Environmental Plant Physiology Facilities and Tools K. Raja Reddy Krreddy@pss.msstate.edu Environmental Plant physiology and Facilities and Tools 9 Facilities: ¾ Field plots Environmental and Cultural Factors Limiting Potential Yields ¾Atmospheric Carbon Dioxide ¾Temperature (Extremes) ¾Solar Radiation ¾Water ¾Wind ¾Nutrients (N and K) ¾Others, ozone etc., ¾Growth Regulators (PIX) Crop Responses to Environment - Tools Field Plots ¾ Free-air carbon dioxide enrichment (FACE) facilities ¾ Indoor plant growth chambers and Greenhouses ¾ Sunlit plant growth chambers 9 Tools: • Environmental factors co-vary at all times. ¾ Crop simulation models Crop Responses to Environment - Tools • Therefore, cause and effects are difficult to understand and quantify. Crop Responses to Environment - Tools Open Top Chambers, USDA-ARS, Beltsville, MD Open Top Chambers, USDA-ARS, Auburn, AL • Some control over certain environmental factors. • Others such as temperature is not controlled; chamber walls etc.. • Some control over certain environmental factors. • Others such as temperature is not controlled; chamber walls etc.. 1 Crop Responses to Environment - Tools Crop Responses to Environment - Tools Free-Air-Carbon Dioxide Enrichment –Soybean T-FACE with infrared heaters Free-Air-Carbon Dioxide-Enrichment • Large study area allows for multiple disciplines. • Some control over certain environmental factors such as CO2 and ozone. • Others such as temperature earlier is not controlled, but recently added infrared heating. Crop Responses to Environment - Tools Solardome – Institute of Terrestrial Ecology, Bangor, UK Crop Responses to Environment - Tools Indoor plant growth chambers and greenhouses • Some control over certain environmental factors. • Some control over certain environmental factors. • Others such as: temperature controlled to certain degree to the ambient levels; chambers walls etc. SPAR - Soil-Plant-Atmosphere-Research Plant Process Quantification and Modeling www.spar.msstate.edu • Suitable for certain studies; however, low light levels, poor control over temperatures, inadequate pot sizes and fertility and irrigation management. SPAR - Soil-Plant-Atmosphere-Research What Can We Control? 9 Temperatures: 10 to 45 °C 9 CO2 concentration: Subambient to 1000 ppm 9 Ultraviolet-B radiation: 0 to three times of ambient UV-B (up to 16 kJ) 9 Water regimes: Can be manipulated based on measured ET nicely A 50 ton cooling & coolant circulating system Two 5.5 kW heating, air circulation & moisture condensing system 9 Fertilization: One or several nutrients can be easily manipulated either alone or in combination Mini-rhizotron system for non-destructive root growth and development 9 Solar radiation: sunlit (>95% passes through the Plexiglas and reaches plant canopy), no artificial light 2 SPAR - Soil-Plant-Atmosphere-Research SPAR - Soil-Plant-Atmosphere-Research What Can We Measure? What can we measure? 9 Growth and developmental processes: 9 Abiotic conditions: - Air, canopy and dew-point temperatures - Solar and ultraviolet-B radiation - Chamber and outside CO2 concentrations - Soil water and temperature by depth - Relative humidity I. Phenological rates: - Similar events: Leaf and internode addition rates, duration rates, etc. - Dissimilar events: seed to emergence, emergence to square, square to flower and flower to open boll. 9 Processes: - Canopy photosynthesis, respiration, and evapotranspiration - Leaf-level physiological, biochemical and molecular processes II. Growth rates: - Leaf, internode (stem), root, and fruiting structures (square, boll, lint, seed/grain etc.). SPAR - Measuring Carbon Fluxes SPAR - Measuring Carbon Fluxes Measuring Photosynthesis: Mass-balance approach Measuring Respiration: During sunlit hours, by maintaining steady or constant CO2 concentration inside the SPAR chamber, we can calculate: During nighttime, by measuring the rise or build up CO2 concentration inside the SPAR chamber, we can calculate, Respiration rate = [(CO2 Conc., at Time 2 - CO2 Conc., at Time 1) + leak rate] Net photosynthesis = Amount of CO2 injected – leak rate Gross Photosynthesis = Net photosynthesis + Respiration Regulators/ Valves/flow meters SPAR DACS or Computer Drying agent Drying agent Regulators/ Valves/flow meters X CO2 analyzer X Compressed CO2 Suction pump SPAR Drying agent Drying agent CO2 analyzer DACS or Computer SPAR – Process Quantification and Modeling Canopy Photosynthesis and Diurnal Trends Canopy Photosynthesis and Light Response Curves 2500 720 ppm 6 Solar Radiation 360 ppm -1 2000 1500 2 1000 0 500 -2 0 0 2 4 6 8 10 12 14 16 18 20 22 24 Time of the Day (Central Standard Time) -2 4 Canopy photosynthesis, mg CO2 m-2 s-1 SPAR – Process Quantification and Modeling PPFD, µmol m s Carbon Exchange Rate, mg CO2 m-2 s-1 Compressed CO2 Suction pump 9 360 ppm and N-sufficient 720 ppm and N-sufficient 720 ppm and N-deficient 360 ppm and N-deficient 8 7 6 5 4 3 2 1 78 DAE and 28 DAT 0 0 500 1000 1500 2000 PPFD, µmol m-2 s-1 3 SPAR – Process Quantification and Modeling SPAR – Process Quantification and Modeling Cotton Canopy Photosynthesis – N and CO2 Cotton Conductance – N and CO2 Temporal Trends in Photosynthesis Processes Temporal Trends in Canopy Conductance 20 8 6 8 0 N-sufficient N-deficient 720 ppm 8 360 ppm 720 ppm 7 6 5 3 2 1 1 2 4 3 Leaf N, % 4 2 30 40 50 60 70 80 5 5 720 ppm N-sufficient N-deficiennt 15 200 5 1 2 3 Leaf N, % 4 5 5 30 40 50 60 70 80 90 100 110 120 130 140 SPAR – Process Quantification and Modeling Leaf-level Gas Exchange and Reflectance Measurements • -2 1995 ambient Temperature We can monitor leaf-level gas exchange: 9 150 • 100 • 50 Boll opening Flower 60 80 100 120 140 160 Day after emergence SPAR – Process Quantification and Modeling Measuring Evapotranspiration (ET) Photosynthesis, stomatal conductance, transpiration, fluorescence, etc. We can monitor leaf-level reflectance measurements: 9 40 10 Days after emergence SPAR – Process Quantification and Modeling Integration of Canopy Net Photosynthesis 0 20 360 ppm 720 ppm 15 0 10 0 20 90 100 110 120 130 140 Days after emergence -1 20 0 4 6 0 20 Nitrogen and Canopy Conductance Relationship 10 -1 2 Canopy photosynthesis (mg CO2 m-2 s-1) 4 Canopy net photosynthesis, g CO2 m d -2 Canopy photosynthesis, mg CO2 m s -1 Nitrogen and Photosynthesis Relationships 360 ppm N-sufficient N-deficient 15 Canopy Conductance (mm s-1) Canopy conductance, mm s N-sufficient N-deficient Canopy conductance, mm s 360 ppm -1 10 Leaf reflectance properties, pigments etc. We can also monitor leaf temperatures and leaf water potentials: 9 Leaf temperatures by infrared thermometers. 9 Leaf water potential by Pressure bomb. SPAR – Process Quantification and Modeling Measuring Evapotranspiration 1. Measuring Evapotranspiration (ET): During day and nighttime periods, by collecting the condensate (moisture in the air) while passing through the cooling coils, ET is measured using a set of values, controllers and pressure transducers every 15 minutes. Inlet – From Condenser Coils in the unit Condensate ET Reservoir 2. Measuring transpiration: By sealing the soil surface and around the plant stems, one can accurately measure transpiration. Solenoid Valves Transducer Outlet - Drain 4 SPAR - Process Quantification and Modeling Cotton - Leaf and Canopy Transpiration and Leaf Area SPAR – Process Quantification and Modeling 0.2 1000 500 . 0 0.0 2 4 6 8 10 12 14 16 18 20 22 24 2.0 1.8 1.6 1.4 1.2 1.0 0.8 0.6 0.4 0.2 Time of the Day (Central Standard Time) Leaf Area Basis y = 36.741 + 0.5098 * X, r ² 0.99 250 200 150 100 100 150 200 250 300 350 400 -2 -1 Transpiration at Ambient [CO2], mg H2O m s 2 0.1 About 250 per sq mm 300 Transpiration at Elevated [CO2], kg H2O m-2 d-1 1500 -1 0.3 0 H20 2000 Solar Radiation Leaf Area at Elevated [CO2], m plant 0.4 CO2 Maize, DAE 36 PPFD, µmol m-2 s-1 Canopy Water Use, g H2O m-2 s-1 2500 -1 720 µmol CO2 mol -1 360 µmol CO2 mol Transpiration at Elevated [CO2], mg H2O m-2 s -1 Maize - Canopy Evopotranspiration – Diurnal Trends 0.5 0.0 0.0 0.2 0.4 0.6 0.8 1.0 1.2 2 Leaf Area at Ambient [CO2], m plant 1.4 1.6 1.8 2.0 40 Canopy Transpiration 30 20 10 y = 2.7799 + 0.9234 * X, R² 0.83 0 0 10 -1 20 30 40 Traspiration at Ambient [CO2], kg H2O m-2 d-1 SPAR – Process Quantification and Modeling SPAR – Process Quantification and Modeling Evapotranpsiration and Two Methods Condensate and TDR - Potato Cotton – Determining Potential Developmental Rates Evapotranspiration -2 -1 from condensate, mm cm d 10 CO2 and Irrigation Irrigation Treatments 370 µmol mol 8 -1 25% 75% 100% 740 µmol mol -1 25% 75% 100% 6 4 2 Leaf addition rates on the mainstem and branches and leaf expansion duration 0 0 Dennis et al. 2007 2 4 6 8 Evapotranspiration from TDR 10 water content, mm cm-2 d-1 SPAR – Process Quantification and Modeling SPAR – Process Quantification and Modeling Cotton – Developmental Rates Cotton – Growth and Developmental Rates Pictorial Representation 0.07 4-week old cotton seedlings Rate of Development,1/d-1 0.06 Emergence to Square = -0.1265 + 0.01142*X - 0.0001949*X2; r2 = 0.98 Square to Flower = -0.1148 + 0.00967*X - 0.0001432*X2; r2 = 0.94 Flower to Open Boll = -0.00583 + 0.000995*X; r2 = 0.92 Square to Flower 0.05 0.04 Emergence to square 0.03 0.02 Flower to open boll 0.01 20/12 25/17 30/22 35/27 Day/night Temperature, °C 40/32 0.00 15 20 25 30 35 40 Temperature, °C 5 SPAR - Plant Responses and Modeling Cotton – Fruit Production Efficiency SPAR - Plant Responses and Modeling Cotton – Square and Boll Growth Rates 0.012 0.12 0.11 0.010 0.10 0.009 0.09 0.008 0.08 Boll 0.007 0.07 0.006 0.06 0.005 15 20 25 30 35 Boll Growth Rate, g d-1 Square growth rate, g d-1 0.011 Fruit Production Efficiency (g kg-1 dry weight) 400 Square 300 200 100 0 24 0.05 40 700 µL L-1 350 µL L-1 26 28 SPAR – Plant Responses and Modeling Cotton Growth Rate Responses to Water Stress 34 1.2 1.0 Photosynthesis Environmental Productivity Indices for Nitrogen Relative Growth Rates 32 SPAR – Plant Responses and Modeling Cotton Growth and Developmental Responses to N 1.2 0.8 0.6 0.4 0.2 30 Temperature, °C Temperature, °C Stem Elongation Leaf Area Expansion 1.0 0.8 0.6 Stem Growth Photosynthesis Leaf Development 0.4 0.2 Leaf Growth 0.0 -1.0 -1.5 -2.0 -2.5 -3.0 -3.5 0.0 1.00 1.25 Midday Leaf Water Potential, MPa SPAR – Plant Responses and Modeling What about Replication? Environment variables 1.75 2.00 2.25 2.50 SPAR – Plant Process Quantification and Modeling 9 Sunlit, but other abiotic factors can be controllable nicely. Plant variables Variable Mean and SD of 12 SPAR Units Variable Mean of 12 SPAR ± Variance Range of variance with in the SPAR Tmax, °C 23.0 ± 0.2 Height, cm 54.9 ± 1.7 2.2 – 18.0 Tmin, °C 18.2 ± 0.7 Leaf area, cm2 141 ± 784 1716 -5120 CO2 - day, ppm 700 ± 90 Total weight, g plant-1 16.9 ± 2.1 22.2 – 84.2 CO2 - night CO2 548 ± 52 Yield, g plant-1 12.2 ± 0.99 13.4 – 46.5 Humidity - day, % 58 ± 5 Humidity - night, % 60 ± 4 1.50 Leaf Nitrogen, g m-2 leaf area 9 Not too expensive if the objectives are to quantify processes and to develop modeling tools. 9 Very well suited for multiple environmental effects on plants either alone or in combination. 9 Particularly very well suited to address omics (genomics, metabomolics, proteomics) questions related environmental controls and responses in crop and plant science area. 9 Space is limited. 6 System Simulation Tools The Cotton simulation model, GOSSYM Timeline for Information Flow Identify knowledge void GOSSYM Months DATES CLYMAT Conceptualize the experiment TMPSOL SOIL FERT RAIN PIX FRTLIZ RUNOFF Months/Years GRAFLO Implementation ET CHEM PREP Months UPTAKE Analyze data CAPFLO PNET Years NITRIF GROWTH ABSCISE PLTMAP PMAP COTPLT FREQ RUTGRO RIMPED Crop model/DSS NITRO MATAL Months Scientists Publication Months/Years Industry Reps Technology transfer OUTPUT Consultants Months/Years Program flow Timeline for Information Flow Ext. Personnel Farmers Farm decisions SPAR and Crop Model Applications Results Researcher DSS Ext. Personnel Box: 1 Box: 1 Box: 1 Reports Specify conditions Reports Box: 1 Industry Reps. Box: 1 Box: 1 User Generalize, Enhance Specify, Refine, Distort Summary - SPAR Capabilities ¾ With cotton as an example crop, we have shown how the SPAR system can be used to generate data needed for understanding the various facets of growth and developmental processes and how this understanding can be used for building process-level models and in learning how to manage the cotton crop. ¾ Operating a SPAR facility to acquire such data will often be more economical than the use of field plot experiments because it allows the scientist to avoid many of the covarying and confounding factors that occur in field experiments. Thus, the basic processes can be related more directly to the environmental variables being studied. Summary - SPAR Capabilities ¾ As we progress in developing systems for understanding plant responses to environment, whether in support of global climatic change research, the application of plants in the remediation of environmental conditions, or the increased application of precision agriculture technologies, the need for diagnostics and management decision aids will become more urgent. ¾ Mechanistic plant models and automated, user-friendly expert systems can facilitate selection of the optimum solutions to problems with many variables. 7 Summary - SPAR Capabilities ¾ Essentially all of the engineering and computing technologies needed to allow the use of variable and sitespecific technologies, such as precision agriculture, are now available. ¾ However, our understanding of the plant ecophysiological responses to the environment as it relates to specific growth and developmental events requires further development. ¾ Modeling forces the organization of known information and concepts. Although we may not know enough to develop a comprehensive model that includes all aspects of plant growth and development at the landscape or even the plot scale, modeling some meaningful portions of the system provides clarity. Summary - SPAR Capabilities ¾ These environment-genotype interactions can be measured and incorporated into a meaningful model. ¾ When a model is based on appropriate concepts and processes, it has predictive capability in new environments and can be used either alone or with other emerging newer technologies to disseminate useful plant growth and development information. Summary - SPAR Capabilities ¾ For a model to correctly predict plant responses to physical conditions, the concepts and the response functions must be appropriately assembled. Critical environment-genotype relations should be incorporated into the model. ¾ These relationships include, but should not be limited to, the phenological responses of specific genotypes to temperature and their responses to environmental stresses. ¾ We would, for example, expect to find quantifiable differences among genotypes in fruit-shed sensitivity to above-optimum temperature and to deficiencies of water and/or nutrients. ¾ One might also find differences in fruit-shed sensitivity to carbon deficiency caused by imbalance between photosynthesis, fruiting rate, and vegetative growth. Summary - SPAR Capabilities ¾ In the past, the SPAR facility has been used extensively for research on only a few species, with a primary purpose of providing functional parameterizations used in crop simulation models, which, in turn, are a component of expert cropmanagement decision-support systems. ¾ There are a variety of approaches and facilities to investigate plant responses to the environment. Among these, the SPAR facilities are optimized for the measurement of plant and canopy-level physiological responses to precisely controlled, but naturally lit, environmental conditions. The data that have been and will be obtained are unique and particularly instructive for applied and basic plant biologists. Facilities Suggested Reading Material • Reddy, K. R., J. J. Read, J. T. Baker, J. M. McKinion, L. Tarpley, H. F. Hodges and V. R. Reddy. 2001. Soil-PlantAtmosphere-Research (SPAR) facility - a tool for plant research and modeling. Biotronics 30: 27-50. • Reddy, K. R. V. G. Kakani, J. M. McKinion and D. N. Baker. 2002. Applications of a cotton simulation model, GOSSYM, for crop management, economic and policy decisions. In: L. R. Ahuja, Liwang Ma and T. A. Howell (Eds.) Agricultural System Models in Field Research and Technology Transfer, CRC Press, LLC, Boca Raton, FL, USA. Pp 33-73. 8