Oxidative phosphorylation
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1 OXIDATIVE PHOSPHORYLATION
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3 Oxidative phosphorylation Oxidative reaction Coupled by phosphorylation to the generation of high energy intermediate (ATP or other high phosphagen) Oxidative phosphorylation at resp chain level via NADH D-ases form 3 mol ATP and via flavoprotein D-ases form 2 mol ATP Phosphorylations at the substrate level captured smaller energy eg:a) High energy phosphates are captured in kreb s cycle during the conversion of succinyl Co- A to succinate. And b) in glycolytic reactions on cytoplasmic.
4 Respiratory chain Enzyme complexes in mitochondria collects and transports reducing equivalents directing them to final reaction with oxygen form water and ATP Reducing equivalents flow through from redox potential negative to positive There are 4 enzyme complexes: - NADH-Q dehydrogenase / I - Succinate-Q dehydrogenase / II - Cytochromes dehydrogenase / III - Cytochrome oxidase / IV
5 DEHYDROGENASE DEHYDROGENASE DEHYDROGENASE OXIDASE AH 2 (Red) Carrier 1 (Ox) Carrier 2 (Red) Carrier 3 (Ox) H 2 O A (Ox) Carrier-H 2 1 (Red) Carrier 2 (Ox) Carrier-H 2 3 (Red) 1 / 2 O 2 Oxidation of a metabolite by dehydrogenases and finally by an oxidase in a respiratory chain
6 AH 2 Substrate A NAD + FpH 2 Flavoprotein 2Fe 3 + Cytochrome s NADH Fp 2Fe 2 + H + H + 2H + 2H+ H 2 O 1 / 2 O 2 Transport of reducing equivalents through the respiratory chain
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8 Mitochondrial Powerhouses of the cell most of energy captured takes place inside it Outer membrane permeable to most metabolites, contain various enzym (acyl Co- A synthetase, glycerolphosphate acyltransferase ) Inner membrane selectively permeable Matrix contain phospholipid cardiolipin together with enzymes of resp chain Intermembrane space has similar composition with cytoplasmic and contain adenylyl kinase and creatine kinase
9 B A Phosphorylating complexes B OUTER MEMBRANE INNER MEMBRANE MATRIX F 1 subunits F 0 subunits MATRIX Cristae Sonication INNER MEMBRANE OUTER MEMBRANE Submitochondrial particel Formed from fragments of the inner membrance
10 Respiratory chain Not all substrates are linked to resp chain through NADH-D-ase Co-Q (ubiquinone) mobile component, collects reducing equivalents from flavoprotein complexes and passes them on to cytochrome b (the lowest redox pot) Cytochrome oxidase has a very high affinity for oxygen resp chain to function at maximum rate until tissue depleted of O 2 irreversible reaction
11 Pyruvat e Lipoate Fp (FAD) Proline 3-Hydroxyacyl- CoA 3- Hydroxybutyrate Glutamate Malate Isocitrate NADH Fp (FMN) FeS Succin ate Cholin e Fp (FAD) FeS Q Cyt b FeS Cyt c 1 Cyt c Cyt aa 3 Cu O 2 - Ketoglutarate Glycerol 3-phosphate Fp (FAD) FeS FeS ETF (FAD) Fp (FAD) Acyl-CoA Sarcosine Dimethylglycin FeS ETF Fp Q Cyt : Iron-sulfur protein : Electrontransferring flavoprotein : Flavoprotein : Ubiquinone : Cytochrome
12 Resp chain & oxd phos inhibitors Inhibitors of resp chain 1. Blocking electrons transfer from Fe-S to co-q, ie: barbiturates, pierisidin-a, rotenon, carboxine, succinate D ase competitive inhibitor: malonate 2. Blocking electrons transfer from cty b to cyt c, ie: dimercaprol, antimycin A 3. Inhibitors of cytochrome oxidase: H 2 S, CO and CN
13 Resp chain & oxd phos inhibitors Inhibitors of oxidative phosphorylation, ie: oligomycine, atractyloside Un-couplers (dissociate oxidation in resp chain from phosphorylation) respiration to become uncontrolled, ie: dinitrophenol, dinitrochressol, pentachlorophenol, chloro carbonyl cyanide phenilhydrazon (cccp)
14 Succinate FAD FeS BAL Antimycin A H 2 S CO CN - Complex I Complex III Complex IV NADH FMN, FeS Q Cyt b, FeS, Cyt c Cyt a Cyt a 3 Cyt C Cu Cu 1 O 2 Piericidin A Amobarbital Rotenone Uncouplers Uncouplers Oligomycin Oligomycin ADP + P 1 ATP ADP + P 1 ATP ADP + P 1 ATP
15 Mechanism of oxidative phosphorylation Mitchell s chemiosmotic theory: - energy from oxidation in resp chain translocation of H + (protons) electrochemical potential difference in matrix and intermembrane space drive the mechanism of responsible for the formation of ATP (ATP synthase)
16 Mechanism of oxidative phosphorylation Complexes I, III and IV of resp chain is a proton pump Pi + ADP ATP, by ATP synthase ATP synthase is a complex enzyme consist of several protein subunits (F 1 ), which attached to membrane protein complex (F 0 ) F 1 project into matrix and contain the phosphorylation mechanism F 0 spans the membrane and forms the proton channel
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19 Exchange metabolites at inner mitochondrial membrane - Exchange of anions against OH - ions and cations against H + ions for transport of ionized metabolites - Freely permeable to uncharged small molecules O 2, H 2 O, CO 2, NH 3 monocarboxylic acids (3 hydroxy butyric, acetoacetic, acetic) - Long chain fatty acids need carnitine system - Symport pyruvate - H +
20 Exchange metabolites at inner mitochondrial membrane Dicarboxylate and tricarboxylate anions require specific carrier linked to inorganic phosphate (H 2 PO 4- ) Exchange ATP / ADP by adenine nucleotide transporter Transport of oxaloacetate need transamination process
21 Oxidation of extramitochondrial NADH - NADH cannot penetrate mitochondrial membrane produced continuously in cytosol by 3 phosphoglyceraldehyde D-ase - Aerobic conditions: not accumulated be oxidized by resp chain - Transfer of reducing equivalents from cytosol to mitochondrial require substrate pairs, linked by suitable D-ase
22 Oxidation of extramitochondrial NADH - The mechanism: 1. Glycerophosphate shuttle only 2 mol ATP are formed per atom oxygen consumed present in brain, muscle, adipose, liver but deficient in heart muscle 2. Malate shuttle more universal utility more complex, due to the impermeability of mitochondrial membrane to oxaloacetate
23 OUTER MEMBRANE INNER MEMBRANE CYTOSOL MITOCHONDRION NAD + Glycerol 3- phosphate Glycerol 3- phosphate FAD GLYCEROL-3- PHOSPHATE DEHYDROGENASE (CYTOSOLIC) GLYCEROL-3- PHOSPHATE DEHYDROGENASE (MITHOCONDRIAL) NADH + H + Dehydroxyacetone phosphate Dehydroxyacetone phosphate FDH 2 Respiratory Chain Glycerophosphate shuttle for transfer of reducing equivalents from the cytosol into the mitochondrion
24 NAD + Malat e CYTOSOL INNER MAMBRAN E 1 MITHOCOND RION Malate NAD + NADH +H + MALATE DEHYDROGENASE Oxaloacetat e -KG -KG MALATE DEHYDROGENASE Oxaloacetat e NADH +H + TRANSAMI NASE TRANSAMI NASE Glutamate Asp Asp Glutamate 2 H + H + Malate shuttle for transfer of reducing equivalents from the cytosol into the mitocondrion. 1. Ketoglutarate transporter, 2. glutamateaspartate transporter (note the proton symport with glutamate)
25 Creatine phosphate shuttle Facilitating transport of high energy phosphat from mitochondria in active tissues Isoenzyme of creatine kinase (CK M ), in intermembrane space catalyzing transfer ~ P (ATP) to creatine: ~ P(ATP) + creatine creatine-p, transported into cytosol via protein pores available for generation of extramitochondrial ATP
26 H P N CREATINE KINASE H 2 N C NH C NH N H 3 C N COO - ΔG O = 12.6 kj/mol COO - Creatine phosphate Creatine
27 Clinical aspects Fatal infantile mitochondrial myopathy and renal dysfunction due to severe diminution / absence of most oxidoreductase MELAS (mitochondrial encephalopathy, lactic acidosis and stroke) due to complex I or complex IV deficiency mutation in mt DNA
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