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Steady-state canopy gas exchange: system design and operation.

This paper describes the use of a commercial growth chamber for canopy photosynthesis, respiration, and transpiration measurements. The system was designed to measure transpiration via water vapor fluxes, and the importance of this measurement is discussed. Procedures for continuous measurement of root-zone respiration are described, and new data is presented to dispel myths about sources of water vapor interference in photosynthesis and in the measurement of CO2 by infrared gas analysis. Mitchell (1992) has described the fundamentals of various approaches to measuring photosynthesis. Because our system evolved from experience with other types of single-leaf and canopy gas-exchange systems, it is useful to review advantages and disadvantages of different systems as they apply to various research objectives.

Air↗

A role for the diazotrophic cyanobacterium, Cyanothece sp. strain ATCC 51142, in nitrogen cycling for CELSS applications.

Simple calculations show that fixed nitrogen regeneration in a CELSS may not be as efficient as stowage and resupply of fixed nitrogen compounds. However, fixed nitrogen regeneration may be important for the sustainability and safety of a deployed CELSS. Cyanothece sp. strain ATCC 51142, a unicellular, aerobic, diazotrophic cyanobacterium, with high growth rates and a robust metabolism, is a reasonable candidate organism for a biological, fixed nitrogen regeneration system. In addition, Cyanothece sp. cultures may be used to balance gas exchange ratio imparities between plants and humans. The regeneration of fixed nitrogen compounds by cyanobacterial cultures was examined in the context of a broad computer model/simulation (called CELSS-3D). When cyanothece sp. cultures were used to balance gas exchange imparities, the biomass harvested could supply as much as half of the total fixed nitrogen needed for plant biomass production.

Biomass↗

Non-conventional approaches to food processing in CELSS. I--Algal proteins; characterization and process optimization.

Protein isolate obtained from green algae (Scenedesmus obliquus) cultivated under controlled conditions was characterized. Molecular weight determination of fractionated algal proteins using SDS-polyacrylamide gel electrophoresis revealed a wide spectrum of molecular weights ranging from 15,000 to 220,000. Isoelectric points of dissociated proteins were in the range of 3.95 to 6.20. Amino acid composition of protein isolate compared favorably with FAO standards. High content of essential amino acids leucine, valine, phenylalanine and lysine makes algal protein isolate a high quality component of CELSS diets. To optimize the removal of algal lipids and pigments supercritical carbon dioxide extraction (with and without ethanol as a co-solvent) was used. Addition of ethanol to supercritical CO2 resulted in more efficient removal of algal lipids and produced protein isolate with a good yield and protein recovery. The protein isolate extracted by the above mixture had an improved water solubility.

Amino Acids↗

Suggestions for crops grown in controlled ecological life-support systems, based on attractive vegetarian diets.

Assuming that crops grown in controlled ecological life-support systems (CELSS) should provide a basis for meals that are both nutritious and attractive (to taste and vision), and that CELSS diets on the moon or Mars or in space-craft during long voyages will have to be mostly vegetarian, a workshop was convened at the Johnson Space Center, Houston, Texas, U.S.A. on 19 to 21 January, 1994. Participants consisted of trained nutritionists and others; many of the approximately 18 presenters who discussed possible diets were practicing vegetarians, some for more than two decades. Considering all the presentations, seven conclusions (or points for discussion) could be formulated: nutritious vegetarian diets are relatively easily to formulate, vegetarian diets are healthy, variety is essential in vegetarian diets, some experiences (e.g., Bios-3 and Biosphere 2) are relevant to planning of CELSS diets, physical constraints will limit the choice of crops, a preliminary list of recommended crops can be formulated, and this line of research has some potential practical spinoffs. The list of crops and the reasons for including specific crops might be of interest to professionals in the field of health and nutrition as well as to those who are designing closed ecological systems.

Crops, Agricultural↗

Evaluation of Cyanothece sp. ATCC 51142 as a candidate for inclusion in a CELSS.

Controlled ecological life support systems (CELSS) have been proposed to make long-duration manned space flights more cost-effective. Higher plants will presumably provide food and a breathable atmosphere for the crew. It has been suggested that imbalances between the CO2/O2 gas exchange ratios of the heterotrophic and autotrophic components of the system will inevitably lead to an unstable system, and the loss of O2 from the atmosphere. Ratio imbalances may be corrected by including a second autotroph with an appropriate CO2/O2 gas exchange ratio. Cyanothece sp. ATCC 51142 is a large unicellular N2-fixing cyanobacterium, exhibiting high growth rates under diverse physiological conditions. A rat-feeding study showed the biomass to be edible. Furthermore, it may have a CO2/O2 gas exchange ratio that theoretically can compensate for ratio imbalances. It is suggested that Cyanothece spp. could fulfill several roles in a CELSS: supplementing atmosphere recycling, generating fixed N from the air, providing a balanced protein supplement, and protecting a CELSS in case of catastrophic crop failure.

Air Conditioning↗

Morphological responses of wheat to blue light.

Blue light significantly increased tillering in wheat (Triticum aestivum L.) plants grown at the same photosynthetic photon flux (PPF). Plants were grown under two levels of blue light (400-500 nm) in a controlled environment with continuous irradiation. Plants received either 50 micromoles m-2 s-1 of blue light or 2 micromoles m-2 s-1 blue light from filtered metal halide lamps at a total irradiance of 200 micromoles m-2 s-1 PPF (400-700 nm). Plants tillered an average of 25% more under the higher level of blue light. Blue light also caused a small, but consistent, increase in main culm development, measured as Haun stage. Leaf length was reduced by higher levels of blue light, while plant dry-mass was not significantly affected by blue light. Applying the principle of equivalent light action, the results suggest that tillering and leaf elongation are mediated by the blue-UV light receptor(s) because phytochrome photoequilibrium for each treatment were nearly identical.

Biomass↗

Utilization of potatoes for life support systems. II. The effects of temperature under 24-h and 12-h photoperiods.

The growth and tuberization of Norland potatoes were studied under five different temperatures and two photoperiods. Treatment levels included 12, 16, 20, 24, and 28 C with either a 24-h (continuous light) or a 12-h photoperiod at 400 micromoles m-2 s-1 PPF. Plants were grown in 6-liter containers and harvested at 56-days-age. Stem length increased with increasing temperature under both photoperiods. The highest tuber yield occurred at 16 C under the 24-h photoperiod (755 g/plant) and at 20 C under the 12-h photoperiod (460 g/plant). Little or no tuber formation occurred at 28 C under either photoperiod or at 24 C under continuous light. As with tuber yield, the highest total plant dry weights also occurred at 16 C under the 24-h photoperiod and at 20 C under the 12-h photoperiod. Harvest index (tuber dry weight to total dry weight ratio) decreased with increasing temperatures and with continuous light. Results indicate that good growth and tuberization can occur under continuous light, and that increasing the photoperiod form 12 to 24 h effectively decreased the optimal temperature for tuber formation from near 20 C to 16 C. Alternatively, the results imply that a cooler temperatures, the potato becomes less obligate for dark period stimulation of tuberization.

Biomass↗

Utilization of potatoes for life support systems in space: III. Productivity at successive harvest dates under 12-h and 24-h photoperiods.

Potatoes are among several crops under consideration for use in controlled ecological life support systems (CELSS) being proposed for space colonies. Efficient crop production for such life support systems will require near-optimal growing conditions with harvests taken when production per unit area per unit time is maximum. To determine this maximum for potato, cv. Norland plants were grown in walk-in growth rooms under 12-h and 24-h photoperiods at 16 C and harvested at 42, 63, 84, 105, 126 and 148 days from planting. At 42 days, plants were encaged in wire fence cylinders with a cross-sectional area of 0.2 m2. The dry weights (dwt) of tubers and of the entire plants increased under both photoperiods until the final harvest date (148 days), reaching 572 g tuber dwt and 704 g total dwt under 12-h, and 791 g tuber dwt and 972 g total dwt under 24-h. At a spacing of 0.2 m2 per plant, the 148-day tuber production from plants under continuous light would equate to nearly 40 t ha-1 dry matter (200 t fresh weight), approximately twice that of exceptionally high field yields. Tuber productivity (g m-2 day-1) under the 24-h photoperiod reached a maximum of 29.4 g dwt m-2 day-1 at 126 days, but continued to rise throughout the experiment under the 12-h photoperiod, reaching 19.5 g dwt m-2 day-1 at 14 days, approximately 25 m2 would continuously provide the daily dietary energy requirements for one human.

Biomass↗

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.

Biomass↗

Quantification and characterization of volatiles evolved during extrusion of rice and soy flours.

NASA-Johnson Space Center is designing and building a habitat (Bioregenerative Planetary Life Support Systems Test Complex, BIO-Plex) intended for evaluating advanced life support systems developed for long-duration missions to the Moon or Mars where all consumables will be recycled and reused. A food system based on raw products obtained from higher plants (such as soybeans, rice, and wheat) may be a central feature of a biologically based Advanced Life Support System. To convert raw crops to edible ingredients or food items, multipurpose processing equipment such as an extruder is ideal. Volatile compounds evolved during the manufacturing of these food products may accumulate and reach toxic levels. Additionally, off-odors often dissipated in open-air environments without consequence may cause significant discomfort in the BIO-Plex. Rice and defatted soy flours were adjusted to 16% moisture, and triplicate samples were extruded using a tabletop single-screw extruder. The extrudate was collected in specially designed Tedlar bags from which air samples could be extracted. The samples were analyzed by GC-MS with special emphasis on compounds with Spacecraft Maximum Allowable Concentrations (SMACs). Results showed a combination of alcohols, aldehydes, ketones, and carbonyl compounds in the different flours. Each compound and its SMAC value, as well as its impact on the air revitalization system, was discussed.

Acetaldehyde↗

Growth, carbon dioxide exchange and mineral accumulation in potatoes grown at different magnesium concentrations.

Plants of Norland potatoes (Solanum tuberosum L.) were maintained for 42 days at Mg concentrations of 0.05, 0.125, 0.25, 1, 2, and 4 mM in a nonrecirculating nutrient film system under controlled environment. With the increased Mg supply from 0.05 to 4 mM, Mg concentrations in the leaves of the 42-day old plants increased significantly from 1.1 to 11.2 mg g-1 dry weight. Plant leaf area and plant and tuber dry weights increased with increased Mg concentrations up to 1 mM in solution or 6.7 mg g-1 in leaves, and then decreased with further increases in Mg concentrations. Rates of CO2 assimilation measured on leaflets in situ at ambient and various intercellular CO2 concentrations were consistently lower at 0.05 and 4 mM Mg than at other Mg treatments, which may indicate decreased photosynthetic activity in mesophyll tissues at the lowest and highest Mg concentrations. Dark respiration rates in leaves were highest at 0.05 and 4 mM Mg, lowest at 0.25 and 1 mM Mg, and intermediate at 0.125 and 2 mM Mg. The different Mg treatments also influenced accumulation of other minerals in leaves. Leaf concentrations of Ca and Mn decreased with increased Mg supply except that Ca and Mn were lower at 0.05 mM than at 0.125 mM Mg. Leaf K concentrations were lower at 1, 2 and 4 mM Mg than at other Mg treatments. Foliar concentrations of P, Fe, Zn, and Cu had small but inconsistent variation with different Mg concentrations. Leaf concentrations of N, S, and B were similar at different Mg concentrations. This study demonstrates that various Mg nutrition, along with altered accumulation of other nutrients, could regulate dry matter production in potatoes by affecting not only leaf area but also leaf carbon dioxide assimilation and respiration.

Carbon Dioxide↗

Varying plant density and harvest time to optimize cowpea leaf yield and nutrient content.

Plant density and harvest time were manipulated to optimize vegetative (foliar) productivity of cowpea [Vigna unguiculata (L.) Walp.] canopies for future dietary use in controlled ecological life-support systems as vegetables or salad greens. Productivity was measured as total shoot and edible dry weights (DW), edible yield rate [(EYR) grams DW per square meter per day], shoot harvest index [(SHI) grams DW per edible gram DW total shoot], and yield-efficiency rate [(YER) grams DW edible per square meter per day per grams DW nonedible]. Cowpeas were grown in a greenhouse for leaf-only harvest at 14, 28, 42, 56, 84, or 99 plants/m2 and were harvested 20, 30, 40, or 50 days after planting (DAP). Shoot and edible dry weights increased as plant density and time to harvest increased. A maximum of 1189 g shoot DW/m2 and 594 g edible DW/m2 were achieved at an estimated plant density of 85 plants/m2 and harvest 50 DAP. EYR also increased as plant density and time to harvest increased. An EYR of 11 g m-2 day-1 was predicted to occur at 86 plants/m2 and harvest 50 DAP. SHI and YER were not affected by plant density. However, the highest values of SHI (64%) and YER (1.3 g m-2 day-1 g-1) were attained when cowpeas were harvested 20 DAP. The average fat and ash contents [dry-weight basis (dwb)] of harvested leaves remained constant regardless of harvest time. Average protein content increased from 25% DW at 30 DAP to 45% DW at 50 DAP. Carbohydrate content declined from 50% DW at 30 DAP to 45% DW at 50 DAP. Total dietary fiber content (dwb) of the leaves increased from 19% to 26% as time to harvest increased from 20 to 50 days.

Biomass↗

Compositional and toxicological evaluation of the diazotrophic cyanobacterium, Cyanothece sp. strain ATCC 51142.

Compositional analyses of Cyanothece sp. strain ATCC 51142 showed high protein (50-60%) and low fat (0.4-1%) content, and the ability to synthesize vitamin B12. The amino acid profile indicated that Cyanothece sp. was a balanced protein source. Fatty acids of the 18:3n-3 type were also present. Mineral analyses indicated that the cellular biomass may be a good source of Fe, Zn and Na. Caloric content was 4.5 to 5.1 kcal g dry weight-1 and the carbon content was approximately 40% on a dry weight basis. Nitrogen content was 8 to 9% on a dry weight basis and total nucleic acids were 1.3% on a dry weight basis. Short-term feeding studies in rats followed by histopathology found no toxicity or dietary incompatibility problems. The level of uric acid and allantoin in urine and tissues was low, suggesting no excess of nucleic acids, as sometimes reported in the past for a cyanobacteria-containing diet. The current work discusses the potential implications of these results for human nutrition applications.

Amino Acids↗

Determining the potential productivity of food crops in controlled environments.

The quest to determine the maximum potential productivity of food crops is greatly benefitted by crop growth models. Many models have been developed to analyze and predict crop growth in the field, but it is difficult to predict biological responses to stress conditions. Crop growth models for the optimal environments of a Controlled Environment Life Support System (CELSS) can be highly predictive. This paper discusses the application of a crop growth model to CELSS; the model is used to evaluate factors limiting growth. The model separately evaluates 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 determine potentially achievable productivity. An analysis of each process suggests that low harvest index is the factor most limiting to yield. PPF absorption by plant canopies and respiration efficiency are also of major importance. Research concerning productivity in a CELSS should emphasize: 1) the development of gas exchange techniques to continuously monitor plant growth rates and 2) environmental techniques to reduce plant height in communities.

Biomass↗

Modeling the effects of light, carbon dioxide, and temperature on the growth of potato.

This study examined the effects of light, temperature and carbon dioxide on the growth of potato (Solanum tuberosum L.) in a controlled environment in order to ascertain the best growing conditions for potato in life support systems in space. 'Norland' and 'Russet Burbank' were grown in 6-L pots of peat-vermiculite for 56 d in growth chambers at the University of Wisconsin Biotron. Environmental factor levels included continuous light (24-h photoperiod) at 250, 400, and 550 micromoles m-2 s-1 PPF; constant temperature at 16, 20, and 24 degrees C; and CO2 at approximately 400, 1000, and 1600 microliters L-1. Separate effects analysis and ridge analysis provided a means to examine the effects of individual environmental factors and to determine combinations of factors that are expected to give the best increases in yields over the central design point. The response surface of Norland indicated that tuber yields were highest with moderately low temperature (18.7 degrees C), low CO2 (400 microliters L-1) and high light (550 micromoles m-2 s-1 PPF). These conditions also favored shorter stem growth. Russet Burbank tuber yields were highest at moderately low temperature (17.5 degrees C), high CO2 (1600 microliters L-1) and medium analyses will be used to project the most efficient conditions for growth of potatoes in closed ecological life support systems (CELSS) in space colonies.

Carbon Dioxide↗

CELSS for advanced manned mission.

An overview of the major concepts of Controlled Ecological Life Support System (CELSS) includes an identification of environmental factors, such as gravity levels, light levels, and growth volume, that influence the type of CELSS system that can be developed. Various plant growth systems are described together with their possible space applications. Life support functions performed by plants include food production, atmosphere regeneration, and water purification. Selected relationships between biological and physical-chemical life support techniques are considered as a part of these functions. Consumers in a CELSS may be humans, animals, or microorganisms, but nutritional, water, and atmosphere requirements of humans are emphasized in this report, as they are the primary requirement drivers for a CELSS design. The human role in waste generation is discussed as it affects plant nutrient availability. The role of waste management systems in recovering nutrients for plant growth and requirements for CELSS are defined for air, water, and food. Both physical and a biological nutrient recovery/waste disposal systems are examined. The separate subsystems of a CELSS are identified and discussed. Nutrient recovery, plant irradiation, automation, and facilities equipment and applications are reviewed with special attention to direct solar irradiation using fiber optics. These subsystems, along with other environmental control systems, such as thermal, humidity, and ventilation, are essential to plant growth in the space environment.

Crops, Agricultural↗

The components of crop productivity: measuring and modeling plant metabolism.

Several investigators in the CELSS program have demonstrated that crop plants can be remarkably productive in optimal environments where plants are limited only by incident radiation. Radiation use efficiencies of 0.4 to 0.7 g biomass per mol of incident photons have been measured for crops in several laboratories. Some early published values for radiation use efficiency (1 g mol-1) were inflated due to the effect of side lighting. Sealed chambers are the basic research module for crop studies for space. Such chambers allow the measurement of radiation and CO2 fluxes, thus providing values for three determinants of plant growth: radiation absorption, photosynthetic efficiency (quantum yield), and respiration efficiency (carbon use efficiency). Continuous measurement of each of these parameters over the plant life cycle has provided a blueprint for daily growth rates, and is the basis for modeling crop productivity based on component metabolic processes. Much of what has been interpreted as low photosynthetic efficiency is really the result of reduced leaf expansion and poor radiation absorption. Measurements and models of short-term (minutes to hours) and long-term (days to weeks) plant metabolic rates have enormously improved our understanding of plant environment interactions in ground-based growth chambers and are critical to understanding plant responses to the space environment.

Biomass↗

Plant productivity in controlled environments.

To assess the cost and area/volume requirements of a farm in a space station or Lunar or Martian base, a few laboratories in the United States, the Soviet Union, France, and Japan are studying optimum controlled environments for the production of selected crops. Temperature, light, photoperiod, CO2, humidity, the root-zone environment, and cultivars are the primary factors being manipulated to increase yields and harvest index. Our best wheat yields on a time basis (24 g m-2 day-1 of edible biomass) are five times good field yields and twice the world record. Similar yields have been obtained in other laboratories with potatoes and lettuce; soybeans are also promising. These figures suggest that approximately 30 m2 under continuous production could support an astronaut with sufficient protein and about 2800 kcal day-1. Scientists under Iosif Gitelzon in Krasnoyarsk, Siberia, have lived in a closed system for up to 5 months, producing 80% of their own food. Thirty square meters for crops were allotted to each of the two men taking part in the experiment. A functional controlled-environment life-support system (CELSS) will require the refined application of several disciplines: controlled-environment agriculture, food preparation, waste disposal, and control-systems technology, to list only the broadest categories. It has seemed intuitively evident that ways could be found to prepare food, regenerate plant nutrients from wastes, and even control and integrate several subsystems of a CELSS. But could sufficient food be produced in the limited areas and with the limited energy that might be available? Clearly, detailed studies of food production were necessary.

Biomass↗