Invited review: gravitational biology of the neuromotor systems: a perspective to the next era.

Earth's gravity has had a significant impact on the designs of the neuromotor systems that have evolved. Early indications are that gravity also plays a key role in the ontogenesis of some of these design features. The purpose of the present review is not to assess and interpret a body of knowledge in the usual sense of a review but to look ahead, given some of the general concepts that have evolved and observations made to date, which can guide our future approach to gravitational biology. We are now approaching an era in gravitational biology during which well-controlled experiments can be conducted for sustained periods in a microgravity environment. Thus it is now possible to study in greater detail the role of gravity in phylogenesis and ontogenesis. Experiments can range from those conducted on the simplest levels of organization of the components that comprise the neuromotor system to those conducted on the whole organism. Generally, the impact of Earth's gravitational environment on living systems becomes more complex as the level of integration of the biological phenomenon of interest increases. Studies of the effects of gravitational vectors on neuromotor systems have and should continue to provide unique insight into these mechanisms that control and maintain neural control systems designed to function in Earth's gravitational environment. A number of examples are given of how a gravitational biology perspective can lead to a clearer understanding of neuromotor disorders. Furthermore, the technologies developed for spaceflight studies have contributed and should continue to contribute to studies of motor dysfunctions, such as spinal cord injury and stroke. Disorders associated with energy support and delivery systems and how these functions are altered by sedentary life styles at 1 G and by space travel in a microgravity environment are also discussed.     

Application of a Theory for Circumnutations to Geotropic Movements

From a previously published theory (Israelsson and Johnsson 1967) for circumnutations in Helianthus annuus it is possible to predict the geotropical curvatures of the hypocotyls. This extension of the theory is given in the present paper and some geotropical experiments are performed and discussed. The agreement between the theory and the experiments has been verified in the case of gravitational stimulation during relatively short stimulation periods, in the case of continuous gravitational stimulation, etc. Restrictions in the proposed model are discussed.     

A gravitational impulse model predicts collision impulse and mechanical work during a step-to-step transition

The simplest walking model, which assumes an instantaneous collision with negligible gravity effect, is limited in its representation of the collision mechanics of human gaits because the actual step-to-step transition occurs over a finite duration of time with finite impulsive ground reaction forces (GRFs) that have the same order of magnitude as the gravitational force. In this study, we propose a new collision model that includes the contribution of the gravitational impulse to the momentum change of the center of mass (COM) during a step-to-step transition. To validate the model, we measured the GRFs of six subjects' over-ground walking at five different gait speeds and calculated the collision impulses and mechanical work. The data showed a significant contribution of the gravitational impulse to the momentum change during collision. To compensate for the gravity, the magnitudes of collision impulse and COM work were estimated to be much greater than in previous predictions. Consistent with the model prediction, push-off propulsion fully compensated for the collision loss, implying the step-to-step transition occurred in an energetically optimal manner. The new model predicted a moderate change in the collision mechanics with gait speed, which seems to be physiologically achievable. The gravitational collision model enables us to better understand collision dynamics during a step-to-step transition.     

Gravitational Symmetry Breaking Leads to a Polar Liquid Crystal Phase of Microtubules In Vitro

Recent space-flight experiments performed by Tabony's team provided further evidence that a microgravity environment strongly affects the spatio-temporal organization of microtubule assemblies. Characteristic time and length scales were found that govern the organization of oriented bundles under Earth's gravitational field (GF). No such organization has been observed in a microgravity environment. This paper discusses physical mechanisms resulting in pattern formation under gravity and its disappearance in microgravity. The subtle interplay between chemical kinetics, diffusion, gravitational drift, thermal fluctuations, electrostatic interactions and liquid crystalline characteristics provides a plausible scenario.     

Sedimentation of a suspension in a centrifugal field.

To model centrifugal sedimentation of biological suspensions, the time history of sedimentation of particles in a centrifugal field was considered for two geometries: a tube and a cylindrical container. The Kynch theory for batch gravitational settling in Cartesian coordinates based on mass conservation was extended to include a centrifugal sedimentation force, cylindrical coordinates, and the Hawksley-Vand hindered settling model. The resulting quasi-linear partial differential equation was solved by the method of characteristics. The combination of radial dependence of the sedimentation force and cylindrical geometry in the centrifugal case results in several differences in the time-position history diagram of the sedimentation process compared to the gravitational case. First, instead of a region of uniform concentration equal to the initial concentration, a region of concentration that is continuously decreasing with time results. Second, in the region of particle accumulation, curved constant concentration contours result instead of straight lines. Finally, a secondary shock that is dependent upon the initial concentration and the radius ratio of the rotating vessel appears in the centrifugal case. The time history of the concentration for a particle suspension with an initial concentration typical of blood is presented.     

The scaling rule for environmental organizing systems in a gravitational field

A recent thermodynamics and information study examined the basis of a scaling rule for simple living organisms. The present paper examines a scaling rule for the relationship between the integrated scaled metabolic energy and the mass of a system for a wide range of masses, from animals to the 4He cores of main-sequence stars, considering the effect of gravitational energy. The expected specific scaled energy for animals and the 4He cores of main-sequence stars is 1600 times greater than the specific scaled energy for fundamental living organisms, such as unicellular organisms. This difference results from their organization in a gravitational field or the lack thereof.     

基于引力模型的林木竞争分析

引力模型是否可以应用于森林群落林木竞争关系分析是值得研究的问题。基于引力模型建立林木相对活力圈能反映竞争木活力大小,基于竞争木的相对活力圈建立引力竞争指数能准确反映林木生长与林木竞争的关系。以浙江省天目山国家级自然保护区针阔混交林为研究对象,将V_Hegyi竞争指数、引力竞争指数分别与胸径进行相关分析,胸高断面积生长量分别与2种竞争指数进行相关分析,胸径生长率与2期引力竞争指数的比值(2021年与2006年的引力竞争指数之比)进行相关分析,此外,对相对活力圈直径与胸径进行相关分析,并比较分析了活立木与枯死木的竞争指数大小。结果表明: 2种竞争指数与胸径均呈显著负相关,且均服从幂函数关系。林木胸高断面积生长量与2种竞争指数均呈显著负相关,但引力竞争指数比V_Hegyi竞争指数更能反映林木生长与林木竞争的关系。相对于V_Hegyi竞争指数的比值,2期引力竞争指数的比值更能说明林木生长与林木竞争的关系。在针阔混交林中,阔叶树种的生长与竞争的相关性>针叶树种生长与竞争的相关性。林木枯损受竞争的显著影响。林木相对活力圈大小与林木胸径大小呈显著负相关。引力模型是反映空间相互作用的重要模型之一,可以应用于林木竞争关系的研究,且基于引力模型建立的引力竞争指数可以作为评价林木竞争和林木活力的一个空间结构指标,比V_Hegyi竞争指数更能反映林木生长与林木竞争的关系。     

Direction finding by hornets in a vertically rotating centrifuge

In a vertically rotating centrifuge, the direction of the resultant gravitational and centrifugal forces is constantly changing. Hornets placed in such a centrifuge will build their combs in the direction of the resultant only if the centrifuge is stopped every day and left in the same position for at least half an hour, because during the cessation of motion, they presumably “learn” some geometrical cues which enable them to determine the preferred angle of building. Hornets can detect and respond to a centrifugal force as small as 0·18% of the earth's gravitational force. At a rotational rate of 1/8 of a revolution per minute there was no comb construction whatsoever and hornet mortality rate was 100% within three days.     

Gravity is a minor determinant of pulmonary blood flow distribution

Regional pulmonary blood flow in dogs under zone 3 conditions was measured in supine and prone postures to evaluate the linear gravitational model of perfusion distribution. Flow to regions of lung that were 1.9 cm3 in volume was determined by injection of radiolabeled microspheres in both postures. There was marked perfusion heterogeneity within isogravitational planes (coefficient of variation = 42.5%) as well as within gravitational planes (coefficient of variation = 44.2 and 39.2% in supine and prone postures, respectively; P = 0.02). On average, vertical height explained only 5.8 and 2.4% of the flow variability in the supine and prone postures, respectively. Whereas the gravitational model predicts that regional flows should be negatively correlated when measured in supine and prone postures, flows in the two postures were positively correlated, with an r2 of 0.708 +/- 0.050. Regional perfusion as a function of distance from the center of a lung explained 13.4 and 10.8% of the flow variability in the supine and prone postures, respectively. A linear combination of vertical height and radial distance from the centers of each lung provided a better-fitting model but still explained only 20.0 and 12.0% of the flow variability in the supine and prone postures, respectively. The entire lung was searched for a region of contiguous lung pieces (22.8 cm3) with high flow. Such a region was found in the dorsal area of the lower lobes in six of seven animals, and flow to this region was independent of posture. Under zone 3 conditions, neither gravity nor radial location is the principal determinant of regional perfusion distribution in supine and prone dogs.     

Gravitaxis in Drosophila melanogaster: a forward genetic screen

Perception of the earth's gravitational force is essential for most forms of animal life. However, little is known of the molecular mechanisms and neuronal circuitry underlying gravitational responses. A forward genetic screen using Drosophila melanogaster that provides insight into these characteristics is described here. Vertical choice mazes combined with additional behavioral assays were used to identify mutants specifically affected in gravitaxic responses. Twenty-three mutants were selected for molecular analysis. As a result, 18 candidate genes are now implicated in the gravitaxic behavior of flies. Many of these genes have orthologs across the animal kingdom, while some are more specific to Drosophila and invertebrates. One gene (yuri) located close to a known locus for gravitaxis has been the subject of more extensive analysis including confirmation by transgenic rescue.     

Continuous optical discharge in a gravitational field

A two-dimensional radiative gas-dynamic model is applied to calculating the parameters of a continuous optical discharge in a vertical focused CO₂ laser beam in air at atmospheric pressure in the Earth’s gravitational field.     

Resistance of plants to gravitational force

Developing resistance to gravitational force is a critical response for terrestrial plants to survive under 1 × g conditions. We have termed this reaction “gravity resistance” and have analyzed its nature and mechanisms using hypergravity conditions produced by centrifugation and microgravity conditions in space. Our results indicate that plants develop a short and thick body and increase cell wall rigidity to resist gravitational force. The modification of body shape is brought about by the rapid reorientation of cortical microtubules that is caused by the action of microtubule-associated proteins in response to the magnitude of the gravitational force. The modification of cell wall rigidity is regulated by changes in cell wall metabolism that are caused by alterations in the levels of cell wall enzymes and in the pH of apoplastic fluid (cell wall fluid). Mechanoreceptors on the plasma membrane may be involved in the perception of the gravitational force. In this review, we discuss methods for altering gravitational conditions and describe the nature and mechanisms of gravity resistance in plants.     

Longitudinal low-frequency waves in the flows of self-gravitating charged dust grains in an electron-ion plasma

A study is made of the features of wave processes in the individual flows of self-gravitating dust grains in a plasma and the electric and gravitational interactions in a system of several dusty plasma flows. It is shown that, in a dusty plasma, Debye screening can substantially weaken the electric coupling between the beams of self-gravitating grains, without affecting the gravitational forces between them, and that the electrostatic perturbations are exchanged between the grain flows via gravitational fields, as happens in vacuum.     

Subject-specific musculoskeletal model of the lower limb in a lying and standing position

Accurate estimation of joint loads implies using subject-specific musculoskeletal models. Moreover, as the lines of action of the muscles are dictated by the soft tissues, which are in turn influenced by gravitational forces, we developed a method to build subject-specific models of the lower limb in a functional standing position. Bones and skin envelope were obtained in a standing position, whereas muscles and a set of bony landmarks were obtained from conventional magnetic resonance images in a lying position. These muscles were merged with the subject-specific skeletal model using a nonlinear transformation, taking into account soft tissue movements and gravitational effects. Seven asymptomatic lower limbs were modelled using this method, and results showed realistic deformations. Comparing the subject-specific skeletal model to a scaled reference model rendered differences in terms of muscle length up to 4% and in terms of moment arm for adductor muscles up to 30%. These preliminary findings enlightened the importance of subject-specific modelling in a functional position.     

Gravity and light effects on the circadian clock of a desert beetle,Trigonoscelis gigas

Circadian function is affected by exposure to altered ambient force environments. Under non-earth gravitational fields, both basic features of circadian rhythms and the expression of the clock responsible for these rhythms are altered. We examined the activity rhythm of the tenebrionid beetle, Trigonoscelis gigas, in conditions of microgravity (microG; spaceflight), earth's gravity (1 G) and 2 G (centrifugation). Data were recorded under a light-dark cycle (LD), constant light (LL), and constant darkness (DD). Free-running period (tau) was significantly affected by both the gravitational field and ambient light intensity. In DD, tau was longer under 2 G than under either 1 G or microG. In addition, tauLL was significantly different from tauDD under microG and 1 G, but not under 2 G.     

A down to earth model of gravisensing or Newton's Law of Gravitation from the apple's perspective

The physiology of gravity perception in plants is examined and a model of gravitational pressure is explained and compared to the statolith model. The gravitational pressure model is based on studies of tension and compression of the plasma membrane against the extracellular matrix. Further studies examine the role of peptides or enzymes that inhibit a compression receptor and calcium channels.     

The effects of a change in gravity on the dynamics of prehension

The objective of this study was to measure the forces applied on an object manipulated in different gravitational fields attained during parabolic flights. Eight subjects participated flights (ES) and four were inexperienced (NES). They had to move continuously an instrumented object up and down in three different gravitational conditions (1 g, 1.8 g, 0 g). In 1 g, the grip force precisely anticipated the fluctuations of load force which was maximum and minimum at the bottom and at the top of the arm trajectory respectively. When the gravity changed (0 g and 1.8 g), the grip-load force coupling persisted for all the subjects from the first parabola. While the ES immediately exerted a grip force appropriate to the gravity, the NES dramatically increased their grip when faced with hyper and microgravity for the first time. Then, they progressively released their grip until a continuous grip-load force relationship with regard to 1 g was established after the fifth parabola. We suggest that each new gravitational field is rapidly incorporated into an internal model within the CNS which can then be reused as required by the occasion.     

Types of biological reactions to the gravitation loads

The nature of adaptation to gravitational loads is reviewed. Topics include an organism's antigravitation function, exposure to gravitational loads, types of physiological reactions, and results of adaptation.     

Simulation of gravitational field variation on fluid-filled biological membranes

The knowledge of the behavior of biological organs in a gravitational field is important to understand the functioning of the human body in the aerospace environment. The disturbances in biological transport processes in microgravity have indicated adverse effects on humans engaged in space operations. The relationship between the deformations in the biological organs and the transport phenomena that take place in them has been long established and widely reported in biological sciences and engineering literature. A number of soft tissue organs such as brain, lungs, heart, kidney, bladder, stomach, and the circulatory system can be modeled as fluid-filled membranes. In this investigation, a mathematical model of a fluid-filled biological membrane is developed, and its deformation and spatial configuration in a variable gravitational field are calculated. The variation in the gravitational field in the range 1g to zero-g is simulated by partial submergence of the fluid-filled membrane which, by virtue of buoyancy, gains an effective density as if it is in a different gravitational field. The equations of motion are derived using the theory of large elastic deformations and numerically solved in conjunction with a constitutive equation suitably selected for the biological membrane.     

On the dynamics of a spherical scaffold in rotating bioreactors

We analyze the dynamics of a spherical scaffold in rotating bioreactors (or clinostats). The idealized clinostat environment consists of a purely rotational flow that is perpendicular to a gravitational field. We confirm through a detailed analytical study that lift effects considerably alter the position of the equilibrium point reached by the scaffolds in the (vertical) direction collinear to the gravitational field. This result holds for small particle and shear Reynolds numbers. Our analysis shows that the inertial lift effect is negligible in the horizontal direction. We show that for all rotations of practical interest, and for the range of particle Reynolds number smaller than unity, the vertical coordinate of the equilibrium point is strongly affected by consideration of lift effects. For light (heavy) particles, inclusion of lift in the formation forces the equilibrium position to be below (above) the horizontal plane that contains the axis of rotation. The equilibrium point for light particles is stable and therefore is observable experimentally. The equilibrium point for heavy particles is unstable. We also estimate the stress level applied to the scaffold and derive an algebraic expression that indicates that the stress level acting on the scaffold decreases with increasing shear Reynolds number.