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Abstract
This paper describes a closed-loop control system for controlling the irradiance and spectral quality generated by fluorescent lamps in a controlled environment chamber. The 400 to 800 nm irradiance and the ratio of the red waveband (600 to 700 nm) to the far-red waveband (700 to 800 nm) were independently controlled and varied as functions of time. A suggested application is to investigate the possibility of synergistic effects of changing irradiance levels and changing spectral distributions on photoperiodism and photomorphogenesis.
Abstract
Many articles have appeared in recent years recommending or defining terminology and units for measuring radiation energy in experiments with plants. A large group of these publications (1, 7, 8, 9, 10, 11, 12, 19, 22, 23) present the units and nomenclature which have been agreed upon, and promoted by, the American Society of Agronomists (ASA), the American Society of Agricultural Engineers (ASAE), the American Society for Horticultural Science (ASHS), and the USDA North Central Regional Growth Chamber Use Committee (NCR-101). The most recent addition to this series has just been published in HortScience (10). The authors advocate adoption of their guidelines by researchers, reviewers, and editors on a worldwide basis. However, there is still both international and interdisciplinary disagreement on the terminology, and to a lesser extent, the units used for radiation measurements in the plant sciences. In many instances, these disagreements have been reduced to para-semantic arguments. On the other hand, there are many discrepancies in the literature which are due to inaccurate etymology, thereby leading to the potential danger that the grammatical roots of the nomenclature will be altered artificially. The purpose of this communication is to indicate these discrepancies before incorrect terminology becomes widely accepted.
Potato (Solanum tuberosum L. cvs. Norland and Denali) plants were grown under high-pressure sodium (HPS), metal halide (MH), and blue-light-enhanced SON-Agro high-pressure sodium (HPS-S) lamps to study the effects of lamp spectral quality on vegetative growth. All plants were initiated from in vitro nodal cultures and grown hydroponically for 35 days at 300 μmol·m–2·s–1 photosynthetic photon flux (PPF) with a 12-hour light/12-hour dark photoperiod and matching 20C/16C thermoperiod. `Denali' main stems and internodes were significantly longer under HPS compared to MH, while under HPS-S, lengths were intermediate relative to those under other lamp types, but not significantly different. `Norland' plants showed no significant differences in stem and internode length among lamp types. Total dry weight of `Denali' plants was unaffected by lamp type, but `Norland' plants grown with HPS had significantly higher dry weight than those under either HPS-S or MH. Spectroradiometer measurements from the various lamps verified the manufacturer's claims of a 30% increase in ultraviolet-blue (350 to 450 nm) output from the HPS-S relative to standard HPS lamps. However, the data from `Denali' suggest that at 300 μmol·m–2·s–1 total PPF, the increased blue from HPS-S lamps is still insufficient to consistently maintain short stem growth typical of blue-rich irradiance environments.
The vegetative growth of potato (Solanum tuberosum L.) cvs. Norland (NL) and Denali (DN) was investigated comparing SONAGRO high-pressure sodium (HPS-S), standard high-pressure sodium (HPS), and metal halide (MH) lamps. Plants were initiated from nodal culture and grown hydroponically in a reach-in growth chamber for 35 d with a 12-hr light/12-hr dark photoperiod and corresponding thermoperiod of 20/16 C. PPF for each treatment was maintained at 300 μmol m-2 s-1 and CO2 levels maintained at l000 μmol mol-1 to promote growth. Results showed that main stem length (SL) and number of internodes (INT) for DN were significantly higher under HPS compared to MH, while HPS-S was not significantly different from the other lamp types. Total dry weight (TDW) of NL plants was significantly higher for HPS than for either HPS-S and MH, however there was no significant difference in SL and INT among lamp types. The data suggest that the 12.6% increase in blue light (400-500 nm) with HPS-S in comparison to conventional HPS lamps may not be sufficient to consistently decrease the stem elongation effects commonly seen with plants grown under HPS.
Requirements for water, nutrient replenishment and acid (for pH control) were monitored for stands of wheat, soybean, potato, and lettuce grown in a recirculating hydroponic culture using a modified 1/2 Hoagland solution with NO3-N. Water use at full canopy cover for all crops ranged from 4 to 5 L m-2 day-1. When averaged over the entire crop cycle, nutrient stock solution (∼0.9 S m-1) use varied from 0.2 L m-2 day-1 (lettuce: to 0.75 L m-2 day-1 (wheat), while acid use varied from 6 mmol m-2 day-1 (lettuce and soybeans) to over 40 mmol m-2 day-1 (wheat). Water-per unit biomass was highest for soybean and lettuce (0.3 to 0.4 L g DW), and least for wheat and potato (0.15 L g DW). Nutrient replenishment per unit biomass was highest for lettuce, 34 mL g-1 DW, with other crops ranging from 21-26 mL g-1 DW. Acid requirements were highest for wheat, 1.2 mmol g-1 DW, and lowest for potato, 0.7 mmol g-1 DW. On a PAR basis, acid needs were highest for wheat, 0.6 mmol mol-1 photons, with all other crops near 0.4 mmol mol-1. Acid data suggest that wheat nutrient uptake favors anions more strongly than the other species, or that more nitrate loss (e.g., denitrification) may occur during wheat growth.
Critical levels of nutrients in leaf tissue are influenced by plant metabolism, environment, and nutrient availability. In this study, we measured the elemental concentrations in fully expanded, upper canopy potato (Solunum tuberosum cv. Norland) leaves throughout growth and development in a controlled environment. Plants were grown hydroponically (NFT) in NASA's Biomass Production Chamber using a complete nutrient solution with the electrical conductivity maintained continuously at 0.12 S m-1. Photoperiod and air and root zone temperatures were changed midseason to promote tuberization, while CO2 levels were maintained at 1000 μmol mol-1 throughout growth. During vegetative growth, leaf nutrient concentrations remained relatively constant, except for a decline in Ca. During tuber enlargement and plant maturation, overall nutrient uptake decreased. Concentrations of the less mobile nutrients such as Ca, Mg, and B increased in the leaf tissue during mature growth, but somewhat surprisingly, highly mobile K also increased. Leaf concentrations of P, Zn, and Cu decreased during maturation.
Soybean plants [Glycine max (L.) Merr. cv. McCall] were grown from seed to harvest (90 days) in NASA's Biomass Production Chamber. The chamber provides approximately 20 m2 of growing area with an atmospheric volume of 113 m3. Photosynthesis and respiration rates of the stand were tracked by monitoring CO2 increase during the 12-h dark period and the subsequent drawdown to controlled set point (1000 ppm) when the lamps were turned on each day. Stand photosynthesis [under 875 μmol m-2 s-1 photosynthetic photon flux (PPF)] peaked at 35 μmol m-2 s-1 at 30 to 35 days after planting (DAP) and averaged 22 μmol m-2 s-1 throughout the life cycle. Dark period respiration peaked near 8 μmol m-2 s-1 at 30 to 35 DAP and averaged nearly 5 μmol m-2 s-1 throughout the life cycle. Prior to full canopy closure near 30 DAP, the light compensation point (LCP) for stand photosynthesis was lass than 100 μmol m-2 s-1 PPF; by 54 DAP the LCP had increasad to 175 μmol m-2 s-1. Stand transpiration rates peaked at 8.2 L m-2 day-1 at 40 to 45 DAP and averaged 4.3 L m-2 day-1 throughout growth.
Lettuce (Lactuca sativa cv. Waldmann's Green) plants were grown in a large, tightly sealed chamber for NASA's Controlled Ecological Life Support Systems (CELSS) program. Plants were started by direct seeding and grown in 64 0.25-m2 trays (six plants per tray) using nutrient film technique. Environmental conditions included: 23°C, 75% relative humidity, 1000 ubar (ppm) CO2, a 16/8 photoperiod, and 300 umol m-2 s-1 PPF from metal halide lamps. Although the chamber was typically opened once each day for cultural activities, atmospheric ethylene levels (measured with GC/PID) increased from near 15 ppb at 23 days after planting (DAP) to 47 ppb at 28 DAP. At harvest (28 DAP), heads averaged 129 g FW or 6.8 g DW per plant, and roots averaged 0.6 g DW per plant. Some tipburn injury was apparent on most of the plants at harvest. By 28 DAP, stand photosynthesis rates for the entire chamber (approx. 20 m2) reached 17.4 umol CO2 m-2 s-1, while dark-period respiration rates reached 5.5 umol CO2 m-2 s-1. Results suggest that good yields can be obtained from lettuce grown in a tightly sealed environment.
A wheat (Triticum aestivum cv. Yecora Rojo) stand was grown using nutrient film culture in the closed conditions of NASA's Biomass Production Chamber. Rates of photosynthesis and respiration of the entire stand (about 20 m2) were determined daily using a regime of 20 hr light/4 hr dark, 20 C light/16 C dark an average PPF of 600 μmol/m2/s from HPS lamps, and a CO2 cone of 1000 ppm. Fractional interception of PPF by the stand reached a maximum of 0.96 at 24 days from planting. Rates of photosynthesis were constant throughout the photoperiod as determined by short term drawdowns of CO2 throughout the photoperiod. Drawdown rates of CO2 were correlated with rates determined by logging of mass flow of CO2 injected during chamber closure. Photosynthetic drawdowns of CO2 indicated that photosynthesis was not saturated at 1000 ppm CO2 and that the CO2 compensation point was about 50 ppm. Whole stand light compensation points were 200 to 250 μmol/m2/s between days 13 and 70 and then increased rapidly during senescence.
Two studies were conducted in which `Waldmann's Green' lettuce (Lactuca sativa L.) was grown hydroponically from seed to harvest in a large (20-m2), atmospherically closed growth chamber for the National Aeronautics and Space Administration's controlled ecological life support system (CELSS) program. The first study used metal-halide (MH) lamps [280 μmol·m-2·s-1 photosynthetic photon flux (PPF)], whereas the second used high-pressure sodium (HPS) lamps (293 μmol·m-2·s-1). Both studies used a 16-hour photoperiod, a constant air temperature (22 to 23C), and 1000 μmol·mol-1 CO2 during the light period. In each study, canopy photosynthesis and evapotranspiration (ET) rates were highly correlated to canopy cover, with absolute rates peaking at harvest (28 days after planting) at 17 μmol CO2/m2 per sec and 4 liters·m-2·day-1, respectively. When normalized for actual canopy cover, photosynthesis and ET rates per unit canopy area decreased with age (between 15 and 28 days after planting). Canopy cover increased earlier during the study with HPS lamps, and final shoot yields averaged 183 g fresh mass (FM)/plant and 8.8 g dry mass (DM)/plant. Shoot yields in the first study with MH lamps averaged 129 g FM/plant and 6.8 g DM/plant. Analysis of leaf tissue showed that ash levels from both studies averaged 22% and K levels ranged from 15% to 17% of tissue DM. Results suggest that lettuce should be easily adaptable to a CELSS with moderate lighting and that plant spacing or transplant schemes are needed to maximize canopy light interception and sustain efficient CO2 removal and water production.