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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.)

Upper- and lower-limb neuromuscular electrical stimulation (NMES) is known to modulate the excitability of the neural motor circuits. However, it remains unclear whether short-duration trunk muscle NMES could achieve similar neuromodulation effects. We assessed motor evoked potentials (MEPs) elicited through transcranial magnetic stimulation of the primary motor cortex representation of the trunk extensor muscles to evaluate corticospinal excitability. Moreover, cervicomedullary motor evoked potentials (CMEPs) were assessed through cervicomedullary junction magnetic stimulation to evaluate subcortical excitability. Twelve able-bodied individuals participated in the MEP study, and another twelve in the CMEP study. During the interventions, NMES was applied bilaterally to activate the erector spinae muscle and produce intermittent contractions (20 s ON/20 s OFF) for a total of 20 min while participants remained seated. Assessments were performed: (i) before; (ii) during (in brief periods when NMES was OFF); and (iii) immediately after the interventions to compare MEP or CMEP excitability. Our results showed that MEP responses were not affected by trunk NMES, while CMEP responses were facilitated for approximately 8 min during the intervention, and returned to baseline before the end of the 20 min stimulating period. Our findings therefore suggest that short-duration NMES of the trunk extensor muscles likely does not affect the corticospinal excitability, but it has a potential to facilitate subcortical neural circuits immediately after starting the intervention. These findings indicate that short-duration application of NEMS may be helpful in rehabilitation to enhance neuromodulation of the trunk subcortical neural motor circuits., (© 2023. The Author(s).)

In recent years, the neural control mechanisms of the arms and legs during human bipedal walking have been clarified. Rhythmic leg stepping leads to suppression of monosynaptic reflex excitability in forearm muscles. However, it is unknown whether and how corticospinal excitability of the forearm muscle is modulated during leg stepping. The purpose of the present study was to investigate the excitability of the corticospinal tract in the forearm muscle during passive and voluntary stepping. To compare the neural effects on corticospinal excitability to those on monosynaptic reflex excitability, the present study also assessed the excitability of the H-reflex in the forearm muscle during both types of stepping. A robotic gait orthosis was used to produce leg stepping movements similar to those of normal walking. Motor evoked potentials (MEPs) and H-reflexes were evoked in the flexor carpi radialis (FCR) muscle during passive and voluntary stepping. The results showed that FCR MEP amplitudes were significantly enhanced during the mid-stance and terminal-swing phases of voluntary stepping, while there was no significant difference between the phases during passive stepping. Conversely, the FCR H-reflex was suppressed during both voluntary and passive stepping, compared to the standing condition. The present results demonstrated that voluntary commands to leg muscles, combined with somatosensory inputs, may facilitate corticospinal excitability in the forearm muscle, and that somatosensory inputs during walking play a major role in monosynaptic reflex suppression in forearm muscle., (© 2023. The Author(s).)

The F-wave is a motor response elicited via the antidromic firings of motor nerves by the electrical stimulation of peripheral nerves, which reflects the motoneuron pool excitability. However, the F-wave generally has low robustness i.e., low persistence and small amplitude. We recently found that motor point stimulation (MPS), which provides the muscle belly with electrical stimulation, shows different neural responses compared to nerve stimulation, e.g., MPS elicits F-waves more robustly than nerve stimulation. Here, we investigated whether F-waves induced by MPS can identify changes in motoneuron pool excitability during handgrip and motor imagery. Twelve participants participated in the present study. We applied MPS on their soleus muscle and recorded F-waves during eyes-open (EO), eyes-closed (EC), handgrip (HG), and motor imagery (MI) conditions. In the EO and EC conditions, participants relaxed with their eyes open and closed, respectively. In the HG, participants matched the handgrip force level to 30% of the maximum voluntary force with visual feedback. In the MI, they performed kinesthetic MI of plantarflexion at the maximal strength with closed eyes. In the HG and MI, the amplitudes of the F-waves induced by MPS were increased compared with those in the EO and EC, respectively. These results indicate that the motoneuron pool excitability was facilitated during the HG and MI conditions, consistent with findings in previous studies. Our findings suggest that F-waves elicited by MPS can be a good tool in human neurophysiology to assess the motoneuron pool excitability during cognitive and motor tasks., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

In humans, peripheral sensory stimulation inhibits subsequent motor evoked potentials (MEPs) induced by transcranial magnetic stimulation; this process is referred to as short- or long-latency afferent inhibition (SAI or LAI, respectively), depending on the inter-stimulus interval (ISI) length. Although upper limb SAI and LAI have been well studied, lower limb SAI and LAI remain under-investigated. Here, we examined the time course of the soleus (SOL) muscle MEP following electrical tibial nerve (TN) stimulation at the popliteal fossa at ISIs of 20-220 ms. When the conditioning stimulus intensity was three-fold the perceptual threshold, MEP amplitudes were inhibited at an ISI of 220 ms, but not at shorter ISIs. TN stimulation just below the Hoffman (H)-reflex threshold intensity inhibited MEP amplitudes at ISIs of 30, 35, 100, 180 and 200 ms. However, the relationship between MEP inhibition and the P30 latency of somatosensory evoked potentials (SEPs) did not show corresponding ISIs at the SEP P30 latency that maximizes MEP inhibition. To clarify whether the site of afferent-induced MEP inhibition occurs at the cortical or spinal level, we examined the time course of SOL H-reflex following TN stimulation. H-reflex amplitudes were not significantly inhibited at ISIs where MEP inhibition occurred but at an ISI of 120 ms. Our findings indicate that stronger peripheral sensory stimulation is required for lower limb than for upper limb SAI and LAI and that lower limb SAI and LAI are of cortical origin. Moreover, the direct pathway from the periphery to the primary motor cortex may contribute to lower limb SAI., (© 2022. The Author(s).)

Non-invasive theta burst stimulation (TBS) can elicit facilitatory or inhibitory changes in the central nervous system when applied intermittently (iTBS) or continuously (cTBS). Conversely, neuromuscular electrical stimulation (NMES) can activate the muscles to send a sensory volley, which is also known to affect the excitability of the central nervous system. We investigated whether cortical iTBS (facilitatory) or cTBS (inhibitory) priming can affect subsequent NMES-induced corticospinal excitability. A total of six interventions were tested, each with 11 able-bodied participants: cortical priming followed by NMES (iTBS + NMES and cTBS + NMES), NMES only (iTBS sham  + NMES and cTBS sham  + NMES), and cortical priming only (iTBS + rest and cTBS + rest). After iTBS or cTBS priming, NMES was used to activate right extensor capri radialis (ECR) muscle intermittently for 10 min (5 s ON/5 s OFF). Single-pulse transcranial magnetic stimulation motor evoked potentials (MEPs) and maximum motor response (M max ) elicited by radial nerve stimulation were compared before and after each intervention for 30 min. Our results showed that associative facilitatory iTBS + NMES intervention elicited greater MEP facilitation that lasted for at least 30 min after the intervention, while none of the interventions alone were effective to produce effects. We conclude that facilitatory iTBS priming can make the central nervous system more susceptible to changes elicited by NMES through sensory recruitment to enhance facilitation of corticospinal plasticity, while cTBS inhibitory priming efficacy could not be confirmed., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Although voluntary muscle contraction modulates spinal reflex excitability of contracted muscles and other muscles located at other segments within a limb (i.e., intra-limb modulation), to what extent corticospinal pathways are involved in intra-limb modulation of spinal reflex circuits remains unknown. The purpose of the present study was to identify differences in the involvement of corticospinal pathways in intra-limb modulation of spinal reflex circuits among lower-limb muscles during voluntary contractions. Ten young males performed isometric plantar-flexion, dorsi-flexion, knee extension, and knee flexion at 10% of each maximal torque. Electromyographic activity was recorded from soleus, tibialis anterior, vastus lateralis, and biceps femoris muscles. Motor evoked potentials and posterior root-muscle reflexes during rest and isometric contractions were elicited from the lower-limb muscles using transcranial magnetic stimulation and transcutaneous spinal cord stimulation, respectively. Motor evoked potential and posterior root-muscle reflex amplitudes of soleus during knee extension were significantly increased compared to rest. The motor evoked potential amplitude of biceps femoris during dorsi-flexion was significantly increased, whereas the posterior root-muscle reflex amplitude of biceps femoris during dorsi-flexion was significantly decreased compared to rest. These results suggest that corticospinal and spinal reflex excitabilities of soleus are facilitated during knee extension, whereas intra-limb modulation of biceps femoris during dorsi-flexion appeared to be inverse between corticospinal and spinal reflex circuits., (© 2021. The Author(s).)

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.

Voluntary contraction facilitates corticospinal and spinal reflex circuit excitabilities of the contracted muscle and inhibits spinal reflex circuit excitability of the antagonist. It has been suggested that modulation of spinal reflex circuit excitability in agonist and antagonist muscles during voluntary contraction differs among lower-limb muscles. However, whether the effects of voluntary contraction on the excitabilities of corticospinal and spinal reflex circuits depend on the tested muscles remains unknown. The purpose of this study was to examine inter-muscle differences in modulation of the corticospinal and spinal reflex circuit excitabilities of multiple lower-limb muscles during voluntary contraction. Eleven young males performed isometric plantar-flexion, dorsi-flexion, knee extension, and flexion at low torque levels. Motor evoked potentials (MEPs) and posterior root-muscle reflexes from seven lower-leg and thigh muscles were evoked by transcranial magnetic stimulation and transcutaneous spinal cord stimulation, respectively, at rest and during weak voluntary contractions. MEP and posterior root-muscle reflex amplitudes of agonists were significantly increased as agonist torque level increased, except for the reflex of the tibialis anterior. MEP amplitudes of antagonists were significantly increased in relation to the agonist torque level, but those of the rectus femoris were slightly depressed during knee flexion. Regarding the posterior root-muscle reflex of the antagonists, the amplitudes of triceps surae and the hamstrings were significantly decreased, but those of the quadriceps femoris were significantly increased as the agonist torque level increased. These results demonstrate that modulation of corticospinal and spinal reflex circuit excitabilities during agonist and antagonist muscle contractions differed among lower-limb muscles.

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.

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.

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.

It is well known that contracting the upper limbs can affect spinal reflexes of the lower limb muscle, via intraneuronal networks within the central nervous system. However, it remains unknown whether neuromuscular electrical stimulation (NMES), which can generate muscle contractions without central commands from the cortex, can also play a role in such inter-limb facilitation. Therefore, the objective of this study was to compare the effects of unilateral upper limb contractions using NMES and voluntary unilateral upper limb contractions on the inter-limb spinal reflex facilitation in the lower limb muscles. Spinal reflex excitability was assessed using transcutaneous spinal cord stimulation (tSCS) to elicit responses bilaterally in multiple lower limb muscles, including ankle and thigh muscles. Five interventions were applied on the right wrist flexors for 70 s: (1) sensory-level NMES; (2) motor-level NMES; (3) voluntary contraction; (4) voluntary contraction and sensory-level NMES; (5) voluntary contraction and motor-level NMES. Results showed that spinal reflex excitability of ankle muscles was facilitated bilaterally during voluntary contraction of the upper limb unilaterally and that voluntary contraction with motor-level NMES had similar effects as just contracting voluntarily. Meanwhile, motor-level NMES facilitated contralateral thigh muscles, and sensory-level NMES had no effect. Overall, our results suggest that inter-limb facilitation effect of spinal reflex excitability in lower limb muscles depends, to a larger extent, on the presence of the central commands from the cortex during voluntary contractions. However, peripheral input generated by muscle contractions using NMES might have effects on the spinal reflex excitability of inter-limb muscles via spinal intraneuronal networks.

Transcutaneous spinal cord stimulation (tSCS) is a useful technique for the clinical assessment of neurological disorders. However, the characteristics of the spinal cord circuits activated by tSCS are not yet fully understood. In this study, we examined whether remote muscle contraction enhances the spinal reflexes evoked by tSCS in multiple lower-limb muscles. Eight healthy men participated in the current experiment, which required them to grip a dynamometer as fast as possible after the presentation of an auditory cue. Spinal reflexes were evoked in multiple lower-limb muscles with different time intervals (50-400 ms) after the auditory signals. The amplitudes of the spinal reflexes in all the recorded leg muscles significantly increased at 50-250 ms after remote muscle activation onset. This suggests that remote muscle contraction simultaneously facilitates the spinal reflexes in multiple lower-limb muscles. In addition, eight healthy men performed five different tasks (i.e., rest, hand grip, pinch grip, elbow flexion, and shoulder flexion). Compared to control values recorded just before each task, the spinal reflexes evoked at 250 ms after the auditory signals were significantly enhanced by the above tasks except for the rest task. This indicates that such facilitatory effects are also induced by remote muscle contractions in different upper-limb areas. The present results demonstrate the existence of a neural interaction between remote upper-limb muscles and spinal reflex circuits activated by tSCS in multiple lower-limb muscles. The combination of tSCS and remote muscle contraction may be useful for the neurological examination of spinal cord circuits.

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.

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 (M max ) was tested in the soleus by stimulating the posterior tibial nerve. Spinal reflexes and M max 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 (M max ). 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.

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.

The study aimed to compare the ability of dance and non-dance subjects to perform fine control of a simple heel-raising/lowering movement, and to determine if there are any differences in motor unit activity in the primary plantar flexor muscles during the movement. Subjects were instructed to accurately track a sinusoidal trace with a heel-raising and lowering movement at four controlled frequencies (1, 0.5, 0.25, and 0.125 Hz). The ankle joint angle was used to characterize movement errors from the target. Surface electromyography was recorded from the soleus and medial gastrocnemius muscles. One trial including five sinusoidal traces was divided into two phases: an up phase and a down phase. To characterize motor unit activity of the plantar flexor muscles, a wavelet transform was applied to electromyographic signals recorded in each phase. For both phases, errors in movement accuracy were lower in dancers than in controls (8.7 ± 4.6 vs. 11.5 ± 6.8%, P < 0.05) regardless of the frequency of the sinusoidal wave traced. During the down phase, peak power of soleus electromyographic signals at ~ 10 Hz was statistically larger in control subjects than in dancers (10.4 ± 0.7 vs. 6.3 ± 0.4% total power, P < 0.05). These results indicate that dancers have a higher degree of motor skill in a heel raise tracking task and exhibit adaptations in the motor unit activity during skilled dynamic movements.

The effect of development and aging on common modulation between bilateral plantarflexors (i.e., the right and left soleus, and the right and left medial gastrocnemius) (bilateral comodulation) and within plantarflexors in one leg (i.e., the right soleus and the right medial gastrocnemius) (unilateral comodulation) was investigated during bipedal quiet standing by comparing electromyography-electromyography (EMG) coherence among three age groups: adult (23-35 years), child (6-8 years), and elderly (60-80 years). The results demonstrate that there was significant coherence between bilateral plantarflexors and within plantarflexors in one leg in the 0- to 4-Hz frequency region in all three age groups. Coherence in this frequency region was stronger in the elderly group than in the adult group, while no difference was found between the adult and child groups. Of particular interest was the finding of significant coherence in bilateral and unilateral EMG recordings in the 8- to 12-Hz frequency region in some subjects in the elderly group, whereas it was not observed in the adult and child groups. These results suggest that aging affects the organization of bilateral and unilateral postural muscle activities (i.e., bilateral and unilateral comodulation) in the plantarflexors during quiet standing.

Unilateral isometric muscle contractions increase motor-evoked potentials (MEPs) produced by transcranial magnetic stimulation not only in the contracting muscle but also in the resting contralateral homologous muscle. Corticospinal excitability in the M1 contralateral to the contracting muscle changes depending on the type of muscle contraction. Here, we investigated the possibility that corticospinal excitability in M1 ipsilateral to the contracting muscle is modulated in a contraction-type-dependent manner. To this end, we evaluated MEPs in the resting left flexor carpi radialis (FCR) during unilateral shortening, lengthening, and isometric muscle contractions of the right wrist flexors at 10, 20, and 30% of maximal isometric contraction force. To compare the effects of different unilateral contractions on MEPs between the contracting and resting sides, MEPs in the right FCR were recorded on two separate days. In a separate experiment, we investigated the contraction specificity of the crossed effect at the spinal level by recording H-reflexes from the resting left FCR during contraction of the right wrist flexors. The results showed that MEPs in the contracting right FCR were the smallest during lengthening contraction. By contrast, MEPs in the resting left FCR were the largest during lengthening contraction, whereas the H-reflex was similar in the resting left FCR during the three types of muscle contraction. These results suggest that different types of unilateral muscle contraction asymmetrically modulate MEP size in the resting contralateral homologous muscle and in the contracting muscle and that this regulation occurs at the supraspinal level.

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.

The purpose of this study was to investigate how the recruitment properties of the corticospinal pathway are modulated in the soleus (SOL) and tibialis anterior (TA) muscles depending on postures. A wide range of stimulus intensities were applied via transcranial magnetic stimulation over the primary motor cortex during standing (STD) and sitting (SIT) with a comparable background activity level in each muscle. The relationship between the stimulation intensities and the size of motor-evoked potentials was assessed by the Boltzmann sigmoid function, which is characterized by a plateau value, maximum slope, and threshold. The plateau value and maximum slope were significantly higher during STD than during SIT in the SOL muscle (STD vs. SIT, plateau value: 50.0 +/- 21.8 vs. 33.9 +/- 12.3 mV ms, maximal slope: 1.6 +/- 0.7 vs. 1.2 +/- 0.5 mV ms/% maximal stimulator output). Similar changes of the parameters were also observed in the TA muscle (STD vs. SIT, plateau value: 71.0 +/- 22.9 vs. 41.4 +/- 16.1 mV ms, maximal slope: 5.0 +/- 2.0 vs. 2.5 +/- 0.7 mV ms/% maximal stimulator output). The threshold did not differ significantly between the two conditions and both muscles. These results indicate that the central nervous system requires a different control for each postural condition; that is, the relative balance of the excitatory and inhibitory inputs to the corticospinal pathways as well as the number of neurons of subliminal fringe in the corticospinal pathway was increased during STD compared with those during SIT.

Lengthening and shortening contractions are the fundamental patterns of muscle activation underlying various movements. It is still unknown whether or not there is a muscle-specific difference in such a fundamental pattern of muscle activation. The purpose of this study was, therefore, to investigate whether or not the relationship between lengthening and shortening contractions in the modulation of corticospinal excitability in the first dorsal interosseus (FDI) muscle is the same as that of previously tested muscles because the hand muscles are anatomically and functionally different from the other muscles. To this end, we investigated the relationship between the input-output curves of the corticospinal pathway (i.e., the relationship between the stimulus intensities vs. the area of motor-evoked potentials) during lengthening and shortening contractions in 17 healthy subjects. The shape of this relationship was sigmoidal and characterized by a plateau value, maximum slope, and threshold. The plateau value was at the same level between lengthening and shortening contractions. However, the maximum slope (P < 0.01) and threshold (P < 0.01) were significantly higher during lengthening contractions than during shortening contractions. These findings were different from the results of other muscles tested in previous studies (i.e., the soleus muscle and the elbow flexors). That is to say, the plateau value and the maximum slope during lengthening contractions were significantly lower than those during shortening contractions in previous studies. This study provides tentative evidence that the relationship between lengthening and shortening contractions in the modulation of corticospinal excitability differs between muscles, indicating that the underlying neural control is not necessarily the same even though the fundamental patterns of muscle activation are carried out.

The purpose of this study was to investigate the effects of hip proprioceptors on soleus stretch reflex excitability in standing humans. A custom-made device to stretch the ankle extensors was mounted on the lower leg portion of a gait orthosis and was used to elicit stretch reflex responses while standing. Six subjects with motor complete spinal cord injury (SCI) and six spinal intact subjects were placed in the orthosis, and stretch reflex responses were elicited when static and/or dynamic hip joint angle changes were imposed. We found that static hip extension significantly enhanced the stretch reflex responses as compared to the neutral position and the hip flexion position only in the SCI group. The EMG magnitude induced by hip extension was 142 +/- 16.6% greater than that induced by the neutral position. When the leg was dynamically swung, the reflex responses also changed with the phase of the hip angle in the SCI group; in particular, the reflex amplitude was enhanced with hip extension and in the transition phase from flexion to extension. Although the magnitude of the changes was less than that in the SCI group, a similar type of modulation was found in the normal group. Given the fact that the persons with SCI had lost the neural connection between higher nervous center and the paralyzed lower limb muscles, the mechanism underlying the present results can be attributed to the peripheral afferent input due to the hip angle changes. We concluded that hip mediated afferent input has a significant influence on the excitability modulation of the soleus stretch reflex pathway. Such neural modulation may play a role in the mechanism responsible for the phase-dependent modulation of the stretch reflex while walking.

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.

This is, to our knowledge, the first report demonstrating the effects of orthotic gait training on the activity of the spinal locomotor neural networks. Three subjects with complete spinal cord injury (SCI) performed 1-h training with reciprocating gait orthosis 5 days/week for 12 weeks. The results showed that after 3 (n=1) or 6 weeks (n=2) of training, EMG activities synchronized with locomotor rhythm appeared in the soleus muscle (SOL) in all subjects, although very little EMG activity accompanied the orthotic gait at the early training stage. Our results suggest that the induced modulation in the SOL EMG waveforms might be attributable to changes in the orthotic gait movement pattern, and/or changes in the interneuronal activities of the spinal locomotor neural networks, as a result of orthotic gait training.

Many studies have demonstrated that the firing behavior of single motor units varies in a nonlinear manner to the exerted torque during gradual muscle contraction and relaxation. However, it is unclear whether corticospinal excitability has such a hysteresis-like feature. In this study, we examined corticospinal excitability using transcranial magnetic stimulation (TMS) during gradual muscle contraction and relaxation for torque regulation in elbow flexor muscles. Eight healthy male subjects performed two different isometric elbow flexion tasks, namely, sinusoidal and tonic torque exertion tasks. In the sinusoidal task, the subjects sinusoidally increased and decreased the isometric elbow flexion torque (range of 0-15% of maximum voluntary contraction) at three different frequencies (0.33, 0.17, and 0.08 Hz). For each ascending (contraction: CON) and descending (relaxation: REL) period of the exerted torque, a single TMS was applied at 5 phases. In the tonic task, the elbow flexion torque was tonically exerted at 7 levels in a similar range as that in the sinusoidal task. EMG activities were recorded from the agonists, the biceps brachii (BB) and brachioradialis (BRD) muscles, and an antagonist, the triceps brachii (TB) muscle. The results demonstrated that the EMG activities of both the agonists and antagonist were larger in the CON period than the REL period, even when the exerted torque was the same. However, there were no significant differences in EMG activation profiles among the different frequencies of contraction. In BB and BRD, the motor-evoked potential (MEP) elicited by the TMS was also greater in the CON period than in the REL period. This CON-REL difference of MEP amplitudes was still observed when corrections were made for the increased EMG activities; that is, the MEP amplitudes to the identical EMG activities were greater in the CON period than in the REL period, and this phenomenon was more pronounced at higher frequencies. In addition, the degree to which sinusoidal MEPs exceeded tonic MEPs in the CON period and were smaller than tonic MEPs in the REL period became more pronounced at higher frequencies. On the other hand, there were significant correlations between the BB and BRD MEP amplitudes and the rate of change of elbow flexion/extension torque. These results indicate that corticospinal excitability during muscle contraction and relaxation has a neural hysteresis to the muscle activity, i.e., spinal motoneuronal activity, according to the rate of change of the exerted torque, i.e., muscle tension. This suggests that corticospinal excitability modulation depends not only on concurrent spinal motoneuronal activity and muscle tension but also on the time-series pattern of their changes during muscle contraction and relaxation.

The purpose of this study was to investigate how gravity level affects the excitability of the soleus muscle (SOL) motoneuron pool to Ia afferent input while erect posture is maintained in humans. Three healthy male subjects participated in an experiment whereby three different gravity conditions [microgravity (MG), normal gravity (NG), and hypergravity (HG)] were imposed using a parabolic flight procedure. The SOL H-reflex was evoked every 2 s while the subjects kept an erect posture. The stimulus intensity was controlled automatically on a real-time basis by personal computer to induce the constant amplitude of M-wave (10+/-5% of maximal M-wave amplitude). The background electromyographic activity (BGA) of the SOL was largest during HG, while it was almost absent during MG. The SOL H-reflex amplitude was significantly larger during HG and MG than during NG ( P<0.05). During NG and HG, there was a linear relationship between the BGA and the H-reflex amplitude; the difference in the SOL H-reflex amplitude between both gravity conditions could be explained in terms of the BGA level. However, during MG, despite the absence of BGA, the SOL H-reflex amplitude was larger than that during NG. Furthermore, when the subjects voluntarily activated the SOL by applying a load to the lower limb joints and spine by pulling a handle upward, this H-reflex enhancement almost disappeared. These results suggest that the somatosensory systems detecting a load at the lower limbs and/or vertebral column might play a role in reducing the excitability of the SOL motoneuron pool to Ia afferent inputs by presynaptic inhibition.

Abstract. The purpose of this study was to investigate the cortical activities during two types of Go/NoGo task with different movement instructions (Push-Go and Release-Go) using transcranial magnetic stimulation (TMS) and event-related potential (ERP) recordings. In the Push-Go condition, ten subjects were instructed either to push a button with their right index finger as fast as possible after a Go signal or not to push it after a NoGo signal. In the Release-Go condition, they were asked beforehand to continually depress a button by pushing, and instructed either to release it as fast as possible after a Go signal or not to release it after a NoGo signal. TMS was applied to the left primary motor cortex at 20--300 ms after each signal. In the Push-Go condition trials, the amplitude of the motor evoked potentials (MEPs) recorded from the right first dorsal interosseous muscle significantly increased at 120--300 ms after the Go signals and decreased at 160--200 ms after the NoGo signals. In contrast, the MEP amplitudes recorded during the Release-Go condition trials significantly decreased at 160--300 ms after the Go signals and significantly increased at 160--180 and 220--300 ms after the NoGo signals. On the other hand, the ERPs recorded in the frontocentral cortex after each signal for five of the subjects were identical in both the Push-Go and Release-Go condition trials. These results indicated that, while the cortical activities related to the Go/NoGo decision were similar in the two task conditions, the corticospinal excitability was modulated so as to suppress or facilitate the required Go responses depending on the given movement instructions. This suggests that the Go/NoGo decision may be separate from the subsequent motor program.

Human quiet standing is often modeled as a single inverted pendulum rotating around the ankle joint, under the assumption that movement around the hip joint is quite small. However, several recent studies have shown that movement around the hip joint can play a significant role in the efficient maintenance of the center of body mass (COM) above the support area. The aim of this study was to investigate how coordination between the hip and ankle joints is controlled during human quiet standing. Subjects stood quietly for 30 s with their eyes either opened (EO) or closed (EC), and we measured subtle angular displacements around the ankle (thetaa) and hip (thetah) joints using three highly sensitive CCD laser displacement sensors. Reliable data were obtained for both angular displacement and angular velocity (the first derivative of the angular displacement). Further, measurement error was not predominant, even among the angular acceleration data, which were obtained by taking the second derivative of the angular displacement. The angular displacement, velocity, and acceleration of the hip were found to be significantly greater (P<0.001) than those of the ankle, confirming that hip-joint motion cannot be ignored, even during quiet standing. We also found that a consistent reciprocal relationship exists between the angular accelerations of the hip and ankle joints, namely positive or negative angular acceleration of ankle joint is compensated for by oppositely directed angular acceleration of the hip joint. Principal component analysis revealed that this relationship can be expressed as: thetah=gammathetaa with gamma=-3.15+/-1.24 and gamma=-3.12+/-1.46 (mean +/-SD) for EO and EC, respectively, where theta is the angular acceleration. There was no significant difference in the values of y for EO and EC, and these values were in agreement with the theoretical value calculated assuming the acceleration of COM was zero. On the other hand, such a consistent relationship was never observed for angular displacement itself. These results suggest that the angular motions around the hip and ankle joints are not to keep the COM at a constant position, but rather to minimize acceleration of the COM.

Recent studies have demonstrated that coordinated stepping movements can be induced in patients with complete para-/tetraplegia, when they were standing on a moving treadmill with their body weight partially unloaded and external assistance. The aim of this study was to determine which part of the spinal cord generated the locomotor pattern. In patients with complete paraplegia due to lesions at different levels of the spinal cord, the locomotor pattern was compared with that of healthy subjects. Any similarities in electromyographic (EMG) activity of gastrocnemius and tibialis anterior muscles between the patients and healthy subjects were reflected by the analysis of the variation ratio and amplitudes of the EMG activity. It was found that the higher the level of spinal cord lesion the more "normal" was the locomotor pattern. This suggests that neuronal circuits underlying locomotor "pattern generation" in man are not restricted to any specific level(s) of the spinal cord, but that an intricate neuronal network contributing to bipedal locomotion extends from thoracolumbal to cervical levels.

We examined the lower-limb electromyographic (EMG) activity from a patient with clinically complete spinal cord injury during orthotic gait. A newly developed gait orthosis was used to obtain bipedal locomotion. The surface EMG data during the gait together with the biomechanical variables were collected by way of a radio EMG system. A cyclic EMG activation pattern corresponding to the gait cycles were observed in each of the paralyzed lower-limb muscles during the orthotic gait. Although the EMG activation did not seem to contribute toward generating the gait, it showed some similarities to that of the infant stepping or immature gait. These results might be regarded as one of the indirect pieces of evidence that suggest the existence of a spinally originating motor mechanism underlying human locomotion.

Stretch reflex responses in three elbow flexor muscles - the brachioradialis and the short and long heads of the biceps brachii - were studied during different motor tasks. The motor tasks were iso-velocity (8 deg/s) elbow flexion movements in which the muscles performed shortening or lengthening contractions, or were isometric contractions. Care was taken to maintain constant background electromyographic (EMG) activity in the brachoradialis muscle at a 50-deg elbow angle across the tasks by changing the magnitude of the initial load. During each task, mechanical perturbations (duration 170 ms) were applied at pseudorandom intervals when the elbow angle was 50 deg. The magnitude of the perturbation was varied across tasks in order to induce an elbow extension velocity of 80 deg/s over the first 50 ms after the onset of perturbation. The stretch reflex EMG responses in all muscles varied across the three tasks, despite a constant EMG level and similar perturbation-induced angular velocity in the direction of elbow extension. In particular, both the short- and long-latency reflex EMG components were reduced during the lengthening contractions. Further, the task-dependent variations in the early (M2) and the late (M3) components of the long-latency reflex were different, i.e., the magnitude of M3 was considerably enhanced during the shortening task as compared with that of M2. These findings suggest that central modification was responsible for the task-dependent modulation of late EMG responses.

In our previous study, 1/f beta-type power spectrum with the spectral exponent beta significantly greater than zero was found in the variability of soleus H-reflex amplitudes. This result indicated that the H-reflex variability was time-correlated owing to fractal characteristics. Furthermore, it was also suggested that the fractal characteristics were generated at the spinal level. The purpose of the present study was to investigate whether the fractal nature of the H-reflex variability was influenced by the loss of supraspinal input. Six healthy normal subjects and seven patients with spinal cord injury participated in this study. Soleus H-reflexes were evoked every 1 s from both legs simultaneously (stimulation intensity: motor threshold) and 1050 successive amplitudes of the H-reflex were recorded. The H-reflex sequence evoked from each leg was analyzed by "coarse graining spectral analysis" to calculate the spectral exponent beta. The value of beta was used to evaluate the level of time-correlation (fractal correlation). Cross-spectral analysis was used to evaluate the degree of synchronization between the H-reflex sequences evoked from both legs. The beta values for normal subjects (0.84 +/- 0.33, left leg; 0.88 +/- 0.34. right leg) were significantly greater (P < 0.001) than those for patients (0.31 +/- 0.18, left leg; 0.32 +/- 0.14, right leg), suggesting that the H-reflex sequences for normal subjects were more time-correlated than for patients. In the frequency range less than 0.2 Hz, the coherence of both legs was high (0.41 +/- 0.14) for normal subjects as compared to 0.20 +/- 0.12 for patients (P < 0.005). In this frequency range, the phase was almost 0 rad for normal subjects, indicating that the H-reflex variabilities of both legs were synchronized. These results suggested that (1) the strong fractal correlation observed in the H-reflex sequences for normal subjects was associated with supraspinal input, and (2) such supraspinal input had equal influence on the reflex arcs of the soleus of both legs.

The amplitude of the H-reflex has been known to have considerable variability even if the intensity of the stimulation is held constant. However, previous studies largely ignored the time-dependent profile of this variability. Recent mathematical analyses have shown that some seemingly irregular biological signals have fractal properties. A fractal time series is characterized by the property of self-similarity (self-affinity), and has long-range time correlation. The aim of this study was to investigate the question of whether the fluctuation of H-reflex was fractal with strong time-correlation. Soleus H-reflexes were evoked in five healthy subjects at two levels of stimulation intensity [1.2 MT (motor threshold) and 0.9 MT] every 1 s and 1050 successive amplitudes of H-wave and M-wave were recorded twice. The sequences of the H-wave and the M-wave amplitudes were analyzed by "coarse graining spectral analysis" to calculate the percentage of random fractal components in the sequences (%Fractal) and the spectral exponent beta. The %Fractal values of both sequences were above 90% [H-wave: 93.3 +/- 2.3% (1.2 MT), 91.6 +/- 3.1% (0.9 MT); M-wave: 94.3 +/- 3.3%; mean +/- SD]. Nonflat power spectra of the fractal components were observed for the H-wave sequences regardless of the stimulation intensity [beta = 0.75 +/- 0.26 (1.2 MT), 0.80 +/- 0.39 (0.9 MT)], indicating that the sequences were strongly time correlated. On the other hand, the M-wave sequences had a flatter spectrum (beta = 0.26 +/- 0.14) which was close to uncorrelated white noise. These results indicated that: (1) the fractal correlation found in the H-wave sequences was caused neither by the conduction through nerve fibers nor by the transmission at the neuromuscular junction, because the M-wave sequence had a significantly weaker time correlation, and (2) antidromic impulses in a motor nerve induced by the stimulation made a minor contribution to the generation of fractal correlation in the H-wave sequences, because it was preserved when the stimulation intensity was below MT. It was suggested that the fractal correlation in human H-reflex was generated at the synaptic connections to alpha-motoneurons in the spinal cord.