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At least 127 records · Page 7

Investigation of otolith responses using ground based vestibular research facility

The general goal was to examine tilt sensitivity of horizontal semicircular canal afferents. Computer programs were tested which controlled the short axis centrifuge at the Vestibular Research Facility, acquired action potentials and produced data reduction analyses including histograms and gain and phase calculations. A pre-amplifier was also developed for the acquisition of action potentials. The data were gathered that can be used to contribute toward the understanding of the tilt sensitivity of semicircular canal afferents in the unanesthetized gerbil preparation.

Correia, Manning J.

Antagonistic otolith-visual units in cat vestibular nuclei

The nature of neural coding of visual (Vis) and vestibular (Vst) information on translational motion in the region of the vestibular nuclei was investigated using extracellular single-unit recordings in alert adult cats. Responses were recorded and averaged over 60 cycles of stimulation in the vertical and horizontal planes, which included the Vst (movement of the animal in the dark), Vis (movement within lighted visual surround), and combined Vis and Vst (movement of the animal within the lighted stationary visual surround). Data are reported on responses to stimulations along the axis showing maximal sensitivity. A small number of units were identified that showed an antagonistic relationship between their Vis and Vst responses (since they were maximally excited by Vis and by Vst stimulations in the same direction). Results suggest that antagonistic units may belong to an infrequently encountered, but functionally distinct, class of neurons.

Daunton, Nancy G.

Ambiguous Tilt and Translation Motion Cues after Space Flight and Otolith Assessment during Post-Flight Re-Adaptation

Adaptive changes during space flight in how the brain integrates vestibular cues with other sensory information can lead to impaired movement coordination, vertigo, spatial disorientation and perceptual illusions following Gtransitions. These studies are designed to examine both the physiological basis and operational implications for disorientation and tilt-translation disturbances following short duration space flights.

Wood, Scott J.

Development of Vestibular Stochastic Resonance as a Sensorimotor Countermeasure: Improving Otolith Ocular and Motor Task Responses

Astronauts experience disturbances in sensorimotor function after spaceflight during the initial introduction to a gravitational environment, especially after long-duration missions. Stochastic resonance (SR) is a mechanism by which noise can assist and enhance the response of neural systems to relevant, imperceptible sensory signals. We have previously shown that imperceptible electrical stimulation of the vestibular system enhances balance performance while standing on an unstable surface. The goal of our present study is to develop a countermeasure based on vestibular SR that could improve central interpretation of vestibular input and improve motor task responses to mitigate associated risks.

Mulavara, Ajitkumar

Vestibular convergence patterns in vestibular nuclei neurons of alert primates

Sensory signal convergence is a fundamental and important aspect of brain function. Such convergence may often involve complex multidimensional interactions as those proposed for the processing of otolith and semicircular canal (SCC) information for the detection of translational head movements and the effective discrimination from physically congruent gravity signals. In the present study, we have examined the responses of primate rostral vestibular nuclei (VN) neurons that do not exhibit any eye movement-related activity using 0.5-Hz translational and three-dimensional (3D) rotational motion. Three distinct neural populations were identified. Approximately one-fourth of the cells exclusively encoded rotational movements (canal-only neurons) and were unresponsive to translation. The canal-only central neurons encoded head rotation in SCC coordinates, exhibited little orthogonal canal convergence, and were characterized with significantly higher sensitivities to rotation as compared to primary SCC afferents. Another fourth of the neurons modulated their firing rates during translation (otolith-only cells). During rotations, these neurons only responded when the axis of rotation was earth-horizontal and the head was changing orientation relative to gravity. The remaining one-half of VN neurons were sensitive to both rotations and translations (otolith + canal neurons). Unlike primary otolith afferents, however, central neurons often exhibited significant spatiotemporal (noncosine) tuning properties and a wide variety of response dynamics to translation. To characterize the pattern of SCC inputs to otolith + canal neurons, their rotational maximum sensitivity vectors were computed using exclusively responses during earth-vertical axis rotations (EVA). Maximum sensitivity vectors were distributed throughout the 3D space, suggesting strong convergence from multiple SCCs. These neurons were also tested with earth-horizontal axis rotations (EHA), which would activate both vertical canals and otolith organs. However, the recorded responses could not be predicted from a linear combination of EVA rotational and translational responses. In contrast, one-third of the neurons responded similarly during EVA and EHA rotations, although a significant response modulation was present during translation. Thus this subpopulation of otolith + canal cells, which included neurons with either high- or low-pass dynamics to translation, appear to selectively ignore the component of otolith-selective activation that is due to changes in the orientation of the head relative to gravity. Thus contrary to primary otolith afferents and otolith-only central neurons that respond equivalently to tilts relative to gravity and translational movements, approximately one-third of the otolith + canal cells seem to encode a true estimate of the translational component of the imposed passive head and body movement.

Non-NASA Center

Spatiotemporal processing of linear acceleration: primary afferent and central vestibular neuron responses

Spatiotemporal convergence and two-dimensional (2-D) neural tuning have been proposed as a major neural mechanism in the signal processing of linear acceleration. To examine this hypothesis, we studied the firing properties of primary otolith afferents and central otolith neurons that respond exclusively to horizontal linear accelerations of the head (0.16-10 Hz) in alert rhesus monkeys. Unlike primary afferents, the majority of central otolith neurons exhibited 2-D spatial tuning to linear acceleration. As a result, central otolith dynamics vary as a function of movement direction. During movement along the maximum sensitivity direction, the dynamics of all central otolith neurons differed significantly from those observed for the primary afferent population. Specifically at low frequencies (</=0.5 Hz), the firing rate of the majority of central otolith neurons peaked in phase with linear velocity, in contrast to primary afferents that peaked in phase with linear acceleration. At least three different groups of central response dynamics were described according to the properties observed for motion along the maximum sensitivity direction. "High-pass" neurons exhibited increasing gains and phase values as a function of frequency. "Flat" neurons were characterized by relatively flat gains and constant phase lags (approximately 20-55 degrees ). A few neurons ("low-pass") were characterized by decreasing gain and phase as a function of frequency. The response dynamics of central otolith neurons suggest that the approximately 90 degrees phase lags observed at low frequencies are not the result of a neural integration but rather the effect of nonminimum phase behavior, which could arise at least partly through spatiotemporal convergence. Neither afferent nor central otolith neurons discriminated between gravitational and inertial components of linear acceleration. Thus response sensitivity was indistinguishable during 0.5-Hz pitch oscillations and fore-aft movements. The fact that otolith-only central neurons with "high-pass" filter properties exhibit semicircular canal-like dynamics during head tilts might have important consequences for the conclusions of previous studies of sensory convergence and sensorimotor transformations in central vestibular neurons.

NASA Discipline Neuroscience

The Influence of Gravito-Inertial Force on Sensorimotor Integration and Reflexive Responses

Sensorimotor responses (e.g.. eye movements, spinal reflexes, etc depend upon the interpretation of the neural signals from the sensory systems. Since neural signals from the otoliths may represent either tilt (gravity) or translation (linear inertial force), sensory signals from the otolith organs are necessarily somewhat ambiguous. Therefore. the neural responses to changing otolith signals depend upon the context of the stimulation (e.g- active vs. passive, relative orientation of gravity, etc.) as well as upon other sensory signals (e.g., vision. canals, etc.). This session will focus upon the -role -played by the sensory signals from the otolith organs in producing efficient sensorimotor and behavioral responses. Curthoys will show the influence of the peripheral anatomy and physiology. Tomko will discuss the influence of tilt and translational otolith signals on eye movements. Merfeld will demonstrate the rate otolith organs play during the interaction of sensory signals from the canals and otoliths. Watt will show the influence of the otoliths on spinal/postural responses. Guedry will discuss the contribution of vestibular information to "path of movement"' perception and to the development of a stable vertical reference. Sensorimotor responses to the ambiguous inertial force stimulation provide an important tool to investigate how the nervous system processes patterns of sensory information and yields functional sensorimotor responses.

Curthoys, Ian S.

Resolution of sensory ambiguities for gaze stabilization requires a second neural integrator

The ability to simultaneously move in the world and maintain stable visual perception depends critically on the contribution of vestibulo-ocular reflexes (VORs) to gaze stabilization. It is traditionally believed that semicircular canal signals drive compensatory responses to rotational head disturbances (rotational VOR), whereas otolith signals compensate for translational movements [translational VOR (TVOR)]. However, a sensory ambiguity exists because otolith afferents are activated similarly during head translations and reorientations relative to gravity (i.e., tilts). Extra-otolith cues are, therefore, necessary to ensure that dynamic head tilts do not elicit a TVOR. To investigate how extra-otolith signals contribute, we characterized the temporal and viewing distance-dependent properties of a TVOR elicited in the absence of a lateral acceleration stimulus to the otoliths during combined translational/rotational motion. We show that, in addition to otolith signals, angular head position signals derived by integrating sensory canal information drive the TVOR. A physiological basis for these results is proposed in a model with two distinct integration steps. Upstream of the well known oculomotor velocity-to-position neural integrator, the model incorporates a separate integration element that could represent the "velocity storage integrator," whose functional role in the oculomotor system has so far remained controversial. We propose that a key functional purpose of the velocity storage network is to temporally integrate semicircular canal signals, so that they may be used to extract translation information from ambiguous otolith afferent signals in the natural and functionally relevant bandwidth of head movements.

Non-NASA Center

Response of pontomedullary reticulospinal neurons to vestibular stimuli in vertical planes. Role in vertical vestibulospinal reflexes of the decerebrate cat

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.

NASA Discipline Neuroscience

Neurophysiological study on visuo-vestibular control of posture and movement in fish during adaptation to weightlessness

We can stand upright and walk smoothly without paying any particular attention to it. This is because we have established in ourselves an integration center that controls our body subconsciously in response to input from eyes, muscles, joints, foot soles, and also from the gravity sensor in the inner ear (the otolith organ). It has been shown that the cerebellum plays an important role for the establishment of the integration center and that the control pattern is comparable to that of a highly sophisticated computer system. The programming for the control, however, may well be acquired for the 1-g ground condition and does not cover the 0-g in space. Although each of the above organs function as it does on the ground, the signal pattern sent to the center must be different under 0-g and, in addition, complementary signals from the otolith organ are missing, leading to confusion in the integration center and causing a variety of symptoms similar to those of car-sickness or sea-sickness. After exposure to microgravity an immediate process of re-programming will begin and be completed in 2-4 days. There is strong supporting evidence for this sensory conflict theory as an explanation for space motion sickness (SMS) episodes. Fish were selected as test organisms for this investigation because they swim around freely in three dimensions and have well-developed organs for vision and gravity detection. They also have an innate nature to orient their back toward a light source. Actually, on the ground, the fish tilts its vertical axis toward the light when illuminated laterally, and the tilt angle is a function of the intensity of light and the magnitude of gravity, while its posture is completely light-dependent in the low-gravity environment produced by aircraft parabolic flight or when the otolith organs are removed. This implies that fish posture is entirely under visual and otolithic control. In this case, the cerebellum will also contribute to the control. In the space shuttle experiment, two fish (one with otolith-removed and the other with intact otoliths) are onboard for 7 days. The arrangement of the experiment is shown. Lamps illuminate each fish alternately from different directions at a duration of 20 seconds for 10 minutes totally, twice a day. The video images and the brain waves from the cerebellum are analyzed later. If the sensory conflict theory is acceptable, then the confusion at the onset and the following recovery process will be manifested both in the light-dependent behavior and also in the cerebellar activity. In addition, there should be some variation in response between the two fish. Even if the results are positive in the present fish experiment, they would not be extensive enough to allow us to understand the complete mechanisms of SMS. However, what is currently needed is a collection of evidence based on animal experiments on which to base future investigations. Countermeasures for SMS are mandatory since SMS-induced vomiting occurring within a space suit could be fatal for a crew member during extravehicular activities. We expect that countermeasures will be developed before the space station era starts, and that the results of these studies will be applied to improving human welfare.

Mori, Shigeo

Vestibular afferent responses to linear accelerations in the alert squirrel monkey

The spontaneous activity of 40 otolith afferents and 44 canal afferents was recorded in 4 alert, intact squirrel monkeys. Polarization vectors and response properties of otolith afferents were determined during static re-orientations relative to gravity and during Earth-horizontal, sinusoidal, linear oscillations. Canal afferents were tested for sensitivity to linear accelerations. For regular otolith afferents, a significant correlation between upright discharge rate and sensitivity to dynamic acceleration in the horizontal plane was observed. This correlation was not present in irregular units. The sensitivity of otolith afferents to both static tilts and dynamic linear acceleration was much greater in irregularly discharging units than in regularly discharging units. The spontaneous activity and static and dynamic response properties of regularly discharging otolith afferents were similar to those reported in barbiturate-anesthetized squirrel monkeys. Irregular afferents also had similar dynamic response properties when compared to anesthetized monkeys. However, this sample of irregular afferents in alert animals had higher resting discharge rates and greater sensitivity to static tilts. The majority of otolith polarization vectors were oriented near the horizontal in the plane of the utricular maculae; however, directions of maximum sensitivity were different during dynamic and static testing. Canal afferents were not sensitive to static tilts or linear oscillations of the head.

Somps, Christopher J.

Formation of otoconia in the Japanese red-bellied newt, Cynops pyrrhogaster

Pre-mated adult female newts and fertilized eggs will be flown on the International Microgravity Laboratory-2 flight, in 1994. One objective of the flight will be to observe the influence of microgravity on the development of the gravity-sensing organs in the inner ear. These organs contain sensory hair cells covered by a layer of dense stones (otoconia). Gravity and linear acceleration exert forces on these masses, leading to excitation of the nerve fibers innervating the hair cells. If the production of the otoliths is regulated to reach an optimal weight, their development might be abnormal in microgravity. Ground-based control experiments are reported describing the developmental sequence in which both the otoliths and their associated sensory epithelium and the semicircular canals appear and develop. Three-dimensional reconstruction of serial sections through the otic vesicle of newt embryos at stages 31 through 58 demonstrate the first appearance, relative position and growth of the otoliths. Reports of experiments in which fertilized frog eggs were flown on a Russian Cosmos mission conclude that the utricular otolith is increased in volume, whereas the saccular otolith maintains normal size, suggesting that at least in the utricle, the weight of the otolith might be regulated.

Wiederhold, M. L.

Development of the gravity-sensing organs in the Japanese red-bellied newt, Cynops pyrrhogaster

Pre-mated adult female newts and fertilized eggs will be flown on the International Microgravity Laboratory-2 flight, schedule for 1994. One objective of the flight will be to observe the influence of microgravity on the development of the gravity-sensing organs in the inner ear. These organs contain sensory hair cells covered by a layer of dense stones (otoliths). Gravity and linear acceleration exert forces on these masses, leading to excitation of the nerve fibers innervating the hair cells. If the production of the otoliths is regulated to reach an optimal weight, their development would be abnormal in microgravity. Ground-based control experiments are reported describing the developmental sequence in which the otoliths and their associated sensory epithelium appear and increase in size. Three-dimensional reconstruction of serial sections through the otic vesicle of newt embryos at stages 31 through 40 demonstrate the first appearance, relative position and growth of the otoliths. In adult newts, the otoconia in the utricle appear similar to mammalian otoconia, which are composed of calcite. The newt saccular otoconia are at least 99% aragonite, as is found in most aquatic species. Reports of experiments in which fertilized frog eggs were flown on a Russian Cosmos mission conclude that the utricular otolith is increased in volume, whereas the saccular otolith maintains normal size, suggesting that at least the utricular weight might be regulated.

Wiederhold, Michael L.

Early Development of Gravity-Sensing Organs in Microgravity

Most animals have organs that sense gravity. These organs use dense stones (called otoliths or statoconia), which rest on the sensitive hairs of specialized gravity- and motion-sensing cells. The weight of the stones bends the hairs in the direction of gravitational pull. The cells in turn send a coded representation of the gravity or motion stimulus to the central nervous system. Previous experiments, in which the eggs or larvae of a marine mollusk (Aplysia californica, the sea hare) were raised on a centrifuge, demonstrated that the size of the stones (or test mass) was reduced in a graded manner as the gravity field was increased. This suggests that some control mechanism was acting to normalize the weight of the stones. The experiments described here were designed to test the hypothesis that, during their initial development, the mass of the stones is regulated to achieve a desired weight. If this is the case, we would expect a larger-than-normal otolith would develop in animals reared in the weightlessness of space. To test this, freshwater snails and swordtail fish were studied after spaceflight. The snails mated in space, and the stones (statoconia) in their statocysts developed in microgravity. Pre-mated adult female swordtail fish were flown on the Space Shuttle, and the developing larvae were collected after landing. Juvenile fish, where the larval development had taken place on the ground, were also flown. In snails that developed in space, the total volume of statoconia forming the test mass was 50% greater than in size-matched snails reared in functionally identical equipment on the ground. In the swordtail fish, the size of otoliths was compared between ground- and flight-reared larvae of the same size. For later-stage larvae, the growth of the otolith was significantly greater in the flight-reared fish. However, juvenile fish showed no significant difference in otolith size between flight- and ground-reared fish. Thus, it appears that fish and snails reared in space do produce larger-than-normal otoliths (or their analogs), apparently in an attempt to compensate for the reduced weight of the stones in space. The fish data suggest that there is a critical period during which altered gravity can affect the size of the test mass, since the larval, but not the juvenile, fish showed the changes.

Wiederhold, Michael L.

Influence of vestibular activation on respiration in humans

The purpose of this study was to determine the effects of the semicircular canals and otolith organs on respiration in humans. On the basis of animal studies, we hypothesized that vestibular activation would elicit a vestibulorespiratory reflex. To test this hypothesis, respiratory measures, arterial blood pressure, and heart rate were measured during engagement of semicircular canals and/or otolith organs. Dynamic upright pitch and roll (15 cycles/min), which activate the otolith organs and semicircular canals, increased respiratory rate (Delta2 +/- 1 and Delta3 +/- 1 breaths/min, respectively; P < 0.05). Dynamic yaw and lateral pitch (15 cycles/min), which activate the semicircular canals, increased respiration similarly (Delta3 +/- 1 and Delta2 +/- 1, respectively; P < 0.05). Dynamic chair rotation (15 cycles/min), which mimics dynamic yaw but eliminates neck muscle afferent, increased respiration (Delta3 +/- 1; P < 0.05) comparable to dynamic yaw (15 cycles/min). Increases in respiratory rate were graded as greater responses occurred during upright (Delta5 +/- 2 breaths/min) and lateral pitch (Delta4 +/- 1) and roll (Delta5 +/- 1) performed at 30 cycles/min. Increases in breathing frequency resulted in increases in minute ventilation during most interventions. Static head-down rotation, which activates otolith organs, did not alter respiratory rate (Delta1 +/- 1 breaths/min). Collectively, these data indicate that semicircular canals, but not otolith organs or neck muscle afferents, mediate increased ventilation in humans and support the concept that vestibular activation alters respiration in humans.

NASA Discipline Neuroscience

Otoconia biogenesis, phylogeny, composition and functional attributes

This work consolidates data about these interesting organic crystals of vertebrate inner ears. It addresses 5 aspects of inner ear otoliths not completely understood to date: 1) embryological data that explains the formation of the crystals, 2) the significance of the organic and the inorganic phase of the otolith and the changing patterns of otoconia formation along the evolutionary tree, 3) otoliths contribution for detecting linear acceleration, 4) the effect that altered gravity and aminoglycosides have on the development and adult shape of the crystals, and the evolutionary significance of a changing shape of the crystals from primitive forms (lamprey) to high vertebrate birds and mammals is discussed, 5) functional attributes of the otolithic organs and morphological modifications of the otoliths by physical and chemical insults are presented with an extensive discussion of the most relevant literature published and available to us.

Non-NASA Center

Development of Gravity-Sensing Organs in Altered Gravity Conditions: Opposite Conclusions From an Amphibian and a Molluscan Preparation

Several components of the systems animals use to orient to gravity might develop differently in micrograms. If the growth of the "test masses" on which gravity acts (otoliths, in vertebrates, statoliths or statoconia in most invertebrates) is controlled on the basis of their weight, larger otoliths (or their analogs) would be expected to develop in micrograms. The vestibular systems in animals reared in altered gravity have been studied in several species, with varied results being reported. Early Russian reports of Xenopus larvae reared in space indicated no qualitative differences in the vestibular organs, compared to ground-reared controls. A similar lack of differences in Xenopus were reported. The ultricular otolith was 30% larger in space-reared Xenopus. No differences in saccular otolith volume between centrifuged and control adult rats were found. A delay in otoconial development in chick embryos reared at 2 grams on a centrifuge was reported but in a later report, no differences in otolith weight between 2 grams and control chicks were found. Increased optokinetic responses in flight-reared Xenopus tadpoles, suggesting that the animals reared in the absence of gravity made greater relative use of their visual system, rather than the vestibular system, in orienting to a moving stimulus was reported. To test early Japanese newt, CYnops pyrrhogaster, were maintained in orbit for 15 days on the IML-2 mission in 1994. All specimens reached orbit before any otoconia were formed and all major components of the inner ear were formed by the end of the flight. In ground-based studies of he Aplysia statocyst, the volume of the statolith in embryos and the number statoconia in post-metamorphic animals were compared between 1-gram controls and specimens reared at 2 to 5.7 grams.

Wiederhold, Michael L.