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Ch 9 Cellular Respiration Extracting usable energy from organic molecules • Energy flows into ecosystems as sunlight • Photosynthesis generates organic molecules and O2 . • Cellular respiration breaks down organic molecules to generate ATP;uses O2. ------>fuels cellular activities LE 9-2 Light energy ECOSYSTEM Photosynthesis in chloroplasts Organic + O molecules 2 CO2 + H2O Cellular respiration in mitochondria ATP powers most cellular work Heat energy If eukaryotes have mitochondria are they capable of of aerobic respiration? Does this include protists, fungi, plants and animals? You bet! Cellular Respiration Net Reaction: C6H12O6 + 6O2 6CO2 + 6H2O + ATP glucose Energy molecule Redox reaction C6H12O6 oxidized to CO2 O2 reduced to H2O Is cellular respiration endergonic or exergonic? Anabolic or catabolic? Exergonic (exothermic) Catabolic (breaks down molecules to release energy) LE 9-6_1 Cytosol Intermembrane space Cristae Glycolysis Matrix Pyruvate Glucose Mitochondrion Cytosol Inner membrane Outer membrane ATP Substrate-level phosphorylation Mitochondrion Three major steps in cellular respiration 1. Glycolysis: the breaking of sugar In cytosol 2. Citric acid cycle (Krebs Cycle) In mitochondria 3. Oxidative phosphorylation LE 9-6_1 Glycolysis Pyruvate Glucose Cytosol Mitochondrion ATP Substrate-level phosphorylation Glycolysis 6C organic molecule-->--> 2(3C) organic molecule 1 glucose --> 2 pyruvates Is the conversion of glucose to pyruvate a simple one step process? NO Requires 10 enzyme catalyzed reactions! LE 9-9a_1 Glucose ATP Hexokinase ADP Glucose-6-phosphate Glycolysis Citric acid cycle ATP ATP Oxidation phosphorylation ATP LE 9-9a_2 Glucose ATP Hexokinase ADP Glucose-6-phosphate Phosphoglucoisomerase Fructose-6-phosphate ATP Phosphofructokinase ADP Fructose1, 6-bisphosphate Aldolase Isomerase Dihydroxyacetone phosphate Glyceraldehyde3-phosphate Glycolysis Citric acid cycle ATP ATP Oxidation phosphorylation ATP LE 9-9b_1 2 NAD+ Triose phosphate dehydrogenase 2 NADH + 2 H+ 1, 3-Bisphosphoglycerate 2 ADP Phosphoglycerokinase 2 ATP 3-Phosphoglycerate Phosphoglyceromutase 2-Phosphoglycerate LE 9-9b_2 2 NAD+ Triose phosphate dehydrogenase 2 NADH + 2 H+ 1, 3-Bisphosphoglycerate 2 ADP Phosphoglycerokinase 2 ATP 3-Phosphoglycerate Phosphoglyceromutase 2-Phosphoglycerate 2 H2O Enolase Phosphoenolpyruvate 2 ADP Pyruvate kinase 2 ATP Pyruvate LE 9-8 Energy investment phase Glucose 2 ATP used 2 ADP + 2 P Glycolysis Citric acid cycle Oxidative phosphorylation Energy payoff phase ATP ATP ATP 4 ADP + 4 P 2 NAD+ + 4 e– + 4 H+ 4 ATP formed 2 NADH + 2 H+ 2 Pyruvate + 2 H2O Net Glucose 4 ATP formed – 2 ATP used 2 NAD+ + 4 e– + 4 H+ 2 Pyruvate + 2 H2O 2 ATP 2 NADH + 2 H+ What is NAD+? NAD+ + e- + H- --> NADH An electron carrier LE 9-6_2 2. Citric acid cycle (aka Krebs cycle) Glycolysis Glucose Pyruvate Citric acid cycle Cytosol ATP Substrate-level phosphorylation ATP Substrate-level phosphorylation Mitochondrion Reactions in the Mitochondrion Conversion of pyruvate (3C) to acetyl coA (2C) In what compartment? Matrix LE 9-10 MITOCHONDRION CYTOSOL NAD+ NADH + H+ Acetyl Co A Pyruvate Transport protein CO2 Coenzyme A LE 9-11 Pyruvate (from glycolysis, 2 molecules per glucose) CO2 NAD+ Glycolysis Citric acid cycle ATP ATP Oxidation phosphorylation CoA Where is this pathway taking place? NADH + H+ Acetyl CoA ATP CoA Citric acid cycle e- carrier FADH2 2 CO2 3 NAD+ 3 NADH + 3 H+ FAD ADP + P i ATP e- carrier Third step Oxidative phosphorylation 1. Electron transport chain 2. Chemiosmosis In what compartment? Inner mitochondrial membrane LE 9-13 NADH 50 Free energy (G) relative to O2 (kcal/mol) FADH2 40 FMN I Multiprotein complexes FAD Fe•S II Fe•S Q III Cyt b 30 Fe•S Cyt c1 Glycolysis Citric acid cycle ATP ATP Oxidative phosphorylation: electron transport and chemiosmosis IV Cyt c Cyt a ATP Cyt a3 20 10 Oxygen final e- acceptor, most electronegative 0 2 H+ O2 + 1/2 H2O LE 9-15 Oxidative phosphorylation Glycolysis Citric acid cycle ATP ATP Oxidative phosphorylation: electron transport and chemiosmosis ATP H+ H+ H+ H+ Intermembrane space Cyt c Protein complex of electron carriers Q IV III I ATP synthase II Inner mitochondrial membrane FADH2 NADH + H+ 2H+ + 1/2 O2 NAD+ (carrying electrons from food) Mitochondrial matrix H2O FAD ATP ADP + P i H+ Electron transport chain Pumping of protons (H+), creates H+ gradient across the membrane Chemiosmosis ATP synthesis powered by diffusion of H+ into matrix LE 9-14 INTERMEMBRANE SPACE H+ H+ H+ H+ H+ H+ A rotor within the membrane spins as shown when H+ flows past it down the H+ gradient. H+ A stator anchored in the membrane holds the knob stationary. A rod (or “stalk”) extending into the knob also spins, activating catalytic sites in the knob. H+ ADP + P ATP i MITOCHONDRAL MATRIX Three catalytic sites in the stationary knob join inorganic phosphate to ADP to make ATP. LE 9-16 Electron shuttles span membrane CYTOSOL 2 NADH Glycolysis Glucose 2 Pyruvate MITOCHONDRION 2 NADH or 2 FADH2 2 NADH 2 Acetyl CoA + 2 ATP 6 NADH Citric acid cycle + 2 ATP by substrate-level phosphorylation by substrate-level phosphorylation Maximum per glucose: About 36 or 38 ATP 2 FADH2 Oxidative phosphorylation: electron transport and chemiosmosis + about 32 or 34 ATP by oxidation phosphorylation, depending on which shuttle transports electrons from NADH in cytosol 3 ATP/NADH 2 ATP/ FADH2 Fermentation production of ATP anaerobically (without O2) How? Glycolysis only (fermentation) • Two fermentation pathways – alcohol fermentation – lactic acid fermentation LE 9-18 Glucose CYTOSOL Pyruvate No O2 present Fermentation O2 present Cellular respiration MITOCHONDRION Ethanol or lactate Acetyl CoA Citric acid cycle LE 9-17a 2 ADP + 2 P i Glucose 2 ATP Glycolysis 2 Pyruvate 2 NAD+ 2 Ethanol Alcohol fermentation 2 NADH + 2 H+ 2 CO2 2 Acetaldehyde LE 9-17b 2 ADP + 2 P i Glucose 2 ATP Glycolysis 2 NAD+ 2 NADH + 2 H+ 2 CO2 2 Pyruvate 2 Lactate Lactic acid fermentation Under what circumstances does anaerobic respiration or fermentation occur? In O2-starved muscles accumulation of lactic acid painful: toxic In some micro-organisms: bacteria, yeast exploited for production of alcoholic beverages cheeses Evolutionary Significance of Glycolysis • Glycolysis occurs in nearly all organisms • May have evolved in ancient prokaryotes before oxygen in atmosphere Regulation of ATP synthesis Negative and positive feedback loops LE 9-20 Glucose AMP Glycolysis Fructose-6-phosphate – Stimulates + Phosphofructokinase – Fructose-1,6-bisphosphate Inhibits Inhibits Pyruvate ATP Citrate Acetyl CoA Citric acid cycle Oxidative phosphorylation The Versatility of Catabolism • Catabolic pathways – funnel electrons from many organic molecules into cellular respiration • Glycolysis – accepts wide range of carbohydrates • Proteins – digested to amino acids; amino groups feed glycolysis or the citric acid cycle • Fats – digested to glycerol (used in glycolysis) and fatty acids (used in generating acetyl CoA) LE 9-19 Proteins Carbohydrates Amino acids Sugars Glycerol Fatty acids Glycolysis Glucose Glyceraldehyde-3- P NH3 Fats Pyruvate Acetyl CoA Citric acid cycle Oxidative phosphorylation