Effect of reduced atmospheric pressure on growth and quality of two lettuce cultivars.

Future space missions will likely include plants to provide fresh foods and bioregenerative life support capabilities. Current spacecraft such as the International Space Station (ISS) operate at 1 atm (101 kPa) pressure, but future missions will likely use reduced pressures to minimize gas leakage and facilitate rapid egress (space walks). Plants for these missions must be able to tolerate and grow reliably at these reduced pressures. We grew two lettuce cultivars, 'Flandria' a green bibb-type and 'Outredgeous,' a red, loose-leaf type, under three pressures: 96 kPa (ambient control), 67 kPa (2/3 atm), and 33 kPa (1/3 atm) for 21 days in rockwool using recirculating nutrient film technique hydroponics. Each treatment was repeated three times using a different hypobaric chamber each time. A daily light integral of 17.2 Moles Photosynthetically Active Radiation per day was provided with metal halide lamps set to deliver 300 µmol m-2s -1 photosynthetic photon flux (PPF) for a 16 h photoperiod at 22 °C. Oxygen was maintained at 21 kPa (equal to 21% at 1 atm) and CO2 at 0.12 kPa (equal to 1200 ppm at 1 atm). Leaf area for 'Outredgeous' was reduced 20% and 38% at 67 kPa and 33 kPa respectively; shoot fresh mass was reduced 22% and 41% at 67 kPa and 33 kPa respectively when compared to control plants at 96 kPa. These trends were not statistically significant at P ≥ 0.05. Leaf area for 'Flandria' showed no difference between 96 and 67 kPa but was reduced 31% at 33 kPa; shoot fresh mass was reduced 6% and 27% at 66 kPa and 33 kPa respectively compared to 96 kPa. There were 10% and 25% increases in anthocyanin concentration at 66 kPa and 33 kPa compared to 96 kPa, potentially increasing the bioprotective capacity of the plant. Previous studies with other cultivars of lettuce showed slight change in growth across this range of pressures, suggesting responses may vary among genotypes, hypobaric exposure treatments, and / or environmental conditions. Collectively, the findings suggest further testing is needed to understand the effects of atmospheric pressure on plant growth.

Response of graywater recycling systems based on hydroponic plant growth to three classes of surfactants.

Anionic (sodium laureth sulfate, SLES), amphoteric (cocamidopropyl betaine, CAPB) and nonionic (alcohol polyethoxylate, AE) surfactants were added to separate nutrient film technique (NFT) hydroponic systems containing dwarf wheat (Triticum aestivum cv. USU Apogee) in a series of 21 day trials. Surfactant was added either in a (1). temporally dynamic mode (1-3 g surfactant m(-2) growing area d(-1)) as effected by automatic addition of a 300 ppm surfactant solution to meet plant water demand, or (2). continuous mode (2 g surfactant m(-2) growing area d(-1)) as effected by slow addition (10 mLh(-1)) of a 2000 ppm surfactant solution beginning at 4d after planting. SLES showed rapid primary degradation in both experiments, with no accumulation 24 h after initial addition. CAPB and AE were degraded less rapidly, with 30-50% remaining 24 h after initial addition, but CAPB and AE levels were below detection limit for the remainder of the study. No reductions in vegetative growth of wheat were observed in response to SLES, but biomass was reduced 20-25% with CAPB and AE. Microbial communities associated with both the plant roots and wetted hardware surfaces actively degraded the surfactants, as determined by monitoring surfactant levels following pulse additions at day 20 (with plants) and day 21 (after plant removal). In order to test whether the biofilm communities could ameliorate phytotoxicity by providing a microbial community acclimated for CAPB and AE decay, the continuous exposure systems were planted with wheat seeds after crop removal at day 21. Acclimation resulted in faster primary degradation (>90% within 24h) and reduced phytotoxicity. Overall, the studies indicate that relatively small areas (3-5m(2)) of hydroponic plant systems can process per capita production of mixed surfactants (5-10 g x person(-1)d(-1)) with minimal effects on plant growth.

Improving spinach, radish, and lettuce growth under red light-emitting diodes (LEDs) with blue light supplementation.

Radish (Raphanus sativus L. cv. Cherriette), lettuce (Lactuca sativa L. cv. Waldmann's Green), and spinach (Spinacea oleracea L. cv. Nordic IV) plants were grown under 660-nm red light-emitting diodes (LEDs) and were compared at equal photosynthetic photon flux (PPF) with either plants grown under cool-white fluorescent lamps (CWF) or red LEDs supplemented with 10% (30 micromoles m-2 s-1) blue light (400-500 nm) from blue fluorescent (BF) lamps. At 21 days after planting (DAP), leaf photosynthetic rates and stomatal conductance were greater for plants grown under CWF light than for those grown under red LEDs, with or without supplemental blue light. At harvest (21 DAP), total dry-weight accumulation was significantly lower for all species tested when grown under red LEDs alone than when grown under CWF light or red LEDs + 10% BF light. Moreover, total dry weight for radish and spinach was significantly lower under red LEDs + 10% BF than under CWF light, suggesting that addition of blue light to the red LEDs was still insufficient for achieving maximal growth for these crops.

Theoretical and practical considerations of staggered crop production in a BLSS.

A functional Bioregenerative Life Support System (BLSS) will generate oxygen, remove excess carbon dioxide, purify water, and produce food on a continuous basis for long periods of operation. In order to minimize fluctuations in gas exchange, water purification, and yield that are inherent in batch systems, staggered planting and harvesting of the crop is desirable. A 418-day test of staggered production of potato cv. Norland (26-day harvest cycles) using nutrients recovered from inedible biomass was conducted at Kennedy Space Center. The results indicate that staggered production can be sustained without detrimental effects on BLSS life support functions. System yields of H2O, O2 and food were higher in staggered than batch plantings. Plants growing in staggered production or batch production on "aged" solution initiated tubers earlier, and were shorter than plants grown on "fresh" solution. This morphological response required an increase in planting density to maintain full canopy coverage. Plants grown in staggered production used available light more efficiently than the batch planting due to increased side lighting.

Effects of CO2 on stomatal conductance: do stomata open at very high CO2 concentrations?

Potato and wheat plants were grown for 50 d at 400, 1000 and 10000 micromoles mol-1 carbon dioxide (CO2). and sweetpotato and soybean were grown at 1000 micromoles mol-1 CO2 in controlled environment chambers to study stomatal conductance and plant water use. Lighting was provided with fluorescent lamps as a 12 h photoperiod with 300 micromoles m-2 s-1 PAR. Mid-day stomatal conductances for potato were greatest at 400 and 10000 micromoles mol-1 and least at 1000 micromoles mol-1 CO2. Mid-day conductances for wheat were greatest at 400 micromoles mol-1 and least at 1000 and 10000 micromoles mol-1 CO2. Mid-dark period conductances for potato were significantly greater at 10000 micromoles mol-1 than at 400 or 1000 micromoles mol-1, whereas dark conductance for wheat was similar in all CO2 treatments. Temporarily changing the CO2 concentration from the native 1000 micromoles mol-1 to 400 micromoles mol-1 increased mid-day conductance for all species, while temporarily changing from 1000 to 10000 micromoles mol-1 also increased conductance for potato and sweetpotato. Temporarily changing the dark period CO2 from 1000 to 10000 micromoles mol-1 increased conductance for potato, soybean and sweetpotato. In all cases, the stomatal responses were reversible, i.e. conductances returned to original rates following temporary changes in CO2 concentration. Canopy water use for potato was greatest at 10000, intermediate at 400, and least at 1000 micromoles mol-1 CO2, whereas canopy water use for wheat was greatest at 400 and similar at 1000 and 10000 micromoles mol-1 CO2. Elevated CO2 treatments (i.e. 1000 and 10000 micromoles mol-1) resulted in increased plant biomass for both wheat and potato relative to 400 micromoles mol-1, and no injurious effects were apparent from the 10000 micromoles mol-1 treatment. Results indicate that super-elevated CO2 (i.e. 10000 micromoles mol-1) can increase stomatal conductance in some species, particularly during the dark period, resulting in increased water use and decreased water use efficiency.

Blue light requirements for crop plants used in bioregenerative life support systems.

As part of NASA's Advanced Life Support Program, the Breadboard Project at Kennedy Space Center is investigating the feasibility of using crop plants in bioregenerative life support systems (BLSS) for long-duration space missions. Several types of electric lamps have been tested to provide radiant energy for plants in a BLSS. These lamps vary greatly in terms of spectral quality resulting in differences in growth and morphology of the plants tested. Broad spectrum or "white" light sources (e.g., metal halide and fluorescent lamps) provide an adequate spectrum for normal growth and morphology; however, they are not as electrically efficient as are low-pressure sodium (LPS) or high-pressure sodium (HPS) lamps. Although LPS and HPS, as well as the newly tested red light-emitting diodes (LEDs), have good photosynthetically active radiation (PAR) efficiencies, they are deficient in blue light. Results with several of the crops tested for BLSS (wheat, potato, soybean, lettuce, and radish) have shown a minimum amount of blue light (approximately 30 micromoles m-2 s-1) is necessary for normal growth and development. For example, the lack of sufficient blue light in these lamps has resulted in increased stem elongation and significant reductions in photosynthesis and yield. To avoid problems with blue-deficient lamps and maximize yield, sufficient intensity of HPS or blue light supplementation with red LEDs or LPS lamps is required to meet spectral requirements of crops for BLSS.

Life cycle experiments with Arabidopsis grown under red light-emitting diodes (LEDs).

Light-emitting diodes (LEDS) are a potential lighting source for space-based plant growth systems because of their small mass, operational longevity, and spectral quality. However, the vegetative and reproductive growth and development of plants grown under narrow spectrum LEDs must be characterized before acceptance of LEDS as an alternative light source for growing plants. The objectives of this study were 1) to determine the feasibility of using red LEDS for growing Arabidopsis thaliana L. through a full seed-bearing generation, and 2) to determine if supplemental blue radiation is necessary for growth and seed production. Arabidopsis grown under red LEDS alone produced viable seed, but these plants had abnormal leaf morphology and delayed flowering in comparison to control plants grown under broad spectrum white light or red LEDS supplemented with blue light.

A recirculating hydroponic system for studying peanut (Arachis hypogaea L.).

Peanut (Arachis hypogaea L.) plants were grown hydroponically, using continuously recirculating nutrient solution. Two culture tray designs were tested; one tray design used only nutrient solution, while the other used a sphagnum-filled pod development compartment just beneath the cover and above the nutrient solution. Both trays were fitted with slotted covers to allow developing gynophores to reach the root zone. Peanut seed yields averaged 350 gm-2 dry mass, regardless of tray design, suggesting that substrate is not required for hydroponic peanut production.

Photomorphogenesis, photosynthesis, and seed yield of wheat plants grown under red light-emitting diodes (LEDs) with and without supplemental blue lighting.

Red light-emitting diodes (LEDs) are a potential light source for growing plants in spaceflight systems because of their safety, small mass and volume, wavelength specificity, and longevity. Despite these attractive features, red LEDs must satisfy requirements for plant photosynthesis and photomorphogenesis for successful growth and seed yield. To determine the influence of gallium aluminium arsenide (GaAlAs) red LEDs on wheat photomorphogenesis, photosynthesis, and seed yield, wheat (Triticum aestivum L., cv. 'USU-Super Dwarf') plants were grown under red LEDs and compared to plants grown under daylight fluorescent (white) lamps and red LEDs supplemented with either 1% or 10% blue light from blue fluorescent (BF) lamps. Compared to white light-grown plants, wheat grown under red LEDs alone demonstrated less main culm development during vegetative growth through preanthesis, while showing a longer flag leaf at 40 DAP and greater main culm length at final harvest (70 DAP). As supplemental BF light was increased with red LEDs, shoot dry matter and net leaf photosynthesis rate increased. At final harvest, wheat grown under red LEDs alone displayed fewer subtillers and a lower seed yield compared to plants grown under white light. Wheat grown under red LEDs+10% BF light had comparable shoot dry matter accumulation and seed yield relative to wheat grown under white light. These results indicate that wheat can complete its life cycle under red LEDs alone, but larger plants and greater amounts of seed are produced in the presence of red LEDs supplemented with a quantity of blue light.

Use of biologically reclaimed minerals for continuous hydroponic potato production in a CELSS.

Plant-derived nutrients were successfully recycled in a Controlled Ecological Life Support System (CELSS) using biological methods. The majority of the essential nutrients were recovered by microbiologically treating the plant biomass in an aerobic bioreactor. Liquid effluent containing the nutrients was then returned to the biomass production component via a recirculating hydroponic system. Potato (Solanum tuberosum L.) cv. Norland plants were grown on those nutrients in either a batch production mode (same age plants on a nutrient solution) or a staggered production mode (4 different ages of plants on a nutrient solution). The study continued over a period of 418 days, within NASA Breadboard Project's Biomass Production Chamber at the Kennedy Space Center. During this period, four consecutive batch cycles (104-day harvests) and 13 consecutive staggered cycles (26-day harvests) were completed using reclaimed minerals and compared to plants grown with standard nutrient solutions. All nutrient solutions were continually recirculated during the entire 418 day study. In general, tuber yields with reclaimed minerals were within 10% of control solutions. Contaminants, such as sodium and recalcitrant organics tended to increase over time in solutions containing reclaimed minerals, however tuber composition was comparable to tubers grown in the control solutions.

Theoretical and practical considerations for staggered production of crops in a BLSS.

A functional Bioregenerative Life Support System (BLSS) will generate oxygen, remove excess carbon dioxide, purify water, and produce food on a continuous basis for long periods of operation. In order to minimize fluctuations in gas exchange, water purification, and yield that are inherent in batch systems, staggered planting and harvesting of the crop is desirable. A 418-d test of staggered production of potato cv. Norland (26-d harvest cycles) using nutrients recovered from inedible biomass was recently completed at Kennedy Space Center. The results indicate that staggered production can be sustained without detrimental effects on life support functions in a CELSS. System yields of H2O, O2 and food were higher in staggered than batch plantings. Plants growing in staggered production or batch production on "aged" solution initiated tubers earlier, and were shorter than plants grown on "fresh" solution. This morphological response required an increase in planting density to maintain full canopy coverage. Plants grown in staggered production used available light more efficiently than the batch planting due to increased sidelighting.

Effect of elevated carbon dioxide on nutritional quality of tomato.

Tomato (Lycopersicon esculentum Mill.) cvs. Red Robin (RR) and Reimann Philipp (RP) were grown hydroponically for 105 d with a 12 h photoperiod, 26 degrees C/22 degrees C thermoperiod, and 500 micromol m-2 s-1 PPF at either 400, 1200, 5000, or 10,000 micromol mol-1 (0.04, 0.12, 0.50, 1.00 kPa) CO2. Harvested fruits were analyzed for proximate composition, total dietary fiber, nitrate, and elemental composition. No trends were apparent with regard to CO2 effects on proximate composition, with fruit from all treatments and both cultivars averaging 18.9% protein, 3.6% fat, 10.2% ash, and 67.2% carbohydrate. In comparison, average values for field-grown fruit are 16.6% protein, 3.8% fat, 8.1% ash, and 71.5% carbohydrate (Duke and Atchely, 1986). Total dietary fiber was highest at 10,000 micromol mol-1 (28.4% and 22.6% for RR and RP) and lowest at 1000 micromol mol-1 (18.2% and 15.9% for RR and RP), but showed no overall trend in response to CO2. Nitrate values ranged from 0.19% to 0.35% and showed no trend with regard to CO2. K, Mg, and P concentrations showed no trend in response to CO2, but Ca levels increased from 198 and 956 ppm in RR and RP at 400 micromol mol-1, to 2537 and 2825 ppm at 10,000 micromol mol-1. This increase in Ca caused an increase in fruit Ca/P ratios from 0.07 and 0.37 for RR and RP at 400 micromol mol-1 to 0.99 and 1.23 for RR and RP at 10,000 micromol mol-1, suggesting that more dietary Ca should be available from high CO2-grown fruit.

Interacting effects of photoperiod and photosynthetic photon flux on net carbon assimilation and starch accumulation in potato leaves.

The effect of photoperiod (PP) on net carbon assimilation rate (Anet) and starch accumulation in newly mature canopy leaves of 'Norland' potato (Solanum tuberosum L.) was determined under high (412 varies as mol m-2s-1) and low (263 varies as mol m-2s-1) photosynthetic photon flux (PPF) conditions. The Anet decreased from 13.9 to 11.6 and 9.3 micromoles m-2s-1, and leaf starch increased from 70 to 129 and 118 mg g-1 drymass (DM) as photoperiod (PP) was increased from 12/12 to 18/6, and 24/0, respectively. Longer PP had a greater effect with high PPF conditions than with low PPF treatments, with high PPF showing greater decline in Anet. Photoperiod did not affect either the CO2 compensation point (50 micromoles mol-1) or CO2 saturation point (1100-1200 micromoles mol-1) for Anet. These results show an apparent limit to the amount of starch that can be stored (approximately 15% DM) in potato leaves. An apparent feedback mechanism exists for regulating Anet under high PPF, high CO2, and long PP, but there was no correlation between Anet and starch concentration in individual leaves. This suggests that maximum Anet cannot be sustained with elevated CO2 conditions under long PP (> or = 12 hours) and high PPF conditions. If a physiological limit exists for the fixation and transport of carbon,then increasing photoperiod and light intensity under high CO2 conditions is not the most appropriate means to maximize the yield of potatoes.

NASA's Biomass Production Chamber: a testbed for bioregenerative life support studies.

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.

Growth and gas exchange by lettuce stands in a closed, controlled environment.

Two studies were conducted in which 'Waldmann's Green' lettuce (Lactuca sativa L.) was grown hydroponically from seed to harvest in a large (20-m2), atmospherically closed growth chamber for the National Aeronautics and Space Administration's controlled ecological life support system (CELSS) program. The first study used metal-halide (MH) lamps [280 micromoles m-2 s-1 photosynthetic photon flux (PPF)], whereas the second used high-pressure sodium (HPS) lamps (293 micromoles m-2 s-1). Both studies used a 16-hour photoperiod, a constant air temperature (22 to 23C), and 1000 micromoles mol-1 CO2 during the light period. In each study, canopy photosynthesis and evapotranspiration (ET) rates were highly correlated to canopy cover, with absolute rates peaking at harvest (28 days after planting ) at 17 micromoles CO2/m2 per sec and 4 liters m-2 day-1, respectively. When normalized for actual canopy cover, photosynthesis and ET rates per unit canopy area decreased with age (between 15 and 28 days after planting). Canopy cover increased earlier during the study with HPS lamps, and final shoot yields averaged 183 g fresh mass (FM)/plant 8.8 g dry mass (DM)/plant. Shoot yields in the first study with MH lamps averaged 129 g FM/plant and 6.8 g DM/plant. Analysis of leaf tissue showed that ash levels from both studies averaged 22% and K levels ranged from 15% to 17% of tissue DM. Results suggest that lettuce should be easily adaptable to a CELSS with moderate lighting and that plant spacing or transplant schemes are needed to maximize canopy light interception and sustained efficient CO2 removal and water production.