真菌病毒的研究进展
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摘要
为了发现和筛选具有生防价值的真菌病毒,通过对真菌病毒的分类、传播以及真菌病毒与寄主、与生物防治关系的综述,详细介绍了真菌病毒的全病毒科(Totiviridae)和产黄青霉病毒科(Chyroviridae),并列举了关于稻瘟病真菌病毒研究方面的实例。真菌营养不亲和性是限制真菌病毒传播和自然扩散的关键因素,在前人研究的基础上提出了几点建议:(1)筛选侵染性极强的真菌病毒;(2)发现和运用真菌病毒的传播介体;(3)运用化学物质抑制营养不亲和性。
In order to find and filtrate mycoviruses with biocontrol value, the taxonomy and transmission of mycovirus and the relationship of mycovirus with host and biocontrol were concluded, the family To tiviridaeand Chyroviridae were presented in detail, and an example for the study on fungal viruses in Magnaportheoryzae was set. The fungal vegetative incompatibility was a key limiting factor in mycovirus transmission andnatural spread, so several suggestions were made to solve this problem:(1) screening mycoviruses with stronginfectivity,(2) finding vectors of mycoviruses for application,(3) inhibiting vegetative in compatibility reaction using chemical compounds.
In order to find and filtrate mycoviruses with biocontrol value, the taxonomy and transmission of mycovirus and the relationship of mycovirus with host and biocontrol were concluded, the family To tiviridaeand Chyroviridae were presented in detail, and an example for the study on fungal viruses in Magnaportheoryzae was set. The fungal vegetative incompatibility was a key limiting factor in mycovirus transmission andnatural spread, so several suggestions were made to solve this problem:(1) screening mycoviruses with stronginfectivity,(2) finding vectors of mycoviruses for application,(3) inhibiting vegetative in compatibility reaction using chemical compounds.
引文
[1]Ghabrial S A.Origin,adaptation and evolutionary pathways of fungal viruses[J].Virus Genes,1998,16:119-131.
[2]Bao X,Roossinck M J.Multiplexed interactions:viruses of endophytic fungi[J].Advance in Virus Research,2013,86:37-58.
[3]Dawe A L,Nuss D L.Hypovirus molecular biology:from Koch'spostulates to host self-recognition genes that restrict virus transmission[J].Advance in Virus Research,2013,86(86):109-147.
[4]Ghabrial S A,Dunn S E,Li H,et al.Viruses of Helminthosporium(Cochlioblus)victoriae[J].Advance in Virus Research,2013,86:289-325.
[5]Hillman B I,Cai G.The family Narnaviridae:simplest of RNA viruses[J].Advance in Virus Research,2013,86(86):149-176.
[6]Hollings M.Viruses associated with a die-back disease of cultivated mushroom[J].Nature,1962,196:962-965.
[7]Xie J T,Jiang D H.New insights into mycoviruses and exploration for the biological control of crop fungal diseases[J].Annual Reviews of Phytopathology,2014,52:45-68
[8]Yu X,Li B,Fu Y P,et al.A geminivirus-related DNA mycovirus that confers hypovirulence to a plant pathogenic fungus[J].National Academy Sciences of USA,2010,107:8387-8392.
[9]Yu X,Li B,Fu Y P,et al.Extracellular transmission of a DNA mycovirus and its use as a natural fungicide[J].National Academy Sciences of USA,2013,110(4):1452-1457.
[10]Yokoi T,Yamashita S,Hibi T.The nucleotide sequence and genome organization of Magnaporthe oryzae virus 1[J].Archive of Virology,2007,152:2265-2269.
[11]Maejima K,Himeno M,Komatsu K,et al.Complete nucleotide sequence of a new double-stranded RNA virus from the rice blast fungus,Magnaporthe oryzae[J].Archive Virology,2008,153:389-391.
[12]Tang L H,Hu Y P,Liu L J,et al.Genomic organization of a novel victorivirus from the rice blast fungus Magnaporthe oryzae[J].Archive of Virology,2015,160(11):2907-2910.
[13]Urayama S,Kato S,Suzuki Y,et al.Mycoviruses related to chrysovirus affect vegetative growth in the rice blast fungus Magnaporthe oryzae[J].Journal of General Virology,2010,91:3085-3094.
[14]Urayama S,Ohta T,Onozuka N,et al.Characterization of Magnaporthe oryzae chrysovirus 1 Structural Proteins and Their Expression in Saccharomyces cerevisiae[J].Journal of General Virology,2012,86(15):8287.
[15]Urayama S,Sakoda H,Takai R,et al.A ds RNA mycovirus,Magnaporthe oryzae chrysovirus1-B,suppresses vegetative growth and development of the rice blast fungus[J].Virology,2014,448:265-273.
[16]Ghabrial S A,Castón J R,Jiang D H,et al.50-plus years of fungal viruses[J].Virology,2015,479-480:356-368
[17]Ghabrial S A,Suzuki N.Viruses of plantpathogenic fungi[J].Annual Reviews of Phytopathology,2009,47:353-384.
[18]Wickner R B,Ghabrial S A,Nibert M L,et al.Totiviridae In:Virus Taxonomy,ninth Report of the International Committee on Taxonomy of Viruses[M].London,UK:Elsevier press,2011,639-650.
[19]Baeza M,Bravo N,Sanhueza M,et al.Molecular characterization of totiviruses in Xanthophyllomyces dendrorhous[J].Virology Journal,2012,9(1):140.
[20]Stielow B,Menzel W.Complete nucleotide sequence of Ta V1,a novel totivirus isolated from a black truffle ascocarp(Tuber aestivum Vittad.)[J].Archive of Virology,2010,155:2075-2078.
[21]Taylor D J,Ballinger M J,Bowman S M,et al.Virus–host coevolution under a modified nuclear genetic code[J].Peer J,2013,1:e50.
[22]Ghabrial S A,Nibert M L.Victorivirus,a new genus of fungal viruses in the family Totiviridae[J].Archive of Virology,2009,154:373-379.
[23]Wickner R B.Double-stranded RNA viruses of Saccharomyces cerevisiae[J].Microbiology Reviews,1996,60:250-265.
[24]Wickner R B,Tang J,Gardner N A,et al.The yeast ds RNA Virus LA resembles mammalian ds RNA virus cores[M].Caister Academic Press,2008,ISBN 978-1-904455-21-9.
[25]Soldevila A,Ghabrial S A.Expression of the totivirus Helminthasporium victoriae 190S Virus RNA dependent RNA polymerase from its down stream open reading frame in dicistronic constructs[J].Journal of General Virology,2000,74:997-1003.
[26]Li H,Havens W M,Nibert M L,et al.RNA sequence determinants of a coupled termination-reinitiation strategy for downstream open reading frame franslation in Helminthosporium victoriae virus 190S and other victoriviruses(family Totiviridae)[J].Journal of General Virology,2011,85:7343-7352.
[27]Jiang D,Ghabrial S A.Molecular characterization of Penicillium chrysogenum virus:reconsideration of the taxonomy of the genus Chrysovirus[J].Journal of General Virology,2004,85:2111-2113.
[28]Castón J R,Ghabrial S A,Jiang D,et al.Three-dimensional structure of Penicillium chrysogenum virus:a double stranded RNA virus with a genuine T=1 capsid[J].Journal of Moleclar Biology,2003,331:417-431.
[29]Nuss D L.Hypovirulence:mycoviruses at tixe fungal-plant interface[J].Nature Reviews of Microbiology,2005,3:632-642.
[30]Choi G H,Dawe A L,Churbanov A,et al.Molecular characterization of vegetative incompatibility genes that restrict hypovirus transmission in the chestnut blight fungus Cryphonectria parasitica[J].Genetics,2012,190:113-270.
[31]Gao K,Xiong Q,Xu J,et al.Cp Birl is required for conidiation,virulence and anti-apoptotic effects and influences hypovirus transmission in Cryphonectria parasitica[J].Fungal Genetics and Biology,2013,51:60-71.
[32]Anagnostakis S L,Chen B,Geletka L M,et al.Hypovirus transmission to ascospore progeny by field-released transgenic hypovirulent strains of Cryphonectria parasitica[J].Phytopathology,1998,88:598-604.
[33]Xie J,Ghabrial S A.Molecular characterizations of two mitoviruses co-infecting a hyovirulent isolate of the plant pathogenic fungus Sclerotinia sclerotiorum[J].Virology,2012,428:77-85.
[34]Yaegashi H,Nakamura H,Sawahata T,et al.Appearance of mycovirus-like double-stranded RNAs in the white root rot fungus,Rosellinia necatrix,in an apple orchard[J].FEMS microbiology ecology,2013,83(1):40–62
[35]Nuss D L.Biological control of chestnut blight:an example of virus-mediated attenuation of fungal pathogenesis[J].Microbiology Reviews,1992,56:561-576.
[36]Kanematsu S,Arakawa M,Oikawa Y,et al.A reovirus causes hypovirulence of Rosellinia necatrix[J].Phytopathology,2004,94:561-568.
[37]Chu Y M,Jeon J J,Yea S J,et al.Double-stranded mycovirus from Fusarium graminearum[J].Applied and Environmental Microbiology,2002,68:2529-2534.
[38]Cho W K,Lee K M,Yu J,et al.Insight into mycoviruses infecting Fusarium species[J].Advance in Virus Research,2013,86(86):273-288.
[39]Nuss D L.Mycoviruses,RNA silencing,and viral RNA recombination.Advances in Virus Research,2001,80:25–48
[40]Dunn S E,Li H,Cardone G,et al.Three-dimensional structure of victorivirus Hv V190S suggests coat proteins in most totiviruses share a conserved core.PLo S Pathogens,2013,9(3):e1003225
[41]Zhu W,Wei W,Fu Y,et al.A secretory protein of necrotrophic fungus Sclerotinia sclerotiorum that suppresses host resistance.PLo S ONE,2013,8(1):e53901
[42]Márquez L M,Redman R S,Rodriguez R J.A virus in a fungus in a plant:three-way symbiosis required for thermal tolerance[J].Science,2007,315:513-515.
[43]Biraghi A.Possible active resistance to Endothia parasitica in Castanea sativa.Reports to Congress of the International Union of Research Organism,1953,11th,Rome
[44]Anagnostakis S L.Biological control of chestnut blight[J].Science,1982,215:466-471.
[45]Shapira R,Choi G H,Nuss D L.Virus-like genetic organization and expression strategy for a double-stranded RNA genetic element associated with biological control of chestnut blight.EMBO Journal,1991,10(4):731-739
[46]Hillman B I,Tian Y,Bedker P J,et al.A North American hypovirulent isolate of the chestnut blight fungus with European isolate-related ds RNA.Journal of General Virology,1992,73:681–686
[47]Smart C D,Yuan W,Foglia R,et al.Cryphonectria hypovirus 3,a virus species in the family Hypoviridae with a single open reading frame.Virology,1999,265:66-73
[48]Linder-Basso D,Dynek J N,Hillman B I.Genome analysis of Cryphonectria hypovirus 4,the most common hypovirus species in North America.Virology,2005,337:192-203
[49]Xiao X Q,Cheng J S,Tang J H,et al.A novel partitivirus that confers hypovirulence on plant pathogenic fungi.Journal of Virology,2014,88(17):10120–10133
[50]Chiba S,Salaipeth L,Lin Y H,et al.A novel bipartite ds RNA mycovirus from the white root rot fungus Rosellinia necatrix:molecular and biological characterization,taxonomic considerations,and potential for biological control.Journal of Virology,2009,83:12801-12812.
[51]Dean R A,Talbot N J,Ebbole D J,et al.The genome sequence of the rice blast fungus Magnaporthe oryzae[J].Nature,2005,434:980-986.
[52]Odenbach D,Breth B,Thines E,et al.The transcription factor Con7p is a central regulator of infection-related morphogenesis in the rice blast fungus Magnaporthe grisea[J].Molecular Microbiology,2007,64(2):293-307.
[53]Hamer J E,Howard R J,Chumley F G,et al.A mechanism for surface attachment in spores of a plant pathogenic fungus[J].Science,1988,239(4837):288-290.
[54]Howard R J,Valent B.Breaking and entering:host penetration by the fungal rice blast pathogen Magnaporthe oryzae[J].Annual Reviews of Microbiology,1996,50(1):491-512.
[55]Yamashita S,Doi Y,Yorak.A polydedral virus found in rice blast fungus,Pyricularia oryzae Cavara[J].Annual phytopathology Soc Jpn,1971,37:356-359.
[56]Hunst P L,Latterell F M,Rossi A E.Variation in double-stranded RNA from isolates of Pyricularia oryzae[J].Phytopathology,1986,76:674-678.
[57]Tuomivirta T T,Hantula J.Two unrelated double-stranded RNA molecule patterns in Gremmeniella abietina type A code for putative viruses of the families Totiviridae and Partitiviridae[J].Archive of Virology,2003,148(12):2293-2305.
[58]Cheng J S,Jiang D H,Fu Y P,et al.Molecular characterization of a ds RNA totivirus infecting the sclerotial parasite Coniothyrium minitans[J].Virus Research,2003,93(1):41-50.
[59]Huang S,Ghabrial S A.Organization and expression of the doublestranded RNA genome of Helminthosporium victoriae 190S virus,a totivirus infecting a plant pathogenic filamentous fungus[J].Proceedings of the National Academy of Sciences of the United states of America,1996,93(22):12541-12546.
[60]Stuart K D,Weeks R,Guilbride L,et al.Molecular organization of Leishmania RNA virus 1[J].Proceedings of the National Academy of Sciences of the United states of America,1992,89(18):8596-8600.
[61]Scheffter S M,Ro Y T,Chung I K,et al.The complete sequence of Leishmania RNA virus LRV2-1,a virus of an old world parasite strain[J].Virology,1995,212(1):84-90.
[62]Goodman R P,Freret T S,Kula T,et al.Clinical isolates of Trichomonas vaginalis concurrently infected by strains of up to four Trichomonasvirus species(family Totiviridae)[J].Journal of Virology,2011,85(9):4258-4270.
[63]Icho T,Wickner R B.The double-stranded RNA genome of yeast virus L-A encodes its own putative RNA polymerase by fusing two open reading frames[J].The Journal of Biological Chemistry,1989,264(12):6716-6723.
[64]Park C M,Lopinski J D,Masuda J,et al.A second double-stranded RNA virus from yeast[J].Virology,1996,216(2):451-454.
[65]Kim J M,Kim J A,Park J A,et al.Molecular diversity of chrysoviruses in Korean isolates of a new fungal species,Cryphonectria nitschkei[J].Journal of Microbiology,2009,47(4):441-447.
[66]Cao Y F,Zhu X W,Xiang Y,et al.Genomic characterization of a novel ds RNA virus detected in the phytopathogenic fungus Verticillium dahliae Kleb[J].Virus Research,2011,159(1):73-78.
[67]Ikeda K,Inoue K,Kida C,et al.Potentiation of mycovirus transmission by zinc compounds via attenuation of heterogenic incompatibility in Rosellinia necatrix[J].Applied and Environmental Microbiology,2013,79(12):3684-3691.
[2]Bao X,Roossinck M J.Multiplexed interactions:viruses of endophytic fungi[J].Advance in Virus Research,2013,86:37-58.
[3]Dawe A L,Nuss D L.Hypovirus molecular biology:from Koch'spostulates to host self-recognition genes that restrict virus transmission[J].Advance in Virus Research,2013,86(86):109-147.
[4]Ghabrial S A,Dunn S E,Li H,et al.Viruses of Helminthosporium(Cochlioblus)victoriae[J].Advance in Virus Research,2013,86:289-325.
[5]Hillman B I,Cai G.The family Narnaviridae:simplest of RNA viruses[J].Advance in Virus Research,2013,86(86):149-176.
[6]Hollings M.Viruses associated with a die-back disease of cultivated mushroom[J].Nature,1962,196:962-965.
[7]Xie J T,Jiang D H.New insights into mycoviruses and exploration for the biological control of crop fungal diseases[J].Annual Reviews of Phytopathology,2014,52:45-68
[8]Yu X,Li B,Fu Y P,et al.A geminivirus-related DNA mycovirus that confers hypovirulence to a plant pathogenic fungus[J].National Academy Sciences of USA,2010,107:8387-8392.
[9]Yu X,Li B,Fu Y P,et al.Extracellular transmission of a DNA mycovirus and its use as a natural fungicide[J].National Academy Sciences of USA,2013,110(4):1452-1457.
[10]Yokoi T,Yamashita S,Hibi T.The nucleotide sequence and genome organization of Magnaporthe oryzae virus 1[J].Archive of Virology,2007,152:2265-2269.
[11]Maejima K,Himeno M,Komatsu K,et al.Complete nucleotide sequence of a new double-stranded RNA virus from the rice blast fungus,Magnaporthe oryzae[J].Archive Virology,2008,153:389-391.
[12]Tang L H,Hu Y P,Liu L J,et al.Genomic organization of a novel victorivirus from the rice blast fungus Magnaporthe oryzae[J].Archive of Virology,2015,160(11):2907-2910.
[13]Urayama S,Kato S,Suzuki Y,et al.Mycoviruses related to chrysovirus affect vegetative growth in the rice blast fungus Magnaporthe oryzae[J].Journal of General Virology,2010,91:3085-3094.
[14]Urayama S,Ohta T,Onozuka N,et al.Characterization of Magnaporthe oryzae chrysovirus 1 Structural Proteins and Their Expression in Saccharomyces cerevisiae[J].Journal of General Virology,2012,86(15):8287.
[15]Urayama S,Sakoda H,Takai R,et al.A ds RNA mycovirus,Magnaporthe oryzae chrysovirus1-B,suppresses vegetative growth and development of the rice blast fungus[J].Virology,2014,448:265-273.
[16]Ghabrial S A,Castón J R,Jiang D H,et al.50-plus years of fungal viruses[J].Virology,2015,479-480:356-368
[17]Ghabrial S A,Suzuki N.Viruses of plantpathogenic fungi[J].Annual Reviews of Phytopathology,2009,47:353-384.
[18]Wickner R B,Ghabrial S A,Nibert M L,et al.Totiviridae In:Virus Taxonomy,ninth Report of the International Committee on Taxonomy of Viruses[M].London,UK:Elsevier press,2011,639-650.
[19]Baeza M,Bravo N,Sanhueza M,et al.Molecular characterization of totiviruses in Xanthophyllomyces dendrorhous[J].Virology Journal,2012,9(1):140.
[20]Stielow B,Menzel W.Complete nucleotide sequence of Ta V1,a novel totivirus isolated from a black truffle ascocarp(Tuber aestivum Vittad.)[J].Archive of Virology,2010,155:2075-2078.
[21]Taylor D J,Ballinger M J,Bowman S M,et al.Virus–host coevolution under a modified nuclear genetic code[J].Peer J,2013,1:e50.
[22]Ghabrial S A,Nibert M L.Victorivirus,a new genus of fungal viruses in the family Totiviridae[J].Archive of Virology,2009,154:373-379.
[23]Wickner R B.Double-stranded RNA viruses of Saccharomyces cerevisiae[J].Microbiology Reviews,1996,60:250-265.
[24]Wickner R B,Tang J,Gardner N A,et al.The yeast ds RNA Virus LA resembles mammalian ds RNA virus cores[M].Caister Academic Press,2008,ISBN 978-1-904455-21-9.
[25]Soldevila A,Ghabrial S A.Expression of the totivirus Helminthasporium victoriae 190S Virus RNA dependent RNA polymerase from its down stream open reading frame in dicistronic constructs[J].Journal of General Virology,2000,74:997-1003.
[26]Li H,Havens W M,Nibert M L,et al.RNA sequence determinants of a coupled termination-reinitiation strategy for downstream open reading frame franslation in Helminthosporium victoriae virus 190S and other victoriviruses(family Totiviridae)[J].Journal of General Virology,2011,85:7343-7352.
[27]Jiang D,Ghabrial S A.Molecular characterization of Penicillium chrysogenum virus:reconsideration of the taxonomy of the genus Chrysovirus[J].Journal of General Virology,2004,85:2111-2113.
[28]Castón J R,Ghabrial S A,Jiang D,et al.Three-dimensional structure of Penicillium chrysogenum virus:a double stranded RNA virus with a genuine T=1 capsid[J].Journal of Moleclar Biology,2003,331:417-431.
[29]Nuss D L.Hypovirulence:mycoviruses at tixe fungal-plant interface[J].Nature Reviews of Microbiology,2005,3:632-642.
[30]Choi G H,Dawe A L,Churbanov A,et al.Molecular characterization of vegetative incompatibility genes that restrict hypovirus transmission in the chestnut blight fungus Cryphonectria parasitica[J].Genetics,2012,190:113-270.
[31]Gao K,Xiong Q,Xu J,et al.Cp Birl is required for conidiation,virulence and anti-apoptotic effects and influences hypovirus transmission in Cryphonectria parasitica[J].Fungal Genetics and Biology,2013,51:60-71.
[32]Anagnostakis S L,Chen B,Geletka L M,et al.Hypovirus transmission to ascospore progeny by field-released transgenic hypovirulent strains of Cryphonectria parasitica[J].Phytopathology,1998,88:598-604.
[33]Xie J,Ghabrial S A.Molecular characterizations of two mitoviruses co-infecting a hyovirulent isolate of the plant pathogenic fungus Sclerotinia sclerotiorum[J].Virology,2012,428:77-85.
[34]Yaegashi H,Nakamura H,Sawahata T,et al.Appearance of mycovirus-like double-stranded RNAs in the white root rot fungus,Rosellinia necatrix,in an apple orchard[J].FEMS microbiology ecology,2013,83(1):40–62
[35]Nuss D L.Biological control of chestnut blight:an example of virus-mediated attenuation of fungal pathogenesis[J].Microbiology Reviews,1992,56:561-576.
[36]Kanematsu S,Arakawa M,Oikawa Y,et al.A reovirus causes hypovirulence of Rosellinia necatrix[J].Phytopathology,2004,94:561-568.
[37]Chu Y M,Jeon J J,Yea S J,et al.Double-stranded mycovirus from Fusarium graminearum[J].Applied and Environmental Microbiology,2002,68:2529-2534.
[38]Cho W K,Lee K M,Yu J,et al.Insight into mycoviruses infecting Fusarium species[J].Advance in Virus Research,2013,86(86):273-288.
[39]Nuss D L.Mycoviruses,RNA silencing,and viral RNA recombination.Advances in Virus Research,2001,80:25–48
[40]Dunn S E,Li H,Cardone G,et al.Three-dimensional structure of victorivirus Hv V190S suggests coat proteins in most totiviruses share a conserved core.PLo S Pathogens,2013,9(3):e1003225
[41]Zhu W,Wei W,Fu Y,et al.A secretory protein of necrotrophic fungus Sclerotinia sclerotiorum that suppresses host resistance.PLo S ONE,2013,8(1):e53901
[42]Márquez L M,Redman R S,Rodriguez R J.A virus in a fungus in a plant:three-way symbiosis required for thermal tolerance[J].Science,2007,315:513-515.
[43]Biraghi A.Possible active resistance to Endothia parasitica in Castanea sativa.Reports to Congress of the International Union of Research Organism,1953,11th,Rome
[44]Anagnostakis S L.Biological control of chestnut blight[J].Science,1982,215:466-471.
[45]Shapira R,Choi G H,Nuss D L.Virus-like genetic organization and expression strategy for a double-stranded RNA genetic element associated with biological control of chestnut blight.EMBO Journal,1991,10(4):731-739
[46]Hillman B I,Tian Y,Bedker P J,et al.A North American hypovirulent isolate of the chestnut blight fungus with European isolate-related ds RNA.Journal of General Virology,1992,73:681–686
[47]Smart C D,Yuan W,Foglia R,et al.Cryphonectria hypovirus 3,a virus species in the family Hypoviridae with a single open reading frame.Virology,1999,265:66-73
[48]Linder-Basso D,Dynek J N,Hillman B I.Genome analysis of Cryphonectria hypovirus 4,the most common hypovirus species in North America.Virology,2005,337:192-203
[49]Xiao X Q,Cheng J S,Tang J H,et al.A novel partitivirus that confers hypovirulence on plant pathogenic fungi.Journal of Virology,2014,88(17):10120–10133
[50]Chiba S,Salaipeth L,Lin Y H,et al.A novel bipartite ds RNA mycovirus from the white root rot fungus Rosellinia necatrix:molecular and biological characterization,taxonomic considerations,and potential for biological control.Journal of Virology,2009,83:12801-12812.
[51]Dean R A,Talbot N J,Ebbole D J,et al.The genome sequence of the rice blast fungus Magnaporthe oryzae[J].Nature,2005,434:980-986.
[52]Odenbach D,Breth B,Thines E,et al.The transcription factor Con7p is a central regulator of infection-related morphogenesis in the rice blast fungus Magnaporthe grisea[J].Molecular Microbiology,2007,64(2):293-307.
[53]Hamer J E,Howard R J,Chumley F G,et al.A mechanism for surface attachment in spores of a plant pathogenic fungus[J].Science,1988,239(4837):288-290.
[54]Howard R J,Valent B.Breaking and entering:host penetration by the fungal rice blast pathogen Magnaporthe oryzae[J].Annual Reviews of Microbiology,1996,50(1):491-512.
[55]Yamashita S,Doi Y,Yorak.A polydedral virus found in rice blast fungus,Pyricularia oryzae Cavara[J].Annual phytopathology Soc Jpn,1971,37:356-359.
[56]Hunst P L,Latterell F M,Rossi A E.Variation in double-stranded RNA from isolates of Pyricularia oryzae[J].Phytopathology,1986,76:674-678.
[57]Tuomivirta T T,Hantula J.Two unrelated double-stranded RNA molecule patterns in Gremmeniella abietina type A code for putative viruses of the families Totiviridae and Partitiviridae[J].Archive of Virology,2003,148(12):2293-2305.
[58]Cheng J S,Jiang D H,Fu Y P,et al.Molecular characterization of a ds RNA totivirus infecting the sclerotial parasite Coniothyrium minitans[J].Virus Research,2003,93(1):41-50.
[59]Huang S,Ghabrial S A.Organization and expression of the doublestranded RNA genome of Helminthosporium victoriae 190S virus,a totivirus infecting a plant pathogenic filamentous fungus[J].Proceedings of the National Academy of Sciences of the United states of America,1996,93(22):12541-12546.
[60]Stuart K D,Weeks R,Guilbride L,et al.Molecular organization of Leishmania RNA virus 1[J].Proceedings of the National Academy of Sciences of the United states of America,1992,89(18):8596-8600.
[61]Scheffter S M,Ro Y T,Chung I K,et al.The complete sequence of Leishmania RNA virus LRV2-1,a virus of an old world parasite strain[J].Virology,1995,212(1):84-90.
[62]Goodman R P,Freret T S,Kula T,et al.Clinical isolates of Trichomonas vaginalis concurrently infected by strains of up to four Trichomonasvirus species(family Totiviridae)[J].Journal of Virology,2011,85(9):4258-4270.
[63]Icho T,Wickner R B.The double-stranded RNA genome of yeast virus L-A encodes its own putative RNA polymerase by fusing two open reading frames[J].The Journal of Biological Chemistry,1989,264(12):6716-6723.
[64]Park C M,Lopinski J D,Masuda J,et al.A second double-stranded RNA virus from yeast[J].Virology,1996,216(2):451-454.
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