Change to Office Hours this Friday and next Monday. Tomorrow (Abel): 8:30 10:30 am. Monday (Katrina): Cancelled (05/04)
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1 Change to Office Hours this Friday and next Monday Tomorrow (Abel): 8:30 10:30 am Monday (Katrina): Cancelled (05/04)
2 Lecture 10 Proton Gradient-dependent ATP Synthesis Oxidative Phosphorylation Photo-Phosphorylation
3 Model of the Electron Transport Chain (ETC) Glycerol-3-P Shuttle Outer Mitochondrial Membrane G3P DHAP 4H + 4H + 2H Cyt c Intermembrane space I Q F 0 II III IV Succinate Fumarate F 1 NADH + H + NAD + TCA-Cycle Fatty acid degradation (β-oxidation) ½O 2 + 2H + H 2 O ADP + Pi ATP H + Matrix p. 55
4 Mechanism of ATP Synthase p. 57
5
6
7 p. 57
8 L ADP + Pi γ O T γadp + Pi H + (P) H + (P) H + (P) ATP γ L O γ T ADP + Pi ATP L γ O T O L T H + (N) H + (N) H + (N) Proton movement through F O causes rotation of γ subunit and conformational changes of β subunits p. 57
9 Model of the Electron Transport Chain (ETC) Glycerol-3-P Shuttle Outer Mitochondrial Membrane G3P DHAP 4H + 4H + 2H Cyt c Intermembrane space I Q F 0 II III IV Succinate Fumarate F 1 NADH + H + NAD + TCA-Cycle Fatty acid degradation (β-oxidation) ½O 2 + 2H + H 2 O ADP + Pi ATP H + Matrix p. 55
10 Bioenergetics of the ETC and ATP Synthesis Do the numbers fit? (yes, more or less )
11 1. What is ΔG o of NADH oxidation? NADH + H O 2 NAD + + H 2 O ΔE o = E o ox E o red ΔE o = 0.82 V ( 0.32 V) ΔE o = 1.14 V ΔG o = nfδe o ΔG o = 220 kj mol -1
12 2. What is ΔG G of H + transport against H + gradient? Experimental data: ΔpH = 0.75 and ΔΨ = 150 mv ΔG = ΔG o + RT ln [H + ] P / [H + ] N + ZFΔΨ ΔG o = 0 ΔG = RT ln [H + ] P / [H + ] N + ZFΔΨ ln = log ΔG = RT log [H + ] P / [H + ] N + ZFΔΨ ΔG = RT (log [H + ] P log[h + ] N ) + ZFΔΨ ph = log [H + ] ΔG = RT (ph N ph P ) + ZFΔΨ ΔG = RT ΔpH + ZFΔΨ ΔG = RT ZF 0.15 V Z = charge of H + (+1) ΔG = ~ + 20 kj per mol H +
13 NADH + H O 2 NAD + ETC 10 H + N 10 H + P + H 2 O 1. ΔG o of NADH oxidation ΔG o = 220 kj mol -1 Energy gained and coupled to proton transport 2. ΔG G of H + transport against electro-chemical chemical gradient ΔG = ~ + 20 kj per mol H + Energy gained is sufficient to pump 10 protons! ΔG = ~ kj per 10 mol H +
14 NADH + H O 2 NAD + ETC 10 H + N 10 H + P + H 2 O 10 H + N ATP + H 2 O F o F 1 10 H + P ADP + Pi 3. How many ATP are be produced per NADH? EMF (NADH) PMF (10 Protons)? ATP ( 220 kjmol - 1 ) ( 200 kjmol - 1 ) ΔG o (ATP Synthesis) = 30.5 kj mol -1 ΔG (ATP Synthesis) ~ 50 kj mol -1 In reality: 6-77 ATP 4 ATP 2-33 ATP
15 Inner Membrane Transporters Tap into the PMF H + H + H + H + Intermembrane Space Pyr H + H + H + H + Pi ADP ATP H + Pyr H + Pi ADP 3- ATP 4- OH - OH - OH - OH - OHŌH- OH - OH - Mitochondrial Matrix
16
17 4. How many ATP are produced per? No complex I (only complex II-IV) O 2 FAD + H 2 O ETC 6 H + N 6 H + P 6 H + N ATP + H 2 O F o F 1 6 H + P ADP + Pi ΔE o = E o ox E o red = 0.82 V (+0.06 V) = 0.76 V ΔG o = nfδe o = 147 kj mol -1 EMF (FADH) PMF (6 Protons) ~2 2 ATP ( 140 kjmol - 1 ) ( 120 kjmol - 1 )
18 Model of the Electron Transport Chain (ETC) Outer Mitochondrial Membrane G3P DHAP 4H + 4H + 2H Cyt c Intermembrane space I Q F 0 II III IV Succinate Fumarate F 1 NADH + H + NAD + ½O 2 + 2H + H 2 O ADP + Pi ATP H + Amytal, Rotenone Antimycin A Cyanide Oligomycin p. 52
19 O 2 Consumption ATP Production No Substrate, but ADP + Pi + Succinate + Cyanide Inhibition of Complex IV
20 Model of the Electron Transport Chain (ETC) Outer Mitochondrial Membrane G3P DHAP 4H + 4H + 2H Cyt c Intermembrane space I Q F 0 II III IV Succinate Fumarate Cyanide F 1 NADH + H + NAD + ½O 2 + 2H + H 2 O ADP + Pi ATP H + Amytal, Rotenone Antimycin A Oligomycin p. 52
21 O 2 ATP Succinate, but no (ADP, Pi) + (ADP, Pi) Inhibition of Fo-F1 F1 ATPase + Oligomycin Acceptor Control [ADP]
22 Model of the Electron Transport Chain (ETC) Outer Mitochondrial Membrane G3P DHAP 4H + 4H + 2H Cyt c Intermembrane space I Q F 0 II III IV Succinate Fumarate Oligomycin F 1 NADH + H + NAD + ½O 2 + 2H + H 2 O ADP + Pi ATP H + Amytal, Rotenone Antimycin A Cyanide p. 52
23 O 2 ATP Succinate, but no (ADP, Pi) + (ADP, Pi) Inhibition of Fo-F1 F1 ATPase + Oligomycin + DNP Uncoupling of PMF and ATP synthesis Acceptor Control [ADP]
24 Dinitrophenol (DNP) Uncouplers (collapse of H + gradient) O O N + N + O - - O G3P DHAP 4H + 4H + 2H Cyt c HO Thermogenin (Proton Channel) I Q F 0 II III IV Succinate Fumarate F 1 NADH + H + NAD + ½O 2 + 2H + H 2 O ADP + Pi ATP H + Amytal, Rotenone Antimycin A Cyanide Oligomycin
25 Feedback Control ( Acceptor Control ) Glucose PFK-1 PK PDH CS IDH KGA-DH ETC ADP Glycogen Glucose-6-P Pyruvate Acetyl-CoA NADH ATP Activation Lactate Inhibition
26 Feedback Control by NADH Glucose Glycogen Glucose-6-P Pyruvate Acetyl-CoA NADH X ATP PK PDH CS IDH KGA-DH ETC Lactate
27 Allosteric Control of the TCA Cycle Pyruvate Pyr Carboxylase (+) Acetyl-CoA PDH Acetyl-CoA (+) AMP, NAD +, CoA (-)) ATP, NADH, Acetyl-CoA (+) Activation (-)) Inhibition CS (+) ADP (-)) ATP, NADH, Citrate, Succinyl-CoA Oxalaoacetate Citrate Malate Isocitrate IDH (+) ADP, NAD + (-)) ATP, NADH Fumarate α-ketoglutarate Succinyl-CoA α-kga (+) AMP (-)) ATP, NADH, Succinyl-CoA p. 49
28 ATP Yield of Respiration Glucose 2 Pyruvate 2 ATP 2 NADH (5%, anaerobic) 2 Pyruvate 2 Acetyl-CoA + 2 CO 2 2 NADH 6 2 Acetyl-CoA 4 CO 2 2 ATP 6 NADH ATP 1 NADH = 3 ATP 1 = 2 ATP 36 ATP (if glycerol-3-p P shuttle) (~65% efficiency, ΔG of Glc Oxidation ATP)
29 Proton Gradient-dependent ATP Synthesis by Photo-phosphorylation
30 Plants and Photosynthetic Bacteria LIGHT O 2 ATP NADPH CO 2 + H 2 O ATP NADH Photosynthesis Respiration NADP + ADP + Pi H 2 O Autotrophic Metabolism Reduced Organic Compounds Heterotrophic Metabolism NAD + ADP + Pi p. 58
31 Plants and Photosynthetic Bacteria LIGHT O 2 ATP NADPH CO 2 + H 2 O ATP NADH Photosynthesis Respiration NADP + ADP + Pi H 2 O Reduced Organic Compounds NAD + ADP + Pi Light Reactions CO 2 Fixation Heterotrophic Metabolism p. 58
32 E o V Chl* e - CO V L I G H T Chl o ΔE ATP NADP + NADPH NADH NAD + Food (Reduced Carbon) ΔE ATP O 2 H 2 O V Chl + p. 58
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