农家种红蚰麦抗白粉病基因的微卫星定位及白粉菌诱导的差异表达谱分析
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摘要
由禾谷类白粉菌Blumeria graminis(DC.)E.O.Speer f. sp.tritici Em.Marchal引起的小麦白粉病是一种世界性病害,在各主要产麦国均有发生。种植抗病品种是防治该病害的有效、安全的手段。由于小麦白粉病菌具有生理小种多,变异速度快的特点,导致生产上种植的小麦品种抗性容易丧失。因此不断鉴定新的抗白粉病基因,深入研究小麦抗白粉病的分子机理是控制该病害的基础性工作。
农家小麦品种中蕴含着丰富的抗病资源,对于发现抗白粉病品种,寻找新的抗病基因具有重要的利用价值。红蚰麦是一农家品种,具有优良的白粉病抗性,遗传分析表明其携带一对显性抗白粉病基因。本文通过SSR标记的方法对该基因进行了染色体定位。同时利用Affymetrix基因芯片结合实时定量PCR验证,分析了红蚰麦／豫麦13F3代纯合抗病家系、感病家系白粉菌诱导条件下的基因表达谱。并根据基因芯片的结果选择一个在抗病家系中诱导后高上调表达的探针进行了电子克隆和表达分析。主要的研究内容和研究结果如下：
1.抗白粉病基因Pmhym的标记定位
根据集团分离分析法在红蚰麦与感病亲本豫麦13的F3代纯合家系中建立抗、感病DNA池。选取位于21条染色体上的250对SSR引物在双亲以及DNA池间进行初筛,筛选出的能扩增出一致性差异片段的引物用F2代群体验证。用软件Mapmaker3.0软件分析结果,最终找到了3个与红蚰麦中抗白粉病基因Pmhym(暂命名)连锁的标记,标记排序如下Xwmc526-31.6cM-Xgwm577-25.6cM-Pmhym-14.3cM-Xwmc232,由于这三个标记都位于7BL染色体上,于是将红蚰麦携带的抗白粉病基因定位在7BL染色体臂上。
2.利用基因芯片筛选白粉菌诱导条件下差异表达基因。
分别以F3 4个纯合抗病家系构建抗病池,4个感病家系构建(?)病池。对两份材料同时设置两种接种处理：未接菌、接菌24h。样品的cRNA与Affymetrix 900493芯片进行杂交,获得的数据均一化处理后,设置了抗病诱导前后、感病诱导前后、诱导后抗病和感病,三组比较。挑选比较后表达倍数有差异的探针序列合成了22对引物,对基因芯片的结果进行了定量PCR验证,结果表明芯片的数据具有可重复性。
以实验组杂交信号为P,change p-value<0.001且ratio值≥2或≤0.5为标准筛选差异表达基因。筛选到2728个在抗病株系诱导前后差异表达的基因,其中已知功能的408个基因中抗病／防御类基因占39.71%。2631个基因在感病诱导前后差异表达,其中已知功能的442个基因中抗病／防御类基因占37.33%,说明病原菌诱导了植物的抗病代谢途径。
通过抗病诱导表达谱和感病诱导表达谱的比较分析、抗病诱导比感病诱导上调表达序列的分析获得了抗病特异诱导转录本,感病特异诱导转录本,以及基础防御转录本以及抗病增强表达转录本。通过差异基因的比较分析推测水杨酸、茉莉酸和乙烯途径协同作用参与了小麦对白粉菌的抗病过程；而感病株系中脱落酸途径可能与以上的三个途径存在拮抗作用,参与了植物-病原物的亲和互作反应。在抗病株系中或活性氧的代谢比感病株系中强,参与了植物的抗病反应过程。许多抗病／防御类基因、信号传导基因上调表达,这些基因可能也参与了抗病反应。
3.一个小麦类萌发素蛋白基因的克隆、表达分析
选取在抗病株系诱导后高上调表达的探针Ta.169.1.S1_x_at,进行电子延伸后合成引物扩增经白粉菌诱导的cDNA模板。结果获得了一个长855 bp,具有完整ORF框的类萌发素蛋白基因,根据ORF框翻译的蛋白序列,在NCBI上进行BLAST分析,表明该片段属于Cupin2超家族成员,有15个相似性在50-64%之间的蛋白序列。用MEGA 4.0软件构建的遗传进化树分析同源关系,本试验中获得的小麦类萌发素蛋白与已知的类萌发素蛋白分属于不同的进化分支,可能是一个新的基因,命名为TaGLP5。序列已提交至Genbank,登录号为FJ594470。用白粉病同时接种抗病、感病株系,发现该基因在感、抗植株中受白粉菌诱导后都上调表达。但上调的模式、程度不同,且在接种24 h内抗病植株中的表达量较感病植株中高。由于病原菌与植物互作的前24 h是决定病原菌能否在植物中定殖的关键时期,推测TaGLP5可能参与了红蚰麦对白粉菌早期的抗病防御反应。
Wheat powdery mildew caused by Blumeria graminis f. sp. tritici is a devastating disease in wheat growing countries worldwide. Resistance cultivars offer the most effective and safe approach to control the disease. However, the use of single or few Pm genes eventually leads to a rapid development of correponding virulence genes in the pathogen, resulting in breakdown of cultivars'resistance. Thus, it is foundational to identify novel Pm genes and dissect resistance mechanism for the control of wheat powdery mildew.
Wheat landrace may contain rich resistance genes, which is very valuable to explore resistant cultivars and identify new resistance genes against powdery mildew. Hongyoumai, one wheat landrace, bears good resistance to powdery mildew. Genetic analysis revealed that it carried one dominant Pm gene. In the present thesis, SSR markers were employed to locate Pmhym gene. Meanwhile, Affymetrix genechip analyses were employed for expression profiling in response to Blumeria graminis f. sp. tritici infection in homologous resistant and susceptible F3 progeny derived from Hongyoumai and Yumai13, respectively, along with verification of differentially expressed genes using real time quantitative RT-PCR (qRT-PCR) analyses. Based on the genechip results, one up-regulated target gene in resistance progeny was further in silico cloned and characterized. The main contents and results were as follows:
1. Chromosome location of Pmhym gene
Bulked segregate analysis (BSA) was used to establish the resistant and susceptible gene pools from F3 homologous progeny of Hongyoumai×Yumai13.250 pairs of primers were employed to screen the two parents and two pools. Primers which can amplify the same polymorphic bands between parents and pools will be further selected to detect an F2 population. Mapmaker/EXP (VERTION3.0b) analysis was then applied, and the results revealed three markers identified to be linked to Pmhym, the order of which was:Xwmc526-31.6cM - Xgwm577 - 25.6cM - Pmhym - 14.3cM - Xwmc232. Three markers were found to be located on the long arm of 7B chromosome, which indicated that gene Pmhym might be also located on chromosome 7BL.
2. Identification of differential expressed genes induced by Blumeria graminis using genechip.
Four F3 homologous resistant progeny of Hongyoumai×Yuma13 were selected as the resistance pool, with four susceptible progeny as the susceptible pool. Two treatments were set for both of the two pools:uninoculated,24 h after inoculation by Blumeria graminis. Data obtained from hybridization of cRNAs of the samples crossed with Affymetrix wheat genechip were normalized, and three groups were categorized including pre- and post-induction of the resistant, pre- and post-induction of the susceptible, and post-induction of the resistant and susceptible, respectively. Base on the differentially expressed genes,22 primer sets were designed and used for qRT-PCR validation. The qRT-PCR confirmed the reliability and reproducibility of the data revealed by genechip.
The gene measure up to the following criterions:P (present) hybridization sigal in tester, change p-value<0.001 and ratio≥2 or≤0.5, it will be considered as differentially expression. 2728 differential genes were identified between uninoculated and infected 24 h in resistant plants,39.71 percent genes are disease/defense ones in 408 genes with known function.2631 differential genes were identified between uninoculated and infected 24 h in susceptible plants,37.33 percent genes are disease/defense ones in 442 genes with known function. It indicated that metabolisms related to resistance were induced by Blumeria graminis.
Comparison of resistance- and susceptibility-related expression profiling in resistant and susceptible wheat infected by Bgt, indicated that three categories of induced transcripts were obtained including resistance-specific, susceptibility-specific and basal defense transcripts. It was found that SA (salicylic acid), JA (jasmonate) and ET (ethylene) signaling pathways related genes were differentially expressed, thus implied that the three signaling pathways might be involved in the wheat defense response against Bgt infection. In the susceptible plants, genes involved in ABA (Abscisic acid) signaling pathway might be inhibitive to the abovementioned three pathways, thus resulting in susceptible reaction. The number of genes involved in ROS (Reactive oxygen species) metabolism was more in the resistant plants, indicating possible involvement in resistance reaction. And genes involved in disease/defense, signal transduction were generally up-regulated, suggesting their implication in defense response.
3. Cloning of a wheat germin-like protein gene
One probes Ta.169.1.S1_x_at up-regulated in the inoculated resistant plants was selected for in silicon extension. Primers were then designed based on in silico cloned sequence to amplify cDNA of the corresponding gene. One germin-like protein encoded gene named TaGLP5 (855 bp) with a full open reading frame was obtained, and deposited in GenBank with accession of FJ594470. BLASTX analysis demonstrated that the encoded TaGLP5 protein belonged to a Cupin2 superfamily, with 15 homologous sequences (50-63%). The analyses of multiple alignment and phylogenetic tree indicated that TaGLP5 belongs to different branches from known germin-like protein. TaGLP5 was located on chromosome 5A with a PCR-based method using a set of Nulli-tetrasomic lines of "Chinese Spring". qRT-PCR analysis showed that TaGLP5 transcript was induced in response to Bgt infection, with higher expression in resistant plants than in the susceptible pre-24 h inoculation. It was speculated that TaGLP5 might be involved in wheat defense response against Bgt.
农家小麦品种中蕴含着丰富的抗病资源,对于发现抗白粉病品种,寻找新的抗病基因具有重要的利用价值。红蚰麦是一农家品种,具有优良的白粉病抗性,遗传分析表明其携带一对显性抗白粉病基因。本文通过SSR标记的方法对该基因进行了染色体定位。同时利用Affymetrix基因芯片结合实时定量PCR验证,分析了红蚰麦／豫麦13F3代纯合抗病家系、感病家系白粉菌诱导条件下的基因表达谱。并根据基因芯片的结果选择一个在抗病家系中诱导后高上调表达的探针进行了电子克隆和表达分析。主要的研究内容和研究结果如下：
1.抗白粉病基因Pmhym的标记定位
根据集团分离分析法在红蚰麦与感病亲本豫麦13的F3代纯合家系中建立抗、感病DNA池。选取位于21条染色体上的250对SSR引物在双亲以及DNA池间进行初筛,筛选出的能扩增出一致性差异片段的引物用F2代群体验证。用软件Mapmaker3.0软件分析结果,最终找到了3个与红蚰麦中抗白粉病基因Pmhym(暂命名)连锁的标记,标记排序如下Xwmc526-31.6cM-Xgwm577-25.6cM-Pmhym-14.3cM-Xwmc232,由于这三个标记都位于7BL染色体上,于是将红蚰麦携带的抗白粉病基因定位在7BL染色体臂上。
2.利用基因芯片筛选白粉菌诱导条件下差异表达基因。
分别以F3 4个纯合抗病家系构建抗病池,4个感病家系构建(?)病池。对两份材料同时设置两种接种处理：未接菌、接菌24h。样品的cRNA与Affymetrix 900493芯片进行杂交,获得的数据均一化处理后,设置了抗病诱导前后、感病诱导前后、诱导后抗病和感病,三组比较。挑选比较后表达倍数有差异的探针序列合成了22对引物,对基因芯片的结果进行了定量PCR验证,结果表明芯片的数据具有可重复性。
以实验组杂交信号为P,change p-value<0.001且ratio值≥2或≤0.5为标准筛选差异表达基因。筛选到2728个在抗病株系诱导前后差异表达的基因,其中已知功能的408个基因中抗病／防御类基因占39.71%。2631个基因在感病诱导前后差异表达,其中已知功能的442个基因中抗病／防御类基因占37.33%,说明病原菌诱导了植物的抗病代谢途径。
通过抗病诱导表达谱和感病诱导表达谱的比较分析、抗病诱导比感病诱导上调表达序列的分析获得了抗病特异诱导转录本,感病特异诱导转录本,以及基础防御转录本以及抗病增强表达转录本。通过差异基因的比较分析推测水杨酸、茉莉酸和乙烯途径协同作用参与了小麦对白粉菌的抗病过程；而感病株系中脱落酸途径可能与以上的三个途径存在拮抗作用,参与了植物-病原物的亲和互作反应。在抗病株系中或活性氧的代谢比感病株系中强,参与了植物的抗病反应过程。许多抗病／防御类基因、信号传导基因上调表达,这些基因可能也参与了抗病反应。
3.一个小麦类萌发素蛋白基因的克隆、表达分析
选取在抗病株系诱导后高上调表达的探针Ta.169.1.S1_x_at,进行电子延伸后合成引物扩增经白粉菌诱导的cDNA模板。结果获得了一个长855 bp,具有完整ORF框的类萌发素蛋白基因,根据ORF框翻译的蛋白序列,在NCBI上进行BLAST分析,表明该片段属于Cupin2超家族成员,有15个相似性在50-64%之间的蛋白序列。用MEGA 4.0软件构建的遗传进化树分析同源关系,本试验中获得的小麦类萌发素蛋白与已知的类萌发素蛋白分属于不同的进化分支,可能是一个新的基因,命名为TaGLP5。序列已提交至Genbank,登录号为FJ594470。用白粉病同时接种抗病、感病株系,发现该基因在感、抗植株中受白粉菌诱导后都上调表达。但上调的模式、程度不同,且在接种24 h内抗病植株中的表达量较感病植株中高。由于病原菌与植物互作的前24 h是决定病原菌能否在植物中定殖的关键时期,推测TaGLP5可能参与了红蚰麦对白粉菌早期的抗病防御反应。
Wheat powdery mildew caused by Blumeria graminis f. sp. tritici is a devastating disease in wheat growing countries worldwide. Resistance cultivars offer the most effective and safe approach to control the disease. However, the use of single or few Pm genes eventually leads to a rapid development of correponding virulence genes in the pathogen, resulting in breakdown of cultivars'resistance. Thus, it is foundational to identify novel Pm genes and dissect resistance mechanism for the control of wheat powdery mildew.
Wheat landrace may contain rich resistance genes, which is very valuable to explore resistant cultivars and identify new resistance genes against powdery mildew. Hongyoumai, one wheat landrace, bears good resistance to powdery mildew. Genetic analysis revealed that it carried one dominant Pm gene. In the present thesis, SSR markers were employed to locate Pmhym gene. Meanwhile, Affymetrix genechip analyses were employed for expression profiling in response to Blumeria graminis f. sp. tritici infection in homologous resistant and susceptible F3 progeny derived from Hongyoumai and Yumai13, respectively, along with verification of differentially expressed genes using real time quantitative RT-PCR (qRT-PCR) analyses. Based on the genechip results, one up-regulated target gene in resistance progeny was further in silico cloned and characterized. The main contents and results were as follows:
1. Chromosome location of Pmhym gene
Bulked segregate analysis (BSA) was used to establish the resistant and susceptible gene pools from F3 homologous progeny of Hongyoumai×Yumai13.250 pairs of primers were employed to screen the two parents and two pools. Primers which can amplify the same polymorphic bands between parents and pools will be further selected to detect an F2 population. Mapmaker/EXP (VERTION3.0b) analysis was then applied, and the results revealed three markers identified to be linked to Pmhym, the order of which was:Xwmc526-31.6cM - Xgwm577 - 25.6cM - Pmhym - 14.3cM - Xwmc232. Three markers were found to be located on the long arm of 7B chromosome, which indicated that gene Pmhym might be also located on chromosome 7BL.
2. Identification of differential expressed genes induced by Blumeria graminis using genechip.
Four F3 homologous resistant progeny of Hongyoumai×Yuma13 were selected as the resistance pool, with four susceptible progeny as the susceptible pool. Two treatments were set for both of the two pools:uninoculated,24 h after inoculation by Blumeria graminis. Data obtained from hybridization of cRNAs of the samples crossed with Affymetrix wheat genechip were normalized, and three groups were categorized including pre- and post-induction of the resistant, pre- and post-induction of the susceptible, and post-induction of the resistant and susceptible, respectively. Base on the differentially expressed genes,22 primer sets were designed and used for qRT-PCR validation. The qRT-PCR confirmed the reliability and reproducibility of the data revealed by genechip.
The gene measure up to the following criterions:P (present) hybridization sigal in tester, change p-value<0.001 and ratio≥2 or≤0.5, it will be considered as differentially expression. 2728 differential genes were identified between uninoculated and infected 24 h in resistant plants,39.71 percent genes are disease/defense ones in 408 genes with known function.2631 differential genes were identified between uninoculated and infected 24 h in susceptible plants,37.33 percent genes are disease/defense ones in 442 genes with known function. It indicated that metabolisms related to resistance were induced by Blumeria graminis.
Comparison of resistance- and susceptibility-related expression profiling in resistant and susceptible wheat infected by Bgt, indicated that three categories of induced transcripts were obtained including resistance-specific, susceptibility-specific and basal defense transcripts. It was found that SA (salicylic acid), JA (jasmonate) and ET (ethylene) signaling pathways related genes were differentially expressed, thus implied that the three signaling pathways might be involved in the wheat defense response against Bgt infection. In the susceptible plants, genes involved in ABA (Abscisic acid) signaling pathway might be inhibitive to the abovementioned three pathways, thus resulting in susceptible reaction. The number of genes involved in ROS (Reactive oxygen species) metabolism was more in the resistant plants, indicating possible involvement in resistance reaction. And genes involved in disease/defense, signal transduction were generally up-regulated, suggesting their implication in defense response.
3. Cloning of a wheat germin-like protein gene
One probes Ta.169.1.S1_x_at up-regulated in the inoculated resistant plants was selected for in silicon extension. Primers were then designed based on in silico cloned sequence to amplify cDNA of the corresponding gene. One germin-like protein encoded gene named TaGLP5 (855 bp) with a full open reading frame was obtained, and deposited in GenBank with accession of FJ594470. BLASTX analysis demonstrated that the encoded TaGLP5 protein belonged to a Cupin2 superfamily, with 15 homologous sequences (50-63%). The analyses of multiple alignment and phylogenetic tree indicated that TaGLP5 belongs to different branches from known germin-like protein. TaGLP5 was located on chromosome 5A with a PCR-based method using a set of Nulli-tetrasomic lines of "Chinese Spring". qRT-PCR analysis showed that TaGLP5 transcript was induced in response to Bgt infection, with higher expression in resistant plants than in the susceptible pre-24 h inoculation. It was speculated that TaGLP5 might be involved in wheat defense response against Bgt.
引文
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