小麦成株抗条锈病差异表达基因的cDNA-AFLP分析及小麦与条锈菌互作相关基因的克隆与特征研究
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摘要
由条形柄锈菌(Puccinia striiformis f. sp. tritici)引起的小麦条锈病是世界各国小麦生产上危害最为严重的一类真菌病害。实践证明,选育并合理利用抗锈品种是控制小麦条锈病最为经济、安全、有效的方法。小麦与条锈菌互作的分子机理研究,可为揭示病原物的致病机理和寄主植物的抗病机制奠定基础,同时也为小麦抗条锈品种的合理利用和遗传改良及病害的持久控制提供理论依据。本论文就小麦与条锈菌的互作主要开展了以下研究工作：
1.采用cDNA-AFLP技术对小麦成株抗锈性品种兴资9104与条锈菌小种CYR32互作中差异表达基因进行分析,通过64对引物分别获得苗期和成株期转录衍生片段约32320个和34880个。其中37对引物检测到成株期差异表达的TDFs (transcripts-derived fragment)2201个、苗期2529个,同时成功回收得到条锈菌诱导的小麦兴资9104的苗期和成株期差异表达的TDF分别为304个和330个。经克隆、测序及Cap3软件聚类分析,在获得509个Unigenes中,包括组成成株期差异表达TDFs文库的259个(其中131个contig,128个singletons)和组成苗期差异表达TDFs文库的250个(其中132个contig,118个singletons)。基因序列已提交GenBank并获得注册号。
所得的Unigenes与NCBI非冗余蛋白质数据库进行BlastX比对分析,经功能注释将苗期和成株期的Unigenes分别分为14类,除占比例最大部分的未知功能蛋白(Unclear classification)和没有显著匹配的序列(Unclassified)即No hits的蛋白外,其余具有很好匹配的Unigenes的功能涉及代谢、能量、细胞生长、转录、蛋白质合成、蛋白质储藏与运输、转运子、胞内运输、细胞结构、信号转导、抗病与防御和转座。明确了条锈菌CYR32与小麦兴资9104互作的差异基因总体表达情况及各类基因所占的比例,同时也为新基因的发现奠定了基础。65条分别于苗期和成株期差异表达的TDFs(涉及能量、代谢、防御、未知功能等方面)在功能上存在重叠,表明这些基因在互作中可能具有不同的表达模式。
2.利用PCR并结合5'RACE技术得到了5个小麦条锈菌吸器分泌蛋白基因的全长cDNA序列,分别命名为PstSP2C7、PstSP11L10、PstSP11P10、PstSP6P1和Pst15a23,全长分别为1094 bp,837 bp,769 bp,1001 bp和568 bp, ORF区域编码89～203个氨基酸。在GenBank中未得到任何同源序列,但与秆锈菌数据库的基因有一定的同源性。生物信息学分析表明,所有的序列中,除了N-端含有一个信号肽序列外,无任何其他的功能结构域、跨膜螺旋及潜在的糖基磷脂酰肌醇锚定位点(potential glycosyl-phosphatidylino-sitol (GPI) anchor sites)的存在,且都为具有信号肽分泌途径的孢外蛋白。
3.对基因PstSP2C7, PstSP11L10和PstSP11P10的实时荧光定量分析表明,各基因在不同生理阶段具有不同的表达模式。PstSP11L10基因,在侵染的叶片中的表达量远远多于孢子和萌发的芽管,推测其可能在吸器、孢间菌丝或是同时在吸器和孢间菌丝中被诱导表达,这也同时说明该基因在条锈菌与小麦互作中可能发挥一定的积极作用。PstSP2C7基因在萌发的孢子中的表达量最少,而在夏孢子中的表达量最大,且在侵染叶片中的表达量并不是远远少于夏孢子,这可能是因为采集的小麦叶片是在夏孢子大量形成时,所以导致该基因在侵染的叶片及夏孢子中的表达量都较萌发的芽管中多很多。PstSP11P10基因可能在互作中的表达受到抑制,导致其在萌发的芽管和侵染的叶片中的表达量都很少。这种不同的表达模式也表明了各基因在互作中可能发挥着不同的功能。
4.采用5'RACE技术获得了条锈菌基因PstAAC9Pl的全长cDNA,其ORF区域编码134个氨基酸,为ADP/ATP载体蛋白。同源性分析表明其与构巢曲霉、烟曲霉、小麦褐斑病菌、苜蓿黄萎病菌、玉蜀黍赤霉菌、葡萄孢盘菌等多种真菌的ADP/ATP载体蛋白的一致性达80%以上。进化树结果显示其与秆锈菌基因PGTG 14813.2的亲缘关系最近。实时荧光定量PCR分析,发现该基因在侵染的叶片中有最大量的表达,表明其在条锈菌侵染的过程中可能发挥着积极作用。
Stripe rust, caused by Puccinia striiformis f. sp. tritici (Pst) is a the most damaging fungal disease of wheat (Triticum aestivum L.) in all over the world. It has been proved that breeding and growing resistant wheat cultivars is the most effective, economical and environmentally friendly way for controlling this disease. Therefore, molecular mechanism of synergism investigation of wheat plant and rust fungi is greatly helpful for revealing the pathogenic mechanism of the rust fungi and the resistant mechanism of the host and providing further information for selection and reasonable use of resistant wheat cultivars. In the present thesis, the main work related to Pst-wheat interaction system were represented.
1. The cDNA-AFLP method is used to analyze the differentially expressed genes of adult plant resistant wheat cultivar'Xingzi9104'infected by PST pathotype CYR32.64 primer combinations had been used and the total number of about 34880 and 32320 differentially expressed transcript derived fragments (DE-TDFs) at seedling stage and adult-plant stage Xingzi9104 respectively were detected, of which 37 primer combinations were selected for producing reliable polymorphic bands. About 2529 DE-TDFs of seedling stage and 2201 DE-TDFs of adult stage were obtained, of which 304 and 330 were successfully purified. The total of 509 Unigenes were produced by cloning, sequencing and assembling analyses by Cap3 software, which consist of adult-plant DE-TDFs (aDE-TDFs) library of 259 unigenes and seedling DE-TDFs (sDE-TDFs) library of 250 unigenes. All the TDF were submitted to GenBank and got the accession Numbers.
All the Unigens were compared to the NCBI non-redundant protein database using the BlastX program. Two libraries were manually classified as fourteen different categories separately based on functional annotations, besides the biggest parts of unclear classification and the unclassified (also as No hits) genes, the other functional genes including metabolism, energy, cell growth, transcription, protein synthesis, protein destination and storage, transporters, intracellular traffic, cell structure, cell signal transduction, disease/defence and transposon. Clarified the variations and proportion of differential genes expression during interaction of Pst-whaet (CYR32 and Xingzi9104), and also laid a foundation for the new gene discovery.65 overlapped DE-TDFs between the two libraries with functions of energy, metabolism, disease/defence and the unclassified implied that these genes showed different expression patterns during interaction of Pst and wheat.
2. Full-length cDNAs were obtained of five putative secreted protein genes from Pst haustoria using the PCR and 5'rapid amplification of cDNA ends (RACE), and designated as PstSP2C7, PstSP11L10, PstSP11P10, PstSP6P1 and Pst15a23. Their transcripts were 1094 bp,837 bp,769 bp,1001 bp and 568 bp, respectively and encoding predicted proteins ranged from 89 to 203 amino acids of open reading frame (ORF) without significant similarities to any accessions in the GenBank protein database, but with some homologies to predicted proteins in P. graminis, the stem rust pathogen. Bioinformatics analysis indicated that all these five genes only had predicted secretary signal peptides in N-terminal and had no conserved functional motifs, trans-membrane helices and potential glycosyl-phosphatidylinositol (GPI) anchor sites. They were the protein which had signal peptide secreted way and located in outside of cells.
3. qRT-PCR study showed that all these three genes of PstSP2C7, PstSP11L10and PstSP11P10 had the different expressed patterns in transcript levels in different developmental and infection stages of the pathogen. PstSP11L10 was expressed at the highest level in the infected leaves and much higher than in the urediniospores and germinated urediniospores, indicating that PstSP11L10 is presumably either in haustoria or the intercellular mycelium or both and more likely to act in the wheat-Pst interaction. PstSP2C7 had the lowest expression in the germinated urediniospores but had the highest expression level in urediniospores, Although not the highest, the PstSP2C7 expression in the infected leaves was not significantly lower than the level in urediniospores, which might be due to the fact that urediniospores were beginning to form in the infected leaves when leaf samples were taken to detect the gene transcripts. Though PstSP11P10 had the highest expression level in urediniospores, however, The PstSP11P10 expression in infected leaves was much lower than that in urediniospores, which indicated this gene's expression maybe in suppression during the interaction. The different expression patterns of the three genes may indicate their different functions.
4. A novel gene which might encode ADP/ATP carrier protein PstAAC9P1 from Pst haustoria was studied by 5'RACE with full length cDNA which contains open reading frame (ORF) of 134 amino acid residues with over 80% homology to many ADP/ATP carrier proteins form fungi like Aspergillus nidulans, Aspergillus fumigatus, Pyrenophora tritici-repentis,Verticillium albo-atrum, Gibberella zea, Botryotinia Fuckeliana and with 99% identity to P. graminis. gene PGTG_14813.2. PstAAC9P1 had the highest transcription level in the infected leaves, indicated that this gene might play the activate role during the infection of Pst.
1.采用cDNA-AFLP技术对小麦成株抗锈性品种兴资9104与条锈菌小种CYR32互作中差异表达基因进行分析,通过64对引物分别获得苗期和成株期转录衍生片段约32320个和34880个。其中37对引物检测到成株期差异表达的TDFs (transcripts-derived fragment)2201个、苗期2529个,同时成功回收得到条锈菌诱导的小麦兴资9104的苗期和成株期差异表达的TDF分别为304个和330个。经克隆、测序及Cap3软件聚类分析,在获得509个Unigenes中,包括组成成株期差异表达TDFs文库的259个(其中131个contig,128个singletons)和组成苗期差异表达TDFs文库的250个(其中132个contig,118个singletons)。基因序列已提交GenBank并获得注册号。
所得的Unigenes与NCBI非冗余蛋白质数据库进行BlastX比对分析,经功能注释将苗期和成株期的Unigenes分别分为14类,除占比例最大部分的未知功能蛋白(Unclear classification)和没有显著匹配的序列(Unclassified)即No hits的蛋白外,其余具有很好匹配的Unigenes的功能涉及代谢、能量、细胞生长、转录、蛋白质合成、蛋白质储藏与运输、转运子、胞内运输、细胞结构、信号转导、抗病与防御和转座。明确了条锈菌CYR32与小麦兴资9104互作的差异基因总体表达情况及各类基因所占的比例,同时也为新基因的发现奠定了基础。65条分别于苗期和成株期差异表达的TDFs(涉及能量、代谢、防御、未知功能等方面)在功能上存在重叠,表明这些基因在互作中可能具有不同的表达模式。
2.利用PCR并结合5'RACE技术得到了5个小麦条锈菌吸器分泌蛋白基因的全长cDNA序列,分别命名为PstSP2C7、PstSP11L10、PstSP11P10、PstSP6P1和Pst15a23,全长分别为1094 bp,837 bp,769 bp,1001 bp和568 bp, ORF区域编码89～203个氨基酸。在GenBank中未得到任何同源序列,但与秆锈菌数据库的基因有一定的同源性。生物信息学分析表明,所有的序列中,除了N-端含有一个信号肽序列外,无任何其他的功能结构域、跨膜螺旋及潜在的糖基磷脂酰肌醇锚定位点(potential glycosyl-phosphatidylino-sitol (GPI) anchor sites)的存在,且都为具有信号肽分泌途径的孢外蛋白。
3.对基因PstSP2C7, PstSP11L10和PstSP11P10的实时荧光定量分析表明,各基因在不同生理阶段具有不同的表达模式。PstSP11L10基因,在侵染的叶片中的表达量远远多于孢子和萌发的芽管,推测其可能在吸器、孢间菌丝或是同时在吸器和孢间菌丝中被诱导表达,这也同时说明该基因在条锈菌与小麦互作中可能发挥一定的积极作用。PstSP2C7基因在萌发的孢子中的表达量最少,而在夏孢子中的表达量最大,且在侵染叶片中的表达量并不是远远少于夏孢子,这可能是因为采集的小麦叶片是在夏孢子大量形成时,所以导致该基因在侵染的叶片及夏孢子中的表达量都较萌发的芽管中多很多。PstSP11P10基因可能在互作中的表达受到抑制,导致其在萌发的芽管和侵染的叶片中的表达量都很少。这种不同的表达模式也表明了各基因在互作中可能发挥着不同的功能。
4.采用5'RACE技术获得了条锈菌基因PstAAC9Pl的全长cDNA,其ORF区域编码134个氨基酸,为ADP/ATP载体蛋白。同源性分析表明其与构巢曲霉、烟曲霉、小麦褐斑病菌、苜蓿黄萎病菌、玉蜀黍赤霉菌、葡萄孢盘菌等多种真菌的ADP/ATP载体蛋白的一致性达80%以上。进化树结果显示其与秆锈菌基因PGTG 14813.2的亲缘关系最近。实时荧光定量PCR分析,发现该基因在侵染的叶片中有最大量的表达,表明其在条锈菌侵染的过程中可能发挥着积极作用。
Stripe rust, caused by Puccinia striiformis f. sp. tritici (Pst) is a the most damaging fungal disease of wheat (Triticum aestivum L.) in all over the world. It has been proved that breeding and growing resistant wheat cultivars is the most effective, economical and environmentally friendly way for controlling this disease. Therefore, molecular mechanism of synergism investigation of wheat plant and rust fungi is greatly helpful for revealing the pathogenic mechanism of the rust fungi and the resistant mechanism of the host and providing further information for selection and reasonable use of resistant wheat cultivars. In the present thesis, the main work related to Pst-wheat interaction system were represented.
1. The cDNA-AFLP method is used to analyze the differentially expressed genes of adult plant resistant wheat cultivar'Xingzi9104'infected by PST pathotype CYR32.64 primer combinations had been used and the total number of about 34880 and 32320 differentially expressed transcript derived fragments (DE-TDFs) at seedling stage and adult-plant stage Xingzi9104 respectively were detected, of which 37 primer combinations were selected for producing reliable polymorphic bands. About 2529 DE-TDFs of seedling stage and 2201 DE-TDFs of adult stage were obtained, of which 304 and 330 were successfully purified. The total of 509 Unigenes were produced by cloning, sequencing and assembling analyses by Cap3 software, which consist of adult-plant DE-TDFs (aDE-TDFs) library of 259 unigenes and seedling DE-TDFs (sDE-TDFs) library of 250 unigenes. All the TDF were submitted to GenBank and got the accession Numbers.
All the Unigens were compared to the NCBI non-redundant protein database using the BlastX program. Two libraries were manually classified as fourteen different categories separately based on functional annotations, besides the biggest parts of unclear classification and the unclassified (also as No hits) genes, the other functional genes including metabolism, energy, cell growth, transcription, protein synthesis, protein destination and storage, transporters, intracellular traffic, cell structure, cell signal transduction, disease/defence and transposon. Clarified the variations and proportion of differential genes expression during interaction of Pst-whaet (CYR32 and Xingzi9104), and also laid a foundation for the new gene discovery.65 overlapped DE-TDFs between the two libraries with functions of energy, metabolism, disease/defence and the unclassified implied that these genes showed different expression patterns during interaction of Pst and wheat.
2. Full-length cDNAs were obtained of five putative secreted protein genes from Pst haustoria using the PCR and 5'rapid amplification of cDNA ends (RACE), and designated as PstSP2C7, PstSP11L10, PstSP11P10, PstSP6P1 and Pst15a23. Their transcripts were 1094 bp,837 bp,769 bp,1001 bp and 568 bp, respectively and encoding predicted proteins ranged from 89 to 203 amino acids of open reading frame (ORF) without significant similarities to any accessions in the GenBank protein database, but with some homologies to predicted proteins in P. graminis, the stem rust pathogen. Bioinformatics analysis indicated that all these five genes only had predicted secretary signal peptides in N-terminal and had no conserved functional motifs, trans-membrane helices and potential glycosyl-phosphatidylinositol (GPI) anchor sites. They were the protein which had signal peptide secreted way and located in outside of cells.
3. qRT-PCR study showed that all these three genes of PstSP2C7, PstSP11L10and PstSP11P10 had the different expressed patterns in transcript levels in different developmental and infection stages of the pathogen. PstSP11L10 was expressed at the highest level in the infected leaves and much higher than in the urediniospores and germinated urediniospores, indicating that PstSP11L10 is presumably either in haustoria or the intercellular mycelium or both and more likely to act in the wheat-Pst interaction. PstSP2C7 had the lowest expression in the germinated urediniospores but had the highest expression level in urediniospores, Although not the highest, the PstSP2C7 expression in the infected leaves was not significantly lower than the level in urediniospores, which might be due to the fact that urediniospores were beginning to form in the infected leaves when leaf samples were taken to detect the gene transcripts. Though PstSP11P10 had the highest expression level in urediniospores, however, The PstSP11P10 expression in infected leaves was much lower than that in urediniospores, which indicated this gene's expression maybe in suppression during the interaction. The different expression patterns of the three genes may indicate their different functions.
4. A novel gene which might encode ADP/ATP carrier protein PstAAC9P1 from Pst haustoria was studied by 5'RACE with full length cDNA which contains open reading frame (ORF) of 134 amino acid residues with over 80% homology to many ADP/ATP carrier proteins form fungi like Aspergillus nidulans, Aspergillus fumigatus, Pyrenophora tritici-repentis,Verticillium albo-atrum, Gibberella zea, Botryotinia Fuckeliana and with 99% identity to P. graminis. gene PGTG_14813.2. PstAAC9P1 had the highest transcription level in the infected leaves, indicated that this gene might play the activate role during the infection of Pst.
引文
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