摘要
致病疫霉(Phytophthora infestans (Mont.) de Bary)引起的马铃薯晚疫病在全球许多马铃薯主产国家发生和流行。晚疫病菌群体结构及其演体规律的变化直接影响着该病害的发生和流行,有关病原菌群体结构的研究再次成为人们关注的焦点。通过对2006-07年采自青海省马铃薯主产区的晚疫病样及分离纯化菌株的表型(交配型、毒性基因、生理小种和抗药性)进行研究,系统地揭示了青海马铃薯主产区晚疫病菌群体的遗传多样性;为马铃薯抗晚疫病育种和晚疫病综合防治技术制订提供了理论依据。主要研究结果如下:
1. 2006年至2007年,采集分离到致病疫霉70株,其中66株采自马铃薯,4株采自番茄。
2.交配型测定和自发卵孢子形成检测,70株纯化菌株均为A1交配型,未发现A2交配型或自育型菌株。
3.利用一套分别携带R1-R11抗性基因标准鉴别寄主,采用人工接种鉴定方法,对分离菌株中的54个进行了生理小种鉴定.,发现青海马铃薯主产区存在可鉴定的晚疫病菌毒性基因谱含有1, 2, 3, 4, 5, 6, 7, 8和10号毒性基因。各地晚疫病菌生理小种组成及其毒性基因构成有差异。经鉴定54株分属于16种不同的生理小种,其中,出现频率最高的小种是3.4.10,发生频率为33.3%.
4.菌株甲霜灵敏感性测定结果:高度抗性、中度抗性和敏感菌株的数量分别为4个、2个和64个,分别占被测菌株总数的5.71%, 2.85%和91.43%;其中中抗菌株分布在互助县境内,高抗菌株均分离自番茄,分布在循化县和西宁城北区。
研究结果表明,青海马铃薯晚疫病菌群体的表型结构具有复杂的多样性,同时也表现出明显的群体结构特点:以A1交配型和复合生理小种为主体,但是对甲霜灵的敏感性处于较高水平。
Late blight caused by the oomycete Phytophthora infestans (Mont.) de Bary occurred and had a rapidly epidemics in many potato countries all over the world. The occurrence and epidemics of this disease are dominated by the stucture and variation of P.infestans population, so the concerning researches on late blight has been focusing on the population stucture of the pathogen again in recent years. The present paper systematically revealed the genetic diversity of P.infestans on potato in Qinghai province, China, by analyzing the phenotypic stucture of local P.infestans population of the major potato production areas from 2006-2007 Qinghai province. The ralative results derived from the work are of significance to the breeding for resistance to late blight, and the management of the disease.Main results were summarized as follows:
1. The 70 isolates of Phytophthora infestans were collected Qinghai Province from 2006 to 2007, four of them whose host were tomatoes, the others were potatoes.
2. The isolates were inspected for mating type and monitoyed the oospores which were producted by self-fertility isolates. The results showed that 70 isolates tested were all subjected to A1 mating,no A2 mating type or relf-fertility isolate was found.
3. 57 isolates out of all collected isolates were tested with 11 monogene R1～R11 potato clones of potato late blight differential host . Except Vir-9 or Vir-11, 9 of 11Virulence genes (Vir1, Vir2, Vir3, Vir4, Vir5, Vir6, Vir7, Vir8, Vir10) had been identified from these collections. The physiological race of P. infestans in Qinghai Province had become much complex. The results showed that there were 16 races in Qinghai Province and the dominant races were race 3.4.10 with frequency of 33.3%.
4. All 70 isolates were tested to sensitivity to Metalaxyl, 64 isolates (91.43% )were sensitive, 2 isolates (2.85% )were intermediate resistance, and 4 isolates(5.71%)were highly resistance. The intermediate resistance isolates were distributed in Huangyuan county, All the highly resistance isolates collected from tomatoes were distribution of Xunhua county and the northland of Xining.
The evidences from above results reflected that the phenotypic structure of P. infestans population in Qinghai shared complex diversity, but also showed some typiccharacteristics including that A1 mating type and complex races were the dominated components of phenotypic structure, most of them were sensitive to Metalaxyl.
引文
[1]Bourke P M. A.Emergence of potato late blight, 1843-46[J]. Nature, 1964, 203: 805-808.
[2]Fry W E, Goodwin S B, Dyer A T, et al. Historical and recent migrations of P.infestans: Chronology, Pathways and implication[J]. Plant Disease, 1993,77:653-661.
[3]李汝刚,伍宁丰,范云六,等.马铃薯抗晚疫病研究进展[J].马铃薯杂志.1997,(4):243-250.
[4]Duncan JM. Phytophthora-anabiding treat to our crops[J]. Microbiology Today, 1999, 26:114-116.
[5]Fry W E, Goodwin S B. Re-emergence of potato and tomato late blight in the United States[J]. P1ant Disease, 1997, 81(12): 1349-1357.
[6]Song J J, Bradeen M, Naess S K,et al.Gene RB clone from Solanum bulbocastanum confers broadspectrum rssistance to potato lat blight[J].Proc.Natl. Acad.Sci.2003,100(16):9128-9133.
[7]Goodwin S B, Sujkowski L S. Direct detection of gene flow and probable sexual reproduction of Phytophthora infestans in northern North America[J]. Phytopathology, 1995, 85: 473-479.
[8]Dick M W. The Straminipillous fungi.A new classification for the biflagellate fungi and their uniflagellate relatives with particular reference to Lagenidiaceous fungi. C.A.B[J]. Internat. Mycol. Pap. 1995, No.168.
[9]Bartnicki-Garcia S, Wang M C.Biochemical aspects of morphogenesis in Phytophthora.In.Erwin D C, Bartnicki-Garcia S,Tsao P H.eds.Phytophthora:its biology taxonomy,ecology a-nd pathology[J].St Paul:Am phyto-path Soc pp121-137.
[10]Vogel H J. Distribution of lysine pathways among fungi:evolutionary implications[J]. Am. Nat., 1964, 98: 435-446.
[11]Pfyffer G, Boraschi-Gaia E, Weber B. A further report on the occurrence of acylic sugar alcohols in fungi[J]. Mycol. Res.,1990,92,219-222.
[12]Hemmes D.E. Cytology of Phytophthora. In: Phytophthora: Its Biology, Taxonomy, Ecology, and Pathology(D C Erwin,S Bartnicki-Garcia and P H Tsao.Eds.). American Phytopathological Society Press, St.Paul, MN. 1983, pp. 9-40.
[13]Hendrix J.M. Sterols in growth and reproduction of fungi[J]. Annu. Rev. Phytopathol., 1970, 8:111-130.
[14]Eliiott C G.Physiology of sexual reproduction in Phytophthora. In: Phytophthora: lts biology, Taxonomy, Ecology, and Pathology (D.C. Erwin, S. Bartnicki-Garcia, and P.H.Tsao.Eds.)[M], American Phytopathological Society Press, St.Paul, MN., 1983, pp,71-80.
[15]Erwin D C, Ribeiro, O.K. Introduction to the genus Phytophthora.In: Phytophthora Diseases Worldwide[M]. American Phytopathological Society Press, St. Paul, MN., 1996, pp. 1-7.
[16]Layton A C,Kuhn D N In plants formation of heterokaryons of Phytophthora megasperma f.sp.glycinea. Phytopathology, 1990,80:602-606.
[17]Judelson H S The genetics and biology of Phytophthora infestans: modern approaches to a historical challenge[J]. Fungal Genet Biol.,1997,21:188-197.
[18]Judelson HS,Yang G. Recombination pathways in Phytophthora infestans: polyploidy resulting from aberrant sexual development and zoospore-mediated heterokaryosis[J]. Mycol.Res.,1998,102:1245-1253.
[19]van West P, Kamoun S, van't Klooster J W, et al. Intemuclear gene silencing in Phytophthara infestans[J]. Mol. Cell, 1999, 3:339-348.
[20]Robertson N.R The challenge of Phytophthora infestans. In: Ingram D S and Williams P H (Ed), Advances in Plant Pathology: Phytophthora infestans, the cause of late blight of potato[M]. 1991.7:1-30.
[21]Turkenatten L J. Durable resistance of potato against Phytophthora infestans. 1n: Jacobs T.H. and Parlevliet J.E. (eds), Durability of disease resistance[M]. Kluwer Academic Publishers, Dordrecht, Boston, London, The Netherlands. 1993, 115-124.
[22]Drenth A, Janssen E M, GoversF. Formation and survival of oospores of Phytophthora infestans under natural conditions[J]. Plant pathology, 1995, 44:86-94.
[23]孙慧生. 马铃薯育种学[M]. 北京: 中国农业出版社, 2003.
[24]Gurtler H. Physiological races of Phytophthora infestans in Denmark and low temperature storage of isolates[J]. Potato Research, 1984,27(1):25-31.
[25]Tooley P W, Therrien D L. Mating type, race composition, nuclear DNA content, and isozyme analysis of Peruvian isolates of Phytophthora infestans[J]. Phytopathology, 1989,79(4):478-481.
[26]Andrivon, D. Races of Phytophthora infestans in France,1991-1993[J]. Potato Research,1994,100(2):97-107.
[27]Peters R D, Platt H W. Changes in race structure of Canadian populations of Phytophthora infestans based on specific virulence to selected potato clones[J]. Potato Research, 1998,41(4):366-370.
[28]Knapova G., Gisi U. Phenotypic and genotypic structure of Phytophthora infestans populations on potato and tomato in France and Switzerland[J]. Plant Pathology, 2002, 51:641-653.
[29]Yigal Cohen. Populations of Phytophthora infestans in Israel underwent three major genetic changes during 1983 to 2000[J]. Phytopathology, 2002,92(3):300-307.
[30]Spielman, L.J.,Drenth A.,Davidse L.C.A second world-wide migration and population displacement of Phytophthora infestans[J].PlantPathology,1991,40:422-430.
[31]Niederhauser J.S. The blight, the blighter, and the blighted[J]. Transactions of the New York Academy of Science.1956,19:55-63.
[32]Hohl,H.R.,Iselin, K. Strains of Phytophthora infestans with A2 mating type behaviour[J].Transactions of the British Mycological Society, 1984, 83: 529-530.
[33]Tantius PM., Fyfe A.M., Sbaw, D.S. Occurrence of the A2 mating type and self-fertile isolates of Phytophthora infestans in England and Wales[J]. Plant Pathology, 1986,35,578-581.
[34]Frisking H D,Davidse L C,Limburg,H.Oospore formation by Phytophthora infestans in host tissue after inoculation with isolates of opposite mating type found in the Netherlands[J]. Netherlands Journal of Plant Pathology, 1987, 93:147-149.
[35]Deahl K L, Goth R W, Young R. Occurrence of the A2 mating type of Phytophthora infestans in potato fields in the United States and Canada[J]. American Potato Journal,1991,68:717-725.
[36]Mosa A A, Kobayashi K, Ogoshi A. Formation of oospores by Phytophthora infestans in potato tissues[J]. Annuals of the Phytopathologial Society of Japan, 1991,57:384-388.
[37]Dagget S S, Gotz E,Therrien C D. Phenotypic change in populations of Phytophthora infestans from eastern Germany[J]. Phytopathology, 1993,83:319-323.
[38]Ludwik S S, Stephen B, Goodwin S B. Increased genotypic diversity via migration and possible occurrence of sexual reproduction of Phytophthora infestans in Poland [J].Phytopathology,1994,84:20l-207.
[39]Barony J, Ersek T. First report of A2 mating type of Phytophthora infestans on potato inHungary[J]. Plant Disease, 1997,81:1094.
[40]Hrestha S K. First report of A2 mating type of Phytophthora infestans on potato and tomato in Nepal[J]. Plant Disease, 1998,82(9):1064.
[41]Sedegui M, Carroll R B, Morehart A L. Genetic structure of the Phytophthora infestans population in Morocco[J]. Plant disease, 2000,84(2):173-176.
[42]Day J P, Hardman N,Shattock R C. Diversity of Phytophthora infestans in the United Kingdom 1996-98: phenotypic and molecular variation in small samples from many sites reveal clonal lineages and related non-clonal variants[C]. Biodiversity in Plant Pathology. Proceedings of the 5th Congress of the European Foundation for Plant Pathology, Taormina, Italy. 2001, Pisa, Italy: SIPV,1-5.
[43]Hermansen A,Hannukkala A,Hafskjold N R. Variation in populations of Phytophthora infestans in Finland and Norway: mating type, metalaxyl resistance and virulence phenotype[J]. Plant Pathology, 2000,49,11-22.
[44]Turkensteen L J, Flier W Q, Wanningen R. Production, survival and infectivity of oospores of Phytophthora infestans[J]. Plant Pathology, 2000,49:688-696.
[45]Davidse L C,Hooijen D,Turkensteen L J.Occurrence of metalaxyl-resistant strains of Phytophthora infestans in Dutch potato-fields[J]. Plant Pathology,1981,87:65-68.
[46]Dowley J, O'Sullivan E. Metalaxyl-resistance in populations of Phytophthora infestans (Mont) De bary in Ireland[J]. Potato Research, 1981,24:417-421.
[47]Garter 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.
[48]Pappas A C. Metalaxyl resistance in Phytophthora infestans on greenhouse tomatoes in Greece[J]. Plant Pathology, 1985,34:293-296.
[49]Cohen Y, Reuveni M. Occurrence of metalaxyl resistant isolates of Phytophthora infestans in potato fields in Israel[J]. Phytopathology,1983,73:925-927.
[50]Kon Y J, Goodwin S B, Dyer A T. Migrations and displacements of Phytophthora infestans populations in east Asian countries[J]. Phytopathology, 1994, 84:922-927.
[51]Matuszak J M, Femandez E J, Gu W K. Sensitivity of Phytophthora infestans populations to metalaxyl in Mexico: Distribution and dynamics[J]. Plant Disease, 1994,78:911-916.
[52]Deahl K L, Demuth S P, Pelter Q. First report of resistance of Phytophthora infestans to metalaxyl in eastern Washington and southwestern British Columbia[J]. Plant Disease,1993a,77:429.
[53]Deahl K.L, lnglis D A, DeMuth S P. Testing for resistance to metalaxyl in Phytophthora infestans isolates from northwestern Washington[J]. American Potato Journal,1993b,70:779-795
[54]Stephen B G , Ludwik S S, William EF. 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.
[55]Kadish D,Cohen Y. Overseasoning of metalaxyl-sensitive and metalaxyl-resistance isolates of Phytophthora infestans in potato tubers[J]. Phytopathology, 1992,82:887-889.
[56]Shattock R C.Studies on the inheritance of resistance to metalaxyl in Phytophthora infestans[J].Plant Pathology, 1988, 37: 4-11.
[57]Marshall-Farrar K D, Mcgrath M, Jame R V.Characterization of Phytophthora infestans in Wisconsin from 1993-1995[J].Plant Disease,1998,82(4):434-436.
[58]Cooke D E L, Young V, Birch P R L. Phenotypic and genotypic diversity of Phytophthora infestans populations in Scotland (1995-97) [J]. Plant Pathology, 2003,52,181-192.
[59]Ailton R, Christine D S, Fry W E. Characterization of isolates of Phytophthora infestans from southern and southeastern Brazil from 1998 to 2000[J]. Plant Disease,2003,87:96-100.
[60]Kadish D, Cohen Y.Estimation of metalaxyl sensitive and resistant isolates of Phytophthora infestans in the absence of metalaxyl[J].Plant Pathology,1988, 37:558-564.
[61]Lee T Y, Fry W E. Genetical analysis of matalaxyl resistance in Phytophthora infestan- s[J].Phytopathology, 1997, 87: 54-57.
[62]Fabritius A L, Judelson H S. Genetic analysis of metalaxyl insensitivity loci in Phytophthora infestans using linked DNA marker[J].Phytopathology,1997, 87:1034-1040.
[63]Judelson H S,Roberts S. Multiple loci determining insensitivity to phenylamide fungicides in Phytophthora infestans[J]. Phytopathology, 1999,89:754-760.
[64]Tooley P W,Therrin R C.Variation in ploidy in Phytophthora infestans[C].1991.In:Lucas J A,R C Shattock, D S Shaw et al.eds.phytophthora.Symposium ofthe British Mycological Society,the British Society for Plant Pathology and the Society of Irish Pathologists held at Trinity College ,Dublin,September,1989.Cambridge:Cambridge University Press.204-217.
[65]Whittake S L,Shattock R C,Shaw D S.Inheritance of DNA comtents in sexual progebies of Phytophthora infesans[J].Mycological Research,1991,(9):1094-1100.
[66]Judelson H S. Chromosomal heteromorphism linked to the mating type locus of oomycete, Phytophthora infestans[J]. Molecular General Genetics ,1996,252:155-161.
[67]Fobers G A, Goodwin S B,Drenth A.et al. A global marker database for Phytophthora infestans[J]. Plant Disease,1998,82:811-818.
[68]Tooley P W, Fry W E, Viliarreal M J. Isozyme characterization of sexual and asexual Phytophthora infestans populations[J]. The Journal of Heredity, 1985,76:431-435.
[69]Tooley P W, Hewings A D, Falkenstein KF. Detection of double-stranded RNA in Phytophthora infestans[J]. Phytopathology,1989,79:470-474.
[70]Fry W E, Drenth A, Pielman L J. Population genetic structure of Phytophthora infestans in the Netherlands[J]. Phytopathology, 1991,81:1330-1336.
[71]Mosa A A,Kobayashi K,Ogoshi A. Isoenzyme polymorphism and segregation in isolates Phytophthora infestans from Japan[J]. Plant Pathology, 1993,42:26-34.
[72]Stephen B,Goodwin S B,Clonal diversity and genetic differentiation of Phytophthora infestans populations in Northern and Central Mexico[J]. Phytopathology,1992,82(9):955-961.
[73]Drenth A, Goodwin S B, Fry W E. Genotypic diversity of Phytophthora infestans in the Netherlands revealed by DNA polymorphisms[J]. Phytopathology,1993,83:1087-1092.
[74]Lionel L, Didier A. French isolates of Phytophthora infestans from potato and tomato differ in phenotype and genotype[J]. European Journal of Plant Pathology, 1998,104:583-594.
[75]Wangsomboondee T, Trout Grovers C, Shoemaker M A,et al. Phytophthora infestans populations from tomato and potato in North Carolina differ in genetic diversity and structure[J]. Phytopathology,2002,92(11):1189-1195.
[76]Griffith G W,Shaw D S. Polymorphism in Phytonhthora infesrans: four mitochondrial haplotypes are detected after PCR amplification of DNA from pure cultures or from host lesions[J]. Applied and Environmental Microbiology, 1998, 64:4007-4014.
[77]Wattier R A M,Gathercole L L,Assinder S J.et al.Sequence variation of intergenic mitochondrial DNA spacers (mt-DNA-IGS) of Phytophthora infestans(Oomycets) and related species[J].Mol Ecol Nates.2003,3:136-138.
[78]Mahuku G, Peters R D, Platt H W. Random amplified polymorphic DNA (RAPD) analysis ofPhyrophthora infestans isolates collected in Canada during 1994 to 1996[J].Plant Pathology,2000,49:252-260.
[79]van der Lee T,De Witte L,Drenth A, et al. AFLP linkage map of the oomycete Phytophthora infestans[J]. Fungal genetics and Biology,1997,21:278-291.
[80]Flier W G,Grunwald N J,Kroon L P N M,et al. The population structure of Phytophthora infestans from the Toluca valley of Central Mexico suggests genetic differentiation between populations from cultivated potato and wild Solanum spp[J].Phytopathology,2003,93:382-390.
[81]Knapova G,Tenzer I, Gessler C. Characterisation of Phytophthora infestans from potato and tomato with molecular markers. Biodiversity in Plant Pathology[C]. Proceedings of the 5th Congerss of the European Foundation for Plant Pathology, Taormina, Italy.2001, Pisa,Italy:SIPV, 6-9.
[82]Wastie R L.Breeding for resistance. In: Advances in Plant Pathology, Vol. 7: Phytophthora infestans, the Cause of Late Blight of Potato (D.S. Ingram and P.H. Williams, eds)[M].Academic Press London,UK 1991, pp 193-223.
[83]Flor H H. Current status of the gene-for-gene concept[J]. Annual Review of Phytopathology,1971,9:275-269.
[84]Kamoun S, Honee C X, Weide R. The fungal gene Avr9 and the oomycete gene Infl confer avirulence to potato virus X on tobacco[J]. Molecular Plant-Microbe Interaction, 1999a,12:459-462.
[85]El-Knarbotly A, Leonards-Schippers C, Huigen D J,et al. Segregation analysis and RFLP mapping of R1 and R3 alleles conferring race-specific resistance to Phytophthora infestans in progeny of dihaploid potato parents[J]. Mol. Gen. Genet,1994,242:749-754.
[86]Leonards-Schippers C, Gieffers W, Salamini E, et al. The R1 gene conferring reca-specific resistance to Phytophthora infestans in potato is located on chromosome V[J]. Mol. Gen.Genet,1992,233:278-283.
[87]Li X., Eck H J, Rouppe van der Voort J N A M, et al. Autotetraploids and genetic ampping using common AFLP markers: the R2 allele conferring resistance to Phytophthora infestans mapped on potato chromosome Ⅳ [J]. TAG, 1998,96:1121-1128.
[88]El-Knarbotly A, Palomino Sanchez C, Salamini,F, et al. R6 and R7 allels of potato conferring race-specific resistance to Phytophthora infestans identified genetic loci clustering with theR3 locus on chromosome Xl[J]. Theor, Appl.Genet,1996,92:880-884.
[89]Huang S. Discovery and characterization of the major late blight resistance complex in potato[D].Wageningen University PHD Thesis. 2005
[90]Ballvora A, Ercolano M R, WeiB J,et al.The R1 gene for potato resistance to late blight (Phytophthora infestans) belongs to the leucinezipper/NBS/LRR class of plant resistance genes[J]. The Plant Journal.,2002, 30:361-371.
[91]van der Vossen E, Sikkema A, Hekkert B T L Positional cloning of a resistance gene from Solanum bulbocastanum conferring race non-specific resistance to Phytophthora infestans[C]. Book of Abstracts G1LB' 02 conference Late blight: managing the global treat, 2002, Abstract 23B,pp38.
[92]Al-Kherb S M, Fininsa C, Shattock R C,et al. The inheritance of virulence of Phyotphthora infestans to potato[J]. Plant Pahol.,1995,44:552-562.
[93]Spielman L J, McMaster,B J,et al. Dominance and recessiveness at loci for virulence against potato and tomato in Phytophthora infestans[J]. Theor.Appl.Genet.,1989,77:832-838.
[94]van der Lee T,Robold A, Testa A, et al. Mapping of avirulence genes in Phytophthora infestans with AFLP markers selected by bulked segregant analysis[J]. Genetics, 2001, 157:949-956.
[95]Whisson S C, Drenth A., Maclean D J,et al. Phytophthora sojae avirulence genes, RAPD, and RFLP markers used to construct a detailed genetic linkage map[J]. Mol. Plant-Microbe Interact, 1995, 8:988-995.
[96]Whisson S C, van der Lee T, Bryan C J,et al. Physical mapping across an avirulence locus of Phytophthora infestans using a high representation,large insert artificial chromosome library[J]. Mol. Genet. Genomics, 2001, 266: 289-295.
[97]Birch ,Kamoun,.Studing interaction transcripion:coordinated analyses of gene expression during plant-microorganism interactions[C] .In:Wood Rceds New technologies for life science : a trend guide Elsevier Science . NewYork. 2000,pp77-82.
[98]Pieterse C M J, Risseeuw E P and Davidse L C. An in planta induced gene of Phytophthora infestans codes for ubiquitin[J]. Plant Mol. Biol.,1991,17:799-811.
[99]Pieterse C M J, Verbakel H M, Spaans J H, et al. Increased expression of the calmodulin gene of the late blight fungus Phytophthora infestans during pathogenesis on potato[J]. Mol.Plant-Microbe Interact., 1993, 6:764-172.
[100]Pieterse C J M.,Derksen A M C E ,Folders J,et al.Expression fo the Phytophthora infestans ipiO and ipiO genes in planta and in vitro[J].Mol.Gem.Genet.1994,244:269-277.
[101]van West P, Kamoun S,de Groot K, et al. Antisense and sense mediated gene silencing as a tool to suppress production of INF1 elicitin of Phytophthora infestans. Pacific Grove CA. 1997[C]. Abstract presented at the 19th Fungal Genetics Conference.
[102]Gornhaldt B, Rouhara I and Schmelzer E. Cyst germination proteins of the potato pathogen Phytnphthora infesrans share homology with human mucins[J]. Mol. Plant-Microbe Interact.,2000,13:32-42.
[103]Dellagi A, Heilbronn J, Avora A O,et al. A potato gene encoding a WRKY-like transcription factor is induced in interactions with Erwinia carotovora subsp.Atroseptica and Phytophthora infestans and is co-regulated with classI endochitinase expression[J]. Mol.Plant-Microbe Interact.2000,13:1092-1101.
[104]Avrora A O, Stewart H E, De Jong, W, et al. A cysteine protease gene is expressed early in resistant potato interactions with Phyotphthora infestans[J]. Mol.Plant-Microbe Interact.,1999,12:1114-1119.
[105]Birch P R J, Avora A O, Lyon G D., et al. Isolation of potato genes thatare induced during an early stage of the hypersensitive response to Phyotphthora infestans[J]. Mol.Plant-Microbe Interact. 1999,12:356-361.
[106]Avrova A O, Venter E, Birch P R J, et al,. Profiling and quantifying differential gene transcription in Phytophthora infestans prior to and during the early stages of potato infection[J].Fungual genetics and biology, 2003,40:4-14.
[107]Kamoun S, Hraber P, Sobral B,et al. Initial assessment of gene diversity for the oomycete Phytophthora infestans based on expressed sequences[J]. Fungal Genet. Biol.,1999b,28:94-106.
[108]Qutob D, Hraber P T, Sobral B W, et al. Comparative analysis of expressed sequences in Phytophthora sojae[J]. Plant Physiol.,2000,123:243-254.
[109]Waugh M., Hraber P, Weller J, et al. The Phytophthora Genome Initiative database: informatics and analysis for distributed pathogenomic research[J]. Nucl. Acids Res.,2000,28:87-90.
[110]Torto T, Styer A and Kamoun S. EST data mining: navel extracellular proteins from theoomycete plant pathogen Phytophthora infestans[J]. Fungal Genet. Newsle.,2001,48.(Suppl.):111.
[111]黄河,程汉清,徐天宇,等.我国北部马铃薯晚疫病菌生理小种的发生和变化[J]. 植物病理学报. 1981, 11(1): 45-49.
[112]张志铭,李玉琴,田世民.中国发现马铃薯晚疫病菌(Phytophthora infestans)A2交配型[J].河北农业大学学报,1996,16(4):63-65.
[113]赵志坚,何云昆,李成云,等.云南省发现马铃薯晚疫病菌(Phytophthora infestans) A2交配型.西南农业学报,1999,12(3):1-3.
[114]朱杰华,张志铭,李玉琴.马铃薯晚疫病菌(Phytophthora infestans)A2交配型的分布[J].植物病理学报,2000,30(4):375.
[115]李纬,张志铭,樊募贞.马铃薯晚疫病菌对瑞毒霉抗性的测定[J].河北农业大学学报, 1998,97:63-65.
[116]王文桥,马志强,张小风.致病疫霉抗药性、交配型和适合度[J].植物病理学报,2002,32(3):278-283.
[117]毕朝位.重庆马铃薯晚疫病菌(Phytophthora infestans)对甲霜灵抗性几抗性水平测定[J].中国马铃薯,2002,24(4):81-83.
[118]刘晓鹏,谢丛华,宋伯符.湖北恩施地区马铃薯晚疫病病菌生理小种的组成及分布[J].马铃薯杂志,1995,9(2):81-83.
[119]杨艳丽,罗文富,国立耘.云南省马铃薯晚疫病菌生理小种的研究[J].植物保护,2001, 27(4):3-5.
[120]金光辉.黑龙江省马铃薯晚疫病菌生理小种的类型分布状况研究[J].中国马铃薯,2003,4:213-215.
[121]朱杰华,杨志辉,邵铁梅,等.中国部分地区马铃薯晚疫病菌生理小种的组成及分布[J].中国农业科学,2003,36(2):169-172.
[122]杨素祥,李灿辉,郝大海,等.云南省德宏州冬作马铃薯晚疫病菌表型特征研究[J].植物病理学报,2007,37(1):18-24.
[123]杨艳丽,胡先奇,鲁绍凤,等.云南省马铃薯晚疫病菌生理小种的组成与分布[J].华中农业大学学报,2007,26(3):297-301.
[124]赵志坚,曹继芬,李灿辉,等.云南致病疫霉交配型、甲霜灵敏感性、mtDNA 单倍型及其群体演替研究[J].中国农业科学,2007,40(4):727-734.
[125]郭军,屈冬玉,巩秀峰,等.内蒙古马铃薯晚疫病菌基因型多样性分析[J]。西北农赫科技大学学报(自然科学版)2007,35(11):120-124.
[126]朱杰华.中国马铃薯晚疫病菌群体遗传结构研究[D].河北农业大学博士论文.2004.
[127]郭军.马铃薯晚疫病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[D].中国农业科学院博士论文,2005.
[128]王军,宋伯符.中国马铃薯学术研讨文集[C].哈尔滨:黑龙江科学技术出版社,1996.
[129]赵志坚,王淑芬,李成云,等.云南省马铃薯晚疫病菌交配型分布及发生频率[J]. 西南农业学报.2001,14(4):55-57.
[130]王英华,国立耘,梁德霖,等.马铃薯晚疫病菌在内蒙古和甘肃的交配型分布及对几种杀菌剂的敏感性[J].中国农业大学学报,2003,8(1):78-82.
[131]朱小琼,车兴碧,国力耘,等.六省市致病疫霉交配型及其对几种杀菌剂的敏感性[J]. 植物保护,2004,30(4):20-23.
[132]方仲达.植病研究方法[M].北京,农业出版社,2001.
[133]Black W, Mastenbrock C, Mills W. R , et al. A proposal for an international nomenclature of races of Phytophthora infestans and of genes controlling immunity in Solanum demissum derivaties[J]. Enphytica, 1953 , 2 (3):173 - 179.
[134]张宝国,朱杰华,彭学文.河北省部分地区晚疫病菌生理小种鉴定[J].河北农业大学学报,2004,27(1):77-79.