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3. Garland Science Vice President:Denise Schanck
AssistantEditor: Sigrid Masson Production Editor and Layout: Emma
leffcock Senior Publisher: JackieHarbor Illustrator: Nigel Orme
Designer:Matthew McClements,Blink Studio,Ltc. Editors:Marjorie
Anderson and SherryGranum Copy Editor: Bruce Goatly Indexer:
Merrall-RossInternational, Ltd. PermissionsCoordinator: Marv
Disoenza Cell Biology Interactiue Artistic and Scientific
Direction: PeterWalter Narrated by: Julie Theriot Production Design
and Development: Michael Morales @2008, 2002 by Bruce Alberts,
Alexander Johnson, Julian Lewis, Martin Rafi Keith Roberts,and
PeterWalter. @ f 983, f 989, 1994by 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 of Biochemistry and Biophysicsat the university of
california, san Francisco.For 12years,he servedas Presidentofthe
u.s. NationalAcademy ofSciences (1993-2005). Alexander Johnson
received his Ph.D. from Harvard University and is professor of
Microbiology and Immunology and Director of the Biochemistry cell
Biology,Genetics, and Developmental Biology Graduate Program at the
University of california, San Francisco. Iulian Lewis received his
D.Phil. from the University of Oxford and is a Principal
Scientistat the London ResearchInstitute of CancerResearchUK.
Martin Raffreceived his M.D. from McGill University and is at the
Medical Research Council Laboratory for Molecular Cell Biology and
the Biology Department at University College London. Keith Roberts
received his Ph.D. from the University of Cambridge and is 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 the Department of Biochemistry and Biophysics at
the University of california, san Francisco, and an Investigator of
the Howard Hughes Medical Institute. All rights reserved. No part
of this book covered by the copyright heron may be reproduced or
used in any format in any form or by any means-graphic, electronic,
or mechanical, including photocopying, recording, taping, or
information storage and retrieval systems-without permission of the
publisher. Library of CongressCataloging-in-Publication Data
Molecularbiologyof the cell / BruceAlberts... [etal.].--5th ed.
p.cm
ISBN978-0-8153-4r05-5(hardcover)---ISBN978-0-8f5g-4t06_Z(paperback)
L Cytology.2. Molecular biology.I. Alberts, Bruce. QHsB1.2.M642008
571.6--dc22 2007005475CIP Published 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 Statesof America 15 14
13 12 lt 1 0 I B 7 6 5 4 3 2 |
4. Preface In many respects,we understandthe structureof the
universebetter than the workingsof living cells.Scientistscan
calculatethe ageof the Sunand predict when it will ceaseto
shine,but we cannotexplainhow it is that a human being may live for
eighty yearsbut a mouse for only two. We know the complete
genomesequencesof theseandmany other species,but we still
cannotpredict how a cell will behaveif we mutate a
previouslyunstudiedgene.Starsmay be l0a3times bigger,but
cellsaremore complex,more intricately structured,and
moreastonishingproductsof
thelawsofphysicsandchemistry.Throughhered- ity and natural
selection,operatingfrom the beginningsof life on Earthto the
presentday-that is,for about20Voof theageof the universe-living
cellshave been progressivelyrefining and extending their molecular
machinery and recordingthe resultsof their experimentsin the
geneticinstructionsthey pass on to their progeny. With eachedition
of this book,we marvelat the new information that cell
biologistshavegatheredin just a fewyears.But we areevenmore
amazedand dauntedatthe sophisticationofthe mechanismsthat
weencounter.Thedeeper weprobeinto the cell,themorewe reafizehowmuch
remainsto beunderstood. In the daysof our innocence,working on the
first edition,we hailedthe identi- fication of asingleprotein-a
signalreceptol say-as agreatstepforward'Now we appreciatethat
eachprotein isgenerallypart of acomplexwith manyothers, working
togetherasa system,regulatingone another'sactivitiesin subtleways,
andheldin specificpositionsbybinding to scaffoldproteinsthat
givethechem- ical factorya definite
spatialstructure.Genomesequencinghasgivenusvirtu- ally
completemolecularparts-listsfor many different
organisms;geneticsand biochemistryhavetold us a greatdeal about
what thoseparts are capableof individually and which onesinteract
with which others;but we haveonly the most primitive graspof the
dynamicsof thesebiochemicalsystems,with all their interlocking
control loops.Therefore,although there are great achieve- mentsto
report,cellbiologistsfaceevengreaterchallengesfor the future. In
this edition,wehaveincludednewmaterialon manytopics,rangingfrom
epigenetics,histonemodifications,smallRNAs,and
comparativegenomics,to
geneticnoise,cytoskeletaldlmamics,cell-cyclecontrol,apoptosis,stemcells,
and novelcancertherapies.As in previouseditions,we havetried
aboveall to givereadersa conceptualframeworkfor the massof
information that we now haveabout cells.This meansgoingbeyondthe
recitationof facts.Thegoalis to learn how to put the facts to
use-to reason,to predict, and to control the behaviorof living
systems. Tohelp readerson the wayto an activeunderstanding,we
havefor the first time incorporatedend-of-chapterproblems,written
by Iohn Wilsonand Tim Hunt. Theseemphasizea quantitativeapproachand
the art of reasoningfrom experiments.A
companionvolume,MolecularBiologyof theCelI,Fifth Edition:
TheProblemsBook0SBN978-0-8153-4110-9),by the sameauthors,givescom-
pleteanswersto theseproblemsand alsocontainsmorethan 1700additional
problemsandsolutions. A further major adjunctto the main book is
the attachedMediaDVD-ROM
disc.Thisprovideshundredsofmoviesandanimations,includingmanythat
are new in this edition, showingcellsand cellularprocessesin action
and bringing thetextto life;thediscalsonowincludesall
thefiguresandtablesfrom themain
5.
book,pre-loadedintoPowerPoint@presentations.Otherancillariesavailablefor
thebookincludeabankof testquestionsandlectureoutlines,availableto
qual- ifiedinstructors,anda setof
200full-coloroverheadtransparencies. Perhapsthe biggestchangeis in
the physicalstructureof the book. In an effort to make the
standardStudentEdition somewhatmore portable,we are providing
chapters 2r-25, coveringmulticellular systems,in electronic (pDF)
form on the accompanyingdisc,while retainingin theprinted
volumechapters l-20, coveringthe core of the usual cell biology
curriculum. But we should emphasizethat the final
chaptershavebeenrevisedand updatedasthoroughly asthe restof the
book andwe sincerelyhopethat theywill beread!A Reference
Edition(ISBN97s-0-8153-4r11-6),containingthefull setof
chaptersasprinred pages,is alsoavailablefor thosewho preferit. Full
detailsof the conventionsadoptedin the book aregivenin the Noteto
the Readerthat followsthis Preface.Asexplainedthere,we havetakena
drastic approachin confrontingthe differentrulesfor thewriting of
genenamesin dif- ferent species:throughout this book, we use the
same style, regardlessof species,andoftenin defianceofthe
usualspecies-specificconventions. As always,we areindebtedto many
people.Full acknowledgmentsfor sci-
entifichelparegivenseparately,but wemustheresingleout
someexceptionally important contributions:Iulie Theriot is almost
entirely responsiblefor chap- ters 16(cytoskeleton)and 24
(Pathogens,Infection,and InnateImmunity), and David
Morganlikewisefor chapter 17(cell cycle).wallaceMarshalland Laura
Attardi provided substantialhelp with chapters 8 and
20,respectively,as did Maynardolsonfor
thegenomicssectionofchapter4,Xiaodongwangfor chap- ter
18,andNicholasHarberdfor theplantsectionof Chapter15. we also owe a
huge debt to the staff of Garlandscienceand otherswho helped
convert writers' efforts into a polished final product.
Deniseschanck directedthe whole enterpriseand shepherdedthe
waywardauthorsalongthe roadwith wisdom,skill,and kindness.Nigelorme
put the artworkinto its final form andsupervisedthevisualaspectsof
thebook,includingthebackcover,with his usual flair. Matthew
Mcclements designedthe book and its front cover. EmmaJeffcocklaid
out its pageswith extraordinaryspeedand unflappableeffi-
ciency,dealingimpeccablywithinnumerablecorrections.MichaelMoralesman-
agedthe transformationof a massof animations,video clips,and
othermateri- als into a user-friendly DVD-ROM. Eleanor Lawrence and
sherry Granum
updatedandenlargedtheglossary.JackieHarborandSigridMassonkeptusorga-
nized.AdamSendroffkeptusawareofour readersandtheir
needsandreactions.
MarjorieAnderson,BruceGoatly,andsherryGranumcombedthetextfor obscu-
rities,infelicities,and errors.we thank them all, not only for
their professional skill and dedicationand for efficiencyfar
surpassingour own,but alsofor their
unfailinghelpftrlnessandfriendship:theyhavemadeit apleasureto work
on the book. Lastly,and with no lessgratitude,we thank our
spouses,families,friends andcolleagues.without
theirpatient,enduringsupport,wecouldnot havepro- ducedanyof the
editionsof thisbook.
6. Contents Speci.alFeatures DetailedContents Acknowledgments A
Noteto theReader PARTI I. 2. 3. PARTII 4. 5. 6. 7. PARTIII B. 9.
PARTIV 10. 11. 12. 13. 14. 15. 16. t7. tB. PARTV 19. 20. 2I, 22.
23, 24. 25. Glossary Index TabIes uiii ix xxui xxxi
INTRODUCTIONTOTHECELL CellsandGenomes CellChemistryandBiosynthesis
Proteins BASICGENETICMECHANISMS DNA,Chromosomes,andGenomes DNA
Replication,Repair,and Recombination HowCellsReadthe
Genome:FromDNAto Protein Controlof GeneExpression METHODS
Manipulating Proteins,DNA,and RNA VisualizingCells
INTERNALORGANIZATIONOFTHECELL MembraneStructure MembraneTransportof
SmallMoleculesandthe Electrical Propertiesof Membranes
IntracellularCompartmentsand ProteinSorting
IntracellularVesicularTraffic
EnergyConversion:MitochondriaandChloroplasts Mechanismsof
CellCommunication TheCytoskeleton The CellCycle Apoptosis
CELLSINTHEIRSOCIALCONTEXT Celllunctions, CellAdhesion,and the
ExtracellularMatrix Cancer Chapters2I-25 availableon MediaDVD-ROM
SexualReproduction:Meiosis,GermCells,andFertilization Developmentof
MulticellularOrganisms
SpecializedTissues,StemCells,andTissueRenewal Pathogens,Infection,
and InnateImmunity TheAdaptiveImmune System The Genetic Code,Amino
Acids 195 263 329 41I I 45 t25 50r 579 6t7 651 695 749 Br3 879 965
1053 1115 1131 I205 1269 r305 L4l7 1485 1539 G-1 L1 T-1
7. SpecialFeatures Table l-l Table l-2 Table2-1 Table2-2 Table
2-3 Table2-4 Panel2-l Panel2-2 Panel 2-3 Panel2-4 Panel 2-5 Panel
2-6 Panel2-7 Panel 2-B Panel2-9 Panel3-l Panel 3-2 Table3-1 Panel
3-3 Table4-l Table 5-3 Table6-l PanelB-l Table 10-l Thble11-l
Panel1l-2 Panelll-3 Table l2-l Table l2-2 Table l4-l Panel l4-l
Thble r5-5 Panel 16-2 Panel 16-3 Table I7-2 Panel l7-l
SomeGenomesThatHaveBeenCompletelySequenced TheNumbersof
GeneFamilies,classifiedby Function,ThatArecommon to All
ThreeDomainsof the LivingWorld CovalentandNoncovalentChemicalBonds
TheTypesof MoleculesThatForma BacterialCell
ApproximateChemicalCompositionsof aTypicalBacteriumandaTypical
MammalianCell RelationshipBetweenthe StandardFree-EnergyChange,AG,
andthe EquilibriumConstant
ChemicalBondsandGroupsCommonlyEncounteredin BiologicalMolecules
WaterandItsInfluenceon theBehaviorof BiologicalMolecules
ThePrincipalTypesofweakNoncovalentBondsthat HoldMacromolecules
Together An Outlineof Someof theTypesof SugarsCommonlyFoundin Cells
FattyAcidsandOtherLipids A Surveyof the Nucleotides
FreeEnergyandBiologicalReactions Detailsof the t0 Stepsof
Glycolysis TheCompleteCitricAcidCycle The20AminoAcidsFoundin
Proteins FourDifferentWaysof Depictinga SmallProtein,the SH2Domain
SomeCommonTypesof Enzymes Someof the MethodsUsedto StudyEnzymes
SomeVitalStatisticsfor theHumanGenome
ThreeMajorClassesofTransposableElements PrincipalTlpesof
RNAsProducedin Cells Reviewof ClassicalGenetics pp.
ApproximateLipid Compositionsof DifferentCellMembranes A
Comparisonof Ion
ConcentrationsInsideandOutsideaTypicalMammalianCell TheDerivationof
the NernstEquation SomeClassicalExperimentson the SquidGiantAxon
RelativeVolumesOccupiedby theMajorIntracellularCompartmentsin a
Liver Cell(Hepatocyre) RelativeAmounts of MembraneTypesin
TwoKindsof Eucaryoticcells ProductYieldsfrom the Oxidationof
SugarsandFats RedoxPotentials TheRasSuperfamilyof MonomericGTpases
The Polymerizationof Actin andTubulin pp. 978-979
AccessoryProteinsthat ControltheAssemblyandpositionof Cvtoskeletal
Filaments pp.994-995 Summaryof theMajorCell-CycleRegulatoryproteins
p. 1066 The Princinle Stases of M Phasp (Mitnsis nnrl Crrfnlrinpcic
in qn Animal /-oll nn rATo rA?a p. 18 p.24 p.53 p.55 p.63 p.77 pp.
106-107 pp. 108-109 pp. pp. pp. pp. pp. pp. pp. pp. pp. r10-111
112-113 I l4-1I5 I 16-1r7 I IB-t 19 r20-I2l I22-t23 tzg-729 132-133
p.159 162-163 p.206 p.318 p.336 554-555 p.624 p.652 p.670 p.679
p.697 p.697 p.824 p.830 p.926 pp.