Table Of ContentEncyclopediaofCancer
(2ndedition)
MANFRED SCHWAB (Ed.)
Encyclopedia of Cancer
(2nd edition)
With 979 Figures* and 210 Tables
*ForcolorfigurespleaseseeourElectronicReferenceonwww.springerlink.com
Editor:
ManfredSchwab
ProfessorforGenetics
DirectorDivisionofTumourGenetics(B030)
GermanCancerResearchCenter(DKFZ)
ImNeuenheimerFeld280
69120Heidelberg
Germany
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Preface To The First Edition
Cancer,althoughadreadfuldisease,isatthesametimeafascinatingbiologicalphenotype.Around1980,cancer
wasfirstattributedtomalfunctioninggenesand,subsequently,cancerresearchhasbecomeamajorareaofscientific
researchsupportingthefoundationsofmodernbiologytoagreatextent.Tounravelthehumangenomesequence
wasoneofthoseextraordinarytasks,whichhaslargelybeenfuelledbycancerresearch,andmanyofthefascinating
insightsintothegeneticcircuitsthatregulatedevelopmentalprocesseshavealsoemergedfromresearchoncancer.
Diverse biological disciplines such as cytogenetics, virology, cell biology, classical and molecular genetics,
epidemiology, biochemistry, together with the clinical sciences, have closed ranks in their search of how cancer
developsandtofindremediestostoptheabnormalgrowththatischaracteristicofcancerouscells.Intheattemptto
establish how, why and when cancer occurs, a plethora of genetic pathways and regulatory circuits have been
discovered that are necessary to maintain general cellular functions such as proliferation, differentiation and
migration. Alterations of this fine-tuned network of cascades and interactions, due to endogenous failure or to
exogenouschallengesbyenvironmentalfactors,maydisableanymemberofsuchregulatorypathways.Thiscould,
for example, induce the death of the affected cell, may mark it for cancerous development or may immediately
provideitwithagrowthadvantagewithinaparticulartissue.
Recentdevelopmentshaveseenthemergerofbasicandclinicalscience.Oftheformer,particularlygeneticshas
providedinstrumentalandanalyticaltoolswithwhichtoassesstheroleofenvironmentalfactorsincancer,torefine
andenablediagnosispriortothedevelopmentofsymptomsandtoevaluatetheprognosisofpatients.Hopefully,
evenbetterstrategiesforcausaltherapywillbecomeavailableinthefuture.Mergingthebasicandclinicalscience
disciplinestowardsthecommongoaloffightingcancer,callsforacomprehensivereferencesourcetoservebothas
atooltoclosethelanguagegapbetweenclinicalandbasicscienceinvestigatorsandasaninformationplatformfor
thestudentandtheinformedlaypersonalike.Obviouslythiswasanextremelyambitiousgoal,andtheimmense
progressinthefieldcannotalwaysbeportrayedinlinewiththelatestdevelopments.TheaimoftheEncyclopediais
toprovidethereaderwith anentrancepointtoaparticular topic.Itshouldbeofvalue tobothbasicandclinical
scientistsworkinginthefieldofcancerresearch.Additionallybothstudentsandlecturersinthelifesciencesshould
benefithighlyfromthisdatabase.IthereforehopethatthisEncyclopediawillbecomeanessentialcomplementto
existingscienceresources.
Theattemptstoidentifythemechanismsunderlyingcancerdevelopmentandprogressionhaveproducedawealth
of facts, and no single individual is capable of addressing the immense breadth of the field with undisputed
authority.Hence,the‘EncyclopedicReferenceofCancer’istheworkofmanyauthors,allofwhomareexpertsin
their fields and reputable members of the international scientific community. Each author contributed a large
number of keyword definitions and in-depth essays and in so doing it was possible to cover the broad field of
cancer-relatedtopicswithinasinglepublication.Obviouslythisapproachentailsaformofpresentation,inwhich
theauthorhasthefreedomtosetprioritiesandtopromoteanindividualpointofview.Thisismostobviouswhenit
comestonomenclature,particularlythatofgenesandproteins.Althoughtheeditorialintentionwastoapplythe
nomenclatureoftheHumanGenomeOrganisation(HUGO),themorevigorousexecutionofthisattempthasbeen
lefttofutureendeavours.
In the early phase of planning the Encyclopedia, exploratory contacts to potential authors produced an
overwhelminglypositiveresponse.Thesubsequentcontactwithalmost300contributoryauthorswasamarvellous
experience,andIamextremelygratefulfortheirexcellentandconstructivecooperation.Animportantelementin
the preparation of the Encyclopedia has been the competent secretarial assistance of Hiltrud Wilbertz of the
Springer-VerlagandofIngridCederlundandCorneliaKirchneroftheDKFZ.Withgreatattentiontodetailthey
helpedtokeeptrackofthetechnicalaspectsinthepreparationofthemanuscript.Itwasapleasuretoworkwiththe
Springercrew,includingDr.RolfLangeastheEditorialDirector(Medicine)andDr.ThomasMager,SeniorEditor
forEncyclopediasandDictionaries.InparticularIwishtothankDr.WalterReuss,whountiringlyhasmasteredall
aspects and problems associated with the management of the numerous manuscripts that were received from
authorsoftheinternationalscientificcommunity.Ithasbeensatisfyingandattimescomfortingtoseehowhemade
illustrationfilescomealive.ThanksalsotoDr.ClaudiaLangewho,beingherselfaknowledgeablecellbiologist,
hasworkedasthescientificeditor.HercommitmentandinteresthavesubstantiallyimprovedthisEncyclopedia.
vi PrefaceToTheFirstEdition
Asafinalword,Iwouldliketostressthatalthoughsubstantialeffortshavebeenmadetocomposefactuallycorrect
andwellunderstandablepresentations,theremaybeplaceswhereadefinitionisincompleteoraphraseinanessay
is flawed. All contributors to this Encyclopedia will be extremely happy to receive possible corrections, or
revisions,inorderforthemtobeincludedinanyfutureeditionsofthe‘EncyclopedicReferenceofCancer’.
Heidelberg,September2001
MANFREDSCHWAB
Preface To The Second Edition
GiventheoverwhelmingsuccessoftheFirstEditionoftheCancerEncyclopedia,whichappearedin2001,andthe
amazingdevelopmentinthedifferentfieldsofcancerresearch,ithasbeendecidedtopublishasecondfullyrevised
andexpandededition,followingtheprincipleconceptofthefirsteditionthathasprovensosuccessful.
Recent developments are seeing a dynamic merging of basic and clinical science, with translational research
increasingly becoming a new paradigm in cancer research. The merging of different basic and clinical science
disciplines towards the common goal of fighting against cancer has long ago called for the establishment of a
comprehensive reference source both as a tool to close the language gap between clinical and basic science
investigatorsandasaplatformofinformationforadvancedstudentsandinformedlaymenalike.Itisintendedtobe
aresourceforallinterestedininformationbeyondtheirspecificownexpertise.
WhiletheFirstEditionhadfeaturedcontributionsfromapproximately300scientists/cliniciansinoneVolume,the
SecondEditionincludesmorethan1000contributorsin4VolumeswithanA–Zformatofmorethan7000entries.
Itprovidesdefinitionsofcommonacronymsandshortdefinitionsofbothrelatedtermsandprocessesintheform
of keyword entries. A major information source are detailed essays that provide comprehensive information
on syndromes, genes and molecules, and processes and methods. Each essay is well-structured, with extensive
cross-referencing between entries. Essays represent original contributions by the corresponding authors, all
distinguishedscientistsintheirownfield,Editorialinputhasbeencarefullyrestrictedtoformalaspects.
ApanelofFieldEditors,eachaneminentinternationalexpertforthecorrespondingfield,hasservedtoensurethe
presentationoftimelyandauthoritativeEncyclopediaentries.Thesenewtraitsarelikelytomeettheexpectancethat
awidecommunityhastowardsacancerreferenceworks.
AnimportantelementinthepreparationoftheEncyclopediahasbeenthecompetentsupportbytheSpringercrew,
Dr.MichaelaBilic,SaskiaEllisand,lately,JanaSimniok.Iamextremelygratefulfortheirexcellentandpleasant
cooperation.
TheCancerEncyclopedia,SecondEdition,willbeaccessiblebothinprintandonlineversions.Clinicians,research
scientistsandadvancedstudentswillfindthisanamazingresourceandahighlyinformativereferencetocancer.
Heidelberg,March15,2008
MANFREDSCHWAB
Editor in Chief
ManfredSchwab
TumorGenetics
GermanCancerResearchCenter,DKFZ
Heidelberg
Germany
[email protected]
Field Editors
STEFAN BARTH Houston,TX
DepartmentofPharmaceuticalProduct USA
Development,Fraunhofer-Institute [email protected]
forMolecularBiologyand
AppliedEcology JOSEPH R. LANDOLPH
Aachen DepartmentsofMolecularMicrobiologyand
Germany Immunology,andPathology;USC/Norris
[email protected] ComprehensiveCancerCenter,KeckSchoolof
Medicine;DepartmentofMolecularPharmacology
andPharmaceuticalSciences,SchoolofPharmacy
PAOLO BOFFETTA HealthSciencesCampus
Lifestyle,EnvironmentandCancerGroup
UniversityofSouthernCalifornia
InternationalAgencyforResearchonCancer
LosAngeles,CA
Lyon
USA
France
[email protected]
[email protected]
HENRY LYNCH
PreventiveMedicineandPublicHealth
GRAHAM A. COLDITZ
HereditaryCancerInstitute
WashingtonUniversityinSt.Louis
CreightonUniversity
St.Louis,MO
Omaha,NE
USA
USA
[email protected]
[email protected]
ROY J. DUHÉ PAUL G. MURRAY
DepartmentofPharmacologyandToxicology CRUKInstituteforCancerStudies
UniversityofMississippiMedicalCenter TheMedicalSchool
Jackson,MS UniversityofBirmingham
USA Birmingham
[email protected] UK
[email protected]
KENT HUNTER SAUL SUSTER
MetastasisSusceptibilitySection
TheOhioStateUniversity
LaboratoryofCancerBiology&Genetics
Columbus,OH
CCR/NCI/NIH
USA
Bethesda,MD
[email protected]
USA
[email protected]
ANDREW THORBURN
UniversityofColoradoatDenverandHealth
JESPER JURLANDER SciencesCenter
DepartmentofHematologyandtheLeukemia Aurora,CO
USA
Laboratory
[email protected]
Rigshospitalet
Copenhagen
Denmark MARTIN TOBI
UniversityofPennsylvaniaSchoolofMedicine
[email protected]
ChiefGISection
PhiladelphiaVAMC
RAKESH KUMAR Philadelphia,PA
MolecularandCellularOncology USA
TheUniversityofTexasMDAndersonCancerCenter [email protected]
ListofContributors xiii
List of Contributors
VESAAALTONEN FARRUKHAFAQ
DepartmentofAnatomy DepartmentofDermatology
InstituteofBiomedicine UniversityofWisconsin
UniversityofTurku MedicalSciencesCenter
Turku Madison,WI
Finland USA
[email protected] [email protected]
CORYABATE-SHEN CHAPLAAGARWAL
CenterforAdvancedBiotechnologyandMedicine DepartmentofPharmaceuticalSciences,SchoolofPharmacy
UMDMJ–RobertWoodJohnsonMedicalSchool UniversityofColoradoHealthSciencesCenter
Piscataway,NJ Denver,CO
USA USA
[email protected] [email protected]
RAJESHAGARWAL
PHILLIPH.ABBOSH DepartmentofPharmaceuticalSciences,SchoolofPharmacy
DepartmentofPathologyandLaboratoryMedicine
UniversityofColoradoHealthSciencesCenter
IndianaUniversitySchoolofMedicine
Denver,CO
Indianapolis,IN
USA
USA
[email protected]
[email protected]
PATRIZIAAGOSTINIS
FRITZABERGER CatholicUniversityofLeuven
DepartmentofMolecularBiology Leuven
UniversityofSalzburg Belgium
Salzburg,Austria [email protected]
[email protected]
TERJEAHLQUIST
HINRICHABKEN DepartmentofCancerPrevention
TumorGenetics,ClinicIInternalMedicineandCenterfor Rikshospitalet-RadiumhospitaletMedicalCentre
MolecularMedicineCologne,UniversityofCologne Oslo
Cologne,Germany Norway
[email protected] [email protected]
AMALM.ABU-GHOSH SHAHIDAHMED
DepartmentsofPediatricHematologyandOncology SaskatoonCancerCenter
LombardiComprehensiveCancerCenter UniversityofSaskatchewan
GeorgetownUniversity Saskatoon,SK
WashingtonDC,NW Canada
USA [email protected]
[email protected]
JOOHONGAHNN
DepartmentofLifeScience
ROSITAACCARDI GwangjuInstituteofScienceandTechnology
InfectionsandCancerBiologyGroup
Buk-Gu,Gwangju
InternationalAgencyforResearchonCancer
Korea
Lyon
[email protected]
France
[email protected]
CEM AKIN
UniversityofMichigan
FILIPPOACCONCIA AnnArbor,MI
MolecularandCellularOncology USA
TheUniversityofTexasM.D.AndersonCancerCenter [email protected]
Houston,TX
USA
ENRIQUE DEALAVA
IFOM CentrodeInvestigacióndelCáncer-IBMCC
TheFIRCInstituteforMolecularOncology Salamanca
Milan Spain
Italy [email protected]