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Feb. 1 st , 2011 B4730/5730 Plant Physiological Ecology. Photosynthesis I. Filled crop, open broadleaf, crosses conifers Medlyn et al. PCE 2002. Light Dependent Reactions. Three fates of photon striking reaction center Heat, fluorescence, photochemistry
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Feb. 1st, 2011B4730/5730Plant Physiological Ecology Photosynthesis I
Filled crop, open broadleaf, crosses conifers Medlyn et al. PCE 2002
Light Dependent Reactions • Three fates of photon striking reaction center • Heat, fluorescence, photochemistry • Linear electron flux (LEF) goes from PSII to PSI to drive H+ pmf, • Cyclic flux around PSI • Low light, efficiency maximized • High light, energy dissipation maximized
Chlorophyll Fluorescence • Absorption (photochemistry), heat, and fluorescence all occur in competition with incoming light • Light absorbed different wavelength than light fluorescing • Light always lost, so all measurements relative • Add to 1 • Measurements made with modulated light • Allows measurement in the field with background light levels • Kautsky effect is rise in fluorescence when leaves go from dark to light conditions • Electrons from PSII to QA, • if QA reduced; reaction center closed • fluorescence when reaction center closed • Quenching causes fluorescence to fall after initial burst • Photochemical quenching-light induction • Nonphotochemical quenching (NPQ)-heat and zeaxanthin cycle
Measuring Fluorescence • F rate of fluoresce • ’ indicates light acclimated (“adapted”) measurements • Fo background fluorescence in dark, Fo’ in light • Fm maximal rate in dark or Fm’ in light • Occurs when QA is maximally reduced • Fv variable fluorescence from dark, Fv’ light • Demonstrates primary photochemistry
Interpreting Fluorescence • Fv/Fm intrinsic quantum yield of PSII • Fv’ = F’-Fm’ • Quantum yield of PSII • ΦPSII = Fv’/Fm’ • Photochemical (qP) and nonphotochemical quenching (NPQ) • qP = (F’m – F’o)/(Fm – Fo) • NPQ = 1 – (F’m – F’o)/(Fm – Fo) • J = ΦPSII · Iabs · β · PPFD • β = 0.5 equal distribution PSII & PSI • Relative without Iabs
Nicotiana tabacum; Kramer et al Photo. Res. 2004 qp puddle model PSII open qL lake model PSII open
Wang & Kellomaki New Phyt. 1997, Pinus sylvestris • Control N & amb CO2, b) Control N & elev CO2, • c) High N & amb CO2, d) High N & elev CO2 • ΔF=Fq’
Fs is steady state Fluorescence (F’) Closed circle F’m amb CO2 Closed triangle Fs amb CO2 Open circle F’m elev CO2 Open triangle Fs elev CO2
Mesophyll conductance • Flux of CO2 to stroma (Cc) limited by gs, gm and gb • Recent work has shown gm to be variable and important • Low gm reduces CC underestimate VCmax • Cc=Ci-A/gm • Estimate gm with curve fitting • Estime gm with fluorescence and A
Many species; Sc= surface area of chloroplasts exposed to intercellular airspace; Evans et al J Exp Bot 2009
Limitations to the Farquhar/Fluorescence model of Photosynthesis • No gene expression controls • Connection to hormones, whole plant controls • Longer time scales • Stomatal physiology almost a black box • Scale of leaf patchiness violates homogeneity • Light dynamics problematic • Respiration models are crude