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1、62MolecularPlantMicrobeInteractionsMPMIVol.13No.12000pp.62–71.Publicationno.M1999110101R.?2000TheAmericanPhytopathologicalSocietyAnalysisofMedicagotruncatulaNoduleExpressedSequenceTagsJnosGyrgyey12DanileVaubert1JosI.Jimn

2、ezZurdo13Celineon1LilianeTroussard1dmKondosi1vaKondosi11InstitutdesSciencesVgtalesCNRS91198GifsurYvetteFrance2InstituteofPlantBiologyBiologicalResearchCenterHungarianAcademyofSciencesH6726SzegedHungary3DepartamentodeMicr

3、obiologadelSueloySistemasSimbiticosEstacinExperimentaldelZadnCSIC18008GranadaSpainAccepted1October1999.Systematicsequencingofexpressedsequencetags(ESTs)cangiveaglobalpictureoftheassemblyofgenesinvolvedinthedevelopmentfun

4、ctionofgans.Indeterminatenodulesrepresentingdifferentstagesofthedevelopmentalprogramareespeciallysuitedtothestudyofganogenesis.WiththevectλHybriZAPacDNAlibrarywasconstructedfromemergingnodulesofMedicagotruncatulainducedb

5、ySinhizobiummeliloti.The5′endsof389cDNAclonesweresequencedthentheseESTswereanalyzedbothbysequencehomologysearchbystudyingtheirexpressioninrootsnodules.TwohundredfiftysixESTsexhibitedsignificantsimilaritiestoacterizeddata

6、baseentries40ofthemrepresented26nodulingeneswhile133hadnosimilaritytosequenceswithknownfunction.Only60outofthe389cDNAclonescrespondedtopreviouslysubmittedM.truncatulaESTsequences.F117cDNAsreverseNthern(RNA)hybridizationw

7、ithrootnoduleRNAprobesrevealedenhancedexpressioninthenodule48clonesarelikelytocodefnovelnodulins33cDNAsareclonesofalreadyknownnodulingenes36clonesexhibitsimilaritytootheracterizedgenes.ThussystematicanalysisoftheESTseque

8、ncestheirexpressionpatternsisapowerfulwaytoidentifynodulespecificnodulationrelatedgenes.Interactionbetweenrhizobiatheirlegumehostplantsresultsinthedevelopmentofnitrogenfixingrootnodules.Thisspecializedplantganhostsitssym

9、bioticbacteriumpartnerswhichconvertmolecularnitrogentoammoniumaccessibleftheplant.Noduleganogenesisiscontrolledbymolecularsignalingbetweentheplantthebacterium.Inthebacteriumhostplantflavonoidsinducethesynthesisexcretiono

10、fthelipochitooligosacideNodsignalsactingashostspecificmphogenscapableofreprogrammingdifferentiatedrootcticalcellsfnoduledevelopment.DistinctrootcelllayersresponddifferentlytoNodfacts.InMedicagospp.roothairsoutercticalcel

11、lsareinvolvedintheinfectionprocessfminginfectionthreadsthattransptthebacteriatowardtheinnerctex.Theselattercellsareinducedfdivisionleadingtothefmationofthenodulemeristem.Thencellsarrestedindivisiondifferentiateintovariou

12、scelltypesresultinginthedevelopmentoftheacteristicperipheralcentralnoduletissues.Medicagospp.nodulesareoftheindeterminatetypehavingapersistentmeristemwhereduetothecontinuousconversionofmeristematiccellsintodistinctnodule

13、celltypesallstagesofnoduledifferentiationarerepresented.Thecentralregionoftheindeterminatenodulesisbuiltupbythreemajzones:theapicalnodulemeristemwherecellsproliferate(zoneI)theinvasionzonewherecellsstoptodivideundergodif

14、ferentiationbecameinfectedwithrhizobia(zoneII)thenitrogenfixationzonewherebacteriaareconvertedintonitrogenfixingbacteroids(zoneIII)(SchultzeKondosi1998Long1996).Understingofthemolecularbasisofnoduledevelopmentisofimptanc

15、etohighlightthemechanismsofsymbioticnitrogenfixation.Meoveritmaygiveabetterinsightintogeneralfeaturesofplantganogenesis.Untilnowonlyafewgenesacterizedbyenhancedexpressioninnoduleshavebeenidentifiedtheirfunctionsinnoduled

16、evelopmentarestilllargelyunknown.Sofardifferentialsubtractivehybridizationsdisplays(Gamasetal.1996Szczyglowskietal.1997)aswellas“coldplaquescreening”(Frugieretal.1998)havebeenusedtoisolatesymbiosisrelatedgenes.Themajityo

17、ftheseapproachesresultedintheisolationofcDNAclonesofhighlyexpressedgenes.Herewepresentsequenceexpressionanalysisof389expressedsequencetags(ESTs)fromanodulecDNAlibraryofM.truncatulawhichhasbecomeamodelplantfstudyingRhizob

18、iumlegumesymbiosis(Barkeretal.1990Blondonetal.1994Cooketal.1997Hoffmannetal.1997Trinhetal.1998).SystematicsequencingofM.truncatulaESTshasbeenreptedsofaronlyfaroothairenrichedcDNAlibrary(Covitzetal.1998).Ourresultsshowtha

19、tsequencingofESTsfromanodulecDNAlibrarygivesaglobalviewofgeneactivitiesinthenodulecomparisonoftheexpressionpatternsofcDNAsintherootnoduleprovidesapowerfulwaytoidentifynewnodulinsnoduleenhancedgenes.Crespondingauth:vaKond

20、osiEmail:cleotideaminoacidsequencedatacanbefoundattheEMBLdatabaseunderaccessionnumbersAJ388667toAJ389056.64MolecularPlantMicrobeInteractionsTable1.ClonesshowingsimilaritytoalreadyacterizeddatabaseentriesPutativegeneprodu

21、ctsbycategyCloneno.RelativetransriptlevelaRelativeinductionbCelldivisioncycleAnkyrinlikeprotein174Low1.1cdc4856Veryhigh22HemCDim1phomolog153Low0.8CellwallstructuremetabolismBetaxylosidase136NDNAEndo14betaglucanase117NDNA

22、Expansin280High1.9Extensinlikeprotein425Medium1.2HydroxyprolinerichglycoproteinHRGP122Medium1.0Pectinacetylesterase106Veryhigh110Pectinacetylesterase(putative)252Veryhigh21Polygalacturonaseprecurs(pectinase)204Low1.0Prol

23、inerichproteinPRP4401Medium0.3Prolinerichprotein(PRP1)403Veryhigh0.7Prolinerichprotein96Medium0.2CytoskeletonActinlikeprotein2397Low3.3Adhesinprotein77Low1.2Annexin152Low1.2Putativemyosinheavychain151Low1.0Tubulinbeta1ch

24、ain367Veryhigh1.8DefensecellrescueAcidicchitinase103Medium7.1Acidicchitinaseprecurs159Medium14AcidicendochitinaseclassIII216Low1.6bateperoxidase156Veryhigh7.1Cadmiuminducedprotein139Medium5.6Cationicperoxidase273Medium0.

25、6Cyanase71Low0.8Ferritin358Medium7.4Isoflavonoidreductase44Veryhigh2.7Metallothionein295High1.6PeroxidaseATP20a246Low0.8Phenylalanineammonialyase371Low0.7RAD6188NMNASuperoxidedismutase213Low1.6Geneexpressionchromatingani

26、sationALYtranionalactivat141Low7.5GlycinerichRNAbindingprotein51High0.6HistoneH1327NMNAHistoneH4391Low1.2PcMYB1protein329Low2.1PcMYB1protein394Medium7.0Poly(A)bindingprotein165Low1.0Poly(ADPribose)polymerase268Low1.9Poly

27、adenylatebindingprotein2186Low1.6Putativepoly(A)bindingprotein43Veryhigh6.8RINGzincfingerprotein76Low1.3RNApolymeraseIcommonsubunit296High9.0RSZp2221splicingfact349Medium0.8Scarecrowlikeprotein307Low0.9SnRNPceprotein270N

28、DNASPF1likeDNAbindingprotein161Low1.0WRKY3DNAbindingprotein332Veryhigh97Zincfingerproteinlike426Low2.2MembranetransptintracellulartraffickingATPase283Low1.4Chloplastouterenvelopeprotein178Veryhigh1.1(continuedonnextpage)

29、aRelativetranlevelswereclassifiedonthebasisofmedianvalueofhybridizationsignalsobtainedfallclones.NM=notmeasuredduetomaskingeffectofneighbingsignals.ND=notdetected.bNodulesperroot.Calculatedasthenmalizedratiooftranlevelso

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