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At least 235 records · Page 13
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NASA NTRS2003

The evolution of concepts of vestibular peripheral information processing: toward the dynamic, adaptive, parallel processing macular model

In a letter to Robert Hooke, written on 5 February, 1675, Isaac Newton wrote "If I have seen further than certain other men it is by standing upon the shoulders of giants." In his context, Newton was referring to the work of Galileo and Kepler, who preceded him. However, every field has its own giants, those men and women who went before us and, often with few tools at their disposal, uncovered the facts that enabled later researchers to advance knowledge in a particular area. This review traces the history of the evolution of views from early giants in the field of vestibular research to modern concepts of vestibular organ organization and function. Emphasis will be placed on the mammalian maculae as peripheral processors of linear accelerations acting on the head. This review shows that early, correct findings were sometimes unfortunately disregarded, impeding later investigations into the structure and function of the vestibular organs. The central themes are that the macular organs are highly complex, dynamic, adaptive, distributed parallel processors of information, and that historical references can help us to understand our own place in advancing knowledge about their complicated structure and functions.

Review, Tutorial↗
NASA NTRS2002

Human otolith-ocular reflexes during off-vertical axis rotation: effect of frequency on tilt-translation ambiguity and motion sickness

The purpose of this study was to examine how the modulation of tilt and translation otolith-ocular responses during constant velocity off-vertical axis rotation varies as a function of stimulus frequency. Eighteen human subjects were rotated in darkness about their longitudinal axis 30 degrees off-vertical at stimulus frequencies between 0.05 and 0.8 Hz. The modulation of torsion decreased while the modulation of horizontal slow phase velocity (SPV) increased with increasing frequency. It is inferred that the ambiguity of otolith afferent information is greatest in the frequency region where tilt (torsion) and translational (horizontal SPV) otolith-ocular responses crossover. It is postulated that the previously demonstrated peak in motion sickness susceptibility during linear accelerations around 0.3 Hz is the result of frequency segregation of ambiguous otolith information being inadequate to distinguish between tilt and translation.

NASA Discipline Neuroscience↗
NASA NTRS2001

Complex vestibular macular anatomical relationships need a synthetic approach

Mammalian vestibular maculae are anatomically organized for complex parallel processing of linear acceleration information. Anatomical findings in rat maculae are provided in order to underscore this complexity, which is little understood functionally. This report emphasizes that a synthetic approach is critical to understanding how maculae function and the kind of information they conduct to the brain.

Non-NASA Center↗
NASA NTRS2000

Ontogeny of vestibular compound action potentials in the domestic chicken

Compound action potentials of the vestibular nerve were measured from the surface of the scalp in 148 chickens (Gallus domesticus). Ages ranged from incubation day 18 (E18) to 22 days posthatch (P22). Responses were elicited using linear acceleration cranial pulses. Response thresholds decreased at an average rate of -0.45 dB/day. The decrease was best fit by an exponential model with half-maturity time constant of 5.1 days and asymptote of approximately -25.9 dB re:1.0 g/ms. Mean threshold approached within 3 dB of the asymptote by ages P6-P9. Similarly, response latencies decreased exponentially to within 3% of mature values at ages beyond P9. The half-maturity time constant for peripheral response peak latencies P1, N1, and P2 was comparable to thresholds and ranged from approximately 4.6 to 6.2 days, whereas central peaks (N2, P3, and N3) ranged from 2.9 to 3.4 days. Latency-intensity slopes for P1, N1, and P2 tended to decrease with age, reaching mature values within approximately 100 hours of hatching. Amplitudes increased as a function of age with average growth rates for response peaks ranging from 0.04 to 0.09 microV/day. There was no obvious asymptote to the growth of amplitudes over the ages studied. Amplitude-intensity slopes also increased modestly with age. The results show that gravity receptors are responsive to transient cranial stimuli as early as E19 in the chicken embryo. The functional response of gravity receptors continues to develop for many days after all major morphological structures are in place. Distinct maturational processes can be identified in central and peripheral neural relays. Functional improvements during maturation may result from refinements in the receptor epithelia, improvements in central and peripheral synaptic transmission, increased neural myelination, as well as changes in the mechanical coupling between the cranium and receptor organ.

NASA Discipline Developmental Biology↗
NASA NTRS1997

Development of gravity-sensing organs in altered gravity

Experiments are described in which the development of the gravity-sensing organs was studied in newt larvae reared in microgravity on the IML-2 mission and in Aplysia embryos and larvae reared on a centrifuge at 1 to 5 g. In Aplysia embryos, the statolith (single dense mass on which gravity and linear acceleration act) was reduced in size in a graded fashion at increasing g. In early post-metamorphic Aplysia or even in isolated statocysts from such animals, the number of statoconia produced is reduced at high g. Newt larvae launched before any of the otoconia were formed and reared for 15 days in microgravity had nearly adult labyrinths at the end of the IML-2 mission. The otoliths of the saccule and utricle were the same size in flight and ground-reared larvae. However, the system of aragonitic otoconia produced in the endolymphatic sac in amphibians was much larger and developed earlier in the flight-reared larvae. At later developmental stages, the aragonitic otoconia enter and fill the saccule. One flight-reared larva was maintained for nine months post-flight and the size of the saccular otolith, as well as the volume of otoconia within the endolymphatic sac, were considerably larger than in age-matched, ground-reared newts. This suggests that rearing in microgravity initiates a process that continues for several months after introduction to 1-g, which greatly increases the volume of otoconia. The flight-reared animal had abnormal posture, pointing its head upward, whereas normal ground-reared newts always keep their head horizontal. This suggests that rearing for even a short period in microgravity can have lasting functional consequences in an animal subsequently reared in 1-g conditions on Earth.

STS-65 Shuttle Project↗
NASA NTRS1990

Vestibular receptor cells and signal detection: bioaccelerometers and the hexagonal sampling of two-dimensional signals

The inner ear contains sensory organs which signal changes in head movement. The vestibular sacs, in particular, are sensitive to linear accelerations. Electron microscopic images have revealed the structure of tiny sensory hair bundles, whose mechanical deformation results in the initiation of neuronal activity and the transmission of electrical signals to the brain. The structure of the hair bundles is shown in this paper to be that of the most efficient two-dimensional phased-array signal processors.

NASA Program Space Physiology and Countermeasures↗
NASA NTRS2000

Changes in ribbon synapses and rough endoplasmic reticulum of rat utricular macular hair cells in weightlessness

This study combined ultrastructural and statistical methods to learn the effects of weightlessness on rat utricular maculae. A principle aim was to determine whether weightlessness chiefly affects ribbon synapses of type II cells, since the cells communicate predominantly with branches of primary vestibular afferent endings. Maculae were microdissected from flight and ground control rat inner ears collected on day 13 of a 14-day spaceflight (F13), landing day (R0) and day 14 postflight (R14) and were prepared for ultrastructural study. Ribbon synapses were counted in hair cells examined in a Zeiss 902 transmission electron microscope. Significance of synaptic mean differences was determined for all hair cells contained within 100 section series, and for a subset of complete hair cells, using SuperANOVA software. The synaptic mean for all type II hair cells of F13 flight rats increased by 100%, and that for complete cells by 200%. Type I cells were less affected, with synaptic mean differences statistically insignificant in complete cells. Synapse deletion began within 8 h upon return to Earth. Additionally, hair cell laminated rough endoplasmic reticulum of flight rats was reversibly disorganized on R0. Results support the thesis that synapses in type II hair cells are uniquely affected by altered gravity. Type II hair cells may be chiefly sensors of gravitational and type I cells of translational linear accelerations.

short duration↗
NASA NTRS1999

Disruption of postural readaptation by inertial stimuli following space flight

Postural instability (relative to pre-flight) has been observed in all shuttle astronauts studied upon return from orbital missions. Postural stability was more closely examined in four shuttle astronaut subjects before and after an 8 day orbital mission. Results of the pre- and post-flight postural stability studies were compared with a larger (n = 34) study of astronauts returning from shuttle missions of similar duration. Results from both studies indicated that inadequate vestibular feedback was the most significant sensory deficit contributing to the postural instability observed post flight. For two of the four IML-1 astronauts, post-flight postural instability and rate of recovery toward their earth-normal performance matched the performance of the larger sample. However, post-flight postural control in one returning astronaut was substantially below mean performance. This individual, who was within normal limits with respect to postural control before the mission, indicated that recovery to pre-flight postural stability was also interrupted by a post-flight pitch plane rotation test. A similar, though less extreme departure from the mean recovery trajectory was present in another astronaut following the same post-flight rotation test. The pitch plane rotation stimuli included otolith stimuli in the form of both transient tangential and constant centripetal linear acceleration components. We inferred from these findings that adaptation on orbit and re-adaptation on earth involved a change in sensorimotor integration of vestibular signals most likely from the otolith organs.

NASA Center JSC↗
NASA NTRS1999

Short latency compound action potentials from mammalian gravity receptor organs

Gravity receptor function was characterized in four mammalian species using far-field vestibular evoked potentials (VsEPs). VsEPs are compound action potentials of the vestibular nerve and central relays that are elicited by linear acceleration ramps applied to the cranium. Rats, mice, guinea pigs, and gerbils were studied. In all species, response onset occurred within 1.5 ms of the stimulus onset. Responses persisted during intense (116 dBSPL) wide-band (50 to 50 inverted question mark omitted inverted question mark000 Hz) forward masking, whereas auditory responses to intense clicks (112 dBpeSPL) were eliminated under the same conditions. VsEPs remained after cochlear extirpation but were eliminated following bilateral labyrinthectomy. Responses included a series of positive and negative peaks that occurred within 8 ms of stimulus onset (range of means at +6 dBre: 1.0 g/ms: P1=908 to 1062 micros, N1=1342 to 1475 micros, P2=1632 to 1952 micros, N2=2038 to 2387 micros). Mean response amplitudes at +6 dBre: 1.0 g/ms ranged from 0.14 to 0.99 microV. VsEP input/output functions revealed latency slopes that varied across peaks and species ranging from -19 to -51 micros/dB. Amplitude-intensity slopes also varied ranging from 0.04 to 0.08 microV/dB for rats and mice. Latency values were comparable to those of birds although amplitudes were substantially smaller in mammals. VsEP threshold values were considerably higher in mammals compared to birds and ranged from -8.1 to -10.5 dBre 1.0 g/ms across species. These results support the hypothesis that mammalian gravity receptors are less sensitive to dynamic stimuli than are those of birds.

Non-NASA Center↗
NASA NTRS1998

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↗
NASA NTRS1998

Three-dimensional organization of otolith-ocular reflexes in rhesus monkeys. III. Responses To translation

The three-dimensional (3-D) properties of the translational vestibulo-ocular reflexes (translational VORs) during lateral and fore-aft oscillations in complete darkness were studied in rhesus monkeys at frequencies between 0.16 and 25 Hz. In addition, constant velocity off-vertical axis rotations extended the frequency range to 0.02 Hz. During lateral motion, horizontal responses were in phase with linear velocity in the frequency range of 2-10 Hz. At both lower and higher frequencies, phase lags were introduced. Torsional response phase changed more than 180 degrees in the tested frequency range such that torsional eye movements, which could be regarded as compensatory to "an apparent roll tilt" at the lowest frequencies, became anticompensatory at all frequencies above approximately 1 Hz. These results suggest two functionally different frequency bandwidths for the translational VORs. In the low-frequency spectrum (<<0.5 Hz), horizontal responses compensatory to translation are small and high-pass-filtered whereas torsional response sensitivity is relatively frequency independent. At higher frequencies however, both horizontal and torsional response sensitivity and phase exhibit a similar frequency dependence, suggesting a common role during head translation. During up-down motion, vertical responses were in phase with translational velocity at 3-5 Hz but phase leads progressively increased for lower frequencies (>90 degrees at frequencies <0.2 Hz). No consistent dependence on static head orientation was observed for the vertical response components during up-down motion and the horizontal and torsional response components during lateral translation. The frequency response characteristics of the translational VORs were fitted by "periphery/brain stem" functions that related the linear acceleration input, transduced by primary otolith afferents, to the velocity signals providing the input to the velocity-to-position neural integrator and the oculomotor plant. The lowest-order, best-fit periphery/brain stem model that approximated the frequency dependence of the data consisted of a second order transfer function with two alternating poles (at 0.4 and 7.2 Hz) and zeros (at 0.035 and 3.4 Hz). In addition to clearly differentiator dynamics at low frequencies (less than approximately 0.5 Hz), there was no frequency bandwidth where the periphery/brain stem function could be approximated by an integrator, as previously suggested. In this scheme, the oculomotor plant dynamics are assumed to perform the necessary high-frequency integration as required by the reflex. The detailed frequency dependence of the data could only be precisely described by higher order functions with nonminimum phase characteristics that preclude simple filtering of afferent inputs and might be suggestive of distributed spatiotemporal processing of otolith signals in the translational VORs.

Non-NASA Center↗
NASA NTRS1998

Vestibular adaptation to space in monkeys

Otolith-induced eye movements of rhesus monkeys were studied before and after the 1989 COSMOS 2044 and the 1992 to 1993 COSMOS 2229 flights. Two animals flew in each mission for approximately 2 weeks. After flight, spatial orientation of the angular vestibulo-ocular reflex was altered. In one animal the time constant of postrotatory nystagmus, which had been shortened by head tilts with regard to gravity before flight, was unaffected by the same head tilts after flight. In another animal, eye velocity, which tended to align with a gravitational axis before flight, moved toward a body axis after flight. This shift of orientation disappeared by 7 days after landing. After flight, the magnitude of compensatory ocular counter-rolling was reduced by about 70% in both dynamic and static tilts. Modulation in vergence in response to naso-occipital linear acceleration during off-vertical axis rotation was reduced by more than 50%. These changes persisted for 11 days after recovery. An up and down asymmetry of vertical nystagmus was diminished for 7 days. Gains of the semicircular canal-induced horizontal and vertical angular vestibulo-ocular reflexes were unaffected in both flights, but the gain of the roll angular vestibulo-ocular reflex was decreased. These data indicate that there are short- and long-term changes in otolith-induced eye movements after adaptation to microgravity. These experiments also demonstrate the unique value of the monkey as a model for studying effects of vestibular adaptation in space. Eye movements can be measured in three dimensions in response to controlled vestibular and visual stimulation, and the results are directly applicable to human beings. Studies in monkeys to determine how otolith afferent input and central processing is altered by adaptation to microgravity should be an essential component of future space-related research.

Cosmos 2229 Project↗
NASA NTRS1998

Physiological principles of vestibular function on earth and in space

Physiological mechanisms underlying vestibular function have important implications for our ability to understand, predict, and modify balance processes during and after spaceflight. The microgravity environment of space provides many unique opportunities for studying the effects of changes in gravitoinertial force on structure and function of the vestibular system. Investigations of basic vestibular physiology and of changes in reflexes occurring as a consequence of exposure to microgravity have important implications for diagnosis and treatment of vestibular disorders in human beings. This report reviews physiological principles underlying control of vestibular processes on earth and in space. Information is presented from a functional perspective with emphasis on signals arising from labyrinthine receptors. Changes induced by microgravity in linear acceleration detected by the vestibulo-ocular reflexes. Alterations of the functional requirements for postural control in space are described. Areas of direct correlation between studies of vestibular reflexes in microgravity and vestibular disorders in human beings are discussed.

Non-NASA Center↗
NASA NTRS1998

The relative roles of the otolith organs and semicircular canals in producing space motion sickness

Inflight and post-landing "immunity" to the "coriolis sickness susceptibility test", observed during the Skylab M131 experiment, suggests that the otolith organs play a major role in space motion sickness (SMS). This view is supported by the report that ocular counter-torsion asymmetries correlate with SMS incidence and severity. Further data indicate that sensory-motor adaptation to microgravity includes a process whereby central interpretation of otolith signals is biased from "tilt" toward translation. However, unexpected responses to linear acceleration suggest the importance of graviceptors distributed throughout the body in addition to the vestibular otolith organs. Research is needed to assess distributed graviceptor effects.

Non-NASA Center↗
NASA NTRS1997

Short latency vestibular evoked potentials in the Japanese quail (Coturnix coturnix japonica)

Short-latency vestibular-evoked potentials to pulsed linear acceleration were characterized in the quail. Responses occurred within 8 ms following the onset of stimuli and were composed of a series of positive and negative peaks. The latencies and amplitudes of the first four peaks were quantitatively characterized. Mean latencies at 1.0 g ms-1 ranged from 1265 +/- 208 microseconds (P1, N = 18) to 4802 +/- 441 microseconds (N4, N = 13). Amplitudes ranged from 3.72 +/- 1.51 microV (P1/N1, N = 18) to 1.49 +/- 0.77 microV (P3/N3, N = 16). Latency-intensity (LI) slopes ranged from -38.7 +/- 7.3 microseconds dB-1 (P1, N = 18) to -71.6 +/- 21.9 microseconds dB-1 (N3, N = 15) and amplitude-intensity (AI) slopes ranged from 0.20 +/- 0.08 microV dB-1 (P1/N1, N = 18) to 0.07 +/- 0.04 microV dB-1 (P3/N3, N = 11). The mean response threshold across all animals was -21.83 +/- 3.34 dB re: 1.0 g ms-1 (N = 18). Responses remained after cochlear extirpation showing that they could not depend critically on cochlear activity. Responses were eliminated by destruction of the vestibular end organs, thus showing that responses depended critically and specifically on the vestibular system. The results demonstrate that the responses are vestibular and the findings provide a scientific basis for using vestibular responses to evaluate vestibular function through ontogeny and senescence in the quail.

NASA Discipline Neuroscience↗
NASA NTRS1996

Modulation of vergence by off-vertical yaw axis rotation in the monkey: normal characteristics and effects of space flight

Horizontal movements of both eyes were recorded simultaneously using scleral search coils in 2 rhesus monkeys before and after the COSMOS 2229 space-flight of 1992-1993. Another 9 monkeys were tested at comparable time intervals and served as controls. Ocular vergence, defined as the difference in horizontal position between the left and right eyes, was measured during off-vertical yaw axis rotation (OVAR) in darkness. Vergence was modulated sinusoidally as a function of head position with regard to gravity during OVAR. The amplitude of peak-to-peak modulation increased with increments in tilt of the angle of the rotational axis (OVAR tilt angle) that ranged from 15 degrees to 90 degrees. Of the 11 monkeys tested, 1 had no measurable modulation in vergence. In the other 10, the mean amplitude of the peak to peak modulation was 5.5 degrees +/- 1.3 degrees at 90 degrees tilt. Each of these monkeys had maximal vergence when its nose was pointed close to upward (gravity back; mean phase: -0.9 degree +/- 26 degrees). After space flight, the modulation in vergence was reduced by over 50% for the two flight monkeys, but the phase of vergence modulation was not altered. The reduction in vergence modulation was sustained for the 11-day postflight testing period. We conclude that changes in vergence are induced in monkeys by the sinusoidal component of gravity acting along the naso-occipital axis during yaw axis OVAR, and that the modulation of the vergence reflex is significantly less sensitive to linear acceleration after space flight.

Flight Experiment↗
NASA NTRS1996

Short latency vestibular evoked potentials in the chicken embryo

Electrophysiological responses to pulsed linear acceleration stimuli were recorded in chicken embryos incubated for 19 or 20 days (E19/E20). Responses occurred within the first 16 ms following the stimulus onset. The evoked potentials disappeared following bilateral labyrinthectomy, but persisted following cochlear destruction alone, thus demonstrating that the responses were vestibular. Approximately 8 to 10 response peaks could be identified. The first 4 positive and corresponding negative components (early peaks with latencies < 6.0 ms) were scored and latencies and amplitudes quantified. Vestibular response latencies were significantly longer (P < 0.01) and amplitudes significantly smaller (P < 0.001) than those observed in 2-week-old birds. Mean response threshold for anesthetized embryos was -15.9dBre 1.0 g/ms, which was significantly higher (P < 0.03) than those observed in 2-week-old birds (-23.0dBre 1.0 g/ms). Latency/intensity functions (that is, slopes) were not significantly different between embryos and 2-week-old animals, but amplitude/intensity functions for embryos were significantly shallower than those for 2-week-old birds (P < 0.001). We presume that these differences reflect the refinement of sensory function that occurs following 19 to 20 days of incubation. The recording of vestibular evoked potentials provides an objective, direct and noninvasive measure of peripheral vestibular function in the embryo and, as such, the method shows promise as an investigative tool. The results of the present study form the definitive basis for using vestibular evoked potentials in the detailed study of avian vestibular ontogeny and factors that may influence it.

Non-NASA Center↗
NASA NTRS1993

A multidimensional model of the effect of gravity on the spatial orientation of the monkey

A "sensory conflict" model of spatial orientation was developed. This mathematical model was based on concepts derived from observer theory, optimal observer theory, and the mathematical properties of coordinate rotations. The primary hypothesis is that the central nervous system of the squirrel monkey incorporates information about body dynamics and sensory dynamics to develop an internal model. The output of this central model (expected sensory afference) is compared to the actual sensory afference, with the difference defined as "sensory conflict." The sensory conflict information is, in turn, used to drive central estimates of angular velocity ("velocity storage"), gravity ("gravity storage"), and linear acceleration ("acceleration storage") toward more accurate values. The model successfully predicts "velocity storage" during rotation about an earth-vertical axis. The model also successfully predicts that the time constant of the horizontal vestibulo-ocular reflex is reduced and that the axis of eye rotation shifts toward alignment with gravity following postrotatory tilt. Finally, the model predicts the bias, modulation, and decay components that have been observed during off-vertical axis rotations (OVAR).

NASA Discipline Neuroscience↗