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胚胎干细胞和成体干细胞标志物
来自 : 蚂蚁淘

Contents

  • EmbryonicStemCellMarkers
  • HematopoieticStemCellMarkers
  • Mesenchymal/StromalStemCellMarkers
  • NeuralStemCellMarkers
  • References

Whilestemcellsarebestdefinedfunctionally,anumberofmolecularmarkershavebeenusedtocharacterizevariousstemcellpopulations.

Althoughfunctionshaveyettobeascertainedformanyoftheseearlymarkers,theiruniqueexpressionpatternandtimingprovideausefultoolforscientiststoinitiallyidentifyaswellasisolatestemcells.Thismini-reviewsummarizesevidenceregardingtherolesofspecificmarkersindefiningembryonic,hematopoietic,mesenchymal/stromal,andneuralstemcellpopulations.Formostofthemoleculesdiscussed,studiesperformedbothinvitroandinvivosupporttheirsignificantroleincharacterizingstemcells.Untilmoreisknownaboutthenovelmarker-negativestemcellpopulation,however,uncertaintystillexistsregardingthebenefitsofusingthesemarkersaloneorinvariouscombinationswhenidentifyingandisolatingcellsforstemcellresearch.

EmbryonicStemCellMarkers

Oct-4:Oct-4(alsotermedOct-3orOct3/4),oneofthePOUtranscriptionfactors,wasoriginallyidentifiedasaDNA-bindingproteinthatactivatesgenetranscriptionviaacis-elementcontainingoctamermotif.1Itisexpressedintotipotentembryonicstemandgermcells.2,3AcriticallevelofOct-4expressionisrequiredtosustainstemcellself-renewalandpluripotency.4Differentiationofembryonicstem(ES)cellsresultsindown-regulationofOct-4,aneventessentialforaproperanddivergentdevelopmentalprogram.5Oct-4isnotonlyamasterregulatorofpluripotencythatcontrolslineagecommitment,butisalsothefirstandmostrecognizedmarkerusedfortheidentificationoftotipotentEScells.

SSEAs(StageSpecificEmbryonicAntigens):SSEAswereoriginallyidentifiedbythreemonoclonalantibodies(Abs)recognizingdefinedcarbohydrateepitopesassociatedwithlacto-andglobo-seriesglycolipids,SSEA-1,-3and-4.6SSEA-1isexpressedonthesurfaceofpreimplantation-stagemurineembryos(i.e.attheeightcellstage)andhasbeenfoundonthesurfaceofteratocarcinomastemcells,butnotontheirdifferentiatedderivatives.7,8Theoviductepithelium,endometriumandepididymis,aswellassomeareasofthebrainandkidneytubulesinadultmicehavealsobeenshowntobereactivewithSSEA-1Abs.9SSEA-3and-4aresynthesizedduringoogenesisandarepresentinthemembranesofoocytes,zygotesandearlycleavage-stageembryos.10,11BIOLOGicalrolesofthesecarbohydrate-associatedmoleculeshavebeensuggestedincontrollingcellsurfaceinteractionsduringdevelopment.6UndifferentiatedprimateEScells,humanECandEScellsexpressSSEA-3andSSEA-4,butnotSSEA-1.UndifferentiatedmouseEScellsexpressSSEA-1,butnotSSEA-3orSSEA-4.12,13

HematopoieticStemCellMarkers

CD34:ThecellsurfacesialomucinCD34
Figure1.AstructuremodelofCD133proposedbyMiragliaS.etal.30ThisproteinhasanextracellularN-terminus,5hydrophobictransmembranedomains,2smallcytoplasmicloops,2largeextracellularloopsandacytoplasmicC-termus.

hasbeenafocusofinteresteversinceitwasfoundexpressedonasmallfractionofhumanbonemarrowcells.14TheCD34+-enrichedcellpopulationfrommarrowormobilizedperipheralbloodappearsresponsIBLeformostofthehematopoieticactivity.14-21CD34hasthereforebeenconsideredtobethemostcriticalmarkerforhematopoieticstemcells(HSCs).CD34expressiononprimitivecellsisdown-regulatedastheydifferentiateintomaturecells.22ItisalsofoundonclonogenicProgenitors,however,andsomelineage-committedcells.23Althoughitsprecisefunctionisstillunknown,thepatternofexpressionofCD34suggeststhatitplaysasignificantroleinearlyhematopoiesis.22ThetheoryofCD34beingthemostprimitiveHSCmarker,however,hasrecentlybeenchallenged.Osawaetal.firstdemonstratedthatmurineHSCscouldbeCD34negative.24Inaddition,alowlevelofengraftmentandhematopoieticcapacityhasbeendemonstratedinhumanCD34-cells.25TransplantationstudiesalsoshowedrepopulatingactivityinaCD34-cellpopulationinfetalsheep.26Additionally,studieshaveshownthatbothmurineandhumanCD34+cellsmaybederivedfromCD34-cells.27,28Collectively,thesereportssuggestthepossibilitythatHSCsmaybeCD34+orCD34-andthatselectionofcellsexpressingCD34mightresultinexclusionofmoreprimitivestemcells.Nevertheless,almostallclinicalandexperimentalprotocolsincludingexvivoculture,genetherapy,andHSCtransplantationarecurrentlydesignedforcellpopulationsenrichedforCD34+cells.

Figure2.ThefamilyofABCtransportersischaracterizedbythepresenceofanATP-bindingcassetteregion,whichhydrolyzesATPtosupportenergy-dependentsubstrateexportationfromtheintracellularcytoplasmtotheextracellularspace.Full-lengthtransporterscontaintwomirrorimagehalvesthatareseparatedbyaflexiblelinkerregion(notshown).Half-transporters,e.g.ABCG2,functionashomo-orheterodimersandmaybelocalizedtotheplasmamembrane.

CD133:CD133,a120kDa,glycosylatedproteincontainingfivetransmembranedomains(Figure1),wasidentifiedinitiallybytheAC133monoclonalAb,whichrecognizesaCD34+subsetofhumanHSCs.29,30ACD133isoform,AC133-2,hasbeenrecentlyclonedandidentifiedastheoriginalsurfaceantigenrecognizedbytheAC133Ab.31CD133mayprovideanalternativetoCD34forHSCselectionandexvivoexpansion.ACD133+enrichedsubsetcanbeexpandedinasimilarmannerasaCD34+enrichedsubset,retainingitsmultilineagecapacity.32RecentstudieshaveofferedevidencethatCD133expressionisnotlimitedtoprimitivebloodcells,butdefinesuniquecellpopulationsinnon-hematopoietictissuesaswell.CD133+progenitorcellsfromperipheralbloodcanbeinducedtodifferentiateintoendothelialcellsinvitro.33Inaddition,humanneuralstemcellscanbedirectlyisolatedbyusingananti-CD133Ab.34

ABCG2:ABCG2(ATP-bindingcassettesuperfamilyGmember2)isadeterminantoftheHoechst-negativephenotypeofsidepopulation(SP)andfoundinawidevarietyofstemcells,includingHSC.35,36ABCG2isamemberofthefamilyofABCtransportersandwasfirstidentifiedinabreastcancercellline.37Itbelongstothehalf-transportergroupandisuniqueasitislocalizedtotheplasmamembrane(Figure2).38TheexpressionofABCG2appearsgreatestonCD34-cellsandisdown-regulatedwiththeacquisitionofCD34onthecellsurface.35Down-regulationinABCG2expressionisalsoobservedinvariouscommittedhematopoieticprogenitors.39ABCG2maythereforeserveasamorepromisingmarkerthanCD34forprimitiveHSCisolationandcharacterization.TheexpressionpatternofABCG2,however,isnotlimitedtoHSC.ABCG2expressionexclusivelycharacterizestheHoechstSPphenotypeincellsfromdiversesources,includingmonkeybonemarrow,mouseskeletalmuscleandEScells.35ThepotentialplasticityofSPcellshasbeendemonstratedbystudiesshowingthatcardiomyocytesandmusclecanberegeneratedfromtransplantedbonemarrow-derivedSPcells.40,41ExclusiveexpressionofABCG2onSPcellssuggeststhatABCG2maybeapotentialmarkerforpositiveselectionofpluripotentstemcellsfromvariousadultsources.ABCG2hasbeenimplicatedinplayingafunctionalroleindevelopmentalstemcellbiology(seereference42forareview).

Sca-1:Sca-1(stemcellantigen1,Ly-6A/E),an18kDaphosphatidylinositol-anchoredprotein,isamemberoftheLy-6antigenfamily.43Sca-1isthemostrecognizedHSCmarkerinmicewithbothLy-6haplotypesasitisexpressedonmultipotentHSCs.44,45Ananti-Sca-1Abisfrequentlyusedincombinationwithnegativeselectionforexpressionofanumberofcellsurfacemarkerscharacteristicofdifferentiatedcellsofhematolymphoidlineages(Lin-)toidentifyandisolatemurineHSCs.Sca-1+HSCscanbefoundintheadultbonemarrow,fetalliverandmobilizedperipheralbloodandspleenwithintheadultanimal.44-49Sca-1hasalsobeendiscoveredinseveralnon-hematopoietictissues,43however,andcanbeusedtoenrichprogenitorcellpopulationsotherthanHSCs.50Sca-1maybeinvolvedinregulatingbothBandTcellactivation.51-54

Mesenchymal/StromalStemCellMarkers

STRO-1:ThemurineIgMmonoclonalAbSTRO-1,producedfromanimmunizationwithapopulationofhumanCD34+bonemarrowcells,canidentifyacellsurfaceantigenexpressedbystromalelementsinhumanbonemarrow.55Frombonemarrowcells,thefrequencyoffibroblastcolony-formingcells(CFU-F)isenrichedapproximately100-foldintheSTRO-1+/GlycophorinA-populationthanintheSTRO-1+/GlycophorinA+population.55ASTRO-1+enrichedsubsetofmarrowcellsiscapableofdifferentiatingintomultiplemesenchymallineagesincludinghematopoiesis-supportivestromalcellswithavascularsmoothmuscle-likephenotype,ADIpocytes,osteoblastsandchondrocytes.56-59STRO-1isavaluableAbfortheidentification,isolationandfunctionalcharacterizationofhumanbonemarrowstromalcellprecursors,whicharequitedistinctfromthoseofprimitiveHSCs.

另外，还有这些marker:CD71,CD105,SH2.SH3,SH4

NeuralStemCellMarkers

Nestin:NestinisaclassVIintermediatefilamentprotein.60,61Althoughitisexpressedpredominantlyinstemcellsofthecentralnervoussystem(CNS),62itsexpressionisabsentfromnearlyallmatureCNScells.63NestinhasbeenthemostextensivelyusedmarkertoidentifyCNSstemcellswithinvariousareasofthedevelopingnervoussystemandinculturedcellsinvitro.34,64-68TheroleofnestininCNSstemcellbiology,however,remainsundefined.Althoughnestindoesnotformintermediatefilamentsbyitselfinvitroa-internexintoformhomo-andheterodimer,coiled-coilcomplexesthatmaythenformintermediatefilaments.69Itstransientexpressionhasbeensuggestedtobeamajorstepintheneuraldifferentiationpathway.61Nestinexpressionhasalsobeendiscoveredinnon-neuralstemcellpopulations,suchaspancreaticisletprogenitors70-72aswellashematopoieticprogenitors.73

PSA-NCAM(Polysialicacid-neuralcelladhesionmolecule):Theregulatedexpressionofneuralcelladhesionmolecule(NCAM)isoformsinthebrainiscriticalformanyneuraldevelopmentalprocesses.TheembryonicformofNCAM,PSA-NCAM,ishighlypolysialylatedandismainlyexpressedinthedevelopingnervoussystem.74PSA-NCAMmayberelatedtosynapticrearrangementandplasticity.75Intheadult,PSA-NCAMexpressionisrestrictedtoregionsthatretainplasticity.76Aneuronal-restrictedprecursoridentifiedbyitshighexpressionofPSA-NCAMcanundergoself-renewalanddifferentiateintomultipleneuronalphenotypes.77PSA-NCAM+neonatalbrainprecursorsarerestrictedtoaglialfateandthyroidhormonecanmodulatethemintoanoligodendrocytefate.78-80PolysialicacidmodificationsignificantlydecreasesNCAMadhesivenessandtherefore,itwasoriginallysuggestedPSA-NCAMworksasapurelyanti-adhesivefactorthatmodulatescell-cellinteractionsinpromotingbrainplasticity.IncreasingevidenceindicatesthatPSA-NCAMmayinteractwithsecretedsignalingmoleculestoperformaninstructiveroleindevelopment.81,82

Figure3.ThestructureofNGFwithamodelofthep75NeurotrophinReceptor.Theextracellulardomainofthereceptoristakenfromthetumornecrosisfactorreceptorstructureandtheintracellularportioncontainsadeathdomain.

p75NeurotrophinR(NTR):p75NTR,alsonamedlowaffinitynervegrowthfactor(NGF)receptor,isatypeItransmembraneproteinthatbelongstothetumornecrosisfactorreceptorsuperfamily(Figure3).83ItbindstoNGF,BDNF,NT-3andNT-4equally(withlowaffinity).p75NTR,whenactivatedinthepresenceofTrk,enhancesresponsestoneurotrophin(seereference84forareview).TrkCreceptorsworkingtogetherwithp75NTRhavebeensuggestedtoservecriticalfunctionsduringthedevelopmentofthenervoussystem.85Neuralcreststemcells(NCSCs)havebeenisolatedbasedontheirsurfaceexpressionofp75NTR.86,87Freshlyisolatedp75NTR+NCSCsfromperipheralnervetissuescanself-renewandgenerateneuronsandgliabothinvitroandinvivo.Inaddition,neuroepithelial-derivedp75NTR+cellsarealsoabletodifferentiateintoneurons,smoothmuscleandSchwanncellsinculture.88Recently,p75NTRhasbeenusedasamarkertoidentifymesenchymalprecursorsaswellashepaticstellatecells.89,90

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