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Adaptation to rotating artificial gravity environments.

A series of pioneering experiments on adaptation to rotating artificial gravity environments was conducted in the 1960s. The results of these experiments led to the general belief that humans with normal vestibular function would not be able to adapt to rotating environments with angular velocities above 3 or 4 rpm. By contrast, our recent work has shown that sensory-motor adaptation to 10 rpm can be achieved relatively easily and quickly if subjects make the same movement repeatedly. This repetition allows the nervous system to gauge how the Coriolis forces generated by movements in a rotating reference frame are deflecting movement paths and endpoints and to institute corrective adaptations. Independent mechanisms appear to underlie restoration of straight movement paths and of accurate movement endpoints. Control of head movements involves adaptation of vestibulo-collic and vestibulo-spinal mechanisms as well as adaptation to motor control of the head as an inertial mass. The vestibular adaptation has a long time constant and the motor adaptation a short one. Surprisingly, Coriolis forces generated by natural turning and reaching movements in our normal environment are typically larger than those elicited in rotating artificial gravity environments. They are not recognized as such because self-generated Coriolis forces during voluntary trunk rotation are perceptually transparent. After adaptation to a rotating environment is complete, the Coriolis forces generated by movements within it also become transparent and are not felt although they are still present.

Adaptation, Physiological↗

The role of reafference in recalibration of limb movement control and locomotion.

The reafference model has frequently been used to explain spatial constancy during eye and head movements. We have found that its basic concepts also form part of the information processing necessary for the control and recalibration of reaching movements. Reaching was studied in a novel force environment--a rotating room that creates centripetal forces of the type that could someday substitute for gravity in space flight, and Coriolis forces which are side effects of rotation. We found that inertial, noncontacting Coriolis forces deviate the path and endpoint of reaching movements, a finding that shows the inadequacy of equilibrium position models of movement control. Repeated movements in the rotating room quickly lead to normal movement patterns and to a failure to perceive the perturbing forces. The first movements made after rotation stops, without Coriolis forces present, show mirror-image deviations and evoke perception of a perturbing force even though none is present. These patterns of sensorimotor control and adaptation can largely be explained on the basis of comparisons of efference copy, reafferent muscle spindle, and cutaneous mechanoreceptor signals. We also describe experiments on human locomotion using an apparatus similar to that which Mittelstaedt used to study the optomotor response of the Eristalis fly. These results show that the reafference principle relates as well to the perception of the forces acting on and exerted by the body during voluntary locomotion.

Arm↗

Motor control and learning in altered dynamic environments.

Dynamic perturbations of reaching movements are an important technique for studying motor learning and adaptation. Adaptation to non-contacting, velocity-dependent inertial Coriolis forces generated by arm movements during passive body rotation is very rapid, and when complete the Coriolis forces are no longer sensed. Adaptation to velocity-dependent forces delivered by a robotic manipulandum takes longer and the perturbations continue to be perceived even when adaptation is complete. These differences reflect adaptive self-calibration of motor control versus learning the behavior of an external object or 'tool'. Velocity-dependent inertial Coriolis forces also arise in everyday behavior during voluntary turn and reach movements but because of anticipatory feedforward motor compensations do not affect movement accuracy despite being larger than the velocity-dependent forces typically used in experimental studies. Progress has been made in understanding: the common features that determine adaptive responses to velocity-dependent perturbations of jaw and limb movements; the transfer of adaptation to mechanical perturbations across different contact sites on a limb; and the parcellation and separate representation of the static and dynamic components of multiforce perturbations.

Animals↗

Sensorimotor aspects of high-speed artificial gravity: III. Sensorimotor adaptation.

As a countermeasure to the debilitating physiological effects of weightlessness, astronauts could live continuously in an artificial gravity environment created by slow rotation of an entire spacecraft or be exposed to brief daily "doses" in a short radius centrifuge housed within a non-rotating spacecraft. A potential drawback to both approaches is that head movements made during rotation may be disorienting and nauseogenic. These side effects are more severe at higher rotation rates, especially upon first exposure. Head movements during rotation generate aberrant vestibular stimulation and Coriolis force perturbations of the head-neck motor system. This article reviews our progress toward distinguishing vestibular and motor factors in side effects of rotation, and presents new data concerning the rates of rotation up to which adaptation is possible. We have studied subjects pointing to targets during constant velocity rotation, because these movements generate Coriolis motor perturbations of the arm but do not involve unusual vestibular stimulation. Initially, reaching paths and endpoints are deviated in the direction of the transient lateral Coriolis forces generated. With practice, subjects soon move in straighter paths and land on target once more. If sight of the arm is permitted, adaptation is more rapid than in darkness. Initial arm movement trajectory and endpoint deviations are proportional to Coriolis force magnitude over a range of rotation speeds from 5 to 20 rpm, and there is rapid, complete motor adaptation at all speeds. These new results indicate that motor adaptation to high rotation rates is possible. Coriolis force perturbations of head movements also occur in a rotating environment but adaptation gradually develops over the course of many head movements.

Adaptation, Physiological↗

Moving objects in a rotating environment: rapid prediction of Coriolis and centrifugal force perturbations.

Grip force adaptation to Coriolis and centrifugal force perturbations was tested in healthy subjects. Eight subjects were seated in a rotating chamber in a rotating axis position. They each grasped an instrumented object resting on the thumb, which was stabilized by the other fingers from above. Subjects performed horizontal point-to-point movements with the grasped object away and towards the trunk. These movements were directed in a nonparallel fashion towards the axis of rotation prior (40 pre-rotational movements), during (80 per-rotational movements) and following (40 post-rotational movements) clockwise body rotation. During pre- and post-rotational movements two load force peaks of similar magnitude occurred during the acceleratory and deceleratory phases of the movements. Accordingly, a Coriolis force, which was orthogonal and proportional to the linear velocity of the moving arm, as well as a centrifugal force proportional to the system's squared angular velocity and movement amplitude developed during per-rotational movements. The load perturbations altered the load force profile in a characteristic way. The first 10 per-rotational movement sequence revealed that there was a less precise coupling between grip and load force magnitudes and a reduced temporo-spatial co-ordination between grip and load force profiles. With increasing number of per-rotational movements, there was significant improvement in the temporo-spatial co-ordination and in the coupling in force magnitude between grip and load force profiles, indicating an ongoing adaptation process. The coupling between grip and load forces proved to be similarly precise for the last 10 per-rotational movements and for pre-rotational movements, suggesting complete adaptation. Significant effects were observed for the first post rotational movements following adaptation to the per-rotational load characteristics both for the temporal co-ordination between grip and load forces and for the coupling in force magnitudes. However, the last 10 post-rotational movements proved to be similarly precise with comparison to pre-rotational performance in terms of grip force regulation with movement-induced loads. The results are discussed within the context of the CNS ability to use internal models when planning and processing anticipatory grip force adjustments during manipulative tasks.

Acoustic Stimulation↗

Aspects of body self-calibration.

The representation of body orientation and configuration is dependent on multiple sources of afferent and efferent information about ongoing and intended patterns of movement and posture. Under normal terrestrial conditions, we feel virtually weightless and we do not perceive the actual forces associated with movement and support of our body. It is during exposure to unusual forces and patterns of sensory feedback during locomotion that computations and mechanisms underlying the ongoing calibration of our body dimensions and movements are revealed. This review discusses the normal mechanisms of our position sense and calibration of our kinaesthetic, visual and auditory sensory systems, and then explores the adaptations that take place to transient Coriolis forces generated during passive body rotation. The latter are very rapid adaptations that allow body movements to become accurate again, even in the absence of visual feedback. Muscle spindle activity interpreted in relation to motor commands and internally modeled reafference is an important component in permitting this adaptation. During voluntary rotary movements of the body, the central nervous system automatically compensates for the Coriolis forces generated by limb movements. This allows accurate control to be maintained without our perceiving the forces generated.

Adaptation, Physiological↗

Coordinated turn-and-reach movements. I. Anticipatory compensation for self-generated coriolis and interaction torques.

When reaching movements involve simultaneous trunk rotation, additional interaction torques are generated on the arm that are absent when the trunk is stable. To explore whether the CNS compensates for such self-generated interaction torques, we recorded hand trajectories in reaching tasks involving various amplitudes and velocities of arm extension and trunk rotation. Subjects pointed to three targets on a surface slightly above waist level. Two of the target locations were chosen so that a similar arm configuration relative to the trunk would be required for reaching to them, one of these targets requiring substantial trunk rotation, the other very little. Significant trunk rotation was necessary to reach the third target, but the arm's radial distance to the body remained virtually unchanged. Subjects reached at two speeds-a natural pace (slow) and rapidly (fast)-under normal lighting and in total darkness. Trunk angular velocity and finger velocity relative to the trunk were higher in the fast conditions but were not affected by the presence or absence of vision. Peak trunk velocity increased with increasing trunk rotation up to a maximum of 200 degrees /s. In slow movements, peak finger velocity relative to the trunk was smaller when trunk rotation was necessary to reach the targets. In fast movements, peak finger velocity was approximately 1.7 m/s for all targets. Finger trajectories were more curved when reaching movements involved substantial trunk rotation; however, the terminal errors and the maximal deviation of the trajectory from a straight line were comparable in slow and fast movements. This pattern indicates that the larger Coriolis, centripetal, and inertial interaction torques generated during rapid reaches were compensated by additional joint torques. Trajectory characteristics did not vary with the presence or absence of vision, indicating that visual feedback was unnecessary for anticipatory compensations. In all reaches involving trunk rotation, the finger movement generally occurred entirely during the trunk movement, indicating that the CNS did not minimize Coriolis forces incumbent on trunk rotation by sequencing the arm and trunk motions into a turn followed by a reach. A simplified model of the arm/trunk system revealed that additional interaction torques generated on the arm during voluntary turning and reaching were equivalent to < or =1.8 g (1 g = 9.81 m/s(2)) of external force at the elbow but did not degrade performance. In slow-rotation room studies involving reaching movements during passive rotation, Coriolis forces as small as 0.2 g greatly deflect movement trajectories and endpoints. We conclude that compensatory motor innervations are engaged in a predictive fashion to counteract impending self-generated interaction torques during voluntary reaching movements.

Adult↗

Sensorimotor adaptation to inertial forces in a multi-force environment does not depend on the number of targets: indirect validation of the altered-proprioception hypothesis.

The ability of our sensorimotor system to adapt to changing and complex environmental demands has been under experimental scrutiny for more than a century. Previous works have shown that aimed arm movements adapt quickly and completely to Coriolis force, but incompletely to the combination of Coriolis and centrifugal forces without visual cues. Two hypotheses may be advanced to explain this discrepancy: the workspace-exploration hypothesis, and the degraded-proprioception hypothesis. The aim of this study was to distinguish between the above two alternatives by comparing adaptive improvement during off-axis rotation in subjects pointing at one, three or seven different targets in complete darkness. Two main results emerge: (a) off-axis rotation led initially to errors in the direction of Coriolis force and in the opposite direction of the centrifugal force; (b) the size of the visited workspace has no effect on the way the subjects adapt to a multi-force environment. The lack of a target-number effect and the persistence of lateral errors in the pointing movements performed during rotation of the platform, support the degraded-proprioception rather than the workspace-exploration hypothesis of adaptation to a multi-force environment.

Acclimatization↗

Toroidal coil counter-current chromatography study of the mass transfer rate of proteins in aqueous-aqueous polymer phase system.

The cause of excessive band broadening of protein samples in polymer phase partitioning by counter-current chromatography (CCC) was investigated. A simple rotary device was constructed to measure the mass transfer rates of five samples including potassium dichromate, methylene blue, lysozyme, ovalbumin and human serum albumin. The results indicated that the mass transfer rates of these samples are closely correlated with their molecular masses: the higher the molecular mass, the lower the mass transfer rate. These findings are also consistent with the partition efficiencies of these samples in the same solvent system by CCC. The beneficial effect of the Coriolis force demonstrated in protein separations by the toroidal coil centrifuge may be reasonably explained on the basis of the mass transfer resistance of protein molecules through the interface: we speculate that when the Coriolis force acts parallel to the effective coil segment it can produce large interfacial areas by dispersing the mobile phase into the stationary phase, thus accelerating the mass transfer rate of protein samples.

Chromatography, Liquid↗

Acoustic streaming in a rotating fluid.

Acoustic streaming theory is derived that is applicable to a fluid that is slow moving in a reference frame that rotates with a constant angular velocity omega. A simplified streaming equation is obtained for the special case in which the acoustic angular frequency omega is large relative to omega, and the change in fluid density due to rotation alone is negligible. For this special case it is shown that the "driving force" for the acoustic streaming is independent of omega. Thus, if no acoustic streaming is present in a fluid system that is stationary, then no steady-state acoustic streaming is predicted for a similar system that rotates with constant angular velocity. For a system in which acoustic streaming is present, the flow behavior depends on the relative magnitudes of the Coriolis forces and the viscous forces. If the Ekman number is large (that is, the viscous force dominates) then the predicted flow is identical to that which would exist in a stationary system. If, on the other hand, the Ekman number is small then the Coriolis force dominates and the component of flow in the direction of the axis of rotation can be much smaller in the rotating system than in a similar system at rest.

Acoustics↗

Frequency-dependent transversal flow control in centrifugal microfluidics.

This work presents a novel flow switch for centrifugal microfluidic platforms which is solely controlled by the Coriolis pseudo force. This Coriolis switch consists of an inverse Y-structure with one common upstream channel and two symmetric outlets on a rotating disk. Above a certain threshold frequency, the Coriolis force becomes dominant that the entire flow is diverted into one of the outlets which is selected by the direction of rotation. The threshold frequency has been measured to be 350 rad s(-1)(approximately 55.7 Hz) for a channel width of 360 microm and a depth of 125 microm. The results are supported by extensive CFD simulations.

Journal Article↗

Extremely non-orthogonal axes in a sense organ for rotation: behavioural analysis of the dipteran haltere system.

Flies acquire information about self-rotation via Coriolis forces detected by their moving halteres. Information processing in the haltere system was analysed by exploiting the method of simulating rotational stimuli by vibrating the fly's body and simultaneously observing compensatory head and wing reactions. Although the force acting on one haltere contains Coriolis terms for rotations about three orthogonal axes, the one-haltered fly has only two measuring axes which are coded in lateral force components. A fly with two halteres has two vertical measuring axes and two horizontal axes, the latter spanning an angle of about 120 degrees. Thus, three-dimensional turning information is acquired by bilateral computation in a highly non-orthogonal system. In the stimulus velocity range up to 1000 degrees/s, comparison of intact and one-haltered flies demonstrates that for the head roll reaction the inputs from both halteres are summated, whereas for the pitch reaction the summated inputs are modified by bilateral inhibition. This non-linear operation results in uniform gains and axis fidelity for all stimulus directions in the case of the head reaction. Response saturation at high velocities takes place after the bilateral summation. The functional consequences of non-orthogonality in the dipteran haltere system is apparently superior sensitivity for pitch compared to roll. Minimization of the "area of confusion", an argument for orthogonality, seems to be of minor importance. The non-orthogonality necessitates a transformation from covariant projections to contravariant motor components. In tensor theory of the vestibulo-ocular reflex of vertebrates, this is widely assumed to be a linear operation performed by a metric tensor. The fly's solution is a linear tensor operation supplemented by a non-linear bilateral inhibition for the pitch reaction.

Acceleration↗

Influence of environmental factors on shot put and hammer throw range.

On the rotating Earth, in addition to the Newtonian gravitational force, two additional relevant inertial forces are induced, the centrifugal and Coriolis forces. Using computer modelling for typical release heights and optimal release angles, we compare the influence of Earth rotation on the range of the male hammer throw and shot put with that of air resistance, wind, air pressure and temperature, altitude and ground obliquity. Practical correction maps are presented, by which the ranges achieved at different latitudes and/or with different release directions can be corrected by a term involving the effect of Earth rotation. Our main conclusion and suggestion is that the normal variations of certain environmental factors can be substantially larger than the smallest increases in the world records as acknowledged by the International Amateur Athletic Federation and, therefore, perhaps these should be accounted for in a normalization and adjustment of the world records to some reference conditions. Although this suggestion has certainly been made before, the comprehensiveness of our study makes it even more compelling. Our numerical calculations contribute to the comprehensive understanding and tabulation of these effects, which is largely lacking today.

Acceleration↗

Drift mechanism caused by a nonlinear wave and the Cassini Division and Uranian rings formation.

The mechanism leading to the observed coexistence of gaps and narrow ringlets in the planetary rings is found. It is based upon the quasi-stationary radial drift of the matter under action of two forces in the disk plane: the Coriolis force and the Reynolds stresses. To an accuracy of the factor of 2 the first force coincides with the Lorentz force, therefore the radial drift in rings is similar to the gradient drift of plasma in the magnetic field. The second force is produced by the wave generated by the nearby satellite in the resonance position. In inertial systems, the second force alone causes a matter flow in its direction, called acoustic streaming. Since the radial drift is caused by nonlinear time-averaged force of high-frequency harmonic interactions in the wave, it exists in the wave propagation zone: from the birth place of the wave-the resonance position, up to the reflection point of the wave, where its group velocity vanishes. Our estimations show that the size of the density wave propagation zone corresponding to the density wave which had been formerly generated the 2:1 orbital resonance with Mimas is consistent with the width of the Cassini Division. In our case the nature of the radial drift is such that first of all it clears out the farthest from the resonance position; later, the closer areas also get affected by the drift. The zone closest to the resonance position itself is the last to be involved in the process. The matter carried away by the drift is partially accumulated near the resonance position forming a narrow dense ringlet. (c) 1996 American Institute of Physics.

Journal Article↗

Experimentally induced motion sickness in fish: possible role of the otolith organs.

OBJECTIVE: Although the occurrence of motion sickness is largely attributed to sensory conflict, the reasons for the highly variable inter-individual susceptibility to motion sickness remain unclear. One cause may be asymmetry between otoconial masses in the right and left inner ear. This hypothesis has been supported by experiments in fish under conditions of prolonged weightlessness and parabolic flight. In swordtail fish (Xiphophorus helleri) we examined the extent to which a Coriolis force environment, which has a strong motion sickness-triggering effect, disturbs swimming behaviour or produces motion sickness. MATERIAL AND METHODS: In a sealed aquarium, fish (n = 22) were subjected to Coriolis stimulation by constant vertical axis rotation, combined with pendular oscillation around a horizontal axis. RESULTS: Initially, at low stimulus intensity, all fish showed active compensatory swimming behaviour. However, while the majority (n = 19) maintained active compensation movements at higher stimulus levels, the others (n = 3) entered a passive uncoordinated state, from which they recovered again after reduction of stimulus intensity. On examination of the otoconial mass, we found asymmetries between the right and left utricles in all three cases and between the saccules in one case. CONCLUSION: We conclude that a lateral preponderance of the otoconial masses in the utricle or saccule promotes motion sickness under experimental conditions. The utricle appears to play a more important role in this than the saccule.

Animals↗

On the origin of interindividual susceptibility to motion sickness.

According to the otolith asymmetry hypothesis for susceptibility to motion sickness, the interlabyrinthine asymmetry in the otoconial mass, which is normally compensated in the circuity of the vestibular system by appropriate neuronal weighting, becomes disrupted as a result of unfamiliar movement patterns or force environments. Measurements in various species have demonstrated a large scatter in the otoconial mass. As the utricle and saccule have distinct functions in the vestibular system it seems appropriate to deal with these organs separately. Results are presented on mass distribution and lateral preponderance of the otoconial mass in the utricles and saccules of the salmon (Salmo salar) and trout (Salmo irideus). The measurements revealed considerably larger dimensions of the saccular otoconia in these species, amounting to twenty-fold. This substantial difference indicates that different regulatory principles underlie the otoconial generation of each of the organs and is presumably related to their specific functions. The lateral preponderance was found to be normally distributed for both organs in both species, with standard deviations of approx. 4% except for the utricular otoconia of the salmon (13%). In a second set of experiments, fish were exposed to a Coriolis force environment. Their sealed aquarium was subjected to constant vertical axis rotation combined with pendular oscillation around the horizontal axis. The aquarium was illuminated by a light source fixed to project through the top, and a video camera recorded the movements of the fish. During combined rotation and pendular oscillation, one group of fish maintained an active compensatory swimming behaviour, whereas the movements of a second group became uncoordinated and often led to passive behaviour. Analysis of the otoconia of these two groups of fish promises to yield further evidence relevant to the otolith asymmetry hypothesis.

Animals↗

Effect of microgravity on spatial orientation and posture regulation during Coriolis stimulation.

OBJECTIVE: To elucidate spatial orientation and posture regulation under conditions of microgravity. MATERIAL AND METHODS: Coriolis stimulation was done with five normal subjects on the ground (1 g) and onboard an aircraft (under conditions of microgravity during parabolic flight). Subjects were asked to tilt their heads forward during rotation at speeds of 0, 50, 100 and 150 degrees/s on the ground and 100 degrees/s during flight. Body sway was recorded using a 3D linear accelerometer and eye movements using an infrared charge-coupled device video camera. Flight experiments were performed on 5 consecutive days, and 11-16 parabolic maneuvers were done during each flight. Two subjects boarded each flight and were examined alternately at least five times. RESULTS: Coriolis stimulation at 1 g caused body sway, nystagmus and a movement sensation in accordance with inertial inputs at 1 g. Neither body sway, excepting a minute sway due to the Coriolis force, nor a movement sensation occurred in microgravity, but nystagmus was recorded. CONCLUSIONS: Posture, eye movement and sensation at 1 g are controlled with reference to spatial coordinates that represent the external world in the brain. Normal spatial coordinates are not relevant in microgravity because there is no Z-axis, and the posture regulation and sensation that depend on them collapse. The discrepancy in responses between posture and eye movement under conditions of microgravity may be caused by a different constitution of the effectors which adjust posture and gaze.

Adult↗

Use of a motion sickness history questionnaire for prediction of simulator sickness.

This research assessed the usefulness of the Motion History Questionnaire (MHQ) for the prediction of simulator sickness, a form of motion sickness experienced by pilots training in ground-based flight simulators. Four MHQ scoring keys were compared: (1) the original MHQ key which had been validated on a sample of U.S. Navy student pilots exposed to Coriolis forces, (2 and 3) two keys which had been validated on a sample of civilian college students exposed to simulated ship motions, and (4) a simulator sickness key empirically derived in the present research and cross-validated. Navy and Marine Corps aviators (N = 456) filled out the MHQ prior to their regularly scheduled flight simulator training and were divided into validation and cross-validation samples. All scoring keys were predictive of reported symptoms of sickness, but highest correlations were obtained with the empirically-derived simulator sickness (SS) key. It is suggested that the SS key be used for self-testing so that pilots may be made aware of their risk for developing simulator sickness.

Aerospace Medicine↗