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Summary Muscular control during walking is believed to be simplified by the coactivation of muscles called muscle synergies. Although significant corticomuscular connectivity during walking has been reported, the level at which the cortical activity is involved in muscle activity (muscle synergy or individual muscle level) remains unclear. Here we examined cortical correlates of muscle activation during walking by brain decoding of activation of muscle synergies and individual muscles from electroencephalographic signals. We demonstrated that the activation of locomotor muscle synergies was decoded from slow cortical waves. In addition, the decoding accuracy for muscle synergies was greater than that for individual muscles and the decoding of individual muscle activation was based on muscle-synergy-related cortical information. These results indicate the cortical correlates of locomotor muscle synergy activation. These findings expand our understanding of the relationships between brain and locomotor muscle synergies and could accelerate the development of effective brain-machine interfaces for walking rehabilitation., Graphical Abstract, Highlights • We examined relationships of brain and locomotor muscle synergies by brain decoding • Locomotor muscle synergy activation was successfully decoded from EEG signals • Single muscle activation was decoded based on muscle-synergy-related EEG signals • The cortical correlates of locomotor muscle synergy may contribute to BMI for gait, Human Physiology; Neuroscience; Biomechanics

Transcutaneous and epidural electrical spinal cord stimulation techniques are becoming more valuable as electrophysiological and clinical tools. Recently, remarkable recovery of the upper limb sensorimotor function during cervical spinal stimulation was demonstrated. In the present study, we sought to elucidate the neural mechanisms underlying the effects of transcutaneous spinal cord stimulation (tSCS) of the cervical spine. We hypothesized that cervical tSCS can be used to selectively activate the sensory route entering the spinal cord and transsynaptically converge on upper limb motor pools. To test this hypothesis, we applied cervical tSCS using paired stimuli (homosynaptic depression) and during passive muscle stretching of the wrist flexor (presynaptic inhibition via Ia afferents), voluntary hand muscle contraction (descending facilitation of motoneuron pool), and muscle-tendon vibration of the wrist (presynaptic inhibition via afferent occlusion). Our results demonstrate significant inhibition of the second evoked response during paired stimulus delivery, inhibition of responses during passive muscle stretching and muscle-tendon vibration, and facilitation during voluntary muscle contraction, which share similarities with responses evoked during lumbosacral tSCS. These results indicate that the route of the stimulation current transmission passes via afferents in the dorsal roots through the spinal cord to activate the motor pools and potentially interneuronal networks projecting to upper limb muscles. Using a novel stimulation paradigm, our study is the first to present evidence of the sensory neuronal pathway of the cervical tSCS propagation. Overall, our work demonstrates the utility and sensitivity of cervical tSCS to engage the sensory pathway projecting to the upper limbs. NEW & NOTEWORTHY Despite therapeutic effects that have been demonstrated previously using noninvasive cervical spinal stimulation, it has been unclear whether, and to what degree, the stimulation can activate the sensory afferent system. Our study presents evidence that cervical transcutaneous spinal cord stimulation can engage the sensory pathways and transsynaptically converge on motor pools projecting to upper limb muscles, demonstrating the utility and sensitivity of cervical spinal stimulation for electrophysiological assessments and neurorehabilitation.

Cervical transcutaneous spinal cord stimulation (tSCS) efficacy for rehabilitation of upper-limb motor function was suggested to depend on recruitment of Ia afferents. However, selectivity and excitability of motor activation with different electrode configurations remain unclear. In this study, activation of upper-limb motor pools was examined with different cathode and anode configurations during cervical tSCS in 10 able-bodied individuals. Muscle responses were measured from six upper-limb muscles simultaneously. First, postactivation depression was confirmed with tSCS paired pulses (50-ms interval) for each cathode configuration (C6, C7, and T1 vertebral levels), with anode on the anterior neck. Selectivity and excitability of activation of the upper-limb motor pools were examined by comparing the recruitment curves (10-100 mA) of first evoked responses across muscles and cathode configurations. Our results showed that hand muscles were preferentially activated when the cathode was placed over T1 compared with the other vertebral levels, whereas there was no selectivity for proximal arm muscles. Furthermore, higher stimulation intensities were required to activate distal hand muscles than proximal arm muscles, suggesting different excitability thresholds between muscles. In a separate protocol, responses were compared between anode configurations (anterior neck, shoulders, iliac crests, and back), with one selected cathode configuration. The level of discomfort was also assessed. Largest muscle responses were elicited with the anode configuration over the anterior neck, whereas there were no differences in the discomfort. Our results therefore inform methodological considerations for electrode configuration to help optimize recruitment of Ia afferents during cervical tSCS.

Paired associative stimulation (PAS) is a non-invasive method to modulate the excitability of the primary motor cortex (M1). PAS involves the combination of peripheral nerve stimulation and transcranial magnetic stimulation (TMS) over the primary motor cortex. However, for lower-limb muscles, PAS has only been applied to the few muscles innervated by peripheral nerves that can easily be stimulated. This study used transcutaneous spinal cord stimulation (tSCS) to the posterior root, stimulating the sensory nerves of multiple lower-limb muscles, and aimed to investigate the effect of PAS consisting of tSCS and TMS on corticospinal excitability. Twelve non-disabled men received 120 paired stimuli on two separate days in (1) an individual-ISI condition, using inter-stimulus intervals (ISIs) of paired stimuli individually calculated to send two signals to M1 with individually-adjusted ISI, and (2) a constant-ISI condition, using a constant ISI of 100 ms. Before and after PAS, corticospinal excitability was assessed in the lower-limb muscles. Facilitation of corticospinal excitability in the lower-leg and hamstring muscles was observed up to 30 min after PAS only in the individual-ISI condition (p 0.05), although there was no significant difference between the individual-ISI and constant-ISI conditions. Additionally, our results revealed a difference in PAS-induced facilitation among lower-limb muscles, suggesting a spatial gradient of PAS-induced facilitation of corticospinal excitability, such that knee flexor muscles have a higher potential for plastic change than knee extensor muscles. These findings will foster a better understanding of the neural mechanisms underlying PAS-induced neuroplasticity, leading to better neurorehabilitation and motor learning strategies.

Objective. Brain decoding of motor imagery (MI) not only is crucial for the control of neuroprosthesis but also provides insights into the underlying neural mechanisms. Walking consists of stance and swing phases, which are associated with different biomechanical and neural control features. However, previous knowledge on decoding the MI of gait is limited to simple information (e.g. the classification of 'walking' and 'rest').Approach. Here, we investigated the feasibility of electroencephalogram (EEG) decoding of the two gait phases during the MI of walking and whether the combined use of MI and action observation (AO) would improve decoding accuracy.Main results. We demonstrated that the stance and swing phases could be decoded from EEGs during MI or AO alone. We also demonstrated the decoding accuracy during MI was improved by concurrent AO. The decoding models indicated that the improved decoding accuracy following the combined use of MI and AO was facilitated by the additional information resulting from the concurrent cortical activations related to sensorimotor, visual, and action understanding systems associated with MI and AO.Significance. This study is the first to show that decoding the stance versus swing phases during MI is feasible. The current findings provide fundamental knowledge for neuroprosthetic design and gait rehabilitation, and they expand our understanding of the neural activity underlying AO, MI, and AO + MI of walking.Novelty and significanceBrain decoding of detailed gait-related information during motor imagery (MI) is important for brain-computer interfaces (BCIs) for gait rehabilitation. This study is the first to show the feasibility of EEG decoding of the stance versus swing phases during MI. We also demonstrated that the combined use of MI and action observation (AO) improves decoding accuracy, which is facilitated by the concurrent and synergistic involvement of the cortical activations for MI and AO. These findings extend the current understanding of neural activity and the combined effects of AO and MI and provide a basis for effective techniques for walking rehabilitation.

Corticospinal excitability in humans can be facilitated during imagination and/or observation of upper-limb motor tasks. However, it remains unclear to what extent facilitation levels may differ from those elicited during execution of the same tasks. Twelve able-bodied individuals were recruited in this study. Motor evoked potentials (MEPs) in extensor carpi radialis (ECR) and flexor carpi radialis (FCR) muscles were elicited through transcranial magnetic stimulation of the primary motor cortex during: (i) rest; (ii) wrist extension; and (iii) wrist flexion. Responses were compared between: (1) motor imagery combined with virtual action observation (MI + AO; first-person virtual wrist movements shown on a computer display, while participants remained at rest and imagined these movements); and (2) motor execution (ME; participants extended or flexed their wrist). During MI + AO, ECR MEPs were facilitated during the extension phase but not the flexion phase, while FCR MEPs were facilitated during the flexion phase but not extension phase, compared to rest. During the ME condition, same, but greater, modulations were shown as those during MI + AO, while background muscle activities were similar in the rest phase as during extension and flexion phase in the MI + AO condition. Our results demonstrated that kinesthetic MI that included imagination and observation of virtual hands can elicit phase-dependent muscles-specific corticospinal facilitation of wrist muscles, consistent to those during actual hand extension and flexion. Moreover, we showed that MI + AO can contribute considerably to the overall corticospinal facilitation (∼20 % of ME) even without muscle contractions. These findings support utility of computer graphics-based motor imagery, which may have implications for rehabilitation and development of brain-computer interfaces.

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.

Electrical stimulation involving visual areas of the brain produces artificial light percepts called phosphenes. These visual percepts have been extensively investigated in previous studies involving intracortical microsimulation (ICMS) and serve as the basis for developing a visual prosthesis for the blind. Although advances have been achieved, many challenges still remain with implementing a functional ICMS for visual rehabilitation purposes. Transcranial magnetic stimulation (TMS) over the primary occipital lobe offers an alternative method to produce phosphenes non-invasively. A main challenge facing blind individuals involves navigation. Within the scientific community, methods to evaluate the ability of a visual prosthesis to facilitate in navigation has been neglected. In this study, we investigate the effectiveness of evoking lateralized phosphenes to navigate a computer simulated virtual environment. More importantly, we demonstrate how virtual environments along with the development of a visual prosthesis share a mutual relationship benefiting both patients and researchers. Using two TMS devices, a pair of 40mm figure-of-eight coils were placed over each occipital hemisphere resulting in lateralized phosphene perception. Participants were tasked with making a series of left and right turns using peripheral devices depending on the visual hemifield in which a phosphene is present. If a participant was able to accurately perceive all ten phosphenes, the simulated target is able to advance and fully exit the virtual environment. Our findings demonstrate that participants can interpret lateralized phosphenes while highlighting the integration of computer based virtual environments to evaluate the capability of a visual prosthesis during navigation.

Despite the well-known tradeoff between speed and accuracy, skilled people often demonstrate the ability to maintain high accuracy during fast movements. We focused on two strategies to improve accuracy, thereby increasing the reproducibility of individual parameters (certain parameters are maintained in low variability) and coordinating covariation among parameters (different parameters compensate each other's variability). The objective of this study was to determine whether coordinated covariation among release parameters is used for high accuracy by skilled baseball pitchers. A model was employed to simulate pitch location after eliminating the coordinated covariation by randomly reshuffling the release parameters, and the variability of simulated and measured pitch locations were compared. The results showed that there was no significant coordinated covariation for any of the release parameters for either the vertical or horizontal pitch location supports strategy of increasing the reproducibility of individual parameter. In addition, for the vertical pitch location, because there was coordinated covariation between the release angle and speed in slow pitching, it was suggested that, the higher speed the task requires, the more important the reproducibility of individual parameter becomes.

Delivering short trains of electric pulses to the muscles and nerves can elicit action potentials resulting in muscle contractions. When the stimulations are sequenced to generate functional movements, such as grasping or walking, the application is referred to as functional electrical stimulation (FES). Implications of the motor and sensory recruitment of muscles using FES go beyond simple contraction of muscles. Evidence suggests that FES can induce short- and long-term neurophysiological changes in the central nervous system by varying the stimulation parameters and delivery methods. By taking advantage of this, FES has been used to restore voluntary movement in individuals with neurological injuries with a technique called FES therapy (FEST). However, long-lasting cortical re-organization (neuroplasticity) depends on the ability to synchronize the descending (voluntary) commands and the successful execution of the intended task using a FES. Brain-computer interface (BCI) technologies offer a way to synchronize cortical commands and movements generated by FES, which can be advantageous for inducing neuroplasticity. Therefore, the aim of this review paper is to discuss the neurophysiological mechanisms of electrical stimulation of muscles and nerves and how BCI-controlled FES can be used in rehabilitation to improve motor function.

The present study was a cross-sectional comparison of probabilistic structure in the distribution of pitching location among baseball pitchers of various age groups (25 elementary school (ES), 20 junior high school (JH), 15 high school (HS), and 18 college students (CL)). In the results, despite the general age-dependent variations in pitching precision, the difference was reflected not only in error 'size' but also in the 'shape' of error as it was shown by fitting 95% confidence ellipse to the two dimensional distribution of pitch location. While the precision measure as a reflection of trial-by-trial variability of release timing (major axis length of the ellipse) was constant, minor axis length of the ellipse as a reflection of variability in the pitching form of each participant demonstrated significant differences among the groups. In the ES group particularly, the trial-by-trial variability in the trajectory angle of the throwing arm was significantly correlated with the minor axis length; this correlation was far greater than those in older groups. The present study is the first to demonstrate the detailed structure of the variability of pitching location of baseball dependent on age.

How do skilled players change their motion patterns depending on motion effort? Pitchers commonly accelerate wrist and elbow joint rotations via proximal joint motions. Contrastingly, they show individually different pitching motions, such as in wind-up or follow-through. Despite the generality of the uniform and diverse features, effort-dependent effects on these features are unclear. Here, we reveal the effort dependence based on muscle activity data in natural three-dimensional pitching performed by skilled players. We extract motor modules and their effort dependence from the muscle activity data via tensor decomposition. Then, we reveal the unknown relations among motor modules, common features, unique features, and effort dependence. The current study clarifies that common features are obvious in distinguishing between low and high effort and that unique features are evident in differentiating high and highest efforts.

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.

Functional electrical stimulation therapy (FEST) can improve motor function after neurological injuries. However, little is known about cortical re-organization after FEST and weather it can improve upper-limb motor function after traumatic brain injury (TBI). Therefore, our study examined cortical and motor changes in a single male participant with chronic TBI suffering from mild motor impairment during 3-months of FEST and at 3-months follow-up. FEST was applied to enable upper-limb grasping and reaching movements during each session, which was performed for 45-60 min, 3 days per week, over 12-weeks. Short-term assessments were examined before and after each session, while long-term assessments were performed at baseline, after 6- and 12-weeks of FEST, and during follow-up 6- and 12-weeks after completing FEST. Short-term assessments carried out using transcranial magnetic stimulation (TMS) showed reduced cortical silent period (CSP), which is related to cortical and/or subcortical inhibition. At the same time, no changes in motor evoked potentials (MEP) were observed, suggesting corticospinal excitability was unaffected. Long-term assessments indicate increased MEP corticospinal excitability after 12-weeks of FEST, which remained during both follow-ups, while no changes in CSP were observed. Similarly, long-term assessments using TMS mapping showed larger hand MEP area in the primary motor cortex (M1) after 12-weeks of FEST as well as during both follow-ups. Corroborating TMS results, fMRI imaging data showed M1, as well as sensory, premotor, parietal area, and supplementary motor area activations increased after 12-weeks of FEST and during both follow-ups. While clinical scores did not change considerably, writing test performance indicates mild improvements after FEST. Our results suggest that FEST can effectively increase cortical activations, while writing tests confirmed functional improvements in fine motor function even after chronic TBI. These results demonstrated long-term recovery mechanisms of FEST, which include cortical re-organization or neuroplasticity to improve motors function after neurological injury.

The execution of cognitive tasks is known to alter postural sway during standing, but the underlying mechanisms are still debated. This study investigated how performing a mental task modified balance control during standing. We required 15 healthy adult males to maintain an upright stance under conditions of simply relaxing and maintaining normal quiet standing (control condition) or while performing a secondary cognitive task (mental arithmetic). Under each condition, we measured the participants’ center of pressure and used kinematic measurements for a quantitative evaluation of postural control modulation. We calculated the standard deviation of the joint angles (ankle, knee, and hip) and the estimated joint stiffness to measure joint mobility changes in postural control. To estimate the kinematic pattern of covariation among these joints, we used uncontrolled manifold analysis, an assessment of the strength of multijoint coordination. Compared to normal standing, executing the cognitive task while standing led to reduced movements of the ankle and hip joints. There were no significant differences in ankle stiffness or uncontrolled manifold ratios between the conditions. Our results suggest that when performing a secondary cognitive task during standing, neither changes in the modification of stiffness nor the strength of multijoint coordination (both of which preserve the center of mass position) explains changes in postural sway.

Motor executions alter sensory processes. Studies have shown that loudness perception changes when a sound is generated by active movement. However, it is still unknown where and how the motor-related changes in loudness perception depend on the task demand of motor execution. We examined whether different levels of precision demands in motor control affects loudness perception. We carried out a loudness discrimination test, in which the sound stimulus was produced in conjunction with the force generation task. We tested three target force amplitude levels. The force target was presented on a monitor as a fixed visual target. The generated force was also presented on the same monitor as a movement of the visual cursor. Participants adjusted their force amplitude in a predetermined range without overshooting using these visual targets and moving cursor. In the control condition, the sound and visual stimuli were generated externally (without a force generation task). We found that the discrimination performance was significantly improved when the sound was produced by the force generation task compared to the control condition, in which the sound was produced externally, although we did not find that this improvement in discrimination performance changed depending on the different target force amplitude levels. The results suggest that the demand for precise control to produce a fixed amount of force may be key to obtaining the facilitatory effect of motor execution in auditory processes.

This study explored the mechanical factors that determine accuracy of a baseball pitching. In particular, we focused on the mechanical parameters at ball release, referred to as release parameters. The aim was to understand which parameter has the most deterministic influence on pitch location by measuring the release parameters during actual pitching and developing a simulation that predicts the pitch location from given release parameters. By comparing the fluctuation of the simulated pitch location when varying each release parameter, it was found that the elevation pitching angle and speed significantly influenced the vertical pitch location, and the azimuth pitching angle significantly influenced the horizontal pitch location. Moreover, a regression model was obtained to predict the pitch location, and it became clear that the significant predictors for the vertical pitch location were the elevation pitching angle, the speed, and spin axis, and those for the horizontal pitch location were the azimuth pitching angle, the spin axis, and horizontal release point. Therefore, it was suggested that the parameter most affecting pitch location weas pitching angle. On the other hand, multiple regression analyses revealed that the relation between release parameters varied between pitchers. The result is expected to contribute to an understanding of the mechanisms underlying accurate ball control skill in baseball pitching.

The main aim of the study was to evaluate how the brain of a Paralympic athlete with severe disability due to cerebral palsy has reorganized after continuous training geared to enhance performance. Both corticospinal excitability of upper-limb muscles and electromyographic activity during swimming were investigated for a Paralympic gold medalist in swimming competitions. Transcranial magnetic stimulation (TMS) to the affected and intact hand motor cortical area revealed that the affected side finger muscle cortical representation area shifted towards the temporal side, and cortico-spinal excitability of the target muscle was prominently facilitated, i.e., the maximum motor evoked potential in the affected side, 6.11 ± 0.19 mV was greater than that in the intact side, 4.52 ± 0.39 mV (mean ± standard error). Electromyographic activities during swimming demonstrated well-coordinated patterns as compared with rather spastic activities observed in the affected side during walking on land. These results suggest that the ability of the brain to reorganize through intensive training in Paralympic athletes can teach interesting lessons to the field neurorehabilitation.

The central nervous system (CNS) achieves a stable gait at several speeds and modes while controlling diverse instability. An essential feature of a gait is the motion of the center of body mass (CoM). CoM motion is at larger risk for trespassing the base of support in the mediolateral direction than in the anteroposterior direction. How the CoM trajectory in the frontal plane changes depending on the speed or mode can thus provide insights about the neural control of stable gaits. Here, we reveal the speed- and mode-dependent modulations of the trajectory by utilizing a Lissajous curve. The current study clarifies that speed-dependent modulations are evident in walking. Between walking and running, there were significant mode-dependent modulations. In contrast, there were no significant speed-dependent modulations during running. Deviations from standard tendencies quantified via Lissajous curve fitting could be a sign of gait impairments and recovery after treatments.

The present study addressed whether motor memory acquired earlier in life through specific training can be maintained through later life with further training. To this end, the present study focused on the training effect of a specific ballet practice and investigated the spinally mediated stretch reflex responses of the soleus muscle in ballet dancers of upper‐middle to old age (60.6 ± 5.4 years old) with experience levels of 28.4 ± 7.4 years (“older ballet” group). Comparisons were conducted with a group of young ballet dancers (“young ballet” group) and groups of both young and older individuals without weekly participation in physical activities (“young sedentary” and “older sedentary” groups). The results revealed natural age‐dependent changes, with reflex responses being larger in older sedentary than in young sedentary individuals. A training‐induced effect was also observed, with responses being smaller in ballet dancers than in sedentary groups of the same age. Furthermore, the responses were surprisingly smaller in the older ballet dancers than in the young sedentary group, at an equivalent level to that of the young ballet dancers. The influence of training, therefore, overcame the natural age‐dependent changes. On the other hand, the onset latencies of the responses showed a solely age‐dependent trend. Taken together, the present is the first to demonstrate that the motor memories in the spinal cord acquired through specific ballet training earlier in life can be maintained and carried forward in later life through further weekly participation in the same training., With the knowledge that human motor system undergoes functional changes after participation in specific motor training, the present study first demonstrated that the motor memories in the spinal cord acquired through participation in specific ballet training earlier in life can be maintained and carried forward in later life through further weekly participation in the training.

Neuromuscular electrical stimulation (NMES) of lower limbs elicits muscle contractions through the activation of efferent fibers and concomitant recruitment of afferent fibers, which can modulate excitability of the central nervous system. However, neural mechanisms of NMES and how unilateral stimulation of the soleus affects spinal reflexes in multiple lower limb muscles bilaterally remains unknown. Twelve able-bodied participants were recruited, and spinal reflex excitability changes were tested after four interventions, each applied for 60 s, on the right plantar flexors: (1) motor-level NMES; (2) sensory-level NMES; (3) voluntary contraction; (4) rest. Spinal reflexes were elicited using single-pulse transcutaneous spinal cord stimulation applied on the lumbar level of the spinal cord to evoke bilateral responses in multiple lower limb muscles, while maximum motor response (Mmax) was tested in the soleus by stimulating the posterior tibial nerve. Spinal reflexes and Mmax before each intervention were compared to immediately after and every 5 min subsequently, for 15 min. Results showed that motor-level NMES inhibited spinal reflexes of the soleus and other studied muscles of the ipsilateral leg, but not the contralateral leg (except vastus medialis) for 15 min, while not affecting soleus muscle properties (Mmax). Voluntary contraction effect lasted less than 5 min, while sensory-level NMES and rest did not produce an effect. Short-term spinal reflex excitability was likely affected because antidromic impulses during motor-level NMES coincided in the spinal cord with afferent inputs to induce spinal neuroplasticity, whereas afferent input alone did not produce short-term effects. Such activation of muscles with NMES could reduce spasticity in individuals with neurological impairments.

The effects of motor imagery on spinal reflexes such as the H-reflex are unclear. One reason for this is that the muscles that can be used to record spinal reflexes are limited to traditional evoking methods Recently, transcutaneous spinal cord stimulation has been used for inducing spinal reflexes from multiple muscles and we aimed to examine the effect of motor imagery on spinal reflexes from multiple muscles. Spinal reflexes evoked by transcutaneous spinal cord stimulation were recorded from six muscles from lower limbs during motor imagery of right wrist extension and ankle plantarflexion with maximum isometric contraction. During both imaginary tasks, facilitation of spinal reflexes was detected in the ankle ipsilateral plantarflexor and dorsiflexor muscles, but not in thigh, toe or contralateral lower limb muscles. These results suggest that motor imagery of isometric contraction facilitates spinal reflex excitability in muscles of the ipsilateral lower leg and the facilitation does not correspond to the imaginary involved muscles.

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.

Animal studies demonstrate that the neural mechanisms underlying locomotion are specific to the modes and/or speeds of locomotion. In line with animal results, human locomotor adaptation studies, particularly those focusing on walking, have revealed limited transfers of adaptation among movement contexts including different locomotion speeds. Running is another common gait that humans utilize in their daily lives and is distinct from walking in terms of the underlying neural mechanisms. The present study employed an adaptation paradigm on a split-belt treadmill to examine the possible independence of neural mechanisms mediating different running speeds. The adaptations learned with split-belt running resulted in aftereffects with magnitudes that varied in a speed-dependent matter. In the two components of the ground reaction force investigated, the anterior braking and posterior propulsive components exhibited different trends. The anterior braking component tended to show larger aftereffect under speeds near the slower side speed of the previously experienced split-belt in contrast to the posterior propulsive component in which the aftereffect size tended to be the largest at a speed that corresponded to the faster side speed of the split-belt. These results show that the neural mechanisms underlying different running speeds in humans may be independent, just as in human walking and animal studies.

Highlights • Static magnetic field stimulation was delivered on cervical spinal cord. • Trans-spinal static magnetic field stimulation (tsSMS) decreased MEP amplitudes. • The suppressive effect of tsSMS was not maintained after the intervention ceased., Objective Transcranial static magnetic field stimulation has recently been demonstrated to modulate cortical excitability. In the present study, we investigated the effect of transspinal static magnetic field stimulation (tsSMS) on excitability of the corticospinal tract. Methods A compact magnet for tsSMS (0.45 Tesla) or a stainless steel cylinder for sham stimulation was positioned over the neck (C8 level) of 24 able-bodied subjects for 15 min. Using 120% of the resting motor threshold transcranial magnetic stimulation intensity, motor evoked potentials (MEPs) were measured from the first digital interosseous muscle before, during, and after the tsSMS or sham intervention. Results Compared with baseline MEP amplitudes were decreased during tsSMS, but not during sham stimulation. Additionally, during the intervention, MEP amplitudes were lower with tsSMS than sham stimulation, although these effects did not last after the intervention ceased. Conclusions The results suggest that static magnetic field stimulation of the spinal cord by a compact magnet can reduce the excitability of the corticospinal tract. Significance Transspinal static magnetic field stimulation may be a new non-invasive neuromodulatory tool for spinal cord stimulation. Its suppressive effect may be applied to patients who have pathological hyperexcitability of the spinal neural network.

Thoracic spinal cord injury (SCI) can have significant negative consequences, which can affect the ability to maintain unsupported sitting. The objectives of this study were to compare postural control of individuals with high- and low-thoracic SCI to able-bodied people and evaluate the effects of upper-limb support on postural control during quiet sitting. Twenty-five individuals were recruited into: (a) high-thoracic SCI; (b) low-thoracic SCI; and (c) able-body subgroups. Participants were seated and asked to maintain a steady balance in the following postures: (1) both hands resting on thighs; (2) both arms crossed over the chest; and (3) both arms extended. Center of pressure (COP) fluctuations were evaluated to compare postural performance between groups and different postures. Results showed that individuals with high- and low-thoracic SCI swayed more compared to the able-bodied group regardless of upper-limb support. No differences between the two SCI groups were observed, but the neurological level of injury was correlated to postural performance implying that those with higher injuries swayed more and faster. Unsupported sitting was more unstable in comparison to supported sitting posture, especially in the anterior-posterior direction. The velocity of postural sway was not different between groups, but the results suggest that postural regulation had unique effect during different postures in different groups. These results imply reduced postural stability after thoracic SCI. Overall, the way individuals with high-thoracic SCI achieved stability was different from that of individuals with low-thoracic SCI, suggesting different postural regulation strategies.

Spinal cord injury (SCI) can result in paralysis of trunk muscles, which can affect sitting balance. The objective of this study was to analyze trunk muscle coordination of individuals with thoracic SCI and compare it to able-body individuals. A total of 27 individuals were recruited and subdivided into: (a) high thoracic SCI; (b) low thoracic SCI; and (c) able-body groups. Participants were seated and asked to lean their trunk in eight directions while trunk muscle activity was recorded. Muscle coordination was assessed using the non-negative matrix factorization (NMF) method to extract muscle modules, which are the synergistic trunk muscle activations, and their directional activation patterns. Our results showed that individuals with SCI used less muscle modules, more co-contractions, and less directional tuning, compared to able-bodied people. These results suggest impaired and simplified muscle coordination due to the loss of supraspinal input after SCI. Observed variability in muscle coordination within SCI groups also suggests that other mechanisms such as spasticity and muscle stretch reflexes or individual factors such as experience and training contributed to the postural muscle synergies. Overall, muscle coordination deficits revealed impaired neuromuscular strategies which provide implications for rehabilitation of trunk muscles during sitting balance after SCI.

Ankle dorsiflexion force control is essential for performing daily living activities. However, the involvement of the corticospinal pathway during different ankle dorsiflexion tasks is not well understood. The objective of this study was to compare the corticospinal excitability during: (1) unilateral and bilateral; and (2) ballistic and tonic ankle dorsiflexion force control. Fifteen healthy young adults (age: 25.2 ± 2.8 years) participated in this study. Participants performed unilateral and bilateral isometric ankle dorsiflexion force-control tasks, which required matching a visual target (10% of maximal effort) as quickly and precisely as possible during ballistic and tonic contractions. Transcranial magnetic stimulation (TMS) was applied over the primary motor cortex to elicit motor-evoked potentials (MEPs) from the right tibialis anterior during: (i) pre-contraction phase; (ii) ascending contraction phase; (iii) plateau phase (tonic tasks only); and (iv) resting phase (control). Peak-to-peak MEP amplitude was computed to compare the corticospinal excitability during each experimental condition. MEP amplitudes significantly increased during unilateral contraction compared to bilateral contraction in the pre-contraction phase. There were no significant differences in the MEP amplitudes between the ballistic tasks and tonic tasks in any parts of the contraction phase. Although different strategies are required during ballistic and tonic contractions, the extent of corticospinal involvement appears to be similar. This could be because both tasks enhance the preparation for precise force control. Furthermore, our results suggest that unilateral muscle contractions may largely facilitate the central nervous system during movement preparation for unilateral force control compared to bilateral muscle contractions.

How the central nervous system (CNS) controls many joints and muscles is a fundamental question in motor neuroscience and related research areas. An attractive hypothesis is the module hypothesis: the CNS controls groups of joints or muscles (i.e., spatial modules) by providing time-varying motor commands (i.e., temporal modules) to the spatial modules rather than controlling each joint or muscle separately. Another fundamental question is how the CNS generates numerous repertoires of movement patterns. One hypothesis is that the CNS modulates the spatial and/or temporal modules depending on the required tasks. It is thus essential to quantify the spatial modules, the temporal modules, and the task-dependent modulation of these modules. Although previous attempts at such quantification have been made, they considered modulation either only in spatial modules or only in temporal modules. These limitations may be attributable to the constraints inherent to conventional methods for quantifying the spatial and temporal modules. Here, we demonstrate the effectiveness of tensor decomposition in quantifying the spatial modules, the temporal modules, and the task-dependent modulation of these modules without such limitations. We further demonstrate that tensor decomposition offers a new perspective on the task-dependent modulation of spatiotemporal modules: in switching from walking to running, the CNS modulates the peak timing in the temporal modules while recruiting more proximal muscles in the corresponding spatial modules.

How the central nervous system (CNS) controls many joints and muscles is a fundamental question in motor neuroscience and related research areas. An attractive hypothesis is the module hypothesis: the CNS controls groups of joints or muscles (i.e., spatial modules) while providing time-varying motor commands (i.e., temporal modules) to the spatial modules rather than controlling each joint or muscle separately. Another fundamental question is how the CNS generates numerous repertories of movement patterns. One hypothesis is that the CNS modulates the spatial and/or temporal modules depending on the required tasks. It is thus essential to quantify the spatial module, the temporal module, and the task-dependent modulation of those modules. Although previous methods attempted to quantify these aspects, they considered the modulation in only the spatial or temporal module. These limitations were possibly due to the constraints inherent to conventional methods for quantifying the spatial and temporal modules. Here, we demonstrate the effectiveness of tensor decomposition in quantifying the spatial module, the temporal module, and the task-dependent modulation of these modules without such limitations. We further demonstrate that the tensor decomposition provides a new perspective on the task-dependent modulation of spatiotemporal modules: in switching from walking to running, the CNS modulates the peak timing in the temporal module while recruiting proximal muscles in the corresponding spatial module.Author summaryThere are at least two fundamental questions in motor neuroscience and related research areas: 1) how does the central nervous system (CNS) control many joints and muscles and 2) how does the CNS generate numerous repertories of movement patterns. One possible answer to question 1) is that the CNS controls groups of joints or muscles (i.e., spatial modules) while providing time-varying motor commands (i.e., temporal modules) to the spatial modules rather than controlling each joint or muscle separately. One possible answer to question 2) is that the CNS modulates the spatial and/or temporal module depending on the required tasks. It is thus essential to quantify the spatial module, the temporal module, and the task-dependent modulation of those modules. Here, we demonstrate the effectiveness of tensor decomposition in quantifying the modules and those task-dependent modulations while overcoming the shortcomings inherent to previous methods. We further show that the tensor decomposition provides a new perspective on how the CNS switches between walking and running. The CNS modulated the peak timing in the temporal module while recruiting proximal muscles in the corresponding spatial module.

Although no damage occurs in the brains of individuals with spinal cord injury, structural and functional reorganization occurs in the sensorimotor cortex because of the deafferentation of afferent signal input from below the injury level. This brain reorganization that is specific to individuals with spinal cord injury is speculated to contribute to the improvement of the motor function of the remaining upper limbs. However, no study has investigated in detail the motor function above the injury level. To clarify this, we designed an experiment using the handgrip force steadiness task, which is a popular technique for evaluating motor function as the index of the variability of common synaptic input to motoneurons. Fourteen complete spinal cord injury (cSCI) individuals in the chronic phase, fifteen individuals with lower limb disabilities, and twelve healthy controls participated in the study. We clarified that the force steadiness in the cSCI group was significantly higher than that in the control groups, and that sports years were significantly correlated with this steadiness. Furthermore, multiple analyses revealed that force steadiness was significantly predicted by sports years. These results suggest that brain reorganization after spinal cord injury can functionally affect the remaining upper limb motor function. These findings may have implications in the clinical rehabilitation field, such as occupational rehabilitation of the upper limbs. They also indicate that individuals with complete spinal cord injury, based on their enhanced force adjustment skills, would excel at fine motor tasks such as manufacturing and handicrafts.

We investigated whether bilateral lower-limb control and leg dominance affect force control ability in 15 healthy young adults (9 males and 6 females, age =26.8 ± 4.1 years). Participants performed isometric ankle dorsiflexion force control tasks, matching a visual target (10% of maximal effort) as quickly and precisely as possible in ballistic and tonic tasks. Performance was evaluated using force error, force steadiness, amount of muscle activity of the tibialis anterior, and response time characteristics. Results showed no significant effects of leg dominance during both ballistic and tonic tasks, while bilateral condition resulted in significantly larger error, less force steadiness, compared to unilateral condition, and only during the tonic task. Consequently, bilateral control, specifically in tasks utilizing feedback control (i.e., tonic task) might affect force control ability, possibly because of the interhemispheric inhibition to meet bilateral task complexity and integrate afferent bilateral sensory information from both right and left legs.

Brain recognization in Paralympic athletes occurs in a manner dependent on both disability type and sport-specific trainings. The mechanisms underlying this reorganization likely include use-dependent plasticity and disability-specific compensations. Studies of the brains of Paralympic athletes and neurorehabilitation aim to better understand the neural mechanisms responsible for brain reorganization following physical rehabilitation interventions and/or athletic trainings.

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.

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.

Trunk muscles are responsible for maintaining trunk stability during sitting. However, the effects of anticipation of perturbation on trunk muscle responses are not well understood. The objectives of this study were to identify the responses of trunk muscles to sudden support surface translations and quantify the effects of anticipation of direction and time of perturbation on the trunk neuromuscular responses. Twelve able-bodied individuals participated in the study. Participants were seated on a kneeling chair and support surface translations were applied in the forward and backward directions with and without direction and time of perturbation cues. The trunk started moving on average approximately 40ms after the perturbation. During unanticipated perturbations, average latencies of the trunk muscle contractions were in the range between 103.4 and 117.4ms. When participants anticipated the perturbations, trunk muscle latencies were reduced by 16.8±10.0ms and the time it took the trunk to reach maximum velocity was also reduced, suggesting a biomechanical advantage caused by faster muscle responses. These results suggested that trunk muscles have medium latency responses and use reflexive mechanisms. Moreover, anticipation of perturbation decreased trunk muscles latencies, suggesting that the central nervous system modulated readiness of the trunk based on anticipatory information.

Highlights • We investigated cortical reorganization in an amputated archer who used his feet. • Lower-limb motor representations were examined using fMRI and TMS mapping. • M1 areas innervating lower-limb muscles were larger in the amputated athlete. • The toe and knee representations were expanded towards the lateral part of the M1. • Paralympic athletes have a unique and dynamic M1 plasticity., Despite their disabilities, top Paralympic athletes have better motor skills than able-bodied athletes. However, the neural underpinnings of these better motor skills remain unclear. We investigated the reorganization of the primary motor cortex (M1) in a Paralympic athlete with congenital amputation of both arms who holds the world record for the farthest accurate shot in archery (Amputee Archer: AA). We recorded brain activity during contraction of right toe, ankle, knee, and hip joint muscles in the AA and 12 able-bodied control subjects using functional magnetic resonance imaging. The results revealed that M1 activation was more widespread in the AA compared with control subjects during all tasks, and shifted towards the lateral part of the M1 during contraction of toe and knee muscles. We also conducted a motor mapping experiment using navigated transcranial magnetic stimulation. The M1 area receiving stimulation elicited motor-evoked potentials from the toe, lower-leg, and thigh muscles, which were larger in the AA compared with 12 control subjects. Furthermore, the AA's motor maps were shifted towards the lateral side of M1. These results suggest an expansion of lower-limb M1 representation towards the lateral side of M1, including the trunk and upper-limb representations, and an expansion of the area of corticomotor neurons innervating the lower limb muscles in the AA. This unique M1 reorganization could underpin the AA's excellent archery performance in the absence of upper limbs. The current results suggest that Paralympic athletes may exhibit extreme M1 plasticity, which could arise through a combination of rigorous long-term motor training and compensatory M1 reorganization for missing body parts.

Walking movements are orchestrated by the activation of a large number of muscles. The control of numerous muscles during walking is believed to be simplified by flexible activation of groups of muscles called muscle synergies. Although significant corticomuscular connectivity during walking has been reported, the level at which the cortex controls locomotor muscle activity (i.e., muscle synergy or individual muscle level) remains unclear. Here, we examined cortical involvement in muscle control during walking by brain decoding of the activation of muscle synergies and individual muscles from electroencephalographic (EEG) signals using linear decoder models. First, we demonstrated that activation of locomotor muscle synergies was decoded from slow cortical waves with significant accuracy. In addition, we found that decoding accuracy for muscle synergy activation was greater than that for individual muscle activation and that decoding of individual muscle activation was based on muscle synergy-related cortical information. Taken together, these results provide indirect evidence that the cerebral cortex hierarchically controls multiple muscles through a few muscle synergies during walking. Our findings extend the current understanding of the role of the cortex in muscular control during walking and could accelerate the development of effective brain-machine interfaces for people with locomotor disabilities.

OBJECTIVE Transcranial static magnetic field stimulation (tSMS) has recently been demonstrated to modulate cortical excitability and perceptual functions in humans, however, the effect of tSMS on motor behavior is still unknown. We investigated whether tSMS over the primary motor cortex (M1) alters voluntary ballistic force control. MATERIALS AND METHODS Twenty healthy participants performed ballistic pinch contractions in both hands alternatively at a predetermined submaximal force level and without visual feedback, before, during and after tSMS and sham interventions. A compact magnet for tSMS and a stainless steel cylinder for sham stimulation were positioned over the either right or left M1 for 15 min. RESULTS The absolute error to the target force level was significantly larger for the tSMS-intervened hand than for the sham-intervened hands during and after intervention (p < 0.05, respectively). Compared with the preintervention session, the absolute error increased in the tSMS-intervened hand during and after intervention (p < 0.05, respectively), but not in the sham-intervened hand. CONCLUSIONS tSMS over M1 can impair the accuracy of submaximal ballistic pinch force control. This suggests that tSMS is strong enough to alter motor behavior in humans.

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.

The capacity for auditory-motor coordination (AMC) is shared by several species, among which humans are most flexible in coordinating with tempo changes. We investigated how humans lose this tempo flexibility at their upper rate limit, and the effect of skill level on this phenomenon. Seven skilled street dancers, including a world champion, and ten non-dancers were instructed to bend their knees according to a metronome beat in a standing position at eight constant beat frequencies (3.8–5 Hz). Although maximum frequency of movement during the task was 4.8 Hz in the non-dancers and 5.0 Hz in the dancers, the rate limit for AMC was 4.1 Hz in the non-dancers and 4.9 Hz in the dancers. These results suggest that the loss of AMC was not due to rate limit of movement execution, but rather due to a constraint on the AMC process. In addition, mediation analysis revealed that a kinematic bias (i.e., the extent of knee flexion during the task) causally affected the extent of phase wandering via mediating factors (e.g., the extent to which movement frequency was reduced relative to the beat frequency). These results add evidence that gravity acts as constraint on AMC involving vertical rhythmic movement.

Regular exercise is effective for improving physical functions and pain relief in patients with lumbar and lowerlimb osteoarthritis. However, some of the patients have difficulty in performing land-based exercises due to disease status. This study focused on defensive style Nordic walking as a land-based exercise the patients can perform and aimed to investigate the effects of defensive style Nordic walking intervention on functional performance and pain in patients with lumbar and lower-limb osteoarthritis with low walking capacity. After 10 weeks of non-intervention, thirteen patients participated in the defensive style Nordic walking intervention for 10 weeks. The sixty-minute Nordic walking exercise was conducted once a week. Self-reported maximum walking distance without a break of the patients was less than 1 km and eight patients usually use a cane while walking. Functional performance measurement was conducted before non-intervention, after non-intervention (before intervention), and after intervention periods. Pain was measured using visual analog scale before and after each exercise session. Indices to estimate mobility, lower-limb muscle strength, and balance ability were improved by the intervention as compared with the non-intervention. Pain score decreased immediately after one session of defensive style Nordic walking and throughout the intervention period. These results suggest that the defensive style Nordic walking is an effective landbased exercise to improve mobility, lower-limb muscle strength, balance, and pain relief in patients with lumbar and lower-limb osteoarthritis with low walking capacity.

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.