Water movement on the convex surfaces of porous media under microgravity

A plant growth system for crop production under microgravity is part of a life supporting system designed for long-duration space missions. A plant growth in soil in space requires the understanding of water movement in soil void spaces under microgravity. Under 1G-force condition, on earth, water movement in porous media is driven by gradients of matric and gravitational potentials. Under microgravity condition, water movement in porous media is supposed to be driven only by a matric potential gradient, but it is still not well understood. We hypothesized that under microgravity water in void spaces of porous media hardly moved comparing in void spaces without obstacles because the concave surfaces of the porous media hindered water movement. The objective of this study was to investigate water movement on the convex surfaces of porous media under microgravity. We conducted parabolic flight experiments that provided 20–25?s of microgravity at the top of a parabolic flight. We observed water movement in void spaces in soil-like porous media made by glass beads and glass spheres (round-bottomed glass flasks) in the different conditions of water injection under microgravity. Without water injection, water did not move much in neither glass beads nor glass spheres. When water was injected during microgravity, water accumulated in contacts between the particles, and the water made thick fluid films on the particles surface. When the water injection was stopped under microgravity, water was held in the contacts between the particles. This study showed that water did not move upward in the void spaces with or without the water injection. In addition, our results suggested that the difficulty of water movement on the convex (i.e. particle surfaces) might result in slower water move in porous media under microgravity than at 1G-force.     

A capillary-driven root module for plant growth in microgravity.

A new capillary-driven root module design for growing plants in microgravity was developed which requires minimal external control. Unlike existing systems, the water supply to the capillary-driven system is passive and relies on root uptake and media properties to develop driving gradients which operate a suction-induced flow control valve. A collapsible reservoir supplies water to the porous membrane which functions to maintain hydraulic continuity. Sheet and tubular membranes consisting of nylon, polyester and sintered porous stainless steel were tested. While finer pore sized membranes allow greater range of operation, they also reduce liquid flux thereby constraining system efficiency. Membrane selection should consider both the maximum anticipated liquid uptake rate and maximum operating matric head (suction) of the system. Matching growth media water retention characteristics to the porous membrane characteristics is essential for supplying adequate liquid flux and gas exchange. A minimum of 10% air-filled porosity (AFP) was necessary for adequate aeration. The capillary-driven module maintained hydraulic continuity and proper gas exchange rates for more than 80 days in a plant growth experiment.     

Seed-to-seed growth of Arabidopsis thaliana on the International Space Station.

The assembly of the International Space Station (ISS) as a permanent experimental outpost has provided the opportunity for quality plant research in space. To take advantage of this orbital laboratory, engineers and scientists at the Wisconsin Center for Space Automation and Robotics (WCSAR), University of Wisconsin-Madison, developed a plant growth facility capable of supporting plant growth in the microgravity environment. Utilizing this Advanced Astroculture (ADVASC) plant growth facility, an experiment was conducted with the objective to grow Arabidopsis thaliana plants from seed-to-seed on the ISS. Dry Arabidopsis seeds were anchored in the root tray of the ADVASC growth chamber. These seeds were successfully germinated from May 10 until the end of June 2001. Arabidopsis plants grew and completed a full life cycle in microgravity. This experiment demonstrated that ADVASC is capable of providing environment conditions suitable for plant growth and development in microgravity. The normal progression through the life cycle, as well as the postflight morphometric analyses, demonstrate that Arabidopsis thaliana does not require the presence of gravity for growth and development.     

Modeling gas exchange in a closed plant growth chamber.

Fluid transport models for fluxes of water vapor and CO2 have been developed for one crop of wheat and three crops of soybean grown in a closed plant growth chamber. Correspondence among these fluxes is discussed. Maximum fluxes of gases are provided for engineering design requirements of fluid recycling equipment in growth chambers. Furthermore, to investigate the feasibility of generalized crop models, dimensionless representations of water vapor fluxes are presented. The feasibility of such generalized models and the need for additional data are discussed.     

Ground performance of air conditioning and water recycle system for a Space Plant Box.

Researchers from 5 Japanese universities have developed a plant growth facility (Space Plant Box) for seed to seed experiments under microgravity. The breadboard model of the Space Plant Box was fabricated by assembling subsystems developed for microgravity. The subsystems include air conditioning and water recycle system, air circulation system, water and nutrient delivery system, lighting system and plant monitoring system. The air conditioning and water recycle system is simply composed of a single heat exchanger, two fans and hydrophilic fibrous strings. The strings allow water movement from the cooler fin in the Cooling Box to root supporting materials in the Plant Growth Chamber driven by water potential deficit. Relative humidity in the Plant Growth Chamber can be changed over a wide range by controlling the ratio of latent heat exchange to sensible heat exchange on the cooling fin of the heat exchanger. The transpiration rate was successfully measured by circulating air inside the Plant Growth Chamber only. Most water was recycled and a small amount of water needed to be added from the outside. The simple, air conditioning and water recycle system for the Space Plant Box showed good performance through a barley (Hordeum vulgare L.) growth experiment.     

Germination and elongation of flax in microgravity.

This experiment was conducted as part of a risk mitigation payload aboard the Space Shuttle Atlantis on STS-101. The objectives were to test a newly developed water delivery system, and to determine the optimal combination of water volume and substrate for the imbibition and germination of flax (Linum usitatissimum) seeds in space. Two different combinations of germination paper were tested for their ability to absorb, distribute, and retain water in microgravity. A single layer of thick germination paper was compared with one layer of thin germination paper under a layer of thick paper. Paper strips were cut to fit snugly into seed cassettes, and seeds were glued to them with the micropyle ends pointing outward. Water was delivered in small increments that traveled through the paper via capillary action. Three water delivery volumes were tested, with the largest (480 microliters) outperforming the 400 microliters and 320 microliters volumes for percent germination (90.6%) and root growth (mean=4.1 mm) during the 34-hour spaceflight experiment. The ground control experiment yielded similar results, but with lower rates of germination (84.4%) and shorter root lengths (mean=2.8 mm). It is not clear if the roots emerged more quickly in microgravity and/or grew faster than the ground controls. The single layer of thick germination paper generally exhibited better overall growth than the two layered option. Significant seed position effects were observed in both the flight and ground control experiments. Overall, the design of the water delivery system, seed cassettes and the germination paper strip concept was validated as an effective method for promoting seed germination and root growth under microgravity conditions.     

The ASTROCULTURE(TM) flight experiment series, validating technologies for growing plants in space.

A flight experiment, ASTROCULTURE(TM)-1 (ASC-1), to evaluate the operational characteristics and hardware performance of a porous tube nutrient delivery system (PTNDS) was flown on STS-50 as part of the U.S. Microgravity Laboratory-1 mission, 25 June to 9 July, 1992. This experiment is the first in a series of planned ASTROCULTURE(TM) flights to validate the performance of subsystems required to grow plants in microgravity environments. Results indicated that the PTNDS was capable of supplying water and nutrients to plants in microgravity and that its performance was similar in microgravity to that in 1g on Earth. The data demonstrated that water transfer rates through a rooting matrix are a function of pore size of the tubes, the degree of negative pressure on the 'supply' fluid, and the pressure differential between the 'supply' and 'recovery' fluid loops. A slightly greater transfer rate was seen in microgravity than in 1g, but differences were likely related to the presence of hydrostatic pressure effects at 1g. Thus, this system can be used to support plant growth in microgravity or in partial gravity as on a lunar or Mars base. Additional subsystems to be evaluated in the ASTROCULTURE(TM) flight series of experiments include lighting, humidity control and condensate recovery, temperature control, nutrient composition control, CO2 and O2 control, and gaseous contaminant control.     

Autonomic straightening of gravitropically curved cress roots in microgravity.

The typical response of plant organs to gravistimulation is differential growth that leads to organ bending. If the gravitropic stimulus is withdrawn, endogenous compensation of the graviresponse and subsequent straightening occur in some plants. For instance, autonomic straightening of Lepidium roots occurs when gravitropically-curved rootsare rotated on a clinostat (Stankovi et al., 1998a). To determine whether endogenous compensation of the graviresponse also occurs in space, microgravity-grown cress roots were laterally centrifuged in-flight and then returned to microgravity using Biorack hardware on a shuttle mission (STS-81). The cress roots were centrifuged at 4 different g-doses (0.1 x g and 1 x g for 15 or 75 min). All four treatments yielded varying degrees of root curvature. Upon removal from the centrifuge, roots in all four treatments underwent subsequent straightening in microgravity. This straightening resulted from a loss of gravitropic curvature in older regions of the root and the coordinated alignment of new growth. These results show that both microgravity and clinostat rotation on Earth are equivalent in stimulus withdrawal with respect to the induction of endogenous compensation of the curvature. Cress roots are the only plant organ shown to undergo compensation of the curvature in both microgravity and on a clinostat. The compensation of graviresponse in space rules out the hypothesis that the endogenous root straightening ("autotropism") represents a commitment to a pre-stimulus orientation with respect to gravity and instead suggests that there is a default tendency towards axiality following a withdrawal of a g-stimulus.     

Cytochemical localization of calcium in soybean root cap cells in microgravity.

The antimonate precipitation technique was used to evaluate the effects of microgravity and ethylene on the cellular and subcellular distribution of free calcium ions in soybean root apices. Soybean (Glycine max L. [Merr.]) dry seeds were launched, activated by hydration, and germinated in the presence of KMnO4 (to remove ethylene) and in its absence onboard the space shuttle Columbia during the STS-87 mission. Primary root apices of 6-day old seedlings were fixed for electron microscopy after landing. Ultrastructural studies indicated that antimonate precipitation appeared as individual electron-dense particles which were more or less round in shape and varied in diameter from 10 nm (minimum size beginning from which the particles were well identified) to 90 nm. It was revealed that analyzed root cap cells varied in both the precipitate particle sizes and the amount particles per unit of the cellular area. In both flight and ground control treatments, antimonate precipitation level increases from apical meristem cells to peripheral (secretory) cells of root apices. In root cap statocytes, subcellular localization of precipitate particles was revealed in the cytoplasm, nucleus and small vacuoles. The quantitative analysis showed a reduction of precipitate density in the cytoplasm and the nucleus, and an increase in precipitate density in the vacuoles from statocytes of both spaceflight treatments in comparison with ground controls.     

Interaction of growth-determining systems with gravity

The experiments have been carried out with lettuce shoots on board the Salyut-7 orbital station, the Kosmos-1667 biological satellite and under ground conditions at 180° plant inversion. By means of the centrifuge Biogravistat-1M the threshold value of gravitational sensitivity of lettuce shoots has been determined on board the Salyut-7 station. It was found to be equal to 2.9 × 10⁻³g for hypocotyls and 1.5 × 10⁻⁴g for roots. The following results have been received in the experiment performed on board the Kosmos-1667 satellite: a) under microgravity the proliferation of the meristem cells and the growth of roots did not differ from the control; b) the growth of hypocotyls in length was significantly enhanced in microgravity; c) under microgravity transverse growth of hypocotyls (increase in cross sectional area) was significantly increased due to enhancement of cortical parenchyma cell growth. At 180° inversion in Earth's gravity root extension growth and rate of cell division in the root apical meristem were decreased. The determination of DNA-fuchsin value in the nuclei of the cell root apexes showed that inversion affected processess of the cell cycle preceeding cytokinesis.     

The effect of gravity on plant germination.

An axis clinostat was constructed to create micro and negative gravity also a rotated flat disk was constructed with different rotation rates to give increased gravity, by centrifugal force up to 48 g. Rice seeds were grown on agar in tubes at the constant air temperature of 20 degrees C under an average light condition of 110 micromol/m2/sec(PPF). Humidity was not controlled but was maintained above 90%. Since the tube containers were not large enough for long cultivation, shoot and root growth were observed every 12 hours until the sixth day from seeding. The lengths of shoots and roots for each individual plant were measured on the last day. The stem lengths were increased by microgravity but the root lengths were not. Under the negative gravity, negative orthogeotropism and under microgravity, diageotropism was observed. No significant effect of increased gravity was observed on shoot and root growth.     

Canopy photosynthesis and transpiration in microgravity: gas exchange measurements aboard Mir.

The SVET Greenhouse on-board the Orbital Station Mir was used to measure canopy photosynthesis and transpiration rates for the first time in space. During the Greenhouse IIB experiment on Mir (June-January 1997), carbon and water vapor fluxes from two wheat (cv. Superdwarf) canopies were measured using the US developed Gas Exchange Measurement System (GEMS). Gas analyzers capable of resolving CO2 concentration differences of 5 micromoles mol-1 against a background of 0.9% CO2, are necessary to measure photosynthetic and respiratory rates on Mir. The ability of the GEMS gas analyzers to measure these CO2 concentration differences was determined during extensive ground calibrations. Similarly, the sensitivity of the analyzers to water vapor was sufficient to accurately measure canopy evapotranspiration. Evapotranspiration, which accounted for over 90% of the water added to the root zone, was estimated using gas exchange and used to estimate substrate moisture content. This paper presents canopy photosynthesis and transpiration data during the peak vegetative phase of development in microgravity.     

Clover development during spaceflight: a model system.

The development of legume root nodules was studied as a model system for the examination of gravitational effects on plant root development. In order to examine whether rhizobial association with clover roots can be achieved in microgravity, experiments were performed aboard the KC-135 parabolic aircraft and aboard the sounding rocket mission Consort 3. Binding of rhizobia to roots and the initial stages of root nodule development successfully occurred in microgravity. Seedling germination experiments were performed in the sliding block device, the Materials Dispersion Apparatus, aboard STS-37. When significant hydration of the seeds was achieved, normal rates of germination and seedling development were observed.     

微重力对拟南芥长势影响的代谢网络流平衡分析

微重力作为典型的空间环境因素,对植物生长发育的影响机制是空间生命科学的研究热点。微重力环境直接或间接影响植物代谢,并引起许多生理适应。 随着系统生物学的发展,代谢网络模型使微重力环境下的植物代谢建模成为可能。采用流平衡分析方法对模式植物拟南芥不同组织的代谢网络进行分析,研究微重力对拟南芥生长发育的影响机制。通过比较空间与地面条件下拟南芥的生物质产量,发现空间条件下拟南芥黄化幼苗、幼苗、芽、根、下胚轴的生物量分别下降了33.00%,51.52%,6.89%,12.53%,11.70%,与空间环境下拟南芥的长势变化趋势一致。代谢通路富集分析发现,微重力使得拟南芥的碳固定等通路下调,而磷酸戊糖途径上调,初步解析了微重力对拟南芥生长发育的影响机制,也验证了流平衡方法用于微重力生物学效应研究中的可行性。      

Embryogenesis and organogenesis of Carausius morosus under spaceflight conditions

The influence of cosmic radiation and/or microgravity on insect development was studied during the 7 day German Spacelab Mission D1. Eggs of Carausius morosus of five stages differing in sensitivity to radiation and in capacity to regeneration were allowed to continue their development in the BIORACK 22°C incubator, either at microgravity conditions or on the 1 g reference centrifuge. Using the Biostack concept - eggs in monolayers were sandwiched between visual track detectors - and the 1 g reference centrifuge, we were able to separate radiation effects from microgravity effects and also from combined effects of these two factors in space. After retrieval, hatching rates, growth kinetics and anomaly frequencies were determined in the different test samples. The early stages of development turned out to be highly sensitive to single hits of cosmic ray particles as well as to the temporary exposure to microgravity during their development. In some cases, the combined action of radiation and microgravity even amplified the effects exerted by the single parameters of space. Hits by single HZE particles caused early effects, such as body anomalies, as well as late effects, such as retarded growth after hatching. Microgravity exposure lead to a reduced hatching rate. A synergistic action of HZE particle hits and microgravity was established in the unexpectedly high frequency of anomal larvae. However, it cannot be excluded, that cosmic background radiation or low LET HZE particles are also causally involved in damage observed in the microgravity samples.     

Possible use of a 3-D clinostat to analyze plant growth processes under microgravity conditions.

A three-dimensional (3-D) clinostat equipped with two rotation axes placed at right angles was constructed, and various growth processes of higher plants grown on this clinostat were compared with ground controls, with plants grown on the conventional horizontal clinostat, and with those under real microgravity in space. On the 3-D clinostat, cress roots developed a normal root cap and the statocytes showed the typical polar organization except a random distribution of statoliths. The structural features of clinostatted statocytes were fundamentally similar to those observed under real microgravity. The graviresponse of cress roots grown on the 3-D clinostat was the same as the control roots. On the 3-D clinostat, shoots and roots exhibited a spontaneous curvature as well as an altered growth direction. Such an automorphogenesis was sometimes exaggerated when plants were subjected to the horizontal rotation, whereas the curvature was suppressed on the vertical rotation. These discrepancies in curvature between the 3-D clinostat and the conventional ones appear to be brought about by the centrifugal force produced. Thus, the 3-D clinostat was proven as a useful device to simulate microgravity.     

连续粒径粉体在浆体中的堆积密度

采用测定相同流动度时浆体需水量的方法,试验证明了不同粒径分布的矿粉其堆积密度有显著的差异.将浆体中的粉体颗粒及其表面包裹着的一层水膜作为复合颗粒,从而将求浆体中粉体颗粒堆积密度的问题转化为求假想复合颗粒体系的堆积密度问题,在Stovall模型的基础上推导了浆体中连续粒径粉体的堆积密度计算公式;通过试验确定了该公式中的待定参数值,并对公式的适用性进行了验证,结果表明:用该公式可以根据粉体的粒径分布计算它在浆体中的堆积密度.      

Induction of hypoxic root metabolism results from physical limitations in O2 bioavailability in microgravity.

Numerous spaceflight experiments have noted changes in the roots that are consistent with hypoxia in the root zone. These observations include general ultrastructure analysis and biochemical measurements to direct measurements of stress specific enzymes. In experiments that have monitored alcohol dehydrogenase (ADH), the data shows this hypoxically responsive gene is induced and is associated with increased ADH activity in microgravity. These changes in ADH could be induced either by spaceflight hypoxia resulting from inhibition of gravity mediated O2 transport, or by a non-specific stress response due to inhibition of gravisensing. We tested these hypotheses in a series of two experiments. The objective of the first experiment was to determine if physical changes in gravity-mediated O2 transport can be directly measured, while the second series of experiments tested whether disruption of gravisensing can induce a non-specific ADH response. To directly measure O2 bioavailability as a function of gravity, we designed a sensor that mimics metabolic oxygen consumption in the rhizosphere. Because of these criteria, the sensor is sensitive to any changes in root O2 bioavailability that may occur in microgravity. In a KC-135 experiment, the sensor was implanted in a moist granular clay media and exposed to microgravity during parabolic flight. The resulting data indicated that root O2 bioavailability decreased in phase with gravity. In experiments that tested for non-specific induction of ADH, we compared the response of transgenic Arabidopsis plants (ADH promoted GUS marker gene) exposed to clinostat, control, and waterlogged conditions. The plants were grown on agar slats in a growth chamber before being exposed to the experimental treatments. The plants were stained for GUS activity localization, and subjected to biochemical tests for ADH, and GUS enzyme activity. These tests showed that the waterlogging treatment induced significant increases in GUS and ADH enzyme activities, while the control and clinostat treatments showed no response. This work demonstrates: (1) the inhibition of gravity-driven convective transport can reduce the O2 bioavailability to the root tip, and (2) the perturbation of gravisensing by clinostat rotation does not induce a nonspecific stress response involving ADH. Together these experiments support the microgravity convection inhibition model for explaining changes in root metabolism during spaceflight.     

Approaches in the determination of plant nutrient uptake and distribution in space flight conditions.

The effective growth and development of vascular plants rely on the adequate availability of water and nutrients. Inefficiency in either the initial absorption, transportation, or distribution of these elements are factors which impinge on plant structure and metabolic integrity. The potential effect of space flight and microgravity conditions on the efficiency of these processes is unclear. Limitations in the available quantity of space-grown plant material and the sensitivity of routine analytical techniques have made an evaluation of these processes impractical. However, the recent introduction of new plant cultivating methodologies supporting the application of radionuclide elements and subsequent autoradiography techniques provides a highly sensitive investigative approach amenable to space flight studies. Experiments involving the use of gel based 'nutrient packs' and the radionuclides calcium-45 and iron-59 were conducted on the Shuttle mission STS-94. Uptake rates of the radionuclides between ground and flight plant material appeared comparable.     

Preliminary evaluation of soil moisture probe for use with Arcillite.

There is a need for reliable methods of measuring the level and distribution of water in the solid substrates that are used for growing plants in space. In a microgravity environment, water distribution is governed generally by capillary forces. Arcillite is the solid substrate used in the ASTROCULTURE (TM) system which was developed for growing plants in space. The goal of this study is to evaluate the applicability of heat pulse moisture sensors for measuring moisture levels in Arcillite. The ASTROCULTURE system uses suction as a means of controlling the moisture level in Arcillite, but the spatial distribution of the moisture is left unknown. Studies of the moisture content in a cell experiment were conducted to calibrate a heat pulse moisture sensor and then the sensor was used in a suction experiment to verify moisture content and distribution. Results of the studies demonstrate that head pulse moisture sensors can be used to monitor moisture content and distribution within the root module of the ASTROCULTURE system.