杂草对AHAS抑制剂的抗药性分子机理研究进展

2009,11(4):399-406ChineseJournalofPesticideScience专论与综述#AHAS隋标峰,张朝贤*,崔海兰,张猛,黄红娟,魏守辉(,,100193):除草剂在田间的重复及不合理使用,导致了杂草抗药性的发生和发展其中AHAS抑制剂由于靶标单一,抗性发展十分迅速截至2009年,已有103种杂草对AHAS抑制剂产生了抗药性,占19类化学除草剂总抗药性杂草生物型的近1/3从AHAS基因突变位点及种类与杂草抗药性水平的关系AHAS基因突变与AHAS酶活性的关系AHAS基因拷贝数与杂草抗药性的关系以及AHAS酶与除草剂结合前后的三维结构等方面,综述了杂草对AHAS抑制剂产生抗药性的机理,旨在为AHAS抑制剂抗性研究提供参考并对自然种群目标基因的等位基因检测技术(ECOTILLING)和衍生型酶切扩增多态性序列(dCAPS)两种通过检测等位基因多态性的手段快速诊断抗药性杂草的新技术进行了介绍,讨论了延缓杂草抗药性发生和发展的策略:AHAS抑制剂;杂草;抗药性;机制;快速检测和诊断DOI:10.3969/.jissn.1008-7303.2009.04.01:S481.4;S482.4:A:1008-7303(2009)04-0399-08AdvancesinMolecularMechanismofAHAS-inhibitingHerbicidesResistanceSUIBiao-feng,ZHANGChao-xian*,CUIHa-ilan,ZHANGMeng,HUANGHong-juan,WEIShou-hui(InstituteofPlantProtection,ChineseAcademyofAgriculturalSciences,KeyLaboratoryofWeedandRodentBiologyandManagement,CAAS,Beijing100193,China)Abstract:Repeatedandunreasonablefieldapplicationofherbicideshaveresultedintheevolutionofresistantweedspecies.AHASisthetargetsiteofAHAS-inhibitingherbicides.By2009,103weedbiotypeshaveshowedresistancetoAHAS-inhibitingherbicides,accountingfornearly1/3oftotalweedbiotypesresistantto19chemicalgroups.TherelationshipbetweenAHASgenemutationsites/typesandthelevelofherbicideresistance,AHASgenemutationsandAHASactivity,AHASgenenumberandherbicideresistance,andthe3DstructureofAHASbeforeandafterbindingwithAHAS-inhibitingherbicideswerereviewed.ECOTILLINGanddCAPS,twonewtechniquesforrapidlydiagnosingherbicideresistancebydetectingSNPwerebroughtforward,andpotentialmanagementstrategyforpreventinganddelayingtheherbicideresistanceevolutioninweedswasalsodiscussed.Keywords:AHAS-inhibitingherbicides;weed;resistance;mechanism;rapiddiagnosisanddetecting:2009-02-27;:2009-06-15.:(1981-),,,,,E-mai:[e-mail];*(Authorforcorrespondence):(1955-),,,,,.:010-62815937;E-mai:[e-mail]:/0(2006BAD08A09).Vo.l11(acetohydroxyacidsynthase,AHAS,EC2.2.1.6)(acetolac-tatesynthase,ALS,EC4.1.3.18)))),[1],,AHASALS,AHAS5:(sulfonylureas,SU)[2](imidazolinones,IMI)[3](pyrimidinylthiobenzoates,PTB)[4](triazolopyrimidines,TP)[5](sulfonylaminocarbonyltriazolinones,SCT)[6],,55WeedScience65WeedResearch65WeedTechnology65WeedBiologyandManagement6,200,AHAS52,,AHAS:AHAS,AHAS,AHAS[7-10],AHASSisymbriumorientale[11-12];[13-14],[15],AHAS,,AHASAHASAHASAHAS,AHASAHAS1AHASAHAS,AHASLactucaserriola,51987[16-17]AHAS2080,,(1)1987200013,70,200910,34103AHAS,,191/339,20,19,9[18]3,MonochoriakorsakowiiSagittariamontevidensis[19-20]Descurainiasophia[7,21]2008AHASCapsellabursa-pastorisAvenasterilis[18]1[18]Fig.1Resistantweedbiotypestovariousherbicides[18]2AHAS,,17AHAS[22](2)AHASAHAS6400No.4:AHAS2AHAS[9,23]Fig.2CodingsequenceoftheAHASgenebasedonthatofArabidopsis[9,23]:17(ArabidopsisthalianaAHAS);Note:Uppernumbersindicatetheaminoacidpositionof17codonsknowntoinfluenceherbicideresistanceamongeukaryoticandprokaryoticorganism(codingsequenceoftheAHASgenebasedonthatofArabidopsis);blackshadingindicatestheregionforphylogeneticanalysis.:Ala122,Pro197,Ala205,Asp376,Trp574Ser653[8-9,24-26],122,197,205,376,574,653AHAS,AHAS[25],Trp574LeuAHAS,Ala122ThrIMI,Ala205ValTrp574Leu[27],Sales[28]OryzasativaGly654GluVal669Me,tAHASAHASAla205ValSolanumptychanthumR,Ashigh[1]R1S2AHAS,,R1AHASS247%;Eberlein[29-30]Lactucasativa(Pro197His)Yu[25]Raphanusraphanistrum,AHAS3~5,NorthernAHASmRNA,AHAS3AHAS,AHASAHAS,AHAS,AHAS,1,1AHAS[31]AHAS,NicotianatabacumAHAS[32],AHAS75%,85%,NAHAS,AHAS1,2[32]AHAS,Rutledge[33]Southern,Brassicanapus5AHAS(AHAS1-5),AHAS,DNA,AHAS1AHAS3,AHAS;AHAS2AHAS1AHAS3DNA,N,AHAS2;AHAS4AHAS5,AHASOhsako[34],Monochoriavaginalis6AHAS,AHAS1AHAS2AHAS3AHAS4,AHAS1AHAS3AHASAHAS,Yu[25]NorthernAHAS,401Vo.l11AHASmRNA,AHASAHASmRNA25,AHAS2,AHASAHASmRNA[35]AHAS[11-12],AHASSouthernPCR,AHASAHASAHAS,4AHASAHASE.coliAHAS,AHAS[36],AHAS[22,37],Duggleby[38-40]X-AHAS3DAHAS(AtAHAS)3D4,3:A(86~280)B(281~451)C(463~639)C(646~668)[41](3),AHASC[28]AHAS,AHAS(ScAHAS)CC[38,41],,AHASC/0C[39,42](4,5)AtAHAS3DAHAS6Trp574SUIMI,Trp574AHAS,Trp574SUIMIPro1978[9,22]Pro1978SU,Pro197Leu,IMI[43]Pro197AHASA,SU,IMI(imazaquin,IQ),IQAHAS,Pro197IMI,Pro1978SUAHAS[41]Ser653AsnAla122ThrIM,SU[44-45]Ala122,Ala122,AHAS;Ala122(chlorimuron-ethy,lCE),SU,Ser653AHASSU,Ser653IMIAHAS,[41]AHAS3D,AHAS[23,46]5AHAS,AHAS,(ECOTILLING),(TILLING):402No.4:AHAS3AHAS[41]Fig.3TheoverallfoldofAtAHAS[41]:(A)()(B)A(86~280),B(281~451),C(463~639)C(646~668)(ThDP)Mg2+(FAD)IQ-,Note:(A)Thetetramericstructurewitheachmonomercoloredseparately.(B)Asinglesubuni.tTheindividualdomainsA(86-280),B(281-451),andC(463-639)arecoloredingold,red,andblue,respectively.TheC-terminaltail(646-668)iscoloredgreen.ThDP,Mg2+,FAD,andIQareshownasbal-land-stickmodelsandarecoloredred,darkblue,cyan,andyellow,respectively.DNADNA1B1,96,,,4AHAS-CEB(a)AHASCE(b)[39]Fig.4StructureofmonomerBintheScAHAS-CEcomplex(a)andmajordifferencesinthestructureofyeastAHASinthepresenceandabsenceofCE(b)[39]:(a)()(C)ThDPFAD-CE-(b)CE,FADThDP;CE,CCENote:(a)Theaminoacidresiduesthatarenotobservedinthestructureofthefreeenzymearecoloredgreen(themobileloop)andbrown(theC-terminaltail).ThDPandFADaredisplayedas