Reflex control of vascular capacitance.
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The influence of the linear acceleration of gravity on the vertical and horizontal vestibulo-ocular reflexes (VVOR, HVOR) as well as the vertical and horizontal optokinetic reflexes (VOKR, HOKR) has been examined in rabbits. Rabbits were mounted in a biaxial rate table in front of a rear projection tangent screen. Eye movements were measured with a light projection technique. The HVOR, VVOR, HOKR and VOKR were measured in rabbits which were maintained both prone and supine. The gain of the HVOR for the supine orientation was reduced at all frequencies tested (0.01-0.80 Hz). Similarly there was a reduction in the gain of the HOKR. By contrast, the gain of the VVOR in the supine orientation was enhanced over a lower range of frequencies (0.02-0.04 Hz) and reduced at higher frequencies (0.10-0.80 Hz). The gain of the VOKR was not reduced in the supine orientation. The range of eye positions over which compensatory eye movements occurred was restricted in the supine orientation. The altered orientation of the medio-laterally polarized hair cells of the utricular maculae with respect to gravity in the supine orientation may cause postural instability and facilitate 'righting reflexes'. A reduction in the gains of the HVOR, VVOR and HOKR caused by linear accelerations in the sagittal plane during locomotion may decrease automatic postural responses during certain movements in which these automatic postural adjustments would not necessarily be adaptive.
A parallel-cascade system identification method was used to identify intrinsic and reflex contributions to dynamic ankle stiffness over a wide range of tonic voluntary contraction levels and ankle positions in healthy human subjects. Intrinsic stiffness dynamics were described well by a linear pathway having elastic, viscous, and inertial properties. A velocity-sensitive pathway comprising a delay, a static non-linearity, resembling a half-wave rectifier, followed by a low-pass filter, described reflex stiffness dynamics. The absolute magnitude of intrinsic and reflex stiffness parameters varied from subject to subject but the relative changes with contraction level and position were consistent. Intrinsic stiffness increased monotonically with contraction level while reflex stiffness was maximal at low contraction levels and then decreased. Intrinsic and reflex stiffness both increased as the ankle was dorsiflexed. As a result, reflex mechanics made their largest relative contributions near the neutral position at low levels of activity. The size of the maximum reflex contribution varied widely among subjects, in some it was so small (ca 1%) that it would be unlikely to have any functional importance; however, in other subjects, reflex contributions were large enough (as high as 55% in one case) to play a significant role in the control of posture and movement. This variability may have arisen because stretch reflexes were not useful for the torque-matching task in these experiments. It will be of interest to examine other tasks where stretch reflexes would have a direct impact on performance.
1. In the crayfish brain, the responses of local spiking interneurons to body roll simulated by bending of statocyst hairs, were investigated with intracellular recording and staining techniques. The neurons had two separate branching portions in the protocerebrum and the deutocerebrum. They were named as type-I local neurons and further classified into 5 types (ac-U, vplc-U, vplc-B, vupc-U, vupc-B). 2. Vupc-U neurons showed excitatory responses and vplc-U neurons showed inhibitory responses to inward hair deflection of the statocyst ipsilateral to their deutocerebral branches. The other 3 types were of mixed populations of the interneurons showing either excitatory or inhibitory responses to the stimulation. 3. Of 10 type-I local neurons showing excitatory responses to inward hair deflection, 6 interneurons had output effects on oculomotor and/or descending neurons. All these 6 interneurons showed large EPSPs and much higher frequency of spikes to the hair stimulation than those of the other 4. All 8 type-I local neurons that showed inhibitory responses had no output effects. 4. Type-I local neurons controlled two equilibrium responses, compensatory eye movement and righting reflex, either simultaneously or independently.
The dynamics of neurons in the vestibular nuclei of canal-plugged, decerebrate cats were studied in response to lateral (roll) tilt. Forelimb and neck extensor reflexes recorded simultaneously develop a progressive phase lag above 0.1 Hz. Neurons which exhibited a muscle-like phase lag were excited during low frequency stimuli by ipsilateral side-up tilt (beta response). Neurons with alpha responses, excited during side-down tilt, exhibited a constant phase, without a high frequency lag. Vestibulospinal neurons were present in both of these response groups, as were units driven at monosynaptic latencies by electrical stimulation of the ipsilateral labyrinth. The phase-lagging beta responses are appropriate for contributing to the reflexes observed in the ipsilateral neck and contralateral forelimb.
In a supine subject, lowering of the foot from heart level to 50 cm below is known to stimulate the local reflex response and the baroreceptor outflow. We lowered and raised the leg of a supine subject periodically, with the leg stationary between movements (square wave). The Traube-Hering-Mayer wave (THM congruent to 0.1 Hz) was captured by or locked on to the leg movement over a certain frequency range, this is usually called the entrainment range. Square wave periodic leg movement in this manner on 10 male subjects, mean age 22 years, demonstrated that the THM frequency can be entrained. The lower limit of the entrainment bandwidth is 0.0841 (SD 0.0030) Hz and the upper limit is 0.1176 (SD 0.0013) Hz. Further examination showed that this phenomenon is independent of the breathing input. Comparison with the Traube-Hering-Mayer entrainment techniques of breathing and periodic neck suction using the Eckberg collar which stimulates the baroreceptors showed similar results. This work supports the hypothesis that the local reflex response and the baroreceptor outflow entrain the THM frequency.
The purpose of this study was to investigate the effects of long-term horizontal body position on baroreceptor reflex control of heart rate. Six male rhesus monkeys (6.2-9.4 kg) were given bolus injections of 4.0 micrograms/kg phenylephrine during each of the following conditions: awake, anesthetized (10 mg/kg ketamine HCl), and after beta-blockade (1 mg/kg propranolol HCl) before, 7, 14, and 28 days after being placed in a horizontal body cast. R-R interval vs. systolic arterial pressure was plotted, and the slope was determined by least-squares-fit linear regression. Baroreceptor slope was significantly reduced by 7 days of horizontal body position and remained attenuated throughout the 28-day restraint period both before and after beta-receptor blockade. These data are consistent with the thesis that prolonged exposure to a zero-gravity environment impairs autonomic reflex regulation of the cardiovascular system.
Under the orthostatic test (OT) pronounced compensation of the initial lowering of the arterial pressure (AP) and insignificant restoration of cardiac output arose in anesthetized cats. An important role of the baroreceptor reflexes from the carotid sinus in the mechanism of AP compensation was established. The response of the resistance anc capacitance vessels to OT was studied by the method of extracorporeal circulation. Quantitative evaluation of the blood pooling in the vascular system under OT was made. Compensatory response of the AP under OT was determined mainly by the reflex vasoconstriction of the resistance vessels, but not by the venomotor reactions.
1. Tonic labyrinth and neck reflexes were studied separately and in combination in the decerebrate cat before and after acute cerebellectomy. Reflex effects were observed as changes in the length of the medial head of triceps in both forelimbs. 2. In the decerebrate cat with an intact cerebellum the tonic labyrinth reflexes, elicited by side-up and side-down head rotations, produced asymmetric length changes in the medial head of triceps in both forelimbs, as described by Lindsay, Roberts & Rosenberg (1976). After cerebellectomy, head movements in either direction produced reflex shortenings in the medial triceps in both forelimbs, in contrast to the normal reciprocal reflex length changes. The presence of the cerebellum is thus required for the occurrence of the normal asymmetric labyrinth reflexes. 3. The direction of the neck reflexes remained unchanged by cerebellectomy. 4. It is suggested that the postural disturbances following cerebellar damage may partly result from the change in the form of the tonic labyrinth reflexes, which in combination with the tonic neck reflexes would no longer act in a stabilizing manner on the trunk.
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In the decapod Crustacea, Palinurus vulgaris and Fasus lalandii, the reflex influences of one particular proprioceptor organ, the coxo-basal chordotonal organ (CB), on all the muscles operating the proximal and distal joints of the same leg, have been analysed. The distal end of CB was clamped in fine forceps mounted on a servo-controlled stretcher, and CB length changes of 2 mm were applied. Motor unit activity of the different muscles was recorded as electromyograms (EMGs). 1. Two types of proprioceptive reflex evoked by CB length changes have been investigated: (a) resistance reflexes of the two levator and two depressor muscles of the same leg segment, the coxopodite, i.e. 'intrasegmental reflexes', (b) 'intersegmental reflexes' induced in the muscles operating the proximal (T-C) joint of the same leg, and in all eight muscles of the limb segments distat to CB. 2. Both levator muscles respond reflexly to imposed CB stretch (which normally occurs with limb 'depression'), while both depressors respond during CB shortening (or passive "elevation" of the leg). 3. Intersegmentally CB stretch reflexly activates the M-C extensor muscle, and sometimes facilitates the T-C remotor and C-P bender muscles. Shortening of the single CB organ of a leg excites one or two tonic motor units of the T-C promotor and M-C flexor muscles, and also facilitates the remotor, I-M reductor, and the single stretcher-opener excitatory motoneurone. 4. Some of the muscles, particularly the M-C flexor and extensor muscles, are also influenced intersegmentally by the resting length of CB, usually but not invariably in the same direction as for the corresponding dynamic reflexes. The role of the CB chordotonal organ is discussed, with particular consideration of its intersegmental reflex influence on the posture of the entire leg, and on the more complex motor behaviour of locomotion, where it may be specially significant in coordination of the limb in lateral walking. A complex picture of both tonic and dynamic, inra- and intersegmental reflex regulation of the positions and movements of the limb segments, thus emerges.
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Vestibular, visual, and proprioceptive influences on muscle activity correcting for backwards body tilt were investigated in normals and patients with bilateral peripheral vestibular deficits. Body tilt was induced by a dorsi-flexion rotation of the feet about the ankle joints while the subject stood on a force measuring platform. Ankle muscle activity and torque were monitored as upright stance was reestablished, and correlated with head angular accelerations and neck muscle activity. In normals with eyes closed, soleus stretch reflex activity at 50-80 ms was followed by two bursts of tibialis anterior (TA) EMG activity at ca 80 and 125 ms from the onset of 36 deg/s, 3 deg amplitude platform rotations. Neck muscle activity rotated the head backwards at the same time as TA activity rotated the body forwards about the ankle joints. Under the influence of vision, i.e. eyes open, slight increases in the second burst of TA activity, and ankle torque were observed. When the subjects sat, and were instructed to activate TA rapidly on onset of the platform movement, TA EMG activity increased gradually at ca. 150 ms and not as a burst. In patients with long-lasting bilateral vestibular deficits, both bursts of TA activity were significantly less than normal with eyes closed. Consequently sway correcting torques were abnormally low and all but one of the patients fell over backwards. With eyes open, TA activity was slightly less than, and ankle torques were approximately equal to normal values. In contrast to normals, TA responses obtained in standing and sitting positions were not significantly different. Neck EMG activity varied from normal, consisting of a long burst 100 ms in duration. The present data indicate that a coordinated pattern of ankle, and neck muscle activity occurs during the first 150 ms following induced backward tilt. Ankle muscle activity corrects for the body sway, and neck muscle activity attempts to stabilise the head with respect to earth fixed coordinates. It is proposed that the vestibulo-spinal reflex system predominantly underlies the genesis and coordination of this muscle activity.
Evoked electromyograms (EMGs) induced by galvanic stimulation of the labyrinth were examined in the muscles of the lower limbs. The labyrinthine-evoked EMGs appeared in the muscles of the femoral and gluteal regions and of the legs, and these muscle activities were changed by the head position in spite of the labyrinthine stimulation being the same. The vestibulo-spinal reflexes fulfil these functions in coordination with the neck proprioceptor.
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