Expressionprofilingofcellsisavaluabletoolinunderstandinggeneregulatorynetworksandinidentifyinggenesnecessaryfornormaldevelopmentalprocesses.Inthehaematopoieticsystem,cellsurfaceMarkersarewelldefinedallowingtheisolationofcellsfromvariousdevelopmentalstagesusingflowcytometry.Howevergeneexpressionstudiesofsolidtissuesareoftenhamperedbythedifficultyinobtainingahomogenouscellpopulation.Oftenappropriatecellmarkersarenotwelldefinedforgivencellstagesthereforemakinggeneexpressionstudiesdifficult.Inadditionstudiesofrarecellpopulationsmeanthattheamountofmaterialobtainedisinsufficientforanalysis.ToovercometheseproblemswehaveemployedasystemwhereagivencellpopulationisGFP-labeledandthensortedbyflowcytometryinordertoobtainapurehomogenouspopulation.RNAwasthenpreparedfromthismaterialandamplifiedbyalinearamplificationmethodallowinglargescalemicroarraygeneexpressionanalysis.Thisapproachwassuccessfullyappliedtogeneexpressionstudiesofthemidbrain-hindbrainorganizerregionofthemousebutcanbeappliedtomanyothersystemstoisolatecellpopulations.
Usingthemouseasamodelorganism,ourgroupinvestigatedtheroleofthepaireddomaintranscriptionfactorPax2inbraindevelopment(Bouchardetal.,2005).Pax2isrequiredfortheformationoftheisthmicorganizer(IsO)atthemidbrain-hindbrainboundary,whereitinitiatesexpressionoftheIsOsignalFgf8.TogainfurtherinsightintotheroleofPax2inmid-hindbrainpatterning,ourgroupsearchedfornovelPax2-regulatedgenesbyCDNAmicroarrayanalysisofFACS-sortedGFP+mid-hindbraincellsfromwild-typeandPax2-/-E8.5embryoscarryingaPax2GFPBACtransgene(Figure1).Oursortingprotocolyielded5,000to10,000GFP+cellsperembryowhichwasinsufficienttogenerateenoughRNAtohybridizetoamicroarray.ThereforeRNAfromthesortedcellswassubjectedtoalinearamplificationasdescribed(Hoffmannetal.,2003)withsomemodifications.Followingthis,theRNAwasusedinamicroarrayscreenasdescribed(Cheungetal.,1999).OurunbiasedchipapproachidentifiedtheEn2,Brn1,(Pou3f3),Sef,Tapp1andnon-codingNcrmsgenesasgeneticPax2targetsthataretotallydependentonPax2functionfortheirexpressioninthemid-hindbrainregion.Notethatgeneswhichshowedonlya2foldregulationonthemicroarraywereconfirmedasPax2targetgenesbyin-situhybridisation.ThisprovesthattheamplificationprotocolislinearandthatgeneswithlowinductionratiosareindeedPax2-regulatedtargetgenes(Figure2).
Backtotop
PreparationoftotalRNAfromlimitingexvivosortedmaterial
Sorting
Cellsweresortedinto1.5mlmicrocentrifugetubescontaining150µLFACSbuffer.Beforesorting,theinnerwallsofthetubewerecoatedwithbuffertopreventcellsfromstickingtothedrysurface.Aftersorting,asmallpercentageofthesortedcellswereusedforreanalysisbyFACStoverifythepurity.Ifnoreanalysisisnecessary,thecellscanbesorteddirectlyintoTRIreagent(Sigma).
RNApreparation
- Centrifugecellsat13,000rpmfor30seconds;
- Removethesupernatantleavingabout20µlremaininginthetube;
- VortexthetubebrieflytobringthecellsintosUSPensionagain,andwhilevortexing,add500µlTRIreagent;
- Leavethetubeatroomtemperaturefor5minutes,andthenadd5mgofcarriertRNA.Add100µlofCHCI3andcentrifugeat13,000rpmfor10minutes;
- Transfertheupperphaseintoanew1.5mltube,andadd700mlofisopropanol;
- PrecipitatetheRNAbyeitherputtingthetubeat-20°Co/norat-80°Cfor1hourfollowedbycentrifugationat13.000rpmfor30minutes;
- Washthepelletwith75%EtOH(preparedwithDEPC-H2O)anddissolveinDEPC-H2O(usuallyRNAfrom50,000-100,000cellsareresuspendedin9ml).
LinearamplificationoftotalRNA
PreparationofdscDNAfromtotalRNA
Firststrandsynthesis(inRNAse-freePCRtubes)
- DissolveRNAfromatleast5,000cells(ideally50,000-100,000cells)in9mlDEPC-H2O;
- Add1µlPAGE-purifiedoligo-dT-T7-primer(500ng/µl),incubatethemixat70°Cfor10minutes,snap-coolonice;
- Preparethefollowingreversetranscriptionmix:4µl5xFirststrandcDNAbuffer,2µl0.1MDTT,1µl10mMdNTPmix,1µlRNAsin,2µlSuperscriptIIRT;
- Add10µlmixtotheRNA/primersampleandincubateat42°Cfor1hour.
Secondstrandsynthesis(inRNAse-freePCRtubes)
- Preparethefollowingmix:86µlDEPC-H2O,30µl5xsecondstrandbuffer,3µl10mMdNTPmix,8µlDNAPolymeraseI,1µlE.coliDNALigase,2µlRNAseH;
- Add130µlmixtothefirststrandcDNA(totalvolume150µl),tapthetubegentlytomix;
- Spinbrieflyandincubateat16°Cfor2hours;
- Add3µlofT4DNApolymerase;
- Incubateat16°Cfor5minutes,putat70°Cfor10minutestostopthereaction.
Clean-upofdouble-strandedcDNA(inRNAse-freemicrocentrifugetubes)
- Extractthesamplewith150µlPhenol/Chloroform;
- AddtheupperaqueousphasefromtheextractionontoaMicrocon-YM50columnalreadypreloadedwith200µlDEPC-H2O;
- Re-extracttheorganicphasewith150µlwater;
- Loadtheupperaqueousphasefromthesecondextractionontothecolumn;
- Centrifugefor8minutesat9,000rpm,discardflow-through;
- Add500µlDEPC-H2Oontothecolumn;
- Spinat9,000rpmfor8minutes,discardflow-through;
- Placethecolumninvertedintoanewtube,centrifugeat2500rpmfor5minutestorelute;
- Bringthevolumeoftheeluateto8µlwithDEPC-H2O,incaseofabiggervolume,speedvacto8µl.
Invitrotranscription(IVT)1(inRNAse-freePCRtubes)
- AddtheIVTmixconsistingof4µl5xIVTreactionbuffer,6µl25mMrNTPmixand2µlT7enzymemixtothedscDNA(roomtemperature);
- Incubateat37°Cfor4hours.
Clean-up(inRNAse-freemicrocentrifugetubes)
- Add130µlDEPC-H2Otobringthetotalvolumeto150µl;
- PurifyaRNAusingaQiagenRNeasykitaccordingtothemanufacturer’sprotocol;
- ConcentrateaRNAto20µlinaspeedvac.
PreparationofdscDNAfromaRNA
Firststrandsynthesis(inRNAse-freePCRtubes)
- UseonlyhalfoftheaRNA,i.e.10µl,forfurthersteps;
- Add1,5µlofpd(N)9(1µg/µl,dissolvedin10mMTRIS,pH7.5)totheaRNA;
- Incubatethemixat70°Cfor10minutes,snap-coolonice.
Reversetranscription
- Preparethefollowingmix:4µl5xFirststrandcDNAbuffer,2µl0.1MDTT,1µl10mMdNTPmix,1µlRNAsin,1µlSuperscriptIIRT;
- Add9µlmixtotheaRNA/primersampleandincubateat42°Cfor1hour;
- Add3µlofRNAseH(removalofDNA/RNAhybrids);
- Incubatefor30minutesat37°C.
Secondstrandsynthesis(inRNAse-freePCRtubes)
- Add1.5µlPAGE-purifiedoligo-dT-T7-primer(500ng/µl);
- Incubateat95°Cfor5minutes,snap-coolonice,incubateat42°Cfor10minutes;
- Addthefollowingmix:92µlDEPC-H2O,15µl10xEco-Polbuffer,4µl10mMdNTPmix,8µlKlenowEnzymeand8µlT4DNAPolymerase;
- Incubateat16°Cfor2hours.
Clean-upofdouble-strandedcDNA(inRNAse-freemicrocentrifugetubes)
- Extractthesamplewith150µlPhenol/Chloroform;
- AddtheupperaqueousphasefromtheextractionontoaMicrocon-YM50columnalreadypreloadedwith200µlDEPC-H2O;
- Re-extracttheorganicphasewith150µlwater;
- Loadtheupperaqueousphasefromthesecondextractionontothecolumn;
- Centrifugefor8minutesat9,000rpm,discardflow-through;
- Add500µlDEPC-H2Oontothecolumn;
- Spinat9,000rpmfor8minutes,discardflow-through;
- Placethecolumninvertedintoanewtube,centrifugedat2500rpmfor5minutesforelution;
- Bringthevolumeoftheeluateto8µlbyaddingDEPC-H2O.Iftheeluateismorethan8µlthenspeedvacthesampleto8µl.
Invitrotranscription(IVT)2
- AddtheIVTmixconsistingof4µl5xIVTreactionbuffer,6µl25mMrNTPmixand2µlT7enzymemixtothedscDNA(roomtemperature);
- Incubateat37°Cfor4hours.
Clean-up(inRNAse-freemicrocentrifugetubes)
- Add130µlDEPC-H2Otobringthetotalvolumeto150µl;
- PurifyaRNAusingaQiagenRNeasykitaccordingtothemanufacturer’sprotocol(seecomment1);
- MeasureaRNAconcentrationonanAgilentBioanalyzer2001accordingtothemanufacturer"sprotocol.
TheaRNAnowcanbeusedformicroarrayhybridizations.Followstandardmicroarrayprotocolsforlabelingandhybridizations.After2roundsofIVT,theamountofaRNAtypicallyvariesbetween60-130µgwhenstartingwith50,000sortedcells.Thisisenoughforatleast20microarrayhybridisations.