Productivity and food value of Amaranthus cruentus under non-lethal salt stress.

Stress effects from the accumulation of metal salts may pose a problem for plants in closed biological systems such as spacecraft. This work examined the effects of salinity on growth, photosynthesis and carbon allocation in the crop plant, Amaranthus. Plants were germinated and grown in modified Hoagland's solution with NaCl concentrations of 0 to 1.0%. Plants received salt treatments at various times in development to assess effects on particular life history phases. For Amaranthus cruentus, germination, vegetative growth, flowering, seed development and yield were normal at salinities from 0 to 0.2%. Inhibition of these phases increased from 0.2 to 0.4% salinity and was total above 0.5%. 1.0% salinity was lethal to all developmental phases. Onset of growth phases were not affected by salinity. Plants could not be adapted by gradually increasing salinity over days or weeks. Water uptake increased, while photosynthetic CO2 uptake decreased with increasing salinity on a dry weight basis during vegetative growth. Respiration was not affected by salinity. After flowering, respiration and photosynthesis decreased markedly, such that 1.0% NaCl inhibited photosynthesis completely. Protein levels were unchanged with increasing salinity. Leaf starch levels were lower at salinities of 0.5% and above, while stem starch levels were not affected by these salinities. The evidence supports salt inhibition arising from changes in primary biochemical processes rather than from effects on water relations. While not addressing the toxic effects of specific ions, it suggests that moderate salinity per se need not be a problem in space systems.     

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.     

Operations of a spaceflight experiment to investigate plant tropisms

Plants will be an important component in bioregenerative systems for long-term missions to the Moon and Mars. Since gravity is reduced both on the Moon and Mars, studies that identify the basic mechanisms of plant growth and development in altered gravity are required to ensure successful plant production on these space colonization missions. To address these issues, we have developed a project on the International Space Station (ISS) to study the interaction between gravitropism and phototropism in Arabidopsis thaliana. These experiments were termed TROPI (for tropisms) and were performed on the European Modular Cultivation System (EMCS) in 2006. In this paper, we provide an operational summary of TROPI and preliminary results on studies of tropistic curvature of seedlings grown in space. Seed germination in TROPI was lower compared to previous space experiments, and this was likely due to extended storage in hardware for up to 8 months. Video downlinks provided an important quality check on the automated experimental time line that also was monitored with telemetry. Good quality images of seedlings were obtained, but the use of analog video tapes resulted in delays in image processing and analysis procedures. Seedlings that germinated exhibited robust phototropic curvature. Frozen plant samples were returned on three space shuttle missions, and improvements in cold stowage and handing procedures in the second and third missions resulted in quality RNA extracted from the seedlings that was used in subsequent microarray analyses. While the TROPI experiment had technical and logistical difficulties, most of the procedures worked well due to refinement during the project.     

Oxygen requirement of germinating flax seeds.

Plant experiments in earth orbit are typically prepared on the ground and germinated in orbit to study gravity effects on the developing seedlings. Germination requires the breakdown of storage compounds, and this metabolism depends upon respiration, making oxygen one of the limiting factors in seed germination. In microgravity lack of run-off of excess water requires careful testing of water dispensation and oxygen availability. In preparation for a shuttle experiment (MICRO on STS-107) we studied germination and growth of flax (Linum usitatissimum L.) seedlings in the developed hardware (Magnetic Field Chamber, MFC). We tested between four to 32 seeds per chamber (air volume=14 mL) and after 36 h measured the root length. At 90 microliters O2 per seed (32 seeds/chamber), the germination decreased from 94 to 69%, and the root length was reduced by 20%, compared to 8 seeds per chamber. Based on the percent germination and root length obtained in controlled gas mixtures between 3.6 and 21.6% O2 we determined the lower limit of reliable germination to be 10 vol. % O2 at atmospheric pressure. Although the oxygen available in the MFC's can support the intended number of seeds, the data show that seed storage and microgravity-related limitations may reduce germination.     

Conceptual design of a bioregenerative life support system containing crops and silkworms

This article summarizes a conceptual design of a bioregenerative life support system for permanent lunar base or planetary exploration. The system consists of seven compartments – higher plants cultivation, animal rearing, human habitation, water recovery, waste treatment, atmosphere management, and storages. Fifteen kinds of crops, such as wheat, rice, soybean, lettuce, and mulberry, were selected as main life support contributors to provide the crew with air, water, and vegetable food. Silkworms fed by crop leaves were designated to produce partial animal nutrition for the crew. Various physical-chemical and biological methods were combined to reclaim wastewater and solid waste. Condensate collected from atmosphere was recycled into potable water through granular activated carbon adsorption, iodine sterilization, and trace element supplementation. All grey water was also purified though multifiltration and ultraviolet sterilization. Plant residue, human excrement, silkworm feces, etc. were decomposed into inorganic substances which were finally absorbed by higher plants. Some meat, ingredients, as well as nitrogen fertilizer were prestored and resupplied periodically. Meanwhile, the same amount and chemical composition of organic waste was dumped to maintain the steady state of the system. A nutritional balanced diet was developed by means of the linear programming method. It could provide 2721 kcal of energy, 375.5 g of carbohydrate, 99.47 g of protein, and 91.19 g of fat per capita per day. Silkworm powder covered 12.54% of total animal protein intakes. The balance of material flows between compartments was described by the system of stoichiometric equations. Basic life support requirements for crews including oxygen, food, potable and hygiene water summed up to 29.68 kg per capita per day. The coefficient of system material closure reached 99.40%.     

A conceptual configuration of the lunar base bioregenerative life support system including soil-like substrate for growing plants

The paper presents a conceptual configuration of the lunar base bioregenerative life support system (LBLSS), including soil-like substrate (SLS) for growing plants. SLS makes it possible to combine the processes of plant growth and the utilization of plant waste. Plants are to be grown on SLS on the basis of 20 kg of dry SLS mass or 100 kg of wet SLS mass per square meter. The substrate is to be delivered to the base ready-made as part of the plant growth subsystem. Food for the crew was provided by prestored stock 24% and by plant growing system 76%. Total dry weight of the food is 631 g per day (2800 kcal/day) for one crew member (CM). The list of candidate plants to be grown under lunar BLSS conditions included 14 species: wheat, rice, soybean, peanuts, sweet pepper, carrots, tomatoes, coriander, cole, lettuce, radish, squash, onion and garlic. From the prestored stock the crew consumed canned fish, iodinated salt, sugar, beef sauce and seafood sauce. Our calculations show that to provide one CM with plant food requires the area of 47.5 m². The balance of substance is achieved by the removal dehydrated urine 59 g, feces 31 g, food waste 50 g, SLS 134 g, and also waters 86 g from system and introduction food 236 g, liquid potassium soap 4 g and mineral salts 120 g into system daily. To reduce system setup time the first plants could be sowed and germinated to a certain age on the Earth.     

Effect of bacterial population density on germination wheat seeds and dynamics of simple artificial ecosystems.

Effect of the size of rhizospheric bacterial populations on germination of seeds and development of simple terrestrial "wheat plants--rhizospheric microorganisms--artificial soil" and "wheat plants-artificial soil" systems has been studied. Experiments demonstrated that within specify ranges in the inoculate, the rhizospheric bacteria are capable of increasing the yield of germinated seeds and stimulate the growth of plantlets. Germination of seeds inoculated with bacteria was either stimulated, or inhibited or remained at control levels depending on the amount of bacteria. Plant biomass growth and total photoassimilation has been found to depend on the amount of bacteria on the plant roots: the higher the amount of bacteria on plant roots, the smaller is the biomass of plants but the total photoassimilation is, higher. Thus, depending on the amount of bacteria on the roots of plants the system either increases the biomass of plants or increases the total photoassimilation, i.e. "pumps" carbon through itself involving bacteria. Grant numbers: N99-04-96017, N15.     

Current concepts and future directions of CELSS.

Studies of bioregenerative life support systems for use in space indicate that they are scientifically feasible. Preliminary data suggest that they would provide cost- and weight-saving benefits for low Earth orbit, long duration space platforms. Concepts of such systems include the use of higher plants and/or micro-algae as sources of food, potable water and oxygen, and as sinks for carbon dioxide and metabolic wastes. Recycling of materials within the system will require processing of food organism and crew wastes using microbiological and/or physical chemical techniques. The dynamics of material flow within the system will require monitoring, control, stabilization and maintenance imposed by computers. Future phases of study will continue investigations of higher plant and algal physiology, environmental responses, and control; flight experiments for testing responses of organisms to weightlessness and increased radiation levels; and development of ground-based facilities for the study of recycling within a bioregenerative life support system.     

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.     

Viability of barley seeds after long-term exposure to outer side of international space station

Barley seeds were exposed to outer space for 13 months in a vented metal container without a climate control system to assess the risk of physiological and genetic mutation during long-term storage in space. The space-stored seeds (S0 generation), with an 82% germination rate in 50 seeds, lost about 20% of their weight after the exposure. The germinated seeds showed normal growth, heading, and ripening. The harvested seeds (S1 generation) also germinated and reproduced (S2 generation) as did the ground-stored seeds. The culm length, ear length, number of seed, grain weight, and fertility of the plants from the space-stored seeds were not significantly different from those of the ground-stored seeds in each of the S0 and S1 generation. Furthermore, the S1 and S2 space-stored seeds respectively showed similar β-glucan content to those of the ground-stored seeds. Amplified fragment length polymorphism analysis with 16 primer combinations showed no specific fragment that appears or disappears significantly in the DNA isolated from the barley grown from the space-stored seeds. Though these data are derived from nine S0 space-stored seeds in a single exposure experiment, the results demonstrate the preservation of barley seeds in outer space for 13 months without phenotypic or genotypic changes and with healthy and vigorous growth in space.     

Aquatic food production modules in bioregenerative life support systems based on higher plants.

Most bioregenerative life support systems (BLSS) are based on gravitropic higher plants which exhibit growth and seed generation disturbances in microgravity. Even when used for a lunar or martian base the reduced gravity may induce a decreased productivity in comparison to Earth. Therefore, the implementation of aquatic biomass production modules in higher plant and/or hybrid BLSS may compensate for this and offer, in addition, the possibility to produce animal protein for human nutrition. It was shown on the SLS-89 and SLS-90 space shuttle missions with the C.E.B.A.S.-MINI MODULE that the edible non gravitropic rootless higher aquatic plant Ceratophyllum demeresum exhibits an undisturbed high biomass production rate in space and that the teleost fish species, Xiphophorus helleri, adapts rapidly to space conditions without loss of its normal reproductive functions. Based on these findings a series of ground-based aquatic food production systems were developed which are disposed for utilization in space. These are plant production bioreactors for the species mentioned above and another suitable candidate, the lemnacean (duckweed) species, Wolffia arrhiza. Moreover, combined intensive aquaculture systems with a closed food loop between herbivorous fishes and aquatic and land plants are being developed which may be suitable for integration into a BLSS of higher complexity. Grant numbers: WS50WB9319-3, IVA1216-00588.     

Growth of sweetpotato cultured in the newly designed hydroponic system for space farming

Life support of crews in long-duration space missions for other planets will be highly dependent on amounts of food, atmospheric O₂ and clean water produced by plants. Therefore, the space farming system with scheduling of crop production, obtaining high yields with a rapid turnover rate, converting atmospheric CO₂ to O₂ and purifying water should be established with employing suitable plant species and cultivars and precisely controlling environmental variables around plants grown at a high density in a limited space. In this study, we developed a new hydroponic method for producing tuberous roots and fresh edible leaves and stems of sweetpotato. In the first experiment, we examined the effects of water contents in the rooting substrate on growth and tuberous root development of sweetpotato. The rooting substrates made with rockwool slabs were inclined in a culture container and absorbed nutrient solution from the lower end of the slabs by capillary action. Tuberous roots developed on the lower surface of the rockwool slabs. The tuberous roots showed better growth and development at locations farther from the water surface on the rockwool slabs, which had lower water content. In the second experiment, three sweetpotato cultivars were cultured in a hydroponic system for five months from June to November under the sun light in Osaka, Japan as a fundamental study for establishing the space farming system. The cultivars employed were ‘Elegant summer’, ‘Kokei-14’ and ‘Beniazuma’. The hydroponic system mainly consisted of culture containers and rockwool slabs. Dry weights of tuberous roots developed in the aerial space between the rockwool slab and the nutrient solution filled at the bottom of the culture container were 0.34, 0.45 and 0.23 kg/plant and dry weights of the top portion (leaves, petioles and stems) were 0.42, 0.29 and 0.61 kg/plant for ‘Elegant summer’, ‘Kokei-14’ and ‘Beniazuma’, respectively. Young stems and leaves as well as tuberous roots of ‘Elegant summer’ are edible and palatable. Therefore ‘Elegant summer’ would be a promising crop to produce large amounts of food with high nutritional values in the present hydroponic system in space farming.     

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.     

Instrumentation for plant health and growth in space.

The present-day plant growth facilities ("greenhouses") for space should be equipped with monitors and controllers of ambient parameters within the chamber because spacecraft environmental variations can be unfavorable to plants. Moreover, little is known about the effects of spaceflight on the greenhouse and rooting media. Lack of information about spaceflight effects on plants necessitates supplying space greenhouses with automatic, non-invasive monitors of, e.g., gas exchange rate, water and nutrient ion uptake, plant mass, temperature and water content of leaves. However, introduction of an environmental or plant sensor into the monitoring system may be reasonable only if it is justified by quantitative evaluation of the influence of a measured parameter on productivity, efficacy of illumination, or some other index of greenhouse efficiency. The multivariate adaptive optimization in terrestrial phytotrons appears to be one of the best methods to assess environmental impacts on crops. Two modifications of greenhouses with the three-dimensional adaptive optimization of crop photosynthetic characteristics include: (1) irradiation, air temperature and carbon dioxide using a modified simplex algorithm; and (2) using irradiation, air temperature, and humidity with sensitivity algorithms with varying frequency of test exposures that have been experimentally developed. As a result, during some stages of plant ontogensis, the photosynthetic productivity of wheat, tomatoes, and Chinese cabbage in these systems was found to increase by a factor of 2-3.     

First flight of the ASTROCULTURE (TM) experiment as a part of the U.S. Shuttle/MIR program.

A number of space-based experiments have been conducted to assess the impact of microgravity on plant growth and development. In general, these experiments did not identify any profound impact of microgravity on plant growth and development, though investigations to study seed development have indicated difficulty in plants completing their reproductive cycle. However, it was not clear whether the lack of seed production was due to gravity effects or some other environmental condition prevailing in the unit used for conducting the experiment. The ASTROCULTURE (TM) flight unit contains a totally enclosed plant chamber in which all the critically important environmental conditions are controlled. Normal wheat (Triticum aestivum L.) growth and development in the ASTROCULTURE (TM) flight unit was observed during a ground experiment conducted prior to the space experiment. Subsequent to the ground experiment, the flight unit was transported to MIR by STS-89, as part of the U.S. Shuttle/MIR program, in an attempt to determine if super dwarf wheat plants that were germinated in microgravity would grow normally and produce seeds. The experiment was initiated on-orbit after the flight unit was transferred from the Space Shuttle to MIR. The ASTROCULTURE (TM) flight unit performed nominally for the first 24 hours after the flight unit was activated, and then the unit stopped functioning abruptly. Since it was not possible to return the unit to nominal operation it was decided to terminate the experiment. On return of the flight unit, it was confirmed that the control computer of the ASTROCULTURE (TM) flight unit sustained a radiation hit that affected the control software embedded in the computer. This experience points out that at high orbital inclinations, such as that of MIR and that projected for the International Space Station, the danger of encountering harmful radiation effects are likely unless the electronic components of the flight hardware are resistant to such impacts.     

Auxin polar transport in Arabidopsis under simulated microgravity conditions — Relevance to growth and development

Activity of auxin polar transport in inflorescence axes of Arabidopsis thaliana grown under simulated microgravity conditions was studied in relation to the growth and development. Seeds were germinated and allowed to grow on an agar medium in test tubes on a horizontal clinostat. Horizontal clinostat rotation substantially reduced the growth of inflorescence axes and the productivity of seeds of Arabidopsis thaliana (ecotypes Landsberg erecta and Columbia), although it little affected seed germination, development of rosette leaves and flowering. The activity of auxin polar transport in inflorescence axes decreased when Arabidopsis plants were grown on a horizontal clinostat from germination stage, being ca. 60% of 1 g control. On the other hand, the auxin polar transport in inflorescence axes of Arabidopsis grown in 1 g conditions was not affected when the segments were exposed to various gravistimuli, including 3-dimensional clinorotation, during transport experiments. Pin-formed mutant of Arabidopsis, having a unique structure of the inflorescence axis with no flower and extremely low levels of the activity of auxin polar transport in inflorescence axes and endogenous auxin, did not continue its vegetative growth under clinostat rotation. These facts suggest that the development of the system of auxin polar transport in Arabidopsis is affected by microgravity, resulting in the inhibition of growth and development, especially during reproductive growth.     

Aquatic modules for bioregenerative life support systems: developmental aspects based on the space flight results of the C.E.B.A.S. MIN-MODULE.

The Closed Equilibrated Biological Aquatic System (C.E.B.A.S.) is an artificial aquatic ecosystem which contains teleost fishes, water snails, ammonia oxidizing bacteria and edible non-gravitropic water plants. It serves as a model for aquatic food production modules which are not seriously affected by microgravity and other space conditions. Its space flight version, the so-called C.E.B.A.S. MINI-MODULE was already successfully tested in the STS-89 and STS-90 (NEUROLAB) missions. It will be flown a third time in space with the STS-107 mission in January 2003. All results obtained so far in space indicate that the basic concept of the system is more than suitable to drive forward its development. The C.E.B.A.S. MINI-MODULE is located within a middeck locker with limited space for additional components. These technical limitations allow only some modifications which lead to a maximum experiment time span of 120 days which is not long enough for scientifically essential multi-generation-experiments. The first necessary step is the development of "harvesting devices" for the different organisms. In the limited space of the plant bioreactor a high biomass production leads to self-shadowing effects which results in an uncontrolled degradation and increased oxygen consumption by microorganisms which will endanger the fishes and snails. It was shown already that the latter reproduce excellently in space and that the reproductive functions of the fish species are not affected. Although the parent-offspring-cannibalism of the ovoviviparous fish species (Xiphophorus helleri) serves as a regulating factor in population dynamics an uncontrolled snail reproduction will also induce an increased oxygen consumption per se and a high ammonia concentration in the water. If harvesting locks can be handled by astronauts in, e. g., 4-week intervals their construction is not very difficult and basic technical solutions are already developed. The second problem is the feeding of the animals. Although C.E.B.A.S.-based aquaculture modules are designed to be closed food loop systems (edible herbivorous fish species and edible water plants) which are already verified on Earth this will not be possible in space without devices in which the animals are fed from a food storage. This has to be done at least once daily which would waste too much crew time when done by astronauts. So, the development of a reliable automated food dispenser has highest priority. Also in this case basic technical solutions are already elaborated. The paper gives a comprehensive overview of the proposed further C.E.B.A.S.-based development of longer-term duration aquatic food production modules.     

Integration of crop production with CELSS waste management.

Lettuce plants were grown utilizing water, inorganic elements, and CO2 inputs recovered from waste streams. The impact of these waste-derived inputs on the growth of lettuce was quantified and compared with results obtained when reagent grade inputs were used. Phytotoxicity was evident in both the untreated wastewater stream and the recovered CO2 stream. The toxicity of surfactants in wastewater was removed using several treatment systems. Harmful effects of gaseous products resulting from incineration of inedible biomass on crop growth were observed. No phytotoxicity was observed when inorganic elements recovered from incinerated biomass ash were used to prepare the hydroponic solution, but the balance of nutrients had to be modified to achieve near optimal growth. The results were used to evaluate closure potential of water and inorganic elemental loops for integrated plant growth and human requirements.     

Bioelectrochemical degradation of urea at platinized boron doped diamond electrodes for bioregenerative systems

The recovery of potable water from space mission wastewater is critical for the life support and environmental health of crew members in long-term missions. NASA estimates reveal that at manned space missions 1.91 kg/person day of urine is produced, with urea and various salts as its main components. In this research we explore the utilization of urease (EC 3.5.1.5, 15,000 U/g) along with a platinized boron doped diamond electrode (Pt-BDD) to degrade urea. Urea is directly degraded to nitrogen by the in situ utilization of the reaction products as a strategy to increase the amount of clean water in future space expeditions. The biochemical reaction of urease produces ammonia and carbon dioxide from urea. Thereafter, ammonia is electrooxidized at the interface of the Pt-BDD producing molecular nitrogen. The herein presented system has been proven to have 20% urea conversion efficiency. This research has potential applications for future long-term space missions since the reaction byproducts could be used for a biomass subsystem (in situ resource recovery), while generating electricity from the same process.     

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.