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Feb. 3rd, 2011 B4730/5730 Plant Physiological Ecology. Respiration I. F s is steady state Fluorescence (F’) Closed circle F’ m amb CO2 Closed triangle F s amb CO2 Open circle F’ m elev CO2 Open triangle F s elev CO2. Leipner et al Env. Exp. Bot. 1999 Maize. Mesophyll conductance.
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Feb. 3rd, 2011B4730/5730Plant Physiological Ecology Respiration I
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
Autotrophic Respiration • Can not describe plant growth from photosynthesis only • High N requirements for enzymes of both • Total Respiration is the sum of • Maintenance • Growth • Transport • All eukaryotes with mitochondria have cytochrome respiratory pathway • Inhibited by cyanide • Plants have cyanide resistant pathway that uses AOX • animal regulation before glycolysis, plants during
Autotrophic Respiration • Respiration can be regulated by energy demand or substrate supply • Photosynthesis and respiration connected through chemiosmosis (AOX) • Ratio of CO2 to O2 (respiratory quotient) varies with tissue type and substrates used • <1 more reduced substrates • >1 more reduced products
Alternative Respiration Pathway • Alternative Respiration pathway (CN resistant) • Respiration production exceeds demand, reactive oxygen species (ROS) activated • Uses alternative oxidase (AOX) instead of cytochrome electron transport chain • Generates heat • Various hypotheses about regulation and purpose • Uncoupling proteins (UCP) connect to photosynthesis • Uncoupling proteins relieve excess pH gradient in mitochondria • Photorespiration inhibited by loss of UCP, ROS problems
Cucumis sativus; Juszcuk et al Phys Plant 2007 MSC16 mutant with lowered respiration/growth rates encoded by mitochondrial genome
Arabidopsis thaliana; Sweetlove et al PNAS 06 ucp1 mutant without uncoupling protein