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ToxboxesfungalsecondarymetabolitesandplantdiseaseAnneEOsbournSainsbur


Seecompanionarticleonpage14174inissue24ofvolumeE-mailannieosbournbbsrcacukpnas261573598PNASDecember42001vol98no25genesregulatedbyTOXEmayencodeatoxineffluxpumpacommonthemecouldbetheregulationofgenesinv

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Document on Subject : "ToxboxesfungalsecondarymetabolitesandplantdiseaseAnneEOsbournSainsbur"— Transcript:

1 Tox-boxes,fungalsecondarymetabolites,and
Tox-boxes,fungalsecondarymetabolites,andplantdiseaseAnneE.Osbourn*SainsburyLaboratory,JohnInnesCentre,NorwichNR47UH,UnitedKingdomanyplantpathogenssecretetoxinsthatdisabletheirhost(1).Someofthesehavegeneralphytotoxicpropertiesandareactivetowardabroadrangeofplantspecies.Othersarehost-selective,affectingonlycertainplantvarietiesorgenotypes.Thesehost-selectivetoxins(HSTs)canactasagentsofvirulenceofpathogenstotoxin-sensitivehosts,sodetermininghostrangeorspecificity(1,2).NearlyallHSTsthathavebeendescribedarelowmolecularweightsecondarymetabolitesproducedbyfungi.Thegenesrequiredforthesynthesisofthesemoleculesareoftentightlyclusteredinthefungalgenomeandarecoordinatelyregu-lated(3).InarecentissueofPNAS,Pedley Seecompanionarticleonpage14174inissue24ofvolume*E-mail:annie.osbourn@bbsrc.ac.uk.pnas.261573598PNASDecember4,2001vol.98no.25 genesregulatedbyTOXEmayencodeatoxineffluxpump,acommonthemecouldbetheregulationofgenesinvolvedinself-protectionmechanisms.AlthoughC.carbonumrace1ishighlyaggressivetosusceptiblemaizelines,mostlinesareresistant.ResistanceisconferredbythedominantgeneIn1991Wal-tonandcoworkersdevelopedamethodforgeneratingradiolabeledHC-toxinwithhighspecificactivitybyfeedingfeeding3H]ala-ninetofungalcultures(17).Whenthissubstratewasincubatedwithshootex-tractsfrommaizeseedlingssegregatingitwasfoundthatonlylinesthatwerehomozygousorheterozygousforwereabletoconvertHC-toxintoasingleproductthatlackedbiologicalac-tivity(8-hydroxy-HC-toxin),suggestingthatdetoxificationwasthebasisofdiseaseresistance(18).TheenzymeresponsibleforthisdetoxificationwasnamedHC-toxinreductase.ThisfindingpavedthewayfortheisolationofthegenebytransposontaggingbyusingascreenforinsertionalinactivationofbasedonHC-toxinreductaseactivity(19).wasthefirstdisease-resistancegenetobeclonedfromplants.However,itisregardedassomewhatunorthodoxbymanyofthoseworkinginthedisease-resistancefield.Thevastmajorityofotherdisease-resistancegenesthathavebeenclonedandcharacterizedoverthelast10yearsareinvolvedinspecificinteractionsofaratherdifferentkind,namelytheclassicalgene-for-geneinteractionsthatinvolverecognitionofpathogensexpress-ingspecificavirulencedeterminants(20).Recognitionleadstotheactivationofplantdefenseresponsesandhencediseaseresistance.Manyavirulencedeterminantsareproteinsorpeptides,althoughinsomecasesitisclearthattheproductsofsomeavirulencegenesareinvolvedinthesyn-thesisoflowmolecularweightsecondarymetabolitesthatcanberecognizedbyplants.Examp

2 lesincludethegeneofPseudomonassyringae,w
lesincludethegeneofPseudomonassyringae,whichisrequiredforthesynthesisoflowmolecularweightmoleculesknownassyringolides(21,22),andthegeneofthericeblastfungusMagnaporthegrisea,whichispredictedtoencodeapolyketidesynthase.Evidenceisemergingtosuggestthatthegenomesofplantpathogenicfungiarerichingenesthatarelikelytobeinvolvedinthesynthesisofsecondarymetabolitessuchasnonribosomalpeptidesynthasesandpolyketidesynthases,whereassapro-phytesappeartobedeficientinsuchgenes(23).Furthergenomeminingislikelytounveilnewclustersofcoregulatedgenesthatarerequiredforthesynthesisofnovelsecondarymetabolites,someofwhichmaybedeterminantsofvirulenceandhostrange.Itseemslikely,then,thatsofarwehaveseenonlythetipoftheiceberg.Secondarymetabolitesmayplayafarmoresignificantroleindeterminingtheoutcomeofplantpathogeninteractionsthanhaspreviouslybeenanticipated.TheSainsburyLaboratoryissupportedbytheGatsbyCharitableFoundation.1.Walton,J.D.(1996)PlantCell2.Idnurm,A.&Howlett,B.J.(2001)Mol.PlantPathol.3.Walton,J.D.(2000)Fung.Genet.Biol.4.Pedley,K.F.&Walton,J.D.(2001)Proc.Natl.Acad.Sci.USA14179.(FirstPublishedNovember6,2001;10.10735.Nelson,R.R.&Ullstrap,A.J.(1961)thology6.Scheffer,R.P.,Nelson,R.R.&Ullstrap,A.J.Phytopathology7.Brosch,G.,Ransom,R.,Lechner,T.,Walton,J.D.&Loidl,P.(1995)PlantCell8.Panaccione,D.G.,Scott-Craig,J.S.,Pocard,J.-A.&Walton,J.D.(1992)Proc.Natl.Acad.Sci.USA9.Pitkin,J.W.,Panaccione,D.G.&Walton,J.D.Microbiology10.Ahn,J.-H.&Walton,J.D.(1997)Mol.Plant-MicrobeInteract.11.Cheng,Y.-Q.,Ahn,J.-H.&Walton,J.D.(1999)Microbiology12.Cheng,Y.-Q.&Walton,J.D.(2000)J.Biol.Chem.13.Ahn,J.-H.&Walton,J.D.(1996)PlantCell14.Ahn,J.-H.,Cheng,Y.-Q.&Walton,J.D.,Genet.Biol.,inpress.15.Ahn,J.-H.&Walton,J.D.(1998)Mol.Gen.Genet.16.Bussink,H.-J.,Clark,A.&Oliver,R.(2001)J.PlantPathol.17.Meeley,R.B.&Walton,J.D.(1991)PlantPhysiol.18.Meeley,R.B.,Johal,G.S.,Briggs,S.P.&Walton,J.D.(1992)PlantCell19.Johal,G.S.&Briggs,S.P.(1992)20.Hulbert,S.H.,Webb,C.A.,Shavannor,S.M.&Sun,Q.(2001)Ann.Rev.Phytopathol.21.Kobayashi,D.Y.,Tamaki,S.J.&Keen,N.T.Mol.Plant-MicrobeInteract.22.Smith,M.J.,Mazzola,E.P.,Sims,J.J.,Midland,S.L.,Keen,N.T.,Burton,V.&Stayton,M.M.TetrahedronLett.24.Yoder,O.C.&Turgeon,B.G.(2001)Curr.Opin.PlantBiol. ¨hnert,H.U.,Fudal,I.,Dioh,W.,Tharreau,D.,Not-teghem,J.-L.&Lebrun,M.-H.,Abstractsofthe10thInter-nationalCongressonMolecularPlant-MicrobeInterac-tions,July10–14,2001,Madison,WI,abstr.362. Fig.1.DiseasesymptomscausedbyaTox2)andaTox2)isolateofC.carbonum