorsiflexion torques (at approximately 5, 10, and 15 Nm) were applied during muscle stretching. Spinal reflexes evoked by transcutaneous spinal cord stimulation were recorded in both the lower-limb muscles before, during, and at 0 and 5 min following muscle stretching. The amplitudes of the spinal reflexes in both the stretched and non-stretched muscles in the right (ipsilateral) leg were smaller during stretching compared to before, and at 0 and 5 min after stretching. Furthermore, the degree of reduction in the amplitude of the spinal reflexes in the right (ipsilateral) leg muscles increased significantly as the dorsiflexion torque (i.e., stretching of the right triceps surae muscles) increased. In contrast, reduction in the amplitude of the spinal reflexes with increasing dorsiflexion torque was not seen in the left (contralateral) leg muscles. Our results clearly indicate that muscle stretching has inhibitory effects on monosynaptic spinal reflexes, not only in stretched muscles, but also in non-stretched muscles of the ipsilateral leg.

Short‐term effect of electrical nerve stimulation on spinal reciprocal inhibition during robot‐assisted passive stepping in humans

The purpose of this study was to investigate the effect of electrical stimulation to the common peroneal nerve (CPN) on the spinal reflex and reciprocal inhibition (RI) during robot‐assisted passive ground stepping (PGS) in healthy subjects. Five interventions were applied for 30 min in healthy subjects: PGS alone; strong CPN stimulation [50% of the maximal tibialis anterior (TA) M‐wave, functional electrical stimulation (FES)] alone; weak CPN stimulation [just above the MT for the TA muscle, therapeutic electrical stimulation (TES)] alone; PGS with FES; and PGS with TES. FES and TES were applied intermittently to the CPN at 25 Hz. The soleus (Sol) H‐reflex and RI, which was assessed by conditioning the Sol H‐reflex with CPN stimulation, were investigated before (baseline), and 5, 15 and 30 min after each intervention. The amplitudes of the Sol H‐reflex were not significantly different after each intervention as compared with the baseline values. The amounts of RI were significantly decreased 5 min after PGS with FES as compared with the baseline values, whereas they were significantly increased 5 and 15 min after PGS with TES. The other interventions did not affect the amount of RI. These results suggest that interventions that combined PGS with CPN stimulation changed the spinal RI in an intensity‐dependent manner.