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doi: 10.3233/ves-2003-134-602pmid: N/A
Embedded in neural and behavioral organization is a structure of sensorimotor space. Both this embedded spatial structure and the structure of physical space inform sensorimotor control. This paper reviews studies in which the gravitational vertical and horizontal are crucial. The mathematical expressions of spatial geometry in these studies indicate methods for investigating sensorimotor control in freefall.In freefall, the spatial structure introduced by gravitation – the distinction between vertical and horizontal – does not exist. However, an astronaut arriving in space carries the physiologically-embedded distinction between horizontal and vertical learned on earth. The physiological organization based on this distinction collapses when the strong otolith activity and other gravitational cues for sensorimotor behavior become unavailable. The mathematical methods in this review are applicable in understanding the changes in physiological organization as an astronaut adapts to sensorimotor control in freefall.Many mathematical languages are available for characterizing the logical structures in physiological organization. Here, group theory is used to characterize basic structure of physical and physiological spaces. Dynamics and topology allow the grouping of trajectory ranges according to the outcomes or attractors. The mathematics of ordered structures express complex orderings, such as in multiphase movements in which different parts of the body are moving in different phase sequences. Conditional dynamics, which combines dynamics with the mathematics of ordered structures, accommodates the parsing of movement sequences into trajectories and transitions.Studies reviewed include those of the sit-to-stand movement and early locomotion, because of the salience of gravitation in those behaviors. Sensorimotor transitions and the conditions leading to them are characterized in conditional dynamic control structures that do not require thinking of an organism as an input-output device. Conditions leading to sensorimotor transitions on earth assume the presence of a gravitational vertical which is lacking in space. Thus, conditions used on earth for sensorimotor transitions may become ambiguous in space. A platform study in which sensorimotor transition conditions are ambiguous and are related to motion sickness is reviewed.
doi: 10.3233/ves-2003-134-603pmid: N/A
Perceptual disturbances in zero-g and 1-g differ. For example, the vestibular coriolis (or "cross-coupled") effect is weaker in zero-g. In 1-g, blindfolded subjects rotating on-axis experience perceptual disturbances upon head tilt, but the effects diminish in zero-g.Head tilts during centrifugation in zero-g and 1-g are investigated here by means of three-dimensional modeling, using a model that was previously used to explain the zero-g reduction of the on-axis vestibular coriolis effect. The model's foundation comprises the laws of physics, including linear-angular interactions in three dimensions.Addressed is the question: In zero-g, will the vestibular coriolis effect be as weak during centrifugation as during on-axis rotation? Centrifugation in 1-g was simulated first, with the subject supine, head toward center. The most noticeable result concerned direction of head yaw. For clockwise centrifuge rotation, greater perceptual effects arose in simulations during yaw counterclockwise (as viewed from the top of the head) than for yaw clockwise. Centrifugation in zero-g was then simulated with the same "supine" orientation. The result: In zero-g the simulated vestibular coriolis effect was greater during centrifugation than during on-axis rotation. In addition, clockwise-counterclockwise differences did not appear in zero-g, in contrast to the differences that appear in 1-g.
doi: 10.3233/ves-2003-134-604pmid: N/A
It has been suggested that, in the fish, the change of otolith mass during development under altered gravity conditions [1,2,3,4,5,6,24,25,36,37] and the growth of otoliths in normal conditions [22,23,26], are determined by feedback between otolith dynamics and the processes that regulate otolith growth. The hypothesis originates from an oscillator model of the otolith [30] in which otolith mass is one of the parameters. However, the validity of this hypothesis is not obvious and has not been experimentally verified. We tested this hypothesis by comparing the oscillator model with a simplified spatially distributed model of the otolith. It was shown that in the case of a spatially distributed fixation of the otolith plate (otoconial layer) to the macular surface, the mechanical sensitivity of the otolith does not depend on the total otolith mass nor on its longitudinal size. It is determined by otolith thickness, the Young's modulus and viscosity of gel layer of the growing otolith. These parameters may change in order to maintain otolith sensitivity under conditions (such as growth or altered gravity) that change the dynamics of otolith movement.
Wiederhold, Michael L.; Harrison, Jeffrey L.; Gao, Wenyuan
doi: 10.3233/ves-2003-134-605pmid: N/A
The otoliths of adult animals do not change significantly during space flight. However, during the period when otoliths are first developing, rearing in space produces significantly larger otoliths. Conversely, animals reared on a centrifuge have smaller than normal otoliths. To identify a critical period during development for gravitational effects on otolith growth, fertilized zebrafish (Danio rerio) eggs were reared on a centrifuge for 1 week. The fine structure of their inner ear during development was studied by both light- and transmission electron microscopy. By 16 hours after fertilization (1-g, at 28.5°C), precursors of the otoliths are seen but no sign of a sensory epithelium is present. Mature hair cells, appearing capable of mechanotransduction, are not seen until between 48 and 72 hours after fertilization. Zebrafish reared at 3-g from 1 to 7 days after fertilization exhibit significantly slower otolith growth than did 1-g controls. Fish exposed to 3-g only from 12–36 h after fertilization had slightly smaller otoliths than 1-g controls, but this difference was not significant. Animals exposed to 3-g from 36h to 7d after fertilization did have significantly smaller otoliths. If the fish use their hair cells to assess otolith weight in a regulatory role, the hair cells would have to be functional. Thus the earliest stage zebrafish, which were not significantly affected by centrifugation, probably did not have an adequate means of sensing otolith weight to "correct" for the excess weight.(Supported by NASA: NAG2-952 and NAG10-0180)
Clarke, A.H.; Schönfeld, U.; Helling, K.
doi: 10.3233/ves-2003-134-606pmid: N/A
Attention is directed towards the recently developed unilateral tests of saccular and utricular function. Together with the now widely used head-thrust test and the standard caloric test for semicircular canal function, these provide for a more comprehensive unilateral examination of labyrinth function.The efficacy of vestibular evoked myogenic potentials (VEMP) as a direct unilateral test of saccular function is currently being demonstrated in an increasing number of reports. Furthermore, the relevant neuronal pathways have been delineated in animal studies, so that all evidence points to the validity of the VEMP as a saccule-mediated response. Concerning utricular function, considerable headway has been made using the unilateral centrifugation paradigm. Testing is performed with a variable radius rotary chair with constant velocity rotation about the earth-vertical axis. Displacing the head by 3.5–4 cm from the rotation axis, the eccentrically positioned utricle is stimulated unilaterally by the resultant centrifugal force. This paradigm can be employed to elicit a utriculo-ocular response (UOR) or to permit measurement of the subjective visual vertical (SVV). More recently, it has also been demonstrated that testing during normal, on-centre yaw axis rotation is often sufficient to localise peripheral otolith dysfunction by means of SVV estimation. This test mode can be easily integrated into routine clinical testing. To illustrate the efficacy of such differential testing, the findings from two patients are presented that demonstrate for the first time an isolated unilateral utricular dysfunction.
L. Wuyts, Floris; Hoppenbrouwers, Mieke; Pauwels, Griet; Van de Heyning, Paul H.
doi: 10.3233/ves-2003-134-607pmid: N/A
Utricular sensitivity and preponderance of the right or left utricle can be assessed by means of the unilateral centrifugation test. In this test, subjects are rotated about an earth vertical axis at a velocity of 400 degrees per second. During the ongoing rotation, the subject is gradually translated 4 cm first to the right, and then to the left, along an interaural axis, to a position at which one utricle becomes aligned with the axis of rotation, and at this point is subjected only to gravitational forces. At this eccentric position, the contralateral utricle is exposed to the combination of gravity and a centrifugal acceleration of 0.4g, corresponding to an apparent roll-tilt of 21.7 degrees. This stimulus induces ocular counterrolling (OCR), which is measured on-line using three-dimensional video-oculography (VOG).We observed that ocular counterrolling appears as a linear function of the gravito-inertial acceleration tilt of the head centre (GIAHC) during the lateral translation. We present a theoretical model for this linear relationship that contains two parameters: 1) the slope of the linear regression is a measure for the utricular sensitivity and 2) the intercept of the linear regression is a measure of the preponderance of the right or left utricle. The strength of the model is supported by data obtained from 28 healthy subjects and 14 patients with unilateral vestibular deafferentiation (UVD) due to acoustic neuroma surgery.
Fraser, Peter J.; Cruickshank, Stuart F.; Shelmerdine, Richard L.
doi: 10.3233/ves-2003-134-608pmid: N/A
Following the discovery of a hydrostatic pressure sensor with no associated gas phase in the crab, and the knowledge that several systems of cells in culture show long term alterations to small changes in hydrostatic pressure, we show here that vestibular type II hair cells in a well known model system (the isolated elasmobranch labyrinth), are sensitive to hydrostatic pressure. This new finding for the vertebrate vestibular system may provide an explanation for low levels of resting activity in vertebrate hair cells and explain how fish without swim bladders sense hydrostatic cues. It could have implications for humans using their balancing systems in hypobaric or hyperbaric environments such as in aircraft or during space exploration. Although lacking the piston mechanism thought to operate in crab thread hairs which sense angular acceleration and hydrostatic pressure, the vertebrate system may use larger numbers of sensory cells with resultant improvement in signal to noise ratio.The main properties of the crab hydrostatic pressure sensing system are briefly reviewed and new experimental work on the isolated elasmobranch labyrinth is presented.
Angelaki, Dora E.; Dickman, J. David
doi: 10.3233/ves-2003-134-609pmid: N/A
The processing and detection of tilts relative to gravity from actual motion (translational accelerations) is one of the most fundamental issues for understanding vestibular sensorimotor control in altered gravity environments. In order to better understand the nature of multisensory signals in detecting motion and tilt, we summarize here our recent studies regarding the central processing of vestibular signals during multi-axis rotational and translational stimuli. Approximately one fourth of the cells in the vestibular nuclei exclusively encoded rotational movements (Canal-Only neurons) and were unresponsive to translation. The Canal-Only central neurons encoded head rotation in canal afferent coordinates, exhibited no orthogonal canal convergence and were characterized by significantly higher sensitivities to rotation as compared to canal afferents. Another fourth of the neurons modulated their firing rates during translation (Otolith-Only cells). During rotations, these neurons typically only responded when the axis of rotation was earth-horizontal and the head was changing orientation relative to gravity. The remaining cells (approximately half of total population) were sensitive to both rotations and translations (Otolith+Canal neurons). Maximum sensitivity vectors to rotation were distributed throughout the 3D space, suggesting strong convergence from multiple semicircular canals. Only a small subpopulation (approximately one third) of these Otolith+Canal neurons seems to encode a true estimate of the translational component of the imposed passive head and body movement. These results provide the first step in further understanding multisensory convergence in normal gravity, as this task is fundamental to our appreciation of neurovestibular adaptation to altered gravity.
doi: 10.3233/ves-2003-134-610pmid: N/A
Prolonged microgravity during orbital flight is a unique way to modify the otolith inputs and to determine the extent of their contribution to the vertical vestibulo-ocular reflex (VOR) and optokinetic nystagmus (OKN). This paper reviews the data collected on 10 astronauts during several space missions and focuses on the changes in the up-down asymmetry. Both the OKN elicited by vertical visual stimulation and the active VOR elicited by voluntary pitch head movements showed an asymmetry before flight, with upward slow phase velocity higher than downward slow phase velocity. Early in-flight, this asymmetry was inverted, and a symmetry of both responses was later observed. An upward shift in the vertical mean eye position in both OKN and VOR suggests that these effects may be related to otolith-dependent changes in eye position which, in themselves, affect slow phase eye velocity.
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