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Photosynthesis: Energy From Sunlight The Ingredients For Photosynthesis Sunlight Sugars Leaf Stem Root Leaves: Plant Solar Collectors Stoma Plant Cell Structure A PLANT CELL Free ribosomes Nucleolus Nucleus Peroxisome Smooth endoplasmic reticulum Rough endoplasmic reticulum Plasma membrane Plasmodesmata Chloroplast Mitochondrion Golgi apparatus The Plant Cell Wall Cell wall of cell 1 Middle lamella Interior of cell 1 Plasma membrane Interior of cell 2 Cell wall of cell 2 Plant Cell Structure Free ribosomes A PLANT CELL Cell wall Vacuole Peroxisome Nucleolus Nucleus Rough endoplasmic reticulum Smooth endoplasmic reticulum Plasma membrane Plasmodesmata Chloroplast Mitochondrion Golgi apparatus The Photosynthetic Reactions Are Carried Out By Chloroplasts Chloroplast Structure Inner membrane Outer membrane Thylakoid Stroma Granum (stack of thylakoids) Photosynthesis Is A Two-Step Process Chloroplast Light (photon) ELECTRON TRANSPORT Thylakoid Light reactions Chlorophyl ATP CYCLE Light‐ independent reactions NADPH CYCLE CALVIN CYCLE Stroma The Light Reactions Light Reactions The Electromagnetic Spectrum Action Spectrum Of Photosynthesis Chlorophyll Figure 10.5 Exciting An Atom Increasing energy Excited state Absorption of photon by molecule Photon Ground state Photon Electron Nucleus Ground state Excited state Photoexcitation Of Chlorophyll Chlorophyll Molecules Are Found In Large Protein Complexes (CHO in chlorophyll b) Chlorophyll molecules Proteins Stroma Thylakoid membrane Thylakoid lumen Photosystems Consist Of Two Parts 1. Photosynthetic Reaction Center 2. Antenna Complex Plants And Cyanobacteria Possess Two Photosystems Far‐red light on Off Red light on Off Both lights on Off Comparative Photosystem Organization Red, photochemical reaction center molecules; green, light harvesting pigments Note: PsaA and PsaB are like D2-Core antenna protein and D1-Core antenna protein fusions, respectively Photosystems Consist Of Two Parts 1. Photosynthetic Reaction Center 2. Antenna Complex Organization Of Pigment Molecules Summary Electron Transfer Reactions Within the Reaction Center Figure 14-45 Molecular Biology of the Cell (© Garland Science 2008) Electron Transfer Reactions Within A Purple Bacterial Photosystem Need to capture 2 photons of light to reduce Quinone Figure 14-46 Molecular Biology of the Cell (© Garland Science 2008) Comparative Photosystem Organization Red, photochemical reaction center molecules; green, light harvesting pigments Note: PsaA and PsaB are like D2-Core antenna protein and D1-Core antenna protein fusions, respectively Plant Photosystem II Structure PSII is a dimer Each monomer contains: 16 Integral membrane protein subunits 3 subunits in the lumen 3 subunits in the lumen 36 chlorohylls 7 carotenoids 2 pheophytins 2 hemes 2 plastoquinones 1 Mn water splitting complex Figure 14-47a,c Molecular Biology of the Cell (© Garland Science 2008) Plant Photosystem II Structure Figure 14-47b Molecular Biology of the Cell (© Garland Science 2008) Light Reactions Take Place On Thylakoid Membranes Chloroplast Thylakoid Electron Transport Pathway Of Photosynthesis Mobile Carriers: Plastoquinone (PQ; organic molecule) and Plastocyanine (PC, Protein) Large Complexes: Photosystem II, Photosystem I, Cytochrome B6-f Complex (Cyt) Thylakoid interior ELECTRON TRANSPORT (high concentration of H+) H2O 1/2 O2 ATP SYNTHESIS PC ATP synthase 2 PQ Cyt Photon Photosystem II Stroma (low concentration of H+) Photon Fd Photosystem I NADP Reductase Topology Of Chloroplast Photosynthetic Components Figure 14-48 Molecular Biology of the Cell (© Garland Science 2008) Energetics Of Non-Cyclic Photophosphorylation Figure 14-49 Molecular Biology of the Cell (© Garland Science 2008) Cyclic And Non-Cyclic Photophosphorylation PS I Cyclic Photophosphorylation PS II Figure 14-73 (part 2 of 3) Molecular Biology of the Cell (© Garland Science 2008) Chemiosmotic ATP Generation ATP Synthase Figure 14-15 Molecular Biology of the Cell (© Garland Science 2008) Chemiosmotic ATP Synthesis Figure 14-19 Molecular Biology of the Cell (© Garland Science 2008) Non-Cyclic Photophosphorylation Summary 4 Stages: 1. 2. 3. 4. Light energy is used to energize an electron that can drive electron transport system Electron transport system generates proton gradient across thylakoid membrane Electron transport system reduces NADP+ Proton gradient is used to chemiosmotically generate ATP Dark Reactions Ribulose 1-5-Bisphosphate Carboxylase-Oxygenase (RuBisCO): The First Step Of The Dark Reactions (RuBP) Figure 14-39 Molecular Biology of the Cell (© Garland Science 2008) (Unstable) The First Stable Product Containing Fixed Carbon From Carbon Dioxide Carboxyl group 3‐Phosphoglycerate (3PG) The Dark Reactions Photon Thylakoid ELECTRON TRANSPORT ATP CYCLE Stroma CALVIN CYCLE NADPH CYCLE The Calvin Cycle The Calvin Cycle START 6 RuBP Carbon fixation 6 RuMP 12 3PG CALVIN CYCLE Regeneration Reduction and of RuBP sugar production 10 G3P 12 G3P 2 G3P Sugars Other carbon compounds Used For All Plant Carbon Metabolism Glyceraldehyde 3‐phosphate (G3P) Overview Of Photosynthetic Reactions Metabolic Interactions In A Plant Cell Lipids (triglycerides) Glycerol 3PG RuBP CALVIN CYCLE RuMP Nucleotides Nucleic acids Fatty acids Pyruvate G3P Acetyl CoA GLYCOLYSIS GLUCONEOGENESIS Hexose‐P Sucrose Polysaccharides (e.g., starch, cellulose) Metabolic Interactions In A Plant Cell Nitrogen from soil Acetyl CoA NH4+ α‐ketoglutarate CITRIC ACID CYCLE Amino acids Proteins Energy Losses During Photosynthesis ENERGY LOSS SUNLIGHT 100% 50% Wavelengths of light not part of absorption spectrum of photosynthetic pigments (e.g., green light) 30% Light energy not absorbed due to plant structure (e.g., leaves not properly oriented to sun) 10% Inefficiency of light reactions converting light to chemical energy Wavelengths of light that can be absorbed by photosynthetic pigments Light energy absorbed by photosynthetic pigments Chemical energy available for CO2 fixation Chemical energy stored Chemical energy stored in carbohydrates in carbohydrates 5% 5% 5% Inefficiency of CO2 fixation pathways Net chemical energy stored in carbohydrates from sunlight: 5% Photorespiration Carbon lost Carbon gained 2‐PG Carboxylase reaction Photorespiration Rubisco Oxygenase reaction Calvin cycle 3‐PGA C3 and C4 Leaves Have Different Anatomy The Anatomy and Biochemistry of C4 Carbon Fixation Mesophyll cell Mesophyll cell Bundle sheath cell The Anatomy and Biochemistry of C4 Carbon Fixation Plasma membrane Cell wall PEP Mesophyll cell Carboxy‐ lation C4 cycle 4C compound Bundle sheath cell Regener‐ ation 3C compound 4C compound 3C compound Decarb‐ oxylation 5C sugar Regener‐ ation Carboxy‐ Calvin Cycle lation Triose‐P 3C sugar Note: the Calvin Cycle that occurs in Bundle Sheath Cells is the standard dark reactions occurring in all plants The CAM C4 Pathway