Tetsuya Ogawa

Effect of sensory inputs on the soleus H-reflex amplitude during robotic passive stepping in humans

We investigated the modulation of the soleus (Sol) Hoffmann (H-) reflex excitability by peripheral sensory inputs during passive stepping using a robotic-driven gait orthosis in healthy subjects and spinal cord-injured patients. The Sol H-reflex was evoked at standing and at six phases during passive stepping in 40 and 100% body weight unloaded conditions. The Sol H-reflex excitability was significantly inhibited during passive stepping when compared with standing posture at each unloaded condition. During passive stepping, the H-reflex amplitude was significantly smaller in the early- and mid-swing phases than in the stance phase, which was similar to the modulation pattern previously reported for normal walking. No significant differences were observed in the H-reflex amplitude between the two unloaded conditions during passive stepping. The reflex depression observed at the early part of the swing phase during passive stepping might be attributed to the sensory inputs elicited by flexion of the hip and knee joints. The present study provides evidence that peripheral sensory inputs have a significant role in phase-dependent modulation of the Sol H-reflex during walking, and that the Sol H-reflex excitability might be less affected by load-related afferents during walking.

Predictive control of ankle stiffness at heel contact is a key element of locomotor adaptation during split-belt treadmill walking in humans

Split-belt treadmill walking has been extensively utilized as a useful model to reveal the adaptability of human bipedal locomotion. While previous studies have clearly identified different types of locomotor adaptation, such as reactive and predictive adjustments, details of how the gait pattern would be adjusted are not fully understood. To gain further knowledge of the strategies underlying split-belt treadmill adaptation, we examined the three-dimensional ground reaction forces (GRF) and lower limb muscle activities during and after split-belt treadmill walking in 22 healthy subjects. The results demonstrated that the anterior component of the GRF (braking force) showed a clear pattern of adaptation and subsequent aftereffects. The muscle activity in the tibialis anterior muscle during the early stance phase was associated with the change of braking force. In contrast, the posterior component of GRF (propulsive force) showed a consistent increase/decrease in the fast/slow leg during the adaptation period and was not followed by subsequent aftereffects. The muscle activity in the gastrocnemius muscle during the stance phase gradually decreased during the adaptation phase and then showed a compensatory reaction during the washout phase. The results indicate that predictive feedforward control is required to set the optimal ankle stiffness in preparation for the impact at the heel contact and passive feedback control is used for the production of reflexively induced propulsive force at the end of the stance phase during split-belt treadmill adaptation. The present study provides information about the detailed mechanisms underlying split-belt adaptation and should be useful for the construction of specific rehabilitation protocols.

Limited Transfer of Newly Acquired Movement Patterns across Walking and Running in Humans

The two major modes of locomotion in humans, walking and running, may be regarded as a function of different speed (walking as slower and running as faster). Recent results using motor learning tasks in humans, as well as more direct evidence from animal models, advocate for independence in the neural control mechanisms underlying different locomotion tasks. In the current study, we investigated the possible independence of the neural mechanisms underlying human walking and running. Subjects were tested on a split-belt treadmill and adapted to walking or running on an asymmetrically driven treadmill surface. Despite the acquisition of asymmetrical movement patterns in the respective modes, the emergence of asymmetrical movement patterns in the subsequent trials was evident only within the same modes (walking after learning to walk and running after learning to run) and only partial in the opposite modes (walking after learning to run and running after learning to walk) (thus transferred only limitedly across the modes). Further, the storage of the acquired movement pattern in each mode was maintained independently of the opposite mode. Combined, these results provide indirect evidence for independence in the neural control mechanisms underlying the two locomotive modes.

Distinct sets of locomotor modules control the speed and modes of human locomotion

Although recent vertebrate studies have revealed that different spinal networks are recruited in locomotor mode- and speed-dependent manners, it is unknown whether humans share similar neural mechanisms. Here, we tested whether speed- and mode-dependence in the recruitment of human locomotor networks exists or not by statistically extracting locomotor networks. From electromyographic activity during walking and running over a wide speed range, locomotor modules generating basic patterns of muscle activities were extracted using non-negative matrix factorization. The results showed that the number of modules changed depending on the modes and speeds. Different combinations of modules were extracted during walking and running, and at different speeds even during the same locomotor mode. These results strongly suggest that, in humans, different spinal locomotor networks are recruited while walking and running, and even in the same locomotor mode different networks are probably recruited at different speeds.

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.

Speed dependency in α-motoneuron activity and locomotor modules in human locomotion: indirect evidence for phylogenetically conserved spinal circuits

Coordinated locomotor muscle activity is generated by the spinal central pattern generators (CPGs). Vertebrate studies have demonstrated the following two characteristics of the speed control mechanisms of the spinal CPGs: (i) rostral segment activation is indispensable for achieving high-speed locomotion; and (ii) specific combinations between spinal interneuronal modules and motoneuron (MN) pools are sequentially activated with increasing speed. Here, to investigate whether similar control mechanisms exist in humans, we examined spinal neural activity during varied-speed locomotion by mapping the distribution of MN activity in the spinal cord and extracting locomotor modules, which generate basic MN activation patterns. The MN activation patterns and the locomotor modules were analysed from multi-muscle electromyographic recordings. The reconstructed MN activity patterns were divided into the following three patterns depending on the speed of locomotion: slow walking, fast walking and running. During these three activation patterns, the proportion of the activity in rostral segments to that in caudal segments increased as locomotion speed increased. Additionally, the different MN activation patterns were generated by distinct combinations of locomotor modules. These results are consistent with the speed control mechanisms observed in vertebrates, suggesting phylogenetically conserved spinal mechanisms of neural control of locomotion.

Rhythmic arm swing enhances patterned locomotor-like muscle activity in passively moved lower extremities

The use of driven gait orthosis (DGO) has drawn attention in gait rehabilitation for patients after central nervous system (CNS) lesions. By imposing a passive locomotor‐like kinematic pattern, the neural mechanisms responsible for locomotion can be activated as in a normal gait. To further enhance this activity, discussions on possible intervention are necessary. Given the possible functional linkages between the upper and lower limbs, we investigated in healthy subjects the degree of modification in the lower limb muscles during DGO‐induced passive gait by the addition of swing movement in the upper extremity. The results clearly showed that muscle activity in the ankle dorsiflexor TA muscle was significantly enhanced when the passive locomotor‐like movement was accompanied by arm swing movement. The modifications in the TA activity were not a general increase through the stride cycles, but were observed under particular phases as in normal gaits. Voluntary effort to swing the arms may have certain effects on the modification of the muscle activity. The results provide clinical implications regarding the usefulness of voluntary arm swing movement as a possible intervention in passive gait training using DGO, since ordinary gait training using DGO does not induce spontaneous arm swing movement despite its known influence on the lower limb movement.

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.

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.

Velocity-dependent suppression of the soleus H-reflex during robot-assisted passive stepping.

The amplitude of the Hoffmann (H)-reflex in the soleus (Sol) muscle is known to be suppressed during passive stepping compared with during passive standing. The reduction of the H-reflex is not due to load-related afferent inputs, but rather to movement-related afferent inputs from the lower limbs. To elucidate the underlying neural mechanisms of this inhibition, we investigated the effects of the stepping velocity on the Sol H-reflex during robot-assisted passive stepping in 11 healthy subjects. The Sol H-reflexes were recorded during passive standing and stepping at five stepping velocities (stride frequencies: 14, 21, 28, 35, and 42 min(-1)) in the air. The Sol H-reflexes were significantly inhibited during passive stepping as compared with during passive standing, and reduced in size as the stepping velocity increased. These results indicate that the extent of H-reflex suppression increases with increasing movement-related afferent inputs from the lower limbs during passive stepping. The velocity dependence suggests that the Ia afferent inputs from lower-limb muscles around the hip and knee joints are most probably related to this inhibition.