四川省马铃薯晚疫病菌生物学特性与遗传多样性研究
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摘要
由致病疫霉Phytophthora infestans(Mont) de Bary引起的马铃薯晚疫病是马铃薯生产中的一种毁灭性病害,在全球范围内绝大多数马铃薯栽培地区广泛传播。四川省是我国马铃薯重要产区之一,2003年的种植面积1500km~2左右,2006年种植面积上升到6326km~2。随着我省马铃薯生产的快速发展,品种调运日益频繁,会造成我省马铃薯晚疫病菌生理小种种群结构和种群数量的变化,会使我省的部分马铃薯品种的抗病性发生改变,给马铃薯晚疫病大发生埋下了隐患。因此,加强对病菌生物学特性和群体结构的研究,对指导马铃薯晚疫病的防治具有重要意义。本研究从晚疫病菌的表现型(交配型、生理小种、抗药性)、生物学特性(不同环境下孢子释放能力等)和病菌群体遗传分析(DNA分子标记)等方面对四川马铃薯主产区晚疫病菌进行了系统的研究。
2006年～2007年对四川省7个马铃薯主产区的马铃薯晚疫病的主要研究结果如下:
1.对158个单孢菌株,利用A1标准菌株进行交配型鉴定,结果均未发现卵孢子形成,表明所测试的菌株均为A1交配型菌株,无A2交配型菌株。
2.利用主效基因R1.R2,R3,R4的4个鉴别寄主对131个单孢菌种进行生理小种鉴定,明确我省有8个生理小种类型,其优势种群为3,4号小种,占56%,次优势种群为3号小种,占17.6%。
3.雷多米尔抗药性测定结果表明:60个单孢菌株中无抗药性菌株,只有8个中度抗性菌株,占被测菌株数的13.33%。彭州市的中度抗性菌株占40%;新都的中度抗性菌株占28.57%;其余地方中度抗性菌株均为0。
4.晚疫病菌生物学特性研究发现:马铃薯晚疫病病原菌菌丝生长对光照很敏感,光照不利于病原菌的生长,适宜在黑暗的条件下生长;不同的糖类及含糖量对病原菌孢子囊的萌发,游动孢子的释放并没有太大的影响;NaCL对孢子囊萌发有一定的抑制作用;偏酸的环境有利于孢子囊的萌发,碱性环境对孢子囊萌发具有一定的抑制作用。
5.利用两个SSR引物Pi4B和Pi4G对来自四川七个不同地区不同年份的91个马铃薯晚疫病病原菌株进行基因型鉴定,共鉴定出8个SSR基因型:D-05、D-03、F-01、F-03、F-06、G-02和H-01。F-01为优势基因型,占参试菌株的32.97%。
Late blight caused by the oomycete Phytophthora infestans (Mont.) de Bary is a disastrous disease in the potato growing areas all over the world. Sichuan is one of the major potato production areas in China. The planting areas is 1,500km~2 in 2003s, It rises to 6,326ks~2 in 2006s . Along with the potato production's fast development, the potato transport becoming more frequently. The structure and variation of P.infestans population are change. It laid down a hidden danger for the occurrence and epidemics of this disease. It is necessary to production of potato to strengthen the research on population structure. This study focuses on the phenotypic(mating type、physiological race、biological traits、chemical resistance) and SSR genotypic diversity of. P.infestans population in the major potato producing areas in Sichuan province.
Main results were summarized as follows:
1. 158 isolates purified from the samples of potato late blight which collected from six major potato-growing areas in Sichuan province were tested for their mating types by using the reference isolates of A1 and A2 mating type. None oospore could be found .All of isolates was subjected to A1 mating type, no A2 mating type was found.
2. Among 131 isolates tested with 4 potato clones of the international late blight differential hosts containing single dominant resistant gene R1,R2,R3,R4, 8 races could be distinguished with different virulence spectrum, and race 3,4 appeared at the highest frequency of 58%,followed by superior race of 3 with the frequency of 17.6% .
3. The results from the chemical resistance test showed that among 60 isolates purified from the samples of potato late blight which collected from six major potato-growing areas in Sichuan province tested, none of them was found resistance to ridomil—M258WP,only found 8 isolates with intermediate resistance. So the ridomil—MZ58WP could continue use to control late blight in the areas.
4. The results from the different illumination time test showed that the P.infestans couldn' t grow, the illumination had inhibitory effect to P.infestans growth. In different ph and nutrient conditions test the sporangium' s release rate. The results show the sμgar types and content had no affected to sporangium' s release rate. NaCL had inhibitory effect to it;PH4 was the optimum ph;PH 10 had suppression.
5. Used 2 SSR-primer: Pi4B and Pi4G to detect 91 isolates from 7 different production areas in Sichuan. Found 8 SSR genotypes .which are D-05、D-03、F-01、F-03、F-06、G-02 and H-01. SSR genotype F-01 appeared at the highest frequency of 39.56%.
2006年～2007年对四川省7个马铃薯主产区的马铃薯晚疫病的主要研究结果如下:
1.对158个单孢菌株,利用A1标准菌株进行交配型鉴定,结果均未发现卵孢子形成,表明所测试的菌株均为A1交配型菌株,无A2交配型菌株。
2.利用主效基因R1.R2,R3,R4的4个鉴别寄主对131个单孢菌种进行生理小种鉴定,明确我省有8个生理小种类型,其优势种群为3,4号小种,占56%,次优势种群为3号小种,占17.6%。
3.雷多米尔抗药性测定结果表明:60个单孢菌株中无抗药性菌株,只有8个中度抗性菌株,占被测菌株数的13.33%。彭州市的中度抗性菌株占40%;新都的中度抗性菌株占28.57%;其余地方中度抗性菌株均为0。
4.晚疫病菌生物学特性研究发现:马铃薯晚疫病病原菌菌丝生长对光照很敏感,光照不利于病原菌的生长,适宜在黑暗的条件下生长;不同的糖类及含糖量对病原菌孢子囊的萌发,游动孢子的释放并没有太大的影响;NaCL对孢子囊萌发有一定的抑制作用;偏酸的环境有利于孢子囊的萌发,碱性环境对孢子囊萌发具有一定的抑制作用。
5.利用两个SSR引物Pi4B和Pi4G对来自四川七个不同地区不同年份的91个马铃薯晚疫病病原菌株进行基因型鉴定,共鉴定出8个SSR基因型:D-05、D-03、F-01、F-03、F-06、G-02和H-01。F-01为优势基因型,占参试菌株的32.97%。
Late blight caused by the oomycete Phytophthora infestans (Mont.) de Bary is a disastrous disease in the potato growing areas all over the world. Sichuan is one of the major potato production areas in China. The planting areas is 1,500km~2 in 2003s, It rises to 6,326ks~2 in 2006s . Along with the potato production's fast development, the potato transport becoming more frequently. The structure and variation of P.infestans population are change. It laid down a hidden danger for the occurrence and epidemics of this disease. It is necessary to production of potato to strengthen the research on population structure. This study focuses on the phenotypic(mating type、physiological race、biological traits、chemical resistance) and SSR genotypic diversity of. P.infestans population in the major potato producing areas in Sichuan province.
Main results were summarized as follows:
1. 158 isolates purified from the samples of potato late blight which collected from six major potato-growing areas in Sichuan province were tested for their mating types by using the reference isolates of A1 and A2 mating type. None oospore could be found .All of isolates was subjected to A1 mating type, no A2 mating type was found.
2. Among 131 isolates tested with 4 potato clones of the international late blight differential hosts containing single dominant resistant gene R1,R2,R3,R4, 8 races could be distinguished with different virulence spectrum, and race 3,4 appeared at the highest frequency of 58%,followed by superior race of 3 with the frequency of 17.6% .
3. The results from the chemical resistance test showed that among 60 isolates purified from the samples of potato late blight which collected from six major potato-growing areas in Sichuan province tested, none of them was found resistance to ridomil—M258WP,only found 8 isolates with intermediate resistance. So the ridomil—MZ58WP could continue use to control late blight in the areas.
4. The results from the different illumination time test showed that the P.infestans couldn' t grow, the illumination had inhibitory effect to P.infestans growth. In different ph and nutrient conditions test the sporangium' s release rate. The results show the sμgar types and content had no affected to sporangium' s release rate. NaCL had inhibitory effect to it;PH4 was the optimum ph;PH 10 had suppression.
5. Used 2 SSR-primer: Pi4B and Pi4G to detect 91 isolates from 7 different production areas in Sichuan. Found 8 SSR genotypes .which are D-05、D-03、F-01、F-03、F-06、G-02 and H-01. SSR genotype F-01 appeared at the highest frequency of 39.56%.
引文
[1]..Goodwin,S.B.,Spielman,et al..Clonal Diversity and Genetic Differentiation of Phytophthora infestans populations in Northern and Gentral Mexico.Phytopathology,1992,82:9-55-961
[2].Reddick,D.Development of Blight Immune varieties.American Potato Journal,1943,20:118-126
[3].李克来,马铃薯晚疫病及抗晚疫病育种进展,晚疫病病原体的生物学,马铃薯科学1984,1:32-36
[4].Austin Bourke P.M.Eergence of potato blight,1843-46 Nature.1964,August,22,203:805-808
[5].林传光,黄河,王高才等.马铃薯晚疫病的田间动态观察及防治试验.植物病理学报.1955,1(1):31-43.
[6].农山渔村文化编李国光、姚方杰译新版蔬菜病虫害诊断原色图谱2001年1月第1版:171
[7].林传光,黄河,霍守祥.关于马铃薯晚疫病的预测和防治研究(摘要)北京农业大学学报,1956,2(1):79-85
[8].林传光,黄河,王道本,等.马铃薯晚疫病中心病株形成的观察.植物病理学报,1957,3(1):19-29
[9].朱杰华,张志铭,李玉琴,等.河北省马铃薯晚疫病病原菌(Phytophthora infestans)A2交配型的分布河北农业大学学报,2000b,23(3):73-75
[10].姚裕琪,巩秀峰,高奇华,等.马铃薯晚疫病抗性鉴定及评价.内蒙古农业科技,2001 C 2＞:8-9
[11].朱杰华,杨志辉,曹克强,等.影响活体内马铃薯晚疫病病原菌卵孢子产生因素的研究.菌物系统,2001,20(2):244-249
[12].Zwankhuizen M J,Govers F,Zadoks J C.Inoculum sources and genotypic diversity of Phytophthora infestans in Southern Flevoland,the Netherlands.European journal of Plant Pathology,2000,106:667-680
[13].郑小波1997疫霉菌及其研究技术中国农业出版社27-28
[14].Gallegly et al.,1958;Smoot et al.,1958;Spielman eral.,1991;Fry et aL,1993
[15].马铃薯晚疫病病原菌遗传多样性及其小种特异无毒基因遗传定位和转录表达谱研究
[16].Goodwin S B,Cohen B A,Deahl KL.Migration from Northern Mexico as the probable cause of recent genetic changes in populations of Phytophthora infestans in the United States and Canada[J].Phytopathology,1994,(84):553-5581
[17].Deahl K L,DeMuth S P,Sinden S L,et al.Identification of mating types and metalaxyl resistance in North American populations of Phytophthora infestans[J].American Potato Journal,1995,(72):35-491
[18].Goodwin S B,Sujkowski L S,Dyer A T,et al.Direct detection of gene flow and probable sexual reproduction of Phytophthora infestans in Northern North America[J]Phytopathology,1995,(85):473-4791
[19].Goodwin S B,Sujkowski L S,Fry W E.Widespread distributionand probable origin of resistance to metalaxyl in clonal genotypesof Phytophthora infestans in the United States and Western Canada[J].Phytopathology,1996,(86):793-8001
[20].Andrivon D Beasse C,Laurent C.Characterization of isolates of Phytophthora infestans collected in Northwestern France from1988 to 1992[J].Plant Pathology,1994,(43):471-4781
[21].Daggett S S,GEtz E,Therrien C D.Phenotypic changes in populations of Phytophthora infestans from Eastern Germany[J].Phytopathology,1993,(83):319-3231
[22].Sujkowski L S,Goodwin S B,Dyer A T,et al.Increased genotypic diversity via migration and possible occurrence of sexual reproduction of Phytophthora infestans in Ploland[J].Phytopathology,1994,(84):201-2071
[23].Forbes G A,Escobar X C,Ayala C C,et al.Populaton genetic structure of Phytophthora infestans in Ecuador[J].Phytopathology,1997,(87):375-3801
[24].Koh Y J,Goodwin S B,Dyer A T,et al.Migrations and displacements of Phytophthora infestans populations in East Asian countries[J].Phytopathology,1994,(84):922-9271
[25].Hohl ILK,Iselin K.Strains of Phytophthora infestans with A2 mating type behaviour.Transactions of the British Mycological Society,1984,83:529-530.
[26].Tantius P.M.,Fyfe AM.,Shaw D.S.Occurrence of the A2 mating type and self-fertile isolates of Phytophthora infestans in England and Wales.Plant Pathology,1986,35,578-581
[27].Deahl K.L,Goth R.W,Young R.Occurrence of the A2 mating type of Phytophthora infestans in potato fields in theUnited States and Canada.American Potato Journal,1991,68:717-725
[28].Mosa A A,Kobayashi K,Ogoshi A.Formation of oospores by Phytophthora infestans in potato tissues.Annuals of the Phytopathologial Society of Japan,1991,57:384-388
[29].Dagget S.S.,Gotz E.,Therrien C.D.Phenotypic change in populations of Phytophthora infestans from eastern Germany.Phytopathology,1993,83:319-323.
[30].Ludwik S.S.,Stephen B.,Goodwin S.B.Increased genotypic diversity via migration and possible occurrence of sexual reproduction of Phytophihora infestans in Poland.Phytopathologg,1994,84:201-207
[31].Hrestha S.K.First report of A2 mating type of Phytophthora infestans on potato and tomato in Nepal.Plant Disease,1998,82(9):1064.
[32].Sedegui M.,Carrol R.B.,Morehart A.L Genetic structure of the Phytophthora infestans population in Morocco.Plant disease,2000,84(2):173-176.
[33].Knapova G,Gisi U.Phenotypic and genotypic structure of Phytophthora infestans populationson potato and tomato in France,and Switzerland[J].Plant Pathology,2002,51,641^53.
[34].黄河,徐田玉,徐大雅,等.马铃薯对晚疫病水平抗性的研究.马铃薯.1983,1:6-15
[35].Flor H H.Current status of the gene-for-gene concept[J].Annu Rev Phytopathol,1971,9:275-296.
[36].Gurtler,H.Physiological races of Phytophthora infestans in Denmark and low temperature storage of isolates.Potato Research,1984,27(1):25-31.
[37].Tooley P.W,Therrien D.L.Mating type,race composition,nuclear DN Acontent,and isozyme analysis of Peruvian isolates of Phytophthora infestans.Phytopathology,1989,79(4):478-481.
[38].Andrivon,D.Races ofPhytophthora infestans in France,1991-1993.Potato Research,1994,100(2):97-107.
[39].Peters R.D.,Plat H.W Changes in race structure of Canadian populations of Phytophthora infestans based on specific virulence to selected potato clones.Potato Research,1998,41(4):366-370.
[40].Knapova G,Gisi U.Phenotypic and genotypic structure of Phytophthora infestans populations on potato and tomato in France and Switzerland.Plant Pathology,2002,51:641-653.
[41].Yigal Cohen.Populations of Phytophthora infestans in Israel underwent three major genetic changes during 1983 to 2000.Phytopathology,2002,92:300-307.
[42].黄河,程汉清,徐天宇,等.我国北部马铃薯晚疫病病原菌生理小种的发生和变化.植物病理学报,1981,11(1):45-49.
[43].刘晓鹏,谢从华,宋伯符.湖北恩施地区马铃薯晚疫病病原菌生理小种的组成及分布.马铃薯杂志,1995,9(2):81-83.
[44].杨艳丽,罗文富,国立耘.云南马铃薯晚疫病病原菌生理小种的研究.植物保护,2001,27(4):3-5.
[45].赵善欢编植物化学保护2003年7月第三版:147
[46].Davidse L C,Looijen D,Turkensteen L J,et al.Occurrence of metalaxyl-resistant strains of Phytophthora infestans in Dutch potato-fields.Netherlands Journal of Plant Pathology,1981,87(2):65-68
[47].ReisA,SmartC D,FryW E,et al.Characterization of isolates ofPhytophthora infestans from southern and southeastern Brazil from 1998 to 2000.Plant Disease,2003,87(8):896-900
[48].Dagget S S,Gotz E,Therrien C D.Phenotypic change in populations of Phytophthora infestans from eastern Germany[J].Phytopathology,1993,83:319323.
[49].Carter G A,Smith R M,Brent K J.Sensitivity to metalaxy of Phylophthora infestans populations in potato crops in south-west England in 1980 and 1981[J].Ann.Appl.Biol.1982,100:433-441.
[50].Pappas A C.Metalaxyl resistance in Phytophthora infestans on greenhouse tomatoes in Greece.Plant Pathology[J],1985,34:293296.
[51].Cohen Y,Reuveni M.Occurrence of metalaxyl resistant isolates of Phytophthora infestans in potato fields in Israel[J].Phytopathology,1983,73:925927.
[52].Kon Y J,Goodwin S B,Dyer A T,et al.Migrations and displacements of Phytophthora infestans populations in east Asian countries[J].Phytopathology,1994,84:922927.
[53].Matuszak J M,Femandez E J,Gu W K.,et al.Sensitivity of Phytophthora infestans populations to metalaxyl in Mexico:Distribution and dynamics[J].Plant Disease,1994,78:911-916.
[54].Deahl K L.Demuth S P,Pelter G,et al.First report of resistance of Phytophthora infestans to metalaxyl in eastern Washington and southwestern British Columbia[J].Plant Disease,1993,77:429.
[55].Deahl,K L,Inglis D A,et al.Testing for resistance to metalaxyl in Phytophthora infestans isolates from northwestem Washington[J].American Potato,1993,70:779-795
[56].Stephen B G,Ludwik S S,William E F,et al.Widespread distribution and probable origin of resistance to metalaxyl in clonal genotypes of Phytophthora infestans in the United States and western Canada[J].Phytopathology,1996,86(7):793-800.
[57].李炜,张志明,樊慕贞.马铃薯晚疫病病原菌对甲霜灵抗性的测定[[J].河北农业大学学报,1998,21(2):63-65
[58].王文桥,马志强,张小风,等.致病疫霉抗药性、交配型和适合度(J7.植物病理学报,2002,32(3):278^283.
[59].林孔勋主编.杀菌剂毒理学.北京.中国农业出版社,1995
[60].遗传学 王亚馥 戴灼华 高等教育出版社 186-190.
[61].邵铁梅,朱杰华,郑秀银.现代生物技术在马铃薯晚疫病病原菌(phytophthora infestans)研究中的应用.中国马铃薯,2002,16(2)87-91
[62].Tootey P W,Therrien D L.Mating type,race composition,nuclear DN Acontent,and isozyme analysis of Peruvian isolates of Phytophthora infestans[J]._Phytopathology,1989,79(4):478-481.
[63].Fry W E,Drenth A,Pielman L J,et al.Population genetic structure ofPhytophora infestans in the Netherlands[J].Phytopathology,1991,81:1330-1336.
[64].Mosa A A,Kobayashi K,Ogoshi A.Isoenzyme polymorphism and segregation in isolates Phvtophthora infestans from Japan[J].Plant Pathology,1993,42:26-34.
[65].Stephen B.,Goodwin S.B.Clonal diversity and genetic differentiation of Phytophthora infestans populations in Northern and Central Mexico.Phytopathology,1992,82(9):955-961.
[66].Drenth A.,Goodwin S.B.,Fry W.E.Genotypic diversity of Phytophthora infestans in the Netherlands revealed by DNA polymorphisms.Phytopathology,1993,83,1087-1092.
[67].Lionel L.,Didier A.French isolates of Phytophthora infestans from potato and tomato differ in phenotype and genotype.European Journal of Plant Pathology,1998,104,583-594.
[68].G.Mahuku,R.D.Peters,H.W.Plattff and F.Daayf.Random amplified polymorphic DNA (RAPD) analysis of Phytophthora infestans Isolate Collection in Canada during 1994 to 1996.Plant Pathology,2000,49:252-260.
[69].Zabeau M et al.European patent[P]0535 858A1,1993-03-31.
[70].Knapova G,Tenzer I,Gesser C.Characterisation of Phytophthora infestans from potato and tomato with molecular markers.Biodiversity in Plant Pathology.Proceedings of the 5th Congerss of the European Foundation for Plant Pathology,Taormina,Italy.2001,Pisa,Itay:SIPV,6-9.
[71].张志铭,李玉琴,田世民,等1中国发生马铃薯晚疫病病原菌(Phytophthora inf eslans)A2交配型[J].河北农业大学学报,1996,19(4):62-651
[72].朱杰华,张志铭,李玉琴.马铃薯晚疫病病原菌(Phytophthora infestans)A2交配型·的分布[J].植物病理学报,2000,30(4):3751
[73].Gurtler,H.Physiological races of Phytophthora infestans in Denmark and low temperature storage of isolates.Potato Research,1984,27(1):25-31.
[74].Andrivon,D.Races of Phytophthora infestans in France,1991-1993.Potato Research,1994,100(2):97-107.
[75].Peters R.D.,Plat H.W.Changes in race structure of Canadian populations of Phytophthora infestans based on specific virulence to selected potato clones.Potato Research,1998,41(4):366-370.
[76].Yigal Cohen.Populations of Phytophthora infestans in Israel underwent three major genetic changes during 1983 to 2000.Phytopathology,2002,92:300-307.
[77].Black W,Mastenbroek C,Mills W R,Peterson L.C.A proposal for an international nomenclature of races of Phytophthora infestans and of genes controlling immunity in Solanum demissum derivatives.Euphytica,1953,2:173-178
[78].胡诚.2003.马铃薯晚疫病水平抗病材料筛选.华中农业大学硕士学位论文.13-14
[79].Paul R J,Birch,Anna O,et al..Isolationb of Potato Gene That Are Induced During an early stage of the hypersensitive response to Phytophthora infestans[J].Mol.Plant-Microbe Inreact,1999,12:356-361.
[80].范娟.2005.马铃薯抗晚疫病资源筛选的研究.东北农业大学硕士学位论文.
[81].郭军.2005.马铃薯晚疫病病原菌遗传多样性及其小种特异无毒基因遗传定位和转录表达谱研究.中国农业科学院博士论文.
[82].孙巍.2003.海岛棉NBS-LRR类抗病基因同源序列的克隆与定位.中国农业科学院硕士研究生学位论文
[83].姚国胜.2006.中国部分地区马铃薯晚疫病病原菌遗传多样性研究.河北农业大学硕士学位论文.
[2].Reddick,D.Development of Blight Immune varieties.American Potato Journal,1943,20:118-126
[3].李克来,马铃薯晚疫病及抗晚疫病育种进展,晚疫病病原体的生物学,马铃薯科学1984,1:32-36
[4].Austin Bourke P.M.Eergence of potato blight,1843-46 Nature.1964,August,22,203:805-808
[5].林传光,黄河,王高才等.马铃薯晚疫病的田间动态观察及防治试验.植物病理学报.1955,1(1):31-43.
[6].农山渔村文化编李国光、姚方杰译新版蔬菜病虫害诊断原色图谱2001年1月第1版:171
[7].林传光,黄河,霍守祥.关于马铃薯晚疫病的预测和防治研究(摘要)北京农业大学学报,1956,2(1):79-85
[8].林传光,黄河,王道本,等.马铃薯晚疫病中心病株形成的观察.植物病理学报,1957,3(1):19-29
[9].朱杰华,张志铭,李玉琴,等.河北省马铃薯晚疫病病原菌(Phytophthora infestans)A2交配型的分布河北农业大学学报,2000b,23(3):73-75
[10].姚裕琪,巩秀峰,高奇华,等.马铃薯晚疫病抗性鉴定及评价.内蒙古农业科技,2001 C 2＞:8-9
[11].朱杰华,杨志辉,曹克强,等.影响活体内马铃薯晚疫病病原菌卵孢子产生因素的研究.菌物系统,2001,20(2):244-249
[12].Zwankhuizen M J,Govers F,Zadoks J C.Inoculum sources and genotypic diversity of Phytophthora infestans in Southern Flevoland,the Netherlands.European journal of Plant Pathology,2000,106:667-680
[13].郑小波1997疫霉菌及其研究技术中国农业出版社27-28
[14].Gallegly et al.,1958;Smoot et al.,1958;Spielman eral.,1991;Fry et aL,1993
[15].马铃薯晚疫病病原菌遗传多样性及其小种特异无毒基因遗传定位和转录表达谱研究
[16].Goodwin S B,Cohen B A,Deahl KL.Migration from Northern Mexico as the probable cause of recent genetic changes in populations of Phytophthora infestans in the United States and Canada[J].Phytopathology,1994,(84):553-5581
[17].Deahl K L,DeMuth S P,Sinden S L,et al.Identification of mating types and metalaxyl resistance in North American populations of Phytophthora infestans[J].American Potato Journal,1995,(72):35-491
[18].Goodwin S B,Sujkowski L S,Dyer A T,et al.Direct detection of gene flow and probable sexual reproduction of Phytophthora infestans in Northern North America[J]Phytopathology,1995,(85):473-4791
[19].Goodwin S B,Sujkowski L S,Fry W E.Widespread distributionand probable origin of resistance to metalaxyl in clonal genotypesof Phytophthora infestans in the United States and Western Canada[J].Phytopathology,1996,(86):793-8001
[20].Andrivon D Beasse C,Laurent C.Characterization of isolates of Phytophthora infestans collected in Northwestern France from1988 to 1992[J].Plant Pathology,1994,(43):471-4781
[21].Daggett S S,GEtz E,Therrien C D.Phenotypic changes in populations of Phytophthora infestans from Eastern Germany[J].Phytopathology,1993,(83):319-3231
[22].Sujkowski L S,Goodwin S B,Dyer A T,et al.Increased genotypic diversity via migration and possible occurrence of sexual reproduction of Phytophthora infestans in Ploland[J].Phytopathology,1994,(84):201-2071
[23].Forbes G A,Escobar X C,Ayala C C,et al.Populaton genetic structure of Phytophthora infestans in Ecuador[J].Phytopathology,1997,(87):375-3801
[24].Koh Y J,Goodwin S B,Dyer A T,et al.Migrations and displacements of Phytophthora infestans populations in East Asian countries[J].Phytopathology,1994,(84):922-9271
[25].Hohl ILK,Iselin K.Strains of Phytophthora infestans with A2 mating type behaviour.Transactions of the British Mycological Society,1984,83:529-530.
[26].Tantius P.M.,Fyfe AM.,Shaw D.S.Occurrence of the A2 mating type and self-fertile isolates of Phytophthora infestans in England and Wales.Plant Pathology,1986,35,578-581
[27].Deahl K.L,Goth R.W,Young R.Occurrence of the A2 mating type of Phytophthora infestans in potato fields in theUnited States and Canada.American Potato Journal,1991,68:717-725
[28].Mosa A A,Kobayashi K,Ogoshi A.Formation of oospores by Phytophthora infestans in potato tissues.Annuals of the Phytopathologial Society of Japan,1991,57:384-388
[29].Dagget S.S.,Gotz E.,Therrien C.D.Phenotypic change in populations of Phytophthora infestans from eastern Germany.Phytopathology,1993,83:319-323.
[30].Ludwik S.S.,Stephen B.,Goodwin S.B.Increased genotypic diversity via migration and possible occurrence of sexual reproduction of Phytophihora infestans in Poland.Phytopathologg,1994,84:201-207
[31].Hrestha S.K.First report of A2 mating type of Phytophthora infestans on potato and tomato in Nepal.Plant Disease,1998,82(9):1064.
[32].Sedegui M.,Carrol R.B.,Morehart A.L Genetic structure of the Phytophthora infestans population in Morocco.Plant disease,2000,84(2):173-176.
[33].Knapova G,Gisi U.Phenotypic and genotypic structure of Phytophthora infestans populationson potato and tomato in France,and Switzerland[J].Plant Pathology,2002,51,641^53.
[34].黄河,徐田玉,徐大雅,等.马铃薯对晚疫病水平抗性的研究.马铃薯.1983,1:6-15
[35].Flor H H.Current status of the gene-for-gene concept[J].Annu Rev Phytopathol,1971,9:275-296.
[36].Gurtler,H.Physiological races of Phytophthora infestans in Denmark and low temperature storage of isolates.Potato Research,1984,27(1):25-31.
[37].Tooley P.W,Therrien D.L.Mating type,race composition,nuclear DN Acontent,and isozyme analysis of Peruvian isolates of Phytophthora infestans.Phytopathology,1989,79(4):478-481.
[38].Andrivon,D.Races ofPhytophthora infestans in France,1991-1993.Potato Research,1994,100(2):97-107.
[39].Peters R.D.,Plat H.W Changes in race structure of Canadian populations of Phytophthora infestans based on specific virulence to selected potato clones.Potato Research,1998,41(4):366-370.
[40].Knapova G,Gisi U.Phenotypic and genotypic structure of Phytophthora infestans populations on potato and tomato in France and Switzerland.Plant Pathology,2002,51:641-653.
[41].Yigal Cohen.Populations of Phytophthora infestans in Israel underwent three major genetic changes during 1983 to 2000.Phytopathology,2002,92:300-307.
[42].黄河,程汉清,徐天宇,等.我国北部马铃薯晚疫病病原菌生理小种的发生和变化.植物病理学报,1981,11(1):45-49.
[43].刘晓鹏,谢从华,宋伯符.湖北恩施地区马铃薯晚疫病病原菌生理小种的组成及分布.马铃薯杂志,1995,9(2):81-83.
[44].杨艳丽,罗文富,国立耘.云南马铃薯晚疫病病原菌生理小种的研究.植物保护,2001,27(4):3-5.
[45].赵善欢编植物化学保护2003年7月第三版:147
[46].Davidse L C,Looijen D,Turkensteen L J,et al.Occurrence of metalaxyl-resistant strains of Phytophthora infestans in Dutch potato-fields.Netherlands Journal of Plant Pathology,1981,87(2):65-68
[47].ReisA,SmartC D,FryW E,et al.Characterization of isolates ofPhytophthora infestans from southern and southeastern Brazil from 1998 to 2000.Plant Disease,2003,87(8):896-900
[48].Dagget S S,Gotz E,Therrien C D.Phenotypic change in populations of Phytophthora infestans from eastern Germany[J].Phytopathology,1993,83:319323.
[49].Carter G A,Smith R M,Brent K J.Sensitivity to metalaxy of Phylophthora infestans populations in potato crops in south-west England in 1980 and 1981[J].Ann.Appl.Biol.1982,100:433-441.
[50].Pappas A C.Metalaxyl resistance in Phytophthora infestans on greenhouse tomatoes in Greece.Plant Pathology[J],1985,34:293296.
[51].Cohen Y,Reuveni M.Occurrence of metalaxyl resistant isolates of Phytophthora infestans in potato fields in Israel[J].Phytopathology,1983,73:925927.
[52].Kon Y J,Goodwin S B,Dyer A T,et al.Migrations and displacements of Phytophthora infestans populations in east Asian countries[J].Phytopathology,1994,84:922927.
[53].Matuszak J M,Femandez E J,Gu W K.,et al.Sensitivity of Phytophthora infestans populations to metalaxyl in Mexico:Distribution and dynamics[J].Plant Disease,1994,78:911-916.
[54].Deahl K L.Demuth S P,Pelter G,et al.First report of resistance of Phytophthora infestans to metalaxyl in eastern Washington and southwestern British Columbia[J].Plant Disease,1993,77:429.
[55].Deahl,K L,Inglis D A,et al.Testing for resistance to metalaxyl in Phytophthora infestans isolates from northwestem Washington[J].American Potato,1993,70:779-795
[56].Stephen B G,Ludwik S S,William E F,et al.Widespread distribution and probable origin of resistance to metalaxyl in clonal genotypes of Phytophthora infestans in the United States and western Canada[J].Phytopathology,1996,86(7):793-800.
[57].李炜,张志明,樊慕贞.马铃薯晚疫病病原菌对甲霜灵抗性的测定[[J].河北农业大学学报,1998,21(2):63-65
[58].王文桥,马志强,张小风,等.致病疫霉抗药性、交配型和适合度(J7.植物病理学报,2002,32(3):278^283.
[59].林孔勋主编.杀菌剂毒理学.北京.中国农业出版社,1995
[60].遗传学 王亚馥 戴灼华 高等教育出版社 186-190.
[61].邵铁梅,朱杰华,郑秀银.现代生物技术在马铃薯晚疫病病原菌(phytophthora infestans)研究中的应用.中国马铃薯,2002,16(2)87-91
[62].Tootey P W,Therrien D L.Mating type,race composition,nuclear DN Acontent,and isozyme analysis of Peruvian isolates of Phytophthora infestans[J]._Phytopathology,1989,79(4):478-481.
[63].Fry W E,Drenth A,Pielman L J,et al.Population genetic structure ofPhytophora infestans in the Netherlands[J].Phytopathology,1991,81:1330-1336.
[64].Mosa A A,Kobayashi K,Ogoshi A.Isoenzyme polymorphism and segregation in isolates Phvtophthora infestans from Japan[J].Plant Pathology,1993,42:26-34.
[65].Stephen B.,Goodwin S.B.Clonal diversity and genetic differentiation of Phytophthora infestans populations in Northern and Central Mexico.Phytopathology,1992,82(9):955-961.
[66].Drenth A.,Goodwin S.B.,Fry W.E.Genotypic diversity of Phytophthora infestans in the Netherlands revealed by DNA polymorphisms.Phytopathology,1993,83,1087-1092.
[67].Lionel L.,Didier A.French isolates of Phytophthora infestans from potato and tomato differ in phenotype and genotype.European Journal of Plant Pathology,1998,104,583-594.
[68].G.Mahuku,R.D.Peters,H.W.Plattff and F.Daayf.Random amplified polymorphic DNA (RAPD) analysis of Phytophthora infestans Isolate Collection in Canada during 1994 to 1996.Plant Pathology,2000,49:252-260.
[69].Zabeau M et al.European patent[P]0535 858A1,1993-03-31.
[70].Knapova G,Tenzer I,Gesser C.Characterisation of Phytophthora infestans from potato and tomato with molecular markers.Biodiversity in Plant Pathology.Proceedings of the 5th Congerss of the European Foundation for Plant Pathology,Taormina,Italy.2001,Pisa,Itay:SIPV,6-9.
[71].张志铭,李玉琴,田世民,等1中国发生马铃薯晚疫病病原菌(Phytophthora inf eslans)A2交配型[J].河北农业大学学报,1996,19(4):62-651
[72].朱杰华,张志铭,李玉琴.马铃薯晚疫病病原菌(Phytophthora infestans)A2交配型·的分布[J].植物病理学报,2000,30(4):3751
[73].Gurtler,H.Physiological races of Phytophthora infestans in Denmark and low temperature storage of isolates.Potato Research,1984,27(1):25-31.
[74].Andrivon,D.Races of Phytophthora infestans in France,1991-1993.Potato Research,1994,100(2):97-107.
[75].Peters R.D.,Plat H.W.Changes in race structure of Canadian populations of Phytophthora infestans based on specific virulence to selected potato clones.Potato Research,1998,41(4):366-370.
[76].Yigal Cohen.Populations of Phytophthora infestans in Israel underwent three major genetic changes during 1983 to 2000.Phytopathology,2002,92:300-307.
[77].Black W,Mastenbroek C,Mills W R,Peterson L.C.A proposal for an international nomenclature of races of Phytophthora infestans and of genes controlling immunity in Solanum demissum derivatives.Euphytica,1953,2:173-178
[78].胡诚.2003.马铃薯晚疫病水平抗病材料筛选.华中农业大学硕士学位论文.13-14
[79].Paul R J,Birch,Anna O,et al..Isolationb of Potato Gene That Are Induced During an early stage of the hypersensitive response to Phytophthora infestans[J].Mol.Plant-Microbe Inreact,1999,12:356-361.
[80].范娟.2005.马铃薯抗晚疫病资源筛选的研究.东北农业大学硕士学位论文.
[81].郭军.2005.马铃薯晚疫病病原菌遗传多样性及其小种特异无毒基因遗传定位和转录表达谱研究.中国农业科学院博士论文.
[82].孙巍.2003.海岛棉NBS-LRR类抗病基因同源序列的克隆与定位.中国农业科学院硕士研究生学位论文
[83].姚国胜.2006.中国部分地区马铃薯晚疫病病原菌遗传多样性研究.河北农业大学硕士学位论文.