水稻纹枯病菌遗传多样性及侵染水稻早期上调表达基因的分析
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摘要
稻纹枯病严重制约水稻的产量和品质,已成为水稻上的首要病害。本论文以分离自我国湖北省、江西省、湖南省、江苏省、广西省等省份的纹枯病菌菌株为材料,研究了其遗传多样性;并以分离自武汉水稻田中的WH-1菌株为材料,分离获得了纹枯菌在侵染水稻早期上调表达的基因。主要结果如下:
自水稻叶鞘发病部位及纹枯病菌核共分离获得1088株菌株,结合形态学和分子生物学等鉴定方法,确认98.2%的菌株为茄丝核菌AG-1ⅠA(Rhizoctonia solaniAG-1ⅠA),另外有19株菌株为稻角担菌(Ceratobasidum oryzae-sativae),1株菌株为茄丝核菌AG-7,表明引发水稻纹枯病的病菌主要为茄丝核菌AG-1融合群中的ⅠA亚群,有性态为担子菌门瓜亡革菌(Thanatephorus cucumeris)。对来自同一田块或来自不同省份的菌株进行了比较,发现它们在生长、菌丝直径、菌落形态和致病力等方面存在丰富的多样性。随机引物扩增多态性分析(RAPD)表明来自同一水稻田块的菌株间出现丰富的遗传分化现象,在0.83相似水平上,可以归于16个亚组;RAPD分析也表明来不同省份的71个菌株可以分为15个亚组,且与菌株来源没有显著关系。自广西桂林和湖北武汉的稻田中分别获得3株和1株表型特异的菌株,它们在生长、菌落形态和致病等方面与正常菌株均有显著性差异。
对分离自广西桂林的菌株GXE4进行了详细的研究。与正常菌株相比,菌株GXE4菌落颜色深,呈黄褐色,菌丝生长速率慢,产菌核数量少且不易产生菌核,对水稻没有致病力。采用真菌rDNA基因ITS区域序列、核型观察、菌丝融合试验,证实菌株GXE4是茄丝核菌AG-1融合群ⅠA亚群。菌株GXE4中存在一条独立于染色体和线粒体DNA外的DNA片段,大小约3.6 kb,命名为3.6-kb frags。3.6-kbfrags可以稳定的存在于菌株内,不能被抗生素、原生质体再生、SDS和溴化乙锭等操作消除掉。DNaseI、RNase、限制性内切酶、外切酶Ⅲ等降解试验表明3.6-kb frags具双链DNA特性,且3'端不含有发夹结构或蛋白质保护性结构。对3.6-kb frags进行了克隆和序列分析,发现在3.6-kb frags至少包含有两个大小类似、序列具有同源性的遗传因子,分别命名为3.6-kb frag-1和3.6-kb frag-2;3.6-kb frag-1没有氨基酸数超过110的阅读框,推定不具有编码功能;不完全序列分析表明3.6-kb frag-2也没有较大的阅读框。DNA杂交结果显示位于约23 kb和7.0 kb处还存在与3.6-kbfrags具明显杂交信号的DNA片段。根据这些研究结果初步推定3.6-kb frags可能是dsDNA病毒基因组的缺陷型DNA,或线性真菌质粒。
菌株RCOL-1分离自湖北武汉华中农业大学试验田。菌株RCOL-1对水稻没有致病力,它与双核菌稻角担菌(C.oryzae-sativa)在生长、菌落形态、菌核大小等表型上非常相似,与茄丝核菌AG-1融合群的菌株则有显著的差异。DAPI荧光染色核型分析表明,菌株RCOL-1的细胞核型没有规律,约95.9%为双核,2.8%为多核,1.3%为单核。通过特异性引物PCR扩增及DNA杂交证实RCOL-1中含有C.oryzae-sativae和T.cucumeris两种真菌的rDNA;进一步研究发现该菌株中存在4种类型的核糖体RNA的内转录间隔区(ITS,internal transcribed space),其中大部分的ITS序列与C.oryzae-sativae的相同,少部分的ITS与T.cucumeris相同;另有2种ITS是这两种真菌ITS的嵌合物,即ITS1与C.oryzae-sativae的ITS1相同,ITS2与T.cucumeris的ITS2相同,或ITS1与T.cucumeris的ITS1相同,ITS2与C.oryzae-sativae的ITS2相同。推定RCOL-1中存在4种类型的ITS是T.cucumeris中的rRNA基因向C.oryzae-sativae水平转移的结果。
以水稻纹枯病菌菌株WH-1和水稻9311为材料,采用抑制差减杂交(suppressionsubtractive hybridization,SSH)的方法,筛选获得了129个病菌侵染水稻初期(12 hpi)表达量明显增强的EST克隆。对表达量明显增强的克隆进行序列分析,结果发现96个克隆的序列与Genbank的序列具有一定程度的相似性,33个序列在数据库中没有相似性序列存在。对克隆了进行MIPS基因功能分类分析,推定这些上调表达基因的功能广泛,涉及到代谢(13%)、蛋白命运(9%)、信号传导(5%)、细胞防御(2%)等11类。其中包括了参与稻瘟病菌(M.grisea)、黑粉菌(U.maydis)等真菌致病过程的促分裂原活化蛋白激酶(Mitogen activated protein kinase,MAPK)、Cyclophilin、cyclin-dependent kinase等基因,它们也有可能参与纹枯病菌致病过程。
Rice sheath blight is a particularly important component of rice disease complex and makes great losses to the yield and quality of rice.In this paper,genetic diversity of rice sheath blight fungus was analyzed among isolates obtained from rice fields of Hubei province,Jiangxi province,Hunan province,Jiangsu province and Guangxi province. Up-regulated genes of strain WH-1 from Wuhan County were screened during the early stage of the infection to rice leaves.
1088 isolates were isolated from typical lesions of sheath blight collected from different counties and identified by morphological and molecular techniques.Out of 1088 isolates,1068 belonged to Rhizoctonia solani AG-1ⅠA,19 were R.oryzae-sativae,and 1 belonged to R.solani AG-7.The results suggested that the anastomosis group 1ⅠA (teleomorph T.cucumeris) of R.solani is the major group infecting rice.Rich genetic diversity existed among isolates from different fields,even from the same field based on the growth rate,clone morphology,hyphal diameter,karyon type and pathogenicity.77 isolates from one field of Wuhan could be classified into 16 RAPD subgroups at the 0.83 similarity level;and 71 stains isolates from different provinces were classified into 15 RAPD subgroups and the polymorphic bands were not closely related with the geographic location.In addition,3 isolates from one rice field of Guilin,Guangxi province and 1 from Wuhan,Hubei province were different significantly from other isolates.
A fungal isolate,GXE4,from Guilin was studied in detail.Compared with normal isolate,isolate GXE4 produced abnormal pigment and less sclerotia,grew more slowly, and was much difficult to produce sclerotia.GXE4 also lost the ability to infect rice when inoculated with hyphal agar.GXE4 was identified to belong to R.solani AG-1ⅠA based on the result of hyphal fusion,the number of nuclei and the nucleotide sequence of ITS. There exsited two extra linear chromosomal elements(named as 3.6-kb frag-1 and 3.6-kb frag-2) which were proved to be very stable when GXE4 was treated with antibiotic, protoplast regeneration and EB respectively.The DNA of 3.6-kb frags was sensitive to DNaseI but not to RNase,and had no protein or knob-shaped structure in 3'end of 3.6-kb frags.Partial sequence of 3.6-kb flags was cloned and analyzed.There was high similarity between 3.6-kb frag-1 and 3.6-kb frag-2.The DNA of 3.6-kb frag-1 was presumed not to encode any protein longer than 110 amino acids.The DNA of 3.6-kb frag-2 did not have a longer ORF either.The result of genomic DNA southern blot showed that there was a significant hybridization signal with DNA fragments of 3.6-kb frags at 23 kb and 7.0 kb. Based on the results,3.6-kb frags could be defected DNA elements from genome of dsDNA virus or linear plasmid in fungi.
Isolate RCOL-1,was obtained from a typical lesion of sheath blight collected from the campus of Huazhong Agricultural University.RCOL-1 grew more slowly than both T. cucumeris and C.oryzae-sativae,and formed abnormal colony on PDA;RCOL-1 also lost the ability to attack rice when inoculated with both hyphal agar and sclerotia.DAPI staining showed that the nuclei pattern in individual cell of RCOL-1 was various,namely some cells contained only one nucleus(1.3%),most contained double nuclei(95.9%), and some cells contained multi-nuclei(2.8%).There existed ITS DNAs of C. oryzae-sativae and T.cucumeris in isolate RCOL-1.Based on the results of PCR amplification using 4 primers specific to T.cucumeris and C.oryzae-sativae,RCOL-1 contained four types ITS DNA,namely ITS DNA of C.oryzae-sativae,ITS DNA of T. cucumeris and two types of chimerical ITS DNA.Two chimerical ITS DNAs were ITS1 of from C.oryzae-sativae and ITS2 of from T.cucumeris,and ITS1 of T.cucumeris and ITS2 of C.oryzae-sativae.The results implied that homologous recombination had occurred at the rDNA region between the two pathogenic fungi.
In order to understand the molecular interaction of R.solani AG-1ⅠA and rice, cDNA library was constructed by suppression subtractive hybridization(SSH) using rice (Oryza sativa) 9311 inoculated with a highly aggressive strain WH-1,and 129 fungal clones expression strengthened significantly during the early stage of infection were obtained.These EST_S were sequenced and analyzed on NCBI non-redundant GenBank database.96 EST_S had significant homology to sequences in the database and 33 had no known homologues in current database.The probable function of the 96 EST_S could be classified into 12 groups according to their putative BLASTX identification.The groups include Metabolism(13%),Protein fate(9%),Protein synthesis(8%),Energy(6%), Signal transduction(5%),Cellular transport(4%),Cell defense(2%),Transcription(1%), Structural(1%),Regulation(1%),Cell cycle(1%) and Unclassified function(49%). Putative proteins of those EST_S included pathogenic signal conduction factor-Mitogen activated protein kinase(MAPK),cyclin-dependent kinase(CDK).These genes may involve in pathogenesis of R.solani and contribute to pathogenesis,and this need further study in the future.
自水稻叶鞘发病部位及纹枯病菌核共分离获得1088株菌株,结合形态学和分子生物学等鉴定方法,确认98.2%的菌株为茄丝核菌AG-1ⅠA(Rhizoctonia solaniAG-1ⅠA),另外有19株菌株为稻角担菌(Ceratobasidum oryzae-sativae),1株菌株为茄丝核菌AG-7,表明引发水稻纹枯病的病菌主要为茄丝核菌AG-1融合群中的ⅠA亚群,有性态为担子菌门瓜亡革菌(Thanatephorus cucumeris)。对来自同一田块或来自不同省份的菌株进行了比较,发现它们在生长、菌丝直径、菌落形态和致病力等方面存在丰富的多样性。随机引物扩增多态性分析(RAPD)表明来自同一水稻田块的菌株间出现丰富的遗传分化现象,在0.83相似水平上,可以归于16个亚组;RAPD分析也表明来不同省份的71个菌株可以分为15个亚组,且与菌株来源没有显著关系。自广西桂林和湖北武汉的稻田中分别获得3株和1株表型特异的菌株,它们在生长、菌落形态和致病等方面与正常菌株均有显著性差异。
对分离自广西桂林的菌株GXE4进行了详细的研究。与正常菌株相比,菌株GXE4菌落颜色深,呈黄褐色,菌丝生长速率慢,产菌核数量少且不易产生菌核,对水稻没有致病力。采用真菌rDNA基因ITS区域序列、核型观察、菌丝融合试验,证实菌株GXE4是茄丝核菌AG-1融合群ⅠA亚群。菌株GXE4中存在一条独立于染色体和线粒体DNA外的DNA片段,大小约3.6 kb,命名为3.6-kb frags。3.6-kbfrags可以稳定的存在于菌株内,不能被抗生素、原生质体再生、SDS和溴化乙锭等操作消除掉。DNaseI、RNase、限制性内切酶、外切酶Ⅲ等降解试验表明3.6-kb frags具双链DNA特性,且3'端不含有发夹结构或蛋白质保护性结构。对3.6-kb frags进行了克隆和序列分析,发现在3.6-kb frags至少包含有两个大小类似、序列具有同源性的遗传因子,分别命名为3.6-kb frag-1和3.6-kb frag-2;3.6-kb frag-1没有氨基酸数超过110的阅读框,推定不具有编码功能;不完全序列分析表明3.6-kb frag-2也没有较大的阅读框。DNA杂交结果显示位于约23 kb和7.0 kb处还存在与3.6-kbfrags具明显杂交信号的DNA片段。根据这些研究结果初步推定3.6-kb frags可能是dsDNA病毒基因组的缺陷型DNA,或线性真菌质粒。
菌株RCOL-1分离自湖北武汉华中农业大学试验田。菌株RCOL-1对水稻没有致病力,它与双核菌稻角担菌(C.oryzae-sativa)在生长、菌落形态、菌核大小等表型上非常相似,与茄丝核菌AG-1融合群的菌株则有显著的差异。DAPI荧光染色核型分析表明,菌株RCOL-1的细胞核型没有规律,约95.9%为双核,2.8%为多核,1.3%为单核。通过特异性引物PCR扩增及DNA杂交证实RCOL-1中含有C.oryzae-sativae和T.cucumeris两种真菌的rDNA;进一步研究发现该菌株中存在4种类型的核糖体RNA的内转录间隔区(ITS,internal transcribed space),其中大部分的ITS序列与C.oryzae-sativae的相同,少部分的ITS与T.cucumeris相同;另有2种ITS是这两种真菌ITS的嵌合物,即ITS1与C.oryzae-sativae的ITS1相同,ITS2与T.cucumeris的ITS2相同,或ITS1与T.cucumeris的ITS1相同,ITS2与C.oryzae-sativae的ITS2相同。推定RCOL-1中存在4种类型的ITS是T.cucumeris中的rRNA基因向C.oryzae-sativae水平转移的结果。
以水稻纹枯病菌菌株WH-1和水稻9311为材料,采用抑制差减杂交(suppressionsubtractive hybridization,SSH)的方法,筛选获得了129个病菌侵染水稻初期(12 hpi)表达量明显增强的EST克隆。对表达量明显增强的克隆进行序列分析,结果发现96个克隆的序列与Genbank的序列具有一定程度的相似性,33个序列在数据库中没有相似性序列存在。对克隆了进行MIPS基因功能分类分析,推定这些上调表达基因的功能广泛,涉及到代谢(13%)、蛋白命运(9%)、信号传导(5%)、细胞防御(2%)等11类。其中包括了参与稻瘟病菌(M.grisea)、黑粉菌(U.maydis)等真菌致病过程的促分裂原活化蛋白激酶(Mitogen activated protein kinase,MAPK)、Cyclophilin、cyclin-dependent kinase等基因,它们也有可能参与纹枯病菌致病过程。
Rice sheath blight is a particularly important component of rice disease complex and makes great losses to the yield and quality of rice.In this paper,genetic diversity of rice sheath blight fungus was analyzed among isolates obtained from rice fields of Hubei province,Jiangxi province,Hunan province,Jiangsu province and Guangxi province. Up-regulated genes of strain WH-1 from Wuhan County were screened during the early stage of the infection to rice leaves.
1088 isolates were isolated from typical lesions of sheath blight collected from different counties and identified by morphological and molecular techniques.Out of 1088 isolates,1068 belonged to Rhizoctonia solani AG-1ⅠA,19 were R.oryzae-sativae,and 1 belonged to R.solani AG-7.The results suggested that the anastomosis group 1ⅠA (teleomorph T.cucumeris) of R.solani is the major group infecting rice.Rich genetic diversity existed among isolates from different fields,even from the same field based on the growth rate,clone morphology,hyphal diameter,karyon type and pathogenicity.77 isolates from one field of Wuhan could be classified into 16 RAPD subgroups at the 0.83 similarity level;and 71 stains isolates from different provinces were classified into 15 RAPD subgroups and the polymorphic bands were not closely related with the geographic location.In addition,3 isolates from one rice field of Guilin,Guangxi province and 1 from Wuhan,Hubei province were different significantly from other isolates.
A fungal isolate,GXE4,from Guilin was studied in detail.Compared with normal isolate,isolate GXE4 produced abnormal pigment and less sclerotia,grew more slowly, and was much difficult to produce sclerotia.GXE4 also lost the ability to infect rice when inoculated with hyphal agar.GXE4 was identified to belong to R.solani AG-1ⅠA based on the result of hyphal fusion,the number of nuclei and the nucleotide sequence of ITS. There exsited two extra linear chromosomal elements(named as 3.6-kb frag-1 and 3.6-kb frag-2) which were proved to be very stable when GXE4 was treated with antibiotic, protoplast regeneration and EB respectively.The DNA of 3.6-kb frags was sensitive to DNaseI but not to RNase,and had no protein or knob-shaped structure in 3'end of 3.6-kb frags.Partial sequence of 3.6-kb flags was cloned and analyzed.There was high similarity between 3.6-kb frag-1 and 3.6-kb frag-2.The DNA of 3.6-kb frag-1 was presumed not to encode any protein longer than 110 amino acids.The DNA of 3.6-kb frag-2 did not have a longer ORF either.The result of genomic DNA southern blot showed that there was a significant hybridization signal with DNA fragments of 3.6-kb frags at 23 kb and 7.0 kb. Based on the results,3.6-kb frags could be defected DNA elements from genome of dsDNA virus or linear plasmid in fungi.
Isolate RCOL-1,was obtained from a typical lesion of sheath blight collected from the campus of Huazhong Agricultural University.RCOL-1 grew more slowly than both T. cucumeris and C.oryzae-sativae,and formed abnormal colony on PDA;RCOL-1 also lost the ability to attack rice when inoculated with both hyphal agar and sclerotia.DAPI staining showed that the nuclei pattern in individual cell of RCOL-1 was various,namely some cells contained only one nucleus(1.3%),most contained double nuclei(95.9%), and some cells contained multi-nuclei(2.8%).There existed ITS DNAs of C. oryzae-sativae and T.cucumeris in isolate RCOL-1.Based on the results of PCR amplification using 4 primers specific to T.cucumeris and C.oryzae-sativae,RCOL-1 contained four types ITS DNA,namely ITS DNA of C.oryzae-sativae,ITS DNA of T. cucumeris and two types of chimerical ITS DNA.Two chimerical ITS DNAs were ITS1 of from C.oryzae-sativae and ITS2 of from T.cucumeris,and ITS1 of T.cucumeris and ITS2 of C.oryzae-sativae.The results implied that homologous recombination had occurred at the rDNA region between the two pathogenic fungi.
In order to understand the molecular interaction of R.solani AG-1ⅠA and rice, cDNA library was constructed by suppression subtractive hybridization(SSH) using rice (Oryza sativa) 9311 inoculated with a highly aggressive strain WH-1,and 129 fungal clones expression strengthened significantly during the early stage of infection were obtained.These EST_S were sequenced and analyzed on NCBI non-redundant GenBank database.96 EST_S had significant homology to sequences in the database and 33 had no known homologues in current database.The probable function of the 96 EST_S could be classified into 12 groups according to their putative BLASTX identification.The groups include Metabolism(13%),Protein fate(9%),Protein synthesis(8%),Energy(6%), Signal transduction(5%),Cellular transport(4%),Cell defense(2%),Transcription(1%), Structural(1%),Regulation(1%),Cell cycle(1%) and Unclassified function(49%). Putative proteins of those EST_S included pathogenic signal conduction factor-Mitogen activated protein kinase(MAPK),cyclin-dependent kinase(CDK).These genes may involve in pathogenesis of R.solani and contribute to pathogenesis,and this need further study in the future.
引文
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