The importance of restoration of the atmospheric electrical environment in closed Bioregenerative Life Supporting Systems.

Earth based Bioregenerative Life Supporting Systems (BLSS) are subject to 4 main physical factors: gravity, light, temperature and electrical environment. The first 3 are obvious to everyone, the Electrical Environment (EE) is not under the majority of prevailing conditions perceived directly by our senses. The EE is one of the important physical factors directly influencing some plants and in a less obvious way also the majority of plants. There are only two long range forces in nature: the electromagnetic and the gravitational forces. Gravity is very much weaker than the electromagnetic forces FG/FEL=10(-38), where FG is the gravitational Force and FEL are the electromagnetic Forces. The atmospheric electric field prevails all the time over the entire Earth with a mean intensity of 130 V/m. It is therefore a potent factor which may be used by some plants exposed throughout their entire life time to the atmospheric electric field. What effect should the normal atmospheric electric field have on plants? All living plants are good electrical conductors for electrostatic fields. The plants distort the normally vertical field lines, which have to be perpendicular to the plant tissue everywhere in order to avoid the extraction of energy from the field. The meristems concentrate the field lines, thus the electrically charged nutrients are supplied to the growing parts of the plant exposed to the field. This results in electrotropism in some plants. It is very well known that plants do have adaptive capabilities as compared to animals, it is important for their survival, because they cannot run away from trouble. It is found by careful observations of the behaviour of different plants that some plants do respond to the presence of the atmospheric electric field while other plants exposed to the same environment are indifferent to the atmospheric electric field. The plants growing in the Biosphere II were shielded by the metal structure of the Biosphere II. Because these plants which do make use of the atmospheric electric field are also to be found in the Biosphere II and must be deprived of this natural atmospheric electric field and in consequence of some of their natural nutrients. I have experimental evidence that at least some plants do respond to the atmospheric electric field. This effect is the most likely cause of the oxygen depletion and of the carbon dioxide accumulation in Biosphere II. Under the micro gravity encountered in space habitats the restoration of electric fields is even more important for plant growth than it is on the Earth.     

Evidence for electrotropism in some plant species.

The ever-present global Atmospheric Electrical Field (AEF) is used by many plant species. There are many natural habitats with electrotropic plants and habitats with no AEF. The plants growing there are not electrotropic, like the plants growing under the canopies of the trees or the Arecibo radio telescope. Examples are given of different plants which belong to one or the other class, and the criteria how to distinguish them. In addition to natural habitat observations, laboratory experiments were run in search of the sensitivity of electrotropic effect to different electric field intensities. During a few years, it was established that in very strong fields (of the order of 1 MV/m) all plants respond immediately to the field. This type of reaction is due to the Coulomb forces, but electrotropism depends on electric field interaction with ions. The "reference field" (130 V/m) was always used with stronger fields in the several kV/m range which enhance plant growth rate and size similar to plant growth hormones. Surprising effects were also observed with reversed and horizontal field polarity. In conclusion electrotropic plants deprived of the electrical field do not develop as expected, as can be seen in Biosphere 2. This is an instructive example of what happens when we forget to provide the plants with this vital natural environmental factor. Electrical fields of different intensity, directions and configurations are cheap and easy to generate.     

Performance of a centrifugal phytotron.

It is possible to cultivate plants under an artificial gravity field generated by a centrifugal device in space. In order to determine an optimal magnitude of gravity, there is a need to investigate the relationship between plant and growth and gravity, including not only reduced gravity but also gravity greater than 1G. A prototype centrifugal phytotron was designed and fabricated in order to investigate the relationship between plant growth and increased gravity. This device enables us to cultivate plants over the long term by controlling environmental conditions in the phytotron such as temperature, relative humidity, CO2 concentration and light intensity. The results of our experiment indicate that plant seeds can germinate and grow even under an artificial gravity which changes sinusoidally from 2G to 4G.     

The use of an electric field in increasing the resistance of plants to the action of unfavorable space flight factors.

The key role in increasing the resistance of plants to unfavorable space flight factors is assigned to biomembranes of root cells. It is these biomembranes in which numerous biochemical and biophysical processes determining the adaptive capacity of plant organisms occur. In the initial period of exposure to unfavorable space flight factors the adaptation reactions of the plant organism undoubtedly increase its resistance. But the intensification of removal of H+ ions through the plasmalemma with an increase of the external influence sharply raises the quantity of cations leaving the cell, which leads to the accumulation of a considerable quantity of intracellular negative charges. These charges together with negative charges built in the membrane force protons to concentrate on the external surface of the membrane. Since protons have a very strong electric field, they form such a charge of which the electric field is about from several to hundreds of V/cm. The concentration of positive charges of protons entails the formation of a double electric field which extremely impedes the diffusion of other ions. Thus, a proton barrier is formed. Its length can be very considerable due to which the whole process of transmembrane energy and mass-transfer is disturbed. The proton barrier is easily destroyed by a weak electric field created in the root zone. In experiments on electrostimulation of different plants under space flight conditions at the orbital station MIR the absorption of nutrient elements by the root system increased to the optimal level, the ratio of physiologically active substances in the rhizosphere was normalized, the content of chlorophyll, carotin, and ascorbic acid in leaves corresponded to the ground-based control. Understanding of the mechanism of formation of a proton barrier on the plasmalemma of root cells as a result of the response of plants to the negative action of external factors (microgravity) is of great importance. It allows the possibility of life support of the vegetable kingdom in extreme conditions to be estimated in a new way.     

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.     

The influence of gravity on the structure and functions of plant material

Growth process generate plant form and relate to most physiological functions. The Earth's gravity force affects plant growth in both obvious and subtle ways. It is a major environmental influence on morphology and physiology of plants. Gravity is less important as an agent for plant stress than as an environmental signal to guide growth. The plant's bioaccelerometers are remarkably sensitive, especially in hypogravity. Simulation (clinostat) studies and experiments in satellite laboratories are needed to understand the sensing, transduction, and response characteristics of g related mechanisms. By examining how plants alter growth processes to accomplish developmental or physiological “objectives” we may find it pragmatically desirable to ask ourselves how we might design a plant to achieve such responses to environmental influences. Examples of this design engineering approach for gravity related effects are described as an aid to experimentation.     

Gravity as an obligatory factor in normal higher plant growth and development

Gravity may influence different aspects of plant activity. The present report deals with two questions: gravity as an ecological factor determining spatial orientation of plant growth; and second, a possible requirement for gravity in the process of normal growth, morphogenesis and generative development of plants.     

Progress in plant research in space.

Progress is reviewed of spaceflight research conducted with plants between 1987 and 1992. Numerous plant experiments have been performed on spacecraft and sounding rockets in the past five years by scientists of the US, the former Soviet Union, Europe, and other areas. The experiments are categorized into three areas: gravity sensing, transduction, and response; development and reproduction; and metabolism, photosynthesis, and transport. The results of these experiments continue to demonstrate that gravity and/or other factors of spaceflight affect plants at the organismal, cellular, subcellular, and molecular levels, resulting in changes in orientation, development, metabolism, and growth. The challenge now is to truly dissect the effects of gravity from those of other spaceflight factors and to identify the basic mechanisms underlying gravity's effects.     

Polarity of root statocytes—Relevance for graviperception

During outgrowth of the radicle of cress ( L.) the statocytes of the root cap develop a structural polarity with the nucleus at the proximal cell pole and a complex of endoplasmic reticulum (ER) at the distal cell pole. Amyloplasts sediment upon this complex of ER. During all stages of development of the cytoskeleton (microtubules, microfilaments) is involved in positioning of the ER. The structural polarity of the statocytes develops independently of gravity, as indicated by corresponding results from fast and slow rotating clinostats and roots grown under microgravity in orbit. Disturbance of the structural polarity is possible by application of drugs, influencing microtubules and microfilaments. If, by rotation of roots on slow rotating clinostats or centrifugation, the structural polarity of the statocytes is changed, the ability of the roots to perceive gravity is affected also.     

Evaluation of experiments involving the study of plant orientation and growth under different gravitational conditions.

The manifestation of gravitropic reaction in plants has been considered from the phylogenetic point of view. A chart has been suggested according to which it is supposed that the first indications of the ability to identify the direction of the gravitational vector were inherent in the most ancient eukaryotes, which gave rise to green, brown, yellow-green, golden and diatomaceous algae as well as fungi. The experiments on the role of gravity in plant ontogenesis are being continued. The sum total of the data obtained in a number of experiments in space shows that under these conditions a structurally modified but normally functioning gravireceptive apparatus is formed. The data confirming the modification, under changed gravity, of the processes of integral and cellullar growth of the axial organs of seedlings as well as of the anatomo-morphological structure and developmental rates of plants during their prolonged growth in space are presented. It is assumed that this fact testifies to the presence of systems interacting with gravity during plant ontogenesis. At the same time the necessity for further experiments in order to differentiate an immediate biological effect of gravity from the ones conditioned by it indirectly due to the changes in the behavior of liquids and gases is pointed out. The methodological aspects of biological experiments in space as the main source of reliable information on the biological role of gravity are discussed.     

Modeling gravity effects on water retention and gas transport characteristics in plant growth substrates

Growing plants to facilitate life in outer space, for example on the International Space Station (ISS) or at planned deep-space human outposts on the Moon or Mars, has received much attention with regard to NASA’s advanced life support system research. With the objective of in situ resource utilization to conserve energy and to limit transport costs, native materials mined on Moon or Mars are of primary interest for plant growth media in a future outpost, while terrestrial porous substrates with optimal growth media characteristics will be useful for onboard plant growth during space missions. Due to limited experimental opportunities and prohibitive costs, liquid and gas behavior in porous substrates under reduced gravity conditions has been less studied and hence remains poorly understood. Based on ground-based measurements, this study examined water retention, oxygen diffusivity and air permeability characteristics of six plant growth substrates for potential applications in space, including two terrestrial analogs for lunar and Martian soils and four particulate substrates widely used in reduced gravity experiments. To simulate reduced gravity water characteristics, the predictions for ground-based measurements (1 − g) were scaled to two reduced gravity conditions, Martian gravity (0.38 − g) and lunar gravity (0.16 − g), following the observations in previous reduced gravity studies. We described the observed gas diffusivity with a recently developed model combined with a new approach that estimates the gas percolation threshold based on the pore size distribution. The model successfully captured measured data for all investigated media and demonstrated the implications of the poorly-understood shift in gas percolation threshold with improved gas percolation in reduced gravity. Finally, using a substrate-structure parameter related to the gaseous phase, we adequately described the air permeability under reduced gravity conditions.     

The mechanism by which an asymmetric distribution of plant growth hormone is attained.

Zea mays (sweet corn) seedlings attain an asymmetric distribution of the growth hormone indole-3-acetic acid (IAA) within 3 minutes following a gravity stimulus. Both free and esterified IAA (that is total IAA) accumulate to a greater extent in the lower half of the mesocotyl cortex of a horizontally placed seedling than in the upper half. Thus, changes in the ratio of free IAA to ester IAA cannot account for the asymmetric distribution. Our studies demonstrate there is no de novo synthesis of IAA in young seedlings. We conclude that asymmetric IAA distribution is attained by a gravity-induced, potential-regulated gating of the movement of IAA from kernel to shoot and from stele to cortex. As a working theory, which we call the Potential Gating Theory, we propose that perturbation of the plant's bioelectric field, induced by gravity, causes opening and closing of transport channels in the plasmodesmata connecting the vascular stele to the surrounding cortical tissues. This results in asymmetric growth hormone distribution which results in the asymmetric growth characteristics of the gravitropic response.     

Plant mineral nutrition,gas exchange and photosynthesis in space: A review

Successful growth and development of higher plants in space rely on adequate availability and uptake of water and nutrients, and efficient energy distribution through photosynthesis and gas exchange. In the present review, literature has been reviewed to assemble the relevant knowledge within space plant research for future planetary missions. Focus has been on fractional gravity, space radiation, magnetic fields and ultimately a combined effect of these factors on gas exchange, photosynthesis and transport of water and solutes.     

Particulated growth media for optimal liquid and gaseous fluxes to plant roots in microgravity.

An important and yet relatively under researched area of plant growth in microgravity, deals with the rooting environment of plants. A comprehensive approach for selecting the physical characteristics of root growth media which optimizes the dynamic availability of water and dissolved nutrients, and gases to plant roots was developed and tested. Physically-based and parametric models describing the relationship between content and fluxes of liquids and gases were used to cast a multi-objective optimization problem. This methodology reveals that a medium's ability to supply liquid and gas fluxes optimally is dependent upon physiological target values, system operation limits and root module design which dictate the medium's range of soil water characteristic and particle size distribution. Optimized media parameters designate a particle size distribution from which a particulated growth media was constructed and matched to the optimized media parameters. This methodology should improve the selection of optimal media properties for plant growth in microgravity as well as other porous media applications.     

带电粒子云的演变过程

根据带电粒子云从破碎点开始向空间扩散过程中粒子云密度和形状的变化规律,以几何形状和起主要作用的因素为特征,定义了球形、椭球形、绳形、螺旋线形、全方位弥漫直至球壳形6个演变阶段.论述了在各个阶段的主要特征和对演变过程起主要作用的因素.分析了在各个阶段电磁场对带电粒子的摄动影响,比较了带电粒子云与不带电粒子云在演化过程上的差异.在球形阶段起主要作用的是分离速度,带电碎片之间的排斥力加快了碎片扩散的速度.从椭球形阶段到球壳形阶段,带电粒子和不带电粒子的演化规律基本一致.带电粒子的演化过程加快或减慢取决于粒子带正电或带负电.将电场摄动力等效于改变地球引力的大小,罗列了阶段转换标志点时刻的计算公式.利用计算机仿真的方法,给出了各个阶段不带电碎片云和带电碎片云分布示意图,验证了演变过程阶段划分和电磁场摄动分析的合理性.      

Possible applications of aquatic bioregenerative life support modules for food production in a Martian base.

Water is the essential precondition of life in general and also for the establishment of a Martian base suitable for long duration stays of humans. It is not yet proven if there is indeed a "frozen ocean" under the surface of Mars but if this could be verified it would open innovative aspects for the construction of bioregenerative life support systems (BLSS). In a general concept higher plants will play the predominant role in a Martian BLSS. It is not clear, however, how these will grow and bring seed in reduced gravity and there may be differences in the productivity in comparison to Earth conditions. Therefore, organisms which are already adapted to low gravity conditions, namely non-gravitropic aquatic plants and also aquatic animals may be used to enhance the functionality of the Martian BLSS as a whole. It has been shown already with the so-called C.E.B.A.S. MINIMODULE in the STS-89 and STS-90 spaceshuttle missions that the water plant Ceratophyllum demersum has an undisturbed and high biomass production under space conditions. Moreover, the teleost fish species Xiphophorus helleri adapted easily to the micro-g environment and maintained its normal reproductive functions. Based on this findings a possible scenario is presented in which aquatic plant production modules and combined animal-plant production systems may be used for human food production and water and air regeneration in a Martian base.     

Effects of a hypogeomagnetic field on gravitropism and germination in soybean

Any plants grown during long-term space missions will inevitably experience an extremely low magnetic field (i.e. a hypogeomagnetic field, HGMF). It is possible that the innate adaptation of plants to the earth’s magnetic field (i.e. the geomagnetic field, GMF) would be disrupted. Effects of the HGMF on plant physiological and metabolic processes are unclear. In this study we established a hypogeomagnetic incubation system on the ground and investigated the effects of the HGMF on the gravitropism and germination of soybean seeds. The gravitropism angle, germination percentage, germination speed, water absorbance ratio, seed weight, radicle length, radicle weight, and radicle weight ratio of soybean seeds grown in the local field and the HGMF were compared. In general, the gravitropism angle in the HGMF was smaller than that in the local field when seeds were positioned before emergence in such a way that the direction of the radicle was opposite to that of gravity. The germination percentage, germination speed, and radicle weight ratio increased in the HGMF compared to the control. Our results indicate that the germination and gravitropism of soybean seeds are affected by elimination of the geomagnetic field.     

Growth of pea epicotyl in low magnetic field implication for space research

A magnetic field is an inescapable environmental factor for plants on the earth. However, its impact on plant growth is not well understood. In order to survey how magnetic fields affect plant, Alaska pea seedlings were incubated under low magnetic field (LMF) and also in the normal geo-magnetic environment. Two-day-old etiolated seedlings were incubated in a magnetic shield box and in a control box. Sedimentation of amyloplasts was examined in the epicotyls of seedlings grown under these two conditions. The elongation of epicotyls was promoted by LMF. Elongation was most prominent in the middle part of the epicotyls. Cell elongation and increased osmotic pressure of cell sap were found in the epidermal cells exposed to LMF. When the gravitational environment was 1G, the epicotyls incubated under both LMF and normal geomagnetic field grew straight upward and amyloplasts sedimented similarly. However, under simulated microgravity (clinostat), epicotyl and cell elongation was promoted. Furthermore, the epicotyls bent and amyloplasts were dispersed in the cells in simulated microgravity. The dispersion of amyloplasts may relate to the posture control in epicotyl growth under simulated microgravity generated by 3D clinorotation, since it was not observed under LMF in 1G. Since enhanced elongation of cells was commonly seen both at LMF and in simulated microgravity, all elongation on the 3D-clinostat could result from pseudo-low magnetic field, as a by-product of clinorotation. (i.e., clinostat results could be based on randomization of magnetic field together with randomization of gravity vector.) Our results point to the possible use of space for studies in magnetic biology. With space experiments, the effects of dominant environmental factors, such as gravity on plants, could be neutralized or controlled for to reveal magnetic effects more clearly.     

Spaceflight hardware for conducting plant growth experiments in space: the early years 1960-2000.

The best strategy for supporting long-duration space missions is believed to be bioregenerative life support systems (BLSS). An integral part of a BLSS is a chamber supporting the growth of higher plants that would provide food, water, and atmosphere regeneration for the human crew. Such a chamber will have to be a complete plant growth system, capable of providing lighting, water, and nutrients to plants in microgravity. Other capabilities include temperature, humidity, and atmospheric gas composition controls. Many spaceflight experiments to date have utilized incomplete growth systems (typically having a hydration system but lacking lighting) to study tropic and metabolic changes in germinating seedlings and young plants. American, European, and Russian scientists have also developed a number of small complete plant growth systems for use in spaceflight research. Currently we are entering a new era of experimentation and hardware development as a result of long-term spaceflight opportunities available on the International Space Station. This is already impacting development of plant growth hardware. To take full advantage of these new opportunities and construct innovative systems, we must understand the results of past spaceflight experiments and the basic capabilities of the diverse plant growth systems that were used to conduct these experiments. The objective of this paper is to describe the most influential pieces of plant growth hardware that have been used for the purpose of conducting scientific experiments during the first 40 years of research.     

IMF polarity effects on the equatorial ionospheric F-region

An exploratory study is made of the influence, during the equinoxes, of the interplanetary magnetic field (IMF) sector structure on the ionospheric F-region using ionosonde data from several equatorial stations for a 3-yr period around the 19th sunspot cycle maximum. It is found that, compared with days having positive IMF polarity, the post-sunset increase of h'F near the dip equator and the depth of the equatorial ionization anomaly (EIA) are reduced during the vernal equinox and enhanced during the antumnal equinox on days with negative IMF polarity. Similar trends are also noted in the data for the 20th sunspot cycle maximum, but with reduced amplitude. The systematic changes in the F-region characteristics suggest a modification of the equatorial zonal electric fields in association with the IMF polarity-related changes in the semi-annual variation of geomagnetic activity.