Часть 1 (B. Alberts, A. Johnson, J. Lewis и др. - Molecular Biology of The Cell (5th edition))

MolecularBiologyofFifth EditionMolecularBiologyofFifth EditionBruceAlbertsJohnsonAlexanderJulianLewisMartinRaffKeithRobertsPeterWalterWithproblemsbyJohnWilsonTimHuntGarlandScienceGroupTaylor& FrancisGarland ScienceVice President:Denise SchanckAssistant Editor: Sigrid MassonProduction Editor and Layout: Emma leffcockSenior Publisher: Jackie HarborIllustrator: Nigel OrmeDesigner: Matthew McClements, Blink Studio, Ltc.Editors: Marjorie Anderson and Sherry GranumCopy Editor: Bruce GoatlyIndexer: Merrall-Ross International, Ltd.Permissions Coordinator: Marv DisoenzaCell Biology InteractiueArtistic and Scientific Direction: PeterWalterNarrated by: Julie TheriotProduction Design and Development: Michael Morales@ 2008, 2002 by Bruce Alberts, Alexander Johnson, Julian Lewis, Martin RafiKeith Roberts, and PeterWalter.@ f 983, f 989, 1994 by Bruce Alberts, Dennis Bray, Iulian Lewis, Martin Raff,Keith Roberts, and lames D.

Watson.Bruce Alberts received his Ph.D. from Harvard university and is professor ofBiochemistry and Biophysics at the university of california, san Francisco.For12 years,he served as President ofthe u.s. NationalAcademy ofSciences (1993-2005).Alexander Johnson received his Ph.D. from Harvard University and is professor ofMicrobiology and Immunology and Director of the Biochemistry cell Biology, Genetics,and Developmental Biology Graduate Program at the University of california, SanFrancisco.

Iulian Lewis received his D.Phil. from the University of Oxford and is aPrincipal Scientist at the London ResearchInstitute of Cancer ResearchUK.Martin Raffreceived his M.D. from McGill University and is at the Medical ResearchCouncil Laboratory for Molecular Cell Biology and the Biology Department at UniversityCollege London.

Keith Roberts received his Ph.D. from the University of Cambridge andis Emeritus Fellow at the John Innes Centre, Norwich. peterWalter received his ph.D.from The Rockefeller University in Newyork and is professor and chairman of theDepartment of Biochemistry and Biophysics at the University of california, sanFrancisco, and an Investigator of the Howard Hughes Medical Institute.All rights reserved. No part of this book covered by the copyright heron may bereproduced or used in any format in any form or by any means-graphic, electronic, ormechanical, including photocopying, recording, taping, or information storage andretrieval systems-without permission of the publisher.Library of CongressCataloging-in-Publication DataMolecularbiology of the cell / BruceAlberts ...

[et al.].-- 5th ed.p.cmISBN 978-0-8153-4r05-5(hardcover)---ISBN978-0-8f5 g-4t06_Z(paperback)L Cytology.2. Molecular biology. I. Alberts, Bruce.QHsB1.2.M642008571.6--dc222007005475CIPPublished by Garland science, Taylor & Francis Group, LLC, an informa business,270 Madison Avenue, NewYork NY f 0016,USA, and 2 park Square,Milton park,Abingdon, OXl4 4RN, UK.Printed in the United States of America15 14 13 12 lt10 IB 7 6 5 4 3 2 |PrefaceIn many respects,we understand the structure of the universebetter than theworkings of living cells.Scientistscan calculatethe age of the Sun and predictwhen it will ceaseto shine,but we cannot explain how it is that a human beingmay live for eighty years but a mouse for only two.

We know the completegenomesequencesof theseand many other species,but we still cannot predicthow a cell will behaveif we mutate a previouslyunstudied gene.Starsmay bel0a3times bigger,but cells are more complex, more intricately structured,andmore astonishingproductsof the laws of physicsand chemistry.Through heredity and natural selection,operating from the beginningsof life on Earth to thepresentday-that is, for about 20Voof the ageof the universe-living cellshavebeen progressivelyrefining and extending their molecular machinery andrecording the results of their experimentsin the genetic instructions they passon to their progeny.With each edition of this book, we marvel at the new information that cellbiologistshave gatheredin just a few years.But we are even more amazedanddaunted at the sophisticationof the mechanismsthat we encounter.The deeperwe probe into the cell,the more we reafizehow much remainsto be understood.In the daysof our innocence,working on the first edition, we hailed the identification of a singleprotein-a signalreceptol say-as a greatstep forward' Nowwe appreciatethat eachprotein is generallypart of a complexwith many others,working togetheras a system,regulatingone another'sactivitiesin subtleways,and held in specificpositionsby binding to scaffoldproteins that givethe chemical factory a definite spatial structure.Genomesequencinghas given us virtually complete molecular parts-listsfor many different organisms;geneticsandbiochemistry have told us a great deal about what those parts are capableofindividually and which ones interact with which others; but we have only themost primitive grasp of the dynamics of these biochemical systems,with alltheir interlocking control loops.

Therefore,although there are great achievements to report, cell biologistsface evengreaterchallengesfor the future.In this edition, we haveincluded new material on many topics,rangingfromepigenetics,histonemodifications,small RNAs,and comparativegenomics,togeneticnoise,cytoskeletaldlmamics,cell-cyclecontrol, apoptosis,stem cells,and novel cancer therapies.As in previous editions, we have tried aboveall togive readersa conceptualframework for the mass of information that we nowhave about cells.This meansgoing beyond the recitation of facts.The goal is tolearn how to put the facts to use-to reason,to predict, and to control thebehavior of living systems.To help readerson the way to an activeunderstanding,we have for the firsttime incorporatedend-of-chapterproblems,written by Iohn Wilson and TimHunt. Theseemphasizea quantitative approach and the art of reasoningfromexperiments.A companion volume, MolecularBiologyof the CelI,Fifth Edition:by the sameauthors,givescomTheProblemsBook0SBN978-0-8153-4110-9),plete answersto theseproblemsand also containsmore than 1700additionalproblemsand solutions.A further major adjunct to the main book is the attachedMedia DVD-ROMdisc.This provideshundredsof moviesand animations,including manythat arenew in this edition, showingcells and cellular processesin action and bringingthe text to life; the disc alsonow includesall the figuresand tablesfrom the mainbook,pre-loadedinto PowerPoint@presentations.Otherancillariesavailableforthe book include a bank of test questionsand lectureoutlines,availableto qualified instructors,and a set of 200full-coloroverheadtransparencies.Perhapsthe biggestchange is in the physical structure of the book.

In aneffort to make the standard Student Edition somewhatmore portable, we areproviding chapters 2r-25, covering multicellular systems,in electronic (pDF)form on the accompanyingdisc,while retaining in the printed volume chaptersl-20, covering the core of the usual cell biology curriculum. But we shouldemphasizethat the final chaptershavebeen revisedand updated as thoroughlyas the rest of the book and we sincerelyhope that they will be read!A ReferenceEdition (ISBN97s-0-8153-4r11-6),containingthe full set of chaptersasprinredpages,is also availablefor thosewho prefer it.Full details of the conventionsadopted in the book are given in the Note tothe Readerthat follows this Preface.As explainedthere,we have taken a drasticapproachin confronting the different rules for the writing of genenamesin different species:throughout this book, we use the same style, regardlessofspecies,and often in defianceofthe usualspecies-specificconventions.As always,we are indebted to many people.

Full acknowledgmentsfor scientific help are given separately,but we must here singleout someexceptionallyimportant contributions: Iulie Theriot is almost entirely responsiblefor chapters 16 (cytoskeleton)and 24 (Pathogens,Infection, and Innate Immunity), andDavid Morgan likewisefor chapter 17 (cell cycle).wallace Marshall and LauraAttardi provided substantialhelp with chapters 8 and 20, respectively,as didMaynardolson for the genomicssectionof chapter4, Xiaodongwangfor chapter 18,and NicholasHarberdfor the plant sectionof Chapter15.we also owe a huge debt to the staff of Garland science and others whohelped convert writers' efforts into a polished final product.

Denise schanckdirected the whole enterpriseand shepherdedthe wayward authors along theroad with wisdom, skill, and kindness.Nigel orme put the artwork into its finalform and supervisedthe visualaspectsof the book,including the backcover,withhis usual flair. Matthew Mcclements designedthe book and its front cover.Emma Jeffcocklaid out its pageswith extraordinaryspeedand unflappableefficiency,dealingimpeccablywith innumerablecorrections.MichaelMoralesmanagedthe transformationof a massof animations,video clips, and other materials into a user-friendly DVD-ROM. Eleanor Lawrence and sherry Granumupdatedand enlargedthe glossary.JackieHarbor and SigridMassonkept us organized.Adam Sendroffkeptus awareofour readersand their needsand reactions.MarjorieAnderson,BruceGoatly,and sherry Granumcombedthe text for obscurities, infelicities, and errors.we thank them all, not only for their professionalskill and dedication and for efficiencyfar surpassingour own, but also for theirunfailing helpftrlnessand friendship:they havemadeit a pleasureto work on thebook.Lastly,and with no less gratitude, we thank our spouses,families, friendsand colleagues.without their patient,enduringsupport,we could not haveproducedany of the editionsof this book.ContentsSpeci.alFeaturesDetailed ContentsAcknowledgmentsA Note to the ReaderPARTIuiiiixxxuixxxiI.2.3.TOTHECELLINTRODUCTIONCellsand GenomesCellChemistryand BiosynthesisProteinsI45t25PARTII4.5.6.7.MECHANISMSBASICGENETICDNA, Chromosomes,and GenomesDNA Replication,Repair,and RecombinationHow CellsReadthe Genome:From DNA to ProteinControlof GeneExpression19526332941IPARTIIIB.9.METHODSManipulating Proteins,DNA, and RNAVisualizingCells50rPARTIV10.11.INTERNALORGANIZATIONOFTHECELLMembrane StructureMembraneTransportof SmallMoleculesand the ElectricalPropertiesof MembranesIntracellular Compartmentsand Protein SortingIntracellularVesicularTrafficEnergyConversion:Mitochondriaand ChloroplastsMechanismsof CellCommunicationThe CytoskeletonThe Cell CycleApoptosis12.13.14.15.16.t7.tB.PARTV19.20.2I,22.23,24.25.GlossaryIndexTabIesCONTEXTCELLSINTHEIRSOCIALCell lunctions, Cell Adhesion,and the ExtracellularMatrixCancerChapters2I-25 availableon Media DVD-ROMSexualReproduction:Meiosis,Germ Cells,and FertilizationDevelopmentof MulticellularOrganismsTissues,StemCells,and TissueRenewalSpecializedPathogens,Infection, and Innate ImmunityThe Adaptive Immune SystemThe Genetic Code,Amino Acids5796t7651695749Br3879965105311151131I2051269r305L4l714851539G-1L1T-1SpecialFeaturesTable l-lTable l-2Table 2-1Table2-2Table 2-3Table2-4Panel 2-lPanel2-2Panel 2-3Panel2-4Panel 2-5Panel 2-6Panel2-7Panel 2-BPanel 2-9Panel 3-lPanel 3-2Table 3-1Panel 3-3Table 4-lTable 5-3Table 6-lPanel B-lTable 10-lThble 11-lPanel 1l-2Panel ll-3Table l2-lTable l2-2Table l4-lPanel l4-lThble r5-5Panel 16-2Panel 16-3Table I7-2Panel l7-lSomeGenomesThat HaveBeenCompletelySequencedp.