Jan. 20, 2011 B4730/5730 Plant Physiological Ecology Biotic and Abiotic Environments Photosynthesis, O2 and H2O • Plants face two major problems – 1) whenever stomata open to allow CO2 to diffuse to the locations of carbon fixation, H2O invariably leaves – 2) Rubisco fixes both CO2 and O2 • Transpiration loss of H2O from plants – Stomatal physiology tries to maximize photosynthesis while minimizing transpiration – Stomatal closure decreases CO2 concentrations and increases O2 concentrations promoting O2 fixation • Photorespiration fixation of O2 by Rubisco – Photorespiration requires light – Photorespiration produces no ATP – Photorespiration uses organic material from the Calvin cycle Alternative Pathways of Photosynthesis • Three major photosynthetic pathways based on which molecule first incorporates CO2 – 1) C3 plants fix CO2 into 3-PGA (3 carbon) – 2) C4 plants initially fix CO2 into a 4 carbon molecule before passing it to the Calvin cycle – 3) CAM plants initially fix CO2 into organic acids • C4 and CAM photosynthetic pathways minimize transpiration and photorespiration at the cost of additional energy for carbon fixation – Temporal or spatial separation – Light reactions same for all pathways Defining Environment • Environment of plants is anything outside of the plant body that influences the plant – Response to present environment due to adaptations/acclimations to previous environments • Biotic and abiotic environmental interactions – Both positive and negative – Stress never completely alleviated Spheres of Plants • Atmosphere – Plants respond to and change the atmosphere – Climate and atmosphere, atmospheric cycles • Hydrosphere – Heat transfer from water evaporation – Water and climate • Lithosphere – Any lithosphere with biological activity is soil – Soil properties and plant response to environment intimately linked • Ecosphere combines all of the above and biological interactions – Rhizosphere most neglected Atmospheric structure http://www.ux1.eiu.edu/~cfjps/1400/atmos_origin.html Boundary Layers • Turbulence is nonparallel, disorderly flow of a fluid – Turbulence intensity is standard deviation of flow divided by mean flow – Increasing turbulence means more fluid moves by eddies • Boundary layers formed by shearing stresses at some surface – Boundary layers form at any solid/liquid interface – Flow must go to zero at interface Visualizing Boundary Layers http://www.cartage.org.lb/en/themes/Sciences/Physics/Mechanics/FluidMechanics/RealFluids/BoundaryLayers/BoundaryLayers.htm Radiation Fundamentals • Total amount of radiation received by a body on earth is a combination of short and longwave radiation • Amount of energy from radiation is a function of the wavelength – Wien’s displacement law – Planck’s law – Stephan-Boltzman law • Shortwave radiation is received directly from the sun from high temperatures • Longwave radiation is given off by bodies on earth Energy emitted by the sun and earth Oke et al. 1987 Atmospheric Effects on Radiation Landsberg and Gower 1997 Oke et al. 1987 Leaf Energy Budgets • The energy budget of a leaf determines its leaf temperature • TL-TA = (RN-λEL)/(ρ·cp·gT) – – – – – – – – TL is leaf temperature TA is air temperature λ latent energy of evaporation EL transpiration per unit leaf area ρ is air density cp is the heat capacity of air gT is the total conductance to water vapor Metabolic heat generation is generally ignored but can be substantial in certain species Impact of canopy structure on temperature and photosynthesis Smith and Carter 1988 Triangles Abies lasiocarpa Diamonds or squaredot Picea engelmannii Squares Pinus contorta Radiation Balance • Radiation balance requires conservation of energy • RN=(1-α)RS+(RLi+RLo) + G – RN is net radiation – α is albedo – RS is solar radiation – RL is longwave radiation (i) incoming (o) outgoing – G is storage Impact of Vegetation on Albedo Landsberg and Gower 1997 Partitioning of Net Radiation • Net radiation and physical and physiological controls on water loss determine the temperature of a stand of vegetation • RN + G = λE + H – G is heat storage – λE is the latent energy or heat of vaporization – H is the sensible energy or heat Examples of Energy Balance Using Eddy Covariance Techniques Wilson et al. Ag. For. Met. 2002 Picea mariana; Goulden et al. 1997 Baldocchi et al. 1997 Fluxes • Molecules move from high concentration to low concentration – Entropy • Flux is the amount substance moving across a planar front per unit time • Flow is the total amount of substance moving per unit time • Ficke’s first and second laws describe fluxes • Flux density is proportional to the driving force – Diffusion coefficient changes flux per driving force • Diffusion coefficient can be converted to resistance or conductance to flux – Resistances sum directly in series – Conductances sum inversely in series Plants respond to environment with fluxes • Plant fluxes – Mass – Energy – Momentum • Soil Plant Atmosphere Continuum (SPAC) defines where fluxes occur – subcellular to global • Deriving flux equations; connecting anatomy – Photosynthesis – Transpiration • Importance of scale SPAC What other fluxes in SPAC besides water? http://www.fsl.orst.edu/~bond/phystalk/Ecohydrology/SPAC%20diagram.jpg