Modulation of flexion reflex induced by hip angle changes in human spinal cord injury

The flexion reflex can be elicited via stimulation of skin, muscle, and high-threshold afferents inducing a generalized flexion of the limb. In spinalized animal models this reflex is quite prominent and is strongly modulated by actions of hip proprioceptors. However, analogous actions on the flexion reflex in spinal cord injured (SCI) humans have not yet been examined. In this study, we investigated the effects of imposed static hip angle changes on the flexion reflex in ten motor incomplete SCI subjects when input from plantar cutaneous mechanoreceptors was also present. Flexion reflexes were elicited by low-intensity stimulation of the sural nerve at the lateral malleolus, and were recorded from the ipsilateral tibialis anterior (TA) muscle. Plantar skin stimulation was delivered through two surface electrodes placed on the metatarsals, and was initiated at different delays ranging from 3 to 90 ms. We found that non-noxious sural nerve stimulation induced two types of flexion reflexes in the TA muscle, an early, and a late response. The first was observed only in three subjects and even in these subjects, it appeared irregularly. In contrast, the second (late) flexion reflex was present uniformly in all ten subjects and was significantly modulated during hip angle changes. Flexion reflexes recorded with hip positioned at different angles were compared to the associated control reflexes recorded with hip flexed at 10°. Hip flexion (30°, 40°) depressed the late flexion reflex, while no significant effects were observed with the hip set in neutral angle (0°). Strong facilitatory effects on the late flexion reflex were observed with the hip extended to 10°. Moreover, the effects of plantar skin stimulation on the flexion reflex were also found to depend on the hip angle. The results suggest that hip proprioceptors and plantar cutaneous mechanoreceptors strongly modulate flexion reflex pathways in chronic human SCI, verifying that this type of sensory afferent feedback interact with spinal interneuronal circuits that have been considered as forerunners of stepping and locomotion. The sensory consequences of this afferent input should be considered in rehabilitation programs aimed to restore movement and sensorimotor function in these patients.     

Control of grip force during restraint of an object held between finger and thumb: responses of cutaneous afferents from the digits

Unexpected pulling and pushing loads exerted by an object held with a precision grip evoke automatic and graded increases in the grip force (normal to the grip surfaces) that prevent escape of the object; unloading elicits a decrease in grip force. Anesthesia of the digital nerves has shown that these grip reactions depend on sensory signals from the digits. In the present study we assessed the capacity of tactile afferents from the digits to trigger and scale the evoked grip responses. Using tungsten microelectrodes inserted percutaneously into the median nerve of awake human subjects, unitary recordings were made from ten FA I and 13 FA II rapidly adapting afferents, and 12 SA I and 18 SA II slowly adapting afferents. While the subject held a manipulandum between a finger and the thumb, tangential load forces were applied to the receptor-bearing digit (index, middle, or ring finger or thumb) as trapezoidal load-force profiles with a plateau amplitude of 0.5 – 2.0 N and rates of loading and unloading at 2 – 8 N/s, or as step-loads of 0.5 N delivered at 32 N/s. Such load trials were delivered in both the distal (pulling) and proximal (pushing) direction. FA I afferents responded consistently to the load forces, being recruited during the loading and unloading phases. During the loading ramp the ensemble discharge of the FA I afferents reflected the first time-derivative of the load force (i.e., the load-force rate). These afferents were relatively insensitive to the subject's grip force responses. However, high static finger forces appeared to suppress excitation of these afferents during the unloading phase. The FA II afferents were largely insensitive to the load trials: only with the step-loads did some afferents respond. Both classes of SA afferents were sensitive to load force and grip force, and discharge rates were graded by the rate of loading. The firing of the SA I afferents appeared to be relatively more influenced by the subject's grip-force response than the discharge of the SA II afferents, which were more influenced by the load-force stimulus. The direction in which the tangential load force was applied to the skin influenced the firing of most afferents and in particular the SA II afferents. Individual afferents within each class (except for the FA IIs) responded to the loading ramp before the onset of the subject's grip response and may thus be responsible for initiating the automatic increase in grip force. However, nearly half of the FA I afferents recruited by the load trials responded to the loading phase early enough to trigger the subject's gripforce response, whereas only ca. one-fifth of the SA Is and SA IIs did so. These observations, together with the high density of FA I receptors in the digits, might place the FA I afferents in a unique position to convey the information required to initiate and scale the reactive gripforce responses to the imposed load forces.     

Modulation of stretch reflex by anticipation of the stimulus through visual information

Summary A stretch stimulus was applied to the right elbow flexor by the free gravitational fall of a weight. The subject was instructed to flex the elbow joint to lift the weight quickly (resist task) or to extend the elbow joint to overtake the falling weight (assist task) as the voluntary reaction to the stretch stimulus. In the resist task, when the subject could predict the time of stretch stimulus onset by watching through the mirror the behavior of the experimenter going to give the stretch stimulus, the integrated EMG (I-EMG) of M 2 (50–80 ms after the stimulus onset) and M3 (80–100 ms) components of stretch reflex were significantly enhanced compared with when the experimenter's movements were invisible to the subject, while no differences were observed in the background discharge before the stretch stimulus onset (BGA) and the short latency reflex (M1). In the assist task, the M2 and M3 were depressed when the visual information about the stimulus presentation was available, while the BGA and M1 were unchanged. From these results, it is suggested that (1) the preparatory set should be classified into two categories, i.e., task-related topographical set and timing-related chronographic set, and (2) visual information about the process of stimulus presentation can modulate the reflex activity of stretched muscle allowing the required task to be executed efficiently by accurately anticipating the stretch stimulus onset.     

Enhanced stretch reflex excitability of the soleus muscle in experienced swimmers

The purpose of this study was to investigate effects of long-term participation to swimming on adaptations of spinal reflex excitability. To this end, mechanically induced stretch reflex (SR) and electrically induced Hoffmann (H-) reflex of the soleus muscle were investigated between swimmers with experience of more than 10 years and non-trained individuals while sitting at rest. The amplitude and the gain (stretch velocity vs. amplitude of the reflex response) of the SR were significantly greater in the swimming group than in the non-trained control group. Similarly, the responses of the H-reflex were also significantly greater in the swimming group than in the non-trained control group. Results of this study demonstrated that the spinal reflex excitability in experienced swimmers was far more enhanced than in non-trained individuals.     

Do muscle afferents contribute to the cervical response evoked by electrical stimulation of the median nerve in man?

Summary The possible contribution of low threshold muscle afferents to the postsynaptic component (N₁₃) of the cervical response evoked by electrical stimulation of the median nerve (MN) was investigated in normal subjects. Electroneurographic (ENG) and electromyographic (EMG) correlates of the reflex motoneuronal discharge (RMND) were recorded simultaneously. A. No reflex activity could be elicited by stimulation of the MN at the wrist, at least in the resting subjects, while well developed ENG (P₂ efferent volley) and EMG (H reflex) monosynaptic responses occurred following stimulation of the MN at the elbow at suitable strengths. In neither case could a surface correlate of interneuronal activity evoked by muscle afferents be demonstrated. B. Recruitment curves showed that at stimulus intensities above maximal for the H reflex both P₂ and H responses started to decrease until they completely disappeared, while N₁₃ showed further enhancement. C. Subthreshold conditioning stimulation of the MN enhanced both P₂ and H responses, while vibratory muscle stimuli provoked a clearcut suppression of these two responses. In contrast, N₁₃ was completely unaffected by either manuvre. D. No cervical evoked activity could be detected following tendon tapping of the anterior forearm muscles in spite of the appearance of well developed cortical responses and the ENG and EMG correlates of the T reflex. E. Conditioning volleys elicited by tendon taps of the anterior forearm muscles suppressed both P₂ and H responses following stimulation of the MN at the elbow without affecting the related N₁₃ component. F. Conditioning supramaximal stimulation of the MN at the wrist suppressed the N₁₃ component of the cervical response evoked by stimulation of the MN at the elbow without affecting the related reflex responses. No component chronologically related to the RMND could be recorded at the posterior neck region during suppression of N₁₃, thus ruling out the possibility that failure to detect the RMND (as well as its interneuronal concomitants) with cervical electrodes is due to a masking effect of the N₁₃ component. G. Conditioning tendon taps of anterior forearm muscles provoked a clearcut reduction of the primary cortical response to finger stimulation without affecting the postsynaptic component of the related cervical response. It is concluded that neither segmental (motoneuronal or interneuronal in origin) nor ascending postsynaptic impulses generated in the spinal cord by stimulation of low threshold muscle afferents contribute to N₁₃, the latter being probably due to activation of both short and long axoned spinal neurons by cutaneous afferents.     

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.     

The suppression of the long-latency stretch reflex in the human tibialis anterior muscle by transcranial magnetic stimulation

The purpose of this study was to investigate the transcortical nature of the long-latency stretch reflex (M3) in the human tibialis anterior muscle. This was achieved by applying a single pulse of subthreshold (90% motor threshold) transcortical magnetic stimulation (subTMS) at the site of the motor cortex. Such a stimulus is able to activate intracortical inhibitory circuits and thereby depress motor cortical output. We hypothesized that it would also suppress a transcortical reflex loop. The stretch reflex was elicited using a pedal attached to an electric motor. SubTMS was applied at several intervals prior to M3. Recordings were repeated 20–40 times. The reflex components were quantified using 20-ms windows in the averaged rectified electromyogram (EMG). SubTMS evoked significantly larger depression of M3 than of the background EMG in the same time frame when applied 55–85 ms prior to M3 (P<0.05, n=10). Furthermore, the effect on M3 was significantly larger than the effect on the spinal M2 (P<0.01, n=7). Our results provide evidence that the long-latency stretch reflex in the tibialis anterior muscle is at least partly transcortical.     

Adaptation of the short latency component of the stretch reflex plays only a minor role in compensating for muscle fatigue induced by spontaneous hopping in humans

The influence of fatigue on the stretch reflex evoked in ankle extensor muscles by hopping was investigated in six healthy men. The men hopped on a force platform, at spontaneous frequency and amplitude, until they were unable to maintain the initial frequency or amplitude of the jumps. This task was done with the knees flexing normally during ground contact or under instructions to straighten the knees. Surface electromyograms (EMG) of soleus (SO), gastrocnemius medialis (GM), and tibialis anterior (TA) muscles were recorded simultaneously with the vertical component of the ground reaction force. Spectrum analysis of the EMG recorded during isometric tests performed immediately before and after the fatiguing hopping task demonstrated the existence of fatigue in SO and GM and often in TA. The stretch reflex was studied during the first and last ten jumps of every hopping series. The long-latency components of the reflex were too variable to be analysed. Whatever the hopping condition, latency and amplitude of the short latency component were not significantly modified by fatigue. Fatigue enhanced the occurrence of this reflex component in SO only. These data suggest that in fatiguing submaximal hopping, the neuromuscular system does not fundamentally change its stiffness regulation before the endurance time has been reached. Accepted: 1 September 2000