根癌农杆菌介导的遗传转化及其在稻瘟病菌中的应用
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摘要
根癌农杆菌介导的遗传转化(ATMT)技术是近20年发展起来的真菌研究中最具变革性的技术之一,它作为一种转化工具已被广泛应用于真菌基因功能研究中,使人们对真菌基因组学的理解有了很大的改变。利用ATMT获得随机的T-DNA插入突变体库,结合突变体菌株表型筛选,使许多过程的遗传基础得到阐明。ATMT法具有操作简单,转化受体广泛,转化效率高以及转化子遗传稳定性高等优势。稻瘟病菌是植物十大重要真菌之一,是引发水稻稻瘟病的病原真菌。目前,培育水稻抗病品种是控制稻瘟病最经济和环保的方法。通过克隆稻瘟病菌相关致病基因并研究其功能,在解释致病机理和生理特性,以及培育抗病品种方面具有重要的理论和实践意义。以根癌农杆菌为介导的遗传转化为稻瘟病菌的插入突变、基因敲除和标记克隆等方面研究提供了一个重要工具。针对根癌农杆菌介导的遗传转化原理、特点、步骤、影响因素以及该技术在稻瘟病菌功能基因组学研究中的应用进行综述,以期能够为稻瘟病菌等真菌的遗传转化方面研究提供相关的参考。
Agrobacterium tumefaciens-mediated transformation(ATMT) is one of the most revolutionary techniques in fungal research in recent 20 years. As a transformation tool, it has been widely used in the study of fungal gene function, which made people have a better understanding of fungal genomics. Combined with phenotype screening of mutants, random T-DNA obtained by ATMT was inserted into mutant library and the genetic basis of many processes was elucidated. ATMT possesses the advantages of simple operation, wide transformation receptors, high transformation efficiency and high stability of transformants etc.Magnaporthe oryzae, one of the top ten important fungi in plants, is the pathogenic fungus that causes rice blast. At present, the most economical and environmentally-friendly method to control rice blast is to cultivate rice resistant varieties. It is of great theoretical and practical significance to explain the pathogenic mechanism and physiological characteristics and cultivate disease-resistant varieties by cloning the pathogenic genes related to Magnaporthe oryzae and studying their functions. ATMT provided an important tool for the study of insertion mutation, gene knockout and marker cloning of Magnaporthe oryzae. The principle, characteristics, steps, and influencing factors of ATMT and its application in functional genomics study of Magnaporthe oryzae were reviewed in order to provide relevant references for the study of genetic transformation of fungi such as Magnaporthe oryzae etc.
Agrobacterium tumefaciens-mediated transformation(ATMT) is one of the most revolutionary techniques in fungal research in recent 20 years. As a transformation tool, it has been widely used in the study of fungal gene function, which made people have a better understanding of fungal genomics. Combined with phenotype screening of mutants, random T-DNA obtained by ATMT was inserted into mutant library and the genetic basis of many processes was elucidated. ATMT possesses the advantages of simple operation, wide transformation receptors, high transformation efficiency and high stability of transformants etc.Magnaporthe oryzae, one of the top ten important fungi in plants, is the pathogenic fungus that causes rice blast. At present, the most economical and environmentally-friendly method to control rice blast is to cultivate rice resistant varieties. It is of great theoretical and practical significance to explain the pathogenic mechanism and physiological characteristics and cultivate disease-resistant varieties by cloning the pathogenic genes related to Magnaporthe oryzae and studying their functions. ATMT provided an important tool for the study of insertion mutation, gene knockout and marker cloning of Magnaporthe oryzae. The principle, characteristics, steps, and influencing factors of ATMT and its application in functional genomics study of Magnaporthe oryzae were reviewed in order to provide relevant references for the study of genetic transformation of fungi such as Magnaporthe oryzae etc.
引文
[1]陈卫华,甘雨,黄宗洪,向关伦,杨占烈,潘建慧,郭慧.健优388在长江上游稻区的主要农艺性状表现分析[J].广东农业科学,2015,42(1):4-7.doi:10.16768/j.issn.1004-874X.2015.01.024.CHEN W H,GAN Y,HUANG Z H,XIANG G L,YANG Z L,PAN JH,GUO H.Analysis of agronomic traits for Jianyou 388 in upper rice region of Yangtze River[J].Guangdong Agricultural Sciences,2015,42(1):4-7.doi:10.16768/j.issn.1004-874X.2015.01.024.
[2]FISHER M C,HENK D A,BRIGGS C J,BROWNSTEIN J S,MADOFFL C,MCCRAW S L,GURR S J.Emerging fungal threats to animal,plant and ecosystem health[J].Nature,2012,484(7393):186-194.
[3]谭明辉,霍光华,彭玉萌,李芳.木荷和无患子等植物组方杀稻瘟剂的药效及其安全性[J].广东农业科学,2014,41(16):85-89.doi:10.16768/j.issn.1004-874X.2014.16.020.TAN M H,HUO G H,PENG Y M,LI F.Efficacy and safety of blasticidin agent prepared by Schima superba,Sapindus mukorossi and other plants[J].Guangdong Agricultural Sciences,2014,41(16):85-89.doi:10.16768/j.issn.1004-874X.2014.16.020.
[4]ZHONG Z H,NORVIENYEKU J,CHEN M L.Directional selection from host plants is a major force driving host specificity in Magnaporthe species[J].Scientific Reports,2016,6:25591.
[5]BAO J D,CHEN M L,ZHONG Z H.PacBio sequencing reveals transposable elements as a key contributor to genomic plasticity and virulence variation in Magnaporthe oryzae[J].Molecular Plant,2017,10(11):1465-1468.
[6]BUNDOCK P,DULK‐RAS A,BEIJERSBERGEN A,HOOYKAAS PJ.Trans‐kingdom T-DNA transfer from Agrobacterium tumefaciens to Saccharomyces cerevisiae[J].The EMBO Journal,1995,14(13):3206-3214.
[7]GROOT M J A,BUNDOCK P,HOOYKAAS P J J,BEIJERSBERGENA G M.Agrobacterium tumefaciens-mediated transformation of filamentous fungi[J].Nature Biotechnology,1998,16(9):839-842.
[8]陈东亮,李纪元,范正琪,范妙华.根癌农杆菌介导真菌遗传转化的影响因素及应用[J].安徽农业科学,2010,38(7):3317-3320.CHEN D L,LI J Y,FAN Z Q,FAN M H.Influencing factors of genetic transformation in fungi mediated by Agrobacterium tumefaciens and its application[J].Journal of Anhui Agriculture science,2010,38(7):3317-3320.
[9]RHO H S,KANG S,LEE Y H.Agrobacterium tumefaciens-mediated transformation of the plant pathogenic fungus,Magnaporthe grisea[J].Molecules and Cells,2001,12(3):407-411.
[10]LACROIX B,TZFIRA T,VAINSTEIN A,CITOVSKY V.A case of promiscuity:Agrobacterium's endless hunt for new partners[J].Trends in Genetics,2006,22(1):29-37.
[11]王栋鑫,彭棣,张爽.农杆菌介导木本植物遗传转化的研究进展[J].北方园艺,2018(2):181-185.doi:10.11937/bfyy.20173329.WANG D X,PENG D,ZHANG S.Advances in Agrobacteriummediated transformation of woody plants[J].Northern Horticulture,2018(2):181-185.doi:10.11937/bfyy.20173329.
[12]邹智.农杆菌vir基因诱导因子研究进展[J].中国生物工程杂志,2011,31(7):126-132.doi:10.13523/j.cb.20110721.ZOU Z.Advances on factors influencing induction of Agrobacterium tumefaciens virulence genes[J].China Biotechnology,2011,31(7):126-132.doi:10.13523/j.cb.20110721.
[13]姜宁,宋春艳,刘建雨,谭琦,章炉军,尚晓冬.农杆菌介导香菇菌丝遗传转化体系的研究[J].菌物学报,2017,36(11):1514-1523.doi:10.13346/j.mycosystema.170041.JIANG N,SONG C Y,LIU J Y,TAN Q,ZHANG L J,SHANG X D.Agrobacterium-mediated transformation of the Lentinula edodes mycelia[J].Mycosystema,2017,36(11):1514-1523.doi:10.13346/j.mycosystema.170041.
[14]高兴喜,杨谦,宋金柱,郭兆奎.根癌农杆菌介导的木霉菌遗传转化方法[J].高技术通讯,2004,5:31-34.GAO X X,YANG Q,SONG J Z,GUO Z K.Agrobacterium tumefaciensmediated transformation of Trichoderma Harzianum[J].High Technology Letters,2004,5:31-34.
[15]MEYER V,MUELLER D,STROWIG T,STAHL U.Comparison of different transformation methods for Aspergillus giganteus[J].Current Genetic,2003,43(5):371-377.
[16]CAMPOY S,PEREZ F,MARTIN J F,GUTIERREZ S,LIRAS P.Stable transformants of the azaphilone pigment-producing monascus purpureus obtained by protoplast transformation and Agrobacteriummediated DNA transfer[J].Current Genetics,2003,43(6):447-452.
[17]LIU J Y,SONG C Y,LI Q Z,XU Z,ZHANG D,ZHANG M Y,TANQ,SHANG X D.A colonized millet grain method for Agrobacteriummediated transformation of the button mushroom Agaricus bisporus[J].Journal of Microbiological Methods,2018,152:148-153.
[18]王宏宇.稻瘟病菌T-DNA插入突变研究[D].福州:福建农林大学,2003.WANG H Y.T-DNA insertional mutagenesis of Magnaporthe grisea[D].Fugzhou:Fujinag Agriculture and Forestry University,2003.
[19]EPSTEIN L,LUSNAK K S.Transformation-mediated developmental mutants of Glomerella graminicola(Collectorichum graminicola)[J].Fungal Genetic Biology,1998,23(2):189-203.
[20]GUO M,ZHU X L,LI H X,TAN L Y,PAN Y M.Development of a novel strategy for fungal transformation based on a mutant locus conferring carboxin-resistance in Magnaporthe oryzae[J].AMB Express,2016,6(1):57.
[21]ZHANG P,LIU T T,ZHOU P P,LI S T,YU L J.Agrobacterium tumefaciens-mediated transformation of a taxol-producing endophytic fungus,Cladosporium cladosporioides MD2[J].Current Microbiology,2011,62(4):1315-1320.
[22]王曦茁,朴春根,李虹,汪来发,郭民伟,李永,刘晓莉.根癌农杆菌介导的淡紫拟青霉遗传转化体系的建立[J].林业科学,2010,46(10):95-102.WANG X Z,PU C G,LI H,WANG L F,GUO M W,LI Y,LIU X L.Establishment of Agrobacterium tumefaciens-mediated transformation system of Paecilomyces lilacinus[J].Scientia Silvae Sinicae,2010,46(10):95-102.
[23]李敏慧,张荣,姜大刚,习平根,庄楚雄,姜子德.根癌农杆菌介导的香蕉枯萎病菌4号生理小种的转化[J].植物病理学报,2009,39(4):405-412.LI M H,ZHANG R,JIANG D G,XI P G,ZHUANG C X,JIANGZ D.Agrobacterium tumefaciens-mediated transformation of Fusarium oxysporum f.sp.cubense race 4[J].Acta Phytopathologica Phytopathologica Sinica,2009,39(4):405-412.
[24]LECLERQUE A,WAN H,ABSCHüTZ A,CHEN S,MITINA GV,ZIMMERMANN G,SCHAIRER H U.Agrobacterium-mediated insertional mutagenesis(AIM)of the entomopathogenic fungus Beauveria bassiana[J].Current Genetics,2004,45(2):111-119.
[25]MULLINS E D,KANG S.Transformation:a tool for studying fungal pathogens of plants[J].Cellular and Molecular Life Sciences,2001,58(14):2043-2052.
[26]TAKAHARA H,TSUJI G,KUBO Y,YAMAMOTO M,TOYODAK,INAGAKI Y,ICHINOSE Y,SHIRAISHI T.Agrobacterium tumefaciens-mediated transformation as a tool for random mutagenesis of Colletotrichum trifolii[J].Journal of General Plant Pathology,2004,70(2):93-96.
[27]SHARMA K K,KUHAD R C.Genetic transformation of lignin degrading fungi facilitated by Agrobacterium tumefaciens[J].BMCBiotechnology,2010,10(1):67.
[28]ZHENG Z,HUANG C,CAO L,XIE C,HAN L.Agrobacterium tumefaciens-mediated transformation as a tool for insertional mutagenesis in medicinal fungus Cordyceps militaris[J].Fungal Biology,2011,115(3):265-274.
[29]LIU N,CHEN G Q,NING G A,SHI H B,ZHANG L C,LU J P,MAO LJ,FENG X X,LIU X H,SU Z Z,LIN F C.Agrobacterium tumefaciensmediated transformation:An efficient tool for insertional mutagenesis and targeted gene disruption in Harpophora oryzae[J].Microbiological Research,2016,182:40-48.
[30]WANG D,HE D,LI G,GAO S,LV H Y,SHAN Q S,WANG L.An efficient tool for random insertional mutagenesis:Agrobacterium tumefaciens-mediated transformation of the filamentous fungus Aspergillus terreus[J].Journal of Microbiological Methods,2014,98:114-118.
[31]LIU T,LIU L,JIANG X,HOU J M,FU K H,ZHOU F H,CHENJ.Agrobacterium-mediated transformation as a useful tool for the molecular genetic study of the phytopathogen Curvularia lunata[J].European Journal of Plant Pathology,2010,126(3):363-371.
[32]李维,张义正.根癌农杆菌介导的白腐丝状真菌-黄孢原毛平革菌的转化[J].微生物学报,2005,45(5):784-787.LI W,ZHANG Y Z.Agrobacterium tumefaciens-mediated transformation of the white-rot basidiomycete,Phanerochaete chrysosporium[J].ActaMicrobiologica Sinica,2005,45(5):784-787.
[33]PARK S Y,JEONG M H,WANG H Y,KIM J A,YU A H,KIM SB,CHEONG Y H,KANG S C,LEE Y H,HUR J S.Agrobacterium tumefaciens-mediated transformation of the lichen fungus,Umbilicaria muehlenbergii[J].PloS One,2013,8(12):e83896.
[34]张俊华,刘烨,韩雨桐,潘春清,王中业,张淋淋,崔凯旋.农杆菌介导稻瘟病菌绿色荧光蛋白(GFP)遗传转化研究[J].东北农业大学学报,2014,45(11):1-7.doi:10.19720/j.cnki.issn.1005-9369.2014.11.001.ZHANG J H,LIU Y,HAN Y T,PAN C Q,WANG Z Y,ZHANG L LCUI K X.GFP genetic transformation of Magnaporthe grisea mediated by Agrobacterium tumefaciens[J].Journal of Northeast Agricultural University,2014,45(11):1-7.doi:10.19720/j.cnki.issn.1005-9369.2014.11.001.
[35]吴毅歆,范成明,周惠萍,久保康之,何月秋.一种农杆菌介导稻瘟病菌的遗传转化[J].植物保护学报,2008,35(5):421-426.doi:10.13802/j.cnki.zwbhxb.2008.05.008.WU Y X,FAN C M,ZHOU H P,YASUYUKI K B,HE Y Q.A genetic of transformation of Magnaporthe grisea by Agrobacterium tumefaciens[J].Acta Phytophylacica Sinica,2008,35(5):421-426.doi:10.13802/j.cnki.zwbhxb.2008.05.008.
[36]LU Y Y,XIAO S Q,WANG F,SUN J Y,ZHAO L K,YAN L B,XUE C S.Agrobacterium tumefaciens-mediated transformation as an efficient tool for insertional mutagenesis of Cercospora zeae-maydis[J].Journal of Microbiological Methods,2017,133:8-13.
[37]BOER P,BRONKHOF J,DUKI?K,KERKMAN R,TOUW H,BERGM,OFFRINGA R.Efficient gene targeting in Penicillium chrysogenum using novel Agrobacterium-mediated transformation approaches[J].Fungal Genetics and Biology,2013,61:9-14.
[38]LIU L,WEI Y M,ZHOU X W.Agrobacterium tumefaciens-mediated genetic transformation of the Taxol-producing endophytic fungus Ozonium sp EFY21[J].Genet Mol Res,2013,12(3):2913.
[39]ZHANG Y P,QU H X,ZHAO P,TANG Y P,ZHOU J S,LUO S C,YINY L,CHEN G Y.Generation and screening of T-DNA insertion mutants mediated by Agrobacterium tumefaciens in the garden asparagus stem blight pathogen Phomopsis asparagi[J].Current Microbiology,2017,74(11):1270-1277.
[40]WANG C X,GUAN X N,WANG H Y,LI G F,DONG X L,WANG GP,Li B H.Agrobacterium tumefaciens-mediated transformation of Valsa mali:an efficient tool for random insertion mutagenesis[J].The Scientific World Journal,2013,2013.
[41]LI G H,ZHOU Z Z,LIU G H,ZHENG F C,HE C Z.Characterization of T-DNA insertion patterns in the genome of rice blast fungus Magnaporthe oryzae[J].Current Genetics,2007,51(4):233-243.
[42]CHEN X L,YANG J,PENG Y L.Large-scale insertional mutagenesis in Magnaporthe oryzae by Agrobacterium tumefaciens-mediated transformation[J].Fungal Genomics,2011,722:213-224.
[43]JEON J,PARK S Y,CHI M H.Genome-wide functional analysis of pathogenicity genes in the rice blast fungus[J].Nature Genetics,2007,39(4):561-565.
[44]PENG H W,FENG Y J,ZHU X H,LAN X Y,TANG M,WANG J Z,DONG H T,CHEN B S.MoDUO1,a Duo1-like gene,is required for full virulence of the rice blast fungus Magnaporthe oryzae[J].Current Genetics,2011,57(6):409-420.
[45]WANG YU,HE D,CHU Y,ZUO Y S,XU X W,CHEN X L,ZHAO W S,ZHANG Y,YANG J,PENG Y L.MoCps1 is important for conidiation,conidial morphology and virulence in Magnaporthe oryzae[J].Current Genetics,2016,62(4):861-871.
[46]MOGGA V,DELVENTHAL R,WEIDENBACH D,LANGER S,BERTRAM P M,ANDRESEN K,THINES E,KROJ T,SCHAFFRATHU.Magnaporthe oryzae effectors MoHEG13 and MoHEG16 interfere with host infection and MoHEG13 counteracts cell death caused by Magnaporthe-NLPs in tobacco[J].Plant Cell Reports,2016,35(5):1169-1185.
[47]ZHANG S L,WANG L,WU W H,HE L Y,YANG X F,PAN Q H.Function and evolution of Magnaporthe oryzae avirulence gene AvrPib responding to the rice blast resistance gene Pib[J].Scientific Reports,2015,5:11642.
[48]吴小燕,王教瑜,张震,井金学,杜新法,柴荣耀,毛雪琴,邱海萍,姜华,王艳丽,孙国昌.稻瘟病菌致病性增强突变体B11的基因分析[J].中国水稻科学,2009,23(6):611-615.doi:10.3969/j.issn.1001-7216.2009.06.08.WU X Y,WANG J Y,ZHANG Z,JING J X,DU X F,CHAI R Y,MAOX Q,QIU H P,JIANG H,WANG Y L,SUN G C.Analysis on gene locus of Magnaporthe oryzae B11,a pathogenicity-enhanced mutant[J].Journal of China Rice Science,2009,23(6):611-615.doi:10.3969/j.issn.1001-7216.2009.06.08.
[49]孔丹丹.稻瘟病菌致病缺陷突变体KD-560的获得及标记基因MoCYP537A3功能的初步研究[D].杭州:浙江大学,2012.KONG D D.The obtaining of a nonpathogenic mutant Kd-560 of Magnaporthe oryzae and a primarily functional study on the tagged gene MoCYP537A3[D].Hangzhou:Zhejiang University,2012.
[2]FISHER M C,HENK D A,BRIGGS C J,BROWNSTEIN J S,MADOFFL C,MCCRAW S L,GURR S J.Emerging fungal threats to animal,plant and ecosystem health[J].Nature,2012,484(7393):186-194.
[3]谭明辉,霍光华,彭玉萌,李芳.木荷和无患子等植物组方杀稻瘟剂的药效及其安全性[J].广东农业科学,2014,41(16):85-89.doi:10.16768/j.issn.1004-874X.2014.16.020.TAN M H,HUO G H,PENG Y M,LI F.Efficacy and safety of blasticidin agent prepared by Schima superba,Sapindus mukorossi and other plants[J].Guangdong Agricultural Sciences,2014,41(16):85-89.doi:10.16768/j.issn.1004-874X.2014.16.020.
[4]ZHONG Z H,NORVIENYEKU J,CHEN M L.Directional selection from host plants is a major force driving host specificity in Magnaporthe species[J].Scientific Reports,2016,6:25591.
[5]BAO J D,CHEN M L,ZHONG Z H.PacBio sequencing reveals transposable elements as a key contributor to genomic plasticity and virulence variation in Magnaporthe oryzae[J].Molecular Plant,2017,10(11):1465-1468.
[6]BUNDOCK P,DULK‐RAS A,BEIJERSBERGEN A,HOOYKAAS PJ.Trans‐kingdom T-DNA transfer from Agrobacterium tumefaciens to Saccharomyces cerevisiae[J].The EMBO Journal,1995,14(13):3206-3214.
[7]GROOT M J A,BUNDOCK P,HOOYKAAS P J J,BEIJERSBERGENA G M.Agrobacterium tumefaciens-mediated transformation of filamentous fungi[J].Nature Biotechnology,1998,16(9):839-842.
[8]陈东亮,李纪元,范正琪,范妙华.根癌农杆菌介导真菌遗传转化的影响因素及应用[J].安徽农业科学,2010,38(7):3317-3320.CHEN D L,LI J Y,FAN Z Q,FAN M H.Influencing factors of genetic transformation in fungi mediated by Agrobacterium tumefaciens and its application[J].Journal of Anhui Agriculture science,2010,38(7):3317-3320.
[9]RHO H S,KANG S,LEE Y H.Agrobacterium tumefaciens-mediated transformation of the plant pathogenic fungus,Magnaporthe grisea[J].Molecules and Cells,2001,12(3):407-411.
[10]LACROIX B,TZFIRA T,VAINSTEIN A,CITOVSKY V.A case of promiscuity:Agrobacterium's endless hunt for new partners[J].Trends in Genetics,2006,22(1):29-37.
[11]王栋鑫,彭棣,张爽.农杆菌介导木本植物遗传转化的研究进展[J].北方园艺,2018(2):181-185.doi:10.11937/bfyy.20173329.WANG D X,PENG D,ZHANG S.Advances in Agrobacteriummediated transformation of woody plants[J].Northern Horticulture,2018(2):181-185.doi:10.11937/bfyy.20173329.
[12]邹智.农杆菌vir基因诱导因子研究进展[J].中国生物工程杂志,2011,31(7):126-132.doi:10.13523/j.cb.20110721.ZOU Z.Advances on factors influencing induction of Agrobacterium tumefaciens virulence genes[J].China Biotechnology,2011,31(7):126-132.doi:10.13523/j.cb.20110721.
[13]姜宁,宋春艳,刘建雨,谭琦,章炉军,尚晓冬.农杆菌介导香菇菌丝遗传转化体系的研究[J].菌物学报,2017,36(11):1514-1523.doi:10.13346/j.mycosystema.170041.JIANG N,SONG C Y,LIU J Y,TAN Q,ZHANG L J,SHANG X D.Agrobacterium-mediated transformation of the Lentinula edodes mycelia[J].Mycosystema,2017,36(11):1514-1523.doi:10.13346/j.mycosystema.170041.
[14]高兴喜,杨谦,宋金柱,郭兆奎.根癌农杆菌介导的木霉菌遗传转化方法[J].高技术通讯,2004,5:31-34.GAO X X,YANG Q,SONG J Z,GUO Z K.Agrobacterium tumefaciensmediated transformation of Trichoderma Harzianum[J].High Technology Letters,2004,5:31-34.
[15]MEYER V,MUELLER D,STROWIG T,STAHL U.Comparison of different transformation methods for Aspergillus giganteus[J].Current Genetic,2003,43(5):371-377.
[16]CAMPOY S,PEREZ F,MARTIN J F,GUTIERREZ S,LIRAS P.Stable transformants of the azaphilone pigment-producing monascus purpureus obtained by protoplast transformation and Agrobacteriummediated DNA transfer[J].Current Genetics,2003,43(6):447-452.
[17]LIU J Y,SONG C Y,LI Q Z,XU Z,ZHANG D,ZHANG M Y,TANQ,SHANG X D.A colonized millet grain method for Agrobacteriummediated transformation of the button mushroom Agaricus bisporus[J].Journal of Microbiological Methods,2018,152:148-153.
[18]王宏宇.稻瘟病菌T-DNA插入突变研究[D].福州:福建农林大学,2003.WANG H Y.T-DNA insertional mutagenesis of Magnaporthe grisea[D].Fugzhou:Fujinag Agriculture and Forestry University,2003.
[19]EPSTEIN L,LUSNAK K S.Transformation-mediated developmental mutants of Glomerella graminicola(Collectorichum graminicola)[J].Fungal Genetic Biology,1998,23(2):189-203.
[20]GUO M,ZHU X L,LI H X,TAN L Y,PAN Y M.Development of a novel strategy for fungal transformation based on a mutant locus conferring carboxin-resistance in Magnaporthe oryzae[J].AMB Express,2016,6(1):57.
[21]ZHANG P,LIU T T,ZHOU P P,LI S T,YU L J.Agrobacterium tumefaciens-mediated transformation of a taxol-producing endophytic fungus,Cladosporium cladosporioides MD2[J].Current Microbiology,2011,62(4):1315-1320.
[22]王曦茁,朴春根,李虹,汪来发,郭民伟,李永,刘晓莉.根癌农杆菌介导的淡紫拟青霉遗传转化体系的建立[J].林业科学,2010,46(10):95-102.WANG X Z,PU C G,LI H,WANG L F,GUO M W,LI Y,LIU X L.Establishment of Agrobacterium tumefaciens-mediated transformation system of Paecilomyces lilacinus[J].Scientia Silvae Sinicae,2010,46(10):95-102.
[23]李敏慧,张荣,姜大刚,习平根,庄楚雄,姜子德.根癌农杆菌介导的香蕉枯萎病菌4号生理小种的转化[J].植物病理学报,2009,39(4):405-412.LI M H,ZHANG R,JIANG D G,XI P G,ZHUANG C X,JIANGZ D.Agrobacterium tumefaciens-mediated transformation of Fusarium oxysporum f.sp.cubense race 4[J].Acta Phytopathologica Phytopathologica Sinica,2009,39(4):405-412.
[24]LECLERQUE A,WAN H,ABSCHüTZ A,CHEN S,MITINA GV,ZIMMERMANN G,SCHAIRER H U.Agrobacterium-mediated insertional mutagenesis(AIM)of the entomopathogenic fungus Beauveria bassiana[J].Current Genetics,2004,45(2):111-119.
[25]MULLINS E D,KANG S.Transformation:a tool for studying fungal pathogens of plants[J].Cellular and Molecular Life Sciences,2001,58(14):2043-2052.
[26]TAKAHARA H,TSUJI G,KUBO Y,YAMAMOTO M,TOYODAK,INAGAKI Y,ICHINOSE Y,SHIRAISHI T.Agrobacterium tumefaciens-mediated transformation as a tool for random mutagenesis of Colletotrichum trifolii[J].Journal of General Plant Pathology,2004,70(2):93-96.
[27]SHARMA K K,KUHAD R C.Genetic transformation of lignin degrading fungi facilitated by Agrobacterium tumefaciens[J].BMCBiotechnology,2010,10(1):67.
[28]ZHENG Z,HUANG C,CAO L,XIE C,HAN L.Agrobacterium tumefaciens-mediated transformation as a tool for insertional mutagenesis in medicinal fungus Cordyceps militaris[J].Fungal Biology,2011,115(3):265-274.
[29]LIU N,CHEN G Q,NING G A,SHI H B,ZHANG L C,LU J P,MAO LJ,FENG X X,LIU X H,SU Z Z,LIN F C.Agrobacterium tumefaciensmediated transformation:An efficient tool for insertional mutagenesis and targeted gene disruption in Harpophora oryzae[J].Microbiological Research,2016,182:40-48.
[30]WANG D,HE D,LI G,GAO S,LV H Y,SHAN Q S,WANG L.An efficient tool for random insertional mutagenesis:Agrobacterium tumefaciens-mediated transformation of the filamentous fungus Aspergillus terreus[J].Journal of Microbiological Methods,2014,98:114-118.
[31]LIU T,LIU L,JIANG X,HOU J M,FU K H,ZHOU F H,CHENJ.Agrobacterium-mediated transformation as a useful tool for the molecular genetic study of the phytopathogen Curvularia lunata[J].European Journal of Plant Pathology,2010,126(3):363-371.
[32]李维,张义正.根癌农杆菌介导的白腐丝状真菌-黄孢原毛平革菌的转化[J].微生物学报,2005,45(5):784-787.LI W,ZHANG Y Z.Agrobacterium tumefaciens-mediated transformation of the white-rot basidiomycete,Phanerochaete chrysosporium[J].ActaMicrobiologica Sinica,2005,45(5):784-787.
[33]PARK S Y,JEONG M H,WANG H Y,KIM J A,YU A H,KIM SB,CHEONG Y H,KANG S C,LEE Y H,HUR J S.Agrobacterium tumefaciens-mediated transformation of the lichen fungus,Umbilicaria muehlenbergii[J].PloS One,2013,8(12):e83896.
[34]张俊华,刘烨,韩雨桐,潘春清,王中业,张淋淋,崔凯旋.农杆菌介导稻瘟病菌绿色荧光蛋白(GFP)遗传转化研究[J].东北农业大学学报,2014,45(11):1-7.doi:10.19720/j.cnki.issn.1005-9369.2014.11.001.ZHANG J H,LIU Y,HAN Y T,PAN C Q,WANG Z Y,ZHANG L LCUI K X.GFP genetic transformation of Magnaporthe grisea mediated by Agrobacterium tumefaciens[J].Journal of Northeast Agricultural University,2014,45(11):1-7.doi:10.19720/j.cnki.issn.1005-9369.2014.11.001.
[35]吴毅歆,范成明,周惠萍,久保康之,何月秋.一种农杆菌介导稻瘟病菌的遗传转化[J].植物保护学报,2008,35(5):421-426.doi:10.13802/j.cnki.zwbhxb.2008.05.008.WU Y X,FAN C M,ZHOU H P,YASUYUKI K B,HE Y Q.A genetic of transformation of Magnaporthe grisea by Agrobacterium tumefaciens[J].Acta Phytophylacica Sinica,2008,35(5):421-426.doi:10.13802/j.cnki.zwbhxb.2008.05.008.
[36]LU Y Y,XIAO S Q,WANG F,SUN J Y,ZHAO L K,YAN L B,XUE C S.Agrobacterium tumefaciens-mediated transformation as an efficient tool for insertional mutagenesis of Cercospora zeae-maydis[J].Journal of Microbiological Methods,2017,133:8-13.
[37]BOER P,BRONKHOF J,DUKI?K,KERKMAN R,TOUW H,BERGM,OFFRINGA R.Efficient gene targeting in Penicillium chrysogenum using novel Agrobacterium-mediated transformation approaches[J].Fungal Genetics and Biology,2013,61:9-14.
[38]LIU L,WEI Y M,ZHOU X W.Agrobacterium tumefaciens-mediated genetic transformation of the Taxol-producing endophytic fungus Ozonium sp EFY21[J].Genet Mol Res,2013,12(3):2913.
[39]ZHANG Y P,QU H X,ZHAO P,TANG Y P,ZHOU J S,LUO S C,YINY L,CHEN G Y.Generation and screening of T-DNA insertion mutants mediated by Agrobacterium tumefaciens in the garden asparagus stem blight pathogen Phomopsis asparagi[J].Current Microbiology,2017,74(11):1270-1277.
[40]WANG C X,GUAN X N,WANG H Y,LI G F,DONG X L,WANG GP,Li B H.Agrobacterium tumefaciens-mediated transformation of Valsa mali:an efficient tool for random insertion mutagenesis[J].The Scientific World Journal,2013,2013.
[41]LI G H,ZHOU Z Z,LIU G H,ZHENG F C,HE C Z.Characterization of T-DNA insertion patterns in the genome of rice blast fungus Magnaporthe oryzae[J].Current Genetics,2007,51(4):233-243.
[42]CHEN X L,YANG J,PENG Y L.Large-scale insertional mutagenesis in Magnaporthe oryzae by Agrobacterium tumefaciens-mediated transformation[J].Fungal Genomics,2011,722:213-224.
[43]JEON J,PARK S Y,CHI M H.Genome-wide functional analysis of pathogenicity genes in the rice blast fungus[J].Nature Genetics,2007,39(4):561-565.
[44]PENG H W,FENG Y J,ZHU X H,LAN X Y,TANG M,WANG J Z,DONG H T,CHEN B S.MoDUO1,a Duo1-like gene,is required for full virulence of the rice blast fungus Magnaporthe oryzae[J].Current Genetics,2011,57(6):409-420.
[45]WANG YU,HE D,CHU Y,ZUO Y S,XU X W,CHEN X L,ZHAO W S,ZHANG Y,YANG J,PENG Y L.MoCps1 is important for conidiation,conidial morphology and virulence in Magnaporthe oryzae[J].Current Genetics,2016,62(4):861-871.
[46]MOGGA V,DELVENTHAL R,WEIDENBACH D,LANGER S,BERTRAM P M,ANDRESEN K,THINES E,KROJ T,SCHAFFRATHU.Magnaporthe oryzae effectors MoHEG13 and MoHEG16 interfere with host infection and MoHEG13 counteracts cell death caused by Magnaporthe-NLPs in tobacco[J].Plant Cell Reports,2016,35(5):1169-1185.
[47]ZHANG S L,WANG L,WU W H,HE L Y,YANG X F,PAN Q H.Function and evolution of Magnaporthe oryzae avirulence gene AvrPib responding to the rice blast resistance gene Pib[J].Scientific Reports,2015,5:11642.
[48]吴小燕,王教瑜,张震,井金学,杜新法,柴荣耀,毛雪琴,邱海萍,姜华,王艳丽,孙国昌.稻瘟病菌致病性增强突变体B11的基因分析[J].中国水稻科学,2009,23(6):611-615.doi:10.3969/j.issn.1001-7216.2009.06.08.WU X Y,WANG J Y,ZHANG Z,JING J X,DU X F,CHAI R Y,MAOX Q,QIU H P,JIANG H,WANG Y L,SUN G C.Analysis on gene locus of Magnaporthe oryzae B11,a pathogenicity-enhanced mutant[J].Journal of China Rice Science,2009,23(6):611-615.doi:10.3969/j.issn.1001-7216.2009.06.08.
[49]孔丹丹.稻瘟病菌致病缺陷突变体KD-560的获得及标记基因MoCYP537A3功能的初步研究[D].杭州:浙江大学,2012.KONG D D.The obtaining of a nonpathogenic mutant Kd-560 of Magnaporthe oryzae and a primarily functional study on the tagged gene MoCYP537A3[D].Hangzhou:Zhejiang University,2012.