Predicting the effects of gas diffusivity on photosynthesis and transpiration of plants grown under hypobaria

As part of a Bio-regenerative Life Support System (BLSS) for long-term space missions, plants will likely be grown at reduced pressure. This low pressure will minimize structural requirements for growth chambers on missions to the Moon or Mars. However, at reduced pressures the diffusivity of gases increases. This will affect the rates at which CO₂ is assimilated and water is transpired through stomata. To understand quantitatively the possible effects of reduced pressure on plant growth, CO₂ and H₂O transport were calculated for atmospheres of various total pressures (101, 66, 33, 22, 11 kPa) and CO₂ concentrations (0.04, 0.1 and 0.18 kPa). The diffusivity of a gas is inversely proportional to total pressure and shows dramatic increases at pressures below 33 kPa (1/3 atm). A mathematical relationship based on the principle of thermodynamics was applied to low pressure conditions and can be used for calculating the transpiration and photosynthesis of plants grown in hypobaria. At 33 kPa total pressure, the stomatal conductance increases by a factor of three with the boundary layer conductance increasing by a factor of ∼1.7, since the leaf conductance is a function of both stomatal and the boundary layer conductance, the overall conductance will increase resulting in significantly higher levels of transpiration as the pressure drops. The conductance of gases is also regulated by stomatal aperture in an inverse relationship. The higher CO₂ concentration inside the leaf air space during low pressure treatments may result in higher CO₂ assimilation and partial stomata closure, resulting in a decrease in transpiration rate. The results of this analysis offer guidelines for experiments in pressure and high CO₂ environments to establish ideal conditions for minimizing transpiration and maximizing the plant biomass yield in BLSS.     

Short-term and long-term effects of low total pressure on gas exchange rates of spinach.

In this study, spinach plants were grown under atmospheric and low pressure conditions with constant O2 and CO2 partial pressures, and the effects of low total pressure on gas exchange rates were investigated. CO2 assimilation and transpiration rates of spinach grown under atmospheric pressure increased after short-term exposure to low total pressure due to the enhancement of leaf conductance. However, gas exchange rates of plants grown at 25 kPa total pressure were not greater than those grown at atmospheric pressure. Stomatal pore length and width were significantly smaller in leaves grown at low total pressure. This result suggested that gas exchange rates of plants grown under low total pressure were not stimulated even with the enhancement of gas diffusion because the stomatal size and stomatal aperture decreased.     

Carbon dioxide exchange of lettuce plants under hypobaric conditions.

Growth of plants in a Controlled Ecological Life Support System (CELSS) may involve the use of hypobaric pressures enabling lower mass requirements for atmospheres and possible enhancement of crop productivity. A controlled environment plant growth chamber with hypobaric capability designed and built at Ames Research Center was used to determine if reduced pressures influence the rates of photosynthesis (Ps) and dark respiration (DR) of hydroponically grown lettuce plants. The chamber, referred to as a plant volatiles chamber (PVC), has a growing area of about 0.2 m2, a total gas volume of about 0.7 m3, and a leak rate at 50 kPa of <0.1%/day. When the pressure in the chamber was reduced from ambient to 51 kPa, the rate of net Ps increased by 25% and the rate of DR decreased by 40%. The rate of Ps increased linearly with decreasing pressure. There was a greater effect of reduced pressure at 41 Pa CO2 than at 81 Pa CO2. This is consistent with reports showing greater inhibition of photorespiration (Pr) in reduced O2 at low CO2 concentrations. When the partial pressure of O2 was held constant but the total pressure was varied between 51 and 101 kPa, the rate of CO2 uptake was nearly constant, suggesting that low pressure enhancement of Ps may be mainly attributable to lowered partial pressure of O2 and the accompanying reduction in Pr. The effects of lowered partial pressure of O2 on Ps and DR could result in substantial increases in the rates of biomass production, enabling rapid throughput of crops or allowing flexibility in the use of mass and energy resources for a CELSS.     

Carbon dioxide exchange of lettuce plants under hypobaric conditions.

Growth of plants in a Controlled Ecological Life Support System (CELSS) may involve the use of hypobaric pressures enabling lower mass requirements for atmospheres and possible enhancement of crop productivity. A controlled environment plant growth chamber with hypobaric capability designed and built at Ames Research Center was used to determine if reduced pressures influence the rates of photosynthesis (Ps) and dark respiration (DR) of hydroponically grown lettuce plants. The chamber, referred to as a plant volatiles chamber (PVC), has a growing area of about 0.2 m2, a total gas volume of about 0.7 m3, and a leak rate at 50 kPa of <0.1%/day. When the pressure in the chamber was reduced from ambient to 51 kPa, the rate of net Ps increased by 25% and the rate of DR decreased by 40%. The rate of Ps increased linearly with decreasing pressure. There was a greater effect of reduced pressure at 41 Pa CO2 than at 81 Pa CO2. This is consistent with reports showing greater inhibition of photorespiration (Pr) in reduced O2 at low CO2 concentrations. When the partial pressure of O2 was held constant but the total pressure was varied between 51 and 101 kPa, the rate of CO2 uptake was nearly constant, suggesting that low pressure enhancement of Ps may be mainly attributable to lowered partial pressure of O2 and the accompanying reduction in Pr. The effects of lowered partial pressure of O2 on Ps and DR could result in substantial increases in the rates of biomass production, enabling rapid throughput of crops or allowing flexibility in the use of mass and energy resources for a CELSS.     

Proximate nutritional composition of CELSS crops grown at different CO2 partial pressures.

Two CELSS candidate crops, soybean (Glycine max) and potato (Solanum tuberosum), were grown hydroponically in controlled environments maintained at carbon dioxide (CO2) partial pressures ranging from 0.05 to 1.00 kPa (500 to 10,000 ppm at 101 kPa atmospheric pressure). Plants were harvested at maturity (90 days for soybean and 105 days for potato) and all tissues analyzed for proximate nutritional composition (i.e. protein, fat, carbohydrate, crude fiber, and ash content). Soybean seed ash and crude fiber were higher and carbohydrate was lower than values reported for field-grown seed. Potato tubers showed little difference from field-grown tubers. With the exception of increased crude fiber of soybean seed with increased CO2, no trends were apparent with regard to CO2 effects on proximate composition of soybean seed and potato tubers. Crude fiber of soybean stems and leaves increased with increased CO2, as did soybean leaf protein (total nitrogen). Potato leaf and stem (combined) protein levels also increased with increased CO2, while leaf and stem carbohydrates decreased. Values for leaf and stem protein and ash were higher than values generally reported for field-grown plants for both species. Results suggest that CO2 partial pressure should have little influence on proximate composition of potato tubers or soybean seed, but that high ash and protein levels might be expected from leaves and stems of crops grown in controlled environments of a CELSS.     

Photosynthesis and respiration of a wheat stand at reduced atmospheric pressure and reduced oxygen.

A 34-day functional test was conducted in Johnson Space Center's Variable Pressure Growth Chamber (VPGC) to determine responses of a wheat stand to reduced pressure (70 kPa) and modified partial pressures of carbon dioxide and oxygen. Reduced pressure episodes were generally six to seven hours in duration, were conducted at reduced ppO2 (14.7 kPa), and were interrupted with longer durations of ambient pressure (101 kPa). Daily measurements of stand net photosynthesis (Pn) and dark respiration (DR) were made at both pressures using a ppCO2 of 121 Pa. Corrections derived from leakage tests were applied to reduced pressure measurements. Rates of Pn at reduced pressure averaged over the complete test were 14.6% higher than at ambient pressure, but rates of DR were unaffected. Further reductions in ppO2 were achieved with a molecular sieve and were used to determine if Pn was enhanced by lowered O2 or by lowered pressure. Decreased ppO2 resulted in enhanced rates of Pn, regardless of pressure, but the actual response was dependent on the ratio of ppO2/ppCO2. Over the range of ppO2/ppCO2 of 80 to 200, the rate of Pn declined linearly. Rate of DR was unaffected over the same range and by dissolved O2 levels down to 3.1 ppm, suggesting that normal rhizosphere and canopy respiration occur at atmospheric ppO2 levels as low as 11 kPa. Partial separation of effects attributable to oxygen and those related to reduced pressure (e.g. enhanced diffusion of CO2) was achieved from analysis of a CO2 drawdown experiment. Results will be used for design and implementation of studies involving complete crop growth tests at reduced pressure.     

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.     

The effects of CO2 on growth and transpiration of radish (Raphanus sativus) in hypobaria

Plants grown on long-term space missions will likely be grown in low pressure environments (i.e., hypobaria). However, in hypobaria the transpiration rates of plants can increase and may result in wilting if the water is not readily replaced. It is possible to reduce transpiration by increasing the partial pressure of CO₂ (pCO₂), but the effects of pCO₂ at high levels (>120 Pa) on the growth and transpiration of plants in hypobaria are not known. Therefore, the effects of pCO₂ on the growth and transpiration of radish (Raphanus sativus var. Cherry Bomb II) in hypobaria were studied. The fresh weight (FW), leaf area, dry weight (DW), CO₂ assimilation rates (C_A), dark respiration rates (DR), and transpiration rates from 26 day-old radish plants that were grown for an additional seven days at different total pressures (33, 66 or 101 kPa) and pCO₂ (40 Pa, 100 Pa and 180 Pa) were measured. In general, the dry weight of plants increased with CO₂ enrichment and with lower total pressure. In limiting pCO₂ (40 Pa) conditions, the transpiration for plants grown at 33 kPa was approximately twice that of controls (101 kPa total pressure with 40 Pa pCO₂). Increasing the pCO₂ from 40 Pa to 180 Pa reduced the transpiration rates for plants grown in hypobaria and in standard atmospheric pressures. However, for plants grown in hypobaria and high pCO₂ (180 Pa) leaf damage was evident. Radish growth can be enhanced and transpiration reduced in hypobaria by enriching the gas phase with CO₂ although at high levels leaf damage may occur.     

Plants survive rapid decompression: Implications for bioregenerative life support

Radish (Raphanus sativus), lettuce (Latuca sativa), and wheat (Triticum aestivum) plants were grown at either 98 kPa (ambient) or 33 kPa atmospheric pressure with constant 21 kPa oxygen and 0.12 kPa carbon dioxide in atmospherically closed pressure chambers. All plants were grown rockwool using recirculating hydroponics with a complete nutrient solution. At 20 days after planting, chamber pressures were pumped down as rapidly as possible, reaching 5 kPa after about 5 min and ∼1.5 kPa after about 10 min. The plants were held at 1.5 kPa for 30 min and then pressures were restored to their original settings. Temperature (22 °C) and humidity (65% RH) controls were engaged throughout the depressurization, although temperatures dropped to near 16 °C for a brief period. CO₂ and O₂ were not detectable at the low pressure, suggesting that most of the 1.5 kPa atmosphere consisted of water vapor. Following re-pressurization, plants were grown for another 7 days at the original pressures and then harvested. The lettuce, radish, and wheat plants showed no visible effects from the rapid decompression, and there were no differences in fresh or dry mass when compared to control plants maintained continuously at 33 or 98 kPa. But radish storage root fresh mass and lettuce head fresh and dry masses were less at 33 kPa compared to 98 kPa for both the controls and decompression treatment. The results suggest that plants are extremely resilient to rapid decompression, provided they do not freeze (from evaporative cooling) or desiccate. The water of the hydroponic system was below the boiling pressure during these tests and this may have protected the plants by preventing pressures from dropping below 1.5 kPa and maintaining humidity near 1.5 kPa. Further testing is needed to determine how long plants can withstand such low pressure, but the results suggest there are at least 30 min to respond to catastrophic pressure losses in a plant production chamber that might be used for life support in space.     

Atmospheric leakage and method for measurement of gas exchange rates of a crop stand at reduced pressure.

The variable pressure growth chamber (VPGC) was used in a 34-day functional test to grow a wheat crop using reduced pressure (70 kPa) episodes totalling 131 hours. Primary goals of the test were to verify facility and subsystem performance at 70 kPa and to determine responses of a wheat stand to reduced pressure and modified partial pressures of carbon dioxide and oxygen. Operation and maintenance of the chamber at 70 kpa involved continuous evacuation of the chamber atmosphere, leading to CO2 influx and efflux. A model for calculating CO2-exchange rates (net photosynthesis and dark respiration) was developed and tested and involved measurements of chamber leakage to determine appropriate corrections. Measurement of chamber leakage was based on the rate of pressure change over a small pressure increment (70.3 to 72.3 kPa) with the pump disabled. Leakage values were used to correct decreases and increases in chamber CO2 concentration resulting from net photosynthesis (Ps) and dark respiration (DR), respectively. Composite leakage corrections (influx and efflux) at day 7 of the test were 9% and 19% of the changes measured for Ps and DR, respectively. On day 33, composite corrections were only 3% for Ps and 4% for DR. During the test, the chamber became progressively tighter; the leak rate at 70.3 kPa decreasing from 2.36 chamber volumes/day pretest, to 1.71 volumes/day at the beginning of the test, and 1.16 volumes/day at the end of the test. Verification of the short-term leakage tests (rate of pressure rise) were made by testing CO2 leakage with the vacuum pump enabled and disabled. Results demonstrate the suitability of the VPGC or conducting gas exhange measurements of a crop stand at reduced pressure.     

Effects of modified atmosphere on crop productivity and mineral content.

Wheat, potato, pea and tomato crops were cultivated from seeding to harvest in a controlled and confined growth chamber at elevated CO2 concentration (3700 microL L-1) to examine the effects on biomass production and edible part yields. Different responses to high CO2 were recorded, ranging from a decline in productivity for wheat, to slight stimulation for potatoes, moderate increase for tomatoes, and very large enhancement for pea. Mineral content in wheat and pea seeds was not greatly modified by the elevated CO2. Short-term experiments (17 d) were conducted on potato at high (3700 microL L-1) and very high (20,000 microL L-1) CO2 concentration and/or low O2 partial pressure (approximately 20,600 microL L-1 or 2 kPa). Low O2 was more effective than high CO2 in total biomass accumulation, but development was affected: Low O2 inhibited tuberization, while high CO2 significantly increased production of tubers.     

Effects of temperature, CO2/O2 concentrations and light intensity on cellular multiplication of microalgae, Euglena gracilis.

Microalgae culture is likely to play an important role in aquatic food production modules in bioregenerative systems for producing feeds for fish, converting CO2 to O2 and remedying water quality as well as aquatic higher plants. In the present study, the effects of culture conditions on the cellular multiplication of microalgae, Euglena gracilis, was investigated as a fundamental study to determine the optimum culture conditions for microalgae production in aquatic food production modules including both microalgae culture and fish culture systems. E. gracilis was cultured under conditions with five levels of temperatures (25-33 degrees C), three levels of CO2 concentrations (2-6%), five levels of O2 concentrations (10-30%), and six levels of photosynthetic photon flux (20-200 micromoles m-2 s-1). The number of Euglena cells in a certain volume of solution was monitored with a microscope under each environmental condition. The multiplication rate of the cells was highest at temperatures of 27-31 degrees C, CO2 concentration of 4%, O2 concentration of 20% and photosynthetic photon flux of about 100 micromoles m-2 s-1. The results demonstrate that E. gracilis could efficiently produce biomass and convert CO2 to O2 under relatively low light intensities in aquatic food production modules.     

Effects of several environmental factors on sweetpotato growth.

Effects of relative humidity, light intensity and photoperiod on growth of 'Ga Jet' and 'TI-155' sweetpotato cultivars, using the nutrient film technique (NFT), have been reported. In this study, the effect of ambient temperature regimes (constant 28 degrees C and diurnal 28:22 degrees C day:night) and different CO2 levels (ambient, 400, 1000 and 10000 microliters/L--400, 1000 and 10000 ppm) on growth of one or both of these cultivars in NFT are reported. For a 24-h photoperiod, no storage roots were produced for either cultivar in NFT when sweetpotato plants were grown at a constant temperature of 28 degrees C. For the same photoperiod, when a 28:22 degrees C diurnal temperature variation was used, there were still no storage roots for 'TI-155' but the cv. 'Ga Jet' produced 537 g/plant of storage roots. For both a 12-h and 24-h photoperiod, 'Ga Jet' storage root fresh and dry weight tended to be higher with a 28:22 degrees C diurnal temperature variation than with a constant 28 degrees C temperature regime. Preliminary results with both 'Ga Jet' and 'TI 155' cultivars indicate a distinctive diurnal stomatal response for sweetpotato grown in NFT under an ambient CO2 level. The stomatal conductance values observed for 'Ga Jet' at elevated CO2 levels indicated that the difference between the light- and dark-period conductance rates persisted at 400, 1000, and 10000 microliters/L.     

Plant responses to short- and long-term exposures to high carbon dioxide levels in closed environments.

When higher plants are exposed to elevated levels of CO2 for both short- and long-term periods photosynthetic C-gain and photoassimilate export from leaves are generally increased. Water use efficiency is increased on a leaf area basis. During long-term exposures, photosynthesis rates on leaf and whole plants bases are altered in a species specific manner. The most common pattern in C3 plants is an enhanced rate of whole plant photosynthesis in a well irradiated canopy. Nevertheless, in some herbaceous species prolonged exposure to high CO2 results in remobilization of nitrogenous reserves (i.e., leaf protein degradation) and reduced rates of mature leaf photosynthesis when assayed at ambient CO2 and O2 levels. Both short- and long-term exposures to those CO2 levels (i.e., 100 to 2,000 microliter l-1) which modify photosynthesis and export, also modify both endogenous ethylene gas (C2H4) release, and substrate, 1-aminocyclopropane-1-carboxylic acid (ACC), saturated C2H4 release rates from irradiated leaves. Photosynthetically active canopy leaves contribute most of the C2H4 released from the canopy. Prolonged growth at high CO2 results in a persistent increase in the rate of endogenous C2H4 release from leaves which can, only in part, be attributed to the increase of the endogenous pools of C2H4 pathway intermediates (e.g., methionine, M-ACC, and ACC). The capacity for increasing the rate of C2H4 release in response to short-term exposures to varying CO2 levels does not decline after prolonged growth at high CO2. When leaves, whole plants, and model canopies of tomato plants are exposed to exogenous C2H4 a reduction in the rate of photosynthesis can, in each case, be attributed to the classical effects of C2H4 on plant development and morphology. The effect of C2H4 on CO2 gas exchange of plant canopies is shown to be dependent on the canopy leaf area index.     

Comparison of aerobically-treated and untreated crop residue as a source of recycled nutrients in a recirculating hydroponic system.

This study compared the growth of potato plants on nutrients recycled from inedible potato biomass. Plants were grown for 105 days in recirculating, thin-film hydroponic systems containing four separate nutrient solution treatments: (1) modified half-strength Hoagland's (control), 2) liquid effluent from a bioreactor containing inedible potato biomass, 3) filtered (0.2 micrometer) effluent, and 4) the water soluble fraction of inedible potato biomass (leachate). Approximately 50% of the total nutrient requirement in treatments 2-4 were provided (recycled) from the potato biomass. Leachate had an inhibitory effect on leaf conductance, photosynthetic rate, and growth (50% reduction in plant height and 60% reduction in tuber yield). Plants grown on bioreactor effluent (filtered or unfiltered) were similar to the control plants. These results indicated that rapidly degraded, water soluble organic material contained in the inedible biomass, i.e., material in leachate, brought about phytotoxicity in the hydroponic culture of potato. Recalcitrant, water soluble organic material accumulated in all nutrient recycling treatments (650% increase after 105 days), but no increase in rhizosphere microbial numbers was observed.     

Proximate composition of CELSS crops grown in NASA's Biomass Production Chamber.

Edible biomass from four crops of wheat (Triticum aestivum L.), four crops of lettuce (Lactuca sativa L.), four crops of potato (Solanum tuberosum L.), and three crops of soybean (Glycine max (L.) Merr.) grown in NASA's CELSS Biomass Production Chamber were analyzed for proximate composition. All plants were grown using recirculating nutrient (hydroponic) film culture with pH and electrical conductivity automatically controlled. Temperature and humidity were controlled to near optimal levels for each species and atmospheric carbon dioxide partial pressures were maintained near 100 Pa during the light cycles. Soybean seed contained the highest percentage of protein and fat, potato tubers and wheat seed contained the highest levels of carbohydrate, and lettuce leaves contained the highest level of ash. Analyses showed values close to data published for field-grown plants with several exceptions: In comparison with field-grown plants, wheat seed had higher protein levels; soybean seed had higher ash and crude fiber levels; and potato tubers and lettuce leaves had higher protein and ash levels. The higher ash and protein levels may have been a result of the continuous supply of nutrients (e.g., potassium and nitrogen) to the plants by the recirculating hydroponic culture.     

The CELSS Test Facility project: an example of a CELSS flight experiment system.

The CELSS Test Facility (CTF) is a device for measuring crop plant productivity in the micro-gravity environment of Space Station Freedom. It will allow us to address questions of crop productivity in space, versus that on the ground. The crop productivity factors that will be measured are rates of: 1) biomass production, 2) food production, 3) O2 and CO2 exchange, and 4) water transpiration. In addition, other productivity factors of specific crops will be determined, such as : 1) the ratio of edible to inedible biomass (harvest index), 2) leaf area exposed to and collecting light (leaf area index), 3) ratio of root mass to total biomass, and 4) photosynthetic efficiency (ratio of moles of CO2 fixed (or O2 produced), per mole of photons of specific energies used). Plant and crop morphology, at several levels, ranging from the community to the sub-cellular, will also be evaluated.     

CO2 crop growth enhancement and toxicity in wheat and rice.

The effects of elevated CO2 on plant growth are reviewed and the implications for crop yields in regenerative systems are discussed. There is considerable theoretical and experimental evidence indicating that the beneficial effects of CO2 are saturated at about 0.12% CO2 in air. However, CO2 can easily rise above 1% of the total gas in a closed system, and we have thus studied continuous exposure to CO2 levels as high as 2%. Elevating CO2 from 340 to 1200 micromoles mol-1 can increase the seed yield of wheat and rice by 30 to 40%; unfortunately, further CO2 elevation to 2500 micromoles mol-1 (0.25%) has consistently reduced yield by 25% compared to plants grown at 1200 micromoles mol-1; fortunately, there was only an additional 10% decrease in yield as the CO2 level was further elevated to 2% (20,000 micromoles mol-1). Yield increases in both rice and wheat were primarily the result of increased number of heads per m2, with minor effects on seed number per head and seed size. Yield increases were greatest in the highest photosynthetic photon flux. We used photosynthetic gas exchange to analyze CO2 effects on radiation interception, canopy quantum yield, and canopy carbon use efficiency. We were surprised to find that radiation interception during early growth was not improved by elevated CO2. As expected, CO2 increased quantum yield, but there was also a small increase in carbon use efficiency. Super-optimal CO2 levels did not reduce vegetative growth, but decreased seed set and thus yield. The reduced seed set is not visually apparent until final yield is measured. The physiological mechanism underlying CO2 toxicity is not yet known, but elevated CO2 levels (0.1 to 1% CO2) increase ethylene synthesis in some plants and ethylene is a potent inhibitor of seed set in wheat.     

Super-optimal CO2 reduces wheat yield in growth chamber and greenhouse environments.

Seven growth chamber trials (six replicate trials using 0.035, 0.12, and 0.25% CO2 in air and one trial using 0.12, 0.80, and 2.0% CO2 in air) and three replicate greenhouse trials (0.035, 0.10, 0.18, 0.26, 0.50, and 1.0% CO2 in air) compare the effects of super-optimal CO2 on the seed yield, harvest index, and vegetative growth rate of wheat (Triticum aestivum L. cvs. USU-Apogee and Veery-10). Plants in the growth chamber trials were grown hydroponically under fluorescent lamps, while the greenhouse trials were grown under sunlight and high pressure sodium lamps and in soilless media. Plants in the greenhouse trials responded similarly to those in the growth chamber trials; maximum yields occurred near 0.10 and 0.12% CO2 and decreased significantly thereafter. This research indicates that the toxic effects of elevated CO2 are not specific to only one environment and has important implications for the design of bio-regenerative life support systems in space, and for the future of terrestrial agriculture.     

Effects of CO2 concentration and light intensity on photosynthesis of a rootless submerged plant, Ceratophyllum demersum L., used for aquatic food production in bioregenerative life support systems.

In addition to green microalgae, aquatic higher plants are likely to play an important role in aquatic food production modules in bioregenerative systems for producing feed for fish, converting CO2 to O2 and remedying water quality. In the present study, the effects of culture conditions on the net photosynthetic rate of a rootless submerged plant, Ceratophyllum demersum L., was investigated to determine the optimum culture conditions for maximal function of plants in food production modules including both aquatic plant culture and fish culture systems. The net photosynthetic rate in plants was determined by the increase in dissolved O2 concentrations in a closed vessel containing a plantlet and water. The water in the vessel was aerated sufficiently with a gas containing a known concentration of CO2 gas mixed with N2 gas before closing the vessel. The CO2 concentrations in the aerating gas ranged from 0.3 to 10 mmol mol-1. Photosynthetic photon flux density (PPFD) in the vessel ranged from 0 (dark) to 1.0 mmol m-2 s-1, which was controlled with a metal halide lamp. Temperature was kept at 28 degrees C. The net photosynthetic rate increased with increasing PPFD levels and was saturated at 0.2 and 0.5 mmol m-2 s-1 PPFD under CO2 levels of 1.0 and 3.0 mmol mol-1, respectively. The net photosynthetic rate increased with increasing CO2 levels from 0.3 to 3.0 mmol mol-1 showing the maximum value, 75 nmol O2 gDW-1 s-1, at 2-3 mmol mol-1 CO2 and gradually decreased with increasing CO2 levels from 3.0 to 10 mmol mol-1. The results demonstrate that C. demersum could be an efficient CO2 to O2 converter under a 2.0 mmol mol-1 CO2 level and relatively low PPFD levels in aquatic food production modules.