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Postflight measurements of Achilles tendon reflex duration on Skylab crewmen indicate a state of disequilibrium between the flexor and extensor muscle groups with an initial decrease in reflex duration. As the muscles regain strength and mass there occurs an overcompensation reflected by increased reflex duration. Finally, when a normal neuromuscular state is reached the reflex duration returns to baseline value.
Impairment of gaze and head stabilization reflexes can lead to disorientation and reduced performance in sensorimotor tasks such as piloting of spacecraft. Transitions between different gravitoinertial force (gif) environments - as during different phases of space flight - provide an extreme test of the adaptive capabilities of these mechanisms. We wish to determine to what extent the sensorimotor skills acquired in one gravity environment will transfer to others, and to what extent gravity serves as a context cue for inhibiting such transfer. We use the general approach of adapting a response (saccades, vestibuloocular reflex: VOR, or vestibulocollic reflex: VCR) to a particular change in gain or phase in one gif condition, adapting to a different gain or phase in a second gif condition, and then seeing if gif itself - the context cue - can recall the previously-learned adapted responses. Previous evidence indicates that unless there is specific training to induce context-specificity, reflex adaptation is sequential rather than simultaneous. Various experiments in this project investigate the behavioral properties, neurophysiological basis, and anatomical substrate of context-specific learning, using otolith (gravity) signals as a context cue. In the following, we outline the methods for all experiments in this project, and provide details and results on selected experiments.
Evidence from animals indicates that skeletal muscle afferents activate the vestibular nuclei and that both vestibular and skeletal muscle afferents have inputs to the ventrolateral medulla. The purpose of the present study was to investigate the interaction between the vestibulosympathetic and skeletal muscle reflexes on muscle sympathetic nerve activity (MSNA) and arterial pressure in humans. MSNA, arterial pressure, and heart rate were measured in 17 healthy subjects in the prone position during three experimental trials. The three trials were 2 min of 1) head-down rotation (HDR) to engage the vestibulosympathetic reflex, 2) isometric handgrip (IHG) at 30% maximal voluntary contraction to activate skeletal muscle afferents, and 3) HDR and IHG performed simultaneously. The order of the three trials was randomized. HDR and IHG performed alone increased total MSNA by 46 +/- 16 and 77 +/- 24 units, respectively (P < 0.01). During the HDR plus IHG trial, MSNA increased 142 +/- 38 units (P < 0.01). This increase was not significantly different from the sum of the individual trials (130 +/- 41 units). This finding was also observed with mean arterial pressure (sum = 21 +/- 2 mmHg and HDR + IHG = 22 +/- 2 mmHg). These findings suggest that there is an additive interaction for MSNA and arterial pressure when the vestibulosympathetic and skeletal muscle reflexes are engaged simultaneously in humans. Therefore, no central modulation exists between these two reflexes with regard to MSNA output in humans.
Head movements in the sagittal pitch plane typically involve off-axis rotation requiring both vertical and horizontal vergence ocular reflexes to compensate for angular and translational motion relative to visual targets of interest. The purpose of this study was to compare passive pitch VOR responses during rotation about an Earth-vertical axis (canal only cues) with off-axis rotation (canal and otolith cues). Methods. Eleven human subjects were oscillated sinusoidally at 0.13, 0.3 and 0.56 Hz while lying left-side down with the interaural axis either aligned with the axis of rotation or offset by 50 cm. In a second set of measurements, twelve subjects were also tested during sinusoidally varying centrifugation over the same frequency range. The modulation of vertical and horizontal vergence ocular responses was measured with a binocular videography system. Results. Off-axis pitch rotation enhanced the vertical VOR at lower frequencies and enhanced the vergence VOR at higher frequencies. During sinusoidally varying centrifugation, the opposite trend was observed for vergence, with both vertical and vergence vestibulo-ocular reflexes being suppressed at the highest frequency. Discussion. These differential effects of off-axis rotation over the 0.13 to 0.56 Hz range are consistent with the hypothesis that otolith-ocular reflexes are segregated in part on the basis of stimulus frequency. At the lower frequencies, tilt otolith-ocular responses compensate for declining canal input. At higher frequencies, translational otolith-ocular reflexes compensate for declining visual contributions to the kinematic demands required for fixating near targets.
Constant velocity Off-Vertical Axis Rotation (OVAR) imposes a continuously varying orientation of the head and body relative to gravity. The ensuing ocular reflexes include modulation of both horizontal and torsional eye velocity as a function of the varying linear acceleration along the lateral plane. The purpose of this study was to examine whether the modulation of these ocular reflexes would be modified by different head-on-trunk positions. Ten human subjects were rotated in darkness about their longitudinal axis 20 deg off-vertical at constant rates of 45 and 180 deg/s, corresponding to 0.125 and 0.5 Hz. Binocular responses were obtained with video-oculography with the head and trunk aligned, and then with the head turned relative to the trunk 40 deg to the right or left of center. Sinusoidal curve fits were used to derive amplitude, phase and bias velocity of the eye movements across multiple cycles for each head-on-trunk position. Consistent with previous studies, the modulation of torsional eye movements was greater at 0.125 Hz while the modulation of horizontal eye movements was greater at 0.5 Hz. Neither amplitude nor bias velocities were significantly altered by head-on-trunk position. The phases of both torsional and horizontal ocular reflexes, on the other hand, shifted towards alignment with the head. These results are consistent with the modulation of torsional and horizontal ocular reflexes during OVAR being primarily mediated by the otoliths in response to the sinusoidally varying linear acceleration along the interaural head axis.
Motor learning was induced in the translational vestibulo-ocular reflex (TVOR) when monkeys were repeatedly subjected to a brief (0.5 sec) head translation while they tried to maintain binocular fixation on a visual target for juice rewards. If the target was world-fixed, the initial eye speed of the TVOR gradually increased; if the target was head-fixed, the initial eye speed of the TVOR gradually decreased. The rate of learning acquisition was very rapid, with a time constant of approximately 100 trials, which was equivalent to <1 min of accumulated stimulation. These learned changes were consolidated over >or=1 d without any reinforcement, indicating induction of long-term synaptic plasticity. Although the learning generalized to targets with different viewing distances and to head translations with different accelerations, it was highly specific for the particular combination of head motion and evoked eye movement associated with the training. For example, it was specific to the modality of the stimulus (translation vs rotation) and the direction of the evoked eye movement in the training. Furthermore, when one eye was aligned with the heading direction so that it remained motionless during training, learning was not expressed in this eye, but only in the other nonaligned eye. These specificities show that the learning sites are neither in the sensory nor the motor limb of the reflex but in the sensory-motor transformation stage of the reflex. The dependence of the learning on both head motion and evoked eye movement suggests that Hebbian learning may be one of the underlying cellular mechanisms.
It has been proposed that a vestibular reflex originating in the otolith organs and other body graviceptors modulates sympathetic activity during changes in posture with regard to gravity. To test this hypothesis, we selectively stimulated otolith and body graviceptors sinusoidally along different head axes in the coronal plane with off-vertical axis rotation (OVAR) and recorded sympathetic efferent activity in the peroneal nerve (muscle sympathetic nerve activity, MSNA), blood pressure, heart rate, and respiratory rate. All parameters were entrained during OVAR at the frequency of rotation, with MSNA increasing in nose-up positions during forward linear acceleration and decreasing when nose-down. MSNA was correlated closely with blood pressure when subjects were within +/-90 degrees of nose-down positions with a delay of 1.4 s, the normal latency of baroreflex-driven changes in MSNA. Thus, in the nose-down position, MSNA was probably driven by baroreflex afferents. In contrast, when subjects were within +/-45 degrees of the nose-up position, i.e., when positive linear acceleration was maximal along the naso-ocipital axis, MSNA was closely related to gravitational acceleration at a latency of 0.4 s. This delay is too short for MSNA changes to be mediated by the baroreflex, but it is compatible with the delay of a response originating in the vestibular system. We postulate that a vestibulosympathetic reflex, probably originating mainly in the otolith organs, contributes to blood pressure maintenance during forward linear acceleration. Because of its short latency, this reflex may be one of the earliest mechanisms to sustain blood pressure upon standing.
1. To investigate the neural substrate of vestibulospinal reflexes in decerebrate cats, we studied the responses of pontomedullary reticulospinal neurons to natural stimulation of the labyrinth in vertical planes. Our principal aim was to determine whether reticulospinal neurons that terminate in, or are likely to give off collaterals to, the upper cervical segments had properties similar to those of the vestibulocollic reflex (VCR). 2. Antidromic stimulation was used to determine whether the neurons projected to the neck, lower cervical, thoracic, or lumbar levels. Dynamics of the responses of spontaneously firing neurons were studied with sinusoidal stimuli delivered at 0.05-1 Hz and aligned to the plane of body rotation, that produced maximal modulation of the neuron (response vector orientation). Each neuron was assigned a vestibular input classification of otolith, vertical canal, otolith + canal, or spatial-temporal convergence (STC). 3. We found, in agreement with previous studies, that the largest fraction of pontomedullary reticulospinal neurons projected to the lumbar cord, and that only a small number ended in the neck segments. Neurons projecting to all levels of the spinal cord had similar responses to labyrinth stimulation. 4. Reticulospinal neurons that received only vertical canal inputs were rare (1 of 67 units). Most reticulospinal neurons (48%) received predominant otolith inputs, 18% received otolith + canal input, and only 9% had STC behavior. These data are in sharp contrast to the results of our previous studies of vestibulospinal neurons. A considerable portion of vestibulospinal neurons receives vertical canal input (38%), fewer receive predominantly otolith input (22%), whereas the proportion that have otolith + canal input or STC behavior is similar to our present reticulospinal data. 5. The response vector orientations of our reticulospinal neurons, particularly those with canal inputs (canal, otolith + canal, STC) were predominantly in the roll quadrants. There was no evidence of convergence of inputs from like canals across the midline (e.g., right anterior + left anterior). 6. Two characteristics of the VCR, STC behavior and bilateral input from symmetric vertical canals (in some muscles), cannot be accounted for by the reticulospinal neurons that we studied. Because these characteristics are also not seen in vestibulocollic neurons, they are likely to be the result of the appropriate convergence of vestibular signals in the spinal cord. 7. Pontomedullary reticulospinal neurons seem particularly well suited to play a role in gravity-dependent postural reflexes of neck and limbs.
An experiment was conducted in order to determine the changes in loudness brought about by electro-cutaneous elicitation of the middle-ear reflex. Subjects were required to judge the relative loudness of the second of three consecutive 30-msec bursts of tone, the second tone being accompanied by an electrical shock to the external auditory meatus, capable of eliciting a contraction of the middle-ear muscles. The difference between these judgments and those of the control condition (shock on the arm) was taken to represent a measure of the attenuation provided by contraction of the middle-ear muscles. Test tones were 500, 1000, 2000, and 3000 Hz at levels of 65, 75, 85, 95, and 105 dB. The results indicate that the middle-ear reflex decreases the middle-ear's transmission mainly for low-frequency sounds. The results fail to lend support to the Loeb-Riopelle hypothesis that the middle-ear reflex acts as a limiter, rather than a linear attenuator.
The contribution of the fastigial nucleus and the vestibular nerves (eighth cranial nerves) to the orthostatic reflexes in anesthetized, paralyzed cats was studied. Bilateral lesions of the rostral fastigial nucleus resulted in impairment of the reflex changes in blood pressure, femoral arterial flow, and resistance evoked by head-up tilting to 30 deg or 60 deg. The rostral fastigial nucleus, which might be triggered by the vestibular apparatus, appears to participate in concert with the baroreceptors in the initiation and possibly the maintenance of the orthostatic reflexes.
A new device has been designed, constructed, and evaluated to characterize the human carotid baroreceptor-cardiac reflex response relation rapidly. This system was designed for study of reflex responses of astronauts before, during, and after space travel. The system comprises a new tightly sealing silicon rubber neck chamber, a stepping motor-driven electrodeposited nickel bellows pressure system, capable of delivering sequential R-wave-triggered neck chamber pressure changes between +40 and -65 mmHg, and a microprocessor-based electronics system for control of pressure steps and analysis and display of responses. This new system provokes classic sigmoid baroreceptor-cardiac reflex responses with threshold, linear, and saturation ranges in most human volunteers during one held expiration.
Reflex responses that depend on human otolith organ sensitivity were measured before, during and after a 10 day space flight. Otolith-spinal reflexes were elicited by means of sudden, unexpected falls. In weightlessness, 'falls' were achieved using elastic cords running from a torso harness to the floor. Electromyographic (EMG) activity was recorded from gastrocnemius-soleus. The EMG response occurring in the first 100-120 ms of a fall, considered to be predominantly otolith-spinal in origin, decreased in amplitude immediately upon entering weightlessness, and continued to decline throughout the flight, especially during the first two mission days. The response returned to normal before the first post-flight testing session. The results suggest that information coming from the otolith organs is gradually ignored by the nervous system during prolonged space flight, although the possibility that otolith-spinal reflexes are decreased independent of other otolith output pathways cannot by ruled out.
The dynamic response properties of horizontal vestibulo-ocular reflex (VOR) and optokinetic reflex (OKR) were characterized in 216 human subjects ranging in age from 7 to 81 years. The object of this cross-sectional study was to determine the effects of aging on VOR and OKR reflex dynamics, and to identify the distributions of parameters which describe VOR and OKR responses to pseudorandom stimuli in a putatively normal population. In general, VOR and OKR response parameters changed in a manner consistent with declining function with increasing age. For the VOR this was reflected in declining response amplitudes, although the magnitude of the decline was small relative to the variability of the data. For the OKR the lag time of the response, probably associated with the time required for visual information processing, increased linearly with age at a rate of about 1 ms per year.
On earth, the functional stretch reflex is an important component in the maintenance of posture and muscle tone. In parabolic flight experiments, it is evaluated whether the functional stretch reflex, as reflected in the tonic vibration reflex, adjusts appropriately for changes in background gravitoinertial force level. Virtually immediate alterations of appropriate sign occurred.
This investigation was designed to determine how a primitive vertebrate, the bullfrog tadpole, is able to sense and process gravitational stimuli. Because of the phylogenetic similarities of the vestibular systems in all vertebrates, the understanding of the gravitational reflexes in this relatively simple vertebrate should elucidate a skeletal framework on a elementary level, upon which the more elaborate reflexes of higher vertebrates may be constructed. The purpose of this study was to understand how the nervous system of the larval amphibian processes gravitational information. This study involved predominantly electrophysiological investigations of the isolated, alert (forebrain removed) bullfrog tadpole head. The focus of these experiments is threefold: (1) to understand from whole extraocular nerve recordings the signals sent to the eye following static gravitational tilt of the head; (2) to localize neuronal centers responsible for generating these signals through reversible pharmacological ablation of these centers; and (3) to record intracellularly from neurons within these centers in order to determine the single neuron's role in the overall processing of the center. This study has provided information on the mechanisms by which a primitive vertebrate processes gravitational reflexes.
The vomiting reflex may be elicited by a number of different types or classes of stimuli involving many varieties of receptor structures and considerable diversity in afferent pathways and central connections. Central relay or mediating structures thus may vary widely according to the type of initial emetic stimulus. The emetic circuits which have been most completely delineated to date are probably those in which the Chemoreceptor Trigger Zone (CTZ) in the Area Postrema (AP) functions as a key mediating structure. Even in this system, however, there are large gaps in our knowledge of the nerve tracts and central nervous connections involved. Knowledge of most other emetic circuits subserving the emetic reflex resulting from many diverse types of stimuli such, for example, as emotional stress (e.g. psychogenic vomiting, Wruble et al. 1982), pain (e.g. testicular trauma), and chemical or mechanical irritation of the gastrointestinal tract or urinary tract is quite incomplete at this time, thus precluding any very adequate description of their central connections at present. One physiological system, however, which has received considerable attention recently in relation to the vomiting reflex elicited by motion stimuli is the vestibular system. Due to the paucity of data on central nervous connections of several or the non-vestibular types of emetic stimuli cited above, we will devote most of our attention in this brief review to the central connections of the vestibular system which seem likely to be involved in the vomiting response to motion stimuli. However, the latter part of the review will be concerned with the concept of the reticular vomiting centre in relation to the ParviCellular Reticular Formation (PCRF), and will thus probably pertain to all of the many classes of emetic stimuli since it will address the question of the final common emetic pathway.
Reflex responses that depend on human otolith organ sensitivity were measured before, during and after a 10 day space flight. Otolith-spinal reflexes were elicited by means of sudden, unexpected falls. In weightlessness, "falls" were achieved using elastic cords running from a torso harness to the floor. Electromyographic (EMG) activity was recorded from gastrocnemius-soleus. The EMG response occurring in the first 100-120 ms of a fall, considered to be predominantly otolith-spinal in origin, decreased in amplitude immediately upon entering weightlessness, and continued to decline throughout the flight, especially during the first two mission days. The response returned to normal before the first post-flight testing session. The results suggest that information coming from the otolith organs is gradually ignored by the nervous system during prolonged space flight, although the possibility that otolith-spinal reflexes are decreased independent of other otolith output pathways cannot be ruled out.
Two joint ESA-NASA studies are examining changes in otolith-ocular reflexes and motion perception following short duration space flights, and the operational implications of post-flight tilt-translation ambiguity for manual control performance. Vibrotactile feedback of tilt orientation is also being evaluated as a countermeasure to improve performance during a closed-loop nulling task. Data is currently being collected on astronaut subjects during 3 preflight sessions and during the first 8 days after Shuttle landings. Variable radius centrifugation is utilized to elicit otolith reflexes in the lateral plane without concordant roll canal cues. Unilateral centrifugation (400 deg/s, 3.5 cm radius) stimulates one otolith positioned off-axis while the opposite side is centered over the axis of rotation. During this paradigm, roll-tilt perception is measured using a subjective visual vertical task and ocular counter-rolling is obtained using binocular video-oculography. During a second paradigm (216 deg/s, less than 20 cm radius), the effects of stimulus frequency (0.15 - 0.6 Hz) are examined on eye movements and motion perception. A closed-loop nulling task is also performed with and without vibrotactile display feedback of chair radial position. Data collection is currently ongoing. Results to date suggest there is a trend for perceived tilt and translation amplitudes to be increased at the low and medium frequencies on landing day compared to pre-flight. Manual control performance is improved with vibrotactile feedback. One result of this study will be to characterize the variability (gain, asymmetry) in both otolith-ocular responses and motion perception during variable radius centrifugation, and measure the time course of post-flight recovery. This study will also address how adaptive changes in otolith-mediated reflexes correspond to one's ability to perform closed-loop nulling tasks following G-transitions, and whether manual control performance can be improved with vibrotactile feedback of orientation.