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Photorespiration & Variations in Photosynthesis Lecture 15 Spring 2014 Variation in Photosynthesis Three types of photosynthesis • C3 • C4 • CAM Why do plants need three different types of photosynthesis? 1 Water Balance in Plants Transpiration & role of stomata 2 Balance between photosynthesis and water loss C3 • 3-carbon sugar (3-phosphoglycerate) in Calvin cycle • Day/Night cycle • In hot temperatures, must close stomata 3 Rubisco • ribulose bisphosphate (RuBP) carboxylase • Rubisco has affinity for both CO2 and O2 Fig. 10.19 4 Photorespiration • A metabolic pathway that consumes O2 and ATP, releases CO2 and reduces photosynthetic output 5 Why Photorespiration? • Rubisco evolved in bacteria before O2 in atmosphere – Enzyme had affinity for both • Modification of active site to reduce access to oxygen may also reduce the access to CO2 • C3 plants compensate by increasing the concentration of rubisco. – Half of the protein in the chloroplast is rubisco • PR may be protective in terms of preventing damage from excessive light 6 7 Alternative Pathways: C4 C4 • First carbon fixation produces a 4-carbon sugar (oxaloacetate) • Can close stomata during day • hot/dry environments – E.g., many grasses, sugar cane, corn Alternative Pathways: C4 • Two types of cells – Mesophyll cells • Carbon fixation – Bundle-sheath cells • Calvin cycle in chloroplasts See Fig. 10.19 • Keeps high CO2 levels in bundle-sheath cells 8 9 Alternative Pathways: C4 In mesophyll cells • CO2 fixed to PEP (phosphoenolpyruvate) • PEP carboxylase – No affinity for O2 Fig. 10.19 10 Cyclic Electron Flow • Linear electron flow produces 6 ATP & 6 NADPH – Calvin cycle requires 9 ATP • Uses cyclic electron flow to generate extra ATP • Occurs in bundle-sheath cells Fig. 10.15 11 Alternative Pathways: CAM CAM (Crassulacean acid metabolism) • Stomata open during night • CO2 stored as 4-carbon compound to be used the next day • Stomata can be closed during the day • Plants found in hot/dry environments – Cactus, pineapple, many orchids – (some are members of plant family Crassulaceae) 12 Alternative Pathways: CAM • Temporal separation of carbon fixation and Calvin cycle • Takes in CO2 at night • Uses C4 pathway to fix CO2 into organic acids – malate • Stored in vacuole – Malic acid – Active transport 13 Alternative Pathways: CAM • Malic acid leaves vacuole • Broken down to form pyruvate and CO2 • CO2 to Calvin cycle • Pyruvate converted to starch and stored 14 Alternative Pathways: CAM Fig. 10.20 15 Trade Offs Type of CO2 fixation Energy to fix one CO2 C3 3 ATP Water Transpired per CO2 fixed 400 – 500 g C4 5 ATP 250-300 g CAM 6.5 ATP 50-100 g 16 Importance of Photosynthesis • Provides the carbon compounds for most organisms on the planet • Changed Earth’s atmosphere to the current one • Important part of the CO2 cycle – Moderates temperature on the planet