10/25/2010 CHAPTER 9 CELLULAR RESPIRATION. Life is Work. Types of cellular respiration. Catabolic pathways = oxidizing fuels

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1 CHAPTER 9 CELLULAR RESPIRATION Life is Work Living cells require transfusions of energy from outside sources to perform their many tasks: Chemical work Transport work Mechanical work Energy stored in the organic molecules of food ultimately comes from the sun Energy flows into an ecosystem as sunlight and leaves as heat In contrast, chemical elements essential to life are recycled Photosynthesis generates oxygen and organic molecules Respiration breaks this fuel down, generating ATP In this chapter, we will focus on the key pathways of aerobic respiration: Glycolysis Citric acid cycle Oxidative phosphorylation Catabolic pathways = oxidizing fuels Catabolic pathways = metabolic pathways that release stored energy by breaking down complex molecules Compounds that can participate in exergonic reactions can act as fuels Types of cellular respiration Aerobic Respiration ( cellular respiration ): Oxygen is consumed as a reactant along with the organic fuel Cells of most eukaryotes and prokaryotes With the help of enzymes, a cell can systematically degrade macromolecules that are rich in potential energy Energy taken out of chemical storage can be used to do work Anaerobic Respiration: Example: fermentation Degradation of organic fuel without the use of oxygen Cells of some prokaryotes and some eukaryotes 1

2 Organic compound Overall process: Oxygen Carbon dioxide Water We will learn the steps of cellular respiration by tracking the degradation of the sugar glucose: C 6 H 12 O O 2 6 CO H 2 O + Energy (ATP, heat) Glucose is the fuel that cells most often use Exergonic process where G = -686 Kcal/mol Energy The Stages of Cellular Respiration Glycolysis: Technically not part of cellular respiration Catabolic pathway that occurs in cytosol Breaks down glucose to two smaller molecules Dehydrogenases transfer e- from NAD + to NADH Citric Acid Cycle Takes place in the mitochondria (matrix) eukaryots Takes place in cytosol in prokaryots Dehydrogenases transfer e- from NAD + to NADH Oxidative phosphorylation: electron transport and chemiosmosis Accepts e- from NADH (H + + e-) Pass through many molecules and the end it is combined with oxygen to form water The energy released makes ATP REDOX REACTIONS Loss of electrons = oxidation Gain of electrons = reduction (reduce + charge) Reducing agent = electron donor Oxidizing agent = electron acceptor NAD + and NADH NAD+ is an electron carrier (a coenzyme) It acts as an oxidizing agent during respiration Oxidation = loss of electrons Oxidizing agent = takes away electrons Dehydrogenase removes a pair of hydrogen atoms (2e - and 2p + ) from the substrate and oxidizes it The enzyme delivers 2 e - and 1p + to the coenzyme (NAD + ) 2 H H H + The other p + (H + ) is released to the surrounding solution 2

3 GLYCOLYSIS Oxidizes glucose to pyruvate Glucose (6 C sugar) is split to two 3 C sugars Pyruvate Two phases: Energy investment Energy payoff Glycolysis Overall: Oxygen is not necessary CO 2 is not released NET ATP YIELD = 2 ATP + 2 NADH 3

4 Citric Acid Cycle Glycolysis releases less than a 1/4 of the chemical energy stored in glucose Most of the energy remains in the two molecules l of pyruvate If oxygen is available, pyruvate enters a mitochondrion (eukaryotes) Active transport Acetyl CoA is very unstable (potential energy? Ender/exergonic?) Citric Acid Cycle = Krebs Cycle (1930s) Oxidizes organic fuel derived from pyruvate The cycle generates: 1 ATP 3 NADH 1 FADH2 1 NADH (junction) 4

5 Oxidative phosphorylation and chemiosmosis So far, glycolysis and the citric acid cycle have produced only 4 ATP molecules per glucose molecule (substrate-level phosphorylation) 2 ATP from glycolysis + 2 NADH 2 ATP from citric acid cycle + 4 NADH + FADH 2 NADH and FADH 2 molecules account for most of the energy extracted from the glucose molecule Electron Transport The electron transport chain is a collection of molecules embedded in the inner membrane of the mitochondrion (eukaryotic cells) Most components are proteins, which exist in complexes (I IV) Other non protein components are essential for the functioning of enzymes, they are called prosthetic groups Electrons from NADH are transferred to first molecule: flavoprotein (flavin mononucleotide) Iron-sulfer protein Coenzyme Q Cytochromes (heme prosthetic groups) Oxygen (very electronegative) Why not release all the energy in just one step? 2 H 2 O 2 2 H 2 O + energy Chemiosmosis: energy-coupling mechanism Many copies of ATP synthase populate the inner membrane of the mitochondrion It uses energy of an existing ion gradient to power ATP synthesis In the mitochondrial inner membrane there is a pump that creates a difference in H+ concentration between the matrix and the intermembrane space Also considered a difference in ph 5

6 Chemiosmosis Process in which energy stored in the form of a hydrogen ion gradient across a membrane is used to drive cellular work, such as the synthesis of ATP ATP synthase is a multisubunit complex with four main parts: Stator Rotor Internal Rod Catalytic knob To read: Fig Inquiry How do we maintain the H+ gradient? That is the function of the electron transport chain The exergonic flow of electrons from NADH (or FADH 2 ) down to oxygen is used to pump H+ across the membrane ATP synthase is the only route through the membrane (for H+) Certain members of the electron transport chain accept and release protons (H+) The H+ gradient that results is referred to as protonmotive force Chemiosmosis is an energy-coupling mechanism that uses energy stored in the form of an H+ gradient across a membrane to drive cellular work Examples: Chloroplasts: use it to generate ATP during phothosynthesis Prokaryotes: use it to make ATP and also to rotate their flagella and pump nutrients and waste across the membrane Peter Mitchell was awarded the Nobel Prize in 1978 for originally proposing the chemiosmotic model 6

7 ATP production? During respiration, most energy flows: Glucose > NADH > ET Chain > Proton MF > ATP Electron shuttles MITOCHONDRION CYTOSOL span membrane 2 NADH or 2 FADH 2 2 NADH 2 NADH 6 NADH 2 FADH 2 1 NADH = 1 pair electrons = pumps 10 H+ 3 4 H+ = 1 ATP 1 NADH = ATP = ~ 3 1 FADH 2 = ATP = ~ 2 Glycolysis 2 Glucose Pyruvate 2 Acetyl CoA Maximum per glucose: Citric acid cycle About 36 or 38 ATP Oxidative phosphorylation: electron transport and chemiosmosis + 2 ATP + 2 ATP + about 32 or 34 ATP READ: p Reasons that affect ATP yield Fermentation and Anaerobic Respiration The estimated ATP yield from aerobic respiration is contingent on an adequate supply of oxygen to the cell However, cells can oxidize organic fuel and generate ATP without the use of oxygen: Anaerobic respiration Fermentation Anaerobic Respiration Takes place in certain prokaryotic organisms (environments without O 2 ) These organisms have electron transport chains, but the end product is a different molecule Ex: SO 2-4 H 2 S is a by-product (instead of water) Fermentation It harvests chemical energy without using oxygen or any electron transport chain Glycolysis oxidizes glucose to two molecules of pyruvate (oxidizing agent NAD + ) Glycolysis is exergonic and makes ATP by substratelevel phosphorylation Fermentation expands glycolysis so that it continuously generates ATP Fermentation There must be a sufficient supply of NAD + Electrons can be transferred to pyruvate instead Types of fermentation (end product): Alcohol Fermentation Lactic Acid Fermentation How?? 7

8 Piruvate is converted to ethanol in two steps: Release of CO 2 = acetaldehyde Acetaldehyde is reduced d by NADH = ethanol Ex: yeast: brewing, winemaking and baking Alcohol fermentation Lactic Acid Fermentation Pyruvate is reduced directly by NADH to form lactate No release of CO 2 Used in the dairy insudstry to make yogurt and cheese Human muscles (when O 2 is scarce) Obligate / Facultative Obligate Anaerobes: Some organisms carry out only fermentation or anaerobic respiration They cannot survive in the presence of oxygen Facultative Anaerobes: Yeasts and many bacteria can make enough ATP to survive using either fermentation or respiration Fermentation and Aerobic Respiration Compared Both processes use glycolysis to oxidize glucose and other organic fuels to pyruvate The processes have different final electron acceptors: an organic molecule (such as pyruvate or acetaldehyde) in fermentation and O 2 in cellular respiration Cellular respiration produces 38 ATP per glucose molecule; fermentation produces 2 ATP per glucose molecule Our muscle cells behave as facultative anaerobes Fig Glucose Obligate anaerobes carry out fermentation or anaerobic respiration and cannot survive in the presence of O 2 Yeast and many bacteria are facultative anaerobes, meaning that they can survive using either fermentation or cellular respiration CYTOSOL No O 2 present: Fermentation Pyruvate Glycolysis O 2 present: Aerobic cellular respiration In a facultative anaerobe, pyruvate is a fork in the metabolic road that leads to two alternative catabolic routes Ethanol or lactate MITOCHONDRION Acetyl CoA Citric acid cycle 8

9 The Evolutionary Significance of Glycolysis Glycolysis occurs in nearly all organisms Oldest fossil bacteria date to ~3.5 bya But atmospheric O 2 only started to accumulate ~ 27bya 2.7 Glycolysis probably evolved in ancient prokaryotes before there was oxygen in the atmosphere Other Catabolic pathways Free glucose molecules are not common in the diets of humans and other animals Most our calories com from fats, proteins, sucrose and starch All these molecules can be used by cellular respiration to make ATP Carbohydrates: Starch can be hydrolyzed Glycogen can also be hydrolyzed Proteins: First they are digested (aa) Aa can be used to build proteins or can be converted to intermediates of glycolysis and the Krebs cycle (deamination + enzymes) Fats: Digested to glycerol and fatty acids Glycerol = glyceraldehyde-3-phosphate (glycolysis) Fatty acids via Beta oxidation = 2 C fragments = enter Krebs cycle as Acetyl CoA (NADH and FADH2 also generated) Anabolic Pathways Cells need substance as well as energy Food must also provide the atoms necessary to make new molecules Intermediates of glycolysis and the citric acid cycle can be diverted into anabolic pathways as precursors for other molecules Humans can make about half of the 20 essential aa, the rest come from diet Example Dihydroxyacetone phosphate (intermediate compound generated during glycolysis) can be converted to one of the precursors of fat So, when we eat more food than we need, we store fat even if our diet is fat-free. Metabolism is remarkably versatile and adaptable 9

10 Feedback Mechanisms Feedback inhibition is the most common control mechanism for anabolic pathways An end product of the anabolic pathway inhibits the enzyme that catalyzes an early step in the pathway Catabolism is also regulated through positive and negative feedback Phosphofructokinase Important switch in metabolic control The cell can speed up or slow down the entire catabolic process by controlling the first step of the glycolytic pathway Phosphofructokinase is an allosteric enzyme with receptor sites for specific inhibitors and activators: Inhibited by ATP Stimulated by AMP Also sensitive to citrate (1 st product Citric Acid Cycle) 10

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