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B J Yates

Publications and source records attributed to B J Yates.

88 records · Page 5Linked to original sources

Neck muscle spindle activity in the decerebrate, unparalyzed cat: dynamics and influence of vestibular stimulation.

1. Using floating electrodes, we recorded from neck-muscle spindle afferents in the C2 dorsal root ganglion of the decerebrate cat. Nerves to dorsal neck muscles were cut so that the afferents presumably originated mainly from ventral and ventrolateral perivertebral muscles and sternocleidomastoid. One goal of our experiments was to study possible vestibular influence exerted on these spindles via the fusimotor system. Unparalyzed preparations were therefore used. 2. Stimuli consisted of sinusoidal rotations in vertical planes. Neck tilt stretched neck muscles, whereas whole-body tilt stimulated vestibular receptors. 3. For each afferent we first determined the most effective direction of neck tilt, then used stimuli oriented close to this direction to study response dynamics, particularly gain of responses to stimuli of different amplitudes (0.5-7.5 degrees). 4. Three-quarters of the afferents failed to respond to 0.5 degrees, 0.2-Hz neck rotations. Stimuli that were effective usually elicited responses that had low gain and were linear over the whole range of amplitudes. Only a few afferents had behavior typical of spindle primary afferents: high-gain responses to small sinusoidal stimuli, gain decreasing as stimulus amplitude increases. This prevalence of static spindle responses in the unparalyzed cat is in striking contrast to results obtained on neck-muscle spindles in paralyzed, decerebrate cats, and on hindlimb extensor muscle spindles in decerebrate, unparalyzed cats. 5. Paralysis produced by injection of Flaxedil changed the behavior of 2/4 spindle afferents tested, causing the appearance of high-gain responses to 0.5 degrees stimuli and of nonlinear behavior.(ABSTRACT TRUNCATED AT 250 WORDS)

Afferent Pathways↗

Responses of spinal cord neurons following stimulation of A beta femoral-saphenous venous afferent fibers.

A population of large (A beta) afferents is known to have endings in the wall of the femoral-saphenous vein. These afferents project to the lower lumbar spinal cord. The purpose of the present study was to identify, localize, and characterize spinal neurons that receive inputs from such afferents. Responses of 50 neurons in the L6 spinal cord segment of decerebrate-spinal cats or intact cats anesthetized using alpha-chloralose were recorded following electrical stimulation of these afferents. Observations were also made on the convergence of muscle and cutaneous afferent inputs onto neurons driven by stimulation of afferents terminating in the femoral-saphenous vein. All recording sites were marked either by intracellularly staining the element characterized with HRP or by extracellularly iontophoresing a small quantity of this tracer. The cells were driven for long durations (mean of 51.5 ms, S.E.M. of 10.0) by single-shock stimulation of femoral-saphenous venous afferents. The recording sites were located in Rexed's laminae IV-VIII and X. Eight of the 50 neurons were activated by venous afferent stimulation at latencies equal to or shorter than that of the first negative wave of the cord dorsum potential; these units were driven at a mean latency of 1.4 ms (S.E.M. of 0.25) following the arrival of the afferent volley at the cord and were assumed to receive monosynaptic, or at least relatively direct, inputs from the primary afferents. Most of these cells (6 of 8) were located in lamina V. The majority of the neurons studied (37 of 50) were activated at latencies longer than 3 ms following the arrival of the afferent volley at the cord; about half (19 of 37) of those activated at longer latencies were located in lamina VII, and the rest were scattered among the other laminae. Twenty-eight of 40 venous afferent-driven cells tested could also be activated by electrical stimulation of either the posterior tibial or sural nerve. In general, the stimulation intensities necessary to activate the neurons were only sufficient to excite large (A alpha or A beta) muscle and cutaneous afferents. Neurons receiving the shortest latency inputs from the femoral-saphenous vein were less likely to receive convergent inputs from muscle or skin than were neurons activated by venous afferents at longer latency.

Action Potentials↗

Peripheral input to L4 neurons whose activity is modulated by neck rotation.

We studied, in decerebrate cats, peripheral input from the ipsilateral hindlimb to L4 neurons whose activity was modulated by neck rotation ('neck-modulated neurons'). Most neurons received convergent input from muscle, cutaneous and mixed nerves. In about half the neurons muscle input consisted of short-latency group I or group II excitation or inhibition, with group II effects far more frequent. Such synaptic actions were produced almost entirely by stimulation of quadriceps or sartorius. In the other neurons muscle afferents produced only late, diffuse excitation or inhibition, with thresholds usually in the group III range.

Animals↗

Three-dimensional sensitivity and caudal projection of neck spindle afferents.

1. We recorded from neck muscle spindle afferents in the C2 dorsal root ganglion of the decerebrate cat using floating electrodes. The afferents presumably innervated mainly ventral and ventrolateral perivertebral muscles, and sternocleidomastoid. Stimuli consisted of combinations of rotatory head movements about the roll/pitch or pitch/yaw axes. An important difference from our earlier experiments (10) was the addition of yaw movement to the stimulus paradigm making possible a three-dimensional analysis of afferent behavior. 2. For each afferent we determined the most effective direction of tilt (orientation of the response vector) in three dimensions by using sinusoidal stimuli that combined pitch and roll, or pitch and yaw, or by measuring the gains to responses to roll, pitch, and yaw rotation. 3. Most afferents were sensitive to rotation around all three axes; pitch and yaw were usually more effective than roll. There was no indication of clustering of response vectors, as might be expected if the receptors were located in a small number of muscles each of which has receptors aligned in a homogeneous direction. 4. The responses of afferents were further studied using sinusoidal and trapezoidal stimuli aligned as closely as possible with the orientation of their response vector. The availability of the yaw stimulus made receptor classification based on response linearity, gain, and dynamic index more reliable than in our earlier experiments (10). 5. Muscle spindle responses were divided into three categories: A, B, and ambiguous. The evidence suggests that category A are probably spindle primary receptors and category B are secondaries. Ambiguous receptors have intermediate properties. 6. The caudal projection of spindle afferents was examined by delivering antidromic stimuli with a movable electrode on the surface of the ipsilateral dorsal column. Eighteen percent of the afferents projected to C4, and 14% as far as C5. Long caudal projections can be found in A, B, and ambiguous receptors with a range of directional sensitivities. 7. The evidence suggests that C2 spindle afferents make synapses in the midcervical segments with interneurons and propriospinal neurons that are part of the intraspinal pathway of the tonic neck reflex.

Action Potentials↗

Tracing of afferent pathways from the femoral-saphenous vein to the dorsal root ganglia using transport of horseradish peroxidase.

The retrograde transport of horseradish peroxidase (HRP) was used to trace afferents from the femoral-saphenous vein to the dorsal root ganglia in the cat. Afferents arising along the entire length of the vein projected to very localized spinal levels; 63% of the labeled cells counted were located in the L6 dorsal root ganglion, 37% were located in the L5 ganglion and less than 1% were located at other levels. Most of the cell bodies labeled by the application of HRP to the femoral-saphenous vein were small in size (diameter less than 35 microns). However, some large cell bodies (diameter greater than 50 microns) were also noted. It was estimated that over two-thirds of the femoral-saphenous venous afferents were C fibers; at least 15% were estimated to be A fibers. The largest venous afferents were predicted to conduct action potentials at approximately 60 m/s.

Afferent Pathways↗

Properties of femoral venous afferent inputs and lumbosacral distribution of spinal evoked activity.

The present studies were done to determine details of the anatomical and physiological characteristics of femoral-saphenous venous afferent input to the lumbar spinal cord. Gross anatomical examination revealed that afferent bundles could be seen coursing from the saphenous nerve to the femoral-saphenous vein. Compound action potentials elicited by femoral-saphenous venous afferent stimulation were recorded from the femoral nerve and in dorsal rootlets of the 6th lumbar cord segment. The compound action potentials included activity correlated with that of fibers conducting impulses at the rate of 31 to 61 m/s. Lumbar cord dorsum potentials elicited by femoral-saphenous venous afferent stimulation were abolished by rhizotomy of the most caudal rootlets of the 6th lumbar cord segment. L6 was also the cord segment from which the largest amplitude cord dorsum negative potentials were recorded, while action potentials with large late positive waves were recorded from more caudal cord segments. These observations suggested that the L6 segment contained the largest number of spinal neurons responding to primary femoral-saphenous venous afferent input, and that input reached the more caudal segments via intersegmental connections. A proposed physiological role of these afferents is briefly described.

Afferent Pathways↗

Activation of spinal cord interneurons which process inputs from the femoral-saphenous vein.

Recordings were made from single spinal cord interneurons which could be activated by electrical stimulation of afferents terminating in the wall of the femoral-saphenous vein. Interneurons were either excited or both excited and inhibited by venous afferent stimulation. Most of the venous afferent-driven interneurons could also be driven by electrical activation of A-alpha beta muscle and cutaneous afferents. Stimulation of several different muscle nerves drove single interneurons.

Animals↗

Properties of spinal cord processing of femoral venous afferent input revealed by analysis of evoked potentials.

Two characteristics of spinal cord interneurons which receive inputs from femoral-saphenous venous afferents were examined. The shortest pathway between primary femoral-saphenous venous afferents and alpha-motoneurons was shown to be a di- or tri-synaptic circuit. In addition, the largest focal synaptic field potentials elicited by venous afferent stimulation at short latency were recorded from Rexed's lamina V. It was thus concluded that most of the first interneurons excited by venous afferents are found in this lamina.

Afferent Pathways↗

Lumbar spinal cord responses to limb vein distention.

The purpose of this study was to determine if central neural responses were elicited by distention of limb veins, and to compare the pattern of these response to those produced in previous studies using electrical stimulation to excite limb venous afferent fibers. Spinal evoked potentials were measured in response to stretch of the wall of a segment of the femoral-saphenous vein by perfusion-distention or by mechanical stretch. These studies revealed that spinal cord evoked potentials were elicited by these procedures, and that the activated venous afferent fibers coursed through the saphenous nerve and entered the sixth lumber spinal cord segment. The minimum stretches which were required to elicit spinal evoked potentials were produced by perfusion pressures starting at 2-3 mm Hg, or by mechanical stretch of the wall of 5 micron/mm. A vein wall proprioceptor hypothesis is proposed and discussed in the light of these findings. In addition to the cord dorsum evoked potentials, distention or stretch of the vein wall elicited ventral root potentials (excitatory postsynaptic population potentials) which are known to be produced by excitatory inputs to motoneurons. A venous afferent mediated muscle-tonus venopressor mechanism hypothesis is proposed and discussed in the light of these and previous findings.

Afferent Pathways↗

Origin and properties of spinal cord field potentials.

Electrical or natural stimulation of cutaneous, muscle, or visceral nerves evokes a field potential in the spinal cord; the field potential elicited by each type of afferent fiber differs in wave form. Cutaneous nerve-evoked spinal cord field potentials consist of a triphasic spike, one to three negative waves, and a slow positive wave. Muscle nerve-evoked spinal cord field potentials are composed of a triphasic spike, a slow positive wave, and one to three negative waves with different relative latencies and durations than the negative waves evoked by cutaneous nerves. Visceral nerve-evoked potentials in the spinal cord comprise only a single negative wave and a slow positive wave. The triphasic spike is a compound action potential propagating through the large primary afferent fibers; the negative waves are generated by dorsal horn interneurons; the positive wave is a reflection of primary afferent depolarization. A knowledge of the origin and properties of spinal cord field potentials has practical usefulness for both the experimental neurophysiologist and the clinician.

Electrodiagnosis↗

Vestibular influences on the sympathetic nervous system.

Studies using both electrical and natural stimulation have established that the vestibular system has prominent effects on sympathetic outflow and blood pressure. Preliminary evidence suggests that receptors in both otolith organs and semicircular canals are involved in producing these effects. Furthermore, vestibulosympathetic reflexes appear to be mediated by the medial vestibular nucleus and slowly conducting projections from the rostral ventrolateral medulla and caudal medullary raphe nuclei to preganglionic neurons in the thoracic spinal cord. However, many details are missing from our knowledge and understanding of the functional significance and neural substrate of vestibular influences on the sympathetic nervous system.

Animals↗

Role of the vestibular system in regulating respiratory muscle activity during movement.

1. Changes in posture can affect the resting length of the diaphragm, which is corrected through increases in both diaphragm and abdominal muscle activity. Furthermore, postural alterations can diminish airway patency, which must be compensated for through increases in firing of particular upper airway muscles. 2. Recent evidence has shown that the vestibular system participates in adjusting the activity of both upper airway muscles and respiratory pump muscles during movement and changes in body position. 3. Vestibulo-respiratory responses do not appear to be mediated through the brainstem respiratory groups; labyrinthine influences on respiratory pump muscles may be relayed through neurons in the medial medullary reticular formation, which have recently been demonstrated to provide inputs to both abdominal and diaphragm motoneurons. 4. Three regions of the cerebellum that receive vestibular inputs, the fastigial nucleus, the nodulus/uvula and the anterior lobe, also influence respiratory muscle activity, although the physiological role of cerebellar regulation of respiratory activity is yet to be determined. 5. It is practical for the vestibular system to participate in the control of respiration, to provide for rapid adjustments in ventilation such that the oxygen demands of the body are continually matched during movement and exercise.

Animals↗

Horizontal rotation responses of medullary reticular neurons in the decerebrate cat.

This study examines the response of neurons in the medullary reticular formation of the decerebrate cat to sinusoidal yaw rotations in the plane of the horizontal semicircular canals. Responsive neurons that could be antidromically activated from the spinal cord appeared to be less sensitive to the rotary stimulus than the rest of the population or responsive neurons. Most neurons had response dynamics similar to those of semicircular canal afferents.

Animals↗

Physiological evidence that the vestibular system participates in autonomic and respiratory control.

Electrical or natural stimulation of the vestibular system results in changes in blood pressure and respiratory motor output. An increase in excitatory drive on the sympathetic nervous system occurs during nose-up vestibular stimulation in cats; this response is appropriate to offset orthostatic hypotension that could result from nose-up body rotations during movements such as vertical climbing. In addition, transection of the vestibular nerves in anesthetized or awake cats compromises the ability to correct decreases in blood pressure that result from nose-up body tilt. The vestibular system also has influences on respiratory muscles; these effects are appropriate to participate in making adjustments in the activity of respiratory muscles that are necessary to offset mechanical constraints on these muscles that occur during changes in body position. These data thus suggest that the influences of the vestibular system on the autonomic and respiratory systems serve to maintain homeostasis during movement.

Animals↗

Horizontal linear and angular responses of neurons in the medial vestibular nucleus of the decerebrate cat.

Responses to linear accelerations in the earth-horizontal plane (typically provoked by tilts of the head or body) are characterized by a stimulus direction that produces the maximal excitation. Although changes in cardiovascular, sympathetic, and respiratory outflow are maximized during pitch, no collection of central vestibular neurons had been identified where pitch responses predominate. In the present study, response properties of neurons in the medial vestibular nucleus were examined in decerebrate cats placed on a turntable. Activation of otolith afferents was provided by constant velocity rotation with the turntable axis tilted 5 degrees from the vertical. Responsive neurons exhibited a sinusoidal modulation in their firing rate; the optimal excitatory stimulus direction was derived from responses to clockwise and counterclockwise rotations. Many of these neurons were also tested for input from horizontal semicircular canals using 0.5 Hz sinusoidal rotation about an earth-vertical axis. Of 22 tilt-sensitive neurons in the medial vestibular nucleus whose optimal stimulus direction was determined, 9 were best stimulated by pitch, 10 by stimuli in one of the two vertical semicircular canal planes, and 3 by roll. Of the 33 neurons in this nucleus tested for possible convergent inputs from the otolith organs and the horizontal semicircular canals, 8 responded to both the constant velocity (otolith) stimulus and to the sinusoidal rotation, 7 appeared to receive otolith, but not horizontal canal, input, while 18 had a canal, but no otolith, response. Thus, besides serving as a relay for horizontal canal signals, the medial vestibular nucleus may also be an important relay for information about orientation within the sagittal (pitch) plane.

Acceleration↗