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Ch 10 Photosynthesis--> To make with light! LE 10-2 Photoautotrophs: Self feeders, producers Use light and inorganic molecules to make own organic molecules. Plants Unicellular protist 10 µm Purple sulfur bacteria 1.5 µm Multicellular algae Cyanobacteria 40 µm • Heterotrophs (food from others): -Consumers -Obtain organic material from other organisms -Dependent on photoautotrophs for food and oxygen Photosynthesis: conversion of light energy into chemical energy Reaction: 6CO2 + 12H2O + light --> C6H12O6 + 6O2 + 6H2O glucose Simplified rxn: 6CO2 + 6H2O + light --> C6H12O6 + 6O2 Simplest rxn: carbohydrate CO2 + H2O + light --> [CH2O] + O2 Enters through roots LE 10-3 6CO2 + 6H2O + light --> C6H12O6 + 6O2 Organic molecule for fuel or other Gas enters through stomata Leaf cross section Vein Stomata CO2 O2 Mesophyll cell Chloroplast 5 µm Granum Thylakoid Thylakoid space or used in respiration Mesophyll Stroma Exits through stomata Outer membrane Intermembrane space Inner membrane 1 µm Two major reactions in photosynthesis Light-dependent (in thylakoid) Light-independent aka dark or Calvin cycle (in stroma) LE 10-7 Chlorophyll in thylakoid membranes Light Reflected light Chloroplast Stroma Absorbed light Granum Transmitted light LE 10-10 CH3 CHO in chlorophyll a in chlorophyll b Porphyrin ring: light-absorbing “head” of molecule; note magnesium atom at center Hydrocarbon tail: interacts with hydrophobic regions of proteins inside thylakoid membranes of chloroplasts; H atoms not shown LE 10-9a Absorption of light by chloroplast pigments Chlorophyll a Chlorophyll b Carotenoids 400 500 600 Wavelength of light (nm) Absorption spectra 700 How do we know that absorption of certain wavelengths of light by plants stimulates a chemical reaction in plants? Specifically how do we know that O2 is a product? LE 10-9c Aerobic bacteria Filament of algae 400 500 600 700 Engelmann’s experiment (1883): Action spectrum What would be an important control experiment? • Chlorophyll a: – main photosynthetic pigment • Accessory pigments – chlorophyll b and carotenoids absorb excessive light that would damage chlorophyll – broaden the spectrum used for photosynthesis Light-Induced Excitation: • When a pigment absorbs light – departs from a ground state to an excited state --> unstable Draw – excited electrons fall back to the ground state, give off photons (glow)-->fluorescence LE 10-11 e– Excited state Heat Photon Chlorophyll molecule Photon (fluorescence) Ground state Excitation of isolated chlorophyll molecule Fluorescence LE 10-5_1 Light-dependent rxn: in thylakoid H2O Light LIGHT REACTIONS Chloroplast LE 10-5_2 H2O Light LIGHT REACTIONS ATP NADPH Chloroplast O2 LE 10-5_3 Calvin cycle: in stroma H2O CO2 Light NADP+ ADP + Pi LIGHT REACTIONS CALVIN CYCLE ATP NADPH Chloroplast O2 [CH2O] (sugar) Photosynthesis as a Redox Process • Water is oxidized (e- are removed). • Carbon dioxide is reduced (e- are gained). Two major reactions in photosynthesis Light dependent (in thylakoid): Creates ATP and an electron carrier, NADPH Electrons supplied through splitting and oxidation of H2O Light -independent (aka dark or Calvin cycle)(in stroma): Synthesis of organic molecules from CO2 Reduction reactions Endergonic: requires ATP Light Reaction: Consists of 2 photosystems Occurs at two different reaction centers each surrounded by light harvesting complexes Light harvesting complex funnels energy to reaction center LE 10-12 Thylakoid Photosystem Photon Thylakoid membrane Light-harvesting complexes Reaction center STROMA Primary electron acceptor e– Transfer of energy Special chlorophyll a molecules Pigment molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID) LE 10-13_1 H2O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH O2 [CH2O] (sugar) Primary acceptor Energy of electrons e– Once P680 is oxidized (gives up e-), is it functional? How is it restored to functionality? Light P680 Photosystem II (PS II) LE 10-13_2 H2O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH O2 [CH2O] (sugar) Energy of electrons Primary acceptor 2 H+ 1/ 2 + O2 Light H2O e– e– e– P680 Photosystem II (PS II) Splitting of H2O yields ethat fill e-”hole” in oxidized P680 LE 10-13_3 H2O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH O2 [CH2O] (sugar) Primary acceptor Energy of electrons Pq 2 H+ + 1/ 2 O 2 Light H2O e– Cytochrome complex Pc e– e– P680 ATP Photosystem II (PS II) LE 10-13_4 H2O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH O2 [CH2O] (sugar) Primary acceptor Primary acceptor e– Energy of electrons Pq 2 H+ 1/ 2 + O2 Light H2O e– Cytochrome complex Pc e– e– P700 P680 Light ATP Photosystem II (PS II) Photosystem I (PS I) LE 10-13_5 H2 O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH O2 [CH2O] (sugar) Primary acceptor Primary acceptor e– Pq Energy of electrons 2 H+ e– H2O Cytochrome complex + 1/2 O2 Light Fd e– e– NADP+ reductase Pc e– e– NADPH + H+ P700 P680 Light ATP Photosystem II (PS II) NADP+ + 2 H+ Photosystem I (PS I) After P700 is oxidized by light energy in PS I are its missing electrons replaced? If so what is the electron source? What would be the effect on photosynthesis if P700 were not reduced to its original state i.e. if the e- hole were not filled? Electron Flow • Noncyclic electron flow – involves both photosystems (II & I) – produces ATP and NADPH LE 10-14 e– ATP e– e– NADPH e– e– e– Mill makes ATP e– Photosystem II Photosystem I Cyclic Electron Flow - Uses only photosystem I - Produces only ATP, no NADPH - Generates surplus ATP – to satisfy demand in the Calvin cycle LE 10-15 Primary acceptor Primary acceptor Fd Fd NADP+ Pq NADP+ reductase Cytochrome complex NADPH Pc Photosystem I Photosystem II ATP How is ATP made? By chemiosmosis LE 10-17 H2 O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH STROMA (Low H+ concentration) O2 [CH2O] (sugar) Cytochrome complex Photosystem II Light 2 Photosystem I Light NADP+ reductase H+ NADP+ + 2H+ Fd NADPH + H+ Pq H2O THYLAKOID SPACE (High H+ concentration) 1/2 Pc O2 +2 H+ 2 H+ To Calvin cycle Thylakoid membrane STROMA (Low H+ concentration) ATP synthase ADP + Pi ATP H+ • Current chemiosmotic model: – H+ (protons) accumulate in thylakoid space • 1. Through splitting of water • 2. By translocation into thylakoid when e- are transported • 3. By removal of H+ from stroma due to bonding with NADPH – H+ diffuses from thylakoid space --> stroma through membrane enzyme, ATP synthase – Movement activates ATP synthase – ATP synthesized on stromal face where the Calvin cycle takes place Products from light reactions power Calvin cycle! What are the light reaction products? ATP: energy carrier NADPH: electron carrier What is the product of the Calvin cycle? Glucose (fuel) What additional molecule must enter the Calvin cycle to make sugar? CO2 • Calvin cycle – Three phases: • Carbon fixation (catalyzed by rubisco) • Reduction Regeneration of the CO2 acceptor (RuBP) LE 10-18_1 H2 O CO2 Input Light (Entering one CO2 at a time) 3 NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP Phase 1: Carbon fixation NADPH Rubisco O2 [CH2O] (sugar) 3 P Short-lived intermediate P P 6 3-Phosphoglycerate 3 P P Ribulose bisphosphate (RuBP) 6 6 ADP CALVIN CYCLE ATP LE 10-18_2 H2O CO2 Input Light (Entering one CO2 at a time) 3 NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP Phase 1: Carbon fixation NADPH Rubisco O2 [CH2O] (sugar) 3 P P Short-lived intermediate 3 P P 6 P 3-Phosphoglycerate Ribulose bisphosphate (RuBP) 6 ATP 6 ADP CALVIN CYCLE 6 P P 1,3-Bisphosphoglycerate 6 NADPH 6 NADP+ 6 Pi 6 P Glyceraldehyde-3-phosphate (G3P) 1 P G3P (a sugar) Output Phase 2: Reduction Glucose and other organic compounds LE 10-18_3 H2O CO2 Input Light (Entering one CO2 at a time) 3 NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP Phase 1: Carbon fixation NADPH Rubisco O2 [CH2O] (sugar) 3 P P Short-lived intermediate 3 P P 6 P 3-Phosphoglycerate Ribulose bisphosphate (RuBP) 6 ATP 6 ADP 3 ADP 3 CALVIN CYCLE 6 P ATP P 1,3-Bisphosphoglycerate 6 NADPH Phase 3: Regeneration of the CO2 acceptor (RuBP) 6 NADP+ 6 Pi P 5 G3P 6 P Glyceraldehyde-3-phosphate (G3P) 1 P G3P (a sugar) Output Glucose and other organic compounds Phase 2: Reduction I had no idea I could do these things! Alternative mechanisms of carbon fixation in hot, dry climates • How to avoid dehydration during day? close stomata Consequences? Positive & Negative conserves water but also blocks CO2 uptake Overall: reduces rate of photosynthesis LE 10-20 CAM: Crassulacean acid metabolism Sugarcane Pineapple CAM C4 CO2 Mesophyll cell Organic acid Bundlesheath cell CO2 CO2 incorporated into four-carbon Organic acid organic acids (carbon fixation) CO2 CALVIN CYCLE Sugar Spatial separation of steps CO2 Organic acids release CO2 to Calvin cycle Night Day CALVIN CYCLE Sugar Temporal separation of steps CAM Plants • CAM plants open stomata at night, incorporating CO2 into organic acids • Stomata closed during the day • CO2 released from organic acids and used in the Calvin cycle • Photosynthesis can occur during day! The Importance of Photosynthesis: A Review • sunlight stored as chemical energy in organic compounds by chloroplasts • Sugar supplies chemical energy and carbon skeletons to synthesize other organic molecules • Production of food and atmospheric oxygen LE 10-21 Light reactions Calvin cycle H2O CO2 Light NADP+ ADP + Pi RuBP Photosystem II Electron transport chain Photosystem I ATP NADPH 3-Phosphoglycerate G3P Starch (storage) Amino acids Fatty acids Chloroplast O2 Sucrose (export)