Overview: Life is Work. General Biology. 6. Cellular Respiration: Harvesting Chemical Energy. Energy
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1 ourse o: BG003 redits: 3.00 General Biology verview: Life is Work Living cells require transfusions of energy from outside sources to perform their many tasks The giant panda obtains energy for its cells by eating plants 6. ellular Respiration: arvesting hemical Energy Prof. Dr. Klaus eese Energy flows into an ecosystem as sunlight and leaves it as heat Light energy atabolic pathways yield energy by oxidizing organic fuels atabolic Pathways and Production of ESYSTEM The breakdown of organic molecules is exergonic ne catabolic process, fermentation + Photosynthesis in chloroplasts ellular respiration in mitochondria rganic molecules + is a partial degradation of sugars that occurs without oxygen ellular respiration is the most prevalent and efficient catabolic pathway eat energy powers most cellular work consumes oxygen and organic molecules such as glucose yields
2 To keep working ells must regenerate Redox Reactions: xidation and Reduction atabolic pathways yield energy due to the transfer of electrons The Principle of Redx Redx reactions Examples of Redx reactions becomes oxidized (loses electron) a + l a + + l becomes reduced (gains electron) Transfer electrons from one reactant to another by oxidation and reduction In oxidation a substance loses electrons, or is oxidized In reduction a substance gains electrons, or is reduced Some Redx reactions do not completely exchange electrons change the degree of electron sharing in covalent bonds xidation of rganic Fuel Molecules During ellular Respiration During cellular respiration Glucose is oxidized and oxygen is reduced Reactants becomes oxidized Products becomes oxidized Energy Energy + becomes reduced becomes reduced Stepwise Energy arvest via AD + and the Electron Transport hain ellular respiration Me t h a n e (reduc ing agent) x y g e n (ox idiz ing agent) arbon dioxide Water xidizes glucose in a series of steps
3 Electrons from organic compounds are usually first transferred to AD +, a coenzyme AD + P P + icotinamide (oxidized form) e + + e + + AD Dehydrogenase + [] Reduction of AD + (from food) xidation of AD + icotinamide (reduced form) AD, the reduced form of AD + passes the electrons to the electron transport chain + If electron transfer is not stepwise a large release of energy occurs as in the reaction of hydrogen and oxygen to form water Free energy, G + / Explosive release of heat and light energy Uncontrolled reaction oxyhydrogen (detonating/explosive) gas The electron transport chain passes electrons in a series of steps instead of in one explosive reaction uses the energy from the electron transfer to form Free energy, G + / (from food via AD) ontrolled release of energy for synthesis of + + e e + ellular respiration / The Stages of ellular Respiration: A Preview Respiration is a cumulative function of three metabolic stages Glycolysis The citric xidative Glycolysis Breaks down glucose into two molecules of pyruvate The citric ompletes the breakdown of glucose xidative is driven by the electron transport chain generates 3
4 An overview of cellular respiration the principle three metabolic stages Electrons carried via AD Electrons c arried via AD and FAD Both glycolysis and the citric can generate by substrate-level Enzy me Enzy me Glycolysis Glucose Py ruvate itric xidative : electron transport and chemiosmosis ytosol Mitochondrion Substrate P + Product Substrate-level Substrate-level xidative Glycolysis harvests energy by oxidizing glucose to pyruvate Glycolysis means splitting of sugar breaks down glucose into pyruvate occurs in the cytoplasm of the cell Glycolysis consists of two major phases Energy investment phase Energy payoff phase it r ic xidat ive phosphor ylat ion Energy investment phase Gl u c o s e + P Energy payoff phase used P 4 formed AD e + 4 AD Gl u c o s e + 4 formed used AD e AD + + 4
5 A closer look at the energy investment phase Gluc os e e x o k i n a s e P Gluc os e -6-ph o spha t e G lycolysis it r ic xidat ive phosphor ylat ion A closer look at the energy payoff phase AD + 6 Tr iose phosphat e dehydr ogenase AD P i + + P P, 3- Bi sphosphogl ycer at e 7 Phosphoglycer okinase Phos phoglucoisomerase P P Fructose-6-ph os p ha t e 3- Phosphogl ycerat e 8 Phosphoglycer om ut ase 3 Phos phofruc tok inase P - Phosphogl ycerat e P P Fructose-, 6-bisphosphate Aldolas e 4 9 Phosphoenol pyruvat e 0 Enolase P Pyr uvat e kinase P Dihydroxy a ce tone phosphate 5 Isomerase P Gly c e ra lde hy de - 3-phosphate 3 Pyruvat e The citric completes the energy-yielding oxidation of organic molecules The citric takes place in the matrix of the mitochondrion Before the citric can begin must first be converted to acetyl oa, which links the to glycolysis YTSL 3 Transport protein AD + AD oenzyme A MI T DRI S 3 o A Acetyle oa An overview of the citric (Krebs ycle) (from glycolysis, molecules per glucose) FAD FAD AD Acetyle oa itric o A o A o A + P i it r ic 3 AD xidat ive phosphor ylat ion 3 AD
6 A closer look at the citric G lycolysis it r ic xidat ive phosphor ylat ion S oa 3 Acetyl oa oa S AD + + AD + x a loa c e tate 8 Ma l a te itra te Isocitrate Figure 9. itric 3 7 AD + AD + + Fumarate oa S α-ke togluta ra te 6 4 oa S FAD 5 FAD AD + Succinate oa P i S AD GTP GDP Succinyl + + oa During oxidative, chemiosmosis couples electron transport to synthesis AD and FAD donate electrons to the electron transport chain, which powers synthesis via oxidative The Pathway of Electron Transport In the electron transport chain electrons from AD and FAD lose energy in several steps At the end of the chain electrons are passed to oxygen, forming water Free energy (G) relative to (kc l/mol) AD I FM Fe S Fe S FAD FAD yt b II Fe S III yt c Mu l t i p ro t e in complexes yt c + + yt a IV yt a3 hemiosmosis and the electron transport chain Intermembrane space Inner mitochondrial membrane Mitochondrial matrix xidat ive phosphor ylat ion. elect ron t ransport and chemiosmosis Protein complex of electron carners AD + I + + II Q FAD III yt c (arrying electrons from, food) Electron transport chain Electron transport and pumping of protons ( + ), which create an + gradient across the membrane IV AD + FA D / + xidative phosphor ylation + P i + + Inner Mitochondrial membrane hemiosmosis synthesis powered by the flow f + back across the membrane An Accounting of Production by ellular Respiration synthase During respiration, most energy flows in this sequence glucose to AD to electron transport chain to proton-motive force to At certain steps along the electron transport chain Electron transfer causes protein complexes to pump + from the mitochondrial matrix to the intermembrane space The resulting + gradient stores energy drives chemiosmosis in synthase is referred to as a proton-motive force (pmf) hemiosmosis Is an energy-coupling mechanism that uses energy in the form of a + gradient across a membrane to drive cellular work 6
7 hemiosmosis: The Energy-oupling Mechanism Aerobic oxidation of pyruvate and fatty s in mitochondria synthase is the enzyme that actually makes The proton motive force (pmf) ITERMEMBRAE SPAE A rotor within the membrane spins clockwise when + flows past it down the + gradient. A stator anchored in the membrane holds the knob stationary. + A rod (for stalk ) extending into the knob also spins, activating catalytic sites in the knob. The outer membrane is freely permeable to all metabolites, but specific transport proteins (colored ovals) in the inner membrane are required to import pyruvate + P i MI T DRI AL MAT RI X Three catalytic sites in the stationary knob join inorganic Phosphate to to make. (y ellow), (green), and Pi (purple) into the matrix and to export (green). AD generated in the c ytos ol is not trans ported direc tly to the matrix b ecause the inner membrane is impermeable to AD + and AD; instead, a shuttle system (red) transports electrons from cytosolic AD to AD + in the matrix. diffuses into the matrix and diffuses out.stage-: faty acyl groups are transferred from fatty acyl oa and transported across the inner membrane via a special carrier (blue oval) and then reatached to oa on the matrix side. is converted to acetyloa with the formation of AD, and fatty s (attac hed to oa) are als o c onv erted to acety l oa with formation of AD and FAD. xidation of acetyl oa in the citric generates AD and FAD. Stage-: electrons from these reduced coenzymes are transferred via electron transport complexes (blue boxes) to concomitant with transport of + ions from the matrix to the intramembrane space, generating the proton-mo t i v e f o rc e. El ec t ro n s f ro m AD f l o w d i re c t l y f ro m c o mp lex I to complex III, bypassing complex II. Stage 3: synthase, the F0F complex (orange), harnesses the proton-mo t i v e f o rc e t o s yn t h e s iz e AT P. Bl u e arro ws i n d i ca t e e le c t ro n fl o w; re d arrows transmembrane movement of protons; and green arrows indicate transport of metabolites. The phosphate and / transport system in the inner mitochondrial membrane There are three main processes in this metabolic enterprise YTSL Electron shuttles span membrane AD or FAD MITDRI AD AD 6 AD FAD Glycolysis Glucose Ac ety l o A itric x ida tiv e : electron transport and chemiosmosis about 3 or 34 by substrate-level by substrate-level by oxidative, depending on which shuttle transports electrons from AD in cytosol Maximum per glucose: About 36 or 38 The coordinated action of two antiporters (purple and green) results in the uptake of one 3- and one P4 - in exchange for one + during e - trans port. The outer membrane is not s hown here because it is permeable to molecules smaller than 5kDa. About 40% of the energy in a glucose molecule is transferred to during cellular respiration, making approximately 38 7
8 Fermentation enables some cells to produce without the use of oxygen ellular respiration relies on oxygen to produce In the absence of oxygen cells can still produce through fermentation Glycolysis can produce with or without oxygen, in aerobic or anaerobic conditions couples with fermentation to produce Types of Fermentati on ( without the use of oxygen) Fermentation consists of - glycolysis plus reactions that regenerate AD +, which can be reused by glyocolysis In alcohol fermentation pyruvate is converted to ethanol in two steps, one of which releases During lactic fermentation pyruvate is reduced directly to AD to form lactate as a waste product Gl u c o s e 3 + P Eth a n o l (a) Alcohol fermentation Gl u c o s e 3 Lactate AD + AD + P AD + AD (b) Lactic fermentation 3 3 Acetaldehyde 3 Fermentation and ellular Respiration ompared Both fermentation and cellular respiration use glycolysis to oxidize glucose and other organic fuels to pyruvate Fermentation and cellular respiration YTSL o present Fermentation Glucose present ellular respiration Glycolysis and the citric connect to many other metabolic pathways The Versatility of atabolism atabolic pathways Funnel electrons from many kinds of organic molecules into cellular respiration 3 Proteins Amino s a rbohydra te s Sugars Gl y c e rol Fatty s Gly c oly s is Glucose Gl y c e ral d e h yd e -3- P Fats differ in their final electron acceptor ellular respiration produces more Ethanol or lactate Acetyl oa MITDRI itric The catabolism of various molecules from food Acetyl oa itric pyruvate is a key juncture in catabolism x ida tiv e 8
9 Biosynthesis (Anabolic Pathways) The body uses small molecules to build other substances These small molecules may come directly from food or through glycolysis or the citric Regulation of ellular Respiration via Feedback Mechanisms ellular respiration is controlled by allosteric enzymes at key points in glycolysis and the citric Inhibits Glucose Gly c oly s is Fructose-6-phosphate Phosphofructokinase Fructose-,6-bi s ph o sp ha t e Acetyl oa itric x ida tiv e AMP Stimulates + Inhibits itrate 9
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