13个稻瘟菌基因的表达谱分析及受稻瘟病诱导的水稻转录因子OsBTB基因的生物学功能初探
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摘要
在前期工作中,对受稻瘟菌生理小种103(PO6-6)侵染2、4、8h的抗性水稻品系C101A51(携带Pi2)叶片mRNA构建的cDNA文库进行EST测序,共获得了12,270条高质量的ESTs并产生5,741条独立基因。应用BLASTn程序将这些序列分别与水稻和稻瘟菌基因组序列进行搜索(E-value≤e~(-15)),结果有36TUTs与稻瘟菌基因组匹配,其中有13条独立基因仅与稻瘟菌基因组同源。利用本实验室保存的受稻瘟菌侵染的水稻叶片cDNA文库中2,171个独立EST克隆制备完成的cDNA阵列,以稻瘟菌侵染8h的水稻近等基因系H7R/H7S的叶片总RNA为探针,通过芯片杂交、信号提取以及数据分析后,得到一批功能未知的基因。对未知基因通过生物信息学分析发现一个编码含有BTB结构域的蛋白的基因(克隆号为J001E09)并命名为OsBTB。为了确证这13个推测的稻瘟菌基因的生物信息学结果和其在稻瘟菌致病性中可能的生物学功能;通过设计特异引物并测序获得了OsBTB基因的全长cDNA。
分别提取稻瘟菌生理小种Y34和粳稻秀水11基因组,运用Southern blot杂交后,结果表明这13个基因确实来源于稻瘟菌,生物信息学分析表明,除2个未知基因外,有2个分别编码环孢菌素(Cyclophilin)和P型ATP酶的基因(GenBank受理号BI808723和BQ907640)与稻瘟病菌侵染早期附着孢形成相关;C-4甲基固醇氧化酶和δ-固醇C-甲基转移酶(BI813122和BQ907428)参与稻瘟病菌致病性相关的膜蛋白——麦角固醇的合成;还有可能与稻瘟病菌侵染过程中信号传导相关的质膜钠离子应答蛋白(BI812033);与蛋白合成相关的核糖体L12蛋白(BM420098)和延伸因子2(BQ907452);与真核细胞细胞体内的蛋白质的运输和分泌相关的外被体蛋白(BI813165);参与脂肪酸生物合成的烯酰还原酶(BQ907177)等。一个编码丝氨酸/苏氨酸蛋白激酶基因(BQ908235)参与了稻瘟菌孢子的产生;从受稻瘟病菌诱导水稻叶组织中筛选到的这些基因可能在病原与寄主的相互作用中起重要作用。Northern blot结果表明,这些基因在稻瘟菌的附着胞阶段都有不同程度的表达。除了环孢菌素基因在菌丝体阶段有表达外,其它基因在该阶段均未检测到。其中,质膜钠离子反应2基因(BI812033)和Nebula相关蛋白(BM419917)仅在幼附着胞阶段表达;而参与麦角固醇的2个基因的表达模式相类似;丝氨酸/苏氨酸蛋白激酶基因在附着胞的发育过程中其表达呈递减趋势。这些结果不仅反映了它们可能的生物学功能,同时也从一个侧面反映了它们是高丰度基因。
根据生物信息学获得的OsBTB基因全长序列设计基因特异性引物,从水稻cDNA文库J001E09克隆中通过PCR扩增到基因的全长序列,测序确证了该基因的全长为1,975bp的cDNA序列,其中
In the previous study, 12,270 ESTs were generated which were 3 -end sequenced from a cDNA library of rice leaf tissues infected with an incompatible isolate of Magnaporthe grisea, 103(PO6-6) at 2, 4, and 8 h after inoculation and 5,741 unigenes were obtained. The unigene sequences were compared to the draft sequences of the rice and M. grisea respectively. Out of the 5,741 TUTs, 13 TUTs were aligned only with M. grisea genome sequence (http://www.genome.wi.mit.edu) (E-value≤e~(-15)) using BLASTn algorithm. A cDNA array containing 2,171 unique genes from rice leaf cDNA library was prepared to compare analysis of expression profiles of incompatible/compatible interactions between a pair of blast resistance near-isogenic rice lines H7R/H7S with race ZB15 at 8 h after inoculation, unknown gene set was obtained and were analyzed further by bioinformatics, a gene (clone No. J001E09) encoding a protein containing BTB domain and named OsBTB. Southern blot and Northern blot were performed respectively in order to conform in silico results of 13 putative M. grisea genes and their possible biological function in M. grisea pathogenicity. Full-length cDNA of OsBTB were generated using specific primer and sequencing.Southern blot analysis of 13 TUTs from M. grisea and two ESTs from Oryza sativa in genomic DNAs isolated from M. grisea mycelia and rice leaf, respectively. Genomic DNA (5μg) was digested with restriction enzymes (EcoK I) and subjected to Southern blot analysis. Hybridizations were performed using probes corresponding to the EST sequences of the 15 cDNAs labeled by random priming. Southern blot hybridization assay was carried out by using the 13 M. grisea ESTs and 2 rice ESTs as probes, the results confirmed that these genes originated from M. grisea. Bioinformatics analysis suggested that these genes were imported to pathogenesis-related development. Some of these genes, such as Cyclophilin (GenBank accession No. BI808723) and P-tpye ATPase (BQ907640), were involved in appressorium formation. C-4 methyl sterol oxidase (BI813122) and delta (24)-sterol C-methyltransferase (BQ907428) were involved in the synthesis of ergosterol which is essential for pathogenicity of M. grisea. Plasma membrane sodium response 2 (BI812033) may be involved in signal transduction during the infection. Ribosomal protein L12 (BM420098) and elongation factor 2 (BQ907452) were important to protein synthesis. Coatomer (BI813165) was attached to the trafficking of proteins and secretion within eukaryotic cells. CipB protein (BQ907177) was involved in fatty acid biosynthesis. All the above genes may be involved in host-pathogen interaction in rice tissue. A gene encoding serine/threonine kinase (BQ908235) was involved in conidia
production. All the above genes may be involved in host-pathogen interaction in rice tissue. Northern blot assay indicated that these genes were expressed at different degree during appressoria. All genes were not detected during mycelia stage except for cyclophilin. Among these, Plasma membrane sodium response 2 and Nebula related protein were only expressed during young appresorium stage;two genes requiring for ergosterol synthesis were similar to each other;the gene encoding serine/threonine kinase was decreased during appresoria stage. All of these reflected not only their biological functions, but also their abundancy.Specific primers were designed according to in silico full-length sequence of OsBTB obtained, the sequence in the clone J001E09 was amplified by PCR prepared from a rice cDNA library as template. The 1,975 bp long sequence phage DNA contained an open eading (ORF) of 1,677 bp was retrieved through sequencing and assembling, which predicts to encode a protein of 558 amino acids a conserved BTB domain. A DNA sequence about 5174 bp was found through BLASTn to rice genome sequence, containing 6 exons and 5 introns. TATA box and CAAT box were at -27 bp and -HObp on the upstream of transcripton initiator respectively. The similarity was 73.34% between OsBTB and GMPOZ, a nuclear protein, after homology search.This gene was mapped on chromosomes 2 by RFLP analysis using 247 F8 population of Zhenshan97B/Miyang46.The polyclonal antibody was obtained using recombinant OsBTB protein expressed in and purified from E. coli as antigen that was immuned in Xinxilan rabbit. Western blot assay suggested that the expression profiles of OsBTB protein were polyphyletic during different rice lines and corresponding M. grisea isolates incomparible/comparible interaction. There was no specific subcellular location after gold labelled immunosorbent assay. OsBTB and nine known genes (such as Pi-ta, Pib, glucosidase, glycine-rich protein, chitinase, beta-l,3-glucanase, PAL and ubiquitin) were detected using quantative real-time RT-PCR assay, trend of expression in OsBTB was similar to Pi-ta, Pib, chitinase and glycine-rich protein, expression were induced rapidly after M. grisea infiltration, had a peak value at 12h in H7R, but expression of OsBTB reached a peak at 24h. All of genes except alpha-glucosidase were induced after M. grisea infection and reached a peak at 24h in H7S. In addition some genes are induced more rapidly in the resistant than in the susceptible cultivar. RNA interference plant expression vector pCAMBIA-HAN-Cte57B was construced, through partical gun-mediated transformation of japonica rice varieties Zhonghual 1 obtained 1 transgenic plant. PCR screening of the regenerated plant showed that itwas positive. Its biological function was in progress.
分别提取稻瘟菌生理小种Y34和粳稻秀水11基因组,运用Southern blot杂交后,结果表明这13个基因确实来源于稻瘟菌,生物信息学分析表明,除2个未知基因外,有2个分别编码环孢菌素(Cyclophilin)和P型ATP酶的基因(GenBank受理号BI808723和BQ907640)与稻瘟病菌侵染早期附着孢形成相关;C-4甲基固醇氧化酶和δ-固醇C-甲基转移酶(BI813122和BQ907428)参与稻瘟病菌致病性相关的膜蛋白——麦角固醇的合成;还有可能与稻瘟病菌侵染过程中信号传导相关的质膜钠离子应答蛋白(BI812033);与蛋白合成相关的核糖体L12蛋白(BM420098)和延伸因子2(BQ907452);与真核细胞细胞体内的蛋白质的运输和分泌相关的外被体蛋白(BI813165);参与脂肪酸生物合成的烯酰还原酶(BQ907177)等。一个编码丝氨酸/苏氨酸蛋白激酶基因(BQ908235)参与了稻瘟菌孢子的产生;从受稻瘟病菌诱导水稻叶组织中筛选到的这些基因可能在病原与寄主的相互作用中起重要作用。Northern blot结果表明,这些基因在稻瘟菌的附着胞阶段都有不同程度的表达。除了环孢菌素基因在菌丝体阶段有表达外,其它基因在该阶段均未检测到。其中,质膜钠离子反应2基因(BI812033)和Nebula相关蛋白(BM419917)仅在幼附着胞阶段表达;而参与麦角固醇的2个基因的表达模式相类似;丝氨酸/苏氨酸蛋白激酶基因在附着胞的发育过程中其表达呈递减趋势。这些结果不仅反映了它们可能的生物学功能,同时也从一个侧面反映了它们是高丰度基因。
根据生物信息学获得的OsBTB基因全长序列设计基因特异性引物,从水稻cDNA文库J001E09克隆中通过PCR扩增到基因的全长序列,测序确证了该基因的全长为1,975bp的cDNA序列,其中
In the previous study, 12,270 ESTs were generated which were 3 -end sequenced from a cDNA library of rice leaf tissues infected with an incompatible isolate of Magnaporthe grisea, 103(PO6-6) at 2, 4, and 8 h after inoculation and 5,741 unigenes were obtained. The unigene sequences were compared to the draft sequences of the rice and M. grisea respectively. Out of the 5,741 TUTs, 13 TUTs were aligned only with M. grisea genome sequence (http://www.genome.wi.mit.edu) (E-value≤e~(-15)) using BLASTn algorithm. A cDNA array containing 2,171 unique genes from rice leaf cDNA library was prepared to compare analysis of expression profiles of incompatible/compatible interactions between a pair of blast resistance near-isogenic rice lines H7R/H7S with race ZB15 at 8 h after inoculation, unknown gene set was obtained and were analyzed further by bioinformatics, a gene (clone No. J001E09) encoding a protein containing BTB domain and named OsBTB. Southern blot and Northern blot were performed respectively in order to conform in silico results of 13 putative M. grisea genes and their possible biological function in M. grisea pathogenicity. Full-length cDNA of OsBTB were generated using specific primer and sequencing.Southern blot analysis of 13 TUTs from M. grisea and two ESTs from Oryza sativa in genomic DNAs isolated from M. grisea mycelia and rice leaf, respectively. Genomic DNA (5μg) was digested with restriction enzymes (EcoK I) and subjected to Southern blot analysis. Hybridizations were performed using probes corresponding to the EST sequences of the 15 cDNAs labeled by random priming. Southern blot hybridization assay was carried out by using the 13 M. grisea ESTs and 2 rice ESTs as probes, the results confirmed that these genes originated from M. grisea. Bioinformatics analysis suggested that these genes were imported to pathogenesis-related development. Some of these genes, such as Cyclophilin (GenBank accession No. BI808723) and P-tpye ATPase (BQ907640), were involved in appressorium formation. C-4 methyl sterol oxidase (BI813122) and delta (24)-sterol C-methyltransferase (BQ907428) were involved in the synthesis of ergosterol which is essential for pathogenicity of M. grisea. Plasma membrane sodium response 2 (BI812033) may be involved in signal transduction during the infection. Ribosomal protein L12 (BM420098) and elongation factor 2 (BQ907452) were important to protein synthesis. Coatomer (BI813165) was attached to the trafficking of proteins and secretion within eukaryotic cells. CipB protein (BQ907177) was involved in fatty acid biosynthesis. All the above genes may be involved in host-pathogen interaction in rice tissue. A gene encoding serine/threonine kinase (BQ908235) was involved in conidia
production. All the above genes may be involved in host-pathogen interaction in rice tissue. Northern blot assay indicated that these genes were expressed at different degree during appressoria. All genes were not detected during mycelia stage except for cyclophilin. Among these, Plasma membrane sodium response 2 and Nebula related protein were only expressed during young appresorium stage;two genes requiring for ergosterol synthesis were similar to each other;the gene encoding serine/threonine kinase was decreased during appresoria stage. All of these reflected not only their biological functions, but also their abundancy.Specific primers were designed according to in silico full-length sequence of OsBTB obtained, the sequence in the clone J001E09 was amplified by PCR prepared from a rice cDNA library as template. The 1,975 bp long sequence phage DNA contained an open eading (ORF) of 1,677 bp was retrieved through sequencing and assembling, which predicts to encode a protein of 558 amino acids a conserved BTB domain. A DNA sequence about 5174 bp was found through BLASTn to rice genome sequence, containing 6 exons and 5 introns. TATA box and CAAT box were at -27 bp and -HObp on the upstream of transcripton initiator respectively. The similarity was 73.34% between OsBTB and GMPOZ, a nuclear protein, after homology search.This gene was mapped on chromosomes 2 by RFLP analysis using 247 F8 population of Zhenshan97B/Miyang46.The polyclonal antibody was obtained using recombinant OsBTB protein expressed in and purified from E. coli as antigen that was immuned in Xinxilan rabbit. Western blot assay suggested that the expression profiles of OsBTB protein were polyphyletic during different rice lines and corresponding M. grisea isolates incomparible/comparible interaction. There was no specific subcellular location after gold labelled immunosorbent assay. OsBTB and nine known genes (such as Pi-ta, Pib, glucosidase, glycine-rich protein, chitinase, beta-l,3-glucanase, PAL and ubiquitin) were detected using quantative real-time RT-PCR assay, trend of expression in OsBTB was similar to Pi-ta, Pib, chitinase and glycine-rich protein, expression were induced rapidly after M. grisea infiltration, had a peak value at 12h in H7R, but expression of OsBTB reached a peak at 24h. All of genes except alpha-glucosidase were induced after M. grisea infection and reached a peak at 24h in H7S. In addition some genes are induced more rapidly in the resistant than in the susceptible cultivar. RNA interference plant expression vector pCAMBIA-HAN-Cte57B was construced, through partical gun-mediated transformation of japonica rice varieties Zhonghual 1 obtained 1 transgenic plant. PCR screening of the regenerated plant showed that itwas positive. Its biological function was in progress.
引文
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