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.     

Developing a vitamin greenhouse for the life support system of the International Space Station and for future interplanetary missions.

In order to evaluate the effects of gravity on growing plants, we conducted ground based long-term experiments with dwarf wheat, cultivar Apogee and Chinese cabbage, cultivar Khibinskaya. The test crops had been grown in overhead position with HPS lamp below root module so gravity and light intensity gradients had been in opposite direction. Plants of the control crop grew in normal position under the same lamp. Both crops were grown on porous metallic membranes with stable -1 kPa matric potential on their surface. Results from these and other studies allowed us to examine the differences in growth and development of the plants as well as the root systems in relation to the value of the gravity force influence. Dry weight of the roots from test group was decreased in 2.5 times for wheat and in 6 times - at the Chinese cabbage, but shoot dry biomass was practically same for both test and control versions. A harvest index of the test plants increased substantially. The data shows, that development of the plants was essentially changed in microgravity. The experiments in the space greenhouse Svet aboard the Mir space station proved that it is possible to compensate the effects of weightlessness on higher plants by manipulating gradients of environmental parameters (i.e. photon flux, matric potential in the root zone, etc.). However, the average productivity of Svet concerning salad crops even in ground studies did not provide more than 14 g fresh biomass per day. This does not provide a sufficient level of supplemental nutrients to the crew of the ISS. A cylindrical design of a space plant growth chamber (SPGC) allows for maximal productivity in presence of very tight energy and volume limitations onboard the ISS and provides a number of operational advantages. Productivity from this type of SPGF with a 0.5 kW energy utilization when salad growing would provide approximately 100 g of edible biomass per day, which would almost satisfy requirements for a crew of two in vitamin C and carotene and partly vitamin B group as well as rough fiber.     

The Spacelab-Mir-1 "Greenhouse-2" experiment.

The Spacelab-Mir-1 (SLM-1) mission is the first docking of the Space Shuttle Atlantis (STS-71) with the Orbital Station Mir in June 1995. The SLM-1 "Greenhouse-2" experiment will utilize the Russian-Bulgarian-developed plant growth unit (Svet). "Greenhouse-2" will include two plantings (1) designed to test the capability of Svet to grow a crop of Superdwarf wheat from seed to seed, and (2) to provide green plant material for post-flight analysis. Protocols, procedures, and equipment for the experiment have been developed by the US-Russian science team. "Greenhouse-2" will also provide the first orbital test of a new Svet Instrumentation System (SIS) developed by Utah State University to provide near real time data on plant environmental parameters and gas-exchange rates. SIS supplements the Svet control and monitoring system with additional sensors for substrate moisture, air temperature, IR leaf temperature, light, oxygen, pressure, humidity, and carbon-dioxide. SIS provides the capability to monitor canopy transpiration and net assimilation of the plants growing in each vegetation unit (root zone) by enclosing the canopy in separate, retractable, ventilated leaf chambers. Six times during the seed-to-seed experiment, plant samples will be collected, leaf area measured, and plant parts fixed and/or dried for ground analysis. A second planting initiated 30 days before the arrival of a U.S. Shuttle [originally planned to be STS-71] is designed to provide green material at the vegetative development stage for ground analysis. [As this paper is being edited, the experiment has been delayed until after the arrival of STS-71.]     

Induced abnormality in Mir- and Earth grown Super Dwarf wheat.

Super-dwarf wheat grown on the Mir space station using the Svet "Greenhouse" exhibited morphological, metabolic and reproductive abnormalities compared with Earth-grown wheat. Of prominent importance were the abnormalities associated with reproductive ontogeny and the total absence of seed formation on Mir. Changes in the apical meristem associated with transition from the vegetative phase to floral initiation and development of the reproductive spike were all typical of 'Super-Dwarf' wheat up to the point of anthesis. Observation of ruptured anthers from the Mir-grown plants revealed what appeared to be normally developed pollen. These pollen gains, however, contained only one nuclei, while normal viable pollen is tri-nucleate. A potentially important difference in the flight experiment, compared with ground reference studies, was the presence of a high level of atmospheric ethylene (1,200 ppb). Ground studies conducted by exposing 'Super-Dwarf' wheat to ethylene just prior to anthesis resulted in manifestation of the same abnormalities observed in the space flight samples.     

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.     

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.     

"From seed-to-seed" experiment with wheat plants under space-flight conditions.

An important goal with plant experiments in microgravity is to achieve a complete life cycle, the "seed-to-seed experiment." Some Soviet attempts to reach this goal are described, notably an experiment with the tiny mustard, Arabidopsis thaliana, in the Phyton 3 device on Salyut 7. Normal seeds were produced although yields were reduced and development was delayed. Several other experiments have shown abnormalities in plants grown in space. In recent work, plants of wheat (Triticum aestivum) were studied on the ground and then in a preliminary experiment in space. Biometric indices of vegetative space plants were 2 to 2.5 times lower than those of controls, levels of chlorophyll a and b were reduced (no change in the ratio of the two pigments), carotenoids were reduced, there was a serious imbalance in major minerals, and membrane lipids were reduced (no obvious change in lipid patterns). Following the preliminary studies, an attempt was made with the Svetoblock-M growth unit to grow a super-dwarf wheat cultivar through a life cycle. The experiment lasted 167 d on Mir. Growth halted from about day 40 to day 100, when new shoots appeared. Three heads had appeared in the boot (surrounded by leaves) when plants were returned to earth. One head was sterile, but 28 seeds matured on earth, and most of these have since produced normal plants and seeds. In principle, a seed-to-seed experiment with wheat should be successful in microgravity.     

Spectral composition of light and plant productivity.

Among other problems the Institute of Biophysics is working on the development of physiological and fundamental aspects of intensive light cultivation of higher plants. These technologies can be used in life support systems for stationary space station such as a Lunar base, a planetary base or a large orbital station. The source of energy may be the Sun or a nuclear reactor. In certain conditions, such sources of energy allow the use of a very broad range of irradiance of plants, in particular in the light energy range up to 2-3 times the solar energy (up to 100-1200 W/m2 PAR). Our Institute was the first to show that under such a high irradiance, some plants (radish, wheat, for example) can actively photosynthesize and exhibit high productivity on a sowing area basis. These results were later confirmed in the laboratory of Prof. Salisbury (USA).     

Canopy photosynthesis and transpiration in microgravity: gas exchange measurements aboard Mir.

The SVET Greenhouse on-board the Orbital Station Mir was used to measure canopy photosynthesis and transpiration rates for the first time in space. During the Greenhouse IIB experiment on Mir (June-January 1997), carbon and water vapor fluxes from two wheat (cv. Superdwarf) canopies were measured using the US developed Gas Exchange Measurement System (GEMS). Gas analyzers capable of resolving CO2 concentration differences of 5 micromoles mol-1 against a background of 0.9% CO2, are necessary to measure photosynthetic and respiratory rates on Mir. The ability of the GEMS gas analyzers to measure these CO2 concentration differences was determined during extensive ground calibrations. Similarly, the sensitivity of the analyzers to water vapor was sufficient to accurately measure canopy evapotranspiration. Evapotranspiration, which accounted for over 90% of the water added to the root zone, was estimated using gas exchange and used to estimate substrate moisture content. This paper presents canopy photosynthesis and transpiration data during the peak vegetative phase of development in microgravity.     

Lipid peroxidation of plants under microgravity and its simulation.

In series of space experiments aboard the biosatellites "Cosmos 1887", "Bion 9", the orbital stations "Salut", "Mir" and under clinostating, changes of lipid peroxidation (LPO) and antioxidation activity (AOA) of Chlorella, Haplopappus tissue culture, wheat and pea roots were determined. The changes had a complex fluctuation character; three steps of response were established: LPO decreasing accompanied by AOA increase; stabilization LPO <==> AOA balance; secondary LPO activation. Most early and highly amplitude decreasing of LPO were fixed in mitochondria. The rate of response have been increased on multicellular level of plants organization.     

Influence of nonuniform magnetic fields on orientation of plant seedlings in microgravity conditions.

Experiments on the spatial behavior of the flax (Linum usitatissimum, L.) seedlings in a nonuniform magnetic field were conducted on the orbital space stations "Salut" and "Mir". This field can displace sensory organelles (statoliths) inside receptor cells and such displacement should cause a physiological reaction of the plant-tropistic curvature. Experiments were conducted in the custom-built "Magnetogravistat" facility, where seeds were germinated and grown for 3-4 days in a magnetic field with the dynamic factor grad (H2/2) approximately equal to 10(7) Oe2/cm, then fixed on orbit and returned to Earth for analysis. It was found, that 93% of the seedlings were oriented in the field consistently with curvature in response to displacement of statoliths along the field gradient by ponderomotive magnetic forces, while control seedlings grew in the direction of the initial orientation of the seed. This suggests, that gravity receptors of plants recognized magnetic forces on statoliths as gravity, and that gravity stimulus can be substituted for plants by a force of a different physical nature.     

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).     

Excess nutrients in hydroponic solutions alter nutrient content of rice, wheat, and potato.

Environment has significant effects on the nutrient content of field-grown crop plants. Little is known, however, about compositional changes caused by controlled environments in which plants receive only artificial radiation and soilless, hydroponic culture. This knowledge is essential for developing a safe, nutritious diet in a Controlled Ecological Life-Support System (CELSS). Three crops that are candidates for inclusion in a CELSS (rice, wheat, and white potato) were grown both in the field and in controlled environments where the hydroponic nutrient solution, photosynthetic photon flux (PPF), and CO2 level were manipulated to achieve rapid growth rates. Plants were harvested at maturity, separated into discrete parts, and dried prior to analysis. Plant materials were analyzed for proximate composition (protein, fat, ash, and carbohydrate), total nitrogen (N), nitrate, minerals, and amino-acid composition. The effect of environment on nutrient content varied by crop and plant part. Total N and nonprotein N (NPN) contents of plant biomass generally increased under controlled-environment conditions compared to field conditions, especially for leafy plant parts and roots. Nitrate levels were increased in hydroponically-grown vegetative tissues, but nitrate was excluded from grains and tubers. Mineral content changes in plant tissue included increased phosphorus and decreased levels of certain micronutrient elements under controlled-environment conditions. These findings suggest that cultivar selection, genetic manipulation, and environmental control could be important to obtain highly nutritious biomass in a CELSS.     

Waste bioregeneration in life support CES: development of soil organic substrate.

An experimental model of matter turnover in the biotic cycle: plants (plant biomass) --> mushrooms (residual substrate + mushroom fruit bodies) --> worms (biohumus) --> microorganisms (soillike substrate) --> plants is presented. The initial mass of soillike substrate was produced from wheat plants grown in a hydroponic system. Three cycles of matter turnover in the biotic cycle were carried out. Grain productivity on soillike substrate was 21.87 g/m2 day. The results obtained were used for designing a CES containing man, plants, soillike substrate, bioregeneration module and aquaculture. It was shown, that the circulating dry mass of the CES is 756 kg. The main part (88%) of the circulating mass accumulates in the soillike substrate and bioregeneration module.     

Plant experiments with light-emitting diode module in Svet space greenhouse

Light is necessary for photosynthesis and shoot orientation in the space plant growth facilities. Light modules (LM) must provide sufficient photosynthetic photon flux for optimal efficiency of photosynthetic processes and also meet the constraints for power, volume and mass. A new LM for Svet space greenhouse using Cree® XLamp® 7090 XR light-emitting diodes (LEDs) was developed. Monochromic LEDs emitting in the red, green, and blue regions of the spectrum were used. The LED-LM contains 36 LED spots – 30 LED spots with one red, green and blue LED and 6 LED spots with three red LEDs. Digital Multiplex Control Unit controls the LED spots and can set 231 levels of light intensity thus achieving Photosynthetic Photon Flux Density (PPFD) in the range 0–400 μmol m⁻² s⁻¹ and different percentages of the red, green and blue light, depending on the experimental objectives. Two one-month experiments with plants – lettuce and radicchio were carried out at 400 μmol m⁻² s⁻¹ PPFD (high light – HL) and 220 μmol m⁻² s⁻¹ PPFD (low light – LL) and 70% red, 20% green and 10% blue light composition. To evaluate the efficiency of photosynthesis, in vivo modulated chlorophyll fluorescence was measured by Pulse Amplitude Modulation (PAM) fluorometer on leaf discs and the following parameters: effective quantum yield of Photosystem II (Φ_PSII) and non-photochemical quenching (NPQ) were calculated. Both lettuce and radicchio plants grown at LL express higher photochemical activity of Photosystem II (PSII) than HL grown plants, evaluated by Φ_PSII. Accelerated rise in NPQ in both LL grown plants was observed, while steady state NPQ values were higher in LL grown lettuce plants and did not differ in LL and HL grown radicchio plants. The extent of photoinhibition process in both plants was evaluated by changes in malonedialdehyde (MDA) concentration, peroxidase (POX) activity and hydrogen peroxide (H₂O₂) content. Accumulation of high levels of MDA and increased POX activity correlating with decreased H₂O₂ content were observed in both HL grown plants. These biochemical indicators revealed higher sensitivity to photodamage in HL grown lettuce and radicchio plants. LL conditions resulted in more effective functioning of PSII than HL when lettuce and radicchio plants were grown at 70% red, 20% green and 10% blue light composition.     

Capillary movement of liquid in granular beds in microgravity.

A more complete understanding of the dynamics of capillary flow through an unsaturated porous medium would be useful for the development of an effective water and nutrient delivery system for the growth of plants in space. An experiment was conducted on the Mir Space Station that used an experimental cuvette called "Capillary Test Bed" to compare fluid migration under terrestrial laboratory conditions by positioning the cuvette such that the hydrostatic force is negated and on Mir under microgravity conditions. Differences in fluid migration in the cuvette were observed with migration being slower in microgravity compared with some ground control experiments.     

Low light intensity effects on the growth,photosynthetic characteristics,antioxidant capacity,yield and quality of wheat (Triticum aestivum L.) at different growth stages in BLSS

Minimizing energy consumption and maximizing crop productivity are major challenges to growing plants in Bioregenerative Life Support System (BLSS) for future long-term space mission. As a primary source of energy, light is one of the most important environmental factors for plant growth. The purpose of this study is to investigate the effects of low light intensity at different stages on growth, pigment composition, photosynthetic efficiency, biological production and antioxidant defence systems of wheat (Triticum aestivum L.) cultivars during ontogenesis. Experiments were divided into 3 intensity-controlled stages according to growth period (a total of 65 days): seedling stage (first 20 days), heading and flowering stage (middle 30 days) and grain filling stage (last 15 days). Initial light condition of the control was 420 μmol m⁻² s⁻¹ and the light intensity increased with the growth of wheat plants. The light intensities of group I and II at the first stage and the last stage were adjusted to the half level of the control respectively. For group III, the first and the last stage were both adjusted to half level of the control. During the middle 30 days, all treatments were kept the same intensity. The results indicated that low-light treatment at seedling stage, biomass, nutritional contents, components of inedible biomass and healthy index (including peroxidase (POD) activity, malondialdehyde (MDA) and proline content) of wheat plants have no significant difference to the control. Furthermore, unit kilojoule yield of group I reached 0.591 × 10⁻³ g/kJ and induced the highest energy efficiency. However, low-light treatment at grain filling stage affected the final production significantly.     

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.     

A conceptual configuration of the lunar base bioregenerative life support system including soil-like substrate for growing plants

The paper presents a conceptual configuration of the lunar base bioregenerative life support system (LBLSS), including soil-like substrate (SLS) for growing plants. SLS makes it possible to combine the processes of plant growth and the utilization of plant waste. Plants are to be grown on SLS on the basis of 20 kg of dry SLS mass or 100 kg of wet SLS mass per square meter. The substrate is to be delivered to the base ready-made as part of the plant growth subsystem. Food for the crew was provided by prestored stock 24% and by plant growing system 76%. Total dry weight of the food is 631 g per day (2800 kcal/day) for one crew member (CM). The list of candidate plants to be grown under lunar BLSS conditions included 14 species: wheat, rice, soybean, peanuts, sweet pepper, carrots, tomatoes, coriander, cole, lettuce, radish, squash, onion and garlic. From the prestored stock the crew consumed canned fish, iodinated salt, sugar, beef sauce and seafood sauce. Our calculations show that to provide one CM with plant food requires the area of 47.5 m². The balance of substance is achieved by the removal dehydrated urine 59 g, feces 31 g, food waste 50 g, SLS 134 g, and also waters 86 g from system and introduction food 236 g, liquid potassium soap 4 g and mineral salts 120 g into system daily. To reduce system setup time the first plants could be sowed and germinated to a certain age on the Earth.     

Production characteristics of the “higher plants–soil-like substrate” system as an element of the bioregenerative life support system

The study addresses the possibility of long-duration operation of a higher plant conveyor, using a soil-like substrate (SLS) as the root zone. Chufa (Cyperus esculentus L.), radish (Raphanus sativus L.), and lettuce (Lactuca sativa L.) were used as study material. A chufa community consisting of 4 age groups and radish and lettuce communities consisting of 2 age groups were irrigated with a nutrient solution, which contained mineral elements extracted from the SLS. After each harvest, inedible biomass of the harvested plants and inedible biomasses of wheat and saltwort were added to the SLS. The amounts of the inedible biomasses of wheat and saltwort to be added to the SLS were determined based on the nitrogen content of the edible mass of harvested plants. CO₂ concentration in the growth chamber was maintained within the range of 1100–1700 ppm. The results of the study show that higher plants can be grown quite successfully using the proposed process of plant waste utilization in the SLS. The addition of chufa inedible biomass to the SLS resulted in species-specific inhibition of growth of both cultivated crops and microorganisms in the “higher plants – SLS” system. There were certain differences between the amounts of some mineral elements removed from the SLS with the harvested edible biomass and those added to it with the inedible biomasses of wheat and saltwort.