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Biomedical subjects

J C Houk

Publications and source records attributed to J C Houk.

At least 73 records · Page 4Linked to original sources

Nonlinear viscosity of human wrist.

Nonlinear viscous properties of stretch and unloading reflexes in the human wrist were examined using constant-velocity ramp stretches and releases in the range between 5 and 500 mm/s. Subjects were asked to oppose an initial flexor preload and were instructed not to intervene voluntarily when the changes in position were applied. Electromyographic (EMG) activity and net force exerted by the wrist were measured. Although subjects were instructed not to intervene to the applied stretches, even well-practiced subjects sometimes showed unintended triggered reactions, which character could be assisting or resisting. A trial comparison method was used to detect and eliminate responses contaminated by unintended reactions. Ramp stretches further loaded the preloaded flexor muscles. Responses of EMG and force increased steeply initially but after about 1-cm displacement, the slope of these responses decreased to a lower value and remained constant during the remainder of the 5-cm ramp. For higher stretch velocities, the magnitudes and slopes of the responses of EMG and force increased but less than proportionally with ramp velocity. Except for the initial transient, EMG in the loaded flexor muscles and force responses could be described by a product relationship between a linear position-related term and a low fractional power of velocity, after a correction was made for delays in the reflex arc. Mean value of the exponent in the power function of velocity was 0.3 for EMG and 0.17 for force. For higher preloads, incremental responses of force to constant-velocity stretches, plotted as a function of wrist position, shifted to higher values and the slope of increase of force with position became somewhat steeper. This upward shift of the force trace reflects a change of apparent threshold of the stretch reflex. Ramp releases shortened and unloaded the preloaded flexor muscles and stretched the initially inactive extensor muscles. Flexor EMG activity declined progressively with a time course that was independent of velocity. Extensor EMG response depended on preload. At high preloads, there was no activity except for some bursting at the highest velocities. At low preloads, EMG activity was initially absent but started part way through the ramp. The increase of activity was somewhat greater for higher ramp velocities. Force responses to shortening ramps depended on preload. At high preloads, force responses superimposed at all of the low velocities but fell to slightly lower forces at the higher velocities. At low preloads, force traces again superimposed for low velocities and at high velocities only during the initial part of the response.(ABSTRACT TRUNCATED AT 400 WORDS)

Axons↗

Somatosensory properties of the inferior olive of the cat.

We examined the somatosensory properties of 391 neurons in the inferior olive in 20 cats that were anesthetized with barbiturate or decerebrated. A response consisted of a single spike with a variable number of wavelets followed by a long refractory period. Neurons responsive to natural somatosensory stimuli were recorded in all olivary subdivisions. The dorsal accessory olive (DAO) contained the highest proportion of responsive units (96%), compared with 66% for the medial accessory olive (MAO) and 43% for the principal olivary (PO) nucleus. Within the rostral DAO we found a refined cutaneous map of the entire contralateral body surface. In the caudal DAO responsiveness to manipulation of deep tissues became prominent, and both individual limbs and bilateral pairs were represented. In the medial region of the PO responsiveness to taps predominated and bilaterally symmetrical fields were frequent. The lateral PO was unresponsive under the conditions of these experiments. The MAO was distinguished by a greater complexity of receptive field and by a preponderance of deep over cutaneous modality. The lateral part of caudal MAO contained cells with interesting spatial patterns of excitation and inhibition, whereas most cells in the rostral MAO had purely excitatory fields. A teleceptive area receiving visual and auditory input was recognized in the medial MAO and nearby structures such as the dorsal cap. Contact and proprioceptive signals arriving via climbing fibers may provide the cerebellum with information necessary to relate the body to external objects.

Afferent Pathways↗

Speed and accuracy of compensatory responses to limb disturbances.

This study examines the speed and accuracy of compensatory responses to flexion-extension perturbations of the wrist in the horizontal plane. In Experiments 1 and 2 the subjects were required to establish an initial flexion or extension force of approximately 15% maximum at a prescribed initial muscle length. The perturbations changed the load force by +/-5% in both simple and choice reaction protocols. The results showed that the latencies to compensate for the perturbation were longer when the direction of disturbance was unknown (i.e., choice effect) and when the perturbation unloaded the muscle (i.e., directional effect). Accuracy constraints on the compensatory response increased movement time and reduced the variability of latency without affecting mean latency. In Experiment 3, a visual stimulus generated a comparable choice effect on latency to that produced by the perturbations, but no directional effect in relation to the preload was apparent. Our behavioral analysis of compensatory responses triggered by wrist perturbations confirms that these responses are susceptible to variables that influence the initiation of voluntary movements. Our analysis also demonstrates a directional preload effect that is stimulus specific.

Adaptation, Physiological↗

Velocity signals related to hand movements recorded from red nucleus neurons in monkeys.

Neural activity of the red nucleus was studied in monkeys trained to operate devices requiring shoulder, elbow, wrist, hand, or finger movements. Single cell activity was more closely related to movements of the hand and fingers than to movements of the other joints. Discharge consistently preceded movements by a constant time interval; duration of discharge was highly correlated with the duration of movement; and discharge rate was highly correlated with movement velocity. These data suggest a role for the rubrospinal pathway in the initiation and control of hand movements.

Action Potentials↗

Mechanisms of the clasp-knife reflex studied in an animal model.

The mechanisms of the clasp-knife reflex were studied in the soleus muscle of an animal model, the decerebrate cat with a dorsal hemisection of the lower thoracic cord. The reflex is shown to be autogenetic, and to depend on muscle length in keeping with previous suggestions. However, the magnitude of the inhibition increases with increasing initial force, and the inhibition is mimicked by gentle manipulation of the muscle and tendon surface. Concurrent muscle afferent recordings showed that the electromyogram (emg) reduction was not a result of a decline in Ia afferent input and was not well related to secondary or tendon organ afferent discharge. It is now known that many group III and some group IV muscle afferents are also activated by muscle stretch and contraction, and we here report limited stretch sensitivity in four non-spindle group II afferents. Since these fiber groups each include afferents that produce inhibition of extensor motoneurons, it is proposed that the clasp-knife reflex may result from the activation of these slowly conducting afferent fibers.

Afferent Pathways↗

Improvement in linearity and regulation of stiffness that results from actions of stretch reflex.

We studied stretch reflexes of soleus muscles of intercollicularly decerebrated cats using a new technique for estimating the component of a stretch reflex that results from the purely mechanical properties of the active muscle (mechanical response). The difference between a net stretch reflex and its underlying mechanical response provided a direct measure of reflex action. 1. The relative contributions of reflex action and the mechanical response are different for stretch and release. With stretch, reflex action is generally large and the mechanical response small. The opposite is true with release. The property that remains relatively constant, when stretch and release are compared, is the net stiffness in opposition to length change. We concluded that reflex action compensates for variations in the inherent stiffness of the muscle. 2. Compensation is effective over a range of intermediate values of initial force, but it fails whenever the reflex force approaches zero or the maximal force at that length. 3. Reflex action is capable of modifying muscular force within 22 ms of the onset of length change. This indicates that even during a fast gallop, there is sufficient time for reflex action. 4. The mechanical properties of the active soleus muscle are highly nonlinear; e.g., muscular stiffness becomes negative transiently during stretch. In stable decerebrate preparations we found that reflex action resulted in a considerable improvement in linearity. 5. Our results support two complementary hypotheses: a) stiffness may be the regulated property of the stretch reflex, and b) the main function of autogenetic reflexes may be to conpensate for variations in the properties of skeletal muscle rather than to oppose changes in load.

Animals↗

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↗

Reflex compensation for variations in the mechanical properties of a muscle.

Soleus muscles of anesthetized cats were stretched and released by different amounts while their motor axons were stimulated. Muscle force increased, then gave way in response to large stretch. In the presence of active stretch reflexes in decerebrate cats, the give in force was absent. We demonstrate that autogenetic reflexes can compensate for variations in muscular stiffness revealed when responses to stretch and release are compared.

Animals↗

Synaptic transmission at single glomeruli in the turtle cerebellum.

We have recorded from the granular layer of the turtle cerebellum extracellular unitary potentials that appear to reflect pre- and postsynaptic events at the synapse between a single swelling of a mossy fiber and the dendritic tips of several granule cells. The presynaptic component is an all-or-none potential. It can be directly activated by spinal stimulation and is unaltered by repetitive activity or by high concentrations of magnesium. The postsynaptic component is a graded potential. It follows the presynaptic component by approximately 1 millisecond and is depressed by repetitive activity and by high concentrations of magnesium. The recording of large potentials produced by the flow of postsynaptic current within a single glomerulus suggests powerful transmission. Electron micrographs demonstrate large cerebellar glomeruli in the turtle and a substantial accumulation of mitochondria in the dendritic tips of granule cells.

Action Potentials↗