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The effect of gravity on the response of ventilation to abrupt change in FICO2.

Recently, Arieli and Farhi (1987) formulated a model for a previous suggestion made by their group that an increased ventilation as gravity increases is due to reduced perfusion of the respiratory center which causes an elevation of tissue PCO2. Extending the model to the dynamic response, we predict a slower ventilatory response to CO2 breathing as gravity increases. To test this prediction, ventilatory response to 5% CO2 was studied in 11 seated subjects at +1 and +2 Gz in a human centrifuge. Five of these subjects were studied at +3 Gz as well. In addition, ventilatory response to 5% CO2, using breath-by-breath analysis, was measured in three subjects in supine and upright postures. The ventilatory response (mainly through tidal volume) was faster as gravity increased from +1 to +2 and to +3 Gz, and from supine to the upright position. These findings disagree with the model prediction. Therefore, an alternative explanation is suggested based on the response of CO2 sensitive stretch receptors in the lung. Increased gravity causes increased ventilation, reduction of cardiac output and increased VA/Q mismatch; all enlarge the part of the lung with low CO2 where responsiveness of the CO2 sensitive stretch receptors is large.

Adult↗

Uvula-nodulus and gravity direction (a study on vertical optokinetic-oculomotor functions).

Some characteristics of vertical optokinetic nystagmus and after-nystagmus, in particular, its up/down gain asymmetry, are known to be influenced by gravity direction, which is basically perceived by otolith organs. However, how such an information of gravity direction is processed within the central nervous system is not well understood, even though the role of cerebellar uvula and nodulus in vertical eye movements has been identified for some time. Accordingly, in this study, vertical optokinetic nystagmus and after-nystagmus were studied in squirrel monkeys both in the upright and side-lateral positions (different gravity direction to the head), before and after uvulonodulectomy. When the optokinetic stimulus was given, the slow phase eye velocity of optokinetic nystagmus was significantly higher in the side-lateral position than that obtained in the upright position. This difference disappeared after uvulonodulectomy. When the initial slow phase eye velocity of optokinetic after-nystagmus was compared, the one obtained in the side-lateral position was significantly higher than that acquired in the upright position. This difference also disappeared after the uvulonodular ablation. Thus, in this study of vertical optokinetic nystagmus and after-nystagmus, the role of uvula and nodulus in relaying the information of gravity direction was confirmed.

Animals↗

Development and altered gravity dependent changes in glucose-6-phosphate dehydrogenase activity in the brain of the cichlid fish Oreochromis mossambicus.

Glucose-6-phosphate dehydrogenase activity was studied in the brain of the cichlid fish Oreochromis mossambicus during early ontogenetic development. In general a slight but continuous decrease in enzyme activity was found (9.5 +/- 0.5 nmol substrate cleaved per mg protein and per min at developmental stage 13 [= 1 day post hatch at 28 degrees C] to a value of 7.9 +/- 0.6 in adult brain). In order to investigate the possible influence of altered gravity during early ontogenetic brain development, fish larvae were exposed to an increased acceleration of three times earth gravity (3 g) or to functional weightlessness in a fast-rotating clinostat for 7 days. A significant increase of brain G6PDH activity of approx. 15% was found after exposure to hyper gravity, whereas a significant decrease of the enzyme activity, approximately 10%, was detected following functional weightlessness in respect to the corresponding 1 g controls. Analyses concerning the regain of normal control enzyme activity of the larvae revealed dramatic fluctuations within the first 5 h after exposure to an increased acceleration of 3 g. Thereafter, between day 1 and day 3 after exposure, brain glucose-6-phosphate dehydrogenase decreased slowly. At day 3 after exposure no further differences of the hyper-g larvae compared to the controls were found. Only slight changes in total brain glucose-6-phosphate dehydrogenase activity occur during ontogenetic development of cichlid fish. This suggests that a more or less constant enzyme activity is important during brain development, but is reacting very sensitively to changes in the environmental factor gravity.

Acceleration↗

Influence of gravity on the circadian timing system.

The circadian timing system (CTS) is responsible for daily temporal coordination of physiological and behavioral functions both internally and with the external environment. Experiments in altered gravitational environments have revealed changes in circadian rhythms of species ranging from fungi to primates. The altered gravitational environments examined included both the microgravity environment of spaceflight and hyperdynamic environments produced by centrifugation. Acute exposure to altered gravitational environments changed homeostatic parameters such as body temperature. These changes were time of day dependent. Exposure to gravitational alterations of relatively short duration produced changes in both the homeostatic level and the amplitude of circadian rhythms. Chronic exposure to a non-earth level of gravity resulted in changes in the period of the expressed rhythms as well as in the phase relationships between the rhythms and between the rhythms and the external environment. In addition, alterations in gravity appeared to act as a time cue for the CTS. Altered gravity also affected the sensitivity of the pacemaker to other aspects of the environment (i.e., light) and to shifts of time cues. Taken together, these studies lead to the conclusion that the CTS is indeed sensitive to gravity and its alterations. This finding has implications for both basic biology and space medicine.

Adaptation, Physiological↗

Experimental concept for examination of biological effects of magnetic field concealed by gravity.

Space is not only a place to study biological effects of gravity, but also provides unique opportunities to examine other environmental factors, where the biological actions are masked by gravity on the ground. Even the earth's magnetic field is steadily acting on living systems, and is known to influence many biological processes. A systematic survey and assessment of its action are difficult to conduct in the presence of dominant factors, such as gravity. Investigation of responses of biological systems against the combined environment of zero-gravity and zero-magnetic field might establish the baseline for the analysis of biological effects of magnetic factors. We propose, in this paper, an experimental concept in this context, together with a practical approach of the experiments, both in orbit and on the ground, with a thin magnetic shielding film. Plant epicotyl growth was taken as an exemplar index to evaluate technical and scientific feasibility of the proposed system concept.

Cotyledon↗

In vitro modeling of human tibial strains during exercise in micro-gravity.

Prolonged exposure to micro-gravity causes substantial bone loss (Leblanc et al., Journal of Bone Mineral Research 11 (1996) S323) and treadmill exercise under gravity replacement loads (GRLs) has been advocated as a countermeasure. To date, the magnitudes of GRLs employed for locomotion in space have been substantially less than the loads imposed in the earthbound 1G environment, which may account for the poor performance of locomotion as an intervention. The success of future treadmill interventions will likely require GRLs of greater magnitude. It is widely held that mechanical tissue strain is an important intermediary signal in the transduction pathway linking the external loading environment to bone maintenance and functional adaptation; yet, to our knowledge, no data exist linking alterations in external skeletal loading to alterations in bone strain. In this preliminary study, we used unique cadaver simulations of micro-gravity locomotion to determine relationships between localized tibial bone strains and external loading as a means to better predict the efficacy of future exercise interventions proposed for bone maintenance on orbit. Bone strain magnitudes in the distal tibia were found to be linearly related to ground reaction force magnitude (R(2)>0.7). Strain distributions indicated that the primary mode of tibial loading was in bending, with little variation in the neutral axis over the stance phase of gait. The greatest strains, as well as the greatest strain sensitivity to altered external loading, occurred within the anterior crest and posterior aspect of the tibia, the sites furthest removed from the neutral axis of bending. We established a technique for estimating local strain magnitudes from external loads, and equations for predicting strain during simulated micro-gravity walking are presented.

Aged↗

Histamine release from the hypothalamus induced by gravity change in rats and space motion sickness.

Freely moving rats were exposed to 2 g hypergravity in an animal centrifuge device to produce motion sickness. Histamine release from the anterior hypothalamus of the rats was measured in vivo with a microdialysis technique. After a 2-h load of 2 g hypergravity, rats ate kaolin. Because pica, eating a nonnutritive substance such as kaolin, is a behavioral index of motion sickness in rats, this finding indicates that the rats suffered from motion sickness. During 2 g hypergravity for 2-h, histamine release from the hypothalamus was transiently increased. In contrast, neither the transient increase of histamine release nor the kaolin consumption were induced by 2 g hypergravity in bilaterally labyrinthectomized rats. Pretreatment with alpha-fluoromethylhistidine, an inhibitor of histamine-synthesizing enzyme, decreased both the basal and hypergravity-induced releases of histamine from the hypothalamus and suppressed the kaolin consumption induced by hypergravity. Taken together, these findings suggest that the vestibular information of changes in gravity activate the histaminergic neuron system, resulting in the development of motion sickness. More prolonged stimulation, a 4-h load of 2 g hypergravity, induced significant increase of kaolin consumption on postdays 1-3, though rats ate kaolin on postdays 1-2 after 2 g hypergravity for 2 h. During 2 g hypergravity for 4 h, the initial transient increase of histamine release was followed by the gradual increase of histamine release after the end of centrifugation. It is suggested that rats adapted to the hypergravity environment after centrifugation for 4 h, but not 2 h, so that the change in gravity from 2 g to 1 g became a provocative stimulation. We, therefore, concluded that motion sickness in rats induced by a negative change in gravity can be used as a simulation of space motion sickness, which is induced by exposure to microgravity. Histaminergic activation in the development of motion sickness induced by negative change in gravity might be an underlying mechanism of space motion sickness.

Animals↗

A new goldfish model to evaluate pharmacokinetic and pharmacodynamic effects of drugs used for motion sickness in different gravity loads.

This paper proposes a new goldfish model to predict pharmacodynamic/pharmacokinetic effects of drugs used to treat motion sickness administered in differing gravity loads. The assumption of these experiments is that the vestibular system is dominant in producing motion sickness and that the visual system is secondary or of small import in the production of motion sickness. Studies will evaluate the parameter of gravity and the contribution of vision to the role of the neurovestibular system in the initiation of motion sickness with and without pharmacologic agents. Promethazine will be studied first. A comparison of data obtained in different groups of goldfish will be done (normal vs. acutely and chronically bilaterally blinded vs. sham operated). Some fish will be bilaterally blinded 10 months prior to initiation of the experiment (designated the chronically bilaterally blinded group of goldfish) to evaluate the neuroplasticity of the nervous system and the associated return of neurovestibular function. Data will be obtained under differing gravity loads with and without a pharmacological agent for motion sickness. Experiments will differentiate pharmacological effects on vision vs. neurovestibular input to motion sickness. Comparison of data obtained in the normal fish and in acutely and chronically bilaterally blinded fish with those obtained in fish with intact and denervated otoliths will differentiate if the visual or neurovestibular system is dominant in response to altered gravity and/or drugs. Experiments will contribute to validation of the goldfish as a model for humans since plasticity of the central nervous system allows astronauts to adapt to the altered visual stimulus conditions of 0-g. Space motion sickness may occur until such an adaptation is achieved.

Animals↗

The effect of gravity on surface temperature and net photosynthetic rate of plant leaves.

To clarify the effects of gravity on heat/gas exchange between plant leaves and the ambient air, the leaf temperatures and net photosynthetic rates of plant leaves were evaluated at 0.01, 1.0, 1.5 and 2.0 G of 20 seconds each during a parabolic airplane flight. Thermal images of leaves were captured using infrared thermography at an air temperature of 26 degrees C, a relative humidity of 15% and an irradiance of 260 W m-2. The net photosynthetic rates were determined by using a chamber method with an infrared gas analyzer at an air temperature of 20 degrees C, a relative humidity of 50% and a photosynthetic photon flux of 0.5 mmol m-2 s-1. The mean leaf temperature increased by 1 degree C and the net photosynthetic rate decreased by 13% with decreasing gravity levels from 1.0 to 0.01 G. The leaf temperature decreased by 0.5 degree C and the net photosynthetic rate increased by 7% with increasing gravity levels from 1.0 to 2.0 G. Heat/gas exchanges between leaves and the ambient air were more retarded at lower gravity levels. A restricted free air convection under microgravity conditions in space would limit plant growth by retarding heat and gas exchanges between leaves and the ambient air.

Gravitation↗

Influence of long-term hyper-gravity on the reactivity of succinic acid dehydrogenase and NADPH-diaphorase in the central nervous system of fish: a histochemical study.

In the course of a densitometric evaluation, the histochemically demonstrated reactivity of succinic acid dehydrogenase (SDH) and of NADPH-diaphorase (NADPHD) was determined in different brain nuclei of two teleost fish (cichlid fish Oreochromis mossambicus, swordtail fish Xiphophorus helleri), which had been kept under 3g hyper-gravity for 8 days. SDH was chosen since it is a rate limiting enzyme of the Krebs cycle and therefore it is regarded as a marker for metabolic and neuronal activity. NADPHD reactivity reflects the activity of nitric oxide synthase. Nitric oxide (NO) is a gaseous intercellular messenger that has been suggested to play a major role in several different in vivo models of neuronal plasticity including learning. Within particular vestibulum-connected brain centers, significant effects of hyper-gravity were obtained, e.g., in the magnocellular nucleus, a primary vestibular relay ganglion of the brain stem octavolateralis area, in the superior rectus subdivision of the oculomotoric nucleus and within cerebellar eurydendroid cells, which in teleosts possibly resemble the deep cerebellar nucleus of higher vertebrates. Non-vestibulum related nuclei did not respond to hyper-gravity in a significant way. The effect of hyper-gravity found was much less distinct in adult animals as compared to the circumstances seen in larval fish (Anken et al., Adv. Space Res. 17, 1996), possibly due to a development correlated loss of neuronal plasticity.

Afferent Pathways↗

Simulation of gaseous diffusion in partially saturated porous media under variable gravity with lattice Boltzmann methods.

Liquid distributions in unsaturated porous media under different gravitational accelerations and corresponding macroscopic gaseous diffusion coefficients were investigated to enhance understanding of plant growth conditions in microgravity. We used a single-component, multiphase lattice Boltzmann code to simulate liquid configurations in two-dimensional porous media at varying water contents for different gravity conditions and measured gas diffusion through the media using a multicomponent lattice Boltzmann code. The relative diffusion coefficients (D rel) for simulations with and without gravity as functions of air-filled porosity were in good agreement with measured data and established models. We found significant differences in liquid configuration in porous media, leading to reductions in D rel of up to 25% under zero gravity. The study highlights potential applications of the lattice Boltzmann method for rapid and cost-effective evaluation of alternative plant growth media designs under variable gravity.

Computer Simulation↗

[Measurement of gravity line position 15-25 years after Harrington-spondylodesis in adolescent idiopathic scoliosis].

AIM: A prospective analysis was performed to evaluate the relation between sagittal spinal morphology, gravity line position and morphological parameters of the pelvis and lumbar spine. METHOD: 25 patients with a mean follow-up of 23 (15-37) years after thoracic Harrington distraction spondylodesis were included. Standing lateral full spine radiographs were performed while patients were standing on a force plate. Pelvic angles according to Duval-Beaupere were measured. RESULTS: Pelvic incidence varied from 30 degrees to 76 degrees (mean 52.1 degrees + 12.5 degrees ). Mean sacral slope was 36.4 degrees (+ 7.8 degrees ). The average thoracic kyphosis (23.6 degrees + 14.3 degrees ) and lumbar lordosis (36.4 degrees + 9.2 degrees ) were diminished. The gravity line position was on average 10.9 mm (+ 21.6mm) posterior to the center of the femoral heads. CONCLUSION: Our method quantifies the relationship between the gravity line position and pelvic parameters according to Duval-Beaupere. The homogenous study group of patients with AIS after Harrington spondylodesis is characterized by decreased lumbar and thoracic profile and posterior displacement of the gravity line position.

Adolescent↗

A test of the gravity lens theory.

Naito and Cole [1994, in Contributions to Mathematical Psychology: Psychometrics and Methodology Eds G H Fischer and D Laming (New York: Springer)] provide a configuration which they describe as the Gravity Lens illusion. In this configuration, four small dots are presented in proximity to four large disks, and one is asked to compare the slope of an imaginary line which connects one pair of dots with the slope of a line which connects the other pair. In fact the slopes are the same, i.e. their axes are parallel, but because of the positioning of the large disks they appear to be at different orientations. Naito and Cole propose that the perceptual bias is analogous to the effects of gravity on the metrics of physical space, such that mental projections in the vicinity of a disk (or an open circle) are distorted just as the path of light is bent as it passes a massive body such as a star. Here we provide a simple test of this concept by having subjects judge alignments of dots which lie near tangents to a circle. Subjects were asked to project straight lines through pairs of stimulus dots, selecting and marking points in open space which were collinear with each pair. As would be predicted by the Gravity Lens theory, the locations selected by subjects were displaced from straight lines. However, the error magnitudes were substantially larger for judgments of dot pairs which had an oblique alignment, as compared with dot pairs which were aligned with a cardinal axis. This differential of effect as a function of stimulus orientation is not predicted by the gravity concept.

Form Perception↗

Muscle activity adapts to anti-gravity posture during pedalling in persons with post-stroke hemiplegia.

With hemiplegia following stroke, a person's movement response to anti-gravity posture often appears rigid and inflexible, exacerbating the motor dysfunction. A major determinant of pathological movement in anti-gravity postures is the failure to adapt muscle-activity patterns automatically to changes in posture. The aim of the present study was to determine whether the impaired motor performance observed when persons with hemiplegia pedal in a horizontal position is exacerbated at more vertical anti-gravity body orientations. Twelve healthy elderly subjects and 17 subjects with chronic (> 6 months) post-stroke hemiplegia participated in the study. Subjects pedalled a modified ergometer at different body orientations (from horizontal to vertical), maintaining the same workload, cadence, and hip and knee kinematics. Pedal reaction forces, and crank and pedal kinematics, were measured and then used to calculate the work done by each leg and their net positive and negative components. The EMG was recorded from four leg muscles (tibialis anterior, medial gastrocnemius, rectus femoris and biceps femoris). The main result from this study was that impaired plegic leg performance, as measured by net negative work done by the plegic leg and abnormal early rectus femoris activity, was exacerbated at the most vertical body orientations. However, contrary to the belief that muscle activity cannot adapt to anti-gravity postures, net positive work increased appropriately and EMG activity in all muscles showed modulated levels of activity similar to those in elderly control subjects. These results support the hypothesis that increased verticality exacerbates the already impaired movement performance. Yet, much of the motor response to verticality was flexible and appropriate, given the mechanics of the task.

Adaptation, Physiological↗

Perioperative halo-gravity traction in the treatment of severe scoliosis and kyphosis.

STUDY DESIGN: A retrospective analysis of patients that underwent perioperative halo-gravity traction as an adjunct to modern instrumentation methods in the treatment of severe scoliosis and kyphosis. OBJECTIVE: To review the clinical and radiographic results of perioperative halo-gravity traction in several time periods. SUMMARY OF BACKGROUND DATA: Few reports to our knowledge review the use of perioperative and intraoperative halo-gravity traction in this patient population. METHODS: A total of 33 patients with severe operative scoliosis, kyphoscoliosis, or kyphosis were studied based on hospital records, standing pretreatment, traction (before anterior/posterior fusion), postoperative (each stage), and final radiographs. Patients were analyzed by age at date of examination (range, 2-20 years; mean, 13.8 years), gender (18 male, 15 female), major coronal curve magnitude (range, 22 degrees-158 degrees; average, 84 degrees), major compensatory coronal curve magnitude (range, 8 degrees-123 degrees; average, 51 degrees), major sagittal curve magnitude (range, 13 degrees-143 degrees; average, 78 degrees), traction protocol, and procedure type. Halo-traction-related, short- and long-term complications were noted in each case. RESULTS: The major coronal curve reduced 38 degrees or 46% after posterior spinal fusion compared to pretreatment radiographs. At an average of 44 months radiographic follow-up (range, 24-107 months), the loss of correction averaged 7 degrees for major coronal curves and 4 degrees of thoracic kyphosis. Clinical complications were noted in the perioperative and long-term time periods. CONCLUSIONS: The treatment of severe scoliosis can be very challenging despite the benefits of modern instrumentation methods, especially if there is a significant kyphosis or a history of intraspinal pathology. Halo-gravity traction is a safe, well-tolerated method of applying gradual, sustained traction to maximize postoperative correction in this difficult population. There were no permanent neurologic deficits in this series.

Adolescent↗

Nonlinear effects due to gravity in a conical Hele-Shaw cell.

In this work we study the viscous fingering instability in a conical Hele-Shaw cell under the presence of gravity. We focus on understanding how the dynamical evolution of the fingering patterns is affected by the combined action of gravity and cell topology. Gravity-induced nonlinear effects are studied by a mode-coupling approach. Our results show that the interplay between gravity and cell topology leads to important effects, and profoundly modifies pattern evolution. We have found that the most dramatic consequences refer to finger tip behavior. Depending on the relative values of fluids' densities and viscosities, finger tip splitting reaches maximum intensity at well defined, preferred values of the cell opening angle. In fact, finger tip splitting can be completely replaced by finger tip sharpening as the cell angle is varied. Finger competition dynamics is also significantly changed: it is considerably enhanced (restrained) if the displaced fluid is more (less) dense.

Journal Article↗

Interface scaling in a two-dimensional porous medium under combined viscous, gravity, and capillary effects.

We have investigated experimentally the competition between viscous, capillary, and gravity forces during drainage in a two-dimensional synthetic porous medium. The displacement of a mixture of glycerol and water by air at constant withdrawal rate has been studied. The setup can be tilted to tune gravity, and pressure is recorded at the outlet of the model. Viscous forces tend to destabilize the displacement front into narrow fingers against the stabilizing effect of gravity. Subsequently, a viscous instability is observed for sufficiently large withdrawal speeds or sufficiently low gravity components on the model. We predict the scaling of the front width for stable situations and characterize it experimentally through analyses of the invasion front geometry and pressure recordings. The front width under stable displacement and the threshold for the instability are shown, both experimentally and theoretically, to be controlled by a dimensionless number F which is defined as the ratio of the effective fluid pressure drop (i.e., average hydrostatic pressure drop minus viscous pressure drop) at pore scale to the width of the fluctuations in the threshold capillary pressures.

Journal Article↗

Quasilocalization of gravity on a brane by resonant modes.

We examine the behavior of gravity in brane theories with extra dimensions in a nonfactorizable background geometry. We find that for metrics which are asymptotically flat far from the brane there is a resonant graviton mode at zero energy. The presence of this resonance ensures quasilocalization of gravity, whereby at intermediate scales the gravitational laws on the brane are approximately four dimensional. However, for scales larger than the lifetime of the graviton resonance the five-dimensional laws of gravity will be reproduced due to the decay of the four-dimensional graviton. We also give a simple classification of effective gravity theories for general background geometries.

Journal Article↗