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Lecture2ThePavementCell:AmodelforplanarcellpolarityandPINpolarityHowdocells“talk”toeachanothertocoordinatetheirpolarity,growth,andmorphogenesis?CoordinatedcellgrowthinstemsPavementcellswithinterdigitatedlobesandindentationsCoordinatedPIN1localizationtothebasalend*ArabidopsisHowislocalcoordinationachieve?Whatistheglobalsignal?Howistheglobalsignalingtolinkedtothelocalcoordination?InterdigitationofLeafPavementCells(PCs)
Aprocessofcellintercalationinvolvingcell-cellcoordinationofmorphogenesisOutgrowthinlobe-formingregionsmustbecoordinatedwithrestrictionofoutgrowthincomplementaryindentingregionsIndentationLobeConvergentextensioninanimalembryosPCintercalationinleavesViewfromtheleafsurfaceIntercalarypavementcells:amodelsystemforthestudyofcytoskeletalsignalingFilamentousactin(F-actin)
Microtubules(MTs)Developmental,Hormonal,Intercellular&EnvironmentalSignalsOutputInputCellpolarity,Polargrowth,CellshapechangesSpatiotemporalcontrolofcelldivisionSubcellularmediator:Thecytoskeleton???Howarealternatinglobesandindentswithinacellcoordinatelyformed?Howarelobingandindentingcoordinatedlocallybetweenneighboringcells?Howistheinterdigitationprocesscoordinatedgloballyintheentireleaf?tipbasetipbaseAuxin:Asmallmiraclemolecule
Stimulatescellelongation/expansionStimulatescelldivisionRegulatecellmorphogenesisandPINpolarizationControlembryogenesisStimulatesdifferentiationofphloemandxylemStimulatesrootinitiationandlateralrootdevelopmentRegulatestropismPromotesapicaldominanceDelaysleafsenescenceModulateleafandfruitabscissionInducefruitsettingandgrowthinsomeplantsInvolvedinassimilatemovementtowardauxinpossiblybyaneffectonphloemtransportDelaysfruitripeningPromotesfloweringinBromeliadsStimulatesgrowthofflowerpartsPromotesfemalenessindioeciousflowersTheTIR1/AFBauxinreceptorsmediatenucleargeneexpressionPolarauxintransportiscriticalforauxinfunctionsandismediatedbypolardistributionofPINproteinsPIN1SomemajorquestionsinauxinbiologyArethereTIR1/AFB-independentauxinsignalingpathways;andifso,whatdotheyregulate?HowisPINproteinpolarizationachieved?DotheTIR1/AFB-dependentand–independentpathwaysworkcoordinately?Howisthemulti-polaritygeneratedtoformalternatinglobesandindentswithinacell?Howdoadjacentcellscoordinatewitheachothertoforminterdigitatedloesandindents?HowistheinterdigitationcoordinatedintheentireleafepidermistipbaseLobeIndentationThepavementcellasapowerfulmodelsystemCoordinationofcellpolarity,expansionandmorphogenesisCytoskeletalorganizationPINtraffickingandpolarizationAuxinactionandsignalingCdc42Rho2ScRhoAHsRhoCHsRhoBHsRho1ScCdc42pHsCdc42bHsCdc42ScRac1HsRac2HsRop4Rop3Rop5Rop2Rop7Rop6Rop1Rop8Rop9Rop10ROPRacRhoRop11Fungi&animalsArabidopsisZheng&Yang.2000.PlantMolBiol.44:1-9RhoGTPasesareconservedmolecularswitchesinpolaritysignalingROP:Rho-likeGTPasefromplantspollenROP2/4promoteslobeformationinpavementcellsrop4-1,ROP2(RNAi)AbnormaloutgrowthinnecksInhibitionoutgrowthinlobesFuetal.2002.PlantCell14:763-776ROP2/ROP4locallyactivateactinassemblyandsuppressmicrotubuleorganizationCRIB153a.a.RIC4CRIB224a.a.RIC1FinecorticalactinMFs
TransverseCorticalMTsROP2/4Fuetal.2005.Cell120:687-800Fuetal.2005.Cell;Fuetal.2009.CurBiolROP6MicrotubuleROP2F-actinCorticalMThoopsintheneckregionFinecorticalF-actininthelobeFuetal.2005.Cell;Fuetal.2009.CurBiolROP2MTRIC4F-ActinLobeIndentRIC1ROP6ROP2RIC4F-ActinPMTheROP2andROP6pathwaysinhibiteachothertogeneratealternatinglobesandindentationsAuxinactivatesROP2andROP6topromoteinterdigitationMultiplelinesofevidence:1)phenotypeAuxinrapidlyactivatesbothROP2andROP6Xuetal.2010.Cell143:99BiochemicalevidenceCol0yuc1/2/4/6abp1-5GFP-RIC1StageIStageIIIRIC1localizationtomicrotubulesisabolishedinbothyuccaandabp1-5mutantsasintherop6-1mutant.StageIStageIIICellbiologicalevidenceBeforetreatment10uMoryzalin
10hrs30hrs
afterwashing0.01%DMSOcontrol20nMNAAAuxincanpromoteMTre-organizationafteroryzalintreatment.YFP-RIC1localizationtocorticalMTsisinducedbyauxininrop2-1rop4-1mutantbutnotinrop6-1mutantrop2-1rop4-1rop6-1Auxin-bindingprotein1(ABP1)isrequiredforauxinactivationofbothROP2andROP6aswellasPCinterdigitationExpressionofABP1antisenseRNAorantibodyalsocausedanabp1-5likephenotypeXuetal.2010.Cell143:99Lobetip-localizedPIN1isrequiredforPCinterdigitationandROP2activation,whichinturnisnecessaryforPIN1polarizationtothePMAuxinPIN1Xuetal.2010.Cell143:99Amodelforauxinactivationofpavementcellinterdigitation
AuxinactivatesactivatesbothROP2andROP6throughABP1ROP2controlsapositivefeedbacklooptoinitiatePIN1polarizationandlocalizedauxineffluxPIN1-exportedauxinactivatestheROP6pathwaytocoordinatelobeformationROP2andROP6pathwaysmutuallyinhibittomaintainrespectivepolarity(Xuetal.2010.Cell143:99)AworkingmodelforauxincoordinationofPCinterdigitationHowdoesROP2signalingregulatePIN1polarization?AuxinAsymmetricendocytosisandrecyclingregulatePINpolarizationAuxinmayinducePINpolarizationbyinhibitingendocytosisEndocytosisofGFP-PIN1occursintheindentationregionbutnottheloberegionDoesROP2,whichisactivatedintheloberegion,inhibitsendocytosis?Nagawaetal.2012.PLOSBiologyPre-conversion(Green+Red)Regionforconversion(Green+Red)Post-conversion(Green+Red)PIN1-Dendra2PIN1-Dendra2405nmPhoto-convertiblePIN1-Dendra2isusedtoassayPIN1endocytosisInactivationofROP2promotesPIN1endocytosisActiveROP2intheloberegioninhibitsPIN1endocytosis,resultingPIN1polarizationtothelobePMregionAmodelfortheregulationofPIN1traffickingbytheROP2-dependentauxinsignalingpathwayNagawaetal.2012.PLOSBiologyTheCdc42-PARpathwaycontrolstheapicalpolarityinepithelialcellsbyinhibitingenodocytosisHarris,TepassJ.CellBiol.2009:183:1129-1143Nagawaetal.2012.PLOSBioTheROP2pathwaypromotesPIN1polarizationbyinhibitingPIN1endocytosisandpromotingitsrecyclingActinDoseROP-basedauxinsignalingprovideacommonmechanismfortheregulationofPINlocalization?LocalizedROP-basedauxinsignalinginhibitsPINendocytosis:Apositivefeedbackloopforself-organizationofPINproteindistributionPavementCellsRootepidermalcells(Linetal.2012.CurBio)F-actin(Nagawaetal.2012.PLOSBio)AmodelforauxincoordinationofPCinterdigitationHowdoesABP1worktoregulateROPactivation?IsABP1acellsurfaceauxinreceptor?ArgumentsforABP1beingacellsurfacereceptorAlthoughABP1isfoundinbothERandthecellsurf
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