Acquisition of a simple motor skill: task-dependent adaptation and long-term changes in the human soleus stretch reflex.

Changing the H reflex through operant conditioning leads to CNS multisite plasticity and can affect previously learned skills. To further understand the mechanisms of this plasticity, we operantly conditioned the initial component (M1) of the soleus stretch reflex. Unlike the H reflex, the stretch reflex is affected by fusimotor control, comprises several bursts of activity resulting from temporally dispersed afferent inputs, and may activate spinal motoneurons via several different spinal and supraspinal pathways. Neurologically normal participants completed 6 baseline sessions and 24 operant conditioning sessions in which they were encouraged to increase (M1up) or decrease (M1down) M1 size. Five of eight M1up participants significantly increased M1; the final M1 size of those five participants was 143 ± 15% (mean ± SE) of the baseline value. All eight M1down participants significantly decreased M1; their final M1 size was 62 ± 6% of baseline. Similar to the previous H-reflex conditioning studies, conditioned reflex change consisted of within-session task-dependent adaptation and across-session long-term change. Task-dependent adaptation was evident in conditioning session 1 with M1up and by session 4 with M1down. Long-term change was evident by session 10 with M1up and by session 16 with M1down. Task-dependent adaptation was greater with M1up than with the previous H-reflex upconditioning. This may reflect adaptive changes in muscle spindle sensitivity, which affects the stretch reflex but not the H reflex. Because the stretch reflex is related to motor function more directly than the H reflex, M1 conditioning may provide a valuable tool for exploring the functional impact of reflex conditioning and its potential therapeutic applications. NEW & NOTEWORTHY Since the activity of stretch reflex pathways contributes to locomotion, changing it through training may improve locomotor rehabilitation in people with CNS disorders. Here we show for the first time that people can change the size of the soleus spinal stretch reflex through operant conditioning. Conditioned stretch reflex change is the sum of task-dependent adaptation and long-term change, consistent with H-reflex conditioning yet different from it in the composition and amount of the two components.

Modulation of soleus stretch reflexes during walking in people with chronic incomplete spinal cord injury.

In people with spasticity due to chronic incomplete spinal cord injury (SCI), it has been presumed that the abnormal stretch reflex activity impairs gait. However, locomotor stretch reflexes across all phases of walking have not been investigated in people with SCI. Thus, to understand modulation of stretch reflex excitability during spastic gait, we investigated soleus stretch reflexes across the entire gait cycle in nine neurologically normal participants and nine participants with spasticity due to chronic incomplete SCI (2.5-11 year post-injury). While the participant walked on the treadmill at his/her preferred speed, unexpected ankle dorsiflexion perturbations (6° at 250°/s) were imposed every 4-6 steps. The soleus H-reflex was also examined. In participants without SCI, spinal short-latency "M1", spinal medium latency "M2", and long-latency "M3" were clearly modulated throughout the step cycle; the responses were largest in the mid-stance and almost completely suppressed during the stance-swing transition and swing phases. In participants with SCI, M1 and M2 were abnormally large in the mid-late-swing phase, while M3 modulation was similar to that in participants without SCI. The H-reflex was also large in the mid-late-swing phase. Elicitation of H-reflex and stretch reflexes in the late swing often triggered clonus and affected the soleus activity in the following stance. In individuals without SCI, moderate positive correlation was found between H-reflex and stretch reflex sizes across the step cycle, whereas in participants with SCI, such correlation was weak to non-existing, suggesting that H-reflex investigation would not substitute for stretch reflex investigation in individuals after SCI.

Afferent contribution to locomotor muscle activity during unconstrained overground human walking: an analysis of triceps surae muscle fascicles.

Plantar flexor series elasticity can be used to dissociate muscle-fascicle and muscle-tendon behavior and thus afferent feedback during human walking. We used electromyography (EMG) and high-speed ultrasonography concomitantly to monitor muscle activity and muscle fascicle behavior in 19 healthy volunteers as they walked across a platform. On random trials, the platform was dropped (8 cm, 0.9 g acceleration) or held at a small inclination (up to +/-3 degrees in the parasagittal plane) with respect to level ground. Dropping the platform in the mid and late phases of stance produced a depression in the soleus muscle activity with an onset latency of about 50 ms. The reduction in ground reaction force also unloaded the plantar flexor muscles. The soleus muscle fascicles shortened with a minimum delay of 14 ms. Small variations in platform inclination produced significant changes in triceps surae muscle activity; EMG increased when stepping on an inclined surface and decreased when stepping on a declined surface. This sensory modulation of the locomotor output was concomitant with changes in triceps surae muscle fascicle and gastrocnemius tendon length. Assuming that afferent activity correlates to these mechanical changes, our results indicate that within-step sensory feedback from the plantar flexor muscles automatically adjusts muscle activity to compensate for small ground irregularities. The delayed onset of muscle fascicle movement after dropping the platform indicates that at least the initial part of the soleus depression is more likely mediated by a decrease in force feedback than length-sensitive feedback, indicating that force feedback contributes to the locomotor activity in human walking.

Neuromodulation of lower limb monoparesis: functional electrical therapy of walking.

After Cerebro-Vascular Accident (CVA), restoration of normal function, such as locomotion, depends on reorganization of existing central nervous system (CNS) circuitry. This capacity for reorganization, generally referred to as plasticity, is thought to underlie many instances of functional recovery after injury as well as learning and memory in the undamaged CNS. Both the reorganization of the supraspinal and spinal circuitry are highly important for the recovery of walking. The neural mechanisms responsible for learning and adapting processes are thought to involve changes both in the efficacy of synaptic function and the pattern of synaptic connections within neural circuits. In the uninjured CNS, these changes occur as a result of alterations in the amount of neural activity within circuits and are, therefore, termed activity-dependent. In this chapter, we will present several therapies of walking that provide effective input for the training of the existing CNS circuitry; thereby, contribute to long term recovery of sensory-motor functions. The focus of this chapter is Functional Electrical Therapy (FET) of walking, that is, the multi-channel electrical stimulation of sensory-motor systems that lead to more normal stance and swing of the paretic leg during the walking exercise.

Symmetry of post-movement beta-ERS and motor recovery from stroke: a low-resolution EEG pilot study.

The inter-hemispheric symmetry of electroencephalographic (EEG) post-movement beta-event-related synchronization (PMBS) after movements on a drawing board was studied in eight acute stroke subjects with mild hemiparesis and eight normal subjects. A follow-up testing was conducted 3 months after the initial recordings with a twofold purpose: (1) to validate the reproducibility of the experimental protocol in normal subjects; and (2) to study changes of inter-hemispheric PMBS-symmetry as a response to recovery of motor function. PMBS values were calculated and their topographic distributions illustrated at various time instances following movement offset. Significant PMBS patterns were present in all normal subjects, with only minor differences within consecutive recordings. The side of hemiparesis in acute stroke subjects could be distinguished (P = 0.04) on the basis of the signed symmetry index, a quantitative measure of lateralization. The follow-up testing on three recovered stroke subjects revealed a trend of changes in the lateralization towards the contralateral side of movement, an indication that the cortical organization of movement following recovery turned out as reported for normal subjects. Further clinical investigations need to be carried out to evaluate the relationship between recovery and PMBS symmetry on a large number of subjects, using the method presented here.

Evidence for transcortical reflex pathways in the lower limb of man.

The existence of transcortical reflex pathways in the control of distal arm and hand muscles in man is now widely accepted. Much more controversy exists regarding a possible contribution of such reflexes to the control of leg muscles. It is often assumed that transcortical reflex pathways play no, or only a minor, role in the control of leg muscles. Transcortical reflex pathways according to this view are reserved for the control of the distal upper limb and are seen in close relation to the evolution of the primate hand. Here we review data, which provide evidence that transcortical reflexes do exist for lower limb muscles and may play a significant role in the control of at least some of these muscles. This evidence is based on animal research, recent experiments combining transcranial magnetic stimulation with peripheral electrical and mechanical stimulation in healthy subjects and neurological patients. We propose that afferent activity from muscle and skin may play a role in the regulation of bipedal gait through transcortical pathways.

Propulsive activity induced by sequential electrical stimulation in the descending colon of the pig.

UNLABELLED: This work was performed to study electrically induced contractions in the descending colon of pigs. Contractions were monitored using impedance planimetry and manometry. The luminal pressure, cross-sectional area (CSA), latency and velocity of CSA decrease were compared when using 3 ms, 9, 12, 15 or 30 mA pulses at 10 Hz for 10 s, and 15 mA, 0.03, 0.3 or 3 ms pulses at 10 Hz for 10 s. Stimulation was performed prior and after the application of N(G)-nitro-L-arginine methyl ester (L-NAME) and atropine. In the untreated colon, contraction was always of an 'off' type. A current increase from 9 to 30 mA increased the pressure. An increase of pulse duration from 0.03 to 3 ms shortened the latency, accelerated contraction and increased pressure. By sequential stimulation, contractions were coordinated to propel semi-fluid and solid luminal contents. L-NAME increased the magnitude of CSA decrease. Atropine induced inhibitory effects on contractions elicited by 3 ms pulses and abolished contractions induced by 0.03 and 0.3 ms pulses. IN CONCLUSION: (i) electrical stimulation evokes'off' colon contractions, which can be coordinated to result in propulsion; (ii) the best combination for current and pulse duration to induce propulsive contractions is 15 mA and 3 ms; (iii) nitrergic and cholinergic pathways mediate responses to electrical stimulation.

Muscle, reflex and central components in the control of the ankle joint in healthy and spastic man.

In understanding the control of the ankle joint during different motor tasks, we have to investigate at least three components, namely the influence of i) the passive and intrinsic properties of the intact and active muscle system around the joint (termed the non-reflex component), ii) the mechanical importance of the stretch reflex in the stretched and unloaded muscles, and iii) the supraspinal control of the stretch reflex. This thesis is dealing with the importance of the three components in healthy and spastic persons during sitting, standing, and walking. The results are based on stretch reflex and H-reflex measurements from the ankle extensor muscles. During stretch reflex experiments the foot was mounted to a platform (portable during walking) from which the ankle joint torque and the position were measured. To elicit a stretch reflex, the ankle joint was rotated by a strong motor connected to the platform. The mechanical importance of the stretch reflex was investigated by measuring the changes in joint torque. Electrically, the stretch reflex was recorded as the compound muscle action potential through bipolar surface EMG electrodes placed over the soleus muscle. During H-reflex experiments, the tibial nerve was stimulated at the popliteal fossa and the H-reflex recorded over the soleus muscle as during stretch reflex experiments. To investigate how the contractile properties of a muscle in humans depend on the history of activation, we investigated the intrinsic stiffness of the ankle extensors in healthy subjects. At matched background contraction in sitting subjects, a prolonged contraction increased the intrinsic muscle stiffness by 49%. Muscle yielding has been considered especially important for understanding the reflex compensation. We found a general lack of muscle yield and a mechanically important non-reflex stiffness of the ankle extensors showing that non-reflex stiffness is a prominent factor in normal movements of the ankle joint. In both healthy and spastic persons, we found a mechanically strong stretch reflex in the isometric, contracted muscles during sitting. This posed the question; how is the reflex regulated during more functional motor tasks. This was dealt with by studying the H-reflex during isometric ramp contractions and during walking in healthy and spastic persons. In the healthy subjects the H-reflex was modulated in consistency with a task dependent control. In the spastic patients the H-reflex lacked a task dependent modulation. In consistency with earlier findings it was suggested that the decreased modulation could have been caused by decreased control of the pre-synaptic inhibition of the Ia terminals or a change in recruitment gain. To test if the stretch reflex behaved as the H-reflex, the short latency stretch reflex was investigated during walking. Here we found that the stretch reflex was strongly modulated during a step in healthy subjects as seen for the H-reflex, but when comparing the stretch reflex at matched excitation levels (same background EMGs) during standing and walking, no task-specific reflex modulation was found except the one relating to the excitation level. Therefore, the results emphasise that at least during walking and standing it is not always possible to draw conclusions about the stretch reflex based on observations of the H-reflex. When investigating the modulation of the short latency stretch reflex during walking in spastic patients, we found that the stretch reflex modulation was impaired in spastic patients at least to the extent demonstrated earlier for the H-reflex. The passive stiffness of the ankle joint was at the same time increased in the patients. At matched ankle extensor contraction levels, stretch responses were compared before and after reversible block of the common peroneal nerve and during an attempted, voluntary, fictive dorsiflexion after common peroneal nerve block. (ABSTRACT TRUNCATED)

Experimental muscle pain increases the human stretch reflex.

In this study we investigated the effect of human experimental muscle pain on H- and stretch reflexes as indicators of changes in muscle spindle sensitivity. Fourteen healthy, male volunteers participated in the study. Muscle pain was produced by infusion of 5% hypertonic saline over a period of 10-15 min in m. soleus and in m. tibialis anterior. Reflexes were elicited in the relaxed and active soleus muscle (10-15 Nm ankle torque) before, during and after muscle pain. Control measurements were made with infusions of 0.9% isotonic saline. Surface electromyograms (EMG) were measured from the soleus muscle, and torque was measured from the ankle joint. With pain in the soleus muscle the mechanical stretch reflex response (ankle torque) increased significantly (P = 0.0007) as compared to before pain. With pain in the tibialis anterior muscle both the mechanical and EMG responses increased significantly (P = 0.001; P = 0.0003) as compared to before pain. The H-reflex showed no significant changes during the infusions in either muscles. This study has demonstrated a muscle pain-related increase in the amplitude of the stretch reflex without a corresponding increase in the H-reflex amplitude. One explanation could be an increased dynamic sensitivity of the muscle spindles during muscle pain caused by an increased firing rate in the dynamic gamma-motoneurones. However, the data could not support the vicious cycle model because the excitability of the alpha-motoneurone pool was unchanged.

Signals from skin mechanoreceptors used in control of a hand grasp neuroprosthesis.

We used signals from tactile mechanoreceptors in the skin of the index finger, recorded with an implanted cuff electrode, to automatically control grasp force in a hand grasp neuroprosthesis. Phasic events in the recorded nerve signal, related to mechanical events on the skin, were used to adjust electrical stimulation of hand muscles without any prior knowledge about muscle strength and properties of a held object. A simulated eating task was used to evaluate the hand grasp neuroprosthesis. When using the neuroprosthesis with feedback from natural sensors, the average grasp force could be reduced in comparison to not using feedback. Reducing grasp force is considered a major factor to decrease muscle fatigue, allowing a prolonged use of the hand grasp neuroprosthesis.

Simultaneous 31P-NMR spectroscopy and EMG in exercising and recovering human skeletal muscle: a correlation study.

A large number of studies have shown amplitude and spectral changes of the electromyogram during exercise, leading to several theories of how these changes might be related to the underlying metabolic changes. The amplitude and spectral changes are generally interpreted as changes in motor unit recruitment and a reduction of the muscle fiber conduction velocity due to proton or lactate accumulation. This study focuses on the causality of spectral changes of the surface electromyogram and proton or lactate accumulation and how the changes in motor unit recruitment are related to the metabolic status of the muscle. Simultaneous 31P-nuclear magnetic resonance spectroscopy and surface electromyography were performed during sustained static exercise and recovery in healthy volunteers and a patient with McArdle's disease. A clear dissociation between the median power frequency of the surface electromyogram and pH was seen in the healthy volunteers during recovery and during exercise in the patient with McArdle's disease. The results indicate that proton or lactate accumulation is not primarily responsible for the spectral changes of the surface electromyogram as previously suggested. The motor unit recruitment (as judged by the root mean square of the surface electromyogram) increased hyperbolically during the submaximal static exercise, with decreasing phosphocreatine-to-P(i) ratio reaching maximum at 0.6 (exhaustion), and seems to constitute a consistent metabolic limit to the exercise. The increased myoelectrical activity seen after exercise is not caused by an incomplete recovery of phosphorous metabolism, pH, or lactate but could probably be an impairment of the excitation-contraction coupling.

A computer-controlled system to perturb the ankle joint of freely standing cats trained to maintain a given force.

A computer-based system was developed to (1) train freely standing cats to match various target forces with the left hindlimb, (2) perturb the left ankle joint when the cat was maintaining a desired force and (3) compare reflex responses before and after decerebration. Cats quickly learned to stand unaided on 4 pedestals. During a training session, a range of target force windows was presented to the cat. A successful trial consisted of maintaining the force applied on the left rear pedestal within the target window for a preset time period. To assist the cat, a light was turned on whenever the force was within the target window. A food pellet reward was delivered by the computer after each successful trial. To test reflex responses, the position of the left hindlimb could be briefly perturbed by activating a servo-controlled printed motor configured to rotate the pedestal about the axis of the ankle joint. Perturbations that either flexed or extended the ankle joint were presented pseudo-randomly by the computer. This approach has been used to quantify the magnitude of muscle afferent volleys and the reflex EMG in ankle extensor muscles of normal and decerebrated cats, in response to similar mechanical perturbations. It has also been used to study dynamic features in the electroneurogram recorded from a cutaneous nerve by implanted nerve cuff electrodes, and the correlations among the electroneurogram, the vertical contact force applied on the pedestal and the force recorded from muscle tendons by implanted transducers. This approach may have general applications in the study of postural control, including the study of the discharge patterns of individual motor, sensory or spinal cord neurons in freely standing cats.

The H-reflex in the passive human soleus muscle is modulated faster than predicted from post-activation depression.

The purpose of the present study was to investigate the influence of afferent activity (mainly homonymous Ia-afferent activity) on the modulation (post-activation depression) of the soleus H-reflex during isolated and passive sinusoidal ankle joint rotations at a speed and amplitude comparable to slow walking. The H-reflex modulation was measured in the relaxed soleus muscle on human subjects during different imposed patterns of 20 degrees haversine ankle joint rotations (0.5-0.6 Hz) while they were sitting comfortably in a chair. Eighteen healthy males and four male patients with clinically complete spinal cord lesion above the soleus motoneuron pool participated in the study. During a single dorsi-plantar flexion rotation the H-reflex was depressed to 27+/-7% (mean+/-S.E.M.) of the initial level within 600 ms. The course of this depression was reversed when the dorsi-flexion velocity started to decrease. At the end of the dorsi-flexion movement the depression was already relieved to a level of 73+/-6% of the initial level. The H-reflex returned more slowly to the initial level within 2 s after the end of the movement cycle. During two consecutive ankle joint rotations and continuous ankle joint rotations both at 0.5 Hz the H-reflex was modulated but also generally depressed while the movement was imposed. The reflex only returned to the reference level after the movements were stopped. These observations indicate the action of a fast and a slow mechanism in the post-activation depression of the soleus H-reflex. The H-reflex modulations observed in the spinal cord injured patients were comparable to the reflex modulations observed in the healthy subjects, except the depressions were smaller. This suggests that a major part of the amplitude of the H-reflex modulation observed in healthy subjects was caused by peripheral and spinal influences. The fast 500 ms recovery of the H-reflex had a time course comparable to presynaptic inhibition. The slow 2 s recovery after the end of a given imposed movement may be explained by a change in the probability of transmitter release from the homonymous soleus Ia-afferent synaptic terminals after repeated activations.

Impaired stretch reflex and joint torque modulation during spastic gait in multiple sclerosis patients.

The modulation of the short latency stretch reflex of the soleus muscle during walking was investigated in seven spastic multiple sclerosis (MS) patients and nine healthy control subjects. Ankle joint stretches were applied by a system which can rotate that ankle joint in any phase of the step cycle during treadmill walking. The torque related to the muscle fibres contracting prior to the stretch and the passive tissues around the ankle joint were measured as the "non-reflex torque". At the same time the short latency stretch reflex-mediated EMG response was measured. The findings show that the stretch reflex modulation was impaired in spastic patients during walking. The stretch reflex modulation was quantified by a modulation index of an average 50% (range -5 to 100%) in the patients and 93% (78-100%) in the control subjects (P < 0.05). The passive stiffness of the ankle joint was at the same time increased in the patients (P < 0.05). It is proposed that the impaired modulation of the stretch reflex along with increased ankle joint stiffness contribute to the impaired walking ability in spastic MS patients.

Soleus long-latency stretch reflexes during walking in healthy and spastic humans.

The present study was carried out to investigate the long-latency soleus stretch reflexes M2 (peak latency of approximately 85 ms) and M3 (peak latency of approximately 115 ms) during walking in healthy and spastic multiple sclerosis (MS) patients. An 8 degrees stretch was applied to the ankle extensors of the left leg in 8 healthy subjects during normal walking speed and 9 spastic MS patients and 10 age-matched healthy subjects during slow walking. When present in walking healthy subjects, M2 and M3 were modulated in a similar way and with the same amplitudes as previously described for the short latency soleus stretch reflex (M1). The spastic patients' soleus M1 was significantly less modulated during walking. The patients' M2 long-latency response was modulated in the same way as the age-matched healthy subjects. All patients' M3 responses were absent or much suppressed during walking. The origin and functional importance of the short- and long-latency stretch reflexes in healthy and spastic persons are discussed in relation to the above findings and the behaviour of the stretch reflexes during matched isometric contractions. M3 is argued to be part of a transcortical reflex in healthy subjects.

The influence of experimental muscle pain on the human soleus stretch reflex during sitting and walking.

OBJECTIVES: The stretch reflex is functionally important during human locomotion. Muscle pain has been found to increase the stretch reflex amplitude during sitting, possibly due to an altered fusimotor drive. To further study the importance of altered fusimotor activity due to muscle pain we investigated the combined effect of muscle pain and motor task on the soleus stretch reflex. METHODS: Stretch reflexes were elicited before, during and after experimentally induced muscle pain in soleus (i.m. infusion of 6% saline) in 3 experiments: (1) in the relaxed soleus muscle and before, during and after an isometric ramp contraction (500 ms, 0-10 Nm), (2) at 3 different time periods during walking, and (3) at matched pain intensity and soleus activity during sitting and walking. RESULTS: Infusion of hypertonic saline into the soleus muscle caused a significant facilitated stretch reflex in the relaxed muscle (P<0.01), but not during walking or during sitting and walking at matched soleus EMG and matched pain levels. The infusion of isotonic saline (non-painful) did not cause any changes (P = 0.75). CONCLUSIONS: The main findings of the present study were that experimental muscle pain facilitated the stretch reflex during pain in the relaxed muscle, but caused no changes in stretch reflex amplitude during sitting and walking at higher "functional" background EMG levels.

Non-noxious stimulation of the glenohumeral joint capsule elicits strong inhibition of active shoulder muscles in conscious human subjects.

Information about the function of joint afferents in humans is scarce. Therefore, we investigated the influence of non-noxious electrical stimulation of the glenohumeral joint capsule on the activity of shoulder muscles in conscious human subjects. Stimulation electrodes were inserted in the joint capsule during arthroscopy. Only when the anterior inferior part of the capsule including the glenohumeral ligament was stimulated a strong and rather long latent general inhibition in voluntarily activated shoulder muscles was observed (average latency and duration: 33 ms and 76 ms, respectively). The inhibition was followed by an excitation and a few but inconsistent oscillations in the muscle activity. The results suggests a potent proprioceptive feedback pathway for the motor control of the human shoulder joint. However, further investigations are needed to clarify the involvement of several possible afferent pathways.

Effect of initial joint position on nerve-cuff recordings of muscle afferents in rabbits.

The objective was to characterize nerve-cuff recordings of muscle afferents to joint rotation over a large part of the physiological joint range. This information is needed to develop control strategies for functional electrical stimulation (FES) systems using muscle afferent signals for sensory feedback. Five acute rabbit experiments were performed. Tripolar cuff electrodes were implanted around the tibial and peroneal divisions of the sciatic nerve in the rabbit's left leg. The electroneurograms (ENG) were recorded during passive ankle rotation, using a ramp-and-hold profile starting at seven different joint positions (excursion = 5 degrees, velocity = 10 degrees/s, initial positions 60 degrees, 70 degrees, 80 degrees, 90 degrees, 100%, 110 , and 120 ). The amplitude of the afferent activity was dependent on the initial joint position. The steady-state sensitivity of both nerve responses increased with increasing joint flexion, whereas the dynamic sensitivity increased initially but then decreased. The results indicate that recordings of the muscle afferents may provide reliable information over only a part of the physiological joint range. Despite this limitation, muscle afferent activity may be useful for motion feedback if the movement to be controlled is within a narrow joint range such as postural sway.

Neural network classification of nerve activity recorded in a mixed nerve.

Whole-nerve cuff electrodes can be used to record electrical nerve activity in peripheral nerves and are suitable for chronic implantation in animals or humans. If the whole nerve innervates multiple target organs or muscles then the recorded activity will be the superposition of the activity of different nerve fibers innervating these organs. In certain cases it is desirable to monitor mixed nerve activity and to determine the origin (modality) of the recorded activity. A method using the autocorrelation function of recorded nerve activity and an artificial neural network was developed to classify the modality of nerve signals. The method works in cases where different end organs are innervated by nerve fibers having different diameter distributions. The electrical activity in the cat S1 sacral spinal root was recorded using a cuff electrode during the activation of cutaneous, bladder, and rectal mechanoreceptors. Using the classification method, 87.5% of nerve signals were correctly classified. This result demonstrates the effectiveness of the neural network classification method to determine the modality of the nerve activity arising from activation of different receptors.

Regulation of wrist stiffness by the stretch reflex.

In restoring the angular position after a displacement, the role of the muscle stretch reflex was investigated by comparing the restored angular torques and angular positions in the wrist under ischaemic and non-ischaemic conditions in normal human subjects. The wrist compliance (COM), defined as the dynamic relation between the angular position and the angular torque of the joint, was calculated to quantify the changes in the restoration of a displacement after abolishing the stretch reflex by ischaemia. The elasticity from the COM-function was found to be single most important factor controlled by the stretch reflex. The elasticity that equals the static stiffness of the system increased by more than 100%, from 0.21 Nm degree-1 with abolished reflex to 0.45 Nm degree-1 with intact reflex. Our results have shown that the stretch reflex assists in the rapid return of the limb to its original position after a mechanical displacement. When the reflex was blocked by ischaemia, the perturbation displaced the limb further away from the initial position.