Chapter 8 Cellular Respiration 1 8.1 Cellular Respiration • Cellular respiration is a cellular process that breaks down nutrient molecules with the concomitant production of ATP • Consumes oxygen and produces carbon dioxide (CO2) Cellular respiration is an aerobic process. • Usually involves the breakdown of glucose to CO2 and H2O Energy is extracted from the glucose molecule: • Released step-wise • Allows ATP to be produced efficiently Oxidation-reduction enzymes include NAD+ and FAD as coenzymes 2 Cellular Respiration NAD+ (nicotinamide adenine dinucleotide) A coenzyme of oxidation-reduction. It can: • Oxidize a metabolite by accepting electrons • Reduce a metabolite by giving up electrons Each NAD+ molecule is used over and over again • FAD (flavin adenine dinucleotide) Also a coenzyme of oxidation-reduction Sometimes used instead of NAD+ Accepts two electrons and two hydrogen ions (H+) to become FADH2 3 Cellular Respiration • Cellular respiration includes four phases: Glycolysis is the breakdown of glucose into two molecules of pyruvate • Occurs in the cytoplasm • ATP is formed • Does not utilize oxygen Preparatory (prep) reaction • Both pyruvates are oxidized and enter the mitochondria • Electron energy is stored in NADH • Two carbons are released as CO2 (one from each pyruvate) 4 Cellular Respiration Citric acid cycle • Occurs in the matrix of the mitochondrion and produces NADH and FADH2 • In series of reactions, it releases 4 carbons as CO2 • Turns twice per glucose molecule (once for each pyruvate) • Produces two immediate ATP molecules per glucose molecule Electron transport chain (ETC) • Extracts energy from NADH & FADH2 • Passes electrons from higher to lower energy states • Produces 32 or 34 molecules of ATP 5 The Four Phases of Complete Glucose Breakdown Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. NADH e– e– NADH e– e– Cytoplasm e– NADH and FADH2 e– e– Glycolysis Citric acid cycle Preparatory reaction glucose Mitochondrion pyruvate Electron transport chain and chemiosmosis 2 ATP 2 ATP 4 ADP 4 ATP total 2 ATP net gain 2 ADP 2 ATP 32 ADP or 34 32 or 34 ATP 6 8.2 Outside the Mitochondria: Glycolysis • Glycolysis occurs in cytoplasm outside mitochondria • Energy Investment Step: Two ATP are used to activate glucose Glucose splits into two G3P molecules • Energy Harvesting Step: Oxidation of G3P occurs by removal of electrons and hydrogen ions Two electrons and one hydrogen ion are accepted by NAD+ resulting in two NADH Four ATP are produced by substrate-level phosphorylation Net gain of two ATP (4 ATP produced - 2 ATP consumed) Both G3Ps converted to pyruvates 7 Outside the Mitochondria: Glycolysis Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Glycolysis inputs outputs 6C glucose 2 NAD+ 2 2 (3C) pyruvate 2 NADH 2 ADP ATP 4 ADP + 4 P 2 4 ATP ATP total net gain 8 Glycolysis Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Glycolysis Energy-investment Step glucose –2 ATP ATP ATP ADP G3P glyceraldehyde-3-phosphate BPG 1,3-bisphosphoglycerate 3PG 3-phosphoglycerate ADP Two ATP are used to get started. Splitting produces two 3-carbon molecules. G3P Energy-harvesting Steps NAD+ G3P NAD+ E1 NADH NADH BPG ADP Oxidation of G3P occurs as NAD+ receives high-energy electrons. BPG ADP E2 Substrate-level ATP synthesis. +2 ATP ATP ATP 3PG 3PG E3 H2O H2O PEP ADP +2 2 ATP PEP ADP E4 ATP ATP (net gain) Oxidation of 3PG occurs by removal of water. Substrate-level ATP synthesis. ATP pyruvate pyruvate Two molecules of pyruvate are the end products of glycolysis. 8.3 Outside the Mitochondria: Fermentation • Pyruvate is a pivotal metabolite in cellular respiration • If O2 is not available to the cell, fermentation, an anaerobic process, occurs in the cytoplasm. During fermentation, glucose is incompletely metabolized to lactate, or to CO2 and alcohol (depending on the organism). • If O2 is available to the cell, pyruvate enters the mitochondria for aerobic respiration. 10 Outside the Mitochondria: Fermentation • Fermentation is an anaerobic process that reduces pyruvate to either lactate or alcohol and CO2 NADH transfers its electrons to pyruvate Alcoholic fermentation, carried out by yeasts, produces carbon dioxide and ethyl alcohol • • • Lactic acid fermentation, carried out by certain bacteria and fungi, produces lactic acid (lactate) • Used in the production of alcoholic spirits and breads. Used commercially in the production of cheese, yogurt, and sauerkraut. Other bacteria produce chemicals anaerobically, including isopropanol, butyric acid, proprionic acid, and acetic acid. 11 Outside the Mitochondria: Fermentation • Advantages Provides a quick burst of ATP energy for muscular activity. • Disadvantages Lactate and alcohol are toxic to cells. Lactate changes pH and causes muscles to fatigue. • Oxygen debt Yeast die from the alcohol they produce by fermentation • Efficiency of Fermentation Only 2 ATP per glucose are produced, compared to 36 or 38 ATP molecules per glucose produced by cellular respiration. 12 Outside the Mitochondria: Fermentation Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Fermentation inputs outputs glucose 2 ADP + 2 2 lactate or 2 alcohol and 2 CO2 P 2 ATP net gain 13 8.4 Inside the Mitochondria • The Preparatory (prep) Reaction Connects glycolysis to the citric acid cycle End product of glycolysis, pyruvate, enters the mitochondrial matrix Pyruvate is converted to a 2-carbon acetyl group • Attached to Coenzyme A to form acetyl-CoA • Electrons are picked up (as hydrogen atom) by NAD+ • CO2 is released and transported out of mitochondria into the cytoplasm 14 Inside the Mitochondria Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 2 NAD+ O OH C 2 C O + 2 CoA CH 3 pyruvate 2 pyruvate + 2 CoA 2 NADH CoA 2 C O + 2 CO2 CH 3 carbon acetyl CoA dioxide 2 acetyl CoA + 2 carbon dioxide 15 Mitochondrion Structure and Function Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Cristae: location of the electron transportchain (ETC) Matrix: location of the prep reaction and the citric acid cycle outer membrane inner membrane cristae intermembrane space matrix 45,0003X © Dr. Donald Fawcett and Dr. Porter/Visuals Unlimited 16 Inside the Mitochondria • Citric Acid Cycle Also called the Krebs cycle Occurs in the matrix of mitochondria Begins with the addition of a two-carbon acetyl group (from acetyl-CoA) to a four-carbon molecule (oxaloacetate), forming a six-carbon molecule (citric acid) NADH and FADH2 capture energy rich electrons ATP is formed by substrate-level phosphorylation Turns twice for one glucose molecule (once for each pyruvate) Produces 4 CO2, 2 ATP, 6 NADH and 2 FADH2 per glucose molecule 17 Citric Acid Cycle Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. NADH e– NADH e– e– e– e– e– NADHand FADH2 e– Glycolysis Preparatory reaction glucose Electron transport chain and chemiosmosis Citric acid cycle pyruvate Matrix 2 ATP 2 ADP 4 ADP 2 4 ATP total ATP net 2ADP 2 ATP 32ADP or 34 32 or 34 ATP CoA 1. The cycle begins when a C2 acetyl group carried by CoA combines with a C4 molecule to form citrate. acetyl CoA C2 citrate C6 NAD NADH oxaloacetate C2 NADH 5. Once again a substrate is oxidized, and NAD+ is reduced to NADH. 2. Twice over, substrates are oxidized as NAD+ is Reduced to NADH, and CO2 is released. Citric acid cycle NAD+ CO2 fumarate C4 ketoglutarate C5 NAD+ succinate C4 FAD 4. Again a substrate is oxidized, but this time FAD is reduced to FADH2 CO2 NADH FADH ATP 3. ATP is produced as an energized phosphate is transferred from a substrate to ADP. Inside the Mitochondria Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Citric acid cycle inputs outputs 2 (2c) acetyl groups 4 CO2 6 NAD+ 6 NADH 2 FADH2 2 FAD 2 ADP +2 P 2 ATP 19 Inside the Mitochondria • Electron Transport Chain (ETC) • Location: Eukaryotes: cristae of the mitochondria Aerobic prokaryotes: plasma membrane • Series of carrier molecules: Pass energy-rich electrons successively from one to another Complex arrays of protein and cytochrome • Cytochromes are proteins with heme groups with central iron atoms • The electron transport chain Receives electrons from NADH & FADH2 Produces ATP by oxidative phosphorylation • Oxygen serves as the final electron acceptor Oxygen combines with hydrogen ions to form water 20 Inside the Mitochondria • The fate of the hydrogens: Hydrogens from NADH deliver enough energy to make 3 ATPs Those from FADH2 have only enough for 2 ATPs “Spent” hydrogens combine with oxygen • Recycling of coenzymes increases efficiency Once NADH delivers hydrogens, it returns (as NAD+) to pick up more hydrogens However, hydrogens must be combined with oxygen to make water If O2 is not present, NADH cannot release H+ No longer recycled back to NAD+ 21 Inside the Mitochondria • The electron transport chain complexes pump H+ from the matrix into the intermembrane space of the mitochondrion • H+ therefore becomes more concentrated in the intermembrane space, creating an electrochemical gradient. • ATP synthase allows H+ to flow down its gradient. • Flow of H+ drives the synthesis of ATP from ADP and inorganic phosphate by ATP synthase. • This process is called chemiosmosis ATP production is linked to the establishment of the H+ gradient • ATP moves out of mitochondria and is used for cellular work It can be broken down to ADP and inorganic phosphate These molecules are returned to the mitochondria for more ATP production 22 Organization and Function of the Electron Transport Chain Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. e- NADH NADH e- e- e- NADH and FADH2 ee- eGlycolysis glucose Electron transport chain and chemiosmosis Citric acid cycle Preparatory reaction pyruvate 2ATP 2ADP 4 ADP 4 ATP total 2 ATP net 2 H+ ATP Electron transport chain NADH-Q reductase H+ 32 or 34 32 ADP or 34 ADP H+ H+ H+ H+ cytochrome reductase H+ H+ H+ cytochrome c coenzyme Q H+ cytochrome oxidase H+ H+ e: H+ FADH2 high energy electron H+ NADH NAD+ FAD + 2 H+ low energy electron e: H+ H+ 2 H+ H+ H+ H+ ATP H2O ADP + P 1O 2 2 H+ H+ H+ Matrix H+ Intermembrane space H+ H+ H+ ATP synthase + complex H H+ H+ ATP channel protein ATP H+ Chemiosmosis H+ H+ H+ H+ Inside the Mitochondria • Energy yield from glucose metabolism: Net yield per glucose: • From glycolysis – 2 ATP • From citric acid cycle – 2 ATP • From electron transport chain – 32 or 34 ATP Energy content: • Reactant (glucose) 686 kcal • Energy yield (36 ATP) 263 kcal • Efficiency is 39% • The rest of the energy from glucose is lost as heat 24 Accounting of Energy Yield per Glucose Molecule Breakdown Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 2 net glycolysis ATP 2 NADH 2 NADH 6 NADH 2 FADH2 2 pyruvate Mitochondrion 2 acetyl CoA 2 CO2 2 ATP 4 or 6 ATP 6 ATP 18 ATP 4 ATP subtotal 32 or 34 ATP Electron transport chain Cytoplasm glucose Citric acid cycle 4 CO2 6 O2 subtotal 4 ATP 36 or 38 total ATP 6 H2O 25 8.5 Metabolic Pool • Foods: Sources of energy rich molecules Carbohydrates, fats, and proteins • Degradative reactions (Catabolism) break down molecules Tend to be exergonic (release energy) • Synthetic reactions (anabolism) build molecules Tend to be endergonic (consume energy) 26 The Metabolic Pool Concept Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. proteins carbohydrates amino acids glucose Glycolysis fats glycerol fatty acids ATP pyruvate acetyl CoA Citric acid cycle ATP Electron transport chain ATP © C Squared Studios/Getty Images. 27 Metabolic Pool • All metabolic compounds are part of the metabolic pool • Intermediates from respiratory pathways can be used for anabolism • Anabolism (synthetic reactions of metabolism): Carbohydrates • Start with acetyl-CoA • Basically reverses glycolysis (but different pathway) Fats • G3P converted to glycerol • Acetyl groups are connected in pairs to form fatty acids 28 Metabolic Pool • Anabolism (cont.): Proteins: • Made up of combinations of 20 different amino acids • Some amino acids (11) can be synthesized by adult humans • However, other amino acids (9) cannot be synthesized by humans – Essential amino acids – Must be present in the diet 29 Photosynthesis vs. Cellular Respiration Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Photosynthesis grana enzymes membrane H2O NADPH O2 CO2 O2 ATP NAD+ H2O NADH Cellular Respiration CH2O membrane CH2O enzyme-catalyzed reactions ATP production via chemiosmosis ADP NADP+ CO2 cristae enzymes 30
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