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Biomedical subjects

Z Hasan

Publications and source records attributed to Z Hasan.

57 records · Page 4Linked to original sources

Regulatory actions of human stretch reflex.

1. The stretch reflex in the elbow flexor musculature was studied in 23 human subjects. The subjects were required to establish an initial force equivalent to 10% maximum at a prescribed initial length; mechanical disturbances delivered at random times increased load force to 15% or reduced it to 5%. We measured arm force, displacement, and EMG (usually biceps); acceleration was calculated from displacement, and average responses from sets of 10 like trials. 2. Modification of the stretch reflex was studied by comparing average responses obtained with different instructions, but with the same disturbance. The usual introductions were "compensate for arm deflection" and "do not intervene voluntarily". The initial response did not depend on instruction; changes in response that depended on instruction began abruptly after a latent period which ranged from 70 to 320 ms (measured from force and acceleration), depending on conditions and subject. The latency became longer (10-50 ms) and more variable when the subject did not know the direction of disturbance in advance. This and other observations indicate that modifications of the stretch reflex are not produced by servo actions. They are produced by triggered reactions, which occur at both short and long latencies and which have properties resembling the movements produced in a reaction-time task. 3. We confirmed that most subjects can suppress triggered reactions when the instruction calls for no intervention, leaving an unmodified reflex response. This response consists of a compliant deflection of the arm in the direction of the disturbance. 4. The compensatory actions associated with unmodified stretch (and unloading) reflexes were assessed from EMG responses of biceps. During a 300-ms transient phase, EMG changes were notably asymmetric when responses to symmetric disturbances were compared. Increased force stretched biceps and produced a prominent increase in EMG, whereas decreased force allowed biceps to shorten and produced either an EMG decrease of smaller magnitude or an actual increase. These asymmetric reflex actions produced quite symmetric mechanical responses (arm displacements and forces), which implies the existence of and compensation for nonlinear muscle mechanical properties. This result is discussed in relation to the hypothesis that the function of the stretch reflex is to compensate for variations in muscle properties, thus maintaining stiffness. 5. Effective control of muscle length or joint position does not result from servo action by the stretch reflex. Errors in position are corrected only when triggered reactions are superimposed on the reflex response.

Adult↗

Analysis of response properties of deefferented mammalian spindle receptors based on frequency response.

Sinusoidal responses of primary and secondary endings in deefferented spindles of anesthetized cats were studied over the low-frequency range 0.001-0.1 Hz. Stretch amplitudes were chosen conservatively small (25-100 mum peak-to-peak) so as to lie within the linear region. 1. At 0.1 Hz average sensitivity was 350 pps/mm for primary endings and 80 pps/mm for secondary endings. Sensitivity fell to lower values at lower frequencies, but even at 0.001 Hz, corresponding to 17 min/cycle, sensitivity remained elevated above static values determined with large stretches. Phase lead varied from 5 to 50 degrees and, in the case of primary endings, tended to be greater at lower frequencies. 2. Except for the different scaling factors, the only apparent difference between the frequency responses of primary and secondary endings was a tendency for primary endings to show a greater phase lead over the range 0.001-0.01 Hz. 3. Dynamic responsiveness was assessed theoretically from frequency-response data by calculating responses to ramps at various velocities. Over most of the velocity range dynamic responses were not proportional to velocity. The greater dynamic responsiveness of primary endings during large (6 mm) ramp stretches might be related to frequency response below 0.01 Hz. 4. Certain aspects of dynamic responsiveness to large ramps (6 mm) were accounted for by assuming all phases of responses were attenuated by 25 dB in the case of primary endings and 20 dB in the case of secondary endings. The nonlinearity responsible for attenuation appears to occur at an early stage in the sensory process. 5. Comparison of individual responses to slow ramps with predictions based on linear theory indicated the presence of abrupt departures from linearity for both primary and secondary endings.

Animals↗

Transition in sensitivity of spindle receptors that occurs when muscle is stretched more than a fraction of a millimeter.

We studied the responses of 34 deefferented spindle receptors to slowly applied ramp stretches (0.01-1 mm/s) of small (0.02-0.2 mm) and intermediate (0.2-1 mm) amplitudes. The afferent discharge from primary and secondary endings was recorded from filaments of dorsal root in anesthetized cats. 1. Responses of most endings to ramps of intermediate amplitude showed abrupt changes in slope (discontinuities) which were highly repeatable. Discontinuities occurred more nearly at constant stretch (in the range 50-400 mum for different receptors) than at constant discharge rate. They were less pronounced in the case of secondary endings. 2. Changes in sensitivity occurred when the degree of stretch exceeded a transitional amplitude which ranged from 50 to 200 mum. These changes were studied by constructing plots based on a family of responses to a family of ramps which were scaled versions of each other. The plots indicated that reductions in sensitivity occurred both during stretch and during adaptation; the reductions were more marked for primary than for secondary endings. 3. Responses were modified considerably by preceding changes in muscle length. When the last change was an increase of a few millimeters, discontinuities became more pronounced and other changes in the appearance of the dynamic response occurred, particularly in the case of primary endings. These changes could last for several minutes, but were abolished by a single test stretch of intermediate amplitude. 4. The resetting of high sensitivity that occurs when muscle length is changed, the discontinuities, the transitions in sensitivity, nonlinear adaptation, and the effects of previous length change appeared to be related phenomena. They can all be accounted for by the hypothesis that polar zones of intrafusal muscle fibers possess a frictionlike property, one analogous to that which has been described for whole muscle. A simple nonlinear model which shows these features is presented. 5. The adequate stimulus for a change in primary ending discharge is a small change in muscle length, relatively independently of its velocity. The dynamic response arises mainly from a changing sensitivity to length itself, which is a nonlinear property.

Animals↗