Sinh học - Chapter 9: Cellular respiration: harvesting chemical energy

Chapter 9Cellular Respiration: Harvesting Chemical EnergyOverview: Life Is WorkLiving cells require energy from outside sources.Some animals, such as the giant panda, obtain energy by eating plants, and some animals feed on other organisms that eat plants.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsEnergy flows into an ecosystem as sunlight and leaves as heat.Photosynthesis generates O2 and organic molecules C-H-O, which are used in cellular respiration.Cells use chemical energy stored in organic molecules C-H-O to regenerate ATP, which powers work.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsEnergy Flow and Chemical Recycling inEcosystems IN: Light EnergyECOSYSTEMPhotosynthesis in chloroplastsCO2 + H2OCellular respirationin mitochondriaOrganicMolecules + O2ATP powers most cellular workOUT: Heat energyATPCatabolic pathways yield energy by oxidizing organic fuels: C-H-O molecules to Produce ATPThe breakdown of organic molecules is exergonic.Fermentation is a partial degradation of sugars that occurs without O2Aerobic respiration consumes organic molecules and O2 and yields ATP.Anaerobic respiration is similar to aerobic respiration but consumes compounds without O2Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsCellular respiration includes both aerobic and anaerobic respiration but is often used to refer to aerobic respiration.Although carbohydrates, fats, and proteins are all consumed as fuel, it is helpful to trace cellular respiration with the sugar glucose: C6H12O6 + 6 O2  6 CO2 + 6 H2O + Energy (ATP + heat)Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsRedox Reactions: Oxidation and ReductionOxidation is a LOSS (of H or electrons).Reduction is a GAIN (of H or electrons).The transfer of electrons during chemical reactions releases energy stored in organic molecules.This released energy is ultimately used to synthesize ATP.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsThe Principle of RedoxChemical reactions that transfer electrons between reactants are called oxidation-reduction reactions, or redox reactions.In oxidation, a substance loses electrons, or is oxidized.In reduction, a substance gains electrons, or is reduced (the amount of positive charge is reduced).Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsOxidation / Reduction Reactionsbecomes oxidized(loses electron)becomes reduced(gains electron)Oxidation / Reduction Reactionsbecomes oxidizedbecomes reducedThe electron donor is oxidized and is called the reducing agentThe electron receptor is reduced and is called the oxidizing agentSome redox reactions do not transfer electrons but change the electron sharing in covalent bonds.An example is the reaction between methane and O2Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsMethane combustion as an energy-yielding redox reactionReactantsbecomes oxidizedbecomes reducedProductsMethane(reducingagent)Oxygen(oxidizingagent)Carbon dioxideWaterOxidation of Organic Fuel Molecules During Cellular RespirationDuring cellular respiration: The fuel C-H-O (such as glucose) is oxidized, looses H’s and O2 is reduced, gains H’sCopyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsDuring Cellular Respiration During cellular respiration, the fuel (such se) is oxidized, and O2 ibecomes oxidizedbecomes reduced NAD+ is Reduced to NADH + H+DehydrogenaseStepwise Energy Harvest via NAD+ and the Electron Transport ChainIn cellular respiration, glucose and other organic molecules are broken down in a series of steps.Electrons from organic compounds are usually first transferred to NAD+ = a coenzyme.As an electron acceptor, NAD+ functions as an oxidizing agent during cellular respiration.NADH = the reduced form of NAD+ . Each NADH represents stored energy that is tapped to synthesize ATP.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsNAD+ as an electron shuttle Dehydrogenase Reduction of NAD+Oxidation of NADH2 e– + 2 H+2 e– + H+NAD++2[H]NADH+H+H+Nicotinamide(oxidized form)Nicotinamide(reduced form)NADH passes the electrons to the electron transport chain ETC.Unlike an uncontrolled reaction, the electron transport chain passes electrons in a series of steps instead of one explosive reaction.O2 pulls electrons down the ETC chain in an energy-yielding tumble.The energy yielded is used to regenerate ATP.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsAn introduction to electron transport chainsFree energy, GFree energy, G(a) Uncontrolled reactionH2OH2 + 1/2 O2 Explosiverelease ofheat and lightenergy(b) Cellular respiration: step-wise controlled reactionControlledrelease ofenergy forsynthesis ofATP2 H+ + 2 e– 2 H+1/2 O2(from food C-H-O via NADH)ATPATPATP1/2 O22 H+2 e–Electron transportchainH2OThe Stages of Cellular Respiration: A PreviewCellular respiration has three stages:Glycolysis (breaks down glucose into two molecules of pyruvate)The citric acid cycle: Krebs Cycle (completes the breakdown of glucose)Oxidative phosphorylation (accounts for most of the ATP synthesis) by chemiosmosis.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsAn overview of cellular respiration - GlycolysisSubstrate-levelphosphorylationATPCytosolGlucosePyruvate GlycolysisElectronscarried via NADHAn overview of cellular respiration: Glycolysis & Krebs CycleMitochondrionSubstrate-levelphosphorylationATPCytosolGlucosePyruvateGlycolysisElectronscarried via NADHSubstrate-levelphosphorylationATPElectrons carriedvia NADH andFADH2KrebsCycleAn overview of Cellular Respiration BasicsMitochondrionSubstrate-levelphosphorylationATPCytosolGlucosePyruvateGlycolysisElectronscarried via NADHSubstrate-levelphosphorylationATPElectrons carriedvia NADH andFADH2OxidativephosphorylationATPKrebsCycleOxidativephosphorylation:electron transportandchemiosmosisThe process in cell respiration that generates most of the ATP is oxidative phosphorylation (powered by redox reactions).Oxidative phosphorylation accounts for almost 90% of the ATP generated by cellular respiration.A smaller amount of ATP is formed in glycolysis and the citric acid / Krebs Cycle by substrate-level phosphorylation.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsSubstrate-level phosphorylationEnzymeADPPSubstrateEnzymeATP+ProductGlycolysis harvests chemical energy by oxidizing glucose to pyruvateGlycolysis (“sugar splitting”) breaks down glucose into two molecules of pyruvate.Glycolysis occurs in the cytoplasm and has two major phases:Energy investment phase = EAEnergy payoff phase = ATP and NADHCopyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsThe energy input and output of glycolysisEnergy investment phaseGlucose2 ADP + 2P2 ATPusedformed4 ATPEnergy payoff phase4 ADP + 4P2 NAD+ + 4 e– + 4 H+2 NADH+ 2 H+2 Pyruvate + 2 H2O2 Pyruvate + 2 H2OGlucoseNet4 ATP formed – 2 ATP used2 ATP2 NAD+ + 4 e– + 4 H+2 NADH + 2 H+A closer look at glycolysisHexokinaseATPADP1Phosphoglucoisomerase2Phosphogluco-isomerase2GlucoseGlucose-6-phosphateFructose-6-phosphateGlucose-6-phosphateFructose-6-phosphate1HexokinaseATPADPPhosphoglucoisomerasePhosphofructokinaseATPADP23ATPADPPhosphofructokinase: allosteric enzymeFructose-1, 6-bisphosphateGlucoseGlucose-6-phosphateFructose-6-phosphateFructose-1, 6-bisphosphate123Fructose-6-phosphate3GlucoseATPADPHexokinaseGlucose-6-phosphatePhosphoglucoisomeraseFructose-6-phosphateATPADPPhosphofructokinaseFructose-1, 6-bisphosphateAldolaseIsomeraseDihydroxyacetonephosphateGlyceraldehyde-3-phosphate12345AldolaseIsomeraseFructose-1, 6-bisphosphateDihydroxyacetonephosphateGlyceraldehyde-3-phosphate452 NAD+NADH2Triose phosphatedehydrogenase+ 2 H+2Pi22 ADP1, 3-BisphosphoglyceratePhosphoglycerokinase2 ATP23-Phosphoglycerate6722 ADP2 ATP1, 3-Bisphosphoglycerate3-PhosphoglyceratePhosphoglycero-kinase27Fig. 9-9-73-PhosphoglycerateTriose phosphatedehydrogenase2 NAD+2NADH+ 2 H+2Pi22 ADPPhosphoglycerokinase1, 3-Bisphosphoglycerate2 ATP3-Phosphoglycerate2Phosphoglyceromutase2-Phosphoglycerate22-Phosphoglycerate22Phosphoglycero-mutase67882 NAD+NADH22222+ 2 H+Triose phosphatedehydrogenase2Pi1, 3-BisphosphoglyceratePhosphoglycerokinase2 ADP2 ATP3-PhosphoglyceratePhosphoglyceromutaseEnolase2-Phosphoglycerate2 H2OPhosphoenolpyruvate987622-PhosphoglycerateEnolase22 H2OPhosphoenolpyruvate9Triose phosphatedehydrogenase2 NAD+NADH2222222 ADP2 ATPPyruvatePyruvate kinasePhosphoenolpyruvateEnolase2 H2O2-PhosphoglyceratePhosphoglyceromutase3-PhosphoglyceratePhosphoglycerokinase2 ATP2 ADP1, 3-Bisphosphoglycerate+ 2 H+67891022 ADP2 ATPPhosphoenolpyruvatePyruvate kinase2Pyruvate102PiThe Citric Acid Cycle = Krebs Cycle: completes the energy-yielding oxidation of organic moleculesIf O2 is present, pyruvates (3 carbon C-H-O) enter the mitochondria.Before the Krebs Cycle can begin, the pyruvate must be converted to acetyl CoAAcetyl CoA = Acetate + CoEnzyme AAcetate = a 2 carbon C-H-O CoEnzyme A = a carrier molecule Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsConversion of pyruvate to acetyl CoA, the junction between glycolysis and the citric acid cycleCYTOSOLMITOCHONDRIONNAD+NADH+ H+213PyruvateTransport proteinCO2Coenzyme AAcetyl CoAThe citric acid cycle / Krebs cycle takes place within the mitochondrial matrix.The Krebs cycle oxidizes organic fuel derived from pyruvate, generating 1 ATP, 3 NADH, and 1 FADH2 per turn (two turns per glucose molecule from glycolysis).Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummingscitric acid /Krebs CyclePyruvateNAD+NADH+ H+Acetyl CoACO2CoACoACoACitricacidcycleFADH2FADCO2233 NAD++ 3 H+ADP + P iATPNADHThe citric acid / Krebs cycle has eight steps, each catalyzed by a specific enzyme.The acetyl group of acetyl CoA joins the cycle by combining with oxaloacetate, OAA, forming citrate (citric acid).The next seven steps break down the citrate and regenerate oxaloacetate,OAA, making the process a cycle.The NADH and FADH2 produced by the Krebs Cycle carry electrons extracted from food (C-H-O) to the electron transport chain in the mitochondrial cristae membrane.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsA closer look at the citric acid / Krebs cycleAcetyl CoACoA—SHCitrateH2OIsocitrateNAD+NADH+ H+CO2-Keto-glutarateCoA—SHCO2NAD+NADH+ H+SuccinylCoACoA—SHPiGTPGDPADPATPSuccinateFADFADH2FumarateCitricAcidCycleH2OMalateOxaloacetate OAANADH+H+NAD+12345678During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesisFollowing glycolysis and the citric acid cycle, NADH and FADH2 account for most of the energy extracted from food.These two electron carriers: NADH and FADH2 donate electrons to the electron transport chain, which powers ATP synthesis via oxidative phosphorylation.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsThe Pathway of Electron TransportThe electron transport chain is in the cristae membrane of the mitochondrion.Most of the chain’s components are proteins, which exist in multiprotein complexes.The carriers alternate reduced and oxidized states as they accept and donate electrons, redox.Electrons drop in free energy as they go down the chain and are finally passed to O2, forming H2O (waste).Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsETC cristaeFree-energy change during electron transportNADHNAD+2FADH22FADMultiproteincomplexesFADFe•SFMNFe•SQFe•SCyt bCyt c1Cyt cCyt aCyt a3IVFree energy (G) relative to O2 (kcal/mol)50403020102(from NADHor FADH2)02 H+ + 1/2O2H2Oe–e–e–wasteElectrons are transferred from NADH or FADH2 to the electron transport chain, ETC.Electrons are passed along the cristae membrane through a number of proteins including cytochromes (each with an iron atom) to O2The electron transport chain generates no ATPThe chain’s function is to break the large free-energy drop from food to O2 into smaller steps that release energy in manageable amounts.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsChemiosmosis The Energy-Coupling MechanismElectron transfer, redox, in the electron transport chain causes proteins to pump H+ from the mitochondrial matrix to the intermembrane space  creating a proton H+ gradient.H+ then moves back across the membrane, passing through channels in ATP synthase. ATP synthase uses the exergonic flow of H+ to drive phosphorylation of ATP.This is an example of chemiosmosis, the use of energy in a H+ gradient to drive ATP synthesis.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsThe energy stored in a H+ (proton) gradient, across a membrane couples the redox reactions of the electron transport chain to ATP synthesis.The H+ gradient is a proton-motive force, emphasizing its capacity to do work.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsChemiosomosisProtein complexof electroncarriersH+H+H+Cyt cQVFADH2FADNAD+NADH(carrying electronsfrom food)Electron transport chain: redox2 H+ + 1/2O2H2OADP +PiChemiosmosisOxidative phosphorylationH+H+ATP synthaseATP21Chemiosmosis: Energy Coupling - couples the electron transport chain to ATP synthesis An Accounting of ATP Production by Cellular RespirationDuring cellular respiration, most energy flows in this sequence: glucose  NADH  electron transport chain  proton-motive force  ATPAbout 40% of the energy in a glucose molecule is transferred to ATP during cellular respiration, making about 38 ATP.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsMaximum per glucose:About36 or 38 ATP+ 2 ATP+ 2 ATP+ about 32 or 34 ATPOxidativephosphorylation:electron transport chemiosmosisCitricAcidCycle2AcetylCoAGlycolysisGlucose2Pyruvate2 NADH2 NADH6 NADH2 FADH22 FADH22 NADHCYTOSOLElectron shuttlesspan membraneorMITOCHONDRIONAerobic Cellular RespirationATP yield per molecule of glucose at each stage of cellular respiration Fermentation and anaerobic respiration enable cells  to produce ATP without the use of oxygenMost cellular respiration requires O2 to produce ATP.Glycolysis produces ATP without O2 (in aerobic or anaerobic conditions).In the absence of O2, glycolysis couples with fermentation or anaerobic respiration to produce ATP.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFermentation: Anaerobic / No Oxygen UsedFermentation consists of glycolysis plus reactions that regenerate NAD+, which can be reused by glycolysis.Two common types are alcohol fermentation and lactic acid fermentation.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsIn alcohol fermentation, pyruvate is converted to ethanol in two steps, with the first releasing CO2Alcohol fermentation by yeast is used in brewing, winemaking, and baking / $$ commercial uses.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFermentation Regenerates NAD+for use in Glycolysis2 ADP + 2Pi2 ATPGlucoseGlycolysis2 NAD+2 NADH2 Pyruvate+ 2 H+2 Acetaldehyde2 Ethanol(a) Alcohol fermentation2 ADP + 2Pi2 ATPGlucoseGlycolysis2 NAD+2 NADH+ 2 H+2 Pyruvate2 Lactate(b) Lactic acid fermentation2CO2In lactic acid fermentation, pyruvate is reduced to NADH, forming lactate as an end product, with no release of CO2Lactic acid fermentation by some fungi and bacteria is used to make cheese and yogurt $$Human muscle cells use lactic acid fermentation to generate ATP when O2 is scarce; meaning there is an O2 debt. This reaction is reversible when O2 is available.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsGlucose2 ADP + 2Pi2 ATPGlycolysis2 NAD+2 NADH+ 2 H+2 Pyruvate2 LactateLactic acid fermentation: reversible if oxygen is availableFermentation and Aerobic Respiration ComparedBoth processes use glycolysis to oxidize glucose and other organic fuels to pyruvateThe processes have different final electron acceptors: an organic molecule (such as pyruvate or acetaldehyde) in fermentation. O2 in aerobic cellular respiration.Aerobic cellular respiration nets 38 ATP per glucose molecule; fermentation nets only 2 ATP per glucose molecule.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsObligate anaerobes carry out fermentation or anaerobic respiration and cannot survive in the presence of O2Facultative anaerobes (yeast and many bacteria) can survive using either fermentation or cellular respiration. In a facultative anaerobe, pyruvate is a fork in the metabolic road that leads to two alternative catabolic routes.Obligate aerobes carry out aerobic cellular respiration and require O2Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsMicro-organismsPyruvate as a key juncture in catabolismGlucoseGlycolysisPyruvateCYTOSOLNo O2 present:FermentationO2 present: Aerobic cellular respirationMITOCHONDRIONAcetyl CoAEthanolorlactateCitricacidcycleThe Evolutionary Significance of GlycolysisGlycolysis occurs in nearly all organisms.Glycolysis probably evolved in ancient prokaryotes before there was oxygen in the atmosphere.Fermentation evolved to recycle NAD+ back to glycolysis so ATP production continues in the absence of O2Gycolysis and the Citric Acid Cycle are major intersections to various catabolic and anabolic pathways.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsGlycolysis and the citric acid cycle connect to many other metabolic pathwaysGycolysis and the citric acid cycle are major intersections to various catabolic and anabolic pathways.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsThe Versatility of Catabolism: breaking downCatabolic pathways funnel electrons from many kinds of organic molecules into cellular respiration.Glycolysis accepts a wide range of carbohydrates.Proteins must be digested to amino acids which can feed glycolysis or the citric acid cycle.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFats are digested to glycerol (used in glycolysis) and fatty acids (used in generating acetyl CoA). Fatty acids are broken down by beta oxidation and yield acetyl CoA.An oxidized gram of fat produces more than twice as much ATP as an oxidized gram of carbohydrate (fat has more calories = unit of energy for cell work).Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsVersatilityProteinsCarbohydratesAminoacidsSugarsFatsGlycerolFattyacidsGlycolysisGlucoseGlyceraldehyde-3-PyruvatePNH3Acetyl CoACitricacidcycleOxidativephosphorylationBiosynthesis - Anabolic Pathways: BuildingThe body uses small molecules to build larger more complex molecules.These small molecules may come directly from food, from glycolysis, or from the citric acid cycle.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsRegulation of Cellular Respiration via Feedback MechanismsFeedback inhibition is the most common mechanism for control, regulation.If ATP concentration begins to drop, respiration speeds up; when there is plenty of ATP, respiration slows down.Control of catabolism is based mainly on regulating the activity of enzymes at strategic points in the catabolic pathway.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsGlucoseGlycolysisFructose-6-phosphatePhosphofructokinaseFructose-1,6-bisphosphateInhibitsAMPStimulatesInhibitsPyruvateCitrateAcetyl CoACitricacidcycleOxidativephosphorylationATP+––Regulation: Feedback InhibitionThe control of cellular respirationInputsGlycolysisOutputs+22ATPNADH2GlucosePyruvateReview Glycolysis:Review: Citric Acid / Krebs CycleInputsOutputsAcetyl CoA22226ATPNADHFADH2OxaloacetateCitric acidcycleS—CoACH3C OO C COOCH2COOReview:Chemiosmosis = Energy Coupling: ATP SynthesisINTER-MEMBRANESPACEH+ATPsynthaseATPADP +PiH+MITO-CHONDRIALMATRIXProton Motive Force = H+ concentration Gradient. As H’s increase, the pH drops in the Intermembrane space; so pH difference Increases across the membraneYou should now be able to:Explain in general terms how redox reactions are involved in energy exchanges.Name the three stages of cellular respiration; for each, state the region of the eukaryotic cell where it occurs and the products that result.In general terms, explain the role of the electron transport chain in cellular respiration.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsExplain where and how the respiratory electron transport chain creates a proton gradient.Distinguish between fermentation and anaerobic respiration.Distinguish between obligate and facultative anaerobes.Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings