Toluene removal from air by Dieffenbachia in a closed environment.

Higher plants are likely to play a major role in bioregeneration systems for food, air and water supplies. Plants may also contribute by the removal of toxic organic substances from the air of a closed environment. Dieffenbachia amoena plants were exposed to 0 to 1.2 x 10(6) micrograms toluene m-3 at light intensities of 35 and 90 micromoles m-2 s-1 in sealed chambers. Toluene removal, photosynthesis and respiration were measured. An increased light intensity increased the rate of toluene removal five-fold over the rate at the lower intensity; the kinetics suggest active regulation by the plant. The removal rate saturated at 2700 micrograms toluene h-1 at the lower intensity and failed to saturate at the higher intensity. Toluene exposure inhibited photosynthesis and respiration only transiently and without correlation to toluene concentration. These plants can act as efficient scavengers of toluene in a contaminated environment.     

Carbon dioxide interactions with irradiance and temperature in potatoes.

Separate controlled environment studies were conducted to determine the interaction of CO2 with irradiance and interaction of CO2 with temperature on growth of three potato cultivars. In the first study, an elevated CO2 concentration of 1000 micromoles mol-1 and an ambient CO2 of 350 micromoles mol-1 were maintained at the photosynthetic photon fluxes (PPF) of 17 and 34 mol m-2 d-1 with 12 h photoperiod, and at the PPF of 34 and 68 mol m-2 d-1 with 24 h photoperiod (400 and 800 micromoles m-2 s-1 PPF at each photoperiod). Tuber and total dry weights of 90-day old potatoes were significantly increased with CO2 enrichment, but the CO2 stimulation was less with higher PPF and longer photoperiod. Shoot dry weight was affected more by photoperiod than by PPF and CO2 concentrations. The elevated CO2 concentration increased leaf CO2 assimilation rates and decreased stomatal conductance with 12 h photoperiod, but had only a marginal effect with 24 h photoperiod. In the second study, four CO2 concentrations of 500, 1000, 1500 and 2000 micromoles mol-1 were combined with two air temperature regimes of 16 and 20 degrees C under a 12 h photoperiod. At harvest, 35 days after transplanting, tuber and total dry weights of potatoes reached a maximum with 1000 micromoles mol-1 CO2 at 16 degrees C, but continued to increase up to 2000 micromoles mol-1 CO2 at 20 degrees C. Plant growth was greater at 20 degrees C than at 16 degrees C under all CO2 concentrations. At 16 degrees C specific leaf weight increased substantially with increasing CO2 concentrations as compared to 500 micromoles mol-1 CO2, but increased only slightly at 20 degrees C. This suggests a carbohydrate build-up in the leaves at 16 degrees C temperature that reduces plant response to increased CO2 concentrations. The data in the two studies indicate that a PPF of 34 mol m-2 d-1, 20 degrees C temperature, and 1000-2000 micromoles mol-1 CO2 produces optimal tuber yield in potatoes.     

Light intensity and production parameters of phytocenoses cultivated on soil-like substrate under controlled [correction of controled] environment conditions.

To increase the degree of closure of biological life support systems of a new generation, we used vermicomposting to involve inedible phytomass in the intra-system mass exchange. The resulting product was a soil-like substrate, which was quite suitable for growing plants (Manukovsky et al. 1996, 1997). However, the soil like substrate can be regarded as a candidate for inclusion in a system only after a comprehensive examination of its physical, chemical, and other characteristics. An important criterion is the ability of the soil-like substrate to supply the necessary mineral elements to the photosynthesizing component under the chosen cultivation conditions. Thus, the purpose of this work was to study the feasibility of enhancing the production activity of wheat and radish crops by varying the intensity of photosynthetically active radiation, without decreasing the harvest index. The increase of light intensity from 920 to 1150 micromoles m-2 s-1 decreased the intensity of apparent photosynthesis of the wheat crops and slightly increased the apparent photosynthesis of the radish crops The maximum total and grain productivity (kg/m2) of the wheat crops was attained at the irradiance of 920 micromoles m-2 s-1. Light intensity of 1150 micromoles m-2 s-1 decreased the productivity of wheat plants and had no significant effect on the productivity of the radish crops (kg/m2) as compared to 920 micromoles m-2 s-1. The qualitative and quantitative composition of microflora of the watering solution and substrate was determined by the condition of plants, developmental phase and light intensity. By the end of wheat growth under 1150 micromoles m-2 s-1 the numbers of bacteria of the coliform family and phytopathogenic bacteria in the watering solution and substrate were an order of magnitude larger than under other illumination conditions. The obtained data suggest that the cultivation of plants in a life support system on soil-like substrate from composts has a number of advantages over the cultivation on neutral substrates, which require continual replenishment of the plant nutrient solution from the system's store to complement the macro- and micro-elements. Yet, a number of problems arise, including those related to the controlling of the production activity of the plants by the intensity of photosynthetically active radiation. It is essential to understand why the intensity of production processes is limited at higher irradiation levels and to overcome the factors responsible for this, so that the soil-like substrate could have an even better chance in the competition for the best plant cultivation technology to be used in biological life support systems.     

Preparations for CELSS flight experiments with wheat.

We are planning a short-term experiment with Superdwarf wheat on the U.S. Space Shuttle and a seed-to-seed experiment on the Russian Space Station Mir. The goals of both experiments are to observe effects of microgravity on developmental steps in the life cycle and to measure photosynthesis, respiration, and transpiration by monitoring gas exchange. This requires somewhat different hardware development for the two experiments. Ground-based research aims to understand plant responses to the environments in the space growth chambers that we will use (after some modification): the Plant Growth Unit (PGU) on the shuttle and units called Svet, Svetoblock 2, or Oasis on Mir. Low irradiance levels (100 to 250 micromoles m-2 s-1 at best) pose a particular problem. Water and nutrient supply are also potentially limiting factors, especially in the long-term experiment. Our ground-based studies emphasize responses to low light levels (50 to 400 micromoles m-2 s-1); results show that all developmental steps are delayed by low light compared with plants at 400 micromoles m-2 s-1. We are also testing various rooting substrates for the shuttle experiment. A 1:1:1 mixture of peat:perlite:vermiculite appears to be the best choice.     

Carbon balance and productivity of Lemna gibba, a candidate plant for CELSS.

The photosynthesis and productivity of Lemna gibba were studied with a view to its use in Controlled Ecological Life Support Systems (CELSS). Photosynthesis of L. gibba floating on the nutrient solution could be driven by light coming from either above or below. Light from below was about 75% as effective as from above when the stand was sparse, but much less so with dense stands. High rates of photosynthesis (ca. 800 nanomoles CO2 g dry weight (DW)-1 s-1) were measured at 750 micromoles m-2 s-1 PPF and 1500 micromoles mol-1 CO2. This was attained at densities up to 660 g fresh weight (FW) m-2 with young cultures. After a few days growth under these conditions, and at higher densities, the rate of photosynthesis dropped to less than 25% of the initial value. This drop was only partly alleviated by thinning the stand or by introducing a short dark period at high temperature (26 degrees C). Despite the drop in the rate of photosynthesis, maximum yields were obtained in batch cultures grown under continuous light, constant temperature and high [CO2]. Plant protein content was less than reported for field grown Lemna. When the plants were harvested daily, maintaining a stand density of 600 g FW m-2, yields of 18 g DW m-2 d-1 were obtained. The total dry weight of L. gibba included 40% soluble material (sugars and amino acids), 15% protein, 5% starch, 5% ash and 35% cellulose and other polymers. We conclude that a CELSS system could be designed around stacked, alternate layers of transparent Lemna trays and lamps. This would allow for 7 tiers per meter height. Based on present data from single layers, the yield of such a system is calculated to be 135 g DW m-3 d-1 of a 100% edible, protein-rich food.     

Current and potential productivity of wheat for a Controlled Environment Life Support System.

The productivity of higher plants is determined by the incident photosynthetic photon flux (PPF) and the efficiency of the following four physiological processes: absorption of PPF by photosynthetic tissue, carbon fixation (photosynthesis), carbon use (respiration), and carbon partitioning (harvest index). These constituent processes are analyzed to determine theoretical and potentially achievable productivity. The effects of optimal environmental and cultural factors on each of these four factors is also analyzed. Results indicate that an increase in the percentage of absorbed photons is responsible for most of the improvement in wheat yields in an optimal controlled environment. Several trials confirm that there is an almost linear increase in wheat yields with increasing PPF. An integrated PPF of 150 mol m-2 d-1 (2.5 times summer sunlight) has produced 60 g m-2 d-1 of grain. Apparently, yield would continue to increase with even higher PPF's. Energy efficiency increased with PPF to about 600 micromoles m-2 s-1, then slowly decreased. We are now seeking to improve efficiency at intermediate PPF levels (1000 micromoles m-2 s-1) before further exploring potential productivity. At intermediate and equal integrated daily PPF levels, photoperiod had little effect on yield per day or energy efficiency. Decreasing temperature from 23 degrees to 17 degrees increased yield per day by 20% but increased the life cycle from 62 to 89 days. We hope to achieve both high productivity and energy efficiency.     

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.     

Utilization of potatoes in bioregenerative life support systems.

Data on the tuberization, harvest index, and morphology of 2 cvs of white potato (Solanum tuberosum L.) grown at 12, 16, 20, 24 and 28 degrees C, 250, 400 and 550 micromoles s-1 m-2 photosynthetic photon flux (PPF), 350, 1000 and 1600 microliters l-1 CO2 will be presented. A productivity of 21.9 g m-2 day-1 of edible tubers from a solid stand of potatoes grown for 15 weeks with continuous irradiation at 400 micromoles s-1 m-2, 16 degrees C and 1000 microliters l-1 CO2 has been obtained. This equates to an area of 34.3 m2 being required to provide 2800 kcal of potatoes per day for a human diet. Separated plants receiving side lighting have produced 32.8 g m-2 day-1 which equates to an area of 23.6 m2 to provide 2800 kcal. Studies with side lighting indicate that productivities in this range should be realized from potatoes. Glycoalkaloid levels in tubers of controlled-environment-grown plants are within the range of levels found in tubers of field grown plants. The use and limitation of recirculating solution cultures for potato growth is discussed.     

Utilization of white potatoes in CELSS.

Potatoes (Solanum tuberosum) have a strong potential as a useful crop species in a functioning CELSS. The cultivar Denali has produced 37.5 g m-2 d-1 when grown for 132 days with the first 40 days under a 12-h photoperiod and a light:dark temperature cycle of 20 degrees C:16 degrees C, and then 92 days under continuous irradiance and a temperature of 16 degrees C. Irradiance was at 725 micromoles m-2 s-1 PPF and carbon dioxide at 1000 micromoles mol-1. The dried tubers had 82% carbohydrates, 9% protein and 0.6% fat. Other studies have shown that carbon dioxide supplementation (1000 micromoles mol-1) is of significant benefit under 12-h irradiance but less benefit under 24 h irradiance. Irradiance cycles of 60 minutes light and 30 minutes dark caused a reduction of more than 50% in tuber weight compared to cycles of 16 h light and 8 h dark. A diurnal temperature change of 22 degrees C for the 12-h light period to 14 degrees C during the 12-h dark period gave increased yields of 30% and 10% for two separate cultivars, compared with plants grown under a constant 18 degrees C temperature. Cultivar screening under continuous irradiance and elevated temperatures (28 degrees C) for 8 weeks of growth indicated that the cvs Haig, Denali, Atlantic, Desiree and Rutt had the best potential for tolerance to these conditions. Harvesting of tubers from plants at weekly intervals, beginning at 8 weeks after planting, did not increase yield over a single final harvest. Spacing of plants on 0.055 centers produced greater yield per m2 than spacing at 0.11 or 0.22 m2. Plants maintained 0.33 meters apart (0.111 m2 per plant) in beds produced the same yields when separated by dividers in the root matrix as when no separation was made.     

Atmospheric dynamics in the "Laboratory Biosphere" with wheat and sweet potato crops.

Laboratory Biosphere is a 40-m3 closed life system equipped with 12,000 W of high pressure sodium lamps over planting beds with 5.37 m2 of soil. Atmospheric composition changes due to photosynthetic fixation of carbon dioxide and corresponding production of oxygen or the reverse, respiration, are observed in short timeframes, e.g., hourly. To focus on inherent characteristics of the crop as distinct from its area or the volume of the chamber, we report fixation and respiration rates in mmol h-1 m-2 of planted area. An 85-day crop of USU Apogee wheat under a 16-h lighted/8-h dark regime peaked in fixation rate at about 100 mmol h-1 m-2 approximately 24 days after planting. Light intensity was about 840 micromoles m-2 s-1. Dark respiration peaked at about 31 mmol h-1 m-2 at the same time. Thereafter, both fixation and respiration declined toward zero as harvest time approached. A residual soil respiration rate of about 1.9 mmol h-1 m-2 was observed in the dark closed chamber for 100 days after the harvest. A 126-day crop of Tuskegee TU-82-155 sweet potato behaved quite differently. Under a 680 micromoles m-2 s-1, 18-h lighted/6-h dark regime, fixation during lighted hours rose to a plateau ranging from about 27 to 48 mmol h-1 m-2 after 42 days and dark respiration settled into a range of 12-23 mmol h-1 m-2. These rates continued unabated until the harvest at 126 days, suggesting that tuber biomass production might have continued at about the same rate for some time beyond the harvest time that was exercised in this experiment. In both experiments CO2 levels were allowed to range widely from a few hundred to about 3000 ppm, which permitted observation of fixation rates both at varying CO2 concentrations and at each number of days after planting. This enables plotting the fixation rate as a function of both variables. Understanding the atmospheric dynamics of individual crops will be essential for design and atmospheric management of more complex CELSS which integrate the simultaneous growth of several crops as in a sustainable remote life support system.     

Manipulating light and temperature to minimize environmental stress in the plant component of bioregenerative life support systems.

Our experiments examined enhancing tolerance of the photosynthesizing component to possible deviations in thermal or illumination conditions inside a bioregenerative life support system (BLSS). In the event of one parameter getting beyond its optimum, the values of other parameters may ensure minimal damage to the plant component during the period of environmental stress. With wheat plants (one of key elements of the plant component) as an example the work considers whether it is possible to enhance thermal tolerance by varying light intensity. Increase of air temperature to 35 degrees C or 45 degrees C with light intensity of 60 W/m2 PAR has been shown to substantially inhibit the photosynthesis processes; at 150 W/m2 PAR photosynthesis decreases from 50% to 100%, respectively; when light intensity is increased to 240 W/m2 PAR photosynthesis increased more than 70% at 35 degrees C and decreased at 45 degrees C by only 20%. Thus, light intensity can be increased to avoid or decrease the inhibiting effect of high temperatures. On the other hand, tolerance of wheat plants to prolonged absence of light can be substantially enhanced by decreasing during this period air temperature to temperatures close to 0 degrees C.     

Wheat production in controlled environments.

Our goal is to optimize conditions for maximum yield and quality of wheat to be used in a controlled-environment, life-support system (CELSS) in a Lunar or Martian base or perhaps in a space craft. With yields of 23 to 57 g m-2 d-1 of edible biomass, a minimum size for a CELSS would be between 12 and 30 m2 per person, utilizing about 600 W m-2 of electrical energy for artificial light. Temperature, irradiance, photoperiod, carbon-dioxide levels, humidity, and wind velocity are controlled in state-of-the-art growth chambers. Nutrient solutions (adjusted for wheat) are supplied to the roots via a recirculating system that controls pH by adding HNO3 and controlling the NO3/NH4 ratio in solution. A rock-wool plant support allows direct seeding and densities up to 10,000 plants per meter2. Densities up to 2000 plants m-2 appear to increase seed yield. Biomass production increases almost linearly with increasing irradiance from 400 to 1700 micromoles m-2 s-1 of photosynthetic photon flux (PPF), but the efficiency of light utilization decreases over this range. Photoperiod and temperature both have a profound influence on floral initiation, spikelet formation, stem elongation, and fertilization. High temperatures (25 to 27 degrees C) and long days shorten the life cycle and promote rapid growth, but cooler temperatures (20 degrees C) and shorter days greatly increase seed number per head and thus yield (g m-2). The life cycle is lengthened in these conditions but yield per day (g m-2 d-1) is still increased. We have evaluated about 600 cultivars from around the world and have developed several breeding lines for our controlled conditions. Some of our ultra-dwarf lines (30 to 50 cm tall) look especially promising with high yields and high harvest indices (percent edible biomass).     

Evaluation of light emitting diode characteristics for a space-based plant irradiation source.

Light emitting diodes (LEDs) are a promising irradiation source for plant growth in space. Improved semiconductor technology has yielded LED devices fabricated with gallium aluminum arsenide (GaAlAs) chips which have a high efficiency for converting electrical energy to photosynthetically active radiation. Specific GaAlAs LEDs are available that emit radiation with a peak wavelength near the spectral peak of maximum quantum action for photosynthesis. The electrical conversion efficiency of installed systems (micromole s-1 of photosynthetic photons per watt) of high output LEDs can be within 10% of that for high pressure sodium lamps. Output of individual LEDs were found to vary by as much as 55% from the average of the lot. LED ratings, in mcd (luminous intensity per solid angle), were found to be proportional to total photon output only for devices with the same dispersion angle and spectral peak. Increasing current through the LED increased output but also increased temperature with a consequent decrease in electrical conversion efficiency. A photosynthetic photon flux as high as 900 micromoles m-2 s-1 has been produced on surfaces using arrays with LEDs mounted 7.6 mm apart, operating as a current of 50 mA device-1 and at an installed density of approximately 17,200 lamps m-2 of irradiated area. Advantages of LEDs over other electric light sources for use in space systems include long life, minimal mass and volume and being a solid state device.     

Effects of air velocity on photosynthesis of plant canopies under elevated CO2 levels in a plant culture system.

To obtain basic data for adequate air circulation for promoting plant growth in closed plant production modules in bioregenerative life support systems in space, effects of air velocities ranging from 0.1 to 0.8 m s-1 on photosynthesis in tomato seedlings canopies were investigated under atmospheric CO2 concentrations of 0.4 and 0.8 mmol mol-1. The canopy of tomato seedlings on a plug tray (0.4 x 0.4 m2) was set in a wind-tunnel-type chamber (0.6 x 0.4 x 0.3 m3) installed in a semi-closed-type assimilation chamber (0.9 x 0.5 x 0.4 m3). The net photosynthetic rate in the plant canopy was determined with the differences in CO2 concentrations between the inlet and outlet of the assimilation chamber multiplied by the volumetric air exchange rate of the chamber. Photosynthetic photon flux (PPF) on the plant canopy was kept at 0.25 mmol m-2 s-1, air temperature at 23 degrees C and relative humidity at 55%. The leaf area indices (LAIs) of the plant canopies were 0.6-2.5 and plant heights were 0.05-0.2 m. The net photosynthetic rate of the plant canopy increased with increasing air velocities inside plant canopies and saturated at 0.2 m s-1. The net photosynthetic rate at the air velocity of 0.4 m s-1 was 1.3 times that at 0.1 m s-1 under CO2 concentrations of 0.4 and 0.8 mmol mol-1. The net photosynthetic rate under CO2 concentrations of 0.8 mmol mol-1 was 1.2 times that under 0.4 mmol mol-1 at the air velocity ranging from 0.1 to 0.8 m s-1. The results confirmed the importance of controlling air movement for enhancing the canopy photosynthesis under an elevated CO2 level as well as under a normal CO2 level in the closed plant production modules.     

Crop yield and light/energy efficiency in a closed ecological system: Laboratory Biosphere experiments with wheat and sweet potato.

Two crop growth experiments in the soil-based closed ecological facility, Laboratory Biosphere, were conducted from 2003 to 2004 with candidate space life support crops. Apogee wheat (Utah State University variety) was grown, planted at two densities, 400 and 800 seeds m-2. The lighting regime for the wheat crop was 16 h of light-8 h dark at a total light intensity of around 840 micromoles m-2 s-1 and 48.4 mol m-2 d-1 over 84 days. Average biomass was 1395 g m-2, 16.0 g m-2 d-1 and average seed production was 689 g m-2 and 7.9 g m-2 d-1. The less densely planted side was more productive than the denser planting, with 1634 g m-2 and 18.8 g m-2 d-1 of biomass vs. 1156 g m-2 and 13.3 g m-2 d-1; and a seed harvest of 812.3 g m-2 and 9.3 g m-2 d-1 vs. 566.5 g m-2 and 6.5 g m-2 d-1. Harvest index was 0.49 for the wheat crop. The experiment with sweet potato used TU-82-155 a compact variety developed at Tuskegee University. Light during the sweet potato experiment, on a 18 h on/6 h dark cycle, totaled 5568 total moles of light per square meter in 126 days for the sweet potatoes, or an average of 44.2 mol m-2 d-1. Temperature regime was 28 +/- 3 degrees C day/22 +/- 4 degrees C night. Sweet potato tuber yield was 39.7 kg wet weight, or an average of 7.4 kg m-2, and 7.7 kg dry weight of tubers since dry weight was about 18.6% wet weight. Average per day production was 58.7 g m-2 d-1 wet weight and 11.3 g m-2 d-1. For the wheat, average light efficiency was 0.34 g biomass per mole, and 0.17 g seed per mole. The best area of wheat had an efficiency of light utilization of 0.51 g biomass per mole and 0.22 g seed per mole. For the sweet potato crop, light efficiency per tuber wet weight was 1.33 g mol-1 and 0.34 g dry weight of tuber per mole of light. The best area of tuber production had 1.77 g mol-1 wet weight and 0.34 g mol-1 of light dry weight. The Laboratory Biosphere experiment's light efficiency was somewhat higher than the USU field results but somewhat below greenhouse trials at comparable light levels, and the best portion of the crop at 0.22 g mol-1 was in-between those values. Sweet potato production was overall close to 50% higher than trials using hydroponic methods with TU-82-155 at NASA JSC. Compared to projected yields for the Mars on Earth life support system, these wheat yields were about 15% higher, and the sweet potato yields averaged over 80% higher.     

The effect of gravity on surface temperature and net photosynthetic rate of plant leaves.

To clarify the effects of gravity on heat/gas exchange between plant leaves and the ambient air, the leaf temperatures and net photosynthetic rates of plant leaves were evaluated at 0.01, 1.0, 1.5 and 2.0 G of 20 seconds each during a parabolic airplane flight. Thermal images of leaves were captured using infrared thermography at an air temperature of 26 degrees C, a relative humidity of 15% and an irradiance of 260 W m-2. The net photosynthetic rates were determined by using a chamber method with an infrared gas analyzer at an air temperature of 20 degrees C, a relative humidity of 50% and a photosynthetic photon flux of 0.5 mmol m-2 s-1. The mean leaf temperature increased by 1 degree C and the net photosynthetic rate decreased by 13% with decreasing gravity levels from 1.0 to 0.01 G. The leaf temperature decreased by 0.5 degree C and the net photosynthetic rate increased by 7% with increasing gravity levels from 1.0 to 2.0 G. Heat/gas exchanges between leaves and the ambient air were more retarded at lower gravity levels. A restricted free air convection under microgravity conditions in space would limit plant growth by retarding heat and gas exchanges between leaves and the ambient air.     

Gravitropism and phototropism of oat coleoptiles: post-tropic autostraightening and tissue shrinkage during tropism.

We measured changes in length on the two opposite sides of the red-light-grown oat (Avena sativa L.) coleoptiles subjected to either gravitropic or phototropic stimulation and subsequently rotated on a horizontal clinostat. The length measurement was conducted using three 5 mm-long zones delimited by ink markers from the tip. Curvature of each zone was analyzed from the length difference between the two sides. Gravitropism was induced by displacing the seedling from the vertical by 30 degrees or 90 degrees for 25 min. Phototropism was induced by exposing the coleoptile to unilateral blue light for 30 s, which provided a fluence (1.0 micromoles m-2) optimal for the pulse-induced positive phototropism or a lower, suboptimal fluence (0.03 micromoles m-2). After negatively gravitropic bending, the upper two zones straightened rapidly at either displacement angle. After positively phototropic bending, straightening occurred, but only in the top zone and at the lower fluence. The upper two zones straightened rapidly, however, when bilateral blue light (30 s; 15 micromoles m-2 from either direction) was applied 25 min after unilateral stimulation at the higher fluence. Bilateral blue light alone induced no curvature. These results confirm that the straightening of gravitropically bent coleoptiles is autonomic, and suggest that a similar autonomic response participates in the straightening of phototropically bent coleoptiles. Suppression of elongation on the concave side of the coleoptile mainly accounted for gravitropic and phototropic curvatures. The concave side of the top zone shrank during both tropisms. This shrinkage progressed at a high rate from the beginning of curvature response, suggesting that a drop in turgor pressure is the main and direct cause of the shrinkage.     

Gravity-dependent reactions of the moss Pohlia nutans protonemata.

In darkness, protonemata of Pohlia nutans (Hedw.) grew negatively gravitropically (upwards). However, not all filaments became gravitropic immediately after transfer to darkness. Some of them (~20%) for several days grew in different directions with respect to gravity. The apical cells of those protonemata predominantly contained multiple chloroplasts. The intensity of chlorophyll fluorescence rapidly decreased in the apical cells of such protonemata while starch content increased in comparison with upright growing protonemata. Light, especially in the red and blue part of the spectrum, inhibited protonemal gravitropism. Red light induced stronger inhibitory effects than blue light. Red light of 1.0 to 1.5 micromoles m-2 s-1 intensity induced bud differentiation in apical cells on almost all side branches of main protonemal filaments. Bright fluorescence of F-actin bundles in the tip of apical protonematal cells and a delicately fluorescing network enclosing plastids basal to the tip in a sedimentation zone were visualized. Bright fluorescence of actin as local patches and fine prominent axially oriented bundles was observed in cells of gametophore buds.     

Red light-induced suppression of gravitropism in moss protonemata.

Moss protonemata are among the few cell types known that both sense and respond to gravity and light. Apical cells of Ceratodon protonemata grow by oriented tip growth which is negatively gravitropic in the dark or positively phototropic in unilateral red light. Phototropism is phytochrome-mediated. To determine whether any gravitropism persists during irradiation, cultures were turned at various angles with respect to gravity and illuminated so that the light and gravity vectors acted either in the same or in different directions. Red light for 24h (> or = l40nmol m-2 s-1) caused the protonemata to be oriented directly towards the light. Similarly, protonemata grew directly towards the light regardless of light position with respect to gravity indicating that all growth is oriented strictly by phototropism, not gravitropism. At light intensities < or = l00nmol m-2 s-1, no phototropism occurs and the mean protonemal tip angle remains above the horizontal, which is the criterion for negative gravitropism. But those protonemata are not as uniformly upright as they would be in the dark indicating that low intensity red light permits gravitropism but also modulates the response. Protonemata of the aphototropic mutant ptr1 that lacks a functional Pfr chromophore, exhibit gravitropism regardless of red light intensity. This indicates that red light acts via Pfr to modulate gravitropism at low intensities and to suppress gravitropism at intensities < or = 140nmol m-2 s-1.     

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.