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.     

Specific modulation of the Hoffmann reflex cutaneous facilitation during a reaction-time task

Summary The Hoffmann (H) reflex and its facilitation produced by electrical stimulation of the sural area were examined before a ballistic extension of the right foot. Modulations of the cutaneous facilitation of the H reflex (CFH) were used to assess the control exerted over the transmission of low threshold cutaneous afferents. The time-course of H and CFH changes were investigated at the end of the foreperiod and during the premotor period, i.e. between the response signal and the onset of the electromyogram (EMG) of the soleus muscle. Four stimulation conditions were set up depending on whether the H reflex was elicited on the contracting or non-contracting limb, and whether cutaneous stimulation was ipsilateral or contralateral to the reflex. During the 100 ms preceding the response signal, the inhibition of the H reflex was more marked in the contracting limb than in the noncontracting limb. At the end of the foreperiod, the CFHs had a symmetric time course: the CFHs evoked by conditioning stimulation of the contracting limb were facilitated just before the response signal, while those produced by conditioning stimulation to the non-contracting limb were depressed. It is suggested that these variations are related to postural adjustments taking place before the movement is performed. As previously reported, the H reflex of the contracting limb exhibited a marked increase in amplitude over the 50 ms preceding the EMG. Reflex facilitations showed specific variations according to the ankle stimulated and the soleus muscle tested. The CFHs produced by stimulation of the non-contracting limb regained, at the end of the premotor period, a value close to their reference level recorded in trials without movement. The CFHs, produced by conditioning stimulation of the contracting limb were modulated differently according to whether the tested soleus muscle was contracting or not: when the CFH was tested on the contracting muscle, it was found to be depressed throughout the premotor period; this contrasted significantly with the isolated depression recorded on the non-contracting muscle. Therefore, only the cutaneous afferents from the mobilized limb, modulating the H reflex of the same limb, were subject to a specific inhibition during the premotor period. Throughout the preparatory and premotor periods, negative correlations were observed between H and CFH amplitude, except just before the EMG onset in the condition where the H reflex was delivered to the contracting muscle and the cutaneous stimulation to the ankle of the non-contracting limb: in this case, CFH amplitude increase paralleled that of the reflex amplitude. The negative correlations between these two variables are discussed in terms of a central control regulating the balance between proprioceptive and cutaneous input.     

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     

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.     

Comparison of external load compensation during rhythmic arm movements and rhythmic jaw movements in humans.

Experiments were performed on human elbow flexor and extensor muscles and jaw-opening and -closing muscles to observe the effect on rhythmic movements of sudden loading. The load was provided by an electromagnetic device, which simulated the appearance of a smoothly increasing spring-like load. The responses to this loading were compared in jaw and elbow movements and between expected and unexpected disturbances. All muscles showed electromyographic responses to unexpected perturbations, with latencies of approximately 65 ms in the arm muscles and 25 ms in the jaw. When loading was predictable, anticipatory responses started in arm muscles approximately 200 ms before and in jaw muscles 100 ms before the onset of loading. The reflex responses relative to the anticipatory responses were smaller for the arm muscles than for the jaw muscles. The reflex responses in the arm muscles were the same with unexpected and expected perturbations, whereas anticipation increased the reflex responses in the jaw muscles. Biceps brachii and triceps brachii showed similar sensory-induced responses and similar anticipatory responses. Jaw muscles differed, however, in that the reflex response was stronger in masseter than in digastric. It was concluded that reflex responses in the arm muscles cannot overcome the loading of the arm adequately, which is compensated by a large centrally programmed response when loading is predictable. The jaw muscles, particularly the jaw-closing muscles, tend to respond mainly through reflex loops, even when loading of the jaw is anticipated. The differences between the responses of the arm and the jaw muscles may be related to physical differences. For example, the jaw was decelerated more strongly by the load than the heavier arm. The jaw was decelerated strongly but briefly, <30 ms during jaw closing, indicating that muscle force increased before the onset of reflex activity. Apparently, the force-velocity properties of the jaw muscles have a stabilizing effect on the jaw and have this effect before sensory induced responses occur. The symmetrical responses in biceps and triceps indicate similar motor control of both arm muscles. The differences in reflex activity between masseter and digastric muscle indicate fundamental differences in sensory feedback to the jaw-closing muscle and jaw-opening muscle.     

Context-dependent modulation of interlimb cutaneous reflexes in arm muscles as a function of stability threat during walking

Cutaneous reflexes evoked in the muscles of the arms with electrical stimulation of nerves of the foot ("interlimb reflexes") are observed during walking. These reflexes have been suggested to coordinate the actions of the legs and arms when walking is disturbed. Recently, we showed that cutaneous reflexes evoked in the leg muscles after stimulation at the foot are modulated according to the level of postural threat during walking. We hypothesized that the amplitude of interlimb cutaneous reflexes would similarly be modulated when subjects walk in unstable environments. Subjects walked on a treadmill under four walking conditions: 1) normal; 2) normal with unpredictable anterior-posterior (AP) perturbations; 3) arms crossed; and 4) arms crossed with unpredictable AP perturbations. Interlimb reflexes evoked from electrical stimulation of the right superficial peroneal or sural nerves were recorded bilaterally, at four points of the step cycle. These reflexes were compared between conditions in which the arms were moving in a similar manner: 1) normal versus AP walking and 2) arms crossed versus arms crossed with AP perturbations. Differences in reflex amplitudes between arms-crossed conditions were observed in most upper limb muscles when subjects were perturbed while walking compared with undisturbed walking. This effect was less apparent when the arms were swinging freely. The results indicate that the strength of interlimb connections is influenced by the level of postural threat (i.e., the context of the behavior), thereby suggesting that these reflexes serve a functional link between the legs and arms during locomotion.     

Finger coordination during discrete and oscillatory force production tasks

We used the framework of the uncontrolled manifold (UCM) hypothesis to analyze the structure of finger force variability in discrete (ramp) and oscillatory force production tasks performed by the index and middle fingers of the right hand acting in parallel. Subjects performed the tasks at fast and slow rates, with and without a visual template presented on the screen. The variance of finger forces was partitioned into two components, compensated variance (V(COMP)), which did not affect total force, and uncompensated variance (V(UN)), which affected total force. Only minor effects of task (discrete or oscillatory) and of template (with or without) were seen on the variance profiles, leading us to conclude that the basic principles of synergy organization are common across discrete and oscillatory tasks. In contrast, the rate of force production had major effects on the structure of force variance. A modification of Goodman's model of motor variability was used to analyze the dependences V(UN) and V(COMP) on the magnitude of force and on the rate of force production. V(UN) showed a strong relation to the rate of force production and only weak dependence on the magnitude of force. In contrast, V(COMP) showed minimal effects of the rate of force production and strong effects of the force magnitude. The findings are interpreted as demonstrations of a limitation in the ability of the central nervous system to organize a two-finger synergy such that errors in the timing of individual finger force profiles are canceling each other's effects on the total force. In contrast, the synergy is efficiently intercompensating errors related to imprecise setting of force magnitudes of the two fingers.     

Muscle metabolism from near infrared spectroscopy during rhythmic handgrip in humans

The rate of metabolism in forearm flexor muscles (MO₂) was derived from near-infrared spectroscopy (NIRS-O₂) during ischaemia at rest rhythmic handgrip at 15% and 30% of maximal voluntary contraction (MVC), post-exercise muscle ischaemia (PEMI), and recovery in seven subjects. The MO₂ was compared with forearm oxygen uptake (O₂) [flow?×?(oxygen saturation in arnterial blood-oxygen saturation in venous blood, S _aO₂?S _vO₂)], and with the ³¹P-magnetic resonance spectroscopy-determined ratio of inorganic phosphate to phosphocreatine (P_I:PCr). During ischaemia at rest, the fall in NIRS-O₂ was more pronounced [76 (SEM 3) to 3 (SEM 1)%] than in S _vO₂ [71 (SEM 3) to 59 (SEM 2)%]. During the handgrip, NIRS-O₂ was lower at 30% compared to 15% MVC [58 (SEM 3) vs 67 (SEM 3)%] while the S _vO₂ was similar [29 (SEM 3) vs 31 (SEM 4)%]. Accordingly, MO₂ as well as P_I:PCr increased twofold, while V˙O₂ increased only 30%. During PEMI after 15% and 30% MVC, NIRS-O₂ fell to 9 (SEM 1)% and “0”, but the use of oxygen by forearm muscles was not reflected in S _vO₂. During reperfusion after PEMI, the peak NIRS-O₂ was lowest after intense exercise, while for S _vO₂ the reverse was seen. The discrepancies between NIRS-O₂ and S _vO₂, and therefore between the estimates of the metabolic rate, would suggest significant limitations in sampling venous blood which is representative of the flexor muscle capillaries. In support of this contention, S _vO₂ and venous pH decreased during the first seconds of reperfusion after PEMI. To conclude, NIRS-O₂ of forearm flexor muscles closely reflected the exercise intensity and the metabolic rate determined by magnetic resonance spectroscopy but not that rate derived from flow and the arterio-venous oxygen difference.     

Relationship between stretch reflex thresholds and voluntary arm muscle activation in patients with spasticity

Previous studies have shown that deficits in agonist–antagonist muscle activation in the single-joint elbow system in patients with spastic hemiparesis are directly related to limitations in the range of regulation of the thresholds of muscle activation. We extended these findings to the double-joint, shoulder-elbow system in these patients. Ten non-disabled individuals and 11 stroke survivors with spasticity in upper limb muscles participated. Stroke survivors had sustained a single unilateral stroke 6–36 months previously, had full pain-free passive range of motion of the affected shoulder and elbow and had some voluntary control of the arm. EMG activity from four elbow and two shoulder muscles was recorded during quasi-static (<5°/s) stretching of elbow flexors/extensors and during slow voluntary elbow flexion/extension movement through full range. Stretches and active movements were initiated from full elbow flexion or extension with the shoulder in three different initial positions (60°, 90°, 145° horizontal abduction). SRTs were defined as the elbow angle at which EMG signals began to exceed 2SD of background noise. SRT angles obtained by passive muscle stretch were compared with the angles at which the respective muscles became activated during voluntary elbow movements. SRTs in elbow flexors were correlated with clinical spasticity scores. SRTs of elbow flexors and extensors were within the biomechanical range of the joint and varied with changes in the shoulder angle in all subjects with hemiparesis but could not be reached in this range in all healthy subjects when muscles were initially relaxed. In patients, limitations in the regulation of SRTs resulted in a subdivision of all-possible shoulder-elbow arm configurations into two areas, one in which spasticity was present (“spatial spasticity zone”) and another in which it was absent. Spatial spasticity zones were different for different muscles in different patients but, taken together, for all elbow muscles, the zones occupied a large part of elbow-shoulder joint space in each patient. The shape of the boundary between the spasticity and no-spasticity zones depended on the state of reflex inter-joint interaction. SRTs in single- and double-joint flexor muscles correlated with the positions at which muscles were activated during voluntary movements, for all shoulder angles, and this effect was greater in elbow flexor muscles (brachioradialis, biceps brachii). Flexor SRTs correlated with clinical spasticity in elbow flexors only when elbow muscles were at mid-length (90°). These findings support the notion that motor impairments after CNS damage are related to deficits in the specification and regulation of SRTs, resulting in the occurrence of spasticity zones in the space of elbow-shoulder configurations. It is suggested that the presence of spatial spasticity zones might be a major cause of motor impairments in general and deficits in inter-joint coordination in particular in patients with spasticity.     

Interindividual differences in H reflex modulation during normal walking

Based on previous studies, at least two different types of soleus Hoffmann (H) reflex modulation were likely to be found during normal human walking. Accordingly, the aim of the present study was to identify different patterns of modulation of the soleus H reflex and to examine whether or not subjects with different H reflex modulation would exhibit different walking mechanics and different EMG activity. Fifteen subjects walked across two force platforms at 4.5 km/h (+/-10%) while the movements were recorded on video. The soleus H reflex and EMG activity were recorded separately during treadmill walking at 4.5 km/h. Using a two-dimensional analysis joint angles, angular velocities, accelerations, linear velocities and accelerations were calculated, and net joint moments about the ankle, knee and hip joint were computed by inverse dynamics from the video and force plate data. Six subjects (group S) showed a suppressed H reflex during the swing phase, and 9 subjects (group LS) showed increasing reflex excitability during the swing phase. The plantar flexor dominated moment about the ankle joint was greater for group LS. In contrast, the extensor dominated moment about the knee joint was greater for the S group. The hip joint moment was similar for the groups. The EMG activity in the vastus lateralis and anterior tibial muscles was greater prior to heel strike for the S group. These data indicate that human walking exhibits at least two different motor patterns as evaluated by gating of afferent input to the spinal cord, by EMG activity and by walking mechanics. Increasing H reflex excitability during the swing phase appears to protect the subject against unexpected perturbations around heel strike by a facilitated stretch reflex in the triceps surae muscle. Alternatively, in subjects with a suppressed H reflex in the swing phase the knee joint extensors seem to form the primary protection around heel strike.     

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.     

Muscle activity and fatigue in the shoulder muscles during repetitive work

Summary The amplitude distribution probability function (ADPF) and the power spectrum of the surface electromyogram from m. deltoideus anterior, m. infraspinatus and m. trapezius pars descendens were analyzed from 7 persons working at a pillar drill. Recordings were performed 6 times during a working day. The ADPF was analyzed from 2–3 work-cycles from each recording. The static contraction level was 11.0% MCV in m. deltoideus anterior, 8.5% MVC in m. infraspinatus, and 20.5% MCV in m. trapezius, without any change occuring throughout the day. When compared to previous suggested upper limits of ADPF levels, both the static and the medium contraction levels were too high in the performance of this particular task. The power spectrum of the EMG was analyzed during isometric contractions of the shoulder muscles. The mean power frequency decreased during the day in m. trapezius only, suggesting muscular fatigue in this area.     

Muscle oxygenation during incremental arm and leg exercise in men and women

The purpose of this study was to compare the rates of muscle deoxygenation in the exercising muscles during incremental arm cranking and leg cycling exercise in healthy men and women. Fifteen men and 10 women completed arm cranking and leg cycling tests to exhaustion in separate sessions in a counterbalanced order. Cardiorespiratory measurements were monitored using an automated metabolic cart interfaced with an electrocardiogram. Tissue absorbency was recorded continuously at 760?nm and 850?nm during incremental exercise and 6?min of recovery, with a near infrared spectrometer interfaced with a computer. Muscle oxygenation was calculated from the tissue absorbency measurements at 30%, 45%, 60%, 75% and 90% of peak oxygen uptake (V˙O₂) during each exercise mode and is expressed as a percentage of the maximal range observed during exercise and recovery (%Mox). Exponential regression analysis indicated significant inverse relationships (P?2 during arm cranking and leg cycling in men (multiple R?=??0.96 and ?0.99, respectively) and women (R?=?0.94 and ?0.99, respectively). No significant interaction was observed for the %Mox between the two exercise modes and between the two genders. The rate of muscle deoxygenation per litre of V˙O₂ was 31.1% and 26.4% during arm cranking and leg cycling, respectively, in men, and 26.3% and 37.4% respectively, in women. It was concluded that the rate of decline in %Mox for a given increase in V˙O₂ between 30% and 90% of the peak V˙O₂ was independent of exercise mode and gender.     

Neural control of rhythmic human arm movement: phase dependence and task modulation of hoffmann reflexes in forearm muscles

Although we move our arms rhythmically during walking, running, and swimming, we know little about the neural control of such movements. Our working hypothesis is that neural mechanisms controlling rhythmic movements are similar in the human lumbar and cervical spinal cord. Thus reflex modulation during rhythmic arm movement should be similar to that seen during leg movement. Our main experimental hypotheses were that the amplitude of H-reflexes in the forearm muscles would be modulated during arm movement (i.e., phase-dependent) and would be inhibited during cycling compared with static contraction (i.e., task-dependent). Furthermore, to determine the locus of any modulation, we tested the effect that active and passive movement of the ipsilateral (relative to stimulated arm) and contralateral arm had on H-reflex amplitude. Subjects performed rhythmic arm cycling on a custom-made hydraulic ergometer in which the two arms could be constrained to move together (180 degrees out of phase) or could rotate independently. Position of the stimulated limb in the movement cycle is described with respect to the clock face. H-reflexes were evoked at 12, 3, 6, and 9 o'clock positions during static contraction as well as during rhythmic arm movements. Reflex amplitudes were compared between tasks at equal M wave amplitudes and similar levels of electromyographic (EMG) activity in the target muscle. Surface EMG recordings were obtained bilaterally from flexor carpi radialis as well as from other muscles controlling the wrist, elbow, and shoulder. Compared with reflexes evoked during static contractions, movement of the stimulated limb attenuated H-reflexes by 50.8% (P < 0.005), 65.3% (P < 0.001), and 52.6% (P < 0.001) for bilateral, active ipsilateral, and passive ipsilateral movements, respectively. In contrast, movement of the contralateral limb did not significantly alter H-reflex amplitude. H-reflexes were also modulated by limb position (P < 0.005). Thus task- and phase-dependent modulation were observed in the arm as previously demonstrated in the leg. The data support the hypothesis that neural mechanisms regulating reflex pathways in the moving limb are similar in the human upper and lower limbs. However, the inhibition of H-reflex amplitude induced by contralateral leg movement is absent in the arms. This may reflect the greater extent to which the arms can be used independently.     

Cutaneous reflexes during rhythmic arm cycling are insensitive to asymmetrical changes in crank length

The neural control of a movement depends upon the motor task performed. To further understand the neural regulation of different variations of the same type of movement, we created three dissimilar bilateral rhythmic arm cycling tasks by unilaterally manipulating crank length (CL). Modulation in the amplitude and sign of cutaneous reflexes was used as an index of neural control. Neurologically intact subjects performed three bilateral cycling trials at ∼1 Hz with the ipsilateral crank arm at one of three different lengths. Cutaneous reflexes were evoked during each trial with trains (5 × 1.0 ms pulses at 300 Hz) of electrical stimulation delivered to the superficial radial nerve at the ipsilateral wrist. EMG recordings were made bilaterally from muscles acting at the shoulder, elbow, and wrist. Analysis was conducted after phase-averaging contingent upon the timing of stimulation in the movement cycle. CL variation created an asymmetrical cycling pattern and produced significant changes in the range of motion at the ipsilateral shoulder and elbow. Background EMG amplitude in muscles of the contralateral arm generally increased significantly as CL decreased. Therefore at a given phase in the movement cycle, the background EMG was different between the three cycling trials. In contrast, cutaneous reflex amplitudes in muscles of both arms were similar at each phase of the movement cycle between the different CLs trials at both early and middle latencies. This was particularly evident in muscles ipsilateral to nerve stimulation. We suggest that variations of arm cycling that primarily yield significant changes in the amplitude of muscle activity do not require significant task-specific change in neural control.     

Short- and long-latency reflex responses during different motor tasks in elbow flexor muscles

 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. Received: 24 April 1996 / Accepted: 24 January 1997     

Flexion reflex modulation during stepping in human spinal cord injury

The flexion reflex modulation pattern was investigated in nine people with a chronic spinal cord injury during stepping using body weight support on a treadmill and manual assistance by therapists. Body weight support was provided by an upper body harness and was adjusted for each subject to promote the best stepping pattern with the least manual assistance required by the therapists. The flexion reflex was elicited by sural nerve stimulation with a 30 ms pulse train at 1.2–2 times the tibialis anterior reflex threshold. During stepping, stimuli were randomly dispersed across the gait cycle which was divided into 16 equal bins. A long latency (>110 ms) flexion reflex was present in all subjects, while a short (>30 ms) and a medium latency (>70 ms) flexion reflex were present only in three subjects. For each response, the non-stimulated EMG was subtracted from the stimulated EMG at identical time windows and bins, normalized to the maximal corresponding EMG, and significant differences were established with a Wilcoxon rank-sum test. The long latency flexion reflex was facilitated at late stance and during the swing-to-stance transition phase. A reflex depression was present from heel strike until mid-stance and during the swing-to-stance transition phase. The short and medium latency flexion reflexes were depressed during mid-stance followed by facilitation during the stance-to-swing transition phase. Regardless of the latency, facilitatory flexion responses during the swing phase coincided with decreased activity of ipsilateral ankle extensors. The flexion reflex was modulated in a phase dependent manner, a behavior that was absent for the soleus H-reflex in most of these patients (Knikou et al. in Exp Brain Res 193:397–407, 2009). We propose that training should selectively target spinal reflex circuits in which extensor muscles and reflexes are involved in order to maximize sensorimotor recovery in these patients.