高等植物病原相关蛋白(英文)

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【作者】 刘陟  杜林方  万波 

【关键词】病原相关蛋白 病原抗性 诱导系统性抗性 

【出版日期】2005-04-30

【摘要】在过去的三十年中,人们对诱导系统性抗性———这一普遍存在于高等植物抗病过程中的现象———进行了深入研究。被真菌、细菌或病毒侵染后,植物表现出广泛的、长时间的系统性抗性。在这一过程中,植物细胞壁组成成分发生改变,表达各种病原相关蛋白(PR蛋白) ,并合成多种植物抗毒素。本文就主要的PR蛋白家族的结构和功能特性,PR蛋白的发现和分类,及PR蛋白的应用作一综述。

【刊名】天然产物研究与开发

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IntroductionIthasbeenwellestablishedthatinplantsthehypersensi tivereaction (HR)isnotonlyrestrictedtotheinfectedtis sues ,butalsomayoccurindistaluninfectedleaves .Thisprocesshasbeentermedsystemicacquiredresistance(SAR) [1] .TheonsetofSARischaracterizedbyinductionofexpressionofadistinctsetofdefensegenes ,theSARgenes.SARgenesprimarilyencodeacidicpathogenesis related (PR)proteinsandareusedasmolecularmarkersforthepathogenesis induceddefensemechanism .PRproteinsareplantspecies specificproteinsproducedinresponsetoinfectionwithviruses ,fungiorbacteria ,andstressconditionslikesenescence ,callusculture ,infection ,exposuretocertainchemicals (includingplasmolysis ,polyacrylicacid ,auxin ,cytokinin ,arachidonicacid ,ozone ,andhydrogenperoxide) ,woundingandenvironmentalconditions[2 ] .CooperationofPRproteinsspatiallyandtemporallydisplaysmultipleeffectswithintheplantandpossessesantimicrobialactivity ,butaccumulationofPRsisnotaprerequisitefortheinductionofresistance .Never theless,becauseoftheantipathogenicactionsofatleastsomeamongthePRs ,theyarelikelytoberesponsibleforthehigherlevelofinducedresistanceassociatedwithSARandtissuenecrosis .However,somePRproteinshavefunctionsrelatedtode velopmentofhealthyplantsinanorgan andtissue spe cificmanner[3] .Theyexpressconstitutivelyandareupreg ulatedduringgermination ,floweringandsenescence[4 ] ,inpollensorfruits ,tissuesthataremorelikelytobeattackedorexposedtoatmosphericconditions (e .g .UVirradia tion) .Both β 1,3 glucanasesandchitinaseshavebeenfoundtobeassociatedwithplantdevelopment[5] .DiscoveryofPRProteinsNearlythreedecadesago ,PRproteinswereshowntobepresentintheleavesofSamsunNNandXanthitobacclantsduringthehypersensitivereactionagainstTMV .Itwasthenfoundsimilarpathogenesis relatedproteinsinwashfluidsfromtheintercellularspacesoftheleaves .In1991,Dorefoundtheacidicchitinases ,β 1,3 glucanasesandthaumatin likeproteinsaccumulateinextracellular“pocket like”vesicleswhilethebasicchitinasesaccumu lateinelectrondenseinclusionbodiesinthevacuoles[6 ] .Later ,manyproteinswithsimilarpropertieshavebeeniso latedby 1 Dand 2 DSDS PAGEfromtobaccoandotherplantspecies ,bothdicotsandmonocots .Themajortobac coPRproteinshavebeenclassifiedinto 5groups :PR1 PR5 .StructualClassesofPRProteinsManyPRproteinsarestableatlowpH .Theirlowmolecu larweightsrangefrom 10kuto 2 0ku .Theydisplaycon siderableresistancetoproteases ,whichisrequirementstoactasfoodallergens .Accordingtosequencecharacteris tics ,serologicorimmunologicrelationshipsandtheirenzy maticorbiologicactivity ,PRproteinscanbedividedinto17groups (Table 1) .Itisknownthattheycomprisefourfamiliesofchitinases(PR 3, 4 , 8and 11) ,oneofβ 1,3 glucanases (PR 2 ) ,oneofproteinaseinhibitors (PR 6 ) ,andonespecificper oxidase (PR 9) ,aswellasthePR 1familywithunknownbiochemicalproperties ,thethaumatin likePR 5family ,andthebirchallergenBetv 1 relatedPR 10family .Sevenofthese 17groupscontainproteinswithallergenicproper ties (membersofPR proteingroups 2 ,3,4 ,5 ,8,10and14 ) [7] ,anothersixgroupscontainfoodallergens[8] .Notallfamiliesrepresentinanyplantspecies ,buteachfamilymaycompriseseveralmembers .Table 1 StructureclassesofknownPRfamiliesFamiliesTypememberPropertiesGenesymbolReferencePR 1tobaccoPR 1aantifungal,1 17kuYpr1PR 2tobaccoPR 2classI ,II,andIIIendo β 1,3 glucanases,2 5 3 5ku Ypr2 ,[Gns2 (‘Gib’) ]PR 3tobaccoP ,QclassI ,II,IV ,V ,VI,andVIIendochitinases,about 3 0kuYpr3 ,ChiaPR 4tobacco‘R’ antifungal,win likeproteins,endochitinaseactivity ,similartoproheveinC terminaldomain ,13 19ku Ypr4,ChidPR 5tobaccoS antifungal,thaumatin likeproteins ,osmotins ,zeamatins ,permeatins,similartoalpha amylase/trypsininhibitors Ypr5PR 6tomatoinhibitorIproteaseinhibitors,6 13kuYpr6,Pis (‘Pirl’)PR 7tomatoP6 9 endoproteinasesYpr7PR 8cucumberchitinaseclassIIIchitinases,chitinase/lysozymeYpr8,ChibPR 9tobacco‘lignin formingperoxidase’peroxidases ,peroxidase likeproteinsYpr9,PrxPR 10parsley‘PR 1’ribonucleases ,Betv 1 relatedproteinsYpr10PR 11tobacco‘classV’chitinaseendochitinasesactivityYpr11,ChicPR 12radishRs AFP3plantdefensinsYpr12PR 13ArabidopsisTHI2 .1thioninsYpr13 ,ThiPR 14barleyLTP4nonspecificlipidtransferproteins (ns LTPs)Ypr14 ,LtpPR 15barleyOxOa (germin)oxalateoxidaseYpr15PR 16barleyOxOLP‘oxalateoxidase like’Ypr16PR 17tobaccoPRp2 7unknownYpr17StructureandBiologicalFunctionsofPRProteinsStructureofPRproteinsTobaccoiscloselyrelatedmembersofSolanaceae ,andhasbeenusedasmodelsystemtoinvestigatephysiologicalandmolecularmechanismsofplant’santi pathogenactivity .Herewereviewstructuralpropertiesofthemainfiveclas sicPRproteingroupsintobacco .Theyhavetwosubclass es :anacidicsubclassthatisusuallysecretedtotheextra cellularspace ,andabasicsubclasswhichisfunctionallysimilarbuthasdifferentmolecularweightandaminoacidsequence[9] ,andisusuallytransportedtothevacuolebyasignalsequencelocatedattheC terminalend .PR 1PR 1familyhasbeenextensivelystudied ,andmostworksfocusedontheacidicforms .PR 1sarehomologoustothesuperfamilyofcysteine richproteins ,andallarecomple mentaryandnecessaryfortheplan

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