Chapter 7: Cellular Respiration and Fermentation Key Concepts 7.1 Catabolic pathways yield energy by oxidizing organic fuels 7.2 Glycolysis harvests chemical energy by oxidizing glucose to pyruvate 7.3 After pyruvate is oxidized, the citric acid cycle completes the energy-yielding oxidation of organic molecules 7.4 During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis 7.5 Fermentation and anaerobic respiration enable cells to produce ATP without oxygen 7.6 Glycolysis and citric acid cycle connect to many other metabolic pathways © 2016 Pearson Education, Inc. Life Is Work Living cells require energy from outside sources Some animals, like this giraffe, obtain energy by eating plants, and some animals feed on other organisms that eat plants How Do These Leaves Power the Work of Life for This Giraffe? © 2016 Pearson Education, Inc. Life Is Work, Continued Concept 7.1: Catabolic pathways yield energy by oxidizing organic fuels Energy flows into an ecosystem as sunlight and leaves as heat Photosynthesis generates oxygen and organic molecules, which are used as fuel for cellular respiration Cells use chemical energy stored in organic molecules to regenerate ATP, which powers work Catabolic pathways involving electron transfer are central processes to cellular respiration Catabolic Pathways and Production of ATP Energy Flow and Chemical Recycling in Ecosystems © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. Catabolic Pathways and Production of ATP, Continued Cellular respiration includes both aerobic and anaerobic processes but is often used to refer to aerobic respiration Although organic compounds like carbohydrates, fats, and proteins are all consumed as fuel, Organic compounds + O2 The breakdown of organic molecules is exergonic Fermentation is a partial degradation of sugars that occurs without oxygen Aerobic respiration consumes organic molecules and oxygen to synthesize ATP Anaerobic respiration consumes organic molecules, but not oxygen to synthesize ATP CO2 + H2O + Energy (ATP + Heat) Cellular Respiration: A Catabolic Pathway and Production of ATP -Complete breakdown of glucose in the presence of oxygen. -Releases energy in the form of ATP - C6H12O6 6 O2 → 6 CO2 6 H2O Energy (ATPheat) It is helpful to trace it with the sugar glucose C6H12O6 + 6O2 6CO2 + 6H2O + Energy (ATP + Heat) C6H12O6 Glucose 6 O2 Oxygen 6 CO2 Carbon dioxide 6 H 2O ATP Water Energy Campbell Essential Biology, 5e Simon/Dickey/Reece Pearson © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. 1 Redox Reactions: Oxidation and Reduction Oxidation of Organic Fuel Molecules During Cellular Respiration In oxidation Glucose is oxidized and oxygen is reduced A substance loses electrons or hydrogen, or is oxidized In reduction Oxidation A substance gains electrons or hydrogen, or is reduced Glucose loses electrons (and hydrogens) C6H12O6 Glucose 6 O2 Oxygen 6 CO2 Carbon dioxide 6 H 2O The electron donor is called the reducing agent The electron receptor is called the oxidizing agent OILRIG Water Reduction Oxygen gains electrons (and hydrogens) Campbell Essential Biology, 5e Simon/Dickey/Reece Pearson © 2016 Pearson Education, Inc. Chemical energy is stored in reduced molecules of carbohydrates and lipids The shared electrons of C-C and C-H bonds of carbohydrates and lipids have high potential energy (chemical energy) So oxidation of glucose releases large amount of energy. In oxidation of glucose, electrons always travel with hydrogen atoms. In biological systems, hydrogen atoms are used as a marker for electrons to follow redox reactions. © 2016 Pearson Education, Inc. Oxidation of Organic Fuel Molecules During cellular respiration © 2016 Pearson Education, Inc. Redox Reactions: Oxidation and Reduction The transfer of electrons during chemical reactions releases energy stored in organic molecules This released energy is ultimately used to synthesize ATP © 2016 Pearson Education, Inc. Figure 9.4 NAD+ as an electron carrier During cellular respiration, fuel (such as glucose) is oxidized, and oxygen is reduced Organic molecules with an abundance of hydrogen, like carbohydrates and fats, are excellent fuels As hydrogen (with its electron) is transferred to oxygen, energy is released that can be used in ATP synthesis 2 e− + 2 H+ NAD+ Dehydrogenase 2 e− + H+ NADH H+ Reduction of NAD+ 2[H] (from food) Oxidation of NADH Nicotinamide (oxidized form) H+ Nicotinamide (reduced form) Enzymes called dehydrogenases facilitate the transfer of two electrons and one hydrogen ion to NAD+ One hydrogen ion is released in this process © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. 2 The Electron Carrier NAD+ empty loaded proton (oxidized) empty goes to pick up more electrons used in later stage of respiration (reduced) used in later stage of respiration Electron or hydrogen carriers: NAD+ and FAD Dehydrogenase Two important electron carriers in the cell are: • NAD+ (Nicotinamide Adenine Dinucleotide) NAD+ + 2H NADH Oxidized form + H+ Reduced form • FAD ( Flavin Adenine Dinucleotide) FAD + 2H FADH2 Oxidized form Reduced form Figure 9.UN04 © 2016 Pearson Education, Inc. th Ed, Pearson Biology, A guide to the natural world. D. Krough, 5 Stepwise energy Harvest via NAD+ and the Electron Transport Chain © 2016 Pearson Education, Inc. Figure 6.5 e e Electrons from food e e NADH NAD Stepwise release of energy used to make Cellular respiration Oxidizes glucose in a series of steps 2 H ATP 2 e Electrons from organic compounds Are first transferred to NAD+ ( Nicotinamide Adenine Dinucleotide), a coenzyme NADH, the reduced form of NAD+ Passes the electrons to the electron transport chain 2 Hydrogen, electrons, and oxygen combine to produce water © 2016 Pearson Education, Inc. Stepwise energy Harvest via NAD+ and the Electron Transport Chain, Continued-1 e 2 H 1 2 O2 H2 O © 2016 Pearson Education, Inc. An Overview of cellular respiration NADH passes the electrons to the electron transport chain Electrons are passed to increasingly electronegative carrier molecules down the chain through a series of redox reactions Electron transfer to oxygen occurs in a series of energyreleasing steps instead of one explosive reaction The energy yielded is used to regenerate ATP © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. 3 Color Code for Stages of cellular respiration The Stages of Cellular Respiration Glycolysis (cytoplasm) 1. 2. 1. Glycolysis (color-coded blue throughout the chapter) 2. PYRUVATE OXIDATION and the CITRIC ACID CYCLE (color-coded orange) Pyruvate Oxidation 3. Citric acid cycle or Krebs cycle 3. OXIDATIVE PHOSPHORYLATION: Electron transport and chemiosmosis (color-coded purple) 4. © 2016 Pearson Education, Inc. Complete breakdown of glucose matrix of mitochondria matrix of mitochondria Oxidative phosphorylation or Electron Transport chain (accounts for most ATP synthesis in the inner membrane of mitochondria) © 2016 Pearson Education, Inc. Mini-Map, Glycolysis What is Glycolysis? Concept 7.2: Glycolysis harvests chemical energy by oxidizing glucose to pyruvate Glycolysis Means “splitting of sugar” Breaks down glucose into two molecules of pyruvate, 2ATP (net) and 2NADH Involves 10 enzyme catalyzed reactions Occurs in the cytosol of the cell in the presence or absence of oxygen How glycolysis works © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. Figure 7.9-1b-s3 Figure 7.9-2a-s3 GLYCOLYSIS: Energy Investment Phase GLYCOLYSIS: Energy Payoff Phase Glyceraldehyde 3-phosphate (G3P) Fructose ATP 6-phosphate Glyceraldehyde 3-phosphate (G3P) Fructose 1,6-bisphosphate Isomerase © 2016 Pearson © 2016 Pearson Education, Inc. 2 ADP 2 H 2 2 ADP Phosphofructokinase 2 ATP 2 NADH 2 NAD Aldolase Dihydroxyacetone phosphate (DHAP) Isomerase Aldolase Dihydroxyacetone phosphate (DHAP) Triose phosphate 2 dehydrogenase Phosphoglycerokinase Pi 1,3-Bisphosphoglycerate 3-Phosphoglycerate © 2016 Pearson © 2016 Pearson Education, Inc. 4 Figure 7.9-2b-s3 The Stages of cellular respiration: A Preview, Continued GLYCOLYSIS: Energy Payoff Phase 2 H2O 2 2 Phosphoglyceromutase 3-Phosphoglycerate 2 2 ADP ATP 2 Pyruvate kinase Enolase 2-Phosphoglycerate 2 Phosphoenolpyruvate (PEP) Pyruvate © 2016 Pearson © 2016 Pearson Education, Inc. Glycolysis: The energy Input and Output Summary Oxidative phosphorylation accounts for almost 90% of the ATP generated by cellular respiration This process involves the transfer of inorganic phosphates to ADP A smaller amount of ATP is formed in Glycolysis and the citric acid cycle by substrate-level phosphorylation In this process, an enzyme transfers a phosphate group directly from a substrate molecule to ADP For each molecule of glucose degraded to CO2 and water by respiration, the cell makes up to 32 molecules of ATP © 2016 Pearson Education, Inc. Substrate-Level Phosphorylation Energy investment phase: ATP provides activation energy by phosphorylating glucose -2 ATP per glucose Energy payoff phase: ATP is produced by substrate-level phosphorylation and NAD+ is reduced to NADH. + 4ATP + 2NADH per glucose _____________________________ Net : 2ATP + 2NADH per glucose © 2016 Pearson Education, Inc. Which statement about glycolysis is true? © 2016 Pearson Education, Inc. Mini-Map, Pyruvate Oxidation A. It splits water. B. It uses oxygen. C. It occurs in the cytoplasm. D. It makes the most ATP compared to the two other steps. E. It makes lipids. © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. 5 An Overview of Pyruvate Oxidation and the Citric Acid Cycle An Overview of Pyruvate Oxidation 2Pyruvates © 2016 Pearson Education, Inc. Concept 7.3: Connecting step: Oxidation of Pyruvate to Acetyl CoA 2CO2 + 2NADH + 2Acetyl CoA © 2016 Pearson Education, Inc. An Overview of the Citric Acid Cycle • In the presence of O2, pyruvate enters the mitochondrion • One carbon is lost in the form of CO2 • Acetyl group (CH3OC-) combines with Coenzyme A • Acetyl Coenzyme A (CoA) links glycolysis to citric acid cycle • Products are 2Acetyl CoA, 2CO2 and 2NADH • Pyruvate Oxidation © 2016 Pearson Education, Inc. Mini-Map, Citric Acid Cycle © 2016 Pearson Education, Inc. Concept 7.3 Citric acid cycle or Krebs cycle • Takes place in the matrix of mitochondria • 2 molecules of Acetyl CoA enter the cycle • Acetyl CoA combines with oxaloacetate and forms citrate • For each glucose there are 2 cycles • Products for 2cycles:2 ATP, 4 CO2, 6 NADH, 2FADH2 • Citric acid cycle © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. 6 Citric acid cycle contd. • Citric acid cycle has 8 steps, each catalyzed by an enzyme • Acetyl CoA combines with oxaloacetate and forms citrate • Citrate then gets converted into oxaloacetate in the next seven step • NADH & FADH2 transfer the electrons extracted from food to the ETC © 2016 Pearson Education, Inc. Which statement about the citric acid cycle is true? A Closer Look at the Citric Acid Cycle and Summary Each cycle produces: • 1 ATP by substrate-level phosphorylation • 3 NADH • 1 FADH2 (electron carrier) • 2 CO2 For each glucose there are two cycle: • 2 ATP by substrate-level phosphorylation • 6 NADH • 2 FADH2 (electron carrier) • 4 CO2 © 2016 Pearson Education, Inc. Mini-Map, Oxidative Phosphorylation A. It occurs during the movement from the cytosol through the mitochondrial membranes. B. It makes ATP through substrate-level phosphorylation. C. It makes the most ATP compared to the two other steps. D. It occurs in the cytoplasm. E. It splits glucose. ©©2016 2016Pearson PearsonEducation, Education,Inc. Inc. Chemiosmosis Couples the Electron Transport Chain to ATP Synthesis The electron transport chain (ETC) is located in the mitochondrial inner membrane. Electrons are passed from electron donors to acceptors until they reach the final electron acceptor, oxygen. © 2016 Pearson Education, Inc. Electron Transport Chain (ETC) • Electron carriers NADH and FADH2 donate electrons to the electron transport chain • Electrons are passed through a number of proteins including cytochromes • Energy released during electron transfer is used to pumps H+ ions (protons) from the matrix into the intermembrane space. Energy is then stored in the H+ ion gradient. • H+ ions move back into the matrix through ATP synthase. The energy of the proton gradient activates the enzyme. • ATP synthase uses the exergonic flow of H+ ions to drive the phosphorylation of ADP to make ATP © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. 7 Free-energy Change During Electron Transport • Most of the chain’s components are proteins, which exist in multiprotein complexes • The carriers alternate reduced and oxidized states as they accept and donate electrons • At the end of ETC, electrons are finally accepted by oxygen, which combines with H+ ions to form water © 2016 Pearson Education, Inc. • Following glycolysis and the citric acid cycle, NADH and FADH2 account for most of the energy extracted from food • These two electron carriers donate electrons to the electron transport chain, which powers ATP synthesis via oxidative phosphorylation Electron Transport Chain © 2016 Pearson Education, Inc. ATP Synthase, A Molecular Mill ATP synthase is the enzyme that synthesizes ATP by adding phosphate to ADP © 2016 Pearson Education, Inc. The energetic electron, taken from glucose or a breakdown product of glucose, is stripped of its energy to A. actively transport H+ into the intermembrane space. B. actively transport NAD+ into the intermembrane space. C. actively transport Na+ into the matrix. D. power facilitated diffusion of H+ into the matrix. E. actively transport H+ into the matrix. © 2016 Pearson Education, Inc. During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis ATP Synthesis in Electron Transport Chain • Chemiosmosis: Energy for ATP synthesis is provided by the diffusion of H+ ions • Oxidative phosphorylation: Energy for ATP synthesis is provided by oxidation of food © 2016 Pearson Education, Inc. ATP Yield Per Molecule of glucose at Each Stage of cellular respiration © 2016 Pearson Education, Inc. 8 ATP production during cellular respiration Pyruvate as a Key Juncture in Catabolism • During cellular respiration, most energy flows in this sequence: Glucose NADH electron transport chain H+ ions ATP – 1 NADH → 2.5 ATP (10NADH → 25ATP) – 1FADH2 → 1.5 ATP (2FADH2 → 3ATP) – 28ATP in ETC in the mitochondria • About 34% of the energy in a glucose molecule – Is transferred to ATP during cellular respiration – About 32 ATP molecules are formed when glucose is completely broken down © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. Concept 7.5: Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen • In the absence of oxygen cells carry out Fermentation of Glucose which consists of - Glycolysis and - Reactions that regenerate NAD+ which can be used by glycolysis © 2016 Pearson Education, Inc. Lactic Acid Fermentation Alcohol Fermentation • In alcohol fermentation, pyruvate is converted to ethanol (alcohol) in two steps, – The first step releases CO2 – The second step produces ethanol • Alcohol fermentation by yeast is used in brewing, winemaking, and baking © 2016 Pearson Education, Inc. Types of Fermentation • In lactic acid fermentation – Pyruvate is converted into lactic acid • Lactic acid fermentation by some fungi and bacteria is used to make cheese and yogurt • Human muscle cells use lactic acid fermentation to generate ATP when O2 is scarce © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. 9 Comparing Fermentation with Anaerobic and Aerobic Respiration Glycolysis Cellular Respiration Fermentation Location Cytoplasm Mitochondria Cytoplasm Electron carriers NAD+ NAD+, FAD NAD+ Oxygen Pyruvate or acetaldehyde 32ATP 2ATP (glycolysis) Final electron acceptor ATP produced 2ATP © 2016 Pearson Education, Inc. Comparing Fermentation with Anaerobic and Aerobic Respiration Obligate anaerobes carry out anaerobic respiration or fermentation and cannot survive in the presence of O2 Facultative anaerobes carry out anaerobic respiration or fermentation and can survive in the presence of O2 , Example - Yeast and many bacteria, they can survive using either fermentation or cellular respiration In a facultative anaerobe, pyruvate is a fork in the metabolic road that leads to two alternative catabolic routes © 2016 Pearson Education, Inc. What is the purpose of fermentation reactions? What is the purpose of fermentation reactions? to regenerate NAD+ so glycolysis can continue to make alcohol or lactic acid that cells can metabolize for energy under anaerobic conditions to make additional ATP when respiration can’t make ATP fast enough to slow down cellular oxygen consumption when oxygen is scarce to make organic molecules that cells can store until oxygen becomes available to regenerate NAD+ so glycolysis can continue to make alcohol or lactic acid that cells can metabolize for energy under anaerobic conditions to make additional ATP when respiration can’t make ATP fast enough to slow down cellular oxygen consumption when oxygen is scarce to make organic molecules that cells can store until oxygen becomes available © 2016 Pearson © 2016 Pearson Education, Inc. The Evolutionary Significance of Glycolysis © 2016 Pearson © 2016 Pearson Education, Inc. The Catabolism of Various Molecules From Food Glycolysis is the most common metabolic pathway among organisms on Earth, indicating that it evolved early in the history of life Early prokaryotes may have generated ATP exclusively through Glycolysis due to the low oxygen content in the atmosphere The location of Glycolysis in the cytosol also indicates its ancient origins Eukaryotic cells with mitochondria evolved much later than prokaryotic cells © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. 10 Concept 7.6: Glycolysis and the citric acid cycle connect to many other metabolic pathways The Versatility of Catabolism Catabolic pathways break down proteins, fats and carbohydrates. Their monomers can be metabolized by glycolysis or another stage of cellular respiration Proteins are digested into amino acids and can then enter glycolysis or the citric acid cycle. Fats are digested to glycerol (used in glycolysis) and fatty acids (used in generating acetyl CoA) Fatty acids are broken down by beta oxidation and yield acetyl CoA © 2016 Pearson Education, Inc. Test Your Understanding, Question 9 (Regulation of Phosphofructokinase) Phosphofructokinase is an enzyme that is part of glycolysis. Under which condition is the enzyme active and why? © 2016 Pearson Education, Inc. Summary of Key Concepts: Citric Acid Cycle Biosynthesis (Anabolic Pathways) The body uses small molecules to build other substances Some of these small molecules come directly from food; others can be produced during Glycolysis or the citric acid cycle © 2016 Pearson Education, Inc. Summary of Key Concepts: Glycolysis © 2016 Pearson Education, Inc. You should now be able to: Explain in general terms how redox reactions are involved in energy exchanges. What reactant is oxidized and what reactant is reduced in cellular respiration 3. Name the three(four) stages of cellular respiration; for each, state the part of the eukaryotic cell where it occurs, the reactants and the products 4. What is substrate level phosphorylation? 5. In general terms, explain the role of the electron transport chain in cellular respiration 6. What is chemiosmosis? 7. Why is the process in the electron transport chain called oxidative phosphorylation? 8. Explain where and how the respiratory electron transport chain creates a proton gradient 9. What is the role of oxygen in cellular respiration? 10. What is the total number of ATP molecules produced when one glucose molecule is broken down? 11. Distinguish between fermentation and anaerobic respiration 12. Distinguish between obligate and facultative anaerobes 1. 2. © 2016 Pearson Education, Inc. © 2016 Pearson Education, Inc. 11