Cellular Respiration: Harvesting Chemical Energy
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1 Lecture 13 9/30/05 I. General Principles Cellular Respiration: arvesting Chemical Energy Chapter 9 Lecture utline 1. Regulation of Enzymes: competitive, allosteric, phosphorylation 2. Equilibrium 3. Digestion vs Metabolism: catabolism and anabolism 4. What is a metabolic pathway? 5. eedback regulation of pathways 6. Catabolic pathways - stepping down the oxidation series of carbon 7. arvesting energy from redox reactions - substrate level phosphorylation ATP reducing equivalent carriers NAD + +, AD 2 8. Example of a catabolic pathway: atty Acid xidation igure eactions that proceed in a closed system Eventually reach equilibrium Can do Useful work Cannot Do work G < 0 G 0 Living systems pen System Must have constant flow of materials in Constant Energy Input Equilibrium to a living system is called. G < 0 (b) An open hydroelectric system. lowing water keeps driving the generator because intake and outflow of water keep the system from reaching equlibrium. ure 8.7 A (a) A closed hydroelectric system. Water flowing downhill turns a turbine that drives a generator providing electricity to a light bulb, but only until the system reaches equilibrium. 3 igure 8.7 4
2 7 8 etabolism totality of all chemical reactions of an organism digestion ydrolysis of polymers to monomers No energy arvested! occurs outside the cell catabolism energy capture capture reactions oxidize substrates, produce energy carriers Metabolism: a series of favorable reactions Inputs igure 8.7 G < 0 G < 0 G < 0 anabolism energy utilizing reactions use energy carriers, build things 5 Note: G<0 Metabolic Pathway: Waste Product The product of each reaction becomes the reactant for a next, so6 no reaction reaches equilibrium Metabolic Pathway Enzymes work in series Each enzyme carries out one reaction Reactions in series constitute a Pathway Enzyme 1 promotes reaction A B Enzyme 2 promotes reaction B C Enzyme 3 promotes reaction C D Enzyme 4 promotes reaction D E Enzyme 5 promotes reaction E Enzyme 6 promotes reaction G o as long as have A, G will be produced Chemistry of Life is organized into Metabolic Pathways Enzyme 1 Enzyme 2 Enzyme 3 A B C D Reaction 1 Reaction 2 Reaction 3 Starting molecule A A A A B C D E Product
3 11 12 eedback Regulation Active site available (threonine) Threonine in active site thr A A A B C Enzymes can be regulated D E Plenty of over ere, Shut it! Product f Pathway Is Allosteric Regulator f irst Enzyme In Pathway Isoleucine used up by cell Isoleucine binds to allosteric site eedback inhibition Active site of enzyme 1 no longer binds threonine; pathway is switched off Intermediate A Enzyme 2 Intermediate B Enzyme 3 Intermediate C Enzyme 4 Intermediate D Enzyme 5 Enzyme 1 (threonine deaminase) Allosteric modulator? modulator? 9 10 igure 8.21 End product (isoleucine) ile Why so many steps in a pathway? or example, oxidation of glucose: (glucose) C gallons f gasoline Many Small Controlled reactions G -686 kcal/mol -673 kcal/mol T S -13 kcal/mol in the cell, this is done in >21 steps! Capture the energy in small packets
4 atabolic pathway xidize in discrete steps tep down the oxidation series of carbon some activation step oxidation step, with energy harvest reorganization step oxidation step, another harvest etc yield product of pathway 13 What is an XIDATIN? or ionic species: Reduced means rich in electrons xidation: loss of e- Reduction : gain of e- e ++ reduced e +++ oxidized xidized means fewer electrons rganic Reductions X + 2e X 2 rganic xidation Y 2 Y + 2e Reduced igh enthalpy few bonds to oxygen many bonds to hydrogen XIDATIN series of carbon ydrocarbon chain Unsaturated hydrocarbon R-CC 2 ighly reduced Ease of Removing electrons electronegativity Alcohol Carbonyl Carboxylic Acid xidized few bonds to oxygen many bonds to hydrogen 15 Carbon Dioxide 16
5 In Metabolism: ighly reduced fully oxidized 3 -C 2 -C 2 -(C 2 ) x -C 2 -C C 2 + energy atty acid (captured) Partially reduced fully oxidized C 2 + energy rbohydrate (captured) 17 R-C 2 -C 3 R-CC 2 R-C 2 -C 2 - R-C 2 -C R-C 2 -C C Catabolic Pathways Progress down the xidation Series f Carbon adding - removed removed 2e removed 18 EDX Reactions xidations always paired with reductions If one thing gets oxidized, another becomes reduced Reactants Products Carriers of Reducing Equivalents CoEnzymes (Coactors) NAD + nicotinamide adenine dinucleotide NAD e- -> NAD becomes oxidized C C 2 + Energy C C becomes reduced Change the degree of electron sharing in covalent bonds NADP + nicotinamide adenine dinucleotide phosphate NADP e- -> NADP AD flavin adenine dinucleotide AD e- -> AD 2 Methane (reducing agent) xygen (oxidizing agent) Carbon dioxide Water igure
6 23 AD to AD 2 24 Electrons from organic compounds Are usually first transferred to NAD +, a coenzyme 2 e e + + NAD + N + C 2 P P C 2 N N C N 2 Nicotinamide (oxidized form) N 2 N N + 2[] (from food) Reduction of NAD + xidation of NAD N NAD C N 2 + Nicotinamide (reduced form) + 2 e- 2 + e- 1 e- 1 + NAD + to NAD Carries 2e- and 1 + igure NADP+ looks like this: AD looks like this: NADP + NADP + + 2e - 1 e e- 1 +
7 ow harvest energy packets upon oxidation? Carriers of Energy potential - high energy phosphate bonds ATP, GTP production substrate level phosphorylation less usual form of energy harvest -Carriers of reducing equivalents xidized form reduced form NAD+ NAD + + AD AD 2 -Can cash in reduced carriers for ATP oxidative phosphorylation ATP common energy currency $$$ phosphorylation igh energy phosphate bonds ubstrate Level Phosphorylation xample: A R- C -C- NAD + B NAD + + Enzyme 1 R- C-P i C P i B R- C-P i + ADP- Enzyme 2 Energy of xidations Captured igh Energy Compound (ATP) ADP -P i xidized to Carbon Dioxide C R- C - 27 ow harvest energy packets upon oxidation? $$$ - high energy phosphate bonds ATP, GTP production substrate level phosphorylation less usual form of energy harvest -Carriers of reducing equivalents xidized form reduced form NAD+ NAD + + AD AD 2 Poker chips -Can cash in reduced carriers for ATP oxidative phosphorylation 28
8 The Regeneration Energy Carriers Energy carriers (ATP, NAD +, AD) present in only minute amounts 2e Captured in catabolism 2e Cashed in Let s put it together Step down oxidation series arvest energy in discrete packets NAD + + atty Acid xidation Pathway Energy from catabolism (exergonic, energy yielding processes) NAD + Energy for cellular work (endergonic, energyconsuming processes) Start of Pathway atty Acid xidation (β-oxidation)( Priming Step Saturated hydrocarbon Priming Step (energy input) C 3 -C 2 -R-C 2 -C 2 -C atty acid - ATP + CoA-S -steps down oxidation states of carbon Ester (acid) unsaturated hydrocarbon 2e- 2 + remove C 3 -C 2 -R-C 2 -C 2 -C atty acyl CoA S-CoA ADP + P i 31 -captures Reducing potential NAD + + AD 2 2e- 2 + Ketone alcohol 32
9 et Result of atty Acid xidation Pathway atty acid shortened by 2 carbon unit 2 carbon acid attached to CoA (acetyl CoA) xidation of Carbon -C 2 - to C to acid S CoA Summary Digestion, Metabolism, Catabolism, Anabolism Biochemical Pathway; feedback regulation Catabolic Pathways - Step down oxidation series of carbon - arvest energy in discrete packets ATP, NAD + +, AD 2 atty Acid xidation Pathway Capture reducing equivalents 2 NAD AD
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