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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↗

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↗

Choosing plants to be grown in a Controlled Environment Life Support System (CELSS) based upon attractive vegetarian diets.

Space explorers on the Moon, Mars, or even in a space craft might grow plants in a CELSS to remove CO2 and provide O2 and food. Selection of crops to be studied has been rather arbitrary but should be based on plants that can provide a balanced and attractive, mostly vegetarian diet. Additional selection criteria include ease of growth in artificial environments and sufficient variety provided over long intervals. This article is based on a workshop convened to study vegetarian diets for use in a CELSS. Participants included nutritional scientists, practicing vegetarians, and interested employees of the Johnson Space Center. It was concluded that diets meeting the criteria could be formulated, and a list of suitable crops was compiled.

Crops, Agricultural↗

Application of photosynthetic N2-fixing cyanobacteria to the CELSS program.

The feasibility of using photosynthetic microalgae (cyanobacteria) as a subsystem component for the CELSS program, with particular emphasis on the manipulation of the biomass (protein/carbohydrate) has been addressed. Using factors which retard growth rates, but not photosynthetic electron flux, the partitioning of photosynthetically derived reductant may be dictated towards CO2 fixation (carbohydrate formation) and away from N2 fixation (protein formation). Cold shock treatment of fairly dense cultures markedly increases the glycogen content from 1% to 35% (dry weight), and presents a useful technique to change the protein/carbohydrate ratio of these organisms to a more nutritionally acceptable form.

Biomass↗

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↗

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↗

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↗

Some challenges in designing a lunar, Martian, or microgravity CELSS.

The design of a bioregenerative life-support system (a Controlled Ecological Life-Support System or CELSS) for long-duration stays on the moon, Mars, or in a space craft poses formidable problems in engineering and in theory. Technological (hardware) problems include: (1) Creation and control of gas composition and pressure, temperature, light, humidity, and air circulation, especially in microgravity to 1/3 xg and in the vacuum of space. Light (energy demanding), CO2 levels, and the rooting media are special problems for plants. (2) Developing specialized equipment for food preparation. (3) Equipment development for waste recycling. (4) Development of computer systems for environmental monitoring and control as well as several other functions. Problems of theory (software) include: (1) Determining crop species and cultivars (some bred especially for CELSS). (2) Optimum environments and growing and harvesting techniques for each crop. (3) Best and most efficient food-preparation techniques and required equipment. (4) Best and most efficient waste-recycling techniques and equipment. This topic includes questions about the extent of closure, resupply, and waste storage. (5) How to achieve long-term stability. (6) How to avoid catastrophic failures--and how to recover from near-catastrophic failures (for example, plant diseases). Many problems must be solved.

Biomass↗

Growing root, tuber and nut crops hydroponically for CELSS.

Among the crops selected by the National Aeronautics and Space Administration for growth in controlled ecological life support systems are four that have subsurface edible parts -- potatoes, sweet potatoes, sugar beets and peanuts. These crops have been produced in open and closed (recirculating), solid media and liquid, hydroponic systems. Fluorescent , fluorescent plus incandescent and high pressure sodium plus metal halide lamps have proven to be effective light sources. Continuous light with 16 degrees C and 28/22 degrees C (day/night) temperatures have produced highest yields for potato and sweet potato, respectively. Dry weight yields of up to 4685, 2541, 1151 and 207 g m-2 for for potatoes, sweet potatoes, sugar beets and peanuts, respectively, have been produced in controlled environment hydroponic systems.

Arachis↗

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↗

Biomass recycle as a means to improve the energy efficiency of CELSS algal culture systems.

Algal cultures can be very rapid and efficient means to generate biomass and regenerate the atmosphere for closed environmental life support systems. However, as in the case of most higher plants, a significant fraction of the biomass produced by most algae cannot be directly converted to a useful food product by standard food technology procedures. This waste biomass will serve as an energy drain on the overall system unless it can be efficiently recycled without a significant loss of its energy content. We report experiments in which cultures of the algae Scenedesmus obliquus were grown in the light and at the expense of an added carbon source, which either replaced or supplemented the actinic light. As part of these experiments we tested hydrolyzed waste biomass from these same algae to determine whether the algae themselves could be made part of the biological recycling process. Results indicate that hydrolyzed algal (and plant) biomass can serve as carbon and energy sources for the growth of these algae, suggesting that the efficiency of the closed system could be significantly improved using this recycling process.

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↗

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).

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↗

Isatin as an auxin source favoring floral and vegetative shoot regeneration from calli produced by thin layer explants of tomato pedicel.

Thin layer explants taken from the pedicels and peduncles of flowering tomato plants yielded calli with great organogenetic potential. Of the 15 cultivars tested, 7 regenerated roots, shoots and eventually entire fruit-bearing plants. Calli grown on modified Murashige-Skoog medium responded to varied auxins and cytokinins with different morphogenetic patterns. Thus, naphthaleneacetic acid yielded root-producing calli, while the auxin precursor isatin (indole 2,3-dione) caused the production of calli with vegetative and floral shoots, rarely yielding roots. This may be related to isatin's slow, steady conversion to an active auxin (Plant Physiol 41:1485-1488, 1966) in contrast with naphthaleneacetic acid's immediate presentation of a high level of active auxin. The highest incidence of vegetative shoot (100%) and flower (50%) formation was obtained with 10 micromoles isatin and 3 micromoles zeatin. A few of the flowers developed into ripe fruits. The high frequency of induction of vegetative shoots and flowers before roots with isatin suggests its utility in micropropagation from plant tissue cultures.

Culture Media↗

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. IV. Effect of CO2 enrichment.

Potatoes (Solanum tuberosum L.) cvs. Norland and Russet Burbank were grown in solid stands in separate controlled environment rooms at two CO2 levels, 365 micromoles mol-1 (ppm) and 1000 micromoles mol-1. Rooms were maintained under continuous fluorescent light (450 micromoles s-1 m-2 PPF), 16 C and 70% relative humidity. Norland plants were grown for 110 days and Russet Burbank plants for 126 days. CO2 assimilation rates (net photosynthetic rates) of exposed, upper canopy leaves were measured at weekly intervals beginning at 21-days-age for Norland and 28-days-age for Russet Burbank. Elevation of CO2 increased CO2 assimilation rates of Norland leaves by approximately 24%, but decreased rates of Russet Burbank leaves by approximately 12%. Assimilation rates of Norland leaves under the high CO2 decreased as plants matured so that their rates were similar to rates under the low CO2 levels after 70-days-age. Assimilation rates of Russet Burbank leaves under high CO2 remained depressed in comparison to low CO2 plants throughout the period of measurements. Yield data showed only marginal benefits from CO2 enrichment: tuber dry weight increased 2% for Norland and 12% for Russet Burbank, total plant dry weight was increased 6% for Norland and 4% for Russet Burbank. The best productivity obtained in this study (21.9 g tuber dry wt m-2 day-1 from Norland at 1000 micromoles mol-1 of CO2) indicates that the dietary energy needs of one human in space could be supplied from 34 m2 of potatoes.

Carbon Dioxide↗

Utilization of potatoes for life support in space. V. Evaluation of cultivars in response to continuous light and high temperature.

Twenty-four potato (Solanum tuberosum L.) cultivars from different regions of the world were evaluated in terms of their responses to continuous light (24 h photoperiod) and to high temperature (30 C) in two separate experiments under controlled environments. In each experiment, a first evaluation of the cultivars was made at day 35 after transplanting, at which time 12 cultivars exhibiting best growth and tuber initiation were selected. A final evaluation of the 12 cultivars was made after an additional 21 days of growth, at which time plant height, total dry weight, tuber dry weight, and tuber number were determined. In the continuous light evaluation, the 12 selected cultivars were Alaska 114, Atlantic, Bintje, Denali, Desiree, Haig, New York 81, Ottar, Rutt, Snogg, Snowchip, and Troll. In the high temperature evaluation, the 12 selected cultivars were Alpha, Atlantic, Bake King, Denali, Desiree, Haig, Kennebec, Norland, Russet Burbank, Rutt, Superior, and Troll. Among the cultivars selected under continuous irradiation, Desiree, Ottar, Haig, Rutt, Denali and Alaska showed the best potential for high productivity whereas New York 81 and Bintje showed the least production capability. Among the cultivars selected under high temperature, Rutt, Haig, Troll and Bake King had best performance whereas Atlantic, Alpha, Kennebec and Russet Burbank exhibited the least production potential. Thus, Haig and Rutt were the two cultivars that performed well under continuous irradiation and high temperature conditions, and could have maximum potential for adaptation to varying stress environments. These two cultivars may have the best potential for use in future space farming in which continuous light and/or high temperature conditions may exist. However, cultivar responses under combined conditions of continuous light and high temperature remains for further validation.

Environment, Controlled↗