Electron transport chain,oxidative phosphorylation & mitochondrial transport systems. M.Kohutiar, B.Sopko

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1 Electron transport chain,oxidative phosphorylation & mitochondrial transport systems. M.Kohutiar, B.Sopko

2 Content 1. Structure of mitochondria Mitochondrial transport systems 2. Electron transport 3. Parts of Electron transport chain 4. Proton gradient and synthesis of ATP 5. Uncoupling

3 Metabolism ANABOLIC CATABOLIC ΔG > 0 ΔG < 0

4 Gain of electron transport chain C H O 6O 6CO 6H O G 2823kJ / mol

5 Gain of electron transport chain 1 NADH H O NAD H O n(adp P ) i n(atp)

6 Gain of electron transport chain C H O 6O 6CO 6H O G 2823kJ / mol C H O 6O 6CO H 24e 6O 24H 24e 12H O 2 2

7 Gain of electron transport chain -0,4 NADH COMPLEX I FADH 2 COMPLEX II CoQ COMPLEX III cytochrome c COMPLEX IV 0,8

8 Gain of electron transport chain -0,4 NADH COMPLEX I FADH 2 COMPLEX II CoQ COMPLEX III cytochrome c COMPLEX IV 0,8

9 Gain of electron transport chain -0,4 NADH COMPLEX I FADH 2 COMPLEX II CoQ COMPLEX III cytochrome c COMPLEX IV 0,8 ΔE > 0

10 Gain of electron transport chain 1 NADH H O NAD H O n(adp P ) i n(atp) -0,4 NADH COMPLEX I FADH 2 COMPLEX II CoQ COMPLEX III cytochrome c COMPLEX IV 0,8

11 GLYCOLYSIS glucose glucose-6-phosphate glyceraldehyde-3-phosphate NAD + NADH 1,3-bisphosphoglycerate pyruvate NAD + acetyl-coa NADH

12 GLYCOLYSIS glucose glucose-6-phosphate glyceraldehyde-3-phosphate NAD + acetyl-coa NADH 1,3-bisphosphoglycerate NADH citrate ocalacetate pyruvate NAD + NAD + isocitrate NAD + acetyl-coa NADH malate fumarate CITRATE CYCLE 2-oxoglutarate NADH FADH 2 sukcinate sukcinyl-coa NAD + FAD NADH

13 Mitochondrial anatomy

14

15 ATP ADP translocase

16 Cofactors structure

17

18

19 Glycerolphosphate shuttle cytosol mitochondria CH 2 OH HC OH CH 2 O P NAD + glycerol-3-phosphate NADH CH 2 OH C O CH 2 O P dihydroxyacetone phosphate

20 Glycerolphosphate shuttle cytosol mitochondria CH 2 OH HC OH CH 2 O P glycerol-3-phosphate NAD + NADH CH 2 OH C O CH 2 O P dihydroxyacetone phosphate CH 2 OH HC OH CH 2 O P glycerol-3-phosphate FAD FADH 2 CH 2 OH C O CH 2 O P dihydroxyaceton phosphate

21 Malate-aspartate shuttle cytosol mitochondria HO NAD + NADH O HC CH 2 COOH COOH MDH C COOH CH 2 COOH malate oxalacetate

22 Malate-aspartate shuttle cytosol mitochondria HO NAD + NADH O HC CH 2 COOH COOH MDH C COOH CH 2 COOH malate oxalacetate malate oxalacetate HO HC COOH CH 2 COOH NAD + MDH NADH O C COOH CH 2 COOH

23 Malate-aspartate shuttle cytosol mitochondria HO NAD + NADH O HC CH 2 COOH COOH MDH C COOH CH 2 COOH malate oxalacetate malate oxalacetate HO HC COOH CH 2 COOH NAD + MDH NADH O C COOH CH 2 COOH AST Glu Asp ketoglutarate O C COOH (CH 2 ) 2 COOH

24 Malate-aspartate shuttle HO NAD + NADH O O HC AST CH 2 COOH COOH MDH C COOH CH 2 COOH Glu Asp C COOH (CH 2 ) 2 COOH cytosol malate oxalacetate a-ketoglutarate mitochondria malate oxalacetate a-ketoglutarate HO HC COOH CH 2 COOH NAD + MDH NADH O AST O C COOH CH 2 COOH Glu Asp C COOH (CH 2 ) 2 COOH

25 Electron transport NAD H 2e NADH E 0 0,32V 1 0 O2 2H 2e H2O E 2 0,82V

26 Electron transport NAD H 2e NADH E 0 0,32V 1 0 O2 2H 2e H2O E 0,82V 2 E 0 0,82-(-0,32) 1,14 V ΔG 0 zfδf 0 2 F1, kj/mol

27 Electron transport ADP P ATP ΔG 0 30,5 kj/mol

28 Electron transport ADP P ATP ΔG 0 30,5 kj/mol 330, %

29 Electron transport ADP P ATP ΔG 0 30,5 kj/mol 330, % ΔE 0 0 ΔG zf F 0,16 V

30 Electron transport NAD CoQ ox NADH CoQ red ΔE 0 0,36 V ΔG 0-69,5 kj/mol

31 Electron transport NADH CoQ ox NAD CoQ red ΔE ΔG 0 0 0,36 V -69,5 kj/mol CoQ red cytc ox CoQ ox cytc red ΔE 0 0,19 V ΔG 0-36,7 kj/mol

32 Electron transport NADH CoQ ox NAD CoQ red ΔE ΔG 0 0 0,36 V -69,5 kj/mol CoQ red cytc ox CoQ ox cytc red ΔE 0 0,19 V ΔG 0-36,7 kj/mol cytc red 1 2 O 2 cytc ox H 2 O ΔE ΔG 0 0 0,58 V -112 kj/mol

33 Electron transport FADH 2 CoQ ox FAD CoQ red ΔE 0 0,015 V ΔG 0-2,9 kj/mol

34 -0,4 NADH COMPLEX I FADH 2 COMPLEX II CoQ COMPLEX III cytochrome c COMPLEX IV 0,8

35 -0,4 FADH 2 COMPLEX II NADH COMPLEX I CoQ COMPLEX III cytochrome c ADP ATP ADP ATP 0,8 COMPLEX IV ADP ATP

36 -0,4 NADH COMPLEX I ADP rotenone ATP FADH 2 COMPLEX II CoQ COMPLEX III ADP antimycin A ATP cytochrome c 0,8 COMPLEX IV ADP ATP cyanide

37 [O 2 ] čas Time

38 [O 2 ] NAD Time

39 [O 2 ] NAD + 1 rotenon čas

40 [O 2 ] NAD + 1 rotenone 2 3 FADH time

41 [O 2 ] NAD + 1 rotenone 2 3 FADH 2 antimycin A Time

42 [O 2 ] NAD + 1 rotenone 2 3 FADH 2 antimycin A 4 5 cyanide 6 Time

43 Components of electron transport chain Complex I: NADH ubiquinonreductase NADH CoQ ox NAD CoQ red ΔE ΔG 0 0 0,36 V -69,5 kj/mol

44 FeS centres

45 Components of electron transport chain Complex II: succinate ubiquinon reductase FADH 2 CoQ ox FAD CoQ red ΔE 0 0,015 V ΔG 0-2,9 kj/mol

46 Components of electron transport chain Complex III:ubiquinol-cytC-reductase CoQ red cytc ox CoQ ox cytc red ΔE 0 0,19 V ΔG 0-36,7 kj/mol

47 Cytochromes

48 Components of electron transport chain Complex IV: cytochrome c oxidase cyt c(fe ) 4H O2 4 cyt c(fe ) 2H2O

49 Oxidative phosphorylation chemiosmotic theory (Mitchell) OXPHOS requires intact mitochondrial membrane Inner membrane is impermeable for some ions Electron transport is accompanied by transport of H +, and development of measurable gradient Compounds which increase the membrane permeability do not affect the electron chain but inhibit ATP synthesis

50 high H + H + H + H + H low H + ADP ATP

51 Proton gradient ΔG RT[pH in ph ex ] zfδ ΔG 0 21,5 kj/mol

52 Proton gradient ΔG RT[pH in ph ex ] zfδ ΔG 0 21,5 kj/mol Cytosolic side

53 Mechanism of redox loop Interimemb. space inner membrane e - matrix QH 2 H + QH.

54 Mechanism of redox loop Interimemb. space inner membrane e - matrix QH 2 QH 2 H + QH.

55 Mechanism of redox loop Interimemb. space inner membrane matrix e - H + QH 2 QH 2 H + e - cyt b k QH. QH.

56 Mechanism of redox loop Mezimemb. prostor inner membrane matrix e - H + QH 2 QH 2 H + e - cyt b k QH. QH. H + e - cyt c Q

57 Mechanism of redox loop Interimemb. space inner membrane matrix e - H + QH 2 QH 2 H + e - cyt b k QH. QH. H + e - Q Q cyt c

58 Mechanism of redox loop Interimemb. space inner membrane matrix e - H + QH 2 QH 2 H + e - cyt b k QH. QH. e - cyt b T H + e - Q Q cyt c

59 Mechanism of redox loop Interimemb. space inner membrane matrix e - H + QH 2 QH 2 H + e - QH. cyt b k e - cyt b T e - QH. H + e - Q Q H + cyt c

60 Structure of Complex V

61 3D Structure of Complex V

62 Schematic Structure of Complex V

63 Dam

64 Dam

65 Protonmotiv force and ATP synthesis

66 ATP synthesis

67 Regulation of oxidative phosphorylation 1 3 ADP P NAD cyt c 2 i NADH cyt c ATP

68 Regulation of oxidative phosphorylation ATP c cyt NAD 2 1 P ADP c cyt NADH i 3 ADP ADP.P cyt c cyt c NADH NAD K i

69 Regulation of oxidative phosphorylation ATP c cyt NAD 2 1 P ADP c cyt NADH i 3 K ATP ADP.P NAD NADH cyt c c cyt i i ADP.P ATP cyt c cyt c NADH NAD K

70 Uncoupling

71 Aerobic vs. Anaerobic ATP production Oxidation of 2 e - NADH. 2,5 mol ATP with consumption of 0,5 mol O 2 1 mol of the substrate is oxidised via Complexes I, III, and IV Oxidation of 2 e - FADH 2 1,5 mol ATP 1 mol of the substrate is oxidised via Complexes II, III a IV Total oxidation of 1 mol of glucose yields mol ATP,

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