The amount of cellular adenine is constant. -It exists as either ATP, ADP, or AMP (the concentration of these vary)
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1 Electron transport chain Final stage of aerobic oxidation! Also known as: -oxidative phosphorylation(when coupled to ATP synthase) -respiration (when coupled to ATP synthase) Purpose: -Recycle reduced molecules -Convert the energy gained from into ATP This occurs by: -Oxidizing NADH and FADH 2 -The e-gained from the above reactions are transported through a membrane bound electron transport system -This generates a membrane gradient (potential) -This potential energy allows for the phosphorylation of ADP (to generate ATP) Recycling The amount of cellular adenine is constant. -It exists as either ATP, ADP, or AMP (the concentration of these vary) - But [Adenine] remains constant. -So we recycle the adenine to whatever energy we need. -Table 10.2 Same is true for NAD and FAD; constant amount -It exist as either NAD or NADH (FAD or FADH 2 ) -But [FAD] or [NAD] remains constant -So we recycle the NAD and FAD to the desired form. NAD + and FAD are in short supply, so we need to recycle. A process has developed to generate energy from this recycling! glycolysis: glucose + 2NAD P i + 2 ADP 2 pyruvate+ 2 ATP + 2 NADH + H + Pyruvate dehydrogenase& TCA: pyruvate+ 4 NAD + + FAD + GDP + 2 H 2 O 3 CO NADH + 4 H + + P i + GTP + FADH 2 Chemistry C483 Fall 2009 Prof Jill Paterson 33-1
2 We have covered much of the background 14.2: ETC occurs in mitochondria Know details about mitochondria ETC proteins are present in the inner mitochondrial membrane Transport occurs from the matrix to the intermembrane space 14.3: Membrane gradients Just remember to switch equation to [from]/[to] May use ln as we have done, or as book does, log, by putting constant of into our equation Use of log allows easier movement to ph, which is a measure of H + We may discuss decouplers later The ETC is: A complex of integral membrane proteins, located in the inner mitochondrial membrane These proteins oxidize NADH and FADH 2 by passing electrons on to O 2, leading to the production of H 2 O The energy from oxidation creates a proton gradient (protons pumped from the matrix to the intermembrane space) This gradient provides energy for production of ATP from ADP (ATP synthase) NADH + H + + ½ O 2 + ADP + P i FADH 2 + ½ O 2 + ADP + P i NAD + + H 2 O + ATP FAD + H 2 O + ATP Chemistry C483 Fall 2009 Prof Jill Paterson 33-2
3 This whole system is a Electrons move from a reducing agent to an oxidizing agent (thus a series of redox reactions) NADH is the strongest reducing agent in biochemistry O 2 is the strongest oxidizing agent in biochemistry Electrons flow from a negative voltage to a positive voltage (high energy) (low energy) This flow of e- is spontaneous and thermodynamically favorable! Chemistry C483 Fall 2009 Prof Jill Paterson 33-3
4 ETC components The ETC is composed of 4 complexes and 2 mobile electron carriers. Complexes: Carriers: Complex I Complex II Complex III Complex IV Ubiquinone(Q) Cytochrome c Chain: I or II Q III cytoc IV O 2 H 2 O See why FADH 2 produces less energy?? Complex I NADH donates its electrons to Complex I Complex I has subunits! NADH-ubiquinone oxidoreductase Contains FMN and proteins with Fe-S clusters This is where redox occurs NADH transfers to FMN FMN transfers e-to Fe-S clusters, releases H+ to matrix Fe-S transfers e-to coenzyme Q ~ 4 H+ are transported for every 2 electrons transferred to Q Chemistry C483 Fall 2009 Prof Jill Paterson 33-4
5 Complex II FADH 2 donates its electrons to Complex II Complex II has 3 multisubunit enzymes Succinate dehydrogenase Contains FAD and proteins with Fe-S clusters This is where redox occurs FADH 2 transfers FAD transfers e-to Fe-S clusters, releases H+ to matrix Fe-S transfers e-to coenzyme Q Complex II does not contribute to H+ transport across membrane! Ubiquinone (Coenzyme Q) Not a protein! Membrane soluble, low molecular weight molecule Contains a long hydrophobic tail that keeps Q in the inner mitochondrial membrane Accepts e-one at a time (not as a hydride) Shuttles e-from complex I OR complex II to complex III Complex III QH 2 donates its electrons to Complex III Complex III has 3 main subunits Ubiquinol-cytochrome c oxidoreductase Contains several cytochromes and Fe-S clusters This is where redox occurs QH 2 transfers Complex III transfers e-to cytochromec ~ 4 H+ are transported for every 1 QH 2 oxidized (2 from matrix & 2 from QH 2 ) Chemistry C483 Fall 2009 Prof Jill Paterson 33-5
6 Cytochrome Cytochrome c Peripheral membrane protein Shuttles e-(and H+) from Complex III to Complex IV Another heme containing protein Complex IV Cytochromec donates e to Complex IV Cytochrome c oxidase 10 subunits Contains 2 cytochromes(a & a3) and proteins with Cu or Fe This is where redox occurs Transfers 4 electrons to O 2, to reduce O 2 to H 2 O 4 H + are transported for every O 2 reduced (2 e / O atom) Summary of ETC BUT it takes 2 NADH to reduce 1 O 2. So for every NADH, we only get 2 H + transported at this step 1. NADH is oxidized by Complex I (Complex I is reduced) 2. Complex I is oxidized by Q (Q is reduced to QH 2 ) 1. FADH 2 is oxidized by Complex II (Complex II is reduced) 2. Complex II is oxidized by Q (Q is reduced to QH 2 ) 1. Succinateis oxidized by Complex II (Complex II is reduced) 2. Complex II is oxidized by Q (Q is reduced to QH 2 ) 1. QH2 is oxidized by Complex III (complex III is reduced) 2. Complex III is oxidized by cytochrome c (cytochrome c is reduced) 1. Cytochrome c is oxidized by Complex IV (Complex IV is reduced) 2. Complex IV is oxidized by O 2 (O 2 is reduced to form H 2 O) Chemistry C483 Fall 2009 Prof Jill Paterson 33-6
7 Generation of a membrane gradient 1. Oxidation of NADH, succinate, and FADH 2 produces energy that allows H + to be pumped across the membrane 2. This generates an electrical AND a chemical (concentration) gradient (electrochemical gradient) 3. End result is the matrix is negative, while the intermembrane space is positive Linkage of gradient to ATP production Chemiosmotic hypothesis (Nobel Prize, 1978) The proton gradient IS the energy source for ATP generation The proton gradient is also referred to as the protonmoticeforce ( p) H + flow out of the matrix due to the ETC H + flow back in via ATP synthase ATP synthesis is driven by the H + gradient This is how NADH and FADH 2 provide additional ATP How does cytoplasmic NADH get into the mitochondria? Chemistry C483 Fall 2009 Prof Jill Paterson 33-7
8 Chemistry C483 Fall 2009 Prof Jill Paterson 33-8
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