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Space adaptation syndrome: multiple etiological factors and individual differences

Space motion sickness is a significant operational concern in the American and Soviet space programs. Nearly 70% of all astronauts and cosmonauts are affected to some degree during their first several days of flight. It is now beginning to appear that space motion sickness like terrestrial motion sickness is the consequence of multiple etiological factors. As we come to understand basic mechanisms of spatial orientation and sensory-motor adaptation we can begin to predict etiological factors in different motion environments. Individuals vary greatly in the extent to which they are susceptible to these different factors. However, individuals seem to be relatively self-consistent in terms of their rates of adaptation to provocative stimulation and their retention of adaptation. Attempts to relate susceptibility to motion sickness during the microgravity phases of parabolic flight maneuvers to vestibular function under 1G and 0G test conditions are described.

short duration↗

Physiological Targets of Artificial Gravity: The Sensory-Motor System

This chapter describes the pros and cons of artificial gravity applications in relation to human sensory-motor functioning in space. Spaceflight creates a challenge for sensory-motor functions that depend on gravity, which include postural balance, locomotion, eye-hand coordination, and spatial orientation. The sensory systems, and in particular the vestibular system, must adapt to weightlessness on entering orbit, and again to normal gravity upon return to Earth. During this period of adaptation, which persists beyond the actual gravity-level transition itself the sensory-motor systems are disturbed. Although artificial gravity may prove to be beneficial for the musculoskeletal and cardiovascular systems, it may well have negative side effects for the neurovestibular system, such as spatial disorientation, malcoordination, and nausea.

Paloski, William↗

Effect of acute exposure to hypergravity (GX vs. GZ) on dynamic cerebral autoregulation

We examined the effects of 30 min of exposure to either +3GX (front-to-back) or +GZ (head-to-foot) centrifugation on cerebrovascular responses to 80 degrees head-up tilt (HUT) in 14 healthy individuals. Both before and after +3 GX or +3 GZ centrifugation, eye-level blood pressure (BP(eye)), end tidal PCO2 (PET(CO2)), mean cerebral flow velocity (CFV) in the middle cerebral artery (transcranial Doppler ultrasound), cerebral vascular resistance (CVR), and dynamic cerebral autoregulatory gain (GAIN) were measured with subjects in the supine position and during subsequent 80 degrees HUT for 30 min. Mean BP(eye) decreased with HUT in both the GX (n = 7) and GZ (n = 7) groups (P < 0.001), with the decrease being greater after centrifugation only in the GZ group (P < 0.05). PET(CO2) also decreased with HUT in both groups (P < 0.01), but the absolute level of decrease was unaffected by centrifugation. CFV decreased during HUT more significantly after centrifugation than before centrifugation in both groups (P < 0.02). However, these greater decreases were not associated with greater increases in CVR. In the supine position after centrifugation compared with before centrifugation, GAIN increased in both groups (P < 0.05, suggesting an autoregulatory deficit), with the change being correlated to a measure of otolith function (the linear vestibulo-ocular reflex) in the GX group (r = 0.76, P < 0.05) but not in the GZ group (r = 0.24, P = 0.60). However, GAIN was subsequently restored to precentrifugation levels during postcentrifugation HUT (i.e., as BP(eye) decreased), suggesting that both types of centrifugation resulted in a leftward shift of the cerebral autoregulation curve. We speculate that this leftward shift may have been due to vestibular activation (especially during +GX) or potentially to an adaptation to reduced cerebral perfusion pressure during +GZ.

NASA Discipline Neuroscience↗

A critical period for functional vestibular development in zebrafish

We have determined a critical period for vestibular development in zebrafish by using a bioreactor designed by NASA to simulate microgravity for cells in culture. A critical period is defined as the briefest period of time during development when stimulus deprivation results in long lasting or permanent sensory deficits. Zebrafish eggs were collected within 3 hours of being laid and fertilized. In experiment 1, eggs were placed in the bioreactor at 3, 24, 30, 36, 48, or 72 hours postfertilization (hPF) and maintained in the bioreactor until 96 hPF. In experiment 2, eggs were placed in the bioreactor immediately after they were collected and maintained in the bioreactor until 24, 36, 48, 60, 66, 72, or 96 hPF. Beginning at 96 hPF, all larvae had their vestibulo-ocular reflexes (VOR) evaluated once each day for 5 days. Only larvae that hatched from eggs that were placed in the bioreactor before 30 hPF in experiment 1 or removed from the bioreactor later than 66 hPF in experiment 2 had VOR deficits that persisted for at least 5 days. These data suggest a critical period for vestibular development in the zebrafish that begins before 30 hPF and ends after 66 hPF. To confirm this, zebrafish eggs were placed in the bioreactor at 24 hPF and removed at 72 hPF. VORs were evaluated in these larvae once each day for 5 days beginning at 96 hPF. These larvae had VOR deficits that persisted for at least 5 days. In addition, larvae that had been maintained in the bioreactor from 24 to 66 hPF or from 30 to 72 hPF, had only temporary VOR deficits. In a final experiment, zebrafish eggs were placed in the bioreactor at 3 hPF and removed at 96 hPF but the bioreactor was turned off from 24 hPF to 72 hPF. These larvae had normal VORs when they were removed from the bioreactor at 96 hPF. Taken as a whole, these data support the idea that there is a critical period for functional maturation of the zebrafish vestibular system. The developmental period identified includes the timeframe during which the vestibular primary afferent neurons are born, innervate their central and peripheral targets, and remodel their central projections. Copyright 2002 Wiley-Liss, Inc.

Non-NASA Center↗

Collaborative research in cardiovascular dynamics and bone elasticity

A collaborative research program covering a variety of topics of biomechanics and biomedical engineering within the fields of cardiovascular dynamics, respiration, bone elasticity and vestibular physiology is described. The goals of the research were to promote: (1) a better understanding of the mechanical behavior of the circulatory system and its control mechanisms; (2) development of noninvasive methods of measuring the changes in the mechanical properties of blood vessels and other cardiovascular parameters in man; (3) application of these noninvasive methods to examine in man the physiological effects of environmental changes, including earth-simulated gravitational changes; and (4) development of in-flight methods for studying the events which lead to post-flight postural hypotension.

Source record↗

Biomedical results of the Skylab Program

Skylab, the fourth in a logical sequence of USA manned space flight projects following Mercury, Gemini and Apollo, presented life scientists with their first opportunity for an in-depth study of man's response to the space environment. Extensive medical investigations were undertaken to increase our understanding of man's adaptation to the space environment and his readaptation to gravity upon return to earth. The flight durations of the three Skylab missions were progressively increased from 28 days to 59 days and, finally, 84 days. The results of these investigations of the various body systems clearly denonstrated that man can adapt to zero gravity and perform useful work during long-duration space flight. However, definite changes (some unexpected) in the vestibular, cardiovascular, musculo-skeletal, renal and electrolyte areas were documented. The most significant were: the occurrence of space motion sickness early in the missions; diminished orthostatic tolerance, both in-flight and post-flight; moderate losses of calcium, phosphorus and nitrogen; and decreased tolerance for exercise post-flight. The mechanisms responsible for these physiological responses must be understood and, if necessary, effective countermeasures developed before man can endure unlimited exposure to space flight.

Michel, E. L.↗

Life sciences space missions. Overview

It has been known for many years that weightlessness induces changes in numerous physiological systems: the cardiovascular system declines in both aerobic capacity and orthostatic tolerance; there is a reduction in fluid and electrolyte balance, hematocrit, and certain immune parameters; bone and muscle mass and strength are reduced; various neurological responses include space motion sickness and posture and gate alterations. These responses are caused by the hypokinesia of weightlessness, the cephalic fluid shift, the unloading of the vestibular system, stress, and the altered temporal environment.

NASA Center HQS↗

Medical Devices Assess, Treat Balance Disorders

You may have heard the phrase as difficult as walking and chewing gum as a joking way of referring to something that is not difficult at all. Just walking, however, is not all that simple physiologically speaking. Even standing upright is an undertaking requiring the complex cooperation of multiple motor and sensory systems including vision, the inner ear, somatosensation (sensation from the skin), and proprioception (the sense of the body s parts in relation to each other). The compromised performance of any of these elements can lead to a balance disorder, which in some form affects nearly half of Americans at least once in their lifetimes, from the elderly, to those with neurological or vestibular (inner ear) dysfunction, to athletes with musculoskeletal injuries, to astronauts returning from space. Readjusting to Earth s gravity has a significant impact on an astronaut s ability to balance, a result of the brain switching to a different "model" for interpreting sensory input in normal gravity versus weightlessness. While acclimating, astronauts can experience headaches, motion sickness, and problems with perception. To help ease the transition and study the effects of weightlessness on the body, NASA has conducted many investigations into post-flight balance control, realizing this research can help treat patients with balance disorders on Earth as well. In the 1960s, the NASA-sponsored Man Vehicle Laboratory at the Massachusetts Institute of Technology (MIT) studied the effects of prolonged space flight on astronauts. The lab s work intrigued MIT doctoral candidate Lewis Nashner, who began conducting NASA-funded research on human movement and balance under the supervision of Dr. Larry Young in the MIT Department of Aeronautics and Astronautics. In 1982, Nashner s work resulted in a noninvasive clinical technique for assessing the cooperative systems that allow the body to balance, commonly referred to as computerized dynamic posturography (CDP). CDP employs a series of dynamic protocols to isolate and assess balance function deficiencies. The technology was based on Nashner s novel, engineering-inspired concept of balance as an adaptable collaboration between multiple sensory and motor systems. CDP proved useful not only for examining astronauts, but for anyone suffering from balance problems. Today, CDP is the standard medical tool for objectively evaluating balance control.

Source record↗

Ambiguous Tilt and Translation Motion Cues in Astronauts after Space Flight

Adaptive changes during space flight in how the brain integrates vestibular cues with visual, proprioceptive, and somatosensory information can lead to impaired movement coordination, vertigo, spatial disorientation, and perceptual illusions following transitions between gravity levels. This joint ESA-NASA pre- and post-flight experiment is designed to examine both the physiological basis and operational implications for disorientation and tilt-translation disturbances in astronauts following short-duration space flights. The first specific aim is to examine the effects of stimulus frequency on adaptive changes in eye movements and motion perception during independent tilt and translation motion profiles. Roll motion is provided by a variable radius centrifuge. Pitch motion is provided by NASA's Tilt-Translation Sled in which the resultant gravitoinertial vector remains aligned with the body longitudinal axis during tilt motion (referred to as the Z-axis gravitoinertial or ZAG paradigm). We hypothesize that the adaptation of otolith-mediated responses to these stimuli will have specific frequency characteristics, being greatest in the mid-frequency range where there is a crossover of tilt and translation. The second specific aim is to employ a closed-loop nulling task in which subjects are tasked to use a joystick to null-out tilt motion disturbances on these two devices. The stimuli consist of random steps or sum-of-sinusoids stimuli, including the ZAG profiles on the Tilt-Translation Sled. We hypothesize that the ability to control tilt orientation will be compromised following space flight, with increased control errors corresponding to changes in self-motion perception. The third specific aim is to evaluate how sensory substitution aids can be used to improve manual control performance. During the closed-loop nulling task on both devices, small tactors placed around the torso vibrate according to the actual body tilt angle relative to gravity. We hypothesize that performance on the closed-loop tilt control task will be improved with this tactile display feedback of tilt orientation. The current plans include testing on eight crewmembers following Space Shuttle missions or short stay onboard the International Space Station. Measurements are obtained pre-flight at L-120 (plus or minus 30), L-90 (plus or minus 30), and L-30, (plus or minus 10) days and post-flight at R+0, R+1, R+2 or 3, R+4 or 5, and R+8 days. Pre-and post-flight testing (from R+1 on) is performed in the Neuroscience Laboratory at the NASA Johnson Space Center on both the Tilt-Translation Device and a variable radius centrifuge. A second variable radius centrifuge, provided by DLR for another joint ESA-NASA project, has been installed at the Baseline Data Collection Facility at Kennedy Space Center to collect data immediately after landing. ZAG was initiated with STS-122/1E and the first post-flight testing will take place after STS-123/1JA landing.

Clement, G.↗

The Turbolift: Linear Sled Hybrid Artificial Gravity Concept

Future crewed space exploration missions into deep space will require enhanced countermeasure technologies to ensure astronaut health. One such hazard is extended exposure to reduced gravity levels (i.e., microgravity, lunar gravity, or Martian gravity). Reduced gravity negatively impacts many physiological systems, leading to hydrostatic intolerance, musculoskeletal atrophy, sensorimotor impairment, bone demineralization, cardiovascular deconditioning, and visual alterations. Various countermeasures have been employed for mitigating these effects, such as exercise, pharmaceuticals, diet, and fluid loading. However, these approaches treat individual symptoms, such that each physiological system is addressed with typically one countermeasure. An alternative to this approach is artificial gravity (AG), which promises to be a holistic, comprehensive countermeasure. The traditional approach to creating AG is through centrifugation. However, centrifugation is not a "pure" form of AG and typically includes the drawbacks of Coriolis forces, gravity gradients, and vestibular cross-coupled illusions.As an alternative, we have proposed a Linear Sled Hybrid (LSH) AG system to mitigate astronauts' physiological deconditioning. This system functions by applying pure linear acceleration to produce footward loading. There is a half rotation (180_) to reorient the rider between acceleration and deceleration phases, such that the loading remains footward, as when standing on Earth. The rotation also provides some footward acceleration to the lower body through centripetal acceleration; hence the "hybrid" aspect of the design. At the end of the deceleration, the rider than accelerates back in the opposite direction and the sequence repeats.This proposed system could be integrated with future crewed space vehicles in a variety of manners.

Gruber, Jason↗

The Effect of Spaceflight on the Ultrastructure of the Cerebellum

In weightlessness, astronauts and cosmonauts may experience postural illusions as well as motion sickness symptoms known as the space adaptation syndrome. Upon return to Earth, they have irregularities in posture and balance. The adaptation to microgravity and subsequent re-adaptation to Earth occurs over several days. At the cellular level, a process called neuronal plasticity may mediate this adaptation. The term plasticity refers to the flexibility and modifiability in the architecture and functions of the nervous system. In fact, plastic changes are thought to underlie not just behavioral adaptation, but also the more generalized phenomena of learning and memory. The goal of this experiment was to identify some of the structural alterations that occur in the rat brain during the sensory and motor adaptation to microgravity. One brain region where plasticity has been studied extensively is the cerebellar cortex-a structure thought to be critical for motor control, coordination, the timing of movements, and, most relevant to the present experiment, motor learning. Also, there are direct as well as indirect connections between projections from the gravity-sensing otolith organs and several subregions of the cerebellum. We tested the hypothesis that alterations in the ultrastructural (the structure within the cell) architecture of rat cerebellar cortex occur during the early period of adaptation to microgravity, as the cerebellum adapts to the absence of the usual gravitational inputs. The results show ultrastructural evidence for neuronal plasticity in the central nervous system of adult rats after 24 hours of spaceflight. Qualitative studies conducted on tissue from the cerebellar cortex (specifically, the nodulus of the cerebellum) indicate that ultrastructural signs of plasticity are present in the cerebellar zones that receive input from the gravity-sensing organs in the inner ear (the otoliths). These changes are not observed in this region in cagematched ground control animals. The specific changes include the formation of lamellar bodies, profoundly enlarged Purkinje cell mitochondria, the presence of inter-neuronal cellular protrusions in the molecular layer, and signs of degeneration in the distal dendrites of the Purkinje cells. Since these morphologic signs are not apparent in the control animals, they are not likely to be due to caging or tissue processing effects. The particular nature of the structural alterations in the nodulus, most notably the formation of lamellar bodies and the presence of degeneration, further suggests that excitotoxicity (damaging overstimulation of neurons) may play a role in the short-term neural response to spaceflight. These findings suggest a structural basis for the neuronal and synaptic plasticity accompanying the central nervous system response to altered gravity and help identify the cellular bases underlying the vestibular abnormalities experienced by astronauts during periods of adaptation and re-adaptation to different gravitational forces. Also, since the short- and long-term changes in neural structure occurring during such periods of adaptation resemble the neuronal alterations that occur in some neurologic disorders such as stroke, these findings may offer guidance in the development of strategies for rehabilitation and treatment of such disorders.

Holstein, Gay R.↗

Short-term adaptation of the VOR: non-retinal-slip error signals and saccade substitution

We studied short-term (30 min) adaptation of the vestibulo-ocular reflex (VOR) in five normal humans using a "position error" stimulus without retinal image motion. Both before and after adaptation a velocity gain (peak slow-phase eye velocity/peak head velocity) and a position gain (total eye movement during chair rotation/amplitude of chair motion) were measured in darkness using search coils. The vestibular stimulus was a brief ( approximately 700 ms), 15 degrees chair rotation in darkness (peak velocity 43 degrees /s). To elicit adaptation, a straight-ahead fixation target disappeared during chair movement and when the chair stopped the target reappeared at a new location in front of the subject for gain-decrease (x0) adaptation, or 10 degrees opposite to chair motion for gain-increase (x1.67) adaptation. This position-error stimulus was effective at inducing VOR adaptation, though for gain-increase adaptation the primary strategy was to substitute augmenting saccades during rotation while for gain-decrease adaptation both corrective saccades and a decrease in slow-phase velocity occurred. Finally, the presence of the position-error signal alone, at the end of head rotation, without any attempt to fix upon it, was not sufficient to induce adaptation. Adaptation did occur, however, if the subject did make a saccade to the target after head rotation, or even if the subject paid attention to the new location of the target without actually looking at it.

Non-NASA Center↗

Postnatal development under conditions of simulated weightlessness and space flight

The adaptability of the developing nervous system to environmental influences and the mechanisms underlying this plasticity has recently become a subject of interest in space neuroscience. Ground studies on neonatal rats using the tail suspension model of weightlessness have shown that the force of gravity clearly influences the events underlying the postnatal development of motor function. These effects depend on the age of the animal, duration of the perturbation and the motor function studied. A nine-day flight study has shown that a dam and neonates can develop under conditions of space flight. The motor function of the flight animals after landing was consistent with that seen in the tail suspension studies, being marked by limb joint extension. However, there were expected differences due to: (1) the unloading of the vestibular system in flight, which did not occur in the ground-based experiments; (2) differences between flight and suspension durations; and (3) the inability to evaluate motor function during the flight. The next step is to conduct experiments in space with the flexibility and rigor that is now limited to ground studies: an opportunity offered by the International Space Station. Copyright 1998 Published by Elsevier Science B.V.

Non-NASA Center↗

Field Test: Results from the One Year Mission

The One Year Mission was designed to aid in determining the effect that extending the duration on orbit aboard the International Space Station (ISS) would have on a number of biological and physiological systems. Two crewmembers were selected to participate in this endeavor, one U.S. On-Orbit Segment (USOS) astronaut and one Russian cosmonaut. The Neuroscience and Cardiovascular and Vision Laboratories at the Johnson Space Center and the Sensory-Motor and Countermeasures Division within the Institute for Biomedical Problems were selected to investigate vestibular, sensorimotor and cardiovascular function with the two long-duration crewmembers using the established methodology developed for the Field Test (FT).

Reschke, M. F.↗

Somatosensory motion after-effect following earth-horizontal rotation about the Z-axis - A new illusion

During rotation about the Z-axis while recumbent one is exposed to a changing pattern of pressure cues over the body surface. If the body is only loosely padded in the experimental apparatus, then apparent motion of part of the body surface may be experienced sometime after rotation has been terminated. This somatosensory motion aftereffect of opposite sign is temporarily abolished if one looks at the affected body area, but is again re-experienced when the gaze is shifted elsewhere. The similarity of this motion aftereffect to those contingent on vestibular and visual stimulation is discussed.

Lackner, J. R.↗

Spatial orientation - Visual-vestibular-somatic interaction

The compensation signals from somatosensory and vestibular receptors, which act to compensate for disturbances produced by the displacement of our eyes relative to other parts of our bodies and for visual disturbances such as tilted frames or moving visual fields, are investigated. Disturbances were evoked by tilting the head and by rotating a large visual display, while compensation signals related to gravity were altered by placing the subjects horizontally on a board or seating them vertically. The first experiment studied the effects of visual disturbance on the ability of supine observers to set a line to the longitudinal body axis while the head was tilted toward one shoulder or while the head was straight. Results showed that the effects of the visual disturbance were greater when the head was tilted than when it was straight, which indicates that the effects of visual disturbance were greater for a task that required compensation. The second experiment compared the performance of supine and erect observers. No differences were found between the performance of observers on a task requiring the use of compensation signals under these two conditions, which suggests that the enrichment of compensatory signals did not reduce the effects of visual disturbancs.

Parker, D. E.↗

Spatial Orientation and Balance Control Changes Induced by Altered Gravito-Inertial Force Vectors

Seventeen healthy and eight vestibular deficient subjects were exposed to an interaural centripetal acceleration of 1 G (resultant 45 deg roll tilt of 1.4 G) on a 0.8 meter radius centrifuge for a period of 90 minutes in the dark. The subjects sat with head fixed upright, except every 4 of 10 minutes when instructed to rotate their head so that their nose and eyes pointed towards a visual point switched on every 3 to 5 seconds at random places (within +/- 30 deg) in the Earth horizontal plane. Motion sickness caused some subjects to limit their head movements during significant portions of the 90 minute period, and led three normal subjects to stop the test earlier. Eye movements, including directed saccades for subjective Earth- and head-referenced planes, were recorded before, during, and immediately after centrifugation using electro-oculography. Postural stability measurements were made before and within ten minutes after centrifugation. In normal subjects, postural sway and multisegment body kinematics were gathered during an eyes-closed head movement cadence (sway-referenced support platform), and in response to translational/rotational platform perturbations. A significant increase in postural sway, segmental motion amplitude and hip frequency was observed after centrifugation. This effect was short-lived, with a recovery time of several postural test trials. There were also asymmetries in the direction of post-centrifugation center of sway and head tilt which depended on the subject's orientation during the centrifugation adaptation period (left ear or right ear out). To delineate the effect of the magnitude of the gravito-inertial vector versus its direction during the adaptive centrifugation period, we tilted eight normal subjects in the roll axis at a 45 deg angle in the dark for 90 minutes without rotational motion. Their postural responses did not change following the period of tilt. Based on verbal reports, normal subjects overestimated roll-tilt during 90 minutes of both tilt and centrifugation stimuli. Subjective estimates of head-horizontal, provided by directed saccades, revealed significant errors after approximately 30 minutes that tended to increase only in the group who underwent centrifugation. Immediately after centrifugation, subjects reported feeling tilted on average 10 degrees in the opposite direction, which was in agreement with the direction of their earth-directed saccades. In vestibular deficient (VD) subjects, postural sway was measured using a sway-referenced or earth-fixed support surface, and with or without a head movement sequence. 'Me protocol was selected for each patient during baseline testing, and corresponded to the most challenging condition in which the patient was able to maintain balance with eyes closed. Bilaterally VD subjects showed no postural decrement after centrifugation, while unilateral VD subjects had varying degrees of decrement. Unilateral VD subjects were tested twice; they underwent centrifugation both with right ear out and left ear out. Their post-centrifuation center of sway shifted at right angles depending on the centrifuge GIF orientation. Bilateral VD subjects bad shifts as well, but no consistent directional trend. VD subjects underestimated roll-tilt during centrifugation, These results suggest that orientation of the gravito-inertial vector and its magnitude arc both used by the central nervous system for calibration of multiple orientation systems. A change in the background gravito-inertial force (otolith input) can rapidly initiate postural and perceptual adaptation in several sensorimotor systems, independent of a structured visual surround.

Kaufman, Galen D.↗

Role of the medial medullary reticular formation in relaying vestibular signals to the diaphragm and abdominal muscles

Changes in posture can affect the resting length of respiratory muscles, requiring alterations in the activity of these muscles if ventilation is to be unaffected. Recent studies have shown that the vestibular system contributes to altering respiratory muscle activity during movement and changes in posture. Furthermore, anatomical studies have demonstrated that many bulbospinal neurons in the medial medullary reticular formation (MRF) provide inputs to phrenic and abdominal motoneurons; because this region of the reticular formation receives substantial vestibular and other movement-related input, it seems likely that medial medullary reticulospinal neurons could adjust the activity of respiratory motoneurons during postural alterations. The objective of the present study was to determine whether functional lesions of the MRF affect inspiratory and expiratory muscle responses to activation of the vestibular system. Lidocaine or muscimol injections into the MRF produced a large increase in diaphragm and abdominal muscle responses to vestibular stimulation. These vestibulo-respiratory responses were eliminated following subsequent chemical blockade of descending pathways in the lateral medulla. However, inactivation of pathways coursing through the lateral medulla eliminated excitatory, but not inhibitory, components of vestibulo-respiratory responses. The simplest explanation for these data is that MRF neurons that receive input from the vestibular nuclei make inhibitory connections with diaphragm and abdominal motoneurons, whereas a pathway that courses laterally in the caudal medulla provides excitatory vestibular inputs to these motoneurons.

Non-NASA Center↗