Cell biology : a short course PDF
Preview Cell biology : a short course
P1:GDZ WY001-Bolsover-FM WY001-Bolsover-v3.cls October22,2003 14:59 CELL BIOLOGY SECOND EDITION i P1:GDZ WY001-Bolsover-FM WY001-Bolsover-v3.cls October22,2003 14:59 CELL BIOLOGY A Short Course SECOND EDITION Stephen R. Bolsover DepartmentofPhysiology UniversityCollegeLondon Jeremy S. Hyams DepartmentofBiology UniversityCollegeLondon Elizabeth A. Shephard DepartmentofBiochemistryandMolecularBiology UniversityCollegeLondon Hugh A. White DepartmentofBiochemistryandMolecularBiology UniversityCollegeLondon Claudia G. Wiedemann DepartmentofPhysiology UniversityCollegeLondon A JOHN WILEY & SONS, INC., PUBLICATION iii P1:GDZ WY001-Bolsover-FM WY001-Bolsover-v3.cls October22,2003 14:59 Copyright(cid:2)C 2004byJohnWiley&Sons,Inc.Allrightsreserved. PublishedbyJohnWiley&Sons,Inc.,Hoboken,NewJersey. PublishedsimultaneouslyinCanada. Nopartofthispublicationmaybereproduced,storedinaretrievalsystem,ortransmittedinanyformorbyany means,electronic,mechanical,photocopying,recording,scanning,orotherwise,exceptaspermittedunder Section107or108ofthe1976UnitedStatesCopyrightAct,withouteitherthepriorwrittenpermissionofthe Publisher,orauthorizationthroughpaymentoftheappropriateper-copyfeetotheCopyrightClearanceCenter, Inc.,222RosewoodDrive,Danvers,MA01923,978-750-8400,fax978-646-8600,oronthewebat www.copyright.com.RequeststothePublisherforpermissionshouldbeaddressedtothePermissions Department,JohnWiley&Sons,Inc.,111RiverStreet,Hoboken,NJ07030,(201)748-6011, fax(201)748-6008. LimitofLiability/DisclaimerofWarranty:Whilethepublisherandauthorhaveusedtheirbesteffortsin preparingthisbook,theymakenorepresentationsorwarrantieswithrespecttotheaccuracyorcompletenessof thecontentsofthisbookandspecificallydisclaimanyimpliedwarrantiesofmerchantabilityorfitnessfora particularpurpose.Nowarrantymaybecreatedorextendedbysalesrepresentativesorwrittensalesmaterials. Theadviceandstrategiescontainedhereinmaynotbesuitableforyoursituation.Youshouldconsultwitha professionalwhereappropriate.Neitherthepublishernorauthorshallbeliableforanylossofprofitoranyother commercialdamages,includingbutnotlimitedtospecial,incidental,consequential,orotherdamages. ForgeneralinformationonourotherproductsandservicespleasecontactourCustomerCareDepartmentwithin theU.S.at877-762-2974,outsidetheU.S.at317-572-3993orfax317-572-4002. Wileyalsopublishesitsbooksinavarietyofelectronicformats.Somecontentthatappearsinprint,however, maynotbeavailableinelectronicformat. LibraryofCongressCataloging-in-PublicationData: Cellbiology:ashortcourse/StephenR.Bolsover...[etal.].—2nded. p. cm. Includesbibliographicalreferencesandindex. ISBN0-471-26393-1(Paper) 1.Cytology. I.Bolsover,StephenR.,1954– QH581.2.C425 2003 571.6—dc21 2003000577 PrintedintheUnitedStatesofAmerica 10 9 8 7 6 5 4 3 2 1 iv P1:GDZ WY001-Bolsover-FM WY001-Bolsover-v3.cls October22,2003 14:59 CONTENTS IN BRIEF 1 CELLSANDTISSUES 1 2 FROMWATERTODNA:THECHEMISTRYOFLIFE 19 3 MEMBRANESANDORGANELLES 51 4 DNASTRUCTUREANDTHEGENETICCODE 65 5 DNAASADATASTORAGEMEDIUM 87 6 TRANSCRIPTIONANDTHECONTROLOFGENEEXPRESSION 105 7 RECOMBINANTDNAANDGENETICENGINEERING 129 8 MANUFACTURINGPROTEIN 163 9 PROTEINSTRUCTURE 183 10 INTRACELLULARPROTEINTRAFFICKING 213 11 HOWPROTEINSWORK 237 12 ENERGYTRADINGWITHINTHECELL 257 13 METABOLISM 281 14 IONSANDVOLTAGES 309 15 THEACTIONPOTENTIAL 325 16 INTRACELLULARSIGNALING 341 17 INTERCELLULARCOMMUNICATION 363 18 MECHANICALMOLECULES 381 19 CELLCYCLEANDCONTROLOFCELLNUMBER 401 20 CASESTUDY:CYSTICFIBROSIS 423 v P1:GDZ WY001-Bolsover-FM WY001-Bolsover-v3.cls October22,2003 14:59 CONTENTS PREFACE, xv BasesAreMoleculesThatTakeH+from Water, 25 ACKNOWLEDGMENTS, xvii IsoelectricPoint, 25 INSTRUCTORNOTES, xix AHydrogenBondFormsWhena HydrogenAtomIsShared, 25 BiologicalMacromolecules, 27 1 CELLSANDTISSUES, 1 Carbohydrates:CandyandCanes, 27 PrinciplesofMicroscopy, 2 AnAssortmentofSweets, 27 TheLightMicroscope, 3 Disaccharides, 28 TheElectronMicroscope, 8 OutoftheSweetComesForth TheScanningElectronMicroscope, 9 Strength, 30 OnlyTwoTypesofCell, 9 ModifiedSugars, 31 SpecialPropertiesofPlantCells, 11 Nucleosides,Phosphate,and Viruses, 11 Nucleotides, 35 OriginofEukaryoticCells, 12 AminoAcids,Polypeptides,andProteins, 37 CellSpecialization, 12 Lipids, 39 Epithelia, 12 Hydrolysis, 44 ConnectiveTissue, 13 Summary, 46 NervousTissue, 13 FurtherReading, 47 Muscle, 14 ReviewQuestions, 47 Plants, 15 AnswerstoReviewQuestions, 48 Summary, 16 ReviewQuestions, 16 3 MEMBRANESAND AnswerstoReviewQuestions, 17 ORGANELLES, 51 BasicPropertiesofCellMembranes, 51 2 FROMWATERTODNA: StraightThroughtheMembrane: THECHEMISTRYOFLIFE, 19 DiffusionThroughtheBilayer, 53 TheChemicalBond:Sharing BeyondtheCellMembrane: Electrons, 19 TheExtracellularMatrix, 53 InteractionswithWater:Solutions, 21 CellJunctions, 54 IonicCompoundsWillDissolveOnlyin OrganellesBoundedbyDouble-Membrane PolarSolvents, 21 Envelopes, 56 AcidsAreMoleculesThatGiveH+to TheNucleus, 56 Water, 21 MitochondriaandChloroplasts, 58 vii P1:GDZ WY001-Bolsover-FM WY001-Bolsover-v3.cls October22,2003 14:59 viii CONTENTS OrganellesBoundedbySingle-Membrane DNAReplication, 87 Envelopes, 58 TheDNAReplicationFork, 88 Peroxisomes, 59 ProteinsOpenuptheDNADoubleHelix EndoplasmicReticulum, 60 DuringReplication, 88 GolgiApparatus, 60 DnaAProtein, 88 Lysosomes, 61 DnaBandDnaCProteins, 90 Summary, 61 Single-StrandBindingProteins, 90 ReviewQuestions, 62 BiochemistryofDNAReplication, 90 AnswerstoReviewQuestions, 63 DNASynthesisRequiresanRNA Primer, 90 4 DNASTRUCTUREANDTHEGENETIC RNAPrimersAreRemoved, 92 CODE, 65 TheSelf-CorrectingDNA Introduction, 65 Polymerase, 92 TheStructureofDNA, 65 DNARepair, 94 TheDNAMoleculeIsaDoubleHelix, 68 SpontaneousandChemicallyInduced BaseChanges, 94 TheTwoDNAChainsAre Complementary, 69 RepairProcesses, 94 DifferentFormsofDNA, 71 GeneStructureandOrganizationin Eukaryotes, 98 DNAastheGeneticMaterial, 71 IntronsandExons—Additional PackagingofDNAMoleculesinto ComplexityinEukaryoticGenes, 98 Chromosomes, 71 TheMajorClassesofEukaryotic EukaryoticChromosomesandChromatin DNA, 99 Structure, 71 GeneNomenclature, 101 ProkaryoticChromosomes, 73 Summary, 101 Plasmids, 74 FurtherReading, 102 Viruses, 74 ReviewQuestions, 102 TheGeneticCode, 75 AnswerstoReviewQuestions, 103 AminoAcidNamesAreAbbreviated, 79 TheCodeIsDegenerateBut 6 Unambiguous, 79 TRANSCRIPTIONANDTHE CONTROLOFGENEEXPRESSION, 105 StartandStopCodonsandtheReading Frame, 79 StructureofRNA, 105 TheCodeIsNearlyUniversal, 80 RNAPolymerase, 106 MissenseMutations, 80 GeneNotation, 106 Summary, 81 BacterialRNASynthesis, 106 FurtherReading, 84 ControlofBacterialGeneExpression, 109 ReviewQuestions, 84 lac,anInducibleOperon, 111 AnswerstoReviewQuestions, 85 trp,aRepressibleOperon, 116 EukaryoticRNASynthesis, 118 5 DNAASADATASTORAGE MessengerRNAProcessing, 118 MEDIUM, 87 ControlofEukaryoticGene Introduction, 87 Expression, 119 P1:GDZ WY001-Bolsover-FM WY001-Bolsover-v3.cls October22,2003 14:59 CONTENTS ix GlucocorticoidsCrosstheCellMembrane Ribosome-BindingSite, 168 toActivateTranscription, 121 ChainInitiation, 169 Summary, 125 The70SInitiationComplex, 171 FurtherReading, 125 ElongationoftheProteinChain, 171 ReviewQuestions, 126 ThePolyribosome, 173 AnswerstoReviewQuestions, 127 TerminationofProteinSynthesis, 174 TheRibosomeIsRecycled, 175 7 RECOMBINANTDNAANDGENETIC EukaryoticProteinSynthesisIsaLittle ENGINEERING, 129 MoreComplex, 175 DNACloning, 129 AntibioticsandProteinSynthesis, 176 CreatingtheClone, 130 Summary, 178 IntroductionofForeignDNAMolecules FurtherReading, 179 intoBacteria, 130 ReviewQuestions, 179 SelectionofcDNAClones, 134 AnswerstoReviewQuestions, 180 GenomicDNAClones, 139 UsesofDNAClones, 143 9 PROTEINSTRUCTURE, 183 DNASequencing, 143 NamingProteins, 184 SouthernBlotting, 146 PolymersofAminoAcids, 184 InsituHybridization, 147 TheAminoAcidBuildingBlocks, 184 NorthernBlotting, 148 TheUniquePropertiesofEachAmino ProductionofMammalianProteinsin Acid, 188 Bacteria, 149 OtherAminoAcidsAreFoundin ProteinEngineering, 149 Nature, 191 PolymeraseChainReaction, 150 TheThree-DimensionalStructuresof IdentifyingtheGeneResponsiblefora Proteins, 192 Disease, 152 HydrogenBonds, 195 ReverseGenetics, 152 ElectrostaticInteractions, 199 TransgenicAnimals, 157 vanderWaalsForces, 199 EthicsofDNATestingforInherited HydrophobicInteractions, 199 Disease, 157 DisulfideBonds, 199 Summary, 158 TertiaryStructure:Domainsand FurtherReading, 159 Motifs, 200 ReviewQuestions, 159 QuaternaryStructure:AssembliesofProtein AnswerstoReviewQuestions, 160 Subunits, 204 ProstheticGroups, 205 8 MANUFACTURINGPROTEIN, 163 ThePrimaryStructureContainsallthe AttachmentofanAminoAcidtoIts InformationNecessarytoSpecify tRNA, 163 Higher-LevelStructures, 206 TransferRNA,theAnticodon,andthe Summary, 209 Wobble, 164 FurtherReading, 209 TheRibosome, 165 ReviewQuestions, 210 BacterialProteinSynthesis, 168 AnswerstoReviewQuestions, 211 P1:GDZ WY001-Bolsover-FM WY001-Bolsover-v3.cls October22,2003 14:59 x CONTENTS 10 TheInitialVelocityofanEnzyme INTRACELLULARPROTEIN Reaction, 242 TRAFFICKING, 213 EffectofSubstrateConcentrationon ThreeModesofIntracellularProtein InitialVelocity, 244 Transport, 213 TheEffectofEnzymeConcentration, 245 TargetingSequences, 215 TheSpecificityConstant, 247 Retention, 215 EnzymeCatalysis, 247 TransporttoandfromtheNucleus, 215 CofactorsandProstheticGroups, 249 TheNuclearPoreComplex, 216 EnzymesCanBeRegulated, 251 GatedTransportThroughtheNuclear Pore, 216 Summary, 254 GTPasesandtheGDP/GTPCycle, 218 FurtherReading, 254 GTPasesinNuclearTransport, 218 ReviewQuestions, 255 TransportAcrossMembranes, 221 AnswerstoReviewQuestions, 256 TransporttoMitochondria, 221 12 ChaperonesandProteinFolding, 221 ENERGYTRADINGWITHINTHE CELL, 257 TransporttoPeroxisomes, 221 CellularEnergyCurrencies, 258 SynthesisontheRoughEndoplasmic Reticulum, 223 ReducedNicotinamideAdenine Dinucleotide(NADH), 259 Glycosylation:TheEndoplasmic ReticulumandGolgiSystem, 225 NucleosideTriphosphates(ATPplus GTP,CTP,TTP,andUTP), 259 VesicularTraffickingBetween IntracellularCompartments, 226 TheHydrogenIonGradientAcrossthe MitochondrialMembrane, 261 ThePrincipleofFissionandFusion, 226 TheSodiumGradientAcrossthePlasma VesicleFormation, 228 Membrane, 262 Coatomer-CoatedVesicles, 228 EnergyCurrenciesAreInterconvertible, 263 Clathrin-CoatedVesicles, 229 ExchangeMechanismsConvertBetween TheTrans-GolgiNetworkandProtein theFourEnergyCurrencies, 263 Secretion, 229 ElectronTransportChain, 265 TargetingProteinstotheLysosome, 230 ATPSynthase, 269 Fusion, 231 Sodium/PotassiumATPase, 270 Summary, 232 ADP/ATPExchanger, 271 FurtherReading, 233 Photosynthesis, 271 ReviewQuestions, 233 AllCarriersCanChangeDirection, 275 AnswerstoReviewQuestions, 234 Summary, 278 11 FurtherReading, 278 HOWPROTEINSWORK, 237 ReviewQuestions, 278 HowProteinsBindOtherMolecules, 237 AnswerstoReviewQuestions, 279 DynamicProteinStructures, 238 AllostericEffects, 238 13 METABOLISM, 281 ChemicalChangesThatShiftthe TheKrebsCycle:TheCentralSwitching PreferredShapeofaProtein, 240 YardofMetabolism, 283 EnzymesAreProteinCatalysts, 241 P1:GDZ WY001-Bolsover-FM WY001-Bolsover-v3.cls October22,2003 14:59 CONTENTS xi FromGlucosetoPyruvate:Glycolysis, 284 ReviewQuestions, 322 GlycolysisWithoutOxygen, 286 AnswerstoReviewQuestions, 324 GlycogenCanProvideGlucosefor Glycolysis, 288 15 THEACTIONPOTENTIAL, 325 GlucoseMayBeOxidizedtoProduce TheCalciumActionPotentialinSeaUrchin PentoseSugars, 289 Eggs, 325 FromFatstoAcetyl-CoA:β Oxidation, 290 EffectofEggTransmembraneVoltageon AminoAcidsasAnotherSourceof SpermFusion, 325 MetabolicEnergy, 292 TheVoltage-GatedCalcium MakingGlucose:Gluconeogenesis, 295 Channel, 327 MakingGlycogen:Glycogenesis, 298 TheCalciumActionPotential, 328 MakingFattyAcidsandGlycerides, 300 TheVoltage-GatedSodiumChannelin SynthesisofAminoAcids, 300 NerveCells, 330 CarbonFixationinPlants, 302 TheVoltage-GatedSodiumChannel, 330 ControlofEnergyProduction, 303 ElectricalTransmissiondownaNerve CellAxon, 332 FeedbackandFeedforward, 303 MyelinationandRapidActionPotential NegativeFeedbackControlof Transmission, 334 Glycolysis, 304 Summary, 337 FeedforwardControlinMuscle Cells, 304 FurtherReading, 338 Summary, 306 ReviewQuestions, 338 FurtherReading, 306 AnswerstoReviewQuestions, 339 ReviewQuestions, 307 16 AnswerstoReviewQuestions, 308 INTRACELLULARSIGNALING, 341 Calcium, 341 14 IONSANDVOLTAGES, 309 CalciumCanEnterfromtheExtracellular Medium, 341 ThePotassiumGradientandtheResting Voltage, 309 CalciumCanBeReleasedfromthe EndoplasmicReticulum, 344 PotassiumChannelsMakethePlasma MembranePermeabletoPotassium ProcessesActivatedbyCytosolic Ions, 310 CalciumAreExtremelyDiverse, 348 ConcentrationGradientsandElectrical ReturnofCalciumtoResting VoltageCanBalance, 311 Levels, 350 TheChlorideGradient, 314 CyclicAdenosineMonophosphate, 350 GeneralPropertiesofChannels, 314 CyclicGuanosineMonophosphate, 353 GeneralPropertiesofCarriers, 316 MultipleMessengers, 353 TheGlucoseCarrier, 316 BiochemicalSignaling, 353 TheSodium–CalciumExchanger, 317 ReceptorTyrosineKinasesandtheMAP KinaseCascade, 353 CarrierswithanEnzymaticAction: TheCalciumATPase, 318 GrowthFactorsCanTriggeraCalcium Signal, 356 Summary, 322 ProteinKinaseBandtheGlucose FurtherReading, 322 Transporter:HowInsulinWorks, 356 P1:GDZ WY001-Bolsover-FM WY001-Bolsover-v3.cls October22,2003 14:59 xii CONTENTS Crosstalk—SignalingPathwaysor FurtherReading, 398 SignalingWebs?, 357 ReviewQuestions, 398 Summary, 359 AnswerstoReviewQuestions, 400 FurtherReading, 360 ReviewQuestions, 360 19 CELLCYCLEANDCONTROLOF AnswerstoReviewQuestions, 361 CELLNUMBER, 401 StagesofMitosis, 402 17 INTERCELLULAR MeiosisandFertilization, 404 COMMUNICATION, 363 Meiosis, 405 ClassifyingTransmittersandReceptors, 363 FertilizationandInheritance, 406 IonotropicCellSurfaceReceptors, 364 DominantGeneticDisease, 408 MetabotropicCellSurface CrossingOverandLinkage, 408 Receptors, 365 ControloftheCellDivisionCycle, 408 IntracellularReceptors, 365 MolecularRegulationoftheG2/M IntercellularCommunicationinAction: (Interphase/Mitosis)CellCycle TheGastrocnemiusMuscle, 365 ControlPoint, 410 TellingtheMuscletoContract: WhatAbouttheG1/SControlPoint?, 412 TheActionofMotoneurones, 367 Apoptosis, 415 ControllingtheBloodSupply:Paracrine InstructedDeath:DeathDomain Transmitters, 368 Receptors, 416 NewBloodVesselsinGrowing DefaultDeath:AbsenceofGrowth Muscle, 371 Factors, 416 SynapsesBetweenNeurons, 372 TheSickAreLefttoDie: Summary, 376 Stress-ActivatedApoptosis, 417 FurtherReading, 377 Summary, 419 ReviewQuestions, 377 FurtherReading, 420 AnswerstoReviewQuestions, 378 ReviewQuestions, 420 AnswerstoReviewQuestions, 421 18 MECHANICALMOLECULES, 381 TheCytoskeletonisBothStrongand 20 CASESTUDY:CYSTIC Motile, 381 FIBROSIS, 423 Microtubules, 381 Introduction, 423 Microtubule-BasedMotility, 386 CysticFibrosisisaSevereGenetic CiliaandFlagella, 386 Disease, 423 IntracellularTransport, 389 TheFundamentalLesioninCysticFibrosis Microfilaments, 390 LiesinChlorideTransport, 424 MuscleContraction, 393 HominginontheCFGene, 425 CellLocomotion, 395 CloningtheGeneforCF, 426 CytoplasmicStreaming, 395 TheCFTRGeneCodesforaChlorideIon Channel, 426 IntermediateFilaments, 396 GeneTherapyforCF, 427 AnchoringCellJunctions, 396 DiagnosticTestsforCF, 431 Summary, 398
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