Charles M. Oman

M.I.T./Canadian vestibular experiments on the Spacelab-1 mission: 1. Sensory adaptation to weightlessness and readaptation to one-g: an overview

SummaryExperiments on human spatial orientation were conducted on four crewmembers of Space Shuttle Spacelab Mission 1. This introductory paper presents the conceptual background of the project, the relationship among the experiments and their relevance to a “sensory reinterpretation hypothesis”. Detailed experiment procedures and results are presented in the accompanying papers in this series. The overall findings are discussed in this article as they pertain to the following aspects of hypothesized sensory reinterpretation in weightlessness: 1) utricular otolith afferent signals are reinterpreted as indicating head translation rather than tilt, 2) sensitivity of reflex responses to footward acceleration is reduced, and 3) increased weighting is given to visual and tactile cues in orientation perception and posture control. Three subjects developed space motion sickness symptoms, which abated after several days. Head movements, as well as visual and tactile cues to orientation influenced symptoms in a manner consistent with the sensory-motor conflict theory of space motion sickness. Six short duration tests of motion sickness susceptibility, conducted pre-flight, failed to predict sickness intensity in weightlessness. An early otolith-spinal reflex, measured by electromyography from the gastrocnemius-soleus muscles during sudden footward acceleration, was inhibited immediately upon entering weightlessness and declined further during the flight, but was unchanged from pre-flight when measured shortly after return to earth. Dynamic visual-vestibular interaction was studied by measuring subjective roll self-motion created by looking into a spinning drum. Results suggest increased weighting of visual cues and reduced weighting of graviceptor signals in weightlessness. Following the 10 day flight, erect posture with eyes closed was disturbed for several days. Somewhat greater visual field dependence post-flight was observed for two of the crew. Post-flight tests using horizontal linear acceleration revealed an increased variance in detection of acceleration. The ability of the returned crew to use non-visual lateral acceleration cues for a manual control task appeared enhanced over their pre-flight ability for a few days after return.

The influence of semicircular canal morphology on endolymph flow dynamics. An anatomically descriptive mathematical model.

The classic Steinhausen/Groen mathematical description of endolymph flow in a toroidal semicircular canal is extended to the case where the size, shape, and curvature of the canal lumen change continuously through the duct, utricle, and ampulla. The resulting second-order differential equation has three coefficients, unlike the equation of a torsion pendulum, which has only two. The salient anatomical parameters which determine endolymph motion are: the length of the central streamline occupying the center of the canal lumen; the area enclosed by this streamline as projected into the plane of rotation; the average inverse cross-sectional area of the lumen (taken around the central streamline); and the average inverse squared cross-sectional area, weighted by a local wall shape factor. These parameters are evaluated and the average displacement of the face of the cupula is estimated for the human, guinea pig, and rat, based on new anatomical data presented in companion papers. The model predicts that the dynamic range of human average cupula motion lies between 520 A and 10 microns.

The Physiological Range of Pressure Difference and Cupula Deflections in the Human Semicircular Canal: Theoretical Considerations

The question of what constitutes the “physiologic” behavior of the cupula has been the subject of debate for some time. Calculations are presented which suggest that the cupula constitutes a biological pressure transducer with a threshold of the same order of magnitude as that of the auditory system. This indicates that it would probably be extremely easy to traumatize the cupula by exposing it to pressure gradients far beyond its normal dynamic range in the course of experimental preparations intended to visualize cupula motion. An estimate of the dynamic range of cupula motion may also be obtained which indicates that the center of the cupula may move as little as ten millimicrons at the 0.1 deg/sec2 acceleration levels commonly associated with behavioral thresholds. Accelerations of 30 deg/sec2 or sudden velocity changes of 600 deg/sec, typical of maximal self-induced head motions should produce cupula midpoint motions no greater than about three microns. Because of the small dynamic range of cupula mo...

M.I.T./Canadian vestibular experiments on the Spacelab-1 mission: 4. Space motion sickness: symptoms, stimuli, and predictability

SummarySpace sickness symptoms were observed by 4 specially trained observers on Spacelab-1. Three reported persistent symptoms, and vomited repeatedly during the first and/or second day of flight. Head movements on all axes were provocative, particularly in pitch and roll. Head acceleration data recorded from 2 symptomatic crewmen showed that after several hours of physical activity in orbit, symptoms appeared, and thereafter both crewmen were compelled to limit head movements. Firm body contact with motionless surfaces helped alleviate symptoms. When crewmembers floated into unfamiliar body orientations in the cabin, inherent ambiguities in static visual orientation cues sometimes produced spatial reorientation episodes which were also provocative. Symptoms largely resembled those of other forms of prolonged motion sickness, superimposed upon other symptoms attributable to fluid shift. All 4 eventually used anti-motion sickness drugs. When they did, vomiting frequency was reduced. By the 4th day, symptoms subsided, and head accelerations again increased in magnitude and variability. Sickness intensity in orbit was not predicted by statistically concordant results of 6 acute preflight susceptibility tests. However, results from a longer duration preflight prism goggles test showed an apparent correlation. All subjects were asymptomatic making head movements in parabolic flight 4 days after the mission, but not 1 year later. Overall, results support the view that space sickness is a motion sickness.

Dimensions of the Horizontal Semicircular Duct, Ampulla and Utricle in the Human

This study provides measurements in individual specimens of the sizes, cross-sectional shapes and areas all around the path of fluid flow through the human horizontal semicircular duct, ampulla and utricle. These data were obtained from multiple measurements on individual specimens which had been fixed by immersion in Karnovskys fixative and microdis-sected. The results are compared with similar measurements in the rat and guinea pig.

Element composition of inner ear lymphs in cats, lizards, and skates determined by electron probe microanalysis of liquid samples

SummarySiliconized, glass micropipets whose tips were filled with oil were used to obtain small (<100 nl) liquid samples from perilymphatic and endolymphatic regions of the inner ears of anesthetized animals: 3 cats, 19 alligator lizards (Gerrhonotus multicarinatus), and 8 skates (Raja erinacea). Samples of cerebrospinal fluid and seawater were also obtained for skates. Electron probe microanalysis was used to measure the concentrations of the following elements in each sample: K, Na, Cl, Ca, Mg, P, S. The Na and K concentrations in cat perilymph (Fig. 1 and Table 2) agree with previous estimates (Table 4) while endolymph samples show relatively low Na and high K concentrations. From a comparison of our results with previous work (Table 3), we infer that contamination of endolymph samples with perilymph is relatively low in our study, and that no large species difference in endolymph content is indicated by present data available for mammals. Our results show that Cl concentration is higher and Ca and Mg concentrations are lower in endolymph than in perilymph. The composition of perilymph in cats and alligator lizards is roughly the same (Figs. 1 and 2, Table 2). Uncontaminated endolymph samples in lizards were apparently difficult to obtain, although the compositions of a few samples suggest that endolymph K concentration is high and Na concentration is low. In skates the concentration of Na is nearly the same in the two inner ear lymphs (Fig. 3 and Table 2), in contrast to the roughly hundredfold ratio of perilymph to endolymph Na concentrations found in the higher vertebrates. The element composition of perilymph is correlated with the composition of seawater in which the skates were kept, whereas the endolymph composition shows no such correlation.

Sensory conflict theory and space sickness: our changing perspective

Motion sickness is a well-known nausea and vomiting syndrome whose physical signs include vomiting and retching, pallor, cold sweating, yawning, belching, flatulence, and decreased gastric tonus. Subjective symptoms include stomach discomfort, nausea, headache, feeling of warmth, and drowsiness. The research literature has been well reviewed (1-7). Over the past hundred years, our perspective on the kinds of sensory stimulation that can cause motion sickness has broadened considerably. Originally it was assumed that motion sickness can only be caused by motion and is due to mechanical stimulation of visceral afferents or changes in cerebral blood flow caused by body motion. It was eventually recognized that people who have no inner ear balance organ function are totally immune, indicating that the malady results specifically from vestibular over-stimulation. However, by 1970 it became apparent that a more comprehensive etiologic framework was needed: some forms of motion sickness were recognized in which head and body motion is normal (that is, spectacle sickness, space sickness) or even absent (flight simulator and Cinerama sickness). We experience vigorous vestibular stimulation when we run, jump, or dance, but these activities almost never make us sick. Claremont (8) had originally suggested that mo-

Dimensions of the Horizontal Semicircular Duct, Ampulla and Utricle in Rat and Guinea Pig

The dimensions of the membranous labyrinth partly determine the mechanical operation of the semicircular canal system. This study provides, for the first time, extensive measures in individual specimens of the sizes, cross-sectional shapes and areas of the horizontal semicircular duct, ampulla and utricle in the rat and the guinea pig. The membranous labyrinths were fixed in Karnovskys fixative, exposed, photographed, sectioned, oriented perpendicular to the line of sight and then measured using a calibrated graticule in the eye piece of an operating microscope. As well as the expected size differences between these species, there are major differences in the shape of the utricle.

Three dimensional spatial memory and learningin real and virtual environments

Human orientation and spatial cognition partlydepends on our ability to remember sets ofvisual landmarks and imagine their relationshipto us from a different viewpoint. We normallymake large body rotations only about a singleaxis which is aligned with gravity. However,astronauts who try to recognize environmentsrotated in 3 dimensions report that theirterrestrial ability to imagine the relativeorientation of remembered landmarks does noteasily generalize. The ability of humansubjects to learn to mentally rotate a simplearray of six objects around them was studied in1-G laboratory experiments. Subjects weretested in a cubic chamber (n = 73) and aequivalent virtual environment (n = 24),analogous to the interior of a space stationnode module. A picture of an object waspresented at the center of each wall. Subjectshad to memorize the spatial relationships amongthe six objects and learn to predict thedirection to a specific object if their bodywere in a specified 3D orientation. Percentcorrect learning curves and response times weremeasured. Most subjects achieved high accuracyfrom a given viewpoint within 20 trials,regardless of roll orientation, and learned asecond view direction with equal or greaterease. Performance of the subject group thatused a head mounted display/head tracker wasqualitatively similar to that of the secondgroup tested in a physical node simulator. Body position with respect to gravity had asignificant but minor effect on performance ofeach group, suggesting that results may alsoapply to weightless situations. A correlationwas found between task performance measures andconventional paper-and-pencil tests of fieldindependence and 2&3 dimensional figurerotation ability.

Spatial Orientation and Navigation in Microgravity

This chapter summarizes the spatial disorientation problems and navigation difficulties described by astronauts and cosmonauts, and relates them to research findings on orientation and navigation in humans and animals. Spacecraft crew are uniquely free to float in any relative orientation with respect to the cabin, and experience no vestibular and haptic cues that directly indicate the direction of “down”. They frequently traverse areas with inconsistently aligned visual vertical cues. As a result, most experience “Visual Reorientation Illusions” (VRIs) where the spacecraft floors, walls and ceiling surfaces exchange subjective identities. The illusion apparently results from a sudden reorientation of the observer’s allocentric reference frame. Normally this frame realigns to local interior surfaces, but in some cases it can jump to the Earth beyond, as with “Inversion Illusions” and EVA height vertigo. These perceptual illusions make it difficult for crew to maintain a veridical perception of orientation and place within the spacecraft, make them more reliant upon landmark and route strategies for 3D navigation, and can trigger space motion sickness. This chapter distinguishes VRIs and Inversion Illusions, based on firsthand descriptions from Vostok, Apollo, Skylab, Mir, Shuttle and International Space Station crew. Theories on human “gravireceptor” and “idiotropic” biases, visual “frame” and “polarity” cues, top-down processing effects on object orientation perception, mental rotation and “direction vertigo” are discussed and related to animal experiments on limbic head direction and place cell responses. It is argued that the exchange in perceived surface identity characteristic of human VRIs is caused by a reorientation of the unseen allocentric navigation plane used by CNS mechanisms coding place and direction, as evidenced in the animal models. Human VRI susceptibility continues even on long flights, perhaps because our orientation and navigation mechanisms evolved to principally support 2D navigation.