J. Wetzig

Oculovestibular interactions under microgravity

SummaryOn a space mission in March 1992 a set of experiments were performed aimed at clarifying the interaction between visual, proprioceptive and vestibular inputs to the equilibrium system. Using the VESTA goggle facility from the European Space Agency we investigated the effect of pure neck receptor stimulation on eye position as measured by the flash afterimage method and on perception of a head-fixed luminous line in space. Space vestibular adaptation processes were measured by rotating pattern perception during prescribed head movements. It was found that static ocular counterrotation does not occur under micro gravity conditions. This result suggests that the neck receptors apparently do not contribute to a measurable extent. The subjective orientation of a vertical line was perceived correctly inflight. Obviously neck receptors on the perception level can fully substitute for the ineffective equilibrium organs of the inner ear within less than 4 days. The rotating pattern perception during different head motion patterns is not influenced by the absence of a gravity reference.

Dorsal light response and changes of its responses under varying acceleration conditions

In order to improve our understanding about functions of the gravity sensors, we have conducted four experiments in goldfish: 1) To define the effect of visual information influx on the static labyrinthine response, the dorsal light response (DLR) which had been proposed by von Holst as a model for postural adjustment in fish was reexamined with a newly designed, rotatory illumination device. The fish responded to illumination from the upper half of the visual field and a narrow range around 180 degrees of the lower half visual field. The maximal tilting angle of normal fish was about 40 degrees under horizontal illumination. 2) Under the changes of the gravito-inertial force level produced by a linear sled, the threshold of the gravity sensors was determined from postural adjustment responses. 3) Under hypogravic conditions during the parabolic flight of an airplane, the light-dependent behavior was investigated in intact and labyrinthectomized goldfish. 4) As one of the most likely candidates of the neural centers for the DLR, the valvula cerebelli, which receives its visual information not through the optic tectum but through the pretectal areas, is confirmed by the brain lesion experiments.

Clinical verification of a unilateral otolith test.

In a previous study we reported promising results for a new test to differentiate in vivo unilateral otolith functions. That study pointed to a need for further validation on known pathological cases. In this presentation we will detail the results gathered on a group of clinically verified vestibular defectives (verum) and a normal (control) group. The subjects in the verum group were former patients of the ENT clinic of the university hospital. These subjects had usually suffered from neurinoma of the VIIth cranial nerve or inner ear infections. All had reguired surgical intervention including removal of the vestibular system. The patients were contacted usually two or more years postoperatively. A group of students from the pre- and clinical phase of medical training served as control. Both groups were subjected to standardized clinical tests. These tests served to reconfirm the intra- or postoperative diagnosis of unilateral vestibular loss in the verum group. In the control group they had to establish the normalcy of the responses of the vestibular system. Both groups then underwent testing on our eccentric rotary chain in the manner described before. Preliminary results of the trials indicate that this test may indeed for the first time offer a chance to look at isolated otolith apparati in vivo.

The European vestibular experiments in spacelab-1

A series of experiments were performed in the Spacelab-1 mission on November/December, 1983, pre-, in-, and postflight. These experiments covered various aspects of the functions of the vestibular system, the inflight tests comprising threshold measurements for linear movements in three orthogonal axes, optokinetic stimulation, vestibulo-ocular reflexes under linear and angular accelerations, caloric stimulation with and without linear accelerations; pre- and postflight tests repeated the inflight protocol with the addition of subjective vertical and eye counter-rotation measurements using a tilt table. One of the most surprising and significant results was the caloric test: strong caloric nystagmus on the two subjects tested was recorded inflight; this was contrary to what was expected from Baranys convection hypothesis for caloric nystagmus.

INFLUENCE OF PROPRIOCEPTIVE INFORMATION ON SPACE ORIENTATION ON THE GROUND AND IN ORBITAL WEIGHTLESSNESS

Conscious space orientation depends on afferent information from different sense organs including the labyrinth, the eyes, tactile cues from the skin, joint receptors, muscle spindles, tendon organs and possibly viscera. An important role is played by impulses from the cervical position receptors in interaction with concomitant information from the otolith system. In order to isolate the effect of cervical position receptors from that of the otolith system, space experiments in orbital weightlessness and in parabolic aircraft flight were performed. It was found that stimulation of the neck receptors in weightlessness markedly influences the perception of the subjective vertical and horizontal and in addition has a weak effect on ocular torsion.

Responses to eccentric rotation in two space-bound subjects

SummaryTwo subjects were rotated eccentrically in the manner described previously. In contrast to a normal control group, settings of a luminous line to the subjective vertical were almost unrelated to the gravitoinertial vector before, and totally so shortly after, space flight. Only 3 days postflight did a clear relation to the gravitoinertial vector re-establish itself in the one subject who actually flew. The correspondence became normal 5 days after the flight. Since there were no clinical abnormalities evident in the subjects, it is suggested that both subjects suppressed their vestibular information, presumably as an effect of vestibular deconditioning training before the flight. In addition, as a consequence of the flight experience one subject continued to ignore it several days after the flight.

Effects of rectilinear acceleration, caloric and optokinetic stimulation of human subjects in the spacelab D-1 mission

A set of vestibular experiments was performed during the course of the German Spacelab D-1 mission from 30 October to 6 November 1985 by a consortium of experimenters from various european countries. Similar to the Spacelab SL-1 mission all of the scientific crew members were theoretically and practically trained for the experiments. Baseline measurements for all tests were collected 113, 86, 44, 30 and 18 days prior to the mission and compared with data taken inflight, on the landing day and the consecutive 7 to 14 days. The hardware comprised mainly a motordriven accelerating platform, the SPACE SLED, and the vestibular helmet, a multi-purpose instrument in support of a variety of vestibular experiments including air-calorisation of the ears, optokinetic stimulation pattern presentation and optical and nystagmographic recording of eye movements. Measurements of the threshold for the perception of detection of whole body movement did not reveal any dramatic changes in the 2 measured axes inflight when compared to preflight values. Early postflight values show a significantly elevated threshold for all axes in 3 out of 4 subjects. The caloric nystagmus, already found during the SL-1 mission, was confirmed on all three tested subjects during the D-1 mission. Its amplitude and in some instances its direction were influenced by horizontal acceleration on the SLED. The amplitude of optokinetic nystagmus increased when subjects were allowed to free-float over that seen when subjects were fixed. Stimulation of the neck receptors by roll movements of the body against the fixated head resulted in illusory object motion to the contralateral side. Torsional movements of the eyes during such neck receptor stimulation was present inflight and postflight, while it had not been observed preflight. Most results point to a reduction of otolithic effects in favour of visual and proprioceptive influences for spatial orientation.

Influence of gravity vector on eye movement elicited by linear acceleration.

When the body/head motion is sensed by otolith organs, they respond not only to the resultant acceleration of the motion but also to the gravitational force. We investigated the influence of the gravity vector on the otolithic-ocular reflex caused by motion in normal subjects. The sled type linear accelerator, moving back and forth with a frequency of 0.25 Hz and an amplitude of 2 m, generated right-left linear acceleration with a maximum magnitude of 0.5 g. We tested every subject under seven different postures: 1) 135 degrees forward tilted (F.T.), 2) 90 degrees F.T., 3) 45 degrees F.T., 4) upright sitting, 5) 45 degrees backward tilted (B.T.), 6) 90 degrees B.T., and 7) 135 degrees B.T. Horizontal eye movements with nystagmic pattern were elicited by these stimulations and were recorded by EOG. The eye movement data were analyzed by using a computer with the following procedure. After extraction of saccadic components from the data, the remaining parts were connected smoothly. The waves reconstructed in this manner, considered to be primary otolithic-ocular responses, were processed with the FFT method for calculating the amplitude of the component at 0.25 Hz. As a result, the responses were larger in the forward tilted postures than in the backward tilted postures.

Ground based eccentric chair experiments

Two subjects were rotated eccentrically in a manner described before. In contrast to a normal control group settings of a luminous line to the subjective vertical were almost unrelated to the gravitoinertial summation vector of gravity and centrifugal forces about four weeks before and totally so shortly after space flight. Only after four days post flight had passed a noticeable relation to the gravitoinertial vector re-established itself in the one subject which actually flew. The correspondence became normal six days after flight. Since there were no clinical abnormalities evident in the subjects, it is suggested, that both subjects suppressed their vestibular information presumably as an effect of vestibular deconditioning training before flight. In addition as a consequence of the flight experience one subject continued to ignore it several days after the flight.

Transition from self tilt to object tilt during maintained lateral tilt in parabolic flight.

19 young healthy subjects were subjected to parabolic rollercoaster flight. A horizontal luminous line was seen by the subjects in a headfixed goggle device. During the hypergravic phases of parabolic flight the luminous line seemed to rotate into and during the hypogravic phase against the direction of static head tilt. Ocular counter rotation and activity of the neck position receptors cannot explain these subjective rotations. We conclude that information from the otolith system, converging with visual information within the brain, dislocated the headfixed visual target line. While the retinal image of the luminous line remains unchanged, loading and unloading the otoliths in parabolic flight changes the sensation of self tilt into object tilt, hereby subjectively rotating visual targets such as the luminous line.