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Cohen, Bernard

Publications and source records attributed to Cohen, Bernard.

Spatial orientation of optokinetic nystagmus and ocular pursuit during orbital space flight

On Earth, eye velocity of horizontal optokinetic nystagmus (OKN) orients to gravito-inertial acceleration (GIA), the sum of linear accelerations acting on the head and body. We determined whether adaptation to micro-gravity altered this orientation and whether ocular pursuit exhibited similar properties. Eye movements of four astronauts were recorded with three-dimensional video-oculography. Optokinetic stimuli were stripes moving horizontally, vertically, and obliquely at 30 degrees/s. Ocular pursuit was produced by a spot moving horizontally or vertically at 20 degrees/s. Subjects were either stationary or were centrifuged during OKN with 1 or 0.5 g of interaural or dorsoventral centripetal linear acceleration. Average eye position during OKN (the beating field) moved into the quick-phase direction by 10 degrees during lateral and upward field movement in all conditions. The beating field did not shift up during downward OKN on Earth, but there was a strong upward movement of the beating field (9 degrees) during downward OKN in the absence of gravity; this likely represents an adaptation to the lack of a vertical 1-g bias in-flight. The horizontal OKN velocity axis tilted 9 degrees in the roll plane toward the GIA during interaural centrifugation, both on Earth and in space. During oblique OKN, the velocity vector tilted towards the GIA in the roll plane when there was a disparity between the direction of stripe motion and the GIA, but not when the two were aligned. In contrast, dorsoventral acceleration tilted the horizontal OKN velocity vector 6 degrees in pitch away from the GIA. Roll tilts of the horizontal OKN velocity vector toward the GIA during interaural centrifugation are consistent with the orientation properties of velocity storage, but pitch tilts away from the GIA when centrifuged while supine are not. We speculate that visual suppression during OKN may have caused the velocity vector to tilt away from the GIA during dorsoventral centrifugation. Vertical OKN and ocular pursuit did not exhibit orientation toward the GIA in any condition. Static full-body roll tilts and centrifugation generating an equivalent interaural acceleration produced the same tilts in the horizontal OKN velocity before and after flight. Thus, the magnitude of tilt in OKN velocity was dependent on the magnitude of interaural linear acceleration, rather than the tilt of the GIA with regard to the head. These results favor a 'filter' model of spatial orientation in which orienting eye movements are proportional to the magnitude of low frequency interaural linear acceleration, rather than models that postulate an internal representation of gravity as the basis for spatial orientation.

Flight Experiment

The critical role of velocity storage in production of motion sickness

We propose that motion sickness is mediated through the orientation properties of velocity storage in the vestibular system that tend to align eye velocity produced by the angular vestibulo-ocular reflex (aVOR) with gravito-inertial acceleration (GIA). (GIA is the sum of the linear accelerations acting on the head. In the absence of translational accelerations, gravity is the GIA.) We further postulate that motion sickness produced by cross-coupled vestibular stimulation can be characterized by a metric composed of the disparity between the axis of eye rotation and the GIA, the strength of the response to angular motion, and the response duration, as determined by the central vestibular time constant, that is, by the time constant of velocity storage. The nodulus and uvula of the vestibulocerebellum are likely to be the central sites where the disparity is sensed, where the vestibular time constants are habituated, and where links are made to the autonomic system to produce the symptoms and signs.

Non-NASA Center

The relation of motion sickness to the spatial-temporal properties of velocity storage

Tilting the head in roll to or from the upright while rotating at a constant velocity (roll while rotating, RWR) alters the position of the semicircular canals relative to the axis of rotation. This produces vertical and horizontal nystagmus, disorientation, vertigo, and nausea. With recurrent exposure, subjects habituate and can make more head movements before experiencing overpowering motion sickness. We questioned whether promethazine lessened the vertigo or delayed the habituation, whether habituation of the vertigo was related to the central vestibular time constant, i.e., to the time constant of velocity storage, and whether the severity of the motion sickness was related to deviation of the axis of eye velocity from gravity. Sixteen subjects received promethazine and placebo in a double-blind, crossover study in two consecutive 4-day test series 1 month apart, termed series I and II. Horizontal and vertical eye movements were recorded with video-oculography while subjects performed roll head movements of approx. 45 degrees over 2 s to and from the upright position while being rotated at 138 degrees /s around a vertical axis. Motion sickness was scaled from 1 (no sickness) to an endpoint of 20, at which time the subject was too sick to continue or was about to vomit. Habituation was determined by the number of head movements that subjects made before reaching the maximum motion sickness score of 20. Head movements increased steadily in each session with repeated testing, and there was no difference between the number of head movements made by the promethazine and placebo groups. Horizontal and vertical angular vestibulo-ocular reflex (aVOR) time constants declined in each test, with the declines being closely correlated to the increase in the number of head movements. The strength of vertiginous sensation was associated with the amount of deviation of the axis of eye velocity from gravity; the larger the deviation of the eye velocity axis from gravity, the more severe the motion sickness. Thus, promethazine neither reduced the nausea associated with RWR, nor retarded or hastened habituation. The inverse relationship between the aVOR time constants and number of head movements to motion sickness, and the association of the severity of motion sickness with the extent, strength, and time of deviation of eye velocity from gravity supports the postulate that the spatiotemporal properties of velocity storage, which are processed between the nodulus and uvula of the vestibulocerebellum and the vestibular nuclei, are likely to represent the source of the conflict responsible for producing motion sickness.

NASA Discipline Neuroscience

Ocular Counter-Rolling During Centrifugation and Static Tilt

Activation of the gravity sensors in the inner ear-the otoliths-generates reflexes that act to maintain posture and gaze. Ocular counter-rolling (OCR) is an example of such a reflex. When the head is tilted to the side, the eyes rotate around the line of sight in the opposite direction (i.e., counter-rolling). While turning comers, undergoing centrifugation, or making side-to-side tilting head movements, the OCR reflex orients the eyes towards the sum of the accelerations from body movements and gravity. Deconditioning of otolith-mediated reflexes following adaptation to microgravity has been proposed as the basis of many of the postural, locomotor, and gaze control problems experienced by returning astronauts. Evidence suggests that OCR is reduced postflight in about 75% of astronauts tested; but the data are sparse, primarily due to difficulties in recording rotational eye movements. During the Neurolab mission, a short-arm human centrifuge was flown that generated sustained sideways accelerations of 0.5-G and one-G to the head and upper body. This produces OCR; and so for the first time, the responses to sustained centrifugation could be studied without the influence of Earth's gravity on the results. This allowed us to determine the relative importance of sideways and vertical acceleration in the generation of OCR. This also provided the first test of the effects of exposure to artificial gravity in space on postflight otolith-ocular reflexes. There was little difference between the responses to centrifugation in microgravity and on Earth. In both conditions, the induced OCR was roughly proportional to the applied acceleration, with the OCR magnitude during 0.5-G centrifugation approximately 60% of that generated during one-G centrifugation. The overall mean OCR from the four payload crewmembers in response to one-G of sideways acceleration was 5.7 plus or minus 1.1 degree (mean and SD) on Earth. Inflight one-G centrifugation generated 5.7 plus or minus 1.1 degree of OCR, which was a small but significant decrease in OCR magnitude. The postflight OCR was 5.9 plus or minus 1.4 degree, which was not significantly different from preflight values. During both 0.5-G and one-G centrifugation in microgravity, where the head vertical gravitational component was absent, the OCR magnitude was not significantly different from that produced by an equivalent acceleration during static tilt on Earth. This suggests that the larger OCR magnitude observed during centrifugation on Earth was due to the larger body vertical linear acceleration component, which may have activated either the otoliths or the body tilt receptors. In contrast to previous studies, there was no decrease in OCR gain postflight. Our findings raise the possibility that inflight exposure to artificial gravity, in the form of intermittent one-G and 0.5-G centripetal acceleration, may have been a countermeasure to deconditioning of otolith-based orientation reflexes.

Cohen, Bernard

Low-Frequency Otolith Function in Microgravity: A Re-Evaluation of the Otolith Tilt-Translation Reinterpretation (OTTR) Hypothesis

On Earth, the low-frequency afferent signal from the otoliths encodes head tilt with respect to the gravitational vertical, and the higher frequency components reflect both tilt and linear acceleration of the head. In microgravity, static tilt of the head does not influence otolith output, and the relationship between sensory input from the vestibular organs, and the visual, proprioceptive and somatosensory systems, would be disrupted. Several researchers have proposed that in 0-g this conflict may induce a reinterpretation of all otolith signals by the brain to encode only linear translation (otolith tilt-translation reinterpretation or OTTR). Ocular counter-rolling (OCR) is a low-frequency otolith-mediated reflex, which generates compensatory torsional eye movements (rotation about the visual axis) towards the spatial vertical during static roll tilt with a gain of approximately 10%. Transient linear acceleration and off-axis centrifugation at a constant angular velocity can also generate OCR. According to the OTTR hypothesis, OCR should be reduced in microgravity, and immediately upon return from a 0-g environment. Results to date have been inconclusive. OCR was reduced following the 10 day Spacelab-1 mission in response to leftward roll tilts (28-56% in 3 subjects and unchanged in one subject), and sinusoidal linear oscillations at 0.4 and 0.8 Hz. OCR gain declined 70% in four monkeys following a 14 day COSMOS mission. Following a 30 day MIR mission OCR gain decreased in one astronaut, but increased in two others following a 180 day mission. We have studied the affect of microgravity on low-frequency otolith function as part of a larger study of the interaction of vision and the vestibular system. This experiment (E-047) involved off-axis centrifugation of payload crewmembers and flew aboard the recent Neurolab mission (STS 90). Presented below are preliminary results focusing on perception and the OCR response during both centrifugation and static tilt.

Moore, Steven T.

Reduction of ocular counter-rolling by adaptation to space

We studied the three-dimensional vestibulo-ocular reflex (VOR) of rhesus monkeys before and after the COSMOS Biosatellite 2229 Mission of 1992-1993. This included tests of ocular counter-rolling (OCR), the gain of the vestibulo-ocular reflex (VOR), and spatial orientation of velocity storage. A four-axis vestibular and oculomotor stimulator was transported to the Institute of Biomedical Problems in Moscow for the pre- and postflight ground-based testing. Twelve normal juvenile male rhesus monkey were implanted surgically with eye coils and tested 60-90 days before spaceflight. Two monkey (7906 and 6151), selected from the twelve as flight animals, flew from 12/29/92 to 1/10/93. Upon recovery, they were tested for 11 days postflight along with three control animals. Compensatory ocular torsion was produced in two ways: (1) Lateral head tilts evoked OCR through otolith-ocular reflexes. OCR was also measured dynamically during off-vertical axis rotation (OVAR). (2) Rotation about a naso-occipital axis that was either vertical of horizontal elicited torsional nystagmus through semicircular canal-ocular reflexes (roll VOR). OCR from the otoliths was substantially reduced (70 percent) for 11 days after reentry on both modes of testing. The gain of the roll VOR was also decreased, but less than OCR. These data demonstrate that there was a long-lasting depression of torsional or roll eye movements after adaptation to microgravity in these monkeys, especially those movements produced by the otolith organs.

Dai, Mingjia

Vestibuloocular reflex of rhesus monkeys after spaceflight

The vestibuloocular reflex (VOR) of two rhesus monkeys was recorded before and after 14 days of spaceflight. The gain (eye velocity/head velocity) of the horizontal VOR, tested 15 and 18 h after landing, was approximately equal to preflight values. The dominant time constant of the animal tested 15 h after landing was equivalent to that before flight. During nystagmus induced by off-vertical axis rotation (OVAR), the latency, rising time constant, steady-state eye velocity, and phase of modulation in eye velocity and eye position with respect to head position were similar in both monkeys before and after flight. There were changes in the amplitude of modulation of horizontal eye velocity during steady-state OVAR and in the ability to discharge stored activity rapidly by tilting during postrotatory nystagmus (tilt dumping) after flight: OVAR modulations were larger, and tilt dumping was lost in the one animal tested on the day of landing and for several days thereafter. If the gain and time constant of the horizontal VOR exchange in microgravity, they must revert to normal soon after landing. The changes that were observed suggest that adaptation to microgravity had caused alterations in way that the central nervous system processes otolith input.

Cohen, Bernard

NASA supporting studies for microgravity research on eye movements

The purpose of the work on this project was to provide support for ground-based studies on the effects of gravity on eye movements. The effects of microgravity on the optokinetic eye movements of humans are investigated. OKN was induced by having subjects watch 3.3 deg stripes moving at 35 deg/s for 45 s in a binocular, head-fixed apparatus. The field (hor., 88 deg; vert., 72 deg), was rotated about axes that were upright or tilted 45 deg or 90 deg. The head was upright or tilted 45 deg on the body. Head-horizontal (yaw axis) and head-vertical (pitch axis) components of OKN were recorded with electro-oculography (EOG). Slow phase velocity vectors were determined relative to gravity. With the head upright, the axis of eye rotation during yaw axis OKN was coincident with the stimulus axis and the spatial vertical. With the head tilted 45 deg on the body, a persistent vertical component of eye velocity developed during yaw axis stimulation, and there was an average shift of the axis of eye rotation toward the spatial vertical of approximately 18 deg in six subjects. During oblique optokinetic stimulation with the head upright, the axis of eye rotation shifted 12 deg toward the spatial vertical. When the head was tilted, the axis of eye rotation rotated to the other side of the spatial vertical by 5.4 deg during the same oblique stimulation. This counter-rotation of the axis of eye rotation is similar to the 'Muller (E) effect', in which the perception of the upright counter-rotates to the opposite side of the spatial vertical when subjects are tilted in darkness. The data were simulated by a model of OKN. Despite the short OKAN time constants, strong horizontal to vertical cross-coupling was produced if the horizontal and vertical time constants were in proper ratio, and there was no suppression of nystagmus orthogonal to the stimulus direction. This shows that the spatial orientation of OKN can be due to a restructuring of the system matrix of velocity storage as a function of gravity. It is concluded that although human OKAN is weak, velocity storage orients the slow phase velocity of OKN towards the spatial vertical.

Cohen, Bernard