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Research at the Utah State University College of Agriculture that is relevant to the space life sciences is reviewed. Specific programs detailed are gravitropism of dicot stems, maximization of wheat yields for use in space exploration, and plant development processes in wheat in microgravity.
If humans are to live permanently in space, regenerative life support systems are an enabling technology and must replace the picnic approach of taking all supplies required for each mission. These systems are classified by technologies as either physical/chemical or bioregenerative. Both of these system-types can recycle water, remove carbon dioxide, produce oxygen, and recover essential elements from waste products. Bioregenerative can also produce food, thus, making it essential if humans are to exist in space independent of earth. A solely bioregenerative life support system includes plants as a biomass production module and microbial organisms in bioreactors as a resource recovery module. In the Advanced Life Support Program, bioregenerative life support systems are being investigated through a research and technology development project which includes large scale testing as part of the Breadboard Project and human tests conducted in the soon to be constructed BioPlex facility. Research and technology development efforts are directed toward optimizing biomass productivity in controlled chambers by developing light weight, energy efficient, and automated systems; recycling liquid and solid wastes; baselining the operation of bioreactors; determining system microbial stability; assessing chemical contamination; and building models required for long term system operations. The program will include space flight studies in the near future to determine if these life support technologies will function in microgravity. When a bioregenerative system is finally incorporated into a mission, the conversion from a picnic and resupply mentality to permanent recycling and independence from earth will be complete.
For long term missions, composters may be an integral part of a life support system that provides edible food crops, extracts nutrients from plant biomass and removes contaminants from the recycling stream.
The effect of root-zone temperature on young tomato plants (Lycopersicon esculentum Mill. cv. Heinz 1350) was evaluated in controlled environments using a recirculating solution culture system. Growth rates were measured at root-zone temperatures of 15 degrees, 20 degrees, 25 degrees, and 30 degrees C in a near optimum foliar environment. Optimum growth occurred at 25 degrees to 30 degrees during the first 4 weeks of growth and 20 degrees to 25 degrees during the 5th and 6th weeks. Growth was severely restricted at 15 degrees. Four concentrations of gibberellic acid (GA3) and kinetin were added to the nutrient solution in a separate trial; root-zone temperature was maintained at 15 degrees and 25 degrees. Addition of 15 micromoles GA3 to solutions increased specific leaf area, total leaf area, and dry weight production of plants in both temperature treatments. GA3-induced growth stimulation was greater at 15 degrees than at 25 degrees. GA3 may promote growth by increasing leaf area, enhancing photosynthesis per unit leaf area, or both. Kinetic was not useful in promoting growth at either temperature.
A 50% increase in total radiation by extending the photoperiod from 16 to 24 hr doubled the weight of all cultivars of loose-leaf lettuce (Lactuca sativa L.) 'Grand Rapids Forcing', 'Waldmanns Green', 'Salad Bowl', and 'RubyConn', but not a Butterhead cultivar, 'Salina'. When total daily radiation (moles of photons) was the same, plants under continuous radiation weighed 30% to 50% more than plants under a 16 hr photoperiod. By using continuous radiation on loose-leaf lettuce, fewer lamp fixtures were required and yield was increased.
The effect of Ca2+ on NO3- assimilation in young barley (Hordeum vulgare L. var CM 72) seedlings in the presence and absence of NaCl was studied. Calcium increased the activity of the NO3- transporter under saline conditions, but had little effect under nonsaline conditions. Calcium decreased the induction period for the NO3- transporter under both saline and nonsaline conditions but had little effect on its apparent Km for NO3- both in the presence and absence of NaCl. The enhancement of NO3- transport by Ca2+ under saline conditions was dependent on the presence of Ca2+ in the uptake solution along with the salt, since Ca2+ had no effect when supplied before or after salinity stress. Although Mn2+ and Mg2+ enhanced NO3- uptake under saline conditions, neither was as effective as Ca2+. In longer studies, increasing the Ca2+ concentration in saline nutrient solutions resulted in increases in NO3- assimilation and seedling growth.
To investigate the effects of environment on plant volatile emissions, 'Waldmann's Green' leaf lettuce was cultivated under different levels of photosynthetic photon flux (PPF), photoperiod, and temperature. A modified growth chamber was used to sample plant volatile emissions nondestructively, over time, and under controlled conditions. Total volatile emission rates were significantly higher from lettuce cultivated under PPF of 360 or 200 micromoles m-2 s-1 compared to 105 micromoles m-2 s-1, and significantly higher under a 16-h photoperiod than an 8-h photoperiod. No differences were detected among emission rates from different temperature treatments. In controlled environments, emissions could be regulated by adjusting environmental conditions accordingly.
Earth benefits of research from the NSCORT in Bioregenerative Life Support will include the following: development of active control mechanisms for light, CO2, and temperature to maximize photosynthesis of crop plants during important phases of crop development; automation of crop culture systems; creation of novel culture systems for optimum productivity; creation of value-added crops with superior nutritional, yield, and waste-process characteristics; environmental control of food and toxicant composition of crops; new process technologies and novel food products for safe, nutritious, palatable vegetarian diets; creation of menus for healthful vegetarian diets with psychological acceptability; enzymatic procedures to degrade recalcitrant crop residues occurring in municipal waste; control-system strategies to ensure sustainabilty of a CELSS that will enable management of diverse complex systems on Earth.
The Biomass Production Chamber (BPC) located at Kennedy Space Center, FL, USA provides a large (20 m2 area, 113 m3 vol.), closed environment for crop growth tests for NASA's Controlled Ecological Life Support System (CELSS) program. Since the summer of 1988, the chamber has operated on a near-continuous basis (over 1200 days) without any major failures (excluding temporary power losses). During this time, five crops of wheat (64-86 days each), three crops of soybean (90 to 97 days), five crops of lettuce (28-30 days), and four crops of potato (90 to 105 days were grown, producing 481 kg of dry plant biomass, 196 kg edible biomass, 540 kg of oxygen, 94,700 kg of condensed water, and fixing 739 kg of carbon dioxide. Results indicate that total biomass yields were close to expected values for the given light input, but edible biomass yields and harvest indices were slightly lower than expected. Stand photosynthesis, respiration, transpiration, and nutrient uptake rates were monitored throughout growth and development of the different crops, along with the build-up of ethylene and other volatile organic compounds in the atmosphere. Data were also gathered on system hardware maintenance and repair, as well as person-hours required for chamber operation. Future tests will include long-term crop production studies, tests in which nutrients from waste treatment systems will be used to grow new crops, and multi-species tests.
Radiation from high-pressure sodium (HPS) lamps provided more than a 50% increased yield (fresh and dry weight of tops) of loose-leaf lettuce cultivars Grand Rapids Forcing and RubyConn, compared to that obtained by radiation from cool-white fluorescent (CWF) lamps at equal photosynthetic photon flux; yet, input wattage was approximately 36% less. It was postulated that the considerable output of 700 to 850 nm radiation from the HPS lamp was a significant factor of the increased yield. Under HPS lamps, the leaves of both cultivars were slightly less green with very little red pigmentation ('RubyConn') and slightly elongated, compared to CWF, but plant productivity per unit electrical energy input was vastly superior with HPS.
To determine if the daily pattern of NO3- and NH4+ uptake is affected by acidity or NO3- : NH4+ ratio of the nutrient solution, non-nodulated soybean plants (Glycine max) were exposed for 21 days to replenished, complete nutrient solutions at pH 6.0, 5.5, 5.0, and 4.5 which contained either 1.0 mM NH4+, 1.0 mM NO3- [correction of NO3+], 0.67 mM NH4+ plus 0.33 mM NO3- (2:1 NH4+ : NO3-) [correction of (2:1 NH3+ : NO4-)], or 0.33 mM NH4+ plus 0.67 mM NO3- (1:2 NH4+ : NO3-). Net uptake rates of NH4+ and NO3- were measured daily by ion chromatography as depletion from the replenished solutions. When NH4+ and NO3- were supplied together, cumulative uptake of total nitrogen was not affected by pH or solution NH4+ : NO3- ratio. The cumulative proportion of nitrogen absorbed as NH4+ decreased with increasing acidity; however, the proportional uptake of NH4+ and NO3- was not constant, but varied day-to-day. This day-to-day variation in relative proportions of NH4+ and NO3- absorbed when NH4+ : NO3- ratio and pH of solution were constant indicates that the regulatory mechanism is not directly competitive. Regardless of the effect of pH on cumulative uptake of NH4+, the specific nitrogen uptake rates from mixed and from individual NH4+ and NO3- sources oscillated between maxima and minima at each pH with average periodicities similar to the expected interval of leaf emergence.
A data set is given describing the daily nutrient uptake, gas exchange, environmental conditions, and carbon (C), and nutrient partitioning at harvest for the entire canopy and root system of a wheat crop (Triticum aestivum, cv. Yecora Rojo). The data were obtained from a 20 m2 stand of wheat plants grown from planting to maturity in a closed, controlled environment, and include daily nutrient uptake [macronutrients, nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), magnesium (Mg), and sulfur (S); and micronutrients, iron (Fe), boron (B), manganese (Mn), zinc (Zn), copper (Cu), and molybdenum (Mo)], canopy carbon dioxide (CO2) exchange rates, and transpiration. Environmental factors such as relative humidity, air temperature, nutrient solution temperature, pH and electrical conductivity, and photoperiod were controlled in the chamber to specific set points. A detailed description of biomass yield for each of the 64 plant growth trays comprising the 20 m2 of growth area is also provided, and includes dry weights of grain, straw, chaff, and roots, along with the concentration of nutrients in different plant tissues and the percent carbohydrate, fat, and protein. To our knowledge, this information represents one of the most extensive data sets available for a canopy of wheat grown from seed to maturity under controlled environmental and nutritional conditions, and thus may provide useful information for model development and validation. A methods section is included to qualify any assumptions that might be required for the use of the data in plant growth models, along with a daily event calendar indicating when adjustments in set points and occasional equipment or sensor failures occurred.
Nitrate and NO2- transport by roots of 8-day-old uninduced and induced intact barley (Hordeum vulgare L. var CM 72) seedlings were compared to kinetic patterns, reciprocal inhibition of the transport systems, and the effect of the inhibitor, p-hydroxymercuribenzoate. Net uptake of NO3- and NO2- was measured by following the depletion of the ions from the uptake solutions. The roots of uninduced seedlings possessed a low concentration, saturable, low Km, possibly a constitutive uptake system, and a linear system for both NO3- and NO2-. The low Km system followed Michaelis-Menten kinetics and approached saturation between 40 and 100 micromolar, whereas the linear system was detected between 100 and 500 micromolar. In roots of induced seedlings, rates for both NO3- and NO2- uptake followed Michaelis-Menten kinetics and approached saturation at about 200 micromolar. In induced roots, two kinetically identifiable transport systems were resolved for each anion. At the lower substrate concentrations, less than 10 micromolar, the apparent low Kms of NO3- and NO2- uptake were 7 and 9 micromolar, respectively, and were similar to those of the low Km system in uninduced roots. At substrate concentrations between 10 and 200 micromolar, the apparent high Km values of NO3- uptake ranged from 34 to 36 micromolar and of NO2- uptake ranged from 41 to 49 micromolar. A linear system was also found in induced seedlings at concentrations above 500 micromolar. Double reciprocal plots indicated that NO3- and NO2- inhibited the uptake of each other competitively in both uninduced and induced seedlings; however, Ki values showed that NO3- was a more effective inhibitor than NO2-. Nitrate and NO2- transport by both the low and high Km systems were greatly inhibited by p-hydroxymercuribenzoate, whereas the linear system was only slightly inhibited.
The nature of the injury and recovery of nitrate uptake (net uptake) from NaCl stress in young barley (Hordeum vulgare L, var CM 72) seedlings was investigated. Nitrate uptake was inhibited rapidly by NaCl, within 1 minute after exposure to 200 millimolar NaCl. The duration of exposure to saline conditions determined the time of recovery of NO3- uptake from NaCl stress. Recovery was dependent on the presence of NO3- and was inhibited by cycloheximide, 6-methylpurine, and cerulenin, respective inhibitors of protein, RNA, and sterol/fatty acid synthesis. These inhibitors also prevented the induction of the NO3- uptake system in uninduced seedlings. Uninduced seedlings exhibited endogenous NO3- transport activity that appeared to be constitutive. This constitutive activity was also inhibited by NaCl. Recovery of constitutive NO3- uptake did not require the presence of NO3-.
To assess the cost and area/volume requirements of a farm in a space station or Lunar or Martian base, a few laboratories in the United States, the Soviet Union, France, and Japan are studying optimum controlled environments for the production of selected crops. Temperature, light, photoperiod, CO2, humidity, the root-zone environment, and cultivars are the primary factors being manipulated to increase yields and harvest index. Our best wheat yields on a time basis (24 g m-2 day-1 of edible biomass) are five times good field yields and twice the world record. Similar yields have been obtained in other laboratories with potatoes and lettuce; soybeans are also promising. These figures suggest that approximately 30 m2 under continuous production could support an astronaut with sufficient protein and about 2800 kcal day-1. Scientists under Iosif Gitelzon in Krasnoyarsk, Siberia, have lived in a closed system for up to 5 months, producing 80% of their own food. Thirty square meters for crops were allotted to each of the two men taking part in the experiment. A functional controlled-environment life-support system (CELSS) will require the refined application of several disciplines: controlled-environment agriculture, food preparation, waste disposal, and control-systems technology, to list only the broadest categories. It has seemed intuitively evident that ways could be found to prepare food, regenerate plant nutrients from wastes, and even control and integrate several subsystems of a CELSS. But could sufficient food be produced in the limited areas and with the limited energy that might be available? Clearly, detailed studies of food production were necessary.
A differential equation model of vegetative growth of the soya bean plant (Glycine max (L.) Merrill cv. Ransom') was developed to account for plant growth in a phytotron system under variation of root temperature and nitrogen concentration in nutrient solution. The model was tested by comparing model outputs with data from four different experiments. Model predictions agreed fairly well with measured plant performance over a wide range of root temperatures and over a range of nitrogen concentrations in nutrient solution between 0.5 and 10.0 mmol NO3- in the phytotron environment. Sensitivity analyses revealed that the model was most sensitive to changes in parameters relating to carbohydrate concentration in the plant and nitrogen uptake rate.
A system was developed in which nutrient flow to plant roots is controlled by a thin (0.98 or 1.18 mm) porous (0.2 or 0.5 microns) stainless steel sheet membrane. The flow of nutrient solution through the membrane is controlled by adjusting the relative negative pressure on the nutrient solution side of the membrane. Thus, the nutrient solution is contained by the membrane and cannot escape from the compartment even under microgravity conditions if the appropriate pressure gradient across the membrane is maintained. Plant roots grow directly on the top surface of the membrane and pull the nutrient solution through this membrane interface. The volume of nutrient solution required by this system for plant growth is relatively small, since the plenum, which contains the nutrient solution in contact with the membrane, needs only to be of sufficient size to provide for uniform flow to all parts of the membrane. Solution not passing through the membrane to the root zone is recirculated through a reservoir where pH and nutrient levels are controlled. The size of the solution reservoir depends on the sophistication of the replenishment system. The roots on the surface of the membrane are covered with a polyethylene film (white on top, black on bottom) to maintain a high relative humidity and also limit light to prevent algal growth. Seeds are sown directly on the stainless steel membrane under the holes in the polyethylene film that allow a pathway for the shoots.
Plants of Norland potatoes (Solanum tuberosum L.) were maintained for 42 days at Mg concentrations of 0.05, 0.125, 0.25, 1, 2, and 4 mM in a nonrecirculating nutrient film system under controlled environment. With the increased Mg supply from 0.05 to 4 mM, Mg concentrations in the leaves of the 42-day old plants increased significantly from 1.1 to 11.2 mg g-1 dry weight. Plant leaf area and plant and tuber dry weights increased with increased Mg concentrations up to 1 mM in solution or 6.7 mg g-1 in leaves, and then decreased with further increases in Mg concentrations. Rates of CO2 assimilation measured on leaflets in situ at ambient and various intercellular CO2 concentrations were consistently lower at 0.05 and 4 mM Mg than at other Mg treatments, which may indicate decreased photosynthetic activity in mesophyll tissues at the lowest and highest Mg concentrations. Dark respiration rates in leaves were highest at 0.05 and 4 mM Mg, lowest at 0.25 and 1 mM Mg, and intermediate at 0.125 and 2 mM Mg. The different Mg treatments also influenced accumulation of other minerals in leaves. Leaf concentrations of Ca and Mn decreased with increased Mg supply except that Ca and Mn were lower at 0.05 mM than at 0.125 mM Mg. Leaf K concentrations were lower at 1, 2 and 4 mM Mg than at other Mg treatments. Foliar concentrations of P, Fe, Zn, and Cu had small but inconsistent variation with different Mg concentrations. Leaf concentrations of N, S, and B were similar at different Mg concentrations. This study demonstrates that various Mg nutrition, along with altered accumulation of other nutrients, could regulate dry matter production in potatoes by affecting not only leaf area but also leaf carbon dioxide assimilation and respiration.