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1、LeadingEdgeReviewHallmarksofCancer:TheNextGenerationDouglasHanahan1,2,*andRobertA.Weinberg3,*1TheSwissInstituteforExperimentalCancerResearch(ISREC),SchoolofLifeSciences,EPFL,LausanneCH-1015,Switzerland2TheDepartmentofBiochemistry&Biophysics,UCSF,SanFrancisco,CA94158,USA3Whitehea
2、dInstituteforBiomedicalResearch,Ludwig/MITCenterforMolecularOncology,andMITDepartmentofBiology,Cambridge,MA02142,USA*Correspondence:dh@epfl.ch(D.H.),weinberg@wi.mit.edu(R.A.W.)DOI10.1016/j.cell.2011.02.013Thehallmarksofcancercomprisesixbiologicalcapabilitiesacquiredduringthemulti
3、stepdevelop-mentofhumantumors.Thehallmarksconstituteanorganizingprincipleforrationalizingthecomplexitiesofneoplasticdisease.Theyincludesustainingproliferativesignaling,evadinggrowthsuppressors,resistingcelldeath,enablingreplicativeimmortality,inducingangiogenesis,andacti-vatingi
4、nvasionandmetastasis.Underlyingthesehallmarksaregenomeinstability,whichgeneratesthegeneticdiversitythatexpeditestheiracquisition,andinflammation,whichfostersmultiplehall-markfunctions.Conceptualprogressinthelastdecadehasaddedtwoemerginghallmarksofpotentialgeneralitytothislist—rep
5、rogrammingofenergymetabolismandevadingimmunedestruction.Inadditiontocancercells,tumorsexhibitanotherdimensionofcomplexity:theycontainarepertoireofrecruited,ostensiblynormalcellsthatcontributetotheacquisitionofhall-marktraitsbycreatingthe‘‘tumormicroenvironment.’’Recognitionofthe
6、widespreadapplicabilityoftheseconceptswillincreasinglyaffectthedevelopmentofnewmeanstotreathumancancer.INTRODUCTIONvatedbythesedevelopments,wenowrevisittheoriginalhall-marks,considernewonesthatmightbeincludedinthisroster,Wehaveproposedthatsixhallmarksofcancertogetherconsti-andex
7、panduponthefunctionalrolesandcontributionsmadetuteanorganizingprinciplethatprovidesalogicalframeworkforbyrecruitedstromalcellstotumorbiology.understandingtheremarkablediversityofneoplasticdiseases(HanahanandWeinberg,2000).ImplicitinourdiscussionwasHALLMARKCAPABILITIESCONCEPTUALP
8、ROGRESSthenotionthatasnormalcellsevolveprogress