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Biomedical subjects

H B Morton

Publications and source records attributed to H B Morton.

At least 19 recordsLinked to original sources

Scope of a technique for electrical stimulation of human brain, spinal cord, and muscle.

Brief high-voltage electrical shocks from a special low-output-resistance stimulator, delivered through electrodes on the skin, can excite human muscle directly (not by way of the nerves) and can also excite the motor cortex, the visual cortex, and the spinal cord. Possible applications of the technique include measurement in muscle disorders of the latency relaxation and of the excitability and contractility of muscle without the interposition of nerve fibres or the neuromuscular junction; measurement of conduction velocity in the pyramidal tract; and the detection of neuropathy in the nerves to the external sphincter ani.

Action Potentials

Reliability and efficacy of the long-latency stretch reflex in the human thumb.

1. The amount of positional compensation afforded by the long-latency reflex in the flexor pollicis longus has been investigated in ten normal human subjects. 2. The interphalangeal joint of the thumb was extended by between 2 and 40 degrees at up to 900 deg/s by suddenly increasing the standing force applied to the lever against which the subject was pressing with the pad of the thumb. 3. Electromyographic (e.m.g.) responses at spinal-latency were very small or absent for stretches of this magnitude. The long-latency stretch reflex produced an average positional correction of about 50% for disturbances in the range of 5-25 degrees. The response began to saturate for disturbances of greater than 25 degrees. 4. The e.m.g. response was pulsatile, lasting only some 50 ms, even during continuously increasing disturbances; frequently it terminated despite a remaining positional error. 5. There was a large variation from subject to subject in the average amount of positional correction provided by the stretch reflex. Examination of single responses to the same stretch in individual subjects showed an even greater variation from trial to trial. 6. Variation in the compensation produced by the long-latency stretch reflex from trial to trial could not be explained by the slight variation in size or maximum velocity of the individual stretches.

Adult

The effect of posterior column lesions on servo responses from the human long thumb flexor.

Lesions of the posterior column pathways, in which muscle spindle afferents run towards the brain, are associated with loss of servo responses in the long flexor of the thumb, in the absence of motor weakness and with tendon jerks preserved. This evidence is consistent with the hypothesis that the long-latency stretch reflex (on which servo responses are based) uses a supraspinal, possibly a transcortical, reflex arc.

Adult

The effect of lesions of the sensorimotor cortex and the capsular pathways on servo responses from the human long thumb flexor.

Lesions of the sensorimotor cortex, or of the capsular pathways beneath it, caused (with one exception out of 14 cases) diminution "r loss of the servo responses in the thumb, which are based on the long-latency stretch reflex. When not absent the long-latency stretch reflex tended to be late in onset. When absent it was often replaced by a large early reflex response at spinal latency. In general the results are consistent with the transcortical theory of the long-latency stretch reflex for the thumb, but, in detail, they indicate that the theory will require elaboration.

Adult

The sensory mechanism of servo action in human muscle.

1. Anaesthesia of the thumb suppresses servo action in the long flexor of the thumb during movements of the terminal phalanx. 2. It needs a greater subjective effort to flex an anaesthetized thumb than a normal one. 3. Anaesthesia of other digits is without effect on the thumb flexor. 4. In an anaesthetic thumb without servo responses the changes in force exerted when a mechanical perturbation is applied which are due purely to the passive mechanical properties of the muscle can be measured. Subtraction of these gives the active components of servo action in the normal thumb and thus an estimate of the mechanical gain of the servo. 5. Giving the subject a visual tracking task can partially restore servo action when the thumb is anaesthetic. 6. After some years subjects become resistant to the effect of peripheral anaesthesia. 7. Peripheral anaesthesia has no detectable effect on servo responses in the long flexor of the great toe, in infraspinatus or in pectoralis major. Servo action in these muscles is presumably based predominantly on muscle receptors. 8. Tendon jerks are not apparently influenced by peripheral anaesthesia. 9. The Discussion considers the possibility that, for the thumb, muscle afferents co-operate with a somatic and a visual input in a servo loop via the cerebral cortex.

Afferent Pathways

Servo action in the human thumb.

1. The servo-like properties of muscle in healthy human subjects have been studied by interfering unexpectedly with flexion movements of the top joint of the thumb. This movement is carried out by the flexor pollicis longus muscle only. 2. The movements were standardized in rate by giving the subject a tracking task. They started off against a constant torque load offered by an electric motor. 3. In some movements the load remained constant, but in others, in mid-course, perturbations were introduced at random. Either the movement was halted, or released and allowed to accelerate by reducing the load, or reversed by suddenly increasing the current in the motor, so stretching the muscle. 4. Usually eight or sixteen responses to each kind of perturbation and a similar number of controls against a constant load were averaged. 5. Muscle activity was recorded as the electromyogram from surface electrodes over the belly of the long flexor in the lower forearm. Action potentials were usually full-wave rectified and integrated. 6. About 50 msec after a perturbation the muscle's activity alters in such a sense as to tend to compensate for the perturbation, i.e. it increases after a halt or a stretch and decreases after a release. The latency is similar in each case. 7. These responses are interpreted as manifestations of automatic servo action based on the stretch reflex. They are considered to be too early to be voluntary. 8. This interpretation was supported by measuring voluntary reaction times to perturbations under tracking conditions. They were found to be 90 msec or longer. 9. When the initial load was increased by a factor of 10, the servo responses were all scaled up likewise. Thus to a first approximation the gain of the servo is proportional to initial load. 10. It follows that in relaxed muscle the gain should be zero. This was confirmed by showing that stretching a relaxed muscle gives no reflex, or only a small one. 11. Gain appears to be determined by the level of muscle activation as determined by the effort made by the subject, rather than by the actual pressure exerted by the thumb. 12. Thus in fatigued muscle gain is boosted as the muscle has to be activated more strongly to keep up the same force output. The net effect is to compensate for fatigue and maintain the performance of the servo. 13. The Discussion centres on the implications of gain control in the servo. For a start, if the gain of the stretch reflex arc is zero in relaxed muscle, contractions cannot be initiated via the stretch reflex by simply causing the spindles to contract, as proposed on the original 'follow-up' servo theory.

Action Potentials

Stretch reflex and servo action in a variety of human muscles.

1. In the long flexor of the thumb the latency of the stretch reflex and of other manifestations of servo action is some 45 msec, roughly double the latency of a finger jerk. 2. Tendon jerks are feeble or absent in the long flexor of the thumb even in subjects with brisk long-latency stretch reflexes in this muscle. This, and other facts, suggests that the nervous mechanism of the tendon jerk is different from that of the stretch reflex. 3. A muscle that has feeble tendon jerks may show a late component in the response to a tendon tap, with a latency similar to that of the long-latency stretch reflex. 4. On the hypothesis that the excess latency of the stretch reflex over that of a tendon jerk is because the stretch reflex employs a cortical rather than a spinal arc, the excess would be expected to be larger in magnitude for the long flexor of the big toe and smaller for the jaw closing muscles. This is confirmed, 5. An alternative hypothesis that the long latency of stretch reflexes in thumb and toe is because they are excited by slow-conducting afferents is made improbable by the finding that stretch reflexes with an equal or greater excess latency are also found in proximal arm muscles. 6. The long-latency stretch reflex in proximal muscles was seen most distinctly in a healthy subject who happened to have feeble or absent tendon jerks. In ordinary subjects there is often a large, short-latency, presumably spinal component of the stretch reflex in proximal muscles; and short-latency responses to halt and release are also seen, The significance of this spinal latency servo action in proximal muscles remains to be explored. 7. The Discussion argues that the available data on conduction time to and from the cerebral cortex are compatible with the hypothesis that the long-latency component of the stretch reflex uses a transcortical reflex arc, and that none of the experiments described in the present paper are inimical to this view.

Biomechanical Phenomena