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Potato growth in a porous tube water and nutrient delivery system.

Potato (Solanum tuberosum L.) cv. 'Norland', vegetative growth and tuber productivity grown in the porous water and nutrient delivery system (PTNDS) developed by the Wisconsin Center for Space Automation and Robotics were compared with the vegetative growth and tuber productivity of plants grown in a peat:vermiculite potting mixture (PT/VR). The plants were grown at 12, 16, and 24-h light periods, 18 degrees C constant temperature, 70% relative humidity, and 300 micromol m-2 s-1 photosynthetic photon flux. Canopy height of plants grown in the PT/VR system was taller than that of plants grown in the PTNDS system. Canopy height differences were greatest when the plants were grown under a 24-h photoperiod. Leaf and stem dry masses were similar for plants grown in the two systems under the 12-h photoperiod. Under the 24-h photoperiod, leaf and stem dry masses of plants grown in the PT/VR system were more than 3 times those of plants grown in the PTNDS system. Tuber dry masses were similar for plants grown in the two systems under the 12-h photoperiod. Under the 24 h-photoperiod, tuber dry weights of plants grown in the PT/VR system were more than twice those of plants grown in the PTNDS system. A slightly higher harvest index (ratio of tuber weight to leaf plus stem weight) was noted for the plants grown in the PTNDS than for the plants grown in the PT/VR system. Plants grown in the PTNDS system at the 24-h photoperiod matured earlier than plants grown at this photoperiod in the PT/VR system. Vegetative growth and tuber productivity of plants grown under the 16-h photoperiod generally were intermediate to those noted for plants grown under the 12 and 24-h photoperiods. These results indicate that potato plants grown in a PTNDS system may require less plant growing volume, mature in a shorter time, and likely produce more tubers per unit area compared with plants grown in the PT/VR system. These plant characteristics are a distinct advantage for a plant growing unit of a CELSS.

Biomass↗

Application of sunlight and lamps for plant irradiation in space bases.

The radiation sources used for plant growth on a space base must meet the biological requirements for photosynthesis and photomorphogenesis. In addition the sources must be energy and volume efficient, while maintaining the required irradiance levels, spectral, spatial and temporal distribution. These requirements are not easily met, but as the biological and mission requirements are better defined, then specific facility designs can begin to accommodate both the biological requirements and the physical limitations of a space based plant growth system.

Ecological Systems, Closed↗

Comparative kinetics and reciprocal inhibition of nitrate and nitrite uptake in roots of uninduced and induced barley (Hordeum vulgare L.) seedlings.

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.

Biological Transport↗

Light-emitting diodes as a radiation source for plants.

Development of a more effective radiation source for use in plant-growing facilities would be of significant benefit for both research and commercial crop production applications. An array of light-emitting diodes (LEDs) that produce red radiation, supplemented with a photosynthetic photon flux (PPF) of 30 micromoles s-1 m-2 in the 400- to 500-nm spectral range from blue fluorescent lamps, was used effectively as a radiation source for growing plants. Growth of lettuce (Lactuca sativa L. Grand Rapids') plants maintained under the LED irradiation system at a total PPF of 325 micromoles s-1 m-2 for 21 days was equivalent to that reported in the literature for plants grown for the same time under cool-white fluorescent and incandescent radiation sources. Characteristics of the plants, such as leaf shape, color, and texture, were not different from those found with plants grown under cool-white fluorescent lamps. Estimations of the electrical energy conversion efficiency of a LED system for plant irradiation suggest that it may be as much as twice that published for fluorescent systems.

Environment, Controlled↗

Responses of soybean to ammonium and nitrate supplied in combination to the whole root system or separately in a split-root system.

To address the questions of whether allocation of carbohydrates to roots is influenced by ionic form of nitrogen absorbed and whether allocation of carbohydrates to roots in turn influences proportionality between NH4+ and NO3- uptake from mixed sources, NH4+ and NO3- were supplied separately to halves of a split-root hydroponic system and were supplied in combination to a whole-root system. Dry matter accumulation in the split-root system was 18% less in the NH4(+)-fed axis than in the NO3(-)-fed axis. This, however, does not indicate that partitioning of carbohydrate between the two axes was different. Most of the reduction in dry matter accumulation in the NH4(+)-fed axis can be accounted for by the retransport of CH2O equivalents from the root back to the shoot with amino acids produced by NH4+ assimilation. Uptake of NH4+ or NO3- by the respective halves of the split-root system was proportional to the estimated allocation of carbohydrate to that half. When NH4+ and NO3- were supplied to separate halves of the split-root system, the cumulative NH4+ to NO3- uptake ratio was 0.81. When supplied in combination to the whole-root system, the cumulative NH4+ to NO3- uptake ratio was 1.67. Thus, while the shoot may affect total nitrogen uptake through the export of carbohydrates to roots, the shoot (common for halves of the split-root system) apparently does not exert a direct effect on proportionality of NH4+ and NO3- uptake by roots. For whole roots supplied with both NH4+ and NO3-, the restriction in uptake of NO3- may involve a stimulation of NO3- efflux rather than an inhibition of NO3- influx. While only the net uptake of NH4+ and NO3- was measured by ion chromatography, monitoring at approximately hourly intervals during the first 3 days of treatment revealed irregularly occurring intervals of both depletion (net influx) and enrichment (net efflux) in solutions. In the case of NH4+, numbers of net efflux events were similar (21 to 24 out of 65 sequential sampling intervals) whether NH4+ was supplied with NO3- to whole-root systems or separately to an axis of the split-root system. In the case of NO3-, however, the number of net efflux events increased from 8 when NO3- was supplied to a separate axis of the split-root system to between 19 and 24 when NO3- was supplied with NH4+ to whole-root systems.

Biological Transport↗

Water relations, gas exchange, and nutrient response to a long term constant water deficit.

Wheat plants (Triticum aestivum) were grown for 43 days in a micro-porous tube nutrient delivery system. Roots were unable to penetrate the microporous tube, but grew on the surface and maintained capillary contact with the nutrient solution on the inside of the tube through the 5-micron pores of the porous tube. Water potential in the system was controlled at -0.4, -0.8, and -3.0 kPa by adjusting the applied pressure (hydrostatic head) to the nutrient solution flowing through the microporous tubes. A relatively small decrease in applied water potential from -0.4 to -3.0 kPa resulted in a 34% reduction of shoot growth but only a moderate reduction in the midday leaf water potential from -1.3 to -1.7 MPa. Carbon dioxide assimilation decreased and water use efficiency increased with the more negative applied water potentials, while intercellular CO2 concentration remained constant. This was associated with a decrease in stomatal conductance to water vapor from 1.90 to 0.98 mol m-2 s-1 and a decrease in total apparent hydraulic conductance from 47 to 12 micromoles s-1 MPa-1. Although the applied water potentials were in the -0.4 to -3.0 kPa range, the actual water potential perceived by the plant roots appeared to be in the range of -0.26 to -0.38 MPa as estimated by the leaf water potential of bagged plants. The amount of K, Ca, Mg, Zn, Cu, and B accumulated with each unit of transpired water increased as the applied water potential became less negative. The increase in accumulation ranged from 1.4-fold for K to 2.2-fold for B. The physiological responses observed in this study in response to small constant differences in applied water potentials were much greater than expected from either the applied water potential or the observed plant water potential. Even though the micro-porous tube may not represent natural conditions and could possibly introduce morphological and physiological artifacts, it enables a high degree of control of water potential that facilitates the investigation of many aspects of water relations not practical with other experimental systems.

Boron↗

A hydroponic system for microgravity plant experiments.

The construction of a permanently manned space station will provide the opportunity to grow plants for weeks or months in orbit for experiments or food production. With this opportunity comes the need for a method to provide plants with a continuous supply of water and nutrients in microgravity. The Capillary Effect Root Environment System (CERES) uses capillary forces to maintain control of circulating plant nutrient solution in the weightless environment of an orbiting spacecraft. The nutrient solution is maintained at a pressure slightly less than the ambient air pressure while it flows on one side of a porous membrane. The root, on the other side of the membrane, is surrounded by a thin film of nutrient solution where it contacts the moist surface of the membrane. The root is provided with water, nutrients and air simultaneously. Air bubbles in the nutrient solution are removed using a hydrophobic/hydrophilic membrane system. A model scaled to the size necessary for flight hardware to test CERES in the space shuttle was constructed.

Capillary Action↗

Dry matter and nitrogen accumulation are not affected by superoptimal concentration of ammonium in flowing solution culture with pH control.

While it is known that superoptimal concentrations of the nitrate (NO3-) ion in solution culture do not increase NO3- uptake or dry matter accumulation, the same is not known for the ammonium (NH4+) ion. An experiment was conducted utilizing flowing solution culture with pH control to investigate the influence of superoptimal NH4+ concentrations on dry matter, nitrogen (N), potassium (K), calcium (Ca), and magnesium (Mg) accumulation by nonnodulated soybean plants. Increasing the NH4+ concentration in solution from 1 to 10 mM did not affect dry matter or N accumulation. Accumulations of K, Ca, and Mg were slightly decreased with increased NH4+ concentration. The NH4+ uptake system, which is saturated at less than 1mM NH4+, is able to regulate uptake of NH4+ at concentrations as high as 10 mM.

Biological Transport↗

Hycrest crested wheatgrass accelerates the degradation of pentachlorophenol in soil.

We investigated the effects of vegetation on the fate of pentachlorophenol (PCP) in soil using a novel high-flow sealed test system. Pentachlorophenol has been widely used as a wood preservative, and this highly toxic biocide contaminates soil and ground water at many sites. Although plants are known to accelerate the rates of degradation of certain soil contaminants, this approach has not been thoroughly investigated for PCP. The fate of [14C]PCP, added to soil at a concentration of 100 mg/kg, was compared in three unplanted and three planted systems. The plant used was Hycrest, a perennial, drought-tolerant cultivar of crested wheatgrass [Agropyron desertorum (Fischer ex Link) Schultes]. The flow-through test system allowed us to maintain a budget for 14C-label as well as monitor mineralization (breakdown to 14CO2) and volatilization of the test compound in a 155-d trial. In the unplanted systems, an average of 88% of the total radiolabel remained in the soil and leachate and only 6% was mineralized. In the planted system, 33% of the radiolabel remained in the soil plus leachate, 22% was mineralized, and 36% was associated with plant tissue (21% with the root fraction and 15% with shoots). Mineralization rates were 23.1 mg PCP mineralized kg-1 soil in 20 wk in the planted system, and for the unplanted system 6.6 mg PCP kg-1 soil for the same time period. Similar amounts of volatile organic material were generated in the two systems (1.5%). Results indicated that establishing crested wheatgrass on PCP-contaminated surface soils may accelerate the removal of the contaminant.

Biodegradation, Environmental↗

Characteristics of injury and recovery of net NO3- transport of barley seedlings from treatments of NaCl.

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-.

Biological Transport↗

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.

Equipment Design↗

Model system for studies of microbial dynamics at exuding surfaces such as the rhizosphere.

An autoclavable all-glass system for studying microbial dynamics at permeable surfaces is described. Standard hydrophobic or hydrophilic membranes (46-mm diameter) of various pore sizes were supported on a glass frit through which nutrient solutions were pumped by a peristaltic pump. The pump provided a precisely controlled flow at speeds of 0.5 to 500 ml of defined or natural cell exudates per h, which passed through the membrane into a receiving vessel. The construction allowed a choice of membranes, which could be modified. The system was tested with a bacterium, isolated from rape plant roots (Brassica napus L.), that was inoculated on a hydrophilic membrane filter and allowed to develop into a biofilm. A defined medium with a composition resembling that of natural rape root exudate was pumped through the membrane at 0.5 ml/h. Scanning electron microscopic examinations indicated that the inoculum formed microcolonies embedded in exopolymers evenly distributed over the membrane surface. The lipid composition and content of poly-beta-hydroxybutyrate in free-living and adhered cells were determined by gas chromatography. The bacterial consumption of amino acids in the exudate was also studied.

Amino Acids↗

A porous stainless steel membrane system for extraterrestrial crop production.

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.

Hydroponics↗

Do the higher oxidation states of the photosynthetic O2-evolving system contain bound H2O?

A modified mass spectrometer was used to determine whether the higher oxidation states of the photosynthetic O2-evolving system contain substrate water that is not freely exchangeable with the external medium. Our data indicated that the higher oxidation states contain no appreciable bound, non-exchangeable H2O. This suggests that H2O oxidation takes place via a rapid, concerted, all-or-none mechanism rather than by a mechanism involving stable, partially oxidized, H2O-derived intermediates. These findings set definite constraints on possible mechanisms of O2 evolution.

Chloroplasts↗

Effect of CO2 levels on nutrient content of lettuce and radish.

Atmospheric carbon-dioxide enrichment is known to affect the yield of lettuce and radish grown in controlled environments, but little is known about CO2 enrichment effects on the chemical composition of lettuce and radish. These crops are useful model systems for a Controlled Ecological Life-Support System (CELSS), largely because of their relatively short production cycles. Lettuce (Lactuca sativa L.) cultivar 'Waldmann's Green' and radish (Raphanus sativus L.) cultivar 'Giant White Globe' were grown both in the field and in controlled environments, where hydroponic nutrient solution, light, and temperature were regulated, and where CO2 levels were controlled at 400, 1000, 5000, or 10,000 ppm. Plants were harvested at maturity, dried, and analyzed for proximate composition (protein, fat, ash, and carbohydrate), total nitrogen (N), nitrate N, free sugars, starch, total dietary fiber, and minerals. Total N, protein N, nonprotein N (NPN), and nitrate N generally increased for radish roots and lettuce leaves when grown under growth chamber conditions compared to field conditions. The nitrate-N level of lettuce leaves, as a percentage of total NPN, decreased with increasing levels of CO2 enrichment. The ash content of radish roots and of radish and lettuce leaves decreased with increasing levels of CO2 enrichment. The levels of certain minerals differed between field- and chamber-grown materials, including changes in the calcium (Ca) and phosphorus (P) contents of radish and lettuce leaves, resulting in reduced Ca/P ratio for chamber-grown materials. The free-sugar contents were similar between the field and chamber-grown lettuce leaves, but total dietary fiber content was much higher in the field-grown plant material. The starch content of growth-chamber lettuce increased with CO2 level.

Atmosphere↗

Optimization of moisture content for wheat seedling germination in a cellulose acetate medium for a space flight experiment.

The Porous Tube Plant Nutrient Delivery System (PTPNDS), a hydrophilic, microporous ceramic tube hydroponic system designed for microgravity, will be tested in a middeck locker of the Space Shuttle. The flight experiment will focus on hardware operation and assess its ability to support seed germination and early seedling growth in microgravity. The water controlling system of the PTPNDS hardware has been successfully tested during the parabolic flight of the KC-135. One challenge to the development of the space flight experiment was to devise a method of holding seeds to the cylindrical porous tube. The seed-holder must provide water and air to the seed, absorb water from the porous tube, withstand sterilization, provide a clear path for shoots and roots to emerge, and be composed of flight qualified materials. In preparation for the flight experiment, a wheat seed-holder has been designed that utilizes a cellulose acetate plug to facilitate imbibition and to hold the wheat seeds in contact with the porous tube in the correct orientation during the vibration of launch and the microgravity environment of orbit. Germination and growth studies with wheat at a range of temperatures showed that optimal moisture was 78% (by weight) in the cellulose acetate seed holders. These and other design considerations are discussed.

Cellulose↗

The Vacuum-Operated Nutrient Delivery System: hydroponics for microgravity.

A nutrient delivery system that may have applicability for growing plants in microgravity is described. The Vacuum-Operated Nutrient Delivery System (VONDS) draws nutrient solution across roots that are under a partial vacuum at approximately 91 kPa. Bean (Phaseolus vulgaris L. cv. Blue Lake 274) plants grown on the VONDS had consistently greater leaf area and higher root, stem, leaf, and pod dry weights than plants grown under nonvacuum control conditions. This study demonstrates the potential applicability of the VONDS for growing plants in microgravity for space biology experimentation and/or crop production.

Equipment Design↗

A dynamic growth model of vegetative soya bean plants: model structure and behaviour under varying root temperature and nitrogen concentration.

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.

Carbohydrate Metabolism↗