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Rolling walkers are common walking aids for individuals with poor physical fitness or balance impairments. There is no doubt that rolling walkers are useful in assisting locomotion. On the other hand, it is arguable that walking with rolling walkers (WW) is effective for maintaining or restoring the nervous systems that are recruited during conventional walking (CW). This is because the differences and similarities of the neural control of these locomotion forms remain unknown. The purpose of the present study was to compare the neural control of WW and CW from the perspective of a split-belt adaptation paradigm and reveal how the adaptations that take place in WW and CW would affect each other. The anterior component of the ground reaction (braking) forces was measured during and after walking on a split-belt treadmill by 10 healthy subjects, and differences in the peak braking forces between the left and right sides were calculated as the index of the split-belt adaptation (the degree of asymmetry). The results demonstrated that (1) WW enabled subjects to respond to the split-belt condition immediately after its start as compared to CW; (2) the asymmetry movement pattern acquired by the split-belt adaptation in one gait mode (i.e., CW or WW) was less transferable to the other gait mode; (3) the asymmetry movement pattern acquired by the split-belt adaptation in CW was not completely washed out by subsequent execution in WW and vice versa. The results suggest unique control of WW and the specificity of neural control between WW and CW; use of the walkers is not necessarily appropriate as training for CW from the perspective of neural control., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Walking with pulling force fields acting at the body center of mass (in the forward or backward directions) is compatible with inclined walking and is used in clinical practice for gait training. From the perspective of known differences in the motor strategies that underlie walking with the respective force fields, the present study elucidated whether the adaptation acquired by walking on a split-belt treadmill with either one of the force fields affects subsequent walking in a force field in the opposite directions. Walking with the force field induced an adaptive and de-adaptive behavior of the subjects, with the aspect evident in the braking and propulsive impulses of the ground reaction force (difference in the peak value between the left and right sides for each stride cycle) as parameters. In the parameters, the adaptation acquired during walking with a force field acting in one direction was transferred to that in the opposite direction only partially. Furthermore, the adaptation that occurred while walking in a force field in one direction was rarely washed out by subsequent walking in a force field in the opposite direction and thus was maintained independently of the other. These results demonstrated possible independence in the neural functional networks capable of controlling walking in each movement task with an opposing force field., (© 2023. The Author(s).)

Action observation (AO) and motor imagery (MI) are used for the rehabilitation of patients who face difficulty walking. Rehabilitation involving AO, MI, and AO combined with MI (AO+MI) facilitates gait recovery after neurological disorders. However, the mechanism by which it positively affects gait function is unclear. We previously examined the neural mechanisms underlying AO and MI of walking, focusing on AO+MI and corticospinal and spinal motor neuron excitability, which play important roles in gait function. Herein, we investigated the effects of a short intervention using AO+MI of walking on the corticospinal and spinal motor neuron excitability and MI ability of participants. Twelve healthy individuals participated in this study, which consisted of a 20 min intervention. Before the experiment, we measured MI ability using the Vividness of Movement Imagery Questionnaire-2 (VMIQ-2). We used motor evoked potential and F-wave measurements to evaluate the corticospinal and spinal motor neuron excitability at rest, pre-intervention, 0 min, and 15 min post-intervention. We also measured corticospinal excitability during MI of walking and the participant's ability to perform MI using a visual analog scale (VAS). There were no significant changes in corticospinal and spinal motor neuron excitability during and after the intervention using AO+MI (p>0.05). The intervention temporarily increased VAS scores, thus indicating clearer MI (p<0.05); however, it did not influence corticospinal excitability during MI of walking (p>0.05). Furthermore, there was no significant correlation between the VMIQ-2 and VAS scores and changes in corticospinal and spinal motor neuron excitability. Therefore, one short intervention using AO+MI increased MI ability in healthy individuals; however, it was insufficient to induce plastic changes at the cortical and spinal levels. Moreover, the effects of intervention using AO+MI were not associated with MI ability. Our findings provide information about intervention using AO+MI in healthy individuals and might be helpful in planning neurorehabilitation strategies., Competing Interests: The authors have declared that no competing interests exist.

Modification of ongoing walking movement to fit changes in external environments requires accurate voluntary control. In cats, the motor and posterior parietal cortices have crucial roles for precisely adjusting limb trajectory during walking. In human walking, however, it remains unclear which cortical information contributes to voluntary gait modification. In this study, we investigated cortical activity changes associated with visually guided precision stepping using electroencephalography source analysis. Our results demonstrated frequency- and gait-event-dependent changes in the cortical power spectrum elicited by voluntary gait modification. The main differences between normal walking and precision stepping were as follows: (a) the alpha, beta or gamma power decrease during the swing phases in the sensorimotor, anterior cingulate and parieto-occipital cortices, and (b) a power decrease in the theta, alpha and beta bands and increase in the gamma band throughout the gait cycle in the parieto-occipital cortex. Based on the previous knowledge of brain functions, the former change was considered to be related to execution and planning of leg movement, while the latter change was considered to be related to multisensory integration and motor awareness. Therefore, our results suggest that the gait modification is achieved by higher cortical involvements associated with different sensorimotor-related functions across multiple cortical regions including the sensorimotor, anterior cingulate and parieto-occipital cortices. The results imply the critical importance of the cortical contribution to voluntary modification in human locomotion. Further, the observed cortical information related to voluntary gait modification would contribute to developing volitional control systems of brain-machine interfaces for walking rehabilitation., (© 2021 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

Underwater walking is one of the most common hydrotherapeutic exercises. Therefore, understanding muscular control during underwater walking is important for optimizing training regimens. The effects of the water environment on walking are mainly related to the hydrostatic and hydrodynamic theories of buoyancy and drag force. To date, muscular control during underwater walking has been investigated at the individual muscle level. However, it is recognized that the human nervous system modularly controls multiple muscles through muscle synergies, which are sets of muscles that work together. We found that the same set of muscle synergies was shared between the two walking tasks. However, some task-dependent modulation was found in the activation combination across muscles and temporal activation patterns of the muscle synergies. The results suggest that the human nervous system modulates activation of lower-limb muscles during water walking by finely tuning basic locomotor muscle synergies that are used during land walking to meet the biomechanical requirements for walking in the water environment., (© 2021. The Author(s).)

Action observation (AO) and motor imagery (MI) are motor simulations which induce cortical activity related to execution of observed and imagined movements. Neuroimaging studies have mainly investigated where the cortical activities during AO and MI of movements are activated and if they match those activated during execution of the movements. However, it remains unclear how cortical activity is modulated; in particular, whether activity depends on observed or imagined phases of movements. We have previously examined the neural mechanisms underlying AO and MI of walking, focusing on the combined effect of AO with MI (AO+MI) and phase dependent modulation of corticospinal and spinal reflex excitability. Here, as a continuation of our previous studies, we investigated cortical activity depending on gait phases during AO and AO+MI of walking by using electroencephalography (EEG); 64-channel EEG signals were recorded in which participants observed walking with or without imagining it, respectively. EEG source and spectral analyses showed that, in the sensorimotor cortex during AO+MI and AO, the alpha and beta power were decreased, and power spectral modulations depended on walking phases. The phase dependent modulations during AO+MI, but not during AO, were like those which occur during actual walking as reported by previous walking studies. These results suggest that combinatory effects of AO+MI could induce parts of the phase dependent activation of the sensorimotor cortex during walking even without any movements. These findings would extend understanding of the neural mechanisms underlying walking and cognitive motor processes and provide clinically beneficial information towards rehabilitation for patients with neurological gait dysfunctions., Competing Interests: Declaration of Competing Interest The authors report no conflict of interest., (Copyright © 2020 The Authors. Published by Elsevier Inc. All rights reserved.)

Pole walking (PW) has received attention not only as a whole-body exercise that can be adapted for elderly people with poor physical fitness but also as a possible intervention for the restoration of gait function in normal walking without the use of poles (i.e., conventional walking CW). However, the characteristics of PW, especially how and why PW training affects CW, remain unclear. The purpose of this study was to examine the characteristics of locomotor adaptation in PW from the perspective of kinematic variables. For this purpose, we compared the locomotor adaptation in PW and CW to that when walking on a split-belt treadmill in terms of spatial and temporal coordination. The result showed that adaptations to the split-belt treadmill in PW and CW were found only in interlimb parameters (step length and double support time ratios (fast/slow limb)), not in intralimb parameters (stride length and stance time ratios). In these interlimb parameters, the movement patterns acquired through split-belt locomotor adaptations (i.e., the aftereffects) were transferred between CW and PW regardless of whether the novel movement patterns were learned in CW or PW. The aftereffects of double support time and step length learned in CW were completely washed out by the subsequent execution in PW. On the other hand, the aftereffect of double support time learned in PW was not completely washed out by the subsequent execution in CW, whereas the aftereffect of step length learned in PW was completely washed out by the subsequent execution in CW. These results suggest that the neural mechanisms related to controlling interlimb parameters are shared between CW and PW, and it is possible that, in interlimb coordination, temporal coordination is preferentially stored in adaptation during PW.

Pole walking (PW), a form of locomotion in which a person holds a pole in each hand, enhances the involvement of alternating upper-limb movement. While this quadruped-like walking increases postural stability for bipedal conventional walking (CW), in terms of the neural controlling mechanisms underlying the two locomotion forms (PW and CW), the similarities and differences remain unknown. The purpose of this study was to compare the neural control of PW and CW from the perspective of locomotor adaptation to a novel environment on a split-belt treadmill. We measured the anterior component of the ground reaction (braking) force during and after split-belt treadmill walking in 12 healthy subjects. The results demonstrated that (1) PW delayed locomotor adaptation when compared with CW; (2) the degrees of transfer of the acquired movement pattern to CW and PW were not different, regardless of whether the novel movement pattern was learned in CW or PW; and (3) the movement pattern learned in CW was washed out by subsequent execution in PW, whereas the movement pattern learned in PW was not completely washed out by subsequent execution in CW. These results suggest that the neural control mechanisms of PW and CW are not independent, and it is possible that PW could be a locomotor behavior built upon a basic locomotor pattern of CW.

Action observation (AO) and motor imagery (MI) are used for motor impairment rehabilitation after neurological disorders, such as spinal cord injuries or strokes. Clinical studies have reported that rehabilitation using AO and MI is effective in restoring motor function. Our previous study showed a difference in the modulation of corticospinal excitability only during action observation (AO) of walking and AO of walking combined with motor imagery (AO + MI). However, it is unclear whether AO and AO + MI can modulate spinal reflex excitability as well as corticospinal excitability. Thus, the purpose of this study was to investigate the modulation of the Hoffmann reflex (H-reflex) during AO and AO + MI. We measured H-reflex in the soleus muscle under the following conditions: (1) control, (2) AO, and (3) AO + MI. Posterior tibial nerve electrical stimulation, which induces H-reflex, was applied at the following four walking phases: 1) mid-stance, 2) terminal-stance, 3) early-swing, and 4) terminal-swing. Our results showed that AO + MI significantly increases H-reflex over AO alone, regardless of phase and that AO significantly modulates H-reflex depending on the observed phase. These findings suggested that spinal reflex excitability can be modulated during AO and AO + MI and that neural effects are different in AO and AO + MI., (Copyright © 2018. Published by Elsevier B.V.)

The adaptability of human bipedal locomotion has been studied using split-belt treadmill walking. Most of previous studies utilized experimental protocol under remarkably different split ratios (e.g. 1:2, 1:3, or 1:4). While, there is limited research with regard to adaptive process under the small speed ratios. It is important to know the nature of adaptive process under ratio smaller than 1:2, because systematic evaluation of the gait adaptation under small to moderate split ratios would enable us to examine relative contribution of two forms of adaptation (reactive feedback and predictive feedforward control) on gait adaptation. We therefore examined a gait behavior due to on split-belt treadmill adaptation under five belt speed difference conditions (from 1:1.2 to 1:2). Gait parameters related to reactive control (stance time) showed quick adjustments immediately after imposing the split-belt walking in all five speed ratios. Meanwhile, parameters related to predictive control (step length and anterior force) showed a clear pattern of adaptation and subsequent aftereffects except for the 1:1.2 adaptation. Additionally, the 1:1.2 ratio was distinguished from other ratios by cluster analysis based on the relationship between the size of adaptation and the aftereffect. Our findings indicate that the reactive feedback control was involved in all the speed ratios tested and that the extent of reaction was proportionally dependent on the speed ratio of the split-belt. On the contrary, predictive feedforward control was necessary when the ratio of the split-belt was greater. These results enable us to consider how a given split-belt training condition would affect the relative contribution of the two strategies on gait adaptation, which must be considered when developing rehabilitation interventions for stroke patients.

The aim of the current study was to clarify how corticospinal excitability (CSE) is modulated during the observation of the action of walking, combined with motor imagery (MI). We examined the difference in phase modulation of CSE with and without MI during the observation of the action of walking. We measured motor-evoked potentials in tibialis anterior and soleus muscles using transcranial magnetic stimulation under the following conditions: (1) baseline, (2) action observation (AO), and (3) AO and MI. Our results showed that, compared with AO alone, AO and MI of walking increased motor-evoked potentials. Our results suggest that CSE may be facilitated during AO and MI of walking, rather than during AO alone. Further, we postulated that AO and MI of walking may facilitate CSE in the tibialis anterior in a manner such as that evoked during actual walking.

Coordinated locomotor muscle activity is generated by the spinal central pattern generators (CPGs), which are modulated by peripheral and supraspinal inputs. The CPGs would consist of multiple motor modules generating basic muscle activity, which are distributed rostrocaudally along the spinal cord. To activate the motor modules in proper sequence, rostrocaudally traveling waves of activation in the spinal cord are important mechanisms in the CPGs. The traveling waves of activation have been observed in nonhuman vertebrates. However, they have not yet been confirmed during human locomotion. Although, rostrocaudal wave-like activations in the spinal cord were observed during walking in humans in a previous study, the propagation shifted rostrally toward the upper lumbar segments at foot contact. Here, using an air stepping task to remove the foot-contact interactions, we examined whether the traveling wave mechanism exists in the human spinal circuits based on the activation sequence of motor modules and their topography. We measured electromyographic activity of lower leg muscles during the air-stepping task. Then, we extracted motor modules (i.e., basic patterns of sets of muscle activations: muscle synergies) from the measured muscle activities using nonnegative matrix factorization method. Next, we reconstructed motoneuron (MN) activity from each module activity based on myotomal charts. We identified four types of motor modules from muscle activities during the air-stepping task. Each motor module represented different sets of synergistic muscle activations. MN clusters innervating each motor module were sequentially activated from the rostral to caudal region in the spinal cord, from the initial flexion to the last extension phase during air-stepping. The rostrocaudally sequential activation of MN clusters suggests the possibility that rostrocaudally traveling waves exist in human locomotor spinal circuits. The present results advance the understanding of human locomotor control mechanisms, and provide important insights into the evolution of locomotor networks in vertebrates., (© 2017 The Authors. Physiological Reports published by Wiley Periodicals, Inc. on behalf of The Physiological Society and the American Physiological Society.)

Studies of robot-assisted passive stepping paradigms have reported that movement-related afferent inputs strongly inhibit the excitability of the Hoffmann (H) reflex in the soleus (Sol) during walking. However, it is unknown if movement-related afferent inputs have the same effect on the excitability of spinal reflexes in the other lower-limb muscles that are involved in normal walking in healthy subjects. The aim of this study was to examine the effects of movement-related afferent inputs on the spinal reflexes in lower-limb muscles during walking. Spinal reflexes that were elicited by transcutaneous spinal cord stimulation (tSCS) were recorded during passive air standing and air stepping at three stepping velocities (stride frequencies: 14, 25, and 36 strides/min). The amplitude of the spinal reflexes was reduced in most of the recorded muscles during passive air stepping compared with air standing. Furthermore, in the Sol and lateral gastrocnemius, the amplitude of the reflexes during air stepping significantly decreased as stride frequency increased. These results demonstrate that movement-related afferent inputs inhibit spinal reflexes in the Sol and other lower-limb muscles during walking., (Copyright © 2016 Elsevier Ireland Ltd. All rights reserved.)

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., (© 2015. Published by The Company of Biologists Ltd.)

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., (© 2015 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

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.

A common source of error when detecting heel-strike moments utilizing split force-plate treadmills is unwillingly stepping on contra-lateral force-plate. In this study, we quantified this error when heel-strike was detected based on such erroneous data and compared three methods to investigate how well the heel-strikes and stride-intervals were detected with erroneous data. Eleven subjects walked on a split force-plate treadmill for more than 20min. We used 20N and 50% body-weight thresholds to detect the heel-strike moments (HS20N and HS50%, respectively). Besides, we used linear approximation to estimate the unaffected force profile from affected force-plate data, and subsequently to detect the heel-strike moments (HSest). We used heel-strike moments detected by a foot-switch as a reference to compare accuracy of HS20N, HS50% and HSest. HS20N and HSest detected heel-strike moments accurately for unaffected force-plate data (median(max) errors for all subjects: 9(23) and 9(37) ms) but HS50% showed significantly larger errors (52(74) ms). Unlike HS50% and HSest, HS20N was considerably affected by the affected force-plate data (23(68) ms). The error in stride-interval measurement was relatively small using any methods for unaffected force-plate data (3(7), 6(8), and 6(12) ms), while stride-interval errors were large for some subjects when using HS20N for affected data (6(175) ms). We concluded that unwillingly stepping on contra-lateral force-plate occurred a few percent and up to 37.7% of all strides (median: 12.9%). Our proposed method (HSest) robustly showed small errors for heel-strike detection and stride-interval calculation consistently among subjects, while HS50% and HS20N showed large errors depending on subjects., (Copyright © 2015 Elsevier B.V. All rights reserved.)

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., (Copyright © 2014 Elsevier Ireland Ltd. All rights reserved.)

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.

Background: Osteoarthritis patients may exhibit different kinematics according to the disease stage. However, changes in the frontal and horizontal planes in each stage remain unclear. The purpose of this study was to investigate changes in the knee kinematic gait variables of osteoarthritis patients, including the frontal and horizontal planes, with respect to the severity of the disease., Methods: Forty-five patients with knee osteoarthritis and 13 healthy young subjects were recruited for the experiment. All subjects were examined while walking on a 10-m walkway at a self-selected speed. In each trial, we calculated the angular displacements of flexion/extension, abduction/adduction, and external/internal tibial rotation. We also measured muscle strength, range of motion (ROM), and alignment. We compared the differences in osteoarthritis severity and knee kinematic variables between osteoarthritis patients and normal subjects., Results: The flexion angle at the time of foot contact was significantly less in patients with severe and moderate osteoarthritis than in normal subjects (both p<0.01). The abduction angle at the 50% stance phase was significantly less in patients with severe osteoarthritis than in normal subjects (p<0.05). The excursion of axial tibial rotation was significantly less in patients with early osteoarthritis than in normal subjects (p<0.05)., Conclusion: Osteoarthritis patients had different knee kinematics during gait, depending on the progress of osteoarthritis. Early-stage patients exhibit decreased axial tibial rotation excursion, while severe-stage patient exhibit increased knee adduction., (Copyright © 2011 Elsevier B.V. All rights reserved.)

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.

We investigated input (stimulus)-output (response) relations of the corticospinal pathway in the lower limb muscles during passive stepping using a robotic driven gait orthosis. Nine healthy adult subjects passively stepped with 40% body weight unloading (ground stepping) and 100% body weight unloading in the air (air stepping). During passive stepping, the motor evoked potentials (MEPs) of the lower limb muscles elicited by transcranial magnetic stimulation (TMS) were recorded at late-stance, early-, and late-swing phases of 2 stepping conditions. The input-output relation at each phase of the stepping conditions was obtained by increasing stimulus intensity in 5% increments from 40% to 70% of maximal stimulator output. The slopes of input-output relations were steeper at the early-swing phase in the rectus femoris muscle and at the late-stance and late-swing phases in the biceps femoris muscle in both stepping conditions. There were no significant differences in the MEP responses of the rectus femoris and biceps femoris muscles at each phase between the 2 conditions. Low muscle activity was seen at the late-stance phase of ground stepping in the soleus muscle and the MEP amplitude at this phase became larger. The slopes in the tibialis anterior muscle were steep at the early- and late-swing phases of ground stepping. There was a significant difference in the MEPs of the tibialis anterior muscle between the late-swing phases in ground and air stepping. The present study indicates that corticospinal excitability to the lower limb muscles is modulated by sensory inputs elicited by passive stepping.

Body fluctuations during both quiet standing and walking exhibit temporal correlations that reflect mechanisms of balance control. However, knowledge about the relationship between the temporal structures observed during standing and walking is limited. The goal of the present study was (1) to investigate temporal correlations in the fluctuations of the center of body mass acceleration (ACC) in standing and walking, and (2) to test the hypothesis that the degree of the temporal correlation for the two tasks is similar and correlated across participants. Seventeen young, healthy participants stood and walked for 10 min on a treadmill equipped with two force platforms. The temporal correlations of the ACC in the anteroposterior (ACC(AP)), mediolateral (ACC(ML)), and two-dimensional (ACC(2D)) directions were evaluated using the scaling index (alpha) as calculated with Detrended Fluctuation Analysis. The scaling indices of ACC fluctuations during standing and walking were categorized as stationary signals which are temporally correlated (0.5

We examined ventilatory response during passive walking-like exercise in the standing posture in complete spinal cord-injured humans and found that ventilatory equivalent for O(2) uptake, which would be related to the sensation of breathlessness, was lower during passive exercise than during quiet standing.

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.

Although phasic modulation of the corticospinal tract excitability to the lower limb muscles has been observed during normal walking, it is unclear to what extent afferent information induced by walking is related to the modulation. The purpose of this study was to test the corticospinal excitability to the lower limb muscles by using transcranial magnetic stimulation (TMS) and transcranial electrical stimulation of the motor cortex while 13 healthy subjects passively stepped in a robotic driven-gait orthosis. Specifically, to investigate the effect of load-related afferent inputs on the corticospinal excitability during passive stepping, motor evoked potentials (MEPs) in response to the stimulation were compared between two passive stepping conditions: 40% body weight unloading on a treadmill (ground stepping) and 100% body weight unloading in the air (air stepping). In the rectus femoris, biceps femoris and tibialis anterior (TA) muscles, electromyographic activity was not observed throughout the step cycle in either stepping condition. However, the TMS-evoked MEPs of the TA muscle at the early- and late-swing phases as well as at the early-stance phase during ground stepping were significantly larger than those observed during air stepping. The modulation pattern of the transcranial electrical stimulation-evoked MEPs was similar to that of the TMS-evoked MEPs. These results suggest that corticospinal excitability to the TA is facilitated by load-related afferent inputs. Thus, these results might be consistent with the notion that load-related afferent inputs play a significant role during locomotor training for gait disorders.

Objective: To examine blood pressure responses during passive walking-like exercise in the standing posture (PWE) in spinal cord-injured (SCI) humans., Methods: Twelve motor-complete SCI individuals (cervical level 6 to thoracic level 12, ASIA grade: A or B) and twelve able-bodied controls (CON) participated in this study. SCI individuals were divided into a group with injury level at or above thoracic (T) 6 (HSCI, n = 7) and a group with injury level at or below T10 (LSCI, n = 5). Subjects carried out 6-minute quiet standing and then 12-minute PWE at 1 Hz using a gait training apparatus that enables subjects to stand and move their legs passively., Results: Mean arterial blood pressures (MAPs) at standing in HSCI, LSCI and CON were 69 +/- 5, 83 +/- 4 and 93 +/- 2 mmHg, respectively. MAP changed significantly during PWE only in HSCI and CON, increasing to 88 +/- 4 (P < 0.001) and 98 +/- 1 mmHg (P < 0.01), respectively. The former group showed a larger increase in MAP (P < 0.001)., Interpretation: Spinal sympathetic reflexes can be induced in a region isolated from the brainstem in response to a stimulus originating below the level of the spinal cord injury, and the magnitude of increase in blood pressure is greater in SCI individuals with lesion level at or above T6 due to loss of supraspinal control of the major sympathetic outflow. This central mechanism may be one of the reasons why greater pressor response to PWE was observed in HSCI.

To address whether the passive observation of walking would induce an increase in motor cortical excitability, we examined the responses of motor-evoked potential elicited by transcranial magnetic stimulation in the tibialis anterior and soleus muscles as the participants observed naturally performed walking. Motor-evoked potentials in these muscles were significantly increased during the observation of walking throughout the entire step-cycle periods, but not during specific step periods. These findings indicate that cortical excitability can be increased not only during the observation of voluntary hand/arm movements, but also during the observation of automatic movements such as walking. It is also suggested that the present results may reflect the increased cortical excitability during the entire walking cycle.

Although cutaneous reflexes are known to be strongly modulated in a phase-dependent manner during walking in both human and cat, it is not clear whether the movement-related or the load-sensitive afferent feedback plays a more important role in regulating this modulation. To address this issue in humans, we investigated modulation of the cutaneous reflex in the tibialis anterior muscles (TA) of 17 subjects during passive walking with a load (0%, 33%, 66% unloading of body weight) and without a load (100% unloading). These walking tasks were performed passively with a robotic gait trainer system. Cutaneous reflexes in TA, elicited by electrical stimulation to the distal tibial (Tib) and superficial peroneal (SP) nerves, were recorded during 10 different phases of the walking cycle, and the middle latency responses (MLR, 70-120 ms) were analysed. During loaded walking, the magnitudes of the MLR induced by Tib nerve stimulation were strongly increased during the late stance-to-early swing phase irrespective of the amount of load (phase modulation), a phenomenon that also occurred without background electromyogram in the TA. Predominant suppression of the MLR following SP nerve stimulation at the early stance phase changed to facilitation at the late stance. By contrast, the MLR following either Tib or SP nerve stimulation was not at all modulated by the stepping phase during both unloaded walking (100% unloading) and standing. These results suggest that phasic changes in the load-related afferent information in concert with rhythmic lower limb movement play a key role in modulating cutaneous reflexes during walking.

We investigated relative activations in the synergistic ankle plantarflexor muscles, the soleus (SOL) and medial gastrocnemius (MG), while subjects were walking in water. Ten healthy volunteers walked in water under three different load- and four different speed-conditions. The depth of water was to the axilla of each subject, which reduced about 80% of his or her body weight. The EMG activities of the SOL and MG were recorded with surface electrodes, and ground reaction forces (GRF) were measured by using a water-resistant force platform. Relations of EMG activity levels between the two muscles during the stance phase were analysed with respect to the load and walking speed. The results revealed that the SOL EMG activity was more dependent on the load than the walking speed in water, while the MG EMG activity was more dependent on the speed. Such dissociated activation of the synergistic ankle plantarflexors might be relevant to the different roles required of these muscles in functional motor tasks.

To investigate the effects of gravity-related somatosensory information on spinal human reflexes, the soleus H-reflex was recorded in ten healthy subjects walking on a treadmill at 2.0 km/h on land and in water. The modulation pattern of the soleus H-reflex was determined in ten different phases of the step cycle. While the subjects were walking in water, the background electromyographic activity (BGA) of the soleus was lower than that on land; on the other hand, the soleus H-reflex amplitude while the subjects were walking in water showed no significant differences throughout the step cycle compared with that while the subjects were on land; the phase-dependent soleus H-reflex modulation pattern was well preserved while walking in water. There was a linear relationship between the BGA and the H-reflex amplitude in each condition; however, the soleus H-reflex gain while walking in water was significantly higher than that on land. These findings suggest that the somatosensory graviception can markedly reduce the spinal reflex excitability. Our findings are discussed in relation to human gait; therefore, further studies are needed to clarify the effect of somatosensory graviception on human neural mechanisms.

Objective: To clarify the functional roles of lower-limb joint moments and their contribution to support and propulsion tasks while walking in water compared with that on land., Design: Sixteen healthy, young subjects walked on land and in water at several different speeds with and without additional loads., Background: Walking in water is a major rehabilitation therapy for patients with orthopedic disorders. However, the functional role of lower-limb joint moments while walking in water is still unclear., Methods: Kinematics, electromyographic activities in biceps femoris and gluteus maximums, and ground reaction forces were measured under the following conditions: walking on land and in water at a self-determined pace, slow walking on land, and fast walking in water with or without additional loads (8 kg). The hip, knee, and ankle joint moments were calculated by inverse dynamics., Results: The contribution of the walking speed increased the hip extension moment, and the additional weight increased the ankle plantar flexion and knee extension moment., Conclusions: The major functional role was different in each lower-limb joint muscle. That of the muscle group in the ankle is to support the body against gravity, and that of the muscle group involved in hip extension is to contribute to propulsion. In addition, walking in water not only reduced the joint moments but also completely changed the inter-joint coordination., Relevance: It is of value for clinicians to be aware that the greater the viscosity of water produces a greater load on the hip joint when fast walking in water.

Purpose: The purpose of this study was to compare the changes in ground reaction forces (GRF), joint angular displacements (JAD), joint moments (JM) and electromyographic (EMG) activities that occur during walking at various speeds in water and on land., Method: Fifteen healthy adults participated in this study. In the water experiments, the water depth was adjusted so that body weight was reduced by 80%. A video-motion analysis system and waterproof force platform was used to obtain kinematics and kinetics data and to calculate the JMs., Results: Results revealed that (1) the anterior-posterior GRF patterns differed between walking in water and walking on land, whereas the medio-lateral GRF patterns were similar, (2) the JAD patterns of the hip and ankle were similar between water- and land-walking, whereas the range of motion at the knee joint was lower in water than on land, (3) the JMs in all three joints were lower in water than on land throughout the stance phase, and (4) the hip joint extension moment and hip extensor muscle EMG activity were increased as walking speed increase during walking in water., Conclusions: Rehabilitative water-walking exercise could be designed to incorporate large-muscle activities, especially of the lower-limb extensor muscles, through full joint range of motion and minimization of joint moments.

Purpose: Walking in water is a widely used rehabilitation method for patients with orthopedic disorders or arthritis, based on the belief that the reduction of weight in water makes it a safer medium and prevents secondary injuries of the lower-limb joints. To our knowledge, however, no experimental data on lower-limb joint moment during walking in water is available. The aim of this study was to quantify the joint moments of the ankle, knee, and hip during walking in water in comparison with those on land., Method: Eight healthy volunteers walked on land and in water at a speed comfortable for them. A video-motion analysis system and waterproof force platform were used to obtain kinematic data and to calculate the joint moments., Results: The hip joint moment was shown to be an extension moment almost throughout the stance phase during walking in water, while it changed from an extension- to flexion-direction during walking on land. The knee joint moment had two extension peaks during walking on land, whereas it had only one extension peak, a late one, during walking in water. The ankle joint moment during walking in water was considerably reduced but in the same direction, plantarflexion, as that during walking on land., Conclusions: The joint moments of the hip, knee, and ankle were not merely reduced during walking in water; rather, inter-joint coordination was totally changed.

Study Design: Comparative study of the effectiveness of walking exercise with a newly developed gait orthosis, the weight-bearing control (WBC) orthosis, for thoracic level of paraplegic patients., Objectives: To test its feasibility as a rehabilitation alternative for paraplegic patients, the energy consumption and cost during walking with WBC were calculated and compared with the values of conventional orthoses given in previous reports., Setting: National Rehabilitation Center for the Disabled, Japan., Methods: Four paraplegic patients with traumatic spinal cord injuries ranging from T8 to T12 participated. Experiments were conducted after 3 months of the orthotic gait training with WBC. The cardiorespiratory parameters were continuously measured at rest and during walking with a telemetric device. The steady-state value of the oxygen uptake (V(O2)), heart rate (HR), the energy consumption (J/kg/s) and energy cost (J/kg/m) were calculated., Results: The average walking speed was 19.0 +/- 2.58 m/min. The steady-state value of the V(O2) and HR were 16.08 +/- 1.93 ml/kg and 147.3 +/- 10.94 b/min, respectively. The energy cost during orthotic walking tended to be better than the values of conventional orthoses, whereas the energy consumption was almost similar., Conclusion: WBC enables thoracic level of paraplegic patients to walk at relatively higher speed than conventional orthoses under similar energy expenditure. The special devices equipped with WBC are therefore considered to lead to improvement of the energy cost of walking. The physical intensity presumed by cardiorespiratory responses during walking with WBC is suited to promote their aerobic capacity. Therefore, it is concluded that the WBC orthosis could be an effective alternative in rehabilitation for thoracic level of paraplegic patients.

During walking in water (WW) the vertical component of ground reaction forces decreases, while the greater propulsive force is required to move forward against the greater resistance of water. In such reduced gravity environment, Hutchison et al. (1989) have demonstrated that the relative activation of rat medial gastrocnemius (MGAS) increased compared to that of the soleus (SOL) during swimming, suggesting different effects of peripheral information on motoneuron excitability of these muscles. It is conceivable that both buoyancy and resistance of water have different effects on the activation patterns of triceps surae muscles during WW, since the reduced weight in water might decrease the peripheral inflow relating load information while greater volitional command might be needed to propel a body forward against the water resistance. The present study was designed to assess each peripheral inflow and efferent input by adjusting the load and walking speed voluntarily during WW. The aim of this study is to investigate the dissociative activation pattern between the SOL and the MGAS during WW.

Study design:Cross-sectional and comparative investigation using quadriplegics (QP) and nondisabled subjects (ND).Objective:To evaluate cardiorespiratory responses during passive walking-like exercise (PWE) in QP.Setting:National Rehabilitation Center for Persons with Disabilities in Japan.Method:The subjects were seven male QP with complete lesion (age: 27.0±5.4, injured level: C6–C7) and six male ND (age: 26.3±4.5). Cardiorespiratory responses were measured until voluntary fatigue during PWE, the rhythmical activity of paralyzed lower limbs synchronized with arm movements.Results:There were no significant differences in oxygen consumption (\[Vdot]O2), pulmonary ventilation (\[Vdot]E), heart rate (HR) and oxygen pulse (O2 pulse) between QP and ND during PWE. ND showed increased ventilatory equivalent for oxygen (\[Vdot]E/\[Vdot]O2 ratio) during exercise, while QP showed a significantly greater respiratory rate (RR) during exercise than ND (P<0.05).Conclusion:PWE elicited an increase in \[Vdot]O2 with workload increment in QP similar to ND. However, higher RR suggested the intrinsic dysfunction of RR control during submaximal exercise in QP. From these results, it was thought that respiratory response would be the restriction factor of efficient oxygen transportation during PWE in QP.Spinal Cord (2006) 44, 480–486. doi:10.1038/sj.sc.3101875; published online 29 November 2005 [ABSTRACT FROM AUTHOR]

Study design:Cross-sectional, experimental research.Objectives:To clarify the effect of lesion level on cardio-respiratory responses and biomechanical characteristics of walking with a reciprocating gait orthosis in complete paraplegia with spinal cord injury (SCI).Setting:National Rehabilitation Center for Persons with Disabilities, Japan.Methods:Ten SCI individuals (age: 20–34 years, injured level: Th5–12) who experienced orthotic gait training at least for 10 weeks participated in two experiments: (1) measurement of the cardiorespiratory responses during 20 min of orthotic gait exercise; and (2) three-dimensional motion analysis and ground reaction force measurement using the VICON system. We calculated the following parameters: pulmonary ventilation, oxygen consumption (VO2), heart rate (HR), gait speed, cadence, stride length, crutch force (CF), hip range of motion (ROM), and hip angular velocity (VEL). Further, energy consumption and energy cost were calculated using the steady-state value of VO2 and gait speed.Results:The steady-state value of the VO2 (18.2±3.80 ml/kg) and HR (133.0±21.63 b/min) tended to be larger in higher thoracic SCI subjects. There were strong positive correlations between the lesion level and walking speed (r=0.74), energy cost (r=0.85), and hip ROM (r=0.78). On the other hand, negative correlation between the lesion level and peak CF (r=−0.78) was clarified.Conclusions:The physiological intensity of the orthotic gait strongly depended on the level of lesion. It seems likely that a limited hip range of motion and excess upper limb load result in the low energy cost of orthotic gait for the higher thoracic level of paraplegic patients.Spinal Cord (2006) 44, 487–494. doi:10.1038/sj.sc.3101916; published online 21 March 2006 [ABSTRACT FROM AUTHOR]