Phase-dependent modulation of soleus H-reflex amplitude induced by rhythmic arm cycling

Rhythmic arm cycling is known to suppress the Hoffmann (H-) reflex amplitudes in the soleus (Sol) muscles of stationary legs. However, it has remained unclear if this suppression is modulated according to the phase of movement in the cycle path or is rather a general setting of excitability level related to rhythmic movement. In the present study we investigated the phase-dependent modulation of the Sol H-reflex induced by rhythmic arm cycling by examining reflex amplitudes at 12 phases of the arm cycle movement. Arm cycling tasks consisted of bilateral, ipsilateral and contralateral movement. Additionally, data were also sampled at 12 static arm positions mimicking those occurring during movement. H-reflexes were evoked and recorded at constant motor wave amplitudes across all conditions. Suppression of Sol H-reflex amplitude was dependent upon the phase of movement (main effect p < 0.0001) during arm cycling, but not during static positioning. Results suggest that locomotor central pattern generators may contribute to the phasic reflex modulation observed in this study. The phasic modulation was more pronounced during bilateral movement, however aspects of the neural control driving this modulation were also present during ipsilateral and contralateral movement.     

Facilitation of soleus H-reflex amplitude evoked by cutaneous nerve stimulation at the wrist is not suppressed by rhythmic arm movement

Neural connections between the cervical and lumbosacral spinal cord may assist in arm and leg coordination during locomotion. Currently the extent to which arm activity can modulate reflex excitability of leg muscles is not fully understood. We showed recently that rhythmic arm movement significantly suppresses soleus H-reflex amplitude probably via modification of presynaptic inhibition of the IA afferent pathway. Further, during walking reflexes evoked in leg muscles by stimulation of a cutaneous nerve at the wrist (superficial radial nerve; SR) are phase and task dependent. However, during walking both the arms and legs are rhythmically active thus it is difficult to identify the locus of such modulation. Here we examined the influence of SR nerve stimulation on transmission through the soleus H-reflex pathway in the leg during static contractions and during rhythmic arm movements. Nerve stimulation was delivered with the right shoulder in flexion or extension. H-reflexes were evoked alone (unconditioned) or with cutaneous conditioning via stimulation of the SR nerve (also delivered alone without H-reflex in separate trials). SR nerve stimulation significantly facilitated H-reflex amplitude during static contractions with the arm extended and countered the suppression of reflex amplitude induced by arm cycling. The results demonstrate that cutaneous feedback from the hand on to the soleus H-reflex pathway in the legs is not suppressed during rhythmic arm movement. This contrasts with the observation that rhythmic arm movement suppresses facilitation of soleus H-reflex when cutaneous nerves innervating the leg are stimulated. In conjunction with other data taken during walking, this suggests that the modulation of transmission through pathways from the SR nerve to the lumbosacral spinal cord is partly determined by rhythmic activity of both the arms and legs.     

Rhythmic leg cycling modulates forearm muscle H-reflex amplitude and corticospinal tract excitability

Rhythmic arm cycling leads to suppression of H-reflexes in both leg and arm muscles, and a reduction in the excitability of corticospinal projections to the forearm flexors. It is unknown, however, whether leg cycling modulates excitability in neural projections to the arms. Here we studied the extent to which rhythmic movement of the legs alters reflex (Experiment 1) and corticospinal (Experiment 2) transmission to arm muscles. In experiment 1, flexor carpi radialis (FCR) H-reflex recruitment curves were recorded with the legs static, and during rhythmic leg movement, while the FCR was both contracted and relaxed. The results indicate that rhythmic leg movement suppresses reflex transmission, both when FCR is at rest and during tonic contraction, but that the effect is not phase-dependent. In experiment 2, we used transcranial magnetic stimulation (TMS) to elicit motor-evoked potentials in the contracted and relaxed FCR during static leg, and leg cycling conditions. Sub-threshold TMS was also used to condition H-reflexes in order to provide specific information about cortical excitability during leg cycling. Both resting and tonically contracting arm muscles showed a greater corticospinal excitability during leg cycling than during the static leg condition. The magnitude of TMS facilitation of H-reflexes was similar during leg cycling and rest, suggesting a considerable sub-cortical component to the increased corticospinal excitability. The results suggest a differential regulation of afferent and descending projections to the arms during leg cycling, and are consistent with the idea that there is a loose, but significant, neural coupling between the arms and legs during rhythmic movement.     

Suppression of soleus H-reflex amplitude is graded with frequency of rhythmic arm cycling

In humans, rhythmic arm cycling has been shown to significantly suppress the soleus H-reflex amplitude in stationary legs. The specific nature of the relationship between frequency of arm cycling and H-reflex modulation in the legs has not been explored. We speculated that the effect of arm cycling on reflexes in leg muscles is related to the neural control of arm movement; therefore, we hypothesized that a graded increase in arm cycling frequency would produce a graded suppression of the soleus H-reflex amplitude. We also hypothesized that a threshold frequency of arm cycling would be identified at which the H-reflex amplitude significantly differed from static control trials (i.e., the arms were stationary). Soleus H-reflexes were evoked in stationary legs with tibial nerve stimulation during both control and rhythmic arm cycling (0.03–2.0 Hz) trials. The results show a significant inverse linear relation between arm cycling frequency and soleus H-reflex amplitude (P < 0.05). Soleus H-reflex amplitude significantly differed from control at an average threshold cycling frequency of 0.8 Hz. The results demonstrate that increased frequency of upper limb movement increases the intensity of interlimb influences on the neural activity in stationary legs. Further, a minimum threshold frequency of arm cycling is required to produce a significant effect. This suggests that achieving a threshold frequency of rhythmic arm movement may be important to incorporate in rehabilitation strategies to engage the appropriate interlimb neural pathways.     

Effect of rhythmic arm movement on reflexes in the legs: modulation of soleus H-reflexes and somatosensory conditioning

During locomotor tasks such as walking, running, and swimming, the arms move rhythmically with the legs. It has been suggested that connections between the cervical and lumbosacral spinal cord may mediate some of this interlimb coordination. However, it is unclear how these interlimb pathways modulate reflex excitability during movement. We hypothesized that rhythmic arm movement would alter the gain of reflex pathways in the stationary leg. Soleus H-reflexes recorded during arm cycling were compared with those recorded at similar positions with the arms stationary. Nerve stimulation was delivered with the right arm at approximately 70 degrees shoulder flexion or 10 degrees shoulder extension. H-reflexes were evoked alone (unconditioned) or with sural or common peroneal nerve (CP) conditioning to decrease or increase soleus IA presynaptic inhibition, respectively. Both conditioning stimuli were also delivered with no H-reflex stimulation. H-reflex amplitudes were compared at similar M-wave amplitudes and activation levels of the soleus. Arm cycling significantly reduced (P < 0.05) unconditioned soleus H-reflexes at shoulder flexion by 21.7% and at shoulder extension by 8.8% compared with static controls. The results demonstrate a task-dependent modulation of soleus H-reflexes between arm cycling and stationary trials. Sural nerve stimulation facilitated H-reflexes at shoulder extension but not at shoulder flexion during static and cycling trials. CP nerve stimulation significantly reduced H-reflex amplitude in all conditions. Reflexes in soleus when sural and CP nerve stimulation were delivered alone, were not different between cycling and static trials; thus the task-dependent change in H reflex amplitude was not due to changes in motoneuron excitability. Therefore modulation occurred at a pre-motoneuronal level, probably by presynaptic inhibition of the IA afferent volley. Results indicate that neural networks coupling the cervical and lumbosacral spinal cord in humans are activated during rhythmic arm movement. It is proposed that activation of these networks may assist in reflex linkages between the arms and legs during locomotor tasks.     

Rhythmic arm cycling produces a non-specific signal that suppresses Soleus H-reflex amplitude in stationary legs

Rhythmic arm cycling significantly suppresses Hoffmann (H-) reflex amplitude in Soleus muscles of stationary legs. The specific parameters of arm cycling contributing to this suppression, however, are unknown. Between the arms or legs, movement results in suppression of the H-reflex that is specifically related to the phase of movement and the locus of limb movement. We speculated that the effects of arm movement features on H-reflexes in the leg would be similar and hypothesized that the Soleus H-reflex suppression evoked by arm movement would therefore be specifically related to: (1) phase of the movement; (2) the locus of the movement (i.e., ipsilateral or contralateral arm); (3) range of arm motion; and (4) frequency of arm cycling. Participants performed bilateral arm cycling at 1 and 2 Hz with short and long-crank lengths. Ipsilateral and contralateral arm cycling was also performed at 1 Hz with a long-crank length. Soleus H-reflexes were evoked at four equidistant phases and comparisons were made while maintaining similar evoked motor waves and Soleus activation. Our results show that comparable suppressive effects were seen at all phases of the arm movement: there was no phase-dependence. Further, bilateral or unilateral (whether ipsi- or contralateral arm) cycling yielded equivalent suppression of the H-reflex amplitude. Cycling at 2 Hz resulted in a significantly larger suppression than with 1 Hz cycling. We conclude that a general, rather than a specific, signal related to the command to produce rhythmic arm muscle activity mediates the suppression of Soleus H-reflex during arm cycling.     

Enhancement of arm and leg locomotor coupling with augmented cutaneous feedback from the hand

Cutaneous feedback from the hand could assist with coordination between the arms and legs during locomotion. Previously we used a reduced walking model of combined arm and leg (ARM&LEG) cycling to examine the separate effects of rhythmic arm (ARM) and leg (LEG) movement. Here we use this same paradigm to test the modulation H-reflexes with and without interlimb cutaneous conditioning evoked by stimulating a nerve innervating the hand (superficial radial, SR). It was hypothesized that both ARM and LEG would contribute significantly to suppression of H-reflex amplitude during ARM&LEG. We also predicted a conservation of interlimb cutaneous conditioning during movement and an interaction between arm and leg rhythmic movement control. Subjects were seated in a recumbent ARM&LEG cycle ergometer and maintained a low-level soleus contraction for all tasks. H-reflex amplitude was facilitated by cutaneous conditioning evoked by stimulation of the SR nerve. H-reflex amplitudes were taken from recruitment curves and included modulation of 50% H max and H max. The suppressive effect of arm was less than that for LEG and ARM&LEG, while suppression during LEG and ARM&LEG were generally equivalent. For H-reflexes conditioned by cutaneous input, amplitudes during ARM&LEG instead were in between those for ARM and LEG modulation. Multiple regression analysis revealed a significant contribution for arm only in trials when SR stimulation was used to condition H-reflex amplitudes. We suggest that there is a measurable interaction between neural activity regulating arm and leg movement during locomotion that is specifically enhanced when cutaneous input from the hand is present.     

How does action resist visual illusion? Uncorrected oculomotor information does not account for accurate pointing in peripersonal space

The amplitudes and signs of cutaneous reflexes are modulated during rhythmic movements of the arms and legs (during walking and arm or leg cycling for instance). This reflex modulation is frequently independent of the background muscle activity and may involve central pattern generator (CPG) circuits. The purpose of the present study was to investigate the nature and degree of coupling between the upper limbs during arm cycling, with regard to the regulation of cutaneous reflexes. Responses to electrical stimulations of the right, superficial radial nerve (five 1 ms pulses, 300 Hz) were recorded bilaterally in six arm muscles of eight participants during arm cycling involving only the limb ipsilateral to the stimulation, only the limb contralateral to the stimulation, and bilateral movement when the limbs were both in-phase and 180° out of phase. The pattern of cutaneous reflex modulation throughout the arm cycle was independent of the functional state of the limb contralateral to the recording site, irrespective of whether recordings were made ipsilateral or contralateral to the stimulation. Furthermore, cutaneous reflexes were significantly (p<0.05) modulated with arm position in only 8% of cases in which the limb containing the responding muscle was either stationary or being moved passively by the experimenter. The results show that there is relatively weak coupling between the arms with regard to the regulation of cutaneous reflexes during rhythmic, cyclical arm movements. This suggests a loose connection between the CPGs for each arm that regulate muscle activity and reflex amplitude during rhythmic movement.     

Corticospinal excitability is lower during rhythmic arm movement than during tonic contraction

Humans perform rhythmic, locomotor movements with the arms and legs every day. Studies using reflexes to probe the functional role of the CNS suggest that spinal circuits are an important part of the neural control system for rhythmic arm cycling and walking. Here, by studying motor-evoked potentials (MEPs) in response to transcranial magnetic stimulation (TMS) of the motor cortex, and H-reflexes induced by electrical stimulation of peripheral nerves, we show a reduction in corticospinal excitability during rhythmic arm movement compared with tonic, voluntary contraction. Responses were compared between arm cycling and tonic contraction at four positions, while participants generated similar levels of muscle activity. Both H-reflexes and MEPs were significantly smaller during arm cycling than during tonic contraction at the midpoint of arm flexion (F = 13.51, P = 0.006; F = 11.83, P = 0.009). Subthreshold TMS significantly facilitated the FCR H-reflex during tonic contractions, but did not significantly modulate H-reflex amplitude during arm cycling. The data indicate a reduction in the responsiveness of cells constituting the fast, monosynaptic, corticospinal pathway during arm cycling and suggest that the motor cortex may contribute less to motor drive during rhythmic arm movement than during tonic, voluntary contraction. Our results are consistent with the idea that subcortical regions contribute to the control of rhythmic arm movements despite highly developed corticospinal projections to the human upper limb.     

Interlimb coupling from the arms to legs is differentially specified for populations of motor units comprising the compound H-reflex during “reduced” human locomotion

Recent experiments have identified neuromechanical interactions between the arms and legs during human locomotor movement. Previous work reported that during the rhythmic movement of all four limbs, the influence of the arms on reflex expression in the legs was superimposed on the dominant effect of the legs. This evidence was based upon studies using cutaneous and H-reflex modulation as indices of neuronal activity related to locomotion. The earlier H-reflex study was restricted to one phase of movement and to only a fixed H-reflex amplitude. Also, all four limbs were actively engaged in locomotor movement, and this led to the speculation that the effect from the arms could be underestimated by “swamping” of the conditioning during movement of the test limb. Work from the cat suggests that descending locomotor drive may be differentially specified for different motor unit populations in the hindlimb. Accordingly, details of interlimb coordination between the arms and legs in humans require further characterization and an examination of different populations of motor units as can be obtained from H-reflex recruitment curve (RC) parameters. Using modulation of H-reflex amplitudes across the entire ascending limb as neural probes for interlimb coupling, the present study evaluated the separated influences of rhythmic activity of the arms and leg on neuronal excitability of a stationary “test leg”. This three-limb “reduced” locomotion approach was applied using a stepping ergometer during the performance of three rhythmic movement tasks: arms (A); contralateral leg (L); and arms and contralateral leg (AL). Data were sampled at four different phases of the stepping cycle (using the moving leg as reference): start power (SP); end power (EP); start recovery (SR); and end recovery (ER). The main result was a large and significant influence of rhythmic AL activity on RC parameters of the H-reflex at EP and SP phases. However, the parameters (and thus motor unit populations) were differentially affected at each phase and task. For instance, a significant contribution of arms movement was noticed for H _max (largest motor units) at EP phase (P < 0.05), but no changes was observed for other parameters related to lower reflex amplitude (e.g., H-reflex evoked with an input that elicited 50% of maximum reflex response during static condition; H@50%). On the other hand, at SR phase, the parameter H@50% was significantly affected during AL compared to L. It is suggested that the remote effect from arms rhythmic activity has been differentially manifested across motor unit populations for each phase of movement. These findings provide definitive evidence for interlimb coupling between cervical and lumbar oscillators in gating the excitability of reflex pathways to a leg muscle for different populations of motorneurons within the pool. This further supports the contention of similar functional organization for locomotor networks in the human when compared to other animals. Additionally, these data provide additional confirmation of the significant role of the output of neural control for rhythmic arm movement in modulating reflex excitability of the legs that is specifically adjusted according to the phase and task.     

Modulation of human cutaneous reflexes during rhythmic cyclical arm movement

The organization and pattern of cutaneous reflex modulation is unknown during rhythmic cyclical movements of the human upper limbs. On the assumption that these cyclic arm movements are central pattern generator (CPG) driven as has been suggested for leg movements such as walking, we hypothesized that cutaneous reflex amplitude would be independent of electromyographic (EMG) muscle activation level during rhythmic arm movement (phase-dependent modulation, as is often the case in the lower limb during locomotion). EMG was recorded from eight muscles crossing the human shoulder, elbow, and wrist joints while whole arm rhythmic cyclical movements were performed. Cutaneous reflexes were evoked with trains of electrical stimulation delivered at non-noxious intensities (approximately 2 x threshold for radiating paresthesia) to the superficial radial nerve innervating the lateral portion of the back of the hand. Phasic bursts of rhythmic muscle activity occurred throughout the movement cycle. Rhythmic EMG and kinematic patterns were similar to what has been seen in the human lower limb during locomotor activities such as cycling or walking: there were extensive periods of reciprocal activation of antagonist muscles. For most muscles, cutaneous reflexes were modulated with the movement cycle and were strongly correlated with the movement-related background EMG amplitude. It is concluded that cutaneous reflexes are primarily modulated by the background muscle activity during rhythmic human upper limb movements, with only some muscles showing phase-dependent modulation.     

Rhythmic arm cycling differentially modulates stretch and H-reflex amplitudes in soleus muscle

During rhythmic arm cycling, soleus H-reflex amplitudes are reduced by modulation of group Ia presynaptic inhibition. This suppression of reflex amplitude is graded to the frequency of arm cycling with a threshold of 0.8 Hz. Despite the data on modulation of the soleus H-reflex amplitude induced by rhythmic arm cycling, comparatively little is known about the modulation of stretch reflexes due to remote limb movement. Therefore, the present study was intended to explore the effect of arm cycling on stretch and H-reflex amplitudes in the soleus muscle. In so doing, additional information on the mechanism of action during rhythmic arm cycling would be revealed. Although both reflexes share the same afferent pathway, we hypothesized that stretch reflex amplitudes would be less suppressed by arm cycling because they are less inhibited by presynaptic inhibition. Failure to reject this hypothesis would add additional strength to the argument that Ia presynaptic inhibition is the mechanism modulating soleus H-reflex amplitude during rhythmic arm cycling. Participants were seated in a customized chair with feet strapped to footplates. Three motor tasks were performed: static control trials and arm cycling at 1 and 2 Hz. Soleus H-reflexes were evoked using single 1 ms pulses of electrical stimulation delivered to the tibial nerve at the popliteal fossa. A constant M-wave and ~6% MVC activation of soleus were maintained across conditions. Stretch reflexes were evoked using a single sinusoidal pulse at 100 Hz given by a vibratory shaker placed over the triceps surae tendon and controlled by a custom-written LabView program. Results demonstrated that rhythmic arm cycling that was effective for conditioning soleus H-reflexes did not show a suppressive effect on the amplitude of the soleus stretch reflex. We suggest this indicates that stretch reflexes are less sensitive to conditioning by rhythmic arm movement, as compared to H-reflexes, due to the relative insensitivity to Ia presynaptic inhibition.     

Quadriceps H-reflex modulation during pedaling

The main aims of this study were 1) to investigate possible phase-, speed-, and task-dependent changes in the quadriceps H-reflex during pedaling, and to achieve this, 2) to develop an optimized H-reflex recording and processing procedure for recording of quadriceps H-reflexes during movement. It was hypothesized that the behavior of the quadriceps H-reflex concerning phase, speed, and task dependency corresponds to the behavior of the soleus H-reflex during rhythmical leg movements. The applied H-reflex procedure appeared to be reliable for obtaining the quadriceps H-reflex modulation during leg movement. The vastus lateralis (VL) and rectus femoris (RF) H-reflexes showed a phase-dependent modulation during pedaling at a frequency of 80 rpm with almost parallel changes in the reflex amplitude and motor recruitment level. However, when the speed of movement was reduced from 80 to 40 revolutions per minute (rpm) and crank load simultaneously increased (i.e., a halving of the movement speed with a constant motor recruitment level), the quadriceps H-reflex modulation pattern changed significantly in relation to the pattern of motor recruitment, i.e., at 40 rpm, the reflex excitability remained high during a gradual derecruitment during power generation in downstroke. Comparison of the "operationally defined H-reflex gain function" obtained during 1) pedaling at 80 rpm and 2) isometric quadriceps contractions in sitting position showed no significant task-dependent changes in the quadriceps H-reflex. Consequently, the hypothesis was only partly corroborated, and the findings indicate differences in the neural control of the soleus and the quadriceps muscle during rhythmical movements.     

Neural coupling between the upper and lower limbs in humans

The aim of this study was to establish the effects of active sinusoidal ipsilateral and contralateral upper limb flexion, extension, abduction, and adduction with elbows extended on the right soleus H-reflex with subjects seated and standing. Reflex effects were also established when both arms moved synchronously in a reciprocal pattern with elbows flexed in seated and standing subjects. Sinusoidal arm movements were timed to a metronome and performed at 0.2 Hz. Soleus H-reflexes were elicited only once (every 4s) in every movement cycle of the upper limbs. Position of arms, and activity of shoulder muscles were recorded through twin-axis goniometers and surface electromyography (EMG), respectively. We found that in seated subjects, regardless the direction of the active movement or the upper limb being moved, the soleus H-reflex was depressed. In standing subjects, a reflex depression was observed during extension, abduction, and adduction of the ipsilateral and contralateral upper limbs. Muscles were active during arm flexion and abduction in all directions of arm movement with subjects either seated or standing. It is suggested that arm movement might be incorporated in the rehabilitation training of people with a supraspinal or spinal cord lesion, since it can benefit motor recovery by decreasing spinal reflex excitability of the legs in these patients.     

Rhythmic arm cycling modulates Hoffmann reflex excitability differentially in the ankle flexor and extensor muscles

Rhythmic arm movement significantly suppresses H-reflex amplitude in the soleus (SOL) muscle. This is evidence of neural linkages between the arms and legs which can be exploited during locomotion and have been ascribed to the descending effects of CPGs for arm cycling. However, the generalizability of the effects of arm movement on reflex excitability within the lower leg musculature has not been confirmed, as findings have been limited solely to the ankle extensor group. Here we tested the hypothesis that rhythmic arm movement similarly modulates H-reflex amplitude in both the ankle flexors and extensors by observing responses in the SOL and tibialis anterior (TA) muscles. SOL and TA H-reflex recruitment curves were recorded bilaterally during control and 1 Hz arm cycling conditions. Our results showed significant suppression in H-reflex amplitude (H_max) in the SOL muscle in both the dominant and non-dominant legs during arm movement. However, results also revealed an unpredicted bidirectional (i.e. either suppression or facilitation) modulation of TA reflex amplitude that was not present in the SOL muscle. These findings suggest a differential regulation of ankle flexor and extensor H-reflex responses during rhythmic arm movement. This may be the result of differences in CPG output to the flexors and extensors during rhythmic movement, as well as increased involvement of cortical drive to the flexors relative to the extensors during rhythmic movement. These findings may be pertinent to future investigation of rehabilitative therapies that involve facilitative modulation of ankle flexor motor responses.     

Neural regulation of rhythmic arm and leg movement is conserved across human locomotor tasks

It has been proposed that different forms of rhythmic human limb movement have a common central neural control ('common core hypothesis'), just as in other animals. We compared the modulation patterns of background EMG and cutaneous reflexes during walking, arm and leg cycling, and arm-assisted recumbent stepping. We hypothesized that patterns of EMG and reflex modulation during cycling and stepping (deduced from mathematical principal components analysis) would be comparable to those during walking because they rely on similar neural substrates. Differences between the tasks were assessed by evoking cutaneous reflexes via stimulation of nerves in the foot and hand in separate trials. The EMG was recorded from flexor and extensor muscles of the arms and legs. Angular positions of the hip, knee and elbow joints were also recorded. Factor analysis revealed that across the three tasks, four principal components explained more than 93% of the variance in the background EMG and middle-latency reflex amplitude. Phase modulation of reflex amplitude was observed in most muscles across all tasks, suggesting activity in similar control networks. Significant correlations between EMG level and reflex amplitude were frequently observed only during static voluntary muscle activation and not during rhythmic movement. Results from a control experiment showed that strong correlation between EMG and reflex amplitudes was observed during discrete, voluntary leg extension but not during walking. There were task-dependent differences in reflex modulation between the three tasks which probably arise owing to specific constraints during each task. Overall, the results show strong correlation across tasks and support common neural patterning as the regulator of arm and leg movement during various rhythmic human movements.     

Short-term plasticity of spinal reflex excitability induced by rhythmic arm movement

Rhythmic arm movement reduces Hoffmann (H)-reflex amplitudes in leg muscles by modulation of presynaptic inhibition in group Ia transmission. To date only the acute effect occurring during arm movement has been studied. We hypothesized that the excitability of soleus H-reflexes would remain suppressed beyond a period of arm cycling conditioning. Subjects used a customized arm ergometer to perform rhythmic 1-Hz arm cycling for 30 min. H-reflexes were evoked before, during, and after arm cycling via stimulation of the tibial nerve in the popliteal fossa. The most important finding was that the H-reflex amplitudes were significantly suppressed during and 相似文献   

Modulation of cutaneous reflexes in arm muscles during walking: further evidence of similar control mechanisms for rhythmic human arm and leg movements

Stimulation of cutaneous nerves innervating the hand evokes prominent reflexes in many arm muscles during arm cycling. We hypothesized that the mechanisms controlling reflex modulation during the rhythmic arm swing of walking would be similar to that documented during arm cycling. Thus, we expected cutaneous reflexes to be modulated by position in the walking cycle (phase dependence) and be different when walking compared to contraction while standing (task dependence). Subjects performed static postures similar to those occurring during walking and also walked on a treadmill while the superficial radial nerve was electrically stimulated pseudorandomly throughout the step cycle. EMG was recorded bilaterally from upper limb muscles and kinematic recordings were obtained from the elbow and shoulder joints. Step cycle information was obtained from force-sensing insoles. Analysis was conducted after averaging contingent upon the occurrence of stimulation in the step cycle. Phase-dependent modulation of cutaneous reflexes at early (~50–80 ms) and middle (~80–120 ms) latencies was observed. Coordinated bilateral reflexes were seen in posterior deltoid and triceps brachii muscles. Task dependency was seen in that reflex amplitude was only correlated with background EMG during static contraction (75% of comparisons for both early and middle latency reflexes). During walking, no significant relationship between reflex amplitude and background EMG level was found. The results show that cutaneous reflex modulation during rhythmic upper limb movement is similar to that seen during arm cycling and to that observed in leg muscles during locomotion. These results add to the evidence that, during cyclical movements of the arms and legs, similar neural mechanisms observed only during movement (e.g. central pattern generators) control reflex output. Electronic Publication     

Excitability changes in human forearm corticospinal projections and spinal reflex pathways during rhythmic voluntary movement of the opposite limb

Rhythmic movements brought about by the contraction of muscles on one side of the body give rise to phase-locked changes in the excitability of the homologous motor pathways of the opposite limb. Such crossed facilitation should favour patterns of  bimanual coordination in which homologous muscles are engaged simultaneously, and disrupt those in which the muscles are activated in an alternating fashion. In order to examine these issues, we obtained responses to transcranial magnetic stimulation (TMS), to stimulation of the cervicomedullary junction (cervicomedullary-evoked potentials, CMEPs), to peripheral nerve stimulation (H-reflexes and f-waves), and elicited stretch reflexes in the relaxed right flexor carpi radialis (FCR) muscle during rhythmic (2 Hz) flexion and extension movements of the opposite (left) wrist. The potentials evoked by TMS in right FCR were potentiated during the phases of movement in which the left FCR was most strongly engaged. In contrast, CMEPs were unaffected by the movements of the opposite limb. These results suggest that there was systematic variation of the excitability of the motor cortex ipsilateral to the moving limb. H-reflexes and stretch reflexes recorded in right FCR were modulated in phase with the activation of left FCR. As the f-waves did not vary in corresponding fashion, it appears that the phasic modulation of the H-reflex was mediated by presynaptic inhibition of Ia afferents. The observation that both H-reflexes and f-waves were depressed markedly during movements of the opposite indicates that there may also have been postsynaptic inhibition or disfacilitation of the largest motor units. Our findings indicate that the patterned modulation of excitability in motor pathways that occurs during rhythmic movements of the opposite limb is mediated primarily by interhemispheric interactions between cortical motor areas.     

Phase-dependent modulation of the soleus H-reflex during rhythmical arm swing in humans

The purpose of this study is to investigate modulation of the soleus H-reflex during rhythmic arm swing in humans. Significant depression of the soleus H-reflex was observed when subjects swung their ipsilateral arms or both arms reciprocally during testing. The degree of soleus H-reflex depression appeared directly proportional to the speed of the arm swing. This depression was observed in the conditioning-testing intervals of 400, 500, and 600 msec during the ipsilateral backward arm swing and at the onset of the ipsilateral arm forward swing. This phase of depression partially overlapped the phase of depression of the soleus H-reflex during walking. However, the pattern of modulation during arm swing was not exactly the same as that during walking. Therefore, we concluded that the ipsilateral arm swing may partially affect the depression of the soleus H-reflex during the arm swing phase of walking but is not responsible for depression of the soleus H-reflex throughout the entire walking cycle.