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The role of gravitation-dependent systems in visual tracking.

The effects of prolonged microgravity conditions on the performance of visual tracking functions such as fixational rotations of the eyes (saccades), smooth tracking of linear and curved movements of a foveal point stimulus, and following a vertical pendulum-like movement of foveoretinal optokinetic stimuli were studied. Experiments were performed on 31 cosmonauts in freefall conditions, in ten cases followed by additional studies after a cycle of head movements and in 14 after resting. These experiments showed that while intrinsic visual functions were retained in microgravity conditions, there were decreases in the precision and speed measures of all types of visual tracking (fixational rotations of the eyes, smooth tracking) and, in some cases, complete degradation of the smooth tracking reflex, an increase in the time taken to fix the gaze on a target (by factors of 2 or more), and decreases in the frequency of stimulus tracking. During the initial period of adaptation to the altered gravitational conditions and periodically during prolonged flight, the system of smooth visual tracking was found to undergo a transition to a strategy of saccadic approximation, in which gaze tracks the movement of the target using a set of macro- or microsaccadic movements. These impairments, seen in virtually all the cosmonauts, resulted from vestibular deprivation (functional deafferentation of the otolith input) in conditions of weightlessness, while in cosmonauts conceptualizing space on the basis of perceiving the positions of the feet and head additionally showed support-tactile deprivation.

Adaptation, Physiological↗

Gravitational haemodynamics and oedema prevention in the giraffe.

Because it is so tall, the giraffe, Giraffa camelopardalis, provides an important animal model for investigating adaptive mechanisms to orthostatic (gravitational) pressure changes. Previous physiological studies of the giraffe have concentrated on arterial blood pressures in the heart and neck. Briefly, these investigations revealed that arterial pressure near the giraffe heart is about twice that in humans, to provide more normal blood pressure and perfusion to the brain. Another important question is that of how giraffes avoid pooling of blood and tissue fluid (oedema) in dependent tissues of their extremities. As monitored by radiotelemetry, the blood and tissue fluid pressures that govern transcapillary exchange vary greatly with exercise. These pressures, combined with a tight skin layer, move fluid upward against gravity. Other mechanisms that prevent oedema include precapillary vasoconstriction and low permeability of capillaries to plasma proteins.

Animals↗

Gravitational scattering as a possible origin for giant planets at small stellar distances.

The recent discoveries of massive planetary companions orbiting several solar-type stars pose a conundrum. Conventional models for the formation of giant planets (such as Jupiter and Saturn) place such objects at distances of several astronomical units from the parent star, whereas all but one of the new objects are on orbits well inside 1 AU; these planets must therefore have originated at larger distances and subsequently migrated inwards. One suggested migration mechanism invokes tidal interactions between the planet and the evolving circumstellar disk. Such a mechanism results in planets with small, essentially circular orbits, which appears to be the case for many of the new planets. But two of the objects have substantial orbital eccentricities, which are difficult to reconcile with a tidal-linkage model. Here we describe an alternative model for planetary migration that can account for these large orbital eccentricities. If a system of three or more giant planets form about a star, their orbits may become unstable as they gain mass by accreting gas from the circumstellar disk; subsequent gravitational encounters among these planets can eject one from the system while placing the others into highly eccentric orbits both closer and farther from the star.

Evolution, Planetary↗

Gravitational evidence for an undifferentiated Callisto.

Before the arrival of the Galileo spacecraft at Jupiter, models for the interior structure of the four galilean satellites--Io, Europa, Ganymede and Callisto-ranged from uniform mixtures of rock and ice (that is, undifferentiated objects) or rocky cores surrounded by a mantle of water ice. Now it appears that Io has a large metallic core and that Ganymede is strongly differentiated, most probably into a three-layer structure consisting of a metallic core, a silicate mantle and a deep outer layer of ice. Direct information on the interior structure of Callisto determined from previous spacecraft fly-bys was essentially limited to an estimate of the mean density being intermediate between pure ice and pure rock. Here we report measurements of Callisto's gravitational field which reveal that, in contrast to Io and Ganymede, this galilean satellite is most probably a homogeneous object consisting of a solar mixture of 40% compressed ice and 60% rock (including iron and iron sulphide). Callisto's undifferentiated state is consistent with the apparent lack of an intrinsic magnetic field, and indicates that the outermost galilean satellite has not experienced a heating phase sufficiently high to separate its rock and metal components from the lighter ices.

Gravitation↗

Redistribution of pulmonary blood flow in pneumonia patients under the influence of gravitation.

In 120 pneumonia patients studied by means of chest radiography, an alteration of density of consolidated areas was detected, depending on the spatial position of the patient during investigation. The pneumonic consolidation shadow grew denser when the radiograph was taken in horizontal position with the damaged lung on the lower side. In contrast to this, the shadow density was markedly diminished when the patient was turned on the opposite (healthy) side. Such quick changing of the radiographic image is explained as a consequence of gravitational influence on the pulmonary circulation. In 20 patients with bronchial carcinoma, changing of the shadow density of the damaged area was very slight. It is believed that this difference may be useful in the identification and diagnosis of these diseases.

Acute Disease↗

The theoretical limits to the power output of a muscle-tendon complex with inertial and gravitational loads.

When a muscle delivers power to an inertial load through a spring, the peak power can exceed the maximum that the muscle alone could produce. Using normalized differential equations relating dimensionless quantities we show, by solving the equations either analytically or numerically, that one dimensionless constant (Xi), representing the inertial load, is sufficient to specify the behaviour during shortening of a muscle-tendon complex with linear force-velocity and force-extension properties. In the presence of gravity, an additional constant (Gamma), representing the gravitational acceleration, is required. Nonlinear force-velocity and force-extension relationships each introduce an additional constant, representing their curvature. In the absence of gravity the power output delivered to an inertial load is limited to approximately 1.4 times the maximum power of the muscle alone, and when gravity is present the power delivered is limited to approximately twice the power of muscle alone. These limits are found for the purely inertial load at Xi ca. 1 and with gravity acting at XiGamma = 0.5 with Xi arbitrarily small. The effects of nonlinear muscle and tendon properties tend to cancel each other out and do not produce large deviations from these optima. A lever system of constant ratio between muscle and load does not alter these limits. Cams and catches are required to exceed these limits and attain the high power outputs sometimes observed during explosive animal movement.

Animals↗

1/f ruffle oscillations in plasma membranes of amphibian epithelial cells under normal and inverted gravitational orientations.

Membrane ruffle fluctuations of amphibian epithelial cells A6 (CCL102) cultured in normal and upside down oriented plates have been analyzed through video microscopy. Our results reveal that their edge ruffle fluctuations exhibit a stochastic dynamics with 1/f(alpha) power spectrum over at least two decades at low frequencies and long range correlated, self-affine lateral border profiles. In a few and small areas of the membrane, probably nearby focal contacts, we found periodic oscillations which could be induced by myosin driven contraction of stress fibers. Furthermore, whereas the different gravitational orientations had none or little effect on the structure (power spectra and surface roughness) of these membrane ruffle fluctuations, their dynamic parameters were differentially affected. Indeed, the decay time of ruffles remained unchanged, but the period of lamellipodia oscillations near the focal adhesion points was significantly altered in A6 cells cultured upside down.

Amphibians↗

Pulmonary blood flow redistribution by increased gravitational force.

This study was undertaken to assess the influence of gravity on the distribution of pulmonary blood flow (PBF) using increased inertial force as a perturbation. PBF was studied in unanesthetized swine exposed to -Gx (dorsal-to-ventral direction, prone position), where G is the magnitude of the force of gravity at the surface of the Earth, on the Armstrong Laboratory Centrifuge at Brooks Air Force Base. PBF was measured using 15-micron fluorescent microspheres, a method with markedly enhanced spatial resolution. Each animal was exposed randomly to -1, -2, and -3 Gx. Pulmonary vascular pressures, cardiac output, heart rate, arterial blood gases, and PBF distribution were measured at each G level. Heterogeneity of PBF distribution as measured by the coefficient of variation of PBF distribution increased from 0.38 +/- 0.05 to 0.55 +/- 0.11 to 0.72 +/- 0.16 at -1, -2, and -3 Gx, respectively. At -1 Gx, PBF was greatest in the ventral and cranial and lowest in the dorsal and caudal regions of the lung. With increased -Gx, this gradient was augmented in both directions. Extrapolation of these values to 0 G predicts a slight dorsal (nondependent) region dominance of PBF and a coefficient of variation of 0.22 in microgravity. Analysis of variance revealed that a fixed component (vascular structure) accounted for 81% and nonstructure components (including gravity) accounted for the remaining 19% of the PBF variance across the entire experiment (all 3 gravitational levels). The results are inconsistent with the predictions of the zone model.

Acceleration↗

Role of the autonomic nervous system in push-pull gravitational stress in anesthetized rats.

Tolerance to +G(z) stress is reduced by preceding exposure to -G(z) (push-pull effect). The mechanism(s) responsible for this effect are not fully understood, although the arterial baroreceptor reflexes have been implicated. We investigated the integrative response of the autonomic nervous system by studying responses to gravitational stress before and after autonomic function was inhibited by hexamethonium in 10 isoflurane-anesthetized male and female Sprague-Dawley rats. Animals were restrained supine and subjected to two rotations imposed about the x-axis: 1) a control G profile consisting of rotation from 0 G(z) (+1 G(y)) to 90 degrees head-up tilt (+1 G(z)) for 10 s and 2) a push-pull G profile consisting of rotation from 0 G(z) to 90 degrees head-down tilt (-1 G(z)) for 2 s immediately preceding 10 s of +1 G(z) stress. Eight G profiles consisting of equal numbers of control and push-pull trials were imposed by using a counterbalanced design. We found that hexamethonium lowered baseline arterial pressure and abolished the push-pull effect. The lack of a push-pull effect after autonomic blockade persisted when arterial pressure was restored to baseline levels by phenylephrine infusion. Lowering baseline arterial pressure by sodium nitroprusside infusion or by hemorrhage when autonomic function was intact also abolished the push-pull effect. We conclude that intact autonomic function and a normal baseline arterial pressure are needed for expression of the push-pull effect in anesthetized rats subjected to tilting.

Animals↗

Gravitational deposition in a rhythmically expanding and contracting alveolus.

In a previous simulation, our laboratory demonstrated that the flow induced by a rhythmically expanding and contracting alveolus is highly complex (Haber S, Butler JP, Brenner H, Emanuel I, and Tsuda A, J Fluid Mech 405: 243-268, 2000). Based on these earlier findings, we hypothesize that the trajectories and deposition of aerosols inside the alveoli differ substantially from those previously predicted. To test this hypothesis, trajectories of fine particles (0.5-2.5 microm in diameter) moving in the foregoing alveolar flow field and simultaneously subjected to the gravity field were simulated. The results show that alveolar wall motion is crucial in determining the enhancement of aerosol deposition inside the alveoli. In particular, 0.5- to 1-microm-diameter particles are sensitive to the detailed alveolar flow structure (e.g., recirculating flow), as they undergo gravity-induced convective mixing and deposition. Accordingly, deposition concentrations within each alveolus are nonuniform, with preferentially higher densities near the alveolar entrance ring, consistent with physiological observations. Deposition patterns along the acinar tree are also nonuniform, with higher deposition in the first half of the acinar generations. This is a result of the combined effects of enhanced alveolar deposition in the proximal region of the acinus due to alveoli expansion and contraction and reduction in the number of particles remaining in the gas phase down the acinar tree. We conclude that the cyclically expanding and contracting motion of alveoli plays an important role in determining gravitational deposition in the pulmonary acinus.

Aerosols↗

Roles of gravitational cues and efference copy signals in the rotational updating of memory saccades.

Primates are able to localize a briefly flashed target despite intervening movements of the eyes, head, or body. This ability, often referred to as updating, requires extraretinal signals related to the intervening movement. With active roll rotations of the head from an upright position it has been shown that the updating mechanism is 3-dimensional, robust, and geometrically sophisticated. Here we examine whether such a rotational updating mechanism operates during passive motion both with and without inertial cues about head/body position in space. Subjects were rotated from either an upright or supine position, about a nasal-occipital axis, briefly shown a world-fixed target, rotated back to their original position, and then asked to saccade to the remembered target location. Using this paradigm, we tested subjects' abilities to update from various tilt angles (0, +/-30, +/-45, +/-90 degrees), to 8 target directions and 2 target eccentricities. In the upright condition, subjects accurately updated the remembered locations from all tilt angles independent of target direction or eccentricity. Slopes of directional errors versus tilt angle ranged from -0.011 to 0.15, and were significantly different from a slope of 1 (no compensation for head-in-space roll) and a slope of 0.9 (no compensation for eye-in-space roll). Because the eyes, head, and body were fixed throughout these passive movements, subjects could not use efference copies or neck proprioceptive cues to assess the amount of tilt, suggesting that vestibular signals and/or body proprioceptive cues suffice for updating. In the supine condition, where gravitational signals could not contribute, slopes ranged from 0.60 to 0.82, indicating poor updating performance. Thus information specifying the body's orientation relative to gravity is critical for maintaining spatial constancy and for distinguishing body-fixed versus world-fixed reference frames.

Adult↗

Independence of bilateral symmetry detection from a gravitational reference frame.

Data were collected during three orbital flights aboard the Russian MIR space station from eight cosmonauts tested on a bilateral symmetry detection task. It is known that on earth subjects' performance is significantly superior for a vertical or horizontal than for an obliquely oriented axis of symmetry, giving a so-called oblique effect. The present results show that this oblique effect did not disappear in microgravity. They confirm that the detection of visual symmetry is not tied to a gravitational reference frame. An unexpected result of the experiments was that practice reduced the reaction time for detection of asymmetrical patterns below that for symmetrical patterns. This result suggests the presence of two separate detection processes. The detection of symmetry, being useful for the efficient encoding of visual information, is well developed in naive subjects. Extensive practice on a symmetry-detection task, however, increases the efficacy of the asymmetry-detection process.

Adult↗

Gravitational effects on structure development in quenched complex fluids.

When binary liquid mixtures are cooled rapidly from a homogeneous phase into a two-phase system, domains of the two equilibrium phases form and grow (coarsen) with time. In the absence of an external forcing due to gravity or an imposed shear flow, a dynamic scaling regime emerges in which the domain morphology is statistically self-similar at different times with a length-scale that grows with time. In the presence of gravity, however, multiple length scales develop, with the system coarsening more rapidly in the direction of the force. The late-time behavior of such a system is characterized in this study by the calculation of anisotropic growth laws. Gravitation effects significantly affect scaling laws, even with small density mismatch, and the growth mechanism has some similarities to the sedimentation process. However, very few numerical studies have been made of such effects; this is one of the first.

Diffusion↗

Shape changes of osteoblastic cells under gravitational variations during parabolic flight--relationship with PGE2 synthesis.

The relationship existing between cell morphology and cell metabolism, and the role of mechanical load in bone remodelling are well-known. In osteoblasts, PGE2 mediates part of the response to mechanical stress and induce cell shape changes. We studied the influence of gravity variations on osteoblast morphology and its relationship with PGE2 synthesis during a parabolic flight. ROS 17/2.8 osteosarcoma cells flew 15 or 30 parabolae. We measured cell area and shape factor after fluorescein staining with a semi-automatic image analyser and PGE2 levels by RIA. Significant flight-induced shape changes consisted in a decrease in cell area and an increase in shape factor (cell irregularity), in some cells, as compared to ground controls. This heterogeneity in cell response might be explained by a cell-cycle sensitivity to mechanical stress. A 45 min pretreatment with indomethacin inhibited the flight-induced increase in cell irregularity whereas cell area remained decreased. PGE2 levels were higher in flight than in ground controls. Linear regression analysis showed a significant negative relationship between cell area and PGE2 synthesis. We concluded that ROS 17/2.8 are highly sensitive to gravitational variations and that PGE2 is partly implicated in cell shape changes observed during parabolic flight. However, other mechanisms than PGE2 synthesis condition ROS 17/2.8 morphology in response to mechanical changes.

Animals↗

The effect of gravitational acceleration on cardiac diastolic function: a biofluid mechanical perspective with initial results.

Echocardiographic measurements of astronaut cardiac function have documented an initial increase, followed by a progressive reduction in both left ventricular end-diastolic volume index and stroke volume with entry into microgravity (micro-G). The investigators hypothesize that the observed reduction in cardiac filling may, in part, be due to the absence of a gravitational acceleration dependent, intraventricular hydrostatic pressure difference in micro-G that exists in the ventricle in normal gravity (1-G) due to its size and anatomic orientation. This acceleration-dependent pressure difference, DeltaP(LV), between the base and the apex of the heart for the upright posture can be estimated to be 6660 dynes/cm(2) ( approximately 5 mm Hg) on Earth. DeltaP(LV) promotes cardiac diastolic filling on Earth, but is absent in micro-G. If the proposed hypothesis is correct, cardiac pumping performance would be diminished in micro-G. To test this hypothesis, ventricular function experiments were conducted in the 1-G environment using an artificial ventricle pumping on a mock circulation system with the longitudinal axis anatomically oriented for the upright posture at 45 degrees to the horizon. Additional measurements were made with the ventricle horizontally oriented to null DeltaP(LV)along the apex-base axis of the heart as would be the case for the supine posture, but resulting in a lesser hydrostatic pressure difference along the minor (anterior-posterior) axis. Comparative experiments were also conducted in the micro-G environment of orbital space flight on board the Space Shuttle. This paper reviews the use of an automated cardiovascular simulator flown on STS-85 and STS-95 as a Get Away Special payload to test this hypothesis. The simulator consisted of a pneumatically actuated, artificial ventricle connected to a closed-loop, fluid circuit with adjustable compliance and resistance elements to create physiologic pressure and flow conditions. Ventricular instrumentation included pressure transducers in the apex and base as well as immediately upstream of the inflow valve and downstream of the outflow valve, and a flow probe downstream of the outflow valve. By varying the circulating fluid volume, ventricular function could be determined for varying preload pressures at a regulated, mean afterload pressure of 95 mm Hg. This variation in preload condition permitted the construction of a ventricular function curve for the micro-G environment for comparison to the same curve for the 1-G environment. Data were collected from both missions at the upper end of the ventricular function curve. Experiment operation in the 1-G, supine orientation or in the micro-G environment eliminated the DeltaP(LV) observed in the 1-G, upright orientation. Consistent with the hypothesis, additional atrial pressure was required in micro-G to obtain stroke volumes and flow rates similar to those measured in 1-G for the upright posture. The necessary increase in atrial pressure was approximately 5 mm Hg in these experiments. In the same range of flow rates and stroke volumes, similar flows were observed in the 1-G supine posture for atrial pressures intermediate to the 1-G upright and micro-G values, also consistent with the hypothesis. Additional experiments on board the Space Shuttle are in preparation to gather data across the rest of the normal physiologic range of the ventricular function curve.

Diastole↗

Radiographic detection of mobilizable lung water: the gravitational shift test.

A radiographic method for detecting excessive lung water in patients with pulmonary infiltrates of uncertain cause is described. The gravitational shift test uses bedside frontal films before and after prolonged lateral decubitus positioning. Excess lung water is identified by detecting a shift in infiltrate to the dependent lung while the opposite side clears or remains stable. The test was evaluated in 33 patients with infiltrates of well defined etiology. Twelve patients had heart failure or fluid overload (edema): 14 had pulmonary infection or parenchymal damage (inflammation); and seven had inflammation plus edema. Infiltrates shifted to the dependent lung in 85% of patients with lung edema, but did not shift in 78% of patients with inflammatory disease. The test also detected excess lung water in six of seven patients with underlying inflammatory disease. Each patient with a positive test showed clinical, physiologic, and radiographic improvement after therapy directed at mobilizing excessive lung water. When the differential diagnosis of a diffuse infiltrate is in question, a positive test represents a strong indication for a trial of diuretic therapy.

Body Water↗

Erythrocytes sedimentation profiles under gravitational field as determined by He-Ne laser. II. Influence of erythrocyte shape.

The erythrocyte sedimentation profiles under gravitational field, by scanning the sample holder along the height and width, containing the blood samples with normal and crenated erythrocytes, are determined. The normal shape of erythrocytes has been altered by the controlled He-Ne laser exposures and this change, as observed microscopically, is similar to that as produced by other methods. At low exposure the erythrocytes have normal appearance, whereas, at 400 mJ/cm2, the percentage of crenated cells is 25 +/- 5 percent. It is observed that the modification of the shape influences the sedimentation characteristics of the erythrocytes. The erythrocytes tend to move faster after being exposed to lower exposure and slower after being exposed to higher exposure compared to that of normal erythrocytes. The possible mechanism associated with this change is discussed.

Blood Sedimentation↗