How Cells Release Stored Energy Chapter 8 8.1 Main Types of Energy-Releasing Pathways Anaerobic pathways Aerobic pathways • Evolved first • Don’t require oxygen • Start with glycolysis in cytoplasm • Completed in cytoplasm • Evolved later • Require oxygen • Start with glycolysis in cytoplasm • Completed in mitochondria Summary Equation for Aerobic Respiration C6H1206 + 6O2 6CO2 + 6H20 glucose carbon oxygen dioxide water CYTOPLASM 2 glucose ATP 4 Glycolysis e- + H+ (2 ATP net) 2 pyruvate 2 NADH e- + H + 2 CO2 Overview of Aerobic Krebs Respiration Cycle 2 NADH 8 NADH 2 FADH2 e- ATP e - + H+ e- + 4 CO2 H+ 2 Electron Transfer Phosphorylation H+ 32 ATP ATP water e- + oxygen Typical Energy Yield: 36 ATP Figure 8.3 Page 135 The Role of Coenzymes • NAD+ and FAD accept electrons and hydrogen • Become NADH and FADH2 • Deliver electrons and hydrogen to the electron transfer chain 8.2 GLYCOLYSIS Glucose • A simple sugar (C6H12O6) • Atoms held together by covalent bonds In-text figure Page 136 Glycolysis Occurs in Two Stages • Energy-requiring steps – ATP energy activates glucose and its six-carbon derivatives • Energy-releasing steps – The products of the first part are split into three- carbon pyruvate molecules – ATP and NADH form Energy-Requiring Steps of Glycolysis 2 ATP invested Energy-Requiring Steps glucose ATP ADP P glucose-6-phosphate P fructose-6-phosphate ATP ADP P P fructose1,6-bisphosphate P PGAL P PGAL Figure 8.4(2) Page 137 P NAD+ Pi P PGAL NADH NAD+ Pi PGAL NADH P P Energy1,3-bisphosphoglycerate ADP Releasing ATP P 3-phosphoglycerate Steps P P 1,3-bisphosphoglycerate ADP ATP P 3-phosphoglycerate P P 2-phosphoglycerate H2 O P 2-phosphoglycerate PEP PEP P ADP ADP ATP ATP pyruvate H2 O pyruvate Figure 8.4 Page 137 Glycolysis: Net Energy Yield Energy requiring steps: 2 ATP invested Energy releasing steps: 2 NADH formed 4 ATP formed Net yield is 2 ATP and 2 NADH 8.3 Second Stage Reactions • Preparatory reactions – Pyruvate is oxidized into two-carbon acetyl units and carbon dioxide – NAD+ is reduced • Krebs cycle – The acetyl units are oxidized to carbon dioxide – NAD+ and FAD are reduced Preparatory Reactions pyruvate coenzyme A (CoA) NAD+ NADH O CoA acetyl-CoA O carbon dioxide Krebs Cycle =CoA acetyl-CoA CoA oxaloacetate citrate NADH H2O NAD+ H2O malate NAD+ H2O FADH2 isocitrate NADH fumarate O a-ketoglutarate FAD NAD+ NADH CoA O succinate succinyl-CoA Figure 8.6 Page 139 ATP O ADP + phosphate group O The Krebs Cycle Overall Reactants Overall Products • • • • • • • • • Acetyl-CoA 3 NAD+ FAD ADP and Pi Coenzyme A 2 CO2 3 NADH FADH2 ATP Results of the Second Stage • All of the carbon molecules in pyruvate end up in carbon dioxide • Coenzymes are reduced (they pick up electrons and hydrogen) • One molecule of ATP forms • Four-carbon oxaloacetate regenerates Coenzyme Reductions during First Two Stages • Glycolysis • Preparatory reactions • Krebs cycle 2 NADH 2 FADH2 + 6 NADH • Total 2 FADH2 + 10 NADH 2 NADH 8.4 Electron Transfer Phosphorylation • Occurs in the mitochondria • Coenzymes deliver electrons to electron transfer chains • Electron transfer sets up H+ ion gradients • Flow of H+ down gradients powers ATP formation Creating an H+ Gradient OUTER COMPARTMENT NADH INNER COMPARTMENT Making ATP: Chemiosmotic Model ATP INNER COMPARTMENT ADP + Pi Importance of Oxygen • Electron transport phosphorylation requires the presence of oxygen • Oxygen withdraws spent electrons from the electron transfer chain, then combines with H+ to form water Summary of Energy Harvest (per molecule of glucose) • Glycolysis – 2 ATP formed by substrate-level phosphorylation • Krebs cycle and preparatory reactions – 2 ATP formed by substrate-level phosphorylation • Electron transport phosphorylation – 32 ATP formed Energy Harvest Varies • NADH formed in cytoplasm cannot enter mitochondrion • It delivers electrons to mitochondrial membrane • Membrane proteins shuttle electrons to NAD+ or FAD inside mitochondrion • Electrons given to FAD yield less ATP than those given to NAD+ Efficiency of Aerobic Respiration • 686 kcal of energy are released • 7.5 kcal are conserved in each ATP • When 36 ATP form, 270 kcal (36 X 7.5) are captured in ATP • Efficiency is 270 / 686 X 100 = 39 percent • Most energy is lost as heat 8.5 Anaerobic Pathways • Do not use oxygen • Produce less ATP than aerobic pathways • Two types – Fermentation pathways – Anaerobic electron transport Fermentation Pathways • Begin with glycolysis • Do not break glucose down completely to carbon dioxide and water • Yield only the 2 ATP from glycolysis • Steps that follow glycolysis serve only to regenerate NAD+ Lactate Fermentation GLYCOLYSIS C6H12O6 2 ATP energy input 2 NAD+ 2 ADP 2 4 NADH ATP energy output 2 pyruvate 2 ATP net LACTATE FORMATION electrons, hydrogen from NADH 2 lactate GLYCOLYSIS Alcoholic Fermentation C6H12O6 2 ATP energy input 2 NAD+ 2 ADP 2 4 NADH ATP 2 pyruvate energy output 2 ATP net ETHANOL FORMATION 2 H2O 2 CO2 2 acetaldehyde electrons, hydrogen from NADH 2 ethanol Anaerobic Electron Transport • Carried out by certain bacteria • Electron transfer chain is in bacterial plasma membrane • Final electron acceptor is compound from environment (such as nitrate), not oxygen • ATP yield is low FOOD fats fatty acids glycogen glycerol complex carbohydrates proteins simple sugars amino acids glucose-6-phosphate NH3 GLYCOLYSIS PGAL pyruvate acetyl-CoA 8.6 ALTERNATIVE ENERGY SOURCES Figure 8.11 Page 145 KREBS CYCLE urea carbon backbones Evolution of Metabolic Pathways • When life originated, atmosphere had little oxygen • Earliest organisms used anaerobic pathways • Later, noncyclic pathway of photosynthesis increased atmospheric oxygen • Cells arose that used oxygen as final acceptor in electron transport 8.7 Processes Are Linked sunlight energy PHOTOSYNTHESIS water + carbon dioxide sugar molecules oxygen AEROBIC RESPIRATION In-text figure Page 146