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1 Electron Transport and Oxidative Phosphorylation Reading Assignment: Start reading Chapter 14: Energy Generation in Mitochondria and Cholorplasts See animation 14.3 on your text CD ATPsynthase animation and lots of other stuff 1

2 What does the citric acid accomplish? Carbons in pyruvate are fully oxidized to CO 2 Some GTPs (which may be later converted to ATPs) are generated by substrate level phosphorylation 8 NADH and 2 FADH 2 are stockpiled NADH and FADH 2 are high energy molecules and they can be used as reducing agents by the cell. But much of the stored energy is not directly accessible to the cell in this form What happens in the mitochondria to convert the potential energy in NADH into the form of ATP? 2

3 NADH and FADH 2 are high energy molecules: As electrons drop from the top to the bottom of the scale, energy is released Substances vary in their tendancy to become oxidized or reduced. This tendancy is expressed as the reduction potential. As electrons drop from the top to the bottom of the scale, energy is released. Glucose has a reduction potential of (-0.43 V) so the exergonic oxidation of glucose is coupled to the endergonic reduction of NAD + 3

4 How to power ATP synthesis build a dam pile up protons on one side poke a hole -- use the rush of protons through the hole to turn a turbine which then makes ATP 4

5 The inner mitochondrial membrane is 70% protein and 30% phospholipid by weight An electron micrograph of the inside surface of the inner mitochondrial membrane in a plant cell. Densely packed particles are visible -- due to protruding portions of ATP synthase and the respiratory enzyme complexes 5

6 A Closer look at the inner mitochondrial membrane ATP synthase ATP Transporter The inner mt membrane is 70% protein and 30% phospholipid by weight many of the proteins belong to the electron transport chain also includes ATP synthase: converts the stored potential energy in the electrochemical proton gradient into chemical energy matrix space between inner membranes 6

7 THE ELECTRON TRANSPORT CHAINS CONSISTS OF MEMBRANE ASSOCIATED ELECTRON CARRIERS These systems have two basic functions: 1. to accept electrons from an electron donor and to transfer them to an electron acceptor 2. to conserve some of the energy released during electron tranasfer for the synthesis of ATP The electron transport chain reoxidizes the coenzymes NADH and FADH and channels the free energy into the synthesis of ATP NADH and FADH gained electrons when oxidizing other compounds they transfer these electrons to the electrontransport chain: electron transport chain reoxidizes the coenzymes NADH and FADH and channels the free energy into the synthesis of ATP 7

8 NADH is a high energy molecule: Brock 5.19 Reduction potential of the components of the electron transport chain of the mitochondrion of eukaryotic cells and the plasma membrane of some bacterial cells. By breaking up the complete oxidaton into a series of discrete steps, energy recapture is possible Substances vary in their tendancy to become oxidized or reduced. This tendancy is expressed as the reduction potential. As electrons drop from the top to the bottom of the scale, energy is released. NOTICE the molecule at the bottom! 8

9 How to power ATP synthesis build a dam pile up protons on one side poke a hole -- use the rush of protons through the hole to turn a turbine which then makes ATP Oxidative Phosphorylation: the production of ATP using energy derived from the redox reactions of an electron transport chain Chemiosmosis: the production of ATP from ADP using the energy of hydrogen ion gradients 9

10 H-bonding is always with us proton wires: net translocation of protons can occur over a long distance through a protein by hopping between pairs of hydrogen bonded donor and acceptor residues; a string of such residues connected by hydrogen bonds can be thought of as a proton wire... Matrix (inside the inner membrane of the mt) is above the membrane (gray bar). The intermembrane space is below the membrane. animation of electron transport How protons can be pumped across membranes: As an electron passes along an electron-transport chain embedded in a lipidbilayer, it can bind and release a proton at each step. In this diagram electron carrier B picks up a proton (H + ) from one side of the membrane when it accepts an electron from carrier A. It releases the proton to the other side of the membrane when it donates its electron to carrier C 10

11 ALBERTS animation 14.2 The energy released is used to transport H + ions across the inner mitochondrial membrane to the space between the two membranes In this way, a gradient of H + ions is maintained across the inner membrane This gradient serves as a source of energy (like a battery) that is tapped to drive a variety of energy-requiring reactions The most prominent of these reactions is the generation of ATP ADP + P i -----> ATP 11

12 How to power ATP synthesis build a dam pile up protons on one side poke a hole -- use the rush of protons through the hole to turn a turbine which then makes ATP 12

13 Potential energy in gradient converted to mechanical energy which via conformtional changes in the cytoplasmic portion of the ATPase is converted to chemical energy in ATP ATP synthase is imbedded in the inner mitochondrial membrane below & left in this figure is the matrix of the mitochondria (compartment contained within the inner mitochondrial membrane) water soluble catalytic domain is in matrix spinning ion transport channel (embedded in lipid bilayer) 13

14 ATP synthase: enzyme that uses energy from the proton gradient to produce ATP from ADP + Pi inner mitochondrial membrane of all eukaryotic cells the thylakoid membrane of chloroplasts of plant cells the plasma membrane of prokaryotic cells 14

15 Now for some serious quaternary structure! ATP synthase energy converter. The enzyme consists of two rotary motors, F0 and F1 which are coupled via their drive shafts. The transmembrane F0 motor has one a, two b and nine to twelve c subunits. The soluble F1 motor has three α and three β subunits, and one each of the other subunits. During ATP synthesis, F0 channels protons across the membrane to drive rotation. Nature 410: 878 4/19/01 The rotating subunits are the c polypeptide in F o and the γ polypeptide in F 1 The rotation of F o (caused by movement of protons) drives the rotation of γ This rotation drives the conformational transitions of the catalytic subunits which, in turn, alters the nucleotide binding site affinities. As a consequence, conformational energy flows from the catalytic subunit into the bound ADP and Pi to promote their dehydration into ATP. ALBERTS animation Step 32 in this animation substrate level ATP synthesis in glycolysis 15

16 MORE optional Stuff on ATP synthase for those who are amused by this protein: Look at First two links ATP synthase do cross section alpha, beta gamma ATP synthase This cartoon is adapted from fig. 2 of Cross. The 3 shades of red represent the 3 different conformational states of the catalytic subunits. The central asymmetric black object represents the gamma subunit which is caused to rotate by themitochondrial proton efflux. This rotation drives the conformational transitions of the catalytic subunits which, in turn,alters the nucleotide binding site affinities. As a consequence, conformational energy flows from the catalytic subunit into the bound ADP and Pi to promote their dehydration into ATP. ATP synthase: the rotory engine in the cell: in vitro rotation of an actin filament attached to ATP synthase 16

17 What fraction of the potential energy can a respiring cell extract from a glucose molecule? A few billion years of evolution have ensured that the aerobic system is 40-54% efficient 17

18 The complete oxidation of 1 mole of glucose generates about 38 moles of ATP (synthesized from ADP) ATP yield from complete oxidation of glucose Process Direct product Final ATP Glycolysis Pyruvate oxidation (2 per glucose) Acetyl CoA oxidation (Citric Acid cycle) two per glucose Total ATP yield per glucose molecule 2 NADH (cytosolic) 2 ATP 2 NADH (mitochondrial matrix) 6 NADH (mitochondrial matrix) 2 FADH 2 2ATP or 2 GTP 3 or 5* 2 5** *** * depends on which "shuttle system" transfers reducing equivalents into the mitochondria ** 3 protons per ATP *** This number varies from reference to reference.. 18

19 Why is ATP high energy ATP has stored potential energy: ATP ADP + P i + energy ΔG o = kcal/mole exergonic reaction (corresponds to an Keq of >10 5 Under cellular conditions, the hydrolysis of ATP creates two molecules of much lower energy and releases a great deal of usable energy The phosphates in ATP can be considered to exist in an activated state: the presence of four negative charges in close proximity destabilizes the molecule -- electrostatic repulsion between negative charges favors hydrolysis Increased hydration ADP and P -- energetically favored Release of a phosphate increases entropy because the PO 4 molecule released is capable of resonance forms (delocalized proton and oxygen binding) not possible when phosphate is bound to another molecule 19

20 20

21 What do cells do with ATP? 21

22 Drives anabolic (endergonic) chemical reactions Used to do work (move stuff around molecular motors) Used for active transport to make/maintain ion and solute gradients Lots of other roles 22

23 Cells drive active transport in three main ways 23

24 Remember lysosomes? Lysosomes contain hydrolytic enzymes that are active under acidic conditions. The interior of this organelle is maintained at an acidic ph (high proton concentration) by a H+ ATPase in the membrane that pumps protons against the concentration gradient 24

25 WHAT HAPPENS DURING GLUCOSE OXIDATION IF NO OXYGEN IS PRESENT? AEROBIC AND ANAEROBIC ORGANISMS One basic metabolic distinction among organisms is whether or not they can use O 2 as an electron acceptor in energy producing pathways AEROBES: CAN USE O 2 Obligate Aerobe: O 2 is obligatory for life ANAEROBES: CAN SUBSIST WITHOUT O 2 Facultative Anaerobe: can adapt to anaerobic conditions by substituting other electron acceptors for O 2 (yeast, bacteria) Obligate Anaerobe: cannot use O 2 and are poisoned by it 25

26 BEAKER CONTAINS YEAST AND SUGAR What is the gas being produced in the beaker? WHAT HAPPENS DURING GLUCOSE OXIDATION IF NO OXYGEN IS PRESENT? Cell has a limited amount of NADH which must be recycled if glycolysis is to continue under anaerobic conditions FERMENTATION: an anaerobic biological reaction process 26

27 FERMENTATION: THE ANAEROBIC FATE OF PYRUVATE Commercially Valuable fermentation reactions: Alcoholic fermentation by yeast used in brewing and winemaking Bacteria also can carry out alcoholic fermentation under anaerobic conditions 27

28 This type of fermentation occurs in some fungi and bacteria (used to make yogurt and cheese) and in human muscle cells when oxygen is limiting 28

29 The three common metabolic fates of pyruvate generated by glycolysis: Under aerobic conditions, the pyruvate is completely oxidized via the citric acid cycle to CO 2 and H 2 O [NADH acts as a high energy compound] Under anaerobic conditions, pyruvate must be converted to a reduced end product in order to reoxidize the NADH produced by the GAPDH reaction alcoholic fermentation: in yeast, pyruvate is converted to ethanol + CO 2 [free energy of NADH oxidation is dissipated as heat] in muscle cells, under anaerobic conditions, pyruvate is reduced to lactate [free energy of NADH oxidation is dissipated as heat] 29

30 FERMENTATION: an anaerobic biological reaction process in which a reduced organic compound (like glucose) acts as an electron donor and another organic compound acts as an electron acceptor An even more formal definition of fermentation fermentation: catabolic reactions producing ATP in which organic compounds serve as both the primary electron donor and ultimate electron acceptor and ATP is produced by substrate level phosophorylation Note: fermentation is extremely inefficient compared to aerobic respiration. From first principles: Why? 30

31 The potential energy drop between glucose and an electron acceptor like pyruvate is a fraction of the potential energy drop that occurs during cellular respiration where O 2 is the electron acceptor 31

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