Cellular Respiration: Harvesting Chemical Energy

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Chapter 9Cellular Respiration:Harvesting Chemical EnergyPowerPoint Lecture Presentations forBiologyEighth EditionNeil Campbell and Jane ReeceLectures by Chris Romero, updated by Erin Barley with contributions from Joan SharpCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Overview: Life Is Work Living cells require energy from outsidesources Some animals, such as the giant panda, obtainenergy by eating plants, and some animalsfeed on other organisms that eat plantsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-1

Energy flows into an ecosystem as sunlightand leaves as heat Photosynthesis generates O2 and organicmolecules, which are used in cellularrespiration Cells use chemical energy stored in organicmolecules to regenerate ATP, which powersworkCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-2LightenergyECOSYSTEMPhotosynthesisin chloroplastsCO2 H2OOrganic Omolecules 2Cellular respirationin mitochondriaATPATP powers most cellular workHeatenergy

Concept 9.1: Catabolic pathways yield energy byoxidizing organic fuels Several processes are central to cellularrespiration and related pathwaysCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Catabolic Pathways and Production of ATP The breakdown of organic molecules isexergonic Fermentation is a partial degradation ofsugars that occurs without O2 Aerobic respiration consumes organicmolecules and O2 and yields ATP Anaerobic respiration is similar to aerobicrespiration but consumes compounds otherthan O2Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Cellular respiration includes both aerobic andanaerobic respiration but is often used to referto aerobic respiration Although carbohydrates, fats, and proteins areall consumed as fuel, it is helpful to tracecellular respiration with the sugar glucose:C6H12O6 6 O2(ATP heat)6 CO2 6 H2O EnergyCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Redox Reactions: Oxidation and Reduction The transfer of electrons during chemicalreactions releases energy stored in organicmolecules This released energy is ultimately used tosynthesize ATPCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Principle of Redox Chemical reactions that transfer electronsbetween reactants are called oxidation-reductionreactions, or redox reactions In oxidation, a substance loses electrons, or isoxidized In reduction, a substance gains electrons, or isreduced (the amount of positive charge isreduced)Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-UN1becomes oxidized(loses electron)becomes reduced(gains electron)

Fig. 9-UN2becomes oxidizedbecomes reduced

The electron donor is called the reducingagent The electron receptor is called the oxidizingagent Some redox reactions do not transfer electronsbut change the electron sharing in covalentbonds An example is the reaction between methaneand O2Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-3ReactantsProductsbecomes oxidizedbecomes )Carbon dioxideWater

Oxidation of Organic Fuel Molecules DuringCellular Respiration During cellular respiration, the fuel (such asglucose) is oxidized, and O2 is reduced:Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-UN3becomes oxidizedbecomes reduced

Fig. 9-UN4Dehydrogenase

Stepwise Energy Harvest via NAD and the ElectronTransport Chain In cellular respiration, glucose and otherorganic molecules are broken down in a seriesof steps Electrons from organic compounds are usuallyfirst transferred to NAD , a coenzyme As an electron acceptor, NAD functions as anoxidizing agent during cellular respiration Each NADH (the reduced form of NAD )represents stored energy that is tapped tosynthesize ATPCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-42 e– 2 H 2 e– H NADHH DehydrogenaseReduction of NAD NAD H 2[H]Oxidation of NADHNicotinamide(reduced form)Nicotinamide(oxidized form)

NADH passes the electrons to the electrontransport chain Unlike an uncontrolled reaction, the electrontransport chain passes electrons in a series ofsteps instead of one explosive reaction O2 pulls electrons down the chain in an energyyielding tumble The energy yielded is used to regenerate ATPCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-5H2 1/2 O22H(from food via NADH)Controlledrelease of –2H 2eenergy forsynthesis ofATP1/2 O2Explosiverelease ofheat and lightenergy1/(a) Uncontrolled reaction(b) Cellular respiration2 O2

The Stages of Cellular Respiration: A Preview Cellular respiration has three stages:– Glycolysis (breaks down glucose into twomolecules of pyruvate)– The citric acid cycle (completes thebreakdown of glucose)– Oxidative phosphorylation (accounts formost of the ATP synthesis)Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-6-1Electronscarriedvia evelphosphorylation

Fig. 9-6-2Electrons carriedvia NADH andFADH2Electronscarriedvia Substrate-levelphosphorylation

Fig. 9-6-3Electrons carriedvia NADH andFADH2Electronscarriedvia ivephosphorylation:electron osphorylationOxidativephosphorylation

The process that generates most of the ATP iscalled oxidative phosphorylation because it ispowered by redox reactionsBioFlix: Cellular RespirationCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Oxidative phosphorylation accounts for almost90% of the ATP generated by cellularrespiration A smaller amount of ATP is formed inglycolysis and the citric acid cycle bysubstrate-level phosphorylationCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-7EnzymeEnzymeADPPSubstrate ProductATP

Concept 9.2: Glycolysis harvests chemical energyby oxidizing glucose to pyruvate Glycolysis (“splitting of sugar”) breaks downglucose into two molecules of pyruvate Glycolysis occurs in the cytoplasm and has twomajor phases:– Energy investment phase– Energy payoff phaseCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-8Energy investment phaseGlucose2 ADP 2 P2 ATPused4 ATPformedEnergy payoff phase4 ADP 4 P2 NAD 4 e– 4 H 2 NADH 2 H 2 Pyruvate 2 H2ONetGlucose4 ATP formed – 2 ATP used2 NAD 4 e– 4 H 2 Pyruvate 2 H2O2 ATP2 NADH 2 H

Fig. phateATP1HexokinaseADPGlucose-6-phosphate

Fig. e

Fig. kinaseADPADPFructose1, 6-bisphosphateFructose1, 6-bisphosphate

Fig. hosphoglucoisomeraseFructose1, sphofructokinaseADP5IsomeraseFructose1, hosphateGlyceraldehyde3-phosphate

Fig. 9-9-52 NAD 2 NADH 2 H 6Triose phosphatedehydrogenase2 Pi2 1, 3-BisphosphoglycerateGlyceraldehyde3-phosphate2 NAD 2 NADH6Triose phosphatedehydrogenase2 Pi 2 H 2 1, 3-Bisphosphoglycerate

Fig. 9-9-62 NAD 2 NADH 2 H 6Triose phosphatedehydrogenase2 Pi2 1, 3-Bisphosphoglycerate2 ADP7Phosphoglycerokinase2 ATP2 1, 3-Bisphosphoglycerate2 ADP23-Phosphoglycerate2 ATP27Phosphoglycerokinase3-Phosphoglycerate

Fig. 9-9-72 NAD 2 NADH 2 H 6Triose phosphatedehydrogenase2 Pi2 1, 3-Bisphosphoglycerate2 ADP7 Phosphoglycerokinase2 te22-Phosphoglycerate

Fig. 9-9-82 NAD 2 NADH 2 H 6Triose phosphatedehydrogenase2 Pi2 1, 3-Bisphosphoglycerate2 ADP7 Phosphoglycerokinase2 lyceromutase922 H2O2-PhosphoglycerateEnolase9Enolase2 H2O2Phosphoenolpyruvate2Phosphoenolpyruvate

Fig. 9-9-92 NAD 6Triose phosphatedehydrogenase2 Pi2 NADH 2 H 2 1, 3-Bisphosphoglycerate2 ADP7 Phosphoglycerokinase2 ATP2Phosphoenolpyruvate2 ADP23-Phosphoglycerate8Phosphoglyceromutase2 ATP210Pyruvatekinase2-Phosphoglycerate92 H2OEnolase2 Phosphoenolpyruvate2 ADP10Pyruvate kinase2 ATP22PyruvatePyruvate

Concept 9.3: The citric acid cycle completes theenergy-yielding oxidation of organic molecules In the presence of O2, pyruvate enters themitochondrion Before the citric acid cycle can begin, pyruvatemust be converted to acetyl CoA, which linksthe cycle to glycolysisCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-10CYTOSOLMITOCHONDRIONNAD NADH H 21PyruvateTransport protein3CO2Coenzyme AAcetyl CoA

The citric acid cycle, also called the Krebscycle, takes place within the mitochondrialmatrix The cycle oxidizes organic fuel derived frompyruvate, generating 1 ATP, 3 NADH, and 1FADH2 per turnCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-11PyruvateCO2NAD CoANADH H Acetyl CoACoACoACitricacidcycleFADH22 CO23 NAD 3 NADHFAD 3 H ADP P iATP

The citric acid cycle has eight steps, eachcatalyzed by a specific enzyme The acetyl group of acetyl CoA joins the cycleby combining with oxaloacetate, forming citrate The next seven steps decompose the citrateback to oxaloacetate, making the process acycle The NADH and FADH2 produced by the cyclerelay electrons extracted from food to theelectron transport chainCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-12-1Acetyl CoACoA—SH1OxaloacetateCitrateCitricacidcycle

Fig. 9-12-2Acetyl cacidcycle

Fig. 9-12-3Acetyl CoACoA—SH1H2OOxaloacetate2CitrateIsocitrateNAD Citricacidcycle3NADH H CO2-Ketoglutarate

Fig. 9-12-4Acetyl CoACoA—SH1H2OOxaloacetate2CitrateIsocitrateNAD CitricacidcycleNADH H 3CO2CoA—SH-Ketoglutarate4NAD SuccinylCoANADH H CO2

Fig. 9-12-5Acetyl CoACoA—SH1H2OOxaloacetate2CitrateIsocitrateNAD CitricacidcycleNADH H 3CO2CoA—SH-Ketoglutarate4CoA—SH5NAD SuccinateGTP GDPADPATPPiSuccinylCoANADH H CO2

Fig. 9-12-6Acetyl CoACoA—SHH2O1Oxaloacetate2CitrateIsocitrateNAD CitricacidcycleNADH H NAD FADSuccinateGTP GDPADPATPPiSuccinylCoANADH H CO2

Fig. 9-12-7Acetyl eNAD Citricacidcycle7H2ONADH H NAD FADSuccinateGTP GDPADPATPPiSuccinylCoANADH H CO2

Fig. 9-12-8Acetyl CoACoA—SHNADH H H2O1NAD 8Oxaloacetate2MalateCitrateIsocitrateNAD Citricacidcycle7H2ONADH H NAD FADSuccinateGTP GDPADPATPPiSuccinylCoANADH H CO2

Concept 9.4: During oxidative phosphorylation,chemiosmosis couples electron transport to ATPsynthesis Following glycolysis and the citric acid cycle,NADH and FADH2 account for most of theenergy extracted from food These two electron carriers donate electrons tothe electron transport chain, which powers ATPsynthesis via oxidative phosphorylationCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Pathway of Electron Transport The electron transport chain is in the cristae ofthe mitochondrion Most of the chain’s components are proteins,which exist in multiprotein complexes The carriers alternate reduced and oxidizedstates as they accept and donate electrons Electrons drop in free energy as they go downthe chain and are finally passed to O2, formingH 2OCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-13NADH502 e–NAD FADH22 e–40FADMultiproteincomplexesFADFMNFe SFe SQCyt b30Fe SCyt c1IVCyt cCyt aCyt a320102 e–(from NADHor FADH2)02 H 1/2 O2H2O

Electrons are transferred from NADH or FADH2to the electron transport chain Electrons are passed through a number ofproteins including cytochromes (each with aniron atom) to O2 The electron transport chain generates no ATP The chain’s function is to break the large freeenergy drop from food to O2 into smaller stepsthat release energy in manageable amountsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Chemiosmosis: The Energy-Coupling Mechanism Electron transfer in the electron transport chaincauses proteins to pump H from themitochondrial matrix to the intermembrane space H then moves back across the membrane,passing through channels in ATP synthase ATP synthase uses the exergonic flow of H todrive phosphorylation of ATP This is an example of chemiosmosis, the use ofenergy in a H gradient to drive cellular workCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-14INTERMEMBRANE SPACEH StatorRotorInternalrodCatalyticknobADP PiATPMITOCHONDRIAL MATRIX

Fig. 9-15EXPERIMENTMagnetic kelplateRESULTSRotation in one directionNumber of photonsdetected ( 103)Rotation in opposite directionNo rotation3025200Sequential trials

Fig. 9-15aEXPERIMENTMagnetic kelplate

Fig. 9-15bRESULTSRotation in one directionRotation in opposite directionNo rotation3025200Sequential trials

The energy stored in a H gradient across amembrane couples the redox reactions of theelectron transport chain to ATP synthesis The H gradient is referred to as a protonmotive force, emphasizing its capacity to doworkCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-16H H H H Protein complexof electroncarriersCyt cVQATPsynthaseFADH2NADH2 H 1/2O2H2OFADNAD ADP P i(carrying electronsfrom food)ATPH 1 Electron transport chainOxidative phosphorylation2 Chemiosmosis

An Accounting of ATP Production by CellularRespiration During cellular respiration, most energy flows inthis sequence:glucoseNADHelectron transport chainproton-motive forceATP About 40% of the energy in a glucose moleculeis transferred to ATP during cellular respiration,making about 38 ATPCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-17Electron shuttlesspan membraneCYTOSOL2 NADHGlycolysisGlucose2PyruvateMITOCHONDRION2 NADHor2 FADH26 NADH2 NADH2AcetylCoA 2 ATPCitricacidcycle 2 ATPMaximum per glucose:About36 or 38 ATP2 FADH2Oxidativephosphorylation:electron transportandchemiosmosis about 32 or 34 ATP

Concept 9.5: Fermentation and anaerobicrespiration enable cells to produce ATP withoutthe use of oxygen Most cellular respiration requires O2 to produceATP Glycolysis can produce ATP with or without O2(in aerobic or anaerobic conditions) In the absence of O2, glycolysis couples withfermentation or anaerobic respiration toproduce ATPCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Anaerobic respiration uses an electrontransport chain with an electron acceptor otherthan O2, for example sulfate Fermentation uses phosphorylation instead ofan electron transport chain to generate ATPCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Types of Fermentation Fermentation consists of glycolysis plusreactions that regenerate NAD , which can bereused by glycolysis Two common types are alcohol fermentationand lactic acid fermentationCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

In alcohol fermentation, pyruvate isconverted to ethanol in two steps, with the firstreleasing CO2 Alcohol fermentation by yeast is used inbrewing, winemaking, and bakingFermentation OverviewCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-182 ADP 2 PiGlucose2 ATPGlycolysis2 Pyruvate2 NAD 2 NADH 2 H 2 CO22 Acetaldehyde2 Ethanol(a) Alcohol fermentation2 ADP 2 PiGlucose2 ATPGlycolysis2 NAD 2 NADH 2 H 2 Pyruvate2 Lactate(b) Lactic acid fermentation

Fig. 9-18a2 ADP 2 P iGlucose2 ATPGlycolysis2 Pyruvate2 NAD 2 Ethanol(a) Alcohol fermentation2 NADH 2 H 2 CO22 Acetaldehyde

In lactic acid fermentation, pyruvate isreduced to NADH, forming lactate as an endproduct, with no release of CO2 Lactic acid fermentation by some fungi andbacteria is used to make cheese and yogurt Human muscle cells use lactic acidfermentation to generate ATP when O2 isscarceCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-18b2 ADP 2 P iGlucose2 ATPGlycolysis2 NAD 2 NADH 2 H 2 Pyruvate2 Lactate(b) Lactic acid fermentation

Fermentation and Aerobic Respiration Compared Both processes use glycolysis to oxidizeglucose and other organic fuels to pyruvate The processes have different final electronacceptors: an organic molecule (such aspyruvate or acetaldehyde) in fermentation andO2 in cellular respiration Cellular respiration produces 38 ATP perglucose molecule; fermentation produces 2ATP per glucose moleculeCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Obligate anaerobes carry out fermentation oranaerobic respiration and cannot survive in thepresence of O2 Yeast and many bacteria are facultativeanaerobes, meaning that they can surviveusing either fermentation or cellular respiration In a facultative anaerobe, pyruvate is a fork inthe metabolic road that leads to two alternativecatabolic routesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 9-19GlucoseCYTOSOLGlycolysisPyruvateNo O2 present:FermentationO2 present:Aerobic etyl CoACitricacidcycle

The Evolutionary Significance of Glycolysis Glycolysis occurs in nearly all organisms Glycolysis probably evolved in ancientprokaryotes before there was oxygen in theatmosphereCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Concept 9.6: Glycolysis and the citric acid cycleconnect to many other metabolic pathways Gycolysis and the citric acid cycle are majorintersections to various catabolic and anabolicpathwaysCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Versatility of Catabolism Catabolic pathways funnel electrons from manykinds of organic molecules into cellularrespiration Glycolysis accepts a wide range ofcarbohydrates Proteins must be digested to amino acids;amino groups can feed glycolysis or the citricacid cycleCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fats are digested to glycerol (used inglycolysis) and fatty acids (used in generatingacetyl CoA) Fatty acids are broken down by beta oxidationand yield acetyl CoA An oxidized gram of fat produces more thantwice as much ATP as an oxidized gram ofcarbohydrateCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. sGlucoseGlyceraldehyde-3- PNH3PyruvateAcetyl erolFattyacids

Biosynthesis

Energy investment phase Glucose 2 ADP 2 P 2 ATP used 4 ATP formed Energy payoff phase 4 ADP 4 P 2 NAD 4 e– 4 H 2 NADH 2 H 2 Pyruvate 2 H 2 O Glucose 2 Pyruvate 2 H 2 O Net 4 ATP formed –2 ATP used 2 ATP 2 NAD 4 e– 4 H 2 NADH 2 H

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