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Overview: The Process That Feeds the Biosphere • Photosynthesis is the process that converts solar energy into chemical energy • Directly or indirectly, photosynthesis nourishes almost the entire living world Autotrophs • Autotrophs sustain themselves without eating anything derived from other organisms • Autotrophs are the producers of the biosphere, producing organic molecules from CO2 and other inorganic molecules • Almost all plants are photoautotrophs, using the energy of sunlight to make organic molecules from H2O and CO2 Photosynthesis occurs in plants, algae, certain other protists, and some prokaryotes (a) Plants (c) Unicellular protist 10 µm (e) Purple sulfur bacteria (b) Multicellular alga (d) Cyanobacteria 40 µm 1.5 µm Heterotrophs • Heterotrophs obtain their organic material from other organisms • Heterotrophs are the consumers of the biosphere • Almost all heterotrophs, including humans, depend on photoautotrophs for food and O2 The Two Stages of Photosynthesis: A Preview • Photosynthesis consists of the light reactions (the photo part) and Calvin cycle (the synthesis part) • The “light” reactions are in the thylakoids • The Calvin cycle is in the stroma Chloroplast Outer membrane Thylakoid Stroma Granum Thylakoid space Intermembrane space Inner membrane 1 µm • The “light” reactions overview – Capture light energy using chlorophyll – Split H2O to electrons, protons and oxygen – Release O2 – Reduce NADP+ to NADPH – Generate ATP from ADP by photophosphorylation • The Calvin cycle overview - begins with carbon fixation, incorporating CO2 into organic molecules - forms sugars from these organic molecules, using ATP and NADPH (from light reactions) H2O Light NADP+ ADP + P i Light Reactions Chloroplast Overview of Photosynthesis 1 H2O Light NADP+ ADP + P i Light Reactions ATP NADPH Chloroplast O2 Overview of Photosynthesis 2 CO2 H2O Light NADP+ ADP + P i Light Reactions Calvin Cycle ATP NADPH Chloroplast O2 Overview of Photosynthesis 3 CO2 H2O Light NADP+ ADP + P i Light Reactions Calvin Cycle ATP NADPH Chloroplast Overview of Photosynthesis 4 O2 [CH2O] (sugar) Chlorophyll is the light-capturing substance Surrounded by other pigment molecules, enzymes and electron carriers A “photosystem” • Anchored to thylakoid membranes by proteins Illustration of important properties of chlorophyll Energy of electron e– Excited state Heat Photon (fluorescence) Photon Chlorophyll molecule Ground state (a) Excitation of isolated chlorophyll molecule (b) Fluorescence If illuminated, a solution of chlorophyll will fluoresce, giving off light and heat Photosystem STROMA Light-harvesting Reaction-center complex complexes Primary electron acceptor Thylakoid membrane Photosynthetic Photon Energy Capture e– Transfer of energy Special pair of chlorophyll a molecules Pigments-chlorophyll and accessory THYLAKOID SPACE (INTERIOR OF THYLAKOID) “Z-Diagram” of photosynthesis 4 Primary acceptor 2 H+ + 1/ O 2 2 H2O e– 2 Primary acceptor e– Pq Cytochrome complex 7 Fd e– e– 8 NADP+ reductase 3 NADPH Pc e– e– P700 5 P680 Light 1 Light 6 ATP Pigment molecules Photosystem II (PS II) NADP+ + H+ Photosystem I (PS I) This illustrates “non-cyclic” electron flowelectrons enter with water and leave with NADPH Primary acceptor Primary acceptor Fd Fd Pq NADP+ reductase Cytochrome complex NADP+ + H+ NADPH Pc Photosystem I Photosystem II ATP Cyclic electron flow uses only photosystem I and produces ATP, but not NADPH • “Light” reactions of PSI and PSII generate ATP and “reducing power” in the form of NADPH • Reducing power allows the conversion of carbon dioxide to carbohydrate • CO2 to CH2O requires NADPH reducing power Input 3 Carbon fixation conversion of carbon dioxide to carbohydrate (Entering one at a time) CO2 Phase 1: Carbon fixation Rubisco 3 P Short-lived intermediate P 6 P 3-Phosphoglycerate 3P P Ribulose bisphosphate (RuBP) 6 ATP 6 ADP Calvin Cycle 3 ADP 3 ATP 6 P P 1,3-Bisphosphoglycerate 6 NADPH Phase 3: Regeneration of the CO2 acceptor (RuBP) 6 NADP+ 6 Pi P 5 G3P Rubisco is the key enzyme of the cycle 6 P Glyceraldehyde-3-phosphate (G3P) 1 Output P G3P (a sugar) Glucose and other organic compounds Phase 2: Reduction Plants that use this cycle exclusively are C3 plants • Dehydration is a problem for plants, sometimes requiring trade-offs with other metabolic processes, especially photosynthesis • Plants have evolved two alternate types of carbon fixation • C4 and Crassulean Acid Metabolism (CAM) fixation H2O CO2 Light NADP+ ADP + P i Light Reactions: Photosystem II Electron transport chain Photosystem I Electron transport chain RuBP ATP G3P Starch (storage) NADPH Chloroplast 3-Phosphoglycerate Calvin Cycle C3 Photosynthesis: a review O2 Sucrose (export) Note Card Questions 1) What is the “reducing power” of NADPH and how does it link the light reactions with the Calvin Cycle? 2) Why are 2 light absorbance events (2 photons) required by oxygenic photosynthesis?