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Chapter 9 Cellular Respiration: Harvesting Chemical Energy Light energy ECOSYSTEM Photosynthesis in chloroplasts Organic molecules CO2 + H2O Cellular respiration in mitochondria ATP powers most cellular work Heat + O2 Ask a Cell Biologist: What is Cellular Respiration? Cellular respiration is the aerobic harvesting of energy from organic molecules It is a catabolic pathway It contains mostly exergonic reactions that release energy Summary Equation for Cellular Respiration C6H12O6 + 6O2 glucose oxygen 6CO2 + 6H2O + ATP + Heat Carbon dioxide Greater number of bonds = Greater potential energy O=C=O carbon dioxide glucose Oxidation-Reduction Reactions Oxidation-Reduction (Redox) reactions Transfer electrons from one reactant to another by oxidation and reduction Oxidation-Reduction Reactions In oxidation A substance LOSES electrons, or is oxidized In reduction A substance GAINS electrons, or is reduced Example of Redox Reaction becomes oxidized (loses electron) becomes reduced (gains electron) Oxidation of Organic Fuel During cellular respiration Glucose is oxidized and oxygen is reduced It is the movement of hydrogen atoms and their electrons from glucose that are important BECOMES OXIDIZED C6H12O6 + 6O2 6CO2 + 6H2O + Energy BECOMES REDUCED Mitochondrion: Site of Cellular Respiration Mitochondrion Intermembrane space Outer membrane Free ribosomes in the mitochondrial matrix Inner membrane Cristae Matrix Mitochondrial DNA 100 µm Stages of Cellular Respiration Glycolysis Breaks down glucose into two molecules of pyruvate (pyruvic acid) Citric acid cycle (Krebs Cycle) Completes the breakdown of energy originally in glucose • Electron transport chain Generates lots of ATP Glycolysis Glucose Pyruvate CYTOSOL ATP Pyruvate oxidation Acetyl CoA Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis MITOCHONDRION ATP ATP Glycolysis Glycolysis Means “splitting of sugar” Breaks down glucose into pyruvate Occurs in the cytoplasm of the cell Ancient pathway No oxygen required! Glycolysis: 2 Phases Glycolysis ATP Citric acid cycle Oxidative phosphorylation ATP ATP Energy investment phase Glucose 2 ADP + 2 P 2 ATP used Energy payoff phase 4 ADP + 4 P 2 NAD+ + 4 e- + 4 H + 4 ATP formed 2 NADH + 2 H+ 2 Pyruvate + 2 H2O Glucose 4 ATP formed – 2 ATP used 2 NAD+ + 4 e– + 4 H + 2 Pyruvate + 2 H2O 2 ATP 2 NADH+ 2 H+ Gradual Harvesting of Energy Electrons from organic compounds Are usually first transferred to NAD+, an electron shuttle NAD+ becomes reduced to NADH as it accepts electrons and H’s from glucose Dehydrogenase is an enzyme which helps move the electrons by removing 2 H atoms (and their electrons) from glucose and giving them to NAD+ NAD+ + 2H NADH + H+ Dehydrogenase Hydrogen Ions = H+ = Proton HYDROGEN ATOM Substrate Level Phosphorylation • Both glycolysis and the citric acid cycle • Can generate ATP by substrate-level phosphorylation ENZYME ENZYME ADP P SUBSTRATE + PRODUCT ATP Citric Acid Cycle The citric acid cycle completes the energy-yielding oxidation of organic molecules The citric acid cycle Takes place in the matrix of the mitochondrion Pre-citric Acid Cycle Before the citric acid cycle can begin Pyruvate must first be converted to acetyl CoA, which links the cycle to glycolysis CYTOSOL MITOCHONDRION NAD+ NADH + H+ O– S CoA C O 2 C C O O 1 3 CH3 Pyruvate Transport protein CH3 Acetyl CoA CO2 Coenzyme A Citric Acid Cycle Pyruvate Glycolysis (from glycolysis, 2 molecules per glucose) ATP Citric acid cycle Oxidative phosphorylation ATP ATP CO2 CoA NADH + 3 H+ Acetyl CoA CoA CoA Oxaloacetate (4 C) FADH2 Citrate (6 C) Citric acid cycle 2 CO2 3 NAD+ 3 NADH FAD+ + 3 H+ ADP + P i ATP Electron Transport Chain Mitochondrial inner membrane proteins (carrier molecules) pass electrons in a series of steps instead of in one explosive reaction Uses the energy from the electron transfer to form ATP Electron Transport Chain At the end of the chain This makes oxygen the final electron acceptor NADH FADH2 Free energy (G) relative to O2 (kcl/mol) Electrons are passed to oxygen, forming water 50 40 FMN I FE•S CARRIER MOLECULES FAD FE•S II O III Cyt b 30 FE•S Cyt c1 IV Cyt c Cyt a Cyt a3 20 10 oxygen 0 2 H + + 12 O2 H2 O Chemiosmosis Chemiosmosis and the electron transport chain Oxidative phosphorylation. electron transport and chemiosmosis Glycolysis ATP Inner Mitochondrial membrane ATP ATP H+ H+ H+ Intermembrane space Protein complex of electron carners Q I Inner mitochondrial membrane IV III ATP synthase II FADH2 FAD+ NAD+ NADH Mitochondrial matrix H+ Cyt c 2 H+ + 1/2 O2 H2O ADP + (Carrying electrons from, food) ATP Pi H+ Chemiosmosis Electron transport chain + ATP synthesis powered by the flow Electron transport and pumping of protons (H ), + + which create an H gradient across the membrane Of H back across the membrane Oxidative phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Chemiosmosis At certain steps along the electron transport chain Electron transfer causes protein complexes to pump H+ from the mitochondrial matrix to the intermembrane space The resulting H+ gradient Stores energy Drives chemiosmosis in ATP synthase But what’s Chemiosmosis really? Chemiosmosis Is an energy-coupling mechanism that uses energy in the form of a H+ gradient across a membrane to drive cellular work Is referred to as a proton-motive force Chemiosmosis ATP synthase INTERMEMBRANE SPACE H + H+ Is the enzyme that actually makes ATP H+ H+ H+ H+ H+ H+ ADP + Pi MITOCHONDRIAL MATRIX ATP Following the Electrons (Energy) During respiration, most energy flows in this sequence Glucose to NADH to electron transport chain to chemiosmosis to ATP About 40% of the energy in a glucose molecule Is transferred to ATP during cellular respiration, making approximately 32 ATP Review of Stages of Cellular Respiration Electron shuttles span membrane 2 NADH Glycolysis 2 Pyruvate Glucose MITOCHONDRION 2 NADH or 2 FADH2 2 NADH Pyruvate oxidation 2 Acetyl CoA 2 ATP Maximum per glucose: CYTOSOL 6 NADH 2 FADH2 Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis 2 ATP about 26 or 28 ATP About 30 or 32 ATP So what happens when there is little or no O2 ? •Without O2, the ETC will cease to operate •BUT glycolysis couples with fermentation to still produce 2 ATP’s •Without the ETC however, fermentation needs an alternate way to generate NAD+ Fermentation • Two common types of fermentation are: • Alcohol fermentation • Lactic acid fermentation Alcohol Fermentation Pyruvate is converted to ethanol in two steps 1. Releases CO2 from pyruvate making acetaldehyde 2. Acetaldehyde reduced by NADH to ethanol which regenerates NAD+ • Many bacteria undergo alcohol fermentation in anaerobic conditions • Alcohol fermentation by yeast is used in brewing, winemaking, and baking Lactic Acid Fermentation • Pyruvate directly reduced by NADH to form lactate (ionized form of lactic acid) •No CO2 released • Fungi and bacteria undergo lactic acid fermentation •Used to make cheese and yogurt • Human muscle cells will undergo lactic acid fermentation when O2 scarce during heavy exercise Summary of Types of Fermentation 2 ADP 2 P Glucose 2 ADP 2 P 2 ATP i Glycolysis Glucose i 2 ATP Glycolysis 2 Pyruvate 2 NAD 2 Ethanol (a) Alcohol fermentation 2 NADH 2 H 2 NAD 2 CO2 2 Acetaldehyde 2 NADH 2 H 2 Pyruvate 2 Lactate (b) Lactic acid fermentation Fermentation Overview Glucose Glycolysis CYTOSOL Pyruvate No O2 present: Fermentation O2 present: Aerobic cellular respiration MITOCHONDRION Ethanol, lactate, or other products Pyruvate as a key juncture in catabolism Acetyl CoA Citric acid cycle