Liposome formation in microgravity.

Liposomes are artificial vesicles with a phospholipid bilayer membrane. The formation of liposomes is a self-assembly process that is driven by the amphipathic nature of phospholipid molecules and can be observed during the removal of detergent from phospholipids dissolved in detergent micelles. As detergent concentration in the mixed micelles decreases, the non-polar tail regions of phospholipids produce a hydrophobic effect that drives the micelles to fuse and form planar bilayers in which phospholipids orient with tail regions to the center of the bilayer and polar head regions to the external surface. Remaining detergent molecules shield exposed edges of the bilayer sheet from the aqueous environment. Further removal of detergent leads to intramembrane folding and membrane folding and membrane vesiculation, forming liposomes. We have observed that the formation of liposomes is altered in microgravity. Liposomes that were formed at 1-g did not exceed 150 nm in diameter, whereas liposomes that were formed during spaceflight exhibited diameters up to 2000 nm. Using detergent-stabilized planar bilayers, we determined that the stage of liposome formation most influenced by gravity is membrane vesiculation. In addition, we found that small, equipment-induced fluid disturbances increased vesiculation and negated the size-enhancing effects of microgravity. However, these small disturbances had no effect on liposome size at 1-g, likely due to the presence of gravity-induced buoyancy-driven fluid flows (e.g., convection currents). Our results indicate that fluid disturbances, induced by gravity, influence the vesiculation of membranes and limit the diameter of forming liposomes.     

Gravity effects on biological systems.

Theoretical investigations of the membrane-solution interface predict different effects of gravity on vertically and horizontally oriented planar membranes. Single channel events of gramicidin incorporated into phosphatidylserine planar bilayer membranes were measured in 0.1 M KCl solution, pH 7, at room temperature. The potential difference across the membrane was set to +/- 70 mV. The mean channel current was observed to be about 20% higher in horizontally oriented membranes compared to vertical membranes. This is in good agreement with the theoretical considerations and demonstrates that gravity does affect membrane processes by interaction with the membrane-solution interface which is a ubiquitous structure in biological systems.     

Cellular responses to endogenous electrochemical gradients in morphological development.

Endogenous electric fields give vectorial direction to morphological development in Zea mays (sweet corn) in response to gravity. Endogenous electrical fields are important because of their ability to influence: 1) intercellular organization and development through their effects on the membrane potential, 2) direct effects such as electrophoresis of membrane components, and 3) both intracellular and extracellular transport of charged compounds. Their primary influence is in providing a vectorial dimension to the progression of one physiological state to another. Gravity perception and transduction in the mesocotyl of vascular plants is a complex interplay of electrical and chemical gradients which ultimately provide the driving force for the resulting growth curvature called gravitropism. Among the earliest events in gravitropism are changes in impedance, voltage, and conductance between the vascular stele and the growth tissues, the cortex, in the mesocotyl of corn shoots. In response to gravistimulation: 1) a potential develops which is vectorial and of sufficient magnitude to be a driving force for transport between the vascular stele and cortex, 2) the ionic conductance changes within seconds showing altered transport between the tissues, and 3) the impedance shows a transient biphasic response which indicates that the mobility of charges is altered following gravistimulation and is possibly the triggering event for the cascade of actions which leads to growth curvature.     

Gravity effects on membrane processes.

Application of the Gouy-Chapman-Debye-Hückel (GCDH) theory to a model membrane in contact with electrolytes of various concentrations and composition predict density variations within an interfacial layer. Assuming that on cellular dimensions hydrodynamics can be applied (the objections are briefly discussed) two types of gravity effects can be defined, 1. convection along the surface of vertically oriented membranes and 2. surface potential variations by layer deformations at horizontally oriented membranes. Both effects should affect transport across the layer to the membrane surface and across the membrane. According to the theoretical predictions first experiments with gramicidin channels incorporated into artificial phosphatidylserine bilayer membranes show a significant difference in single channel currents in vertical and horizontal membranes. The complexity of biological membrane functions requires investigation of isolated membrane surface reactions and transport systems to study the gravisensitivity for each process separately.     

Calcium signaling in plant cells in altered gravity.

Changes in the intracellular Ca2+ concentration in altered gravity (microgravity and clinostating) evidence that Ca2+ signaling can play a fundamental role in biological effects of microgravity. Calcium as a second messenger is known to play a crucial role in stimulus-response coupling for many plant cellular signaling pathways. Its messenger functions are realized by transient changes in the cytosolic ion concentration induced by a variety of internal and external stimuli such as light, hormones, temperature, anoxia, salinity, and gravity. Although the first data on the changes in the calcium balance in plant cells under the influence of altered gravity have appeared in 80th, a review highlighting the performed research and the possible significance of such Ca2+ changes in the structural and metabolic rearrangements of plant cells in altered gravity is still lacking. In this paper, an attempt was made to summarize the available experimental results and to consider some hypotheses in this field of research. It is proposed to distinguish between cell gravisensing and cell graviperception; the former is related to cell structure and metabolism stability in the gravitational field and their changes in microgravity (cells not specialized to gravity perception), the latter is related to active use of a gravitational stimulus by cells presumebly specialized to gravity perception for realization of normal space orientation, growth, and vital activity (gravitropism, gravitaxis) in plants. The main experimental data concerning both redistribution of free Ca2+ ions in plant cell organelles and the cell wall, and an increase in the intracellular Ca2+ concentration under the influence of altered gravity are presented. Based on the gravitational decompensation hypothesis, the consequence of events occurring in gravisensing cells not specialized to gravity perception under altered gravity are considered in the following order: changes in the cytoplasmic membrane surface tension --> alterations in the physicochemical properties of the membrane --> changes in membrane permeability, --> ion transport, membrane-bound enzyme activity, etc. --> metabolism rearrangements --> physiological responses. An analysis of data available on biological effects of altered gravity at the cellular level allows one to conclude that microgravity environment appears to affect cytoskeleton, carbohydrate and lipid metabolism, cell wall biogenesis via changes in enzyme activity and protein expression, with involvement of regulatory Ca2+ messenger system. Changes in Ca2+ influx/efflux and possible pathways of Ca2+ signaling in plant cell biochemical regulation in altered gravity are discussed.     

Evaluation of gravity-dependent membrane potential shift in Paramecium.

It is still debated whether or not gravity can stimulate unicellular organisms. This question may be settled by revealing changes in the membrane potential in a manner depending on the gravitational forces imposed on the cell. We estimated the gravity-dependent membrane potential shift to be about 1 mV G⁻¹ for Paramecium showing gravikinesis at 1–5 G, on the basis of measurements of gravity-induced changes in active propulsion and those of propulsive velocity in solutions, in which the membrane potential has been measured electrophysiologically. The shift in membrane potential to this extent may occur from mechanoreceptive changes in K⁺ or Ca²⁺ conductance by about 1% and might be at the limit of electrophysiological measurement using membrane potential-sensitive dyes. Our measurements of propulsive velocity vs membrane potential also suggested that the reported propulsive force of Paramecium measured in a solution of graded densities with the aid of a video centrifuge microscope at 350 G was 11 times as large as that for −29 mV, i.e., the resting membrane potential at [K⁺]_o = 1 mM and [Ca²⁺]_o = 1 mM, and, by extrapolation, that Paramecium was hyperpolarized to −60 mV by gravity stimulation of 100- G equivalent, the value corrected by considering the reduction of density difference between the interior and exterior of the cell in the graded density solution. The estimated shift of the membrane potential from −29 mV to −60 mV by 100- G equivalent stimulation, i.e., 0.3 mV G⁻¹, could reach the magnitude entirely feasible to be measured more directly.     

Long-range transport by chemical waves

A space experiment is proposed to study a transport mechanism that can become effective in chemically reacting liquids. In certain reaction systems that are kept far from thermodynamic equilibrium chemical waves can be generated spontaneously or triggered artificially. They do not involve transport of matter over long distances, but under appropriate conditions fluid particles encountered by a wave should be moved. The resulting transport depends critically on size and surface properties of the particles. The observation is greatly facilitated by low gravity.     

Physical parameters affecting living cells in space

The question is posed: Why does a living cell react to the absence of gravity? What sensors may it have? Does it note pressure, sedimentation, convection, or other parameters?

If somewhere in a liquid volume sodium ions are replaced by potassium ions, the density of the liquid changes locally: the heavier regions sink, the lighter regions rise. This may contribute to species transport, to the metabolism. Under microgravity this mechanism is strongly reduced. On the other hand, other reasons for convection like thermal and solutal interface convection are left. Do they affect species transport?

Another important effect of gravity is the hydrostatic pressure. On the macroscopic side, the pressure between our head and feet changes by 0.35 atmospheres. On the microscopic level the hydrostatic pressure on the upper half of a cell membrane is lower than on the lower half. This, by affecting the ion transport through the membrane, may change the surrounding electric potential. It has been suggested to be one of the reasons for graviperception.

Following the discussion of these and other effects possibly important in life sciences in space, an order of magnitude analysis of the residual accelerations tolerable during experiments in materials sciences is outlined. In the field of life sciences only rough estimates are available at present.     

Evaluation of gravity-dependent membrane potential shift in Paramecium

It is still debated whether or not gravity can stimulate unicellular organisms. This question may be settled by revealing changes in the membrane potential in a manner depending on the gravitational forces imposed on the cell. We estimated the gravity-dependent membrane potential shift to be about 1 mV G⁻¹ for Paramecium showing gravikinesis at 1–5 G, on the basis of measurements of gravity-induced changes in active propulsion and those of propulsive velocity in solutions, in which the membrane potential has been measured electrophysiologically. The shift in membrane potential to this extent may occur from mechanoreceptive changes in K⁺ or Ca²⁺ conductance by about 1% and might be at the limit of electrophysiological measurement using membrane potential-sensitive dyes. Our measurements of propulsive velocity vs membrane potential also suggested that the reported propulsive force of Paramecium measured in a solution of graded densities with the aid of a video centrifuge microscope at 350 G was 11 times as large as that for −29 mV, i.e., the resting membrane potential at [K⁺]_o = 1 mM and [Ca²⁺]_o = 1 mM, and, by extrapolation, that Paramecium was hyperpolarized to −60 mV by gravity stimulation of 100- G equivalent, the value corrected by considering the reduction of density difference between the interior and exterior of the cell in the graded density solution. The estimated shift of the membrane potential from −29 mV to −60 mV by 100- G equivalent stimulation, i.e., 0.3 mV G⁻¹, could reach the magnitude entirely feasible to be measured more directly.     

Recent development of Marangoni flows theory and experimental results

Microgravity fluid dynamics deals with the thermodynamics and dynamics of fluid systems in low-gravity environments and, thus, covers a rather large variety of subjects /1/. Among them, free convection in systems with fluid-fluid interfaces has received renewed attention because these systems are found in many materials sciences processes and exhibit a number of new, interesting phenomena. This paper surveys some recent studies by the author and his coworkers on the free convection occurring in a system of two interfacing, immiscible fluids subject to an imposed temperature gradient in a gravity environment. In such a system, three classes of free convection may be distinguished: natural, Marangoni, and combined free convection, according to whether it is caused by thermally induced buoyant forces, Marangoni stresses, or both. For each class one may further distinguish different regimes according to the relative importance of convective and diffusive transport of momentum and internal energy.First, the use of order of magnitude analysis is illustrated to identify and determine the types of flow regimes that can be established for each class of free convection. Subsequently, the attention is focused on the study of the main features of the two classes of flow regimes that can be established when the leading driving action is the Marangoni stress: Marangoni boundary layers and Marangoni-Stokes flow. Finally, a brief review of the experimental experience of the author's team is reported, looking in particular at the space experiments performed on board the missions SL1 and D1 of the Spacelab.     

Magnetic levitation-based Martian and Lunar gravity simulator.

Missions to Mars will subject living specimens to a range of low gravity environments. Deleterious biological effects of prolonged exposure to Martian gravity (0.38 g), Lunar gravity (0.17 g), and microgravity are expected, but the mechanisms involved and potential for remedies are unknown. We are proposing the development of a facility that provides a simulated Martian and Lunar gravity environment for experiments on biological systems in a well controlled laboratory setting. The magnetic adjustable gravity simulator will employ intense, inhomogeneous magnetic fields to exert magnetic body forces on a specimen that oppose the body force of gravity. By adjusting the magnetic field, it is possible to continuously adjust the total body force acting on a specimen. The simulator system considered consists of a superconducting solenoid with a room temperature bore sufficiently large to accommodate small whole organisms, cell cultures, and gravity sensitive bio-molecular solutions. It will have good optical access so that the organisms can be viewed in situ. This facility will be valuable for experimental observations and public demonstrations of systems in simulated reduced gravity.     

含滚阻摩擦平面多体系统的建模和数值方法

针对含滚阻和摩擦的平面多体系统,给出了通用的力学模型,利用第一类Lagrange方程建立了系统的动力学方程,为便于约束力的计算,利用局部方法列写约束方程,并以矩阵形式给出了该系统滚阻力偶和摩擦力的广义力.该系统动力学方程是非光滑的,通过建立滚阻定律的互补条件以及光滑铰链约束力的互补条件,给出了具有约束稳定化的基于事件驱动法的数值算法,将非光滑约束力、铰链约束力的求解以及滚轮滑动与粘滞的判断问题统一转化为一个线性互补问题进行求解.对两个算例进行了数值仿真,算例数值结果表明了该方法的可行性和有效性.     

The role of gravity in the evolutionary emergence of multicellular complexity: Microgravity effects on arthropod development and aging

While experiments carried out in Space with isolated cells have shown that eucaryotic cells are able to sense and respond to the absence of gravity by modifying their reactions, experiments in which more complex processes have been investigated, such as Biological Systems undergoing development under Microgravity, have been surprisingly unaffected by the space environment. This can be considered a curious result since all organisms are evolutionarily adapted to the current level of the gravity force in our planet and should eventually change if this parameter will vary in a permanent manner. In fact, the small effects of the modifications in gravity on development in short term experiments may be equivalent to the difficulties in detecting the involvement of other basic physical processes such as diffusion-controled auto-organizative reactions in currently developing biological systems. An apparent exception to this lack of effect is experiments where brine shrimp dormant gastrulae directly exposed to the space environment accumulate developmental defects as a consequence of cosmic irradiation. In this article we discuss the idea that at a certain stage during the evolutionary emergence of multicellular organisms the cues laid by generic forces such as gravity were involved in the evolutionary organization of these primitive organisms. As evolution proceed, these early mechanisms may have been obscured and/or made redundant by the appearance of new internal, environment-independent biological regulatory mechanisms. On the other hand, behavioral responses that may be important, for example, in setting the life-spans of organisms may still be more readily susceptible to manipulation by external cues as experiments carried out by our group in Space and on the ground with Drosophila melanogaster indicate.     

Attitude control actuator scaling laws for orbiting solar reflectors

The rapid evolution of in-orbit manufacturing will enable the fabrication of low-cost, large-scale space structures. In particular, the use of 3D printing technologies will remove traditional payload constraints associated with launch vehicles, due to fairing size and launch loads, thus allowing the construction of larger and lighter structures, such as orbiting solar reflectors. These structures will require efficient attitude control systems, able to provide the necessary torque for maneuvers and to counteract perturbations, such as gravity gradient and solar radiation pressure. In this paper, a top-level overview of actuator performances for orbiting solar reflectors is provided, and scaling laws associated with the required actuator mass and input power are developed. For each class of actuator, upper bounds on the maximum size of the structure which can be effectively controlled are presented. The results can also be extended to other classes of large planar Earth-pointing structures such as solar power satellites, solar sails, or large antennae.     

变重力场中散体颗粒堆的安息角及接触力分布

为研究重力与散体颗粒堆安息角的关系,针对颗粒系统的随机性和离散性特点,通过三维离散单元法建立颗粒的运动模型,对颗粒在不同重力场中的堆积过程进行了模拟,得出了颗粒堆的安息角及接触力概率分布.结果表明:颗粒堆内接触力构成的力链呈非规则网络状,其中,接触力近似对数正态分布,约65%接触力低于平均值;接触点中约70%处于临界滑移,其余接触点切向力与法向力的比值为均匀分布;不同重力场中颗粒堆接触力的分布规律具有相似性,即接触力相对重力无量纲化之后,其分布函数高度相符;颗粒堆的微观结构具有随机性差异,但是安息角不受重力大小的影响.     

Dust grain orbital behavior around ceres

Many asteroids show indications they have undergone impacts with meteoroid particles having radii between 0.01 m and 1 m. During such impacts, small dust grains will be ejected at the impact site. The possibility of these dust grains (with radii greater than 2.2 μm) forming a halo around a spherical asteroid (such as Ceres) is investigated using standard numerical integration techniques. The orbital elements, positions, and velocities are determined for particles with varying radii taking into account both the influence of gravity, radiation pressure, and the interplanetary magnetic field (for charged particles). Under the influence of these forces it is found that dust grains (under the appropriate conditions) can be injected into orbits with lifetimes in excess of one year. The lifetime of the orbits is shown to be highly dependent on the location of the ejection point as well as the angle between the surface normal and the ejection path. It is also shown that only particles ejected within 10° relative to the surface tangential survive more than a few hours and that the longest-lived particles originate along a line perpendicular to the Ceres-Sun line.     

Changes in membrane lipid composition in azuki bean epicotyls under hypergravity conditions: Possible role of membrane sterols in gravity resistance

Seedlings of azuki bean (Vigna angularis Ohwi et Ohashi) were cultivated under hypergravity conditions, and changes in membrane lipid composition in their epicotyls were analyzed. Under hypergravity conditions at 300g, the levels of total sterols, phospholipids, and fatty acids per fresh weight were kept higher, as compared with 1g controls. In particular, sterol levels were prominently increased by hypergravity. On the other hand, hypergravity did not clearly influence the levels of each phospholipid and glycolipid class, or their fatty acid compositions. Thus, the effect of hypergravity on membrane lipid metabolism was specific for sterol biosynthesis. In various regions of azuki epicotyls, high growth rate was associated with high sterol levels. Hypergravity suppressed elongation growth and stimulated lateral expansion of azuki epicotyls. In the presence of lovastatin, an inhibitor of sterol biosynthesis, at 30 μM, such changes in growth parameters occurred even under 1g conditions, suggesting that lovastatin made epicotyls hypersensitive to the gravitational force. These results support the hypothesis that membrane sterols are involved in maintenance of normal growth capacity of plant organs against gravity.     

Microtubule self-organisation depends upon gravity.

The molecular processes by which gravity is transduced into biological systems are poorly, if at all, understood. Under equilibrium conditions, chemical and biochemical structures do not depend upon gravity. It has been proposed that biological systems might show a gravity dependence by way of the bifurcation properties of certain types of non-linear chemical reactions that are far-from-equilibrium. We have found that in-vitro preparations of microtubules, an important element of the cellular cytoskeleton, show this type of behaviour. On earth, the solutions show macroscopic self-ordering, and the morphology of the structures that form depend upon the orientation of the sample with respect to gravity at a critical moment at an early stage in the development of the self-organised state. An experiment carried out in a sounding rocket, showed that as predicted by theories of this type, no self-organisation occurs when the microtubules are assembled under low gravity conditions. This is an experimental demonstration of how a very simple biochemical system, containing only two molecules, can be gravity sensitive. At a molecular level this behaviour results from an interaction of gravity with macroscopic concentration and density fluctuations that arise from the processes of microtubule contraction and elongation.     

Super-helix model: a physiological model for gravitaxis of Paramecium.

A new model explaining the gravitactic behavior of Paramecium is derived on the basis of its mechanism of gravity sensing. Paramecium is know to have depolarizing mechanoreceptor ion channels in the anterior and hyperpolarizing channels in the posterior of the cell. This arrangement may lead to bidirectional changes of the membrane potential due to the selective deformation of the anterior and posterior cell membrane responding to the orientation of the cell with respect to the gravity vector; i.e., negative- and positive-going shifts of the potential due to the upward and downward orientation, respectively. The orientation dependent changes in membrane potential, in combination with the close coupling between the membrane potential and ciliary locomotor activity, may allow the changes in swimming direction along the otherwise simple helical swimming path in the following manner: an upward shift of the axis of helical swimming occurs by decreasing the pitch angle due to channel-dependent hyperpolarization in upward-orienting cells, and an upward shift of the swimming helix occurs by increasing the cell's pitch angle due to depolarization in downward-orienting cells. Computer simulation of the model demonstrated that the cell can swim upward along the "super-helical" trajectory consisting of a small helix winding helically along an axis parallel to the gravity vector.     

颗粒气体的类气液相变

颗粒体系是远离平衡态的复杂耗散体系,在外力驱动下,能表现出类似于固体、液体或气体的特性.在稀疏的颗粒气体中,由于耗散,体系中常常会形成颗粒在局部的凝聚.这种凝聚行为具有自发对称性破缺的特征,使得颗粒气体中自动形成高密度区域和低密度区域共存的非均匀状态,类似于气体中液滴的形成.本文利用零重力条件下的三维颗粒气体理论模型进行计算和分析,揭示了颗粒气体这一相变行为的不稳定性根源及类气液相变本质,给出了颗粒气体中这种相分离发生的临界条件,并通过分子动力学模拟进行检验,结果从定性上和定量上都能够很好地吻合,为进一步的空间实验提供了理论依据和相关实验参数.