玉米抗大斑病Ht2相关基因的差异表达分析
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摘要
玉米大斑病是玉米生产中最重要的病害之一,近年来大斑病的发生程度逐年加重,导致局部玉米减产,严重威胁玉米的生产。分析其主要原因是抗病品种抗病性丧失,而栽培抗性品种是防治病害发生最经济有效的方法。迄今对大斑病抗性基因的研究多局限于染色体定位和分子标记研究,有关大斑病抗性基因的生物学功能并不清楚,抗性表达过程中的重要事件不了解,因此给深入研究Ht基因的工作带来障碍。本研究旨在利用cDNA-AFLP技术,分析Ht2基因背景下玉米与HT毒素及大斑病菌互作后的差异表达谱,寻找与Ht2相关的基因,为深入研究抗大斑病基因及抗病分子机制提供依据,并最终服务于玉米抗大斑病的分子育种。本研究获得以下结果:
鉴定了东北春玉米区的48个大斑病菌分离物,共鉴定出15个小种类型,1、0号小种较其它小种出现比例高,N和1N号小种有上升趋势,玉米大斑病病菌的优势小种已不明显,病菌呈现小种变异复杂化和小种类型多样化的群体格局。毒力分析表明,能克服1、2、3和4个抗性基因大斑病菌分别占41.7%、29.1%、10.4%和2.1%。
利用玉米离体叶片法对48个分离物的毒素粗提液进行了生物测定,其中43个的鉴定结果与生理小种鉴定结果基本一致,筛选出3个了对黄早四和黄早四Ht1严重致病,而对黄早四Ht2完全不致病的HT毒素,J15、J22和H7;2个对黄早四和黄早四Ht2严重致病,而对黄早四Ht1完全不致病的HT毒素,J13和L5。
建立一套适用于玉米的cDNA-AFLP技术体系,从1024对MseI+3和PstI+2的引物组合中筛选了70对指纹图谱清晰,多态性好,重复性好的引物组合。
对接种毒素后的黄早四和黄早四Ht2进行cDNA-AFLP分析,发现21个转录表达片段在Ht2背景下特异表达。按同源基因的功能分为3类,其中接种3 h表达的基因主要和基础能量代谢以及抗逆相关基因中的信号物质有关,6 h表达的基因主要是转录因子类基因和部分抗逆基因,接种12–72 h之间只有少数基因表达。利用MaizeGDB Blast对其进行了染色体定位分析,并发现了1个基因簇,位于bin5.03。克隆了H8的全长基因,命名为ZmHP,登录号FJ600319。ZmHP基因的表达量在接毒素后6 h达到高峰,其余时间与未接种对照植株的表达水平基本一致。
构建了Ht2基因背景下玉米对大斑病菌非亲和性1号小种的基因表达谱,识别了一批与Ht2相关的高表达基因。按同源基因的功能分为8类,在非亲和互作前期主要是一些信号相关基因的表达,接种后12 h表达的基因大多是各种抗病反应初期相关基因,与抗病有关的基因大多数在接种后48–72 h表达,生长发育类基因在非亲和互作的不同阶段均表现作用,后期主要是蛋白质代谢基因的表达。染色体定位分析发现了4个基因簇,并发现5个TDFs位于已报道的Ht2定位区域内bin8.04–8.05/8.06。
对病原菌处理和毒素处理后在Ht2背景下特异表达的片段进行了比较,发现:1)玉米受到大斑病菌胁迫后的抗性反应速度较受毒素胁迫时的慢,但抗性机制较复杂。2)两种互作模式中的抗病过程基本相同。3)bin 3.04是黄早四Ht2和病原菌非亲和小种互作的差异条带特有的基因簇,同时发现了两种互作模式的差异条带在bin 5.03处都含有基因簇。4)发现3对功能或序列相似的TDFs。
利用5' RACE技术克隆了ZmQM基因,登录号FJ600320。RT-PCR分析发现,与对照相比, ZmQM在接种病原菌12~24 h和接种毒素6~12 h的黄早四Ht2中表达量均上调。因此,推测ZmQM在Ht2基因介导的抗性反应过程中发挥重要作用。
构建了ZmQM基因的表达载体,为进一步研究ZmQM基因在抗玉米大斑病中的作用做好了准备。
Northern corn leaf blight (NCLB), caused by Exserohilum turcicum (Pass.) Leonard et Suggs, is a prevalent foliar disease occurred in most maize (Zea mays L.) production regions worldwide. Yield losses of susceptible cultivars due to occurrence of NCLB can be as high as 30% when the disease is severe. The most effective measure for controlling NCLB is application of host genetic resistance. Since 1961, six do minant genesHt1,Ht3 , HtN,HtM andHtP have been reported. Understanding of resistance to NCLB conferred byHt genes is important for improving maize against the disease.However, the information of the molecular characters ofHt genes orHt-relative genes is not available. This study was carried out to identify theHt2-related genes in RNA level and to provide the basis on studying geneHt2 and understanding the molecular mechanism of resistance to north corn leaf blight. The main results were as follow:
1. Using single spore isolating technique, 48 E.turcicum strains were purified from northeast provinces of China and the physiological races were analyzed by using differential host. Fifteen physiological races including 0、1、2、3、N、12、13、1N、23、2N、3N、12N、123N、123、23N were found. Race 0 and 1 were identified more frequently than others and the ratio of N and 1N are increasing, but there are no obvious do minant races in northeast provinces of China. About 41.7 percent E.turcicum can infect corn with one geneHt and 29.1percent, 10.4 percent and 2.1percent can infect corn with two, three and four geneHt respectively.
2. Based on the biological assay in vitro, the virulence of 48HT-toxins were identified. The result of virulence analysis fundamentally agreed with that of physiological races identification. At the same time J15, J22 andH7HT-toxins which induced necrotic spot on leaves ofHZS andHZSHt1, but no spot was present onHZSHt2 were selected. J13 and L5HT-toxins induced necrotic spot on leaves ofHZS andHZSHt2, but no spot was present onHZSHt1 were also selected.
3. cDNA-AFLP reaction system of corn was established. Seventy out of 1024 prime combinations with clear polymorphic AFLP fingerprints were screened
4. cDNA-AFLP was conducted to analyze differential expression ofHt2-related genes between near-isogenic lines (NILs) Huangzaosi (HZS) andHuangzaosiHt2 (HZSHt2) under crudeHT-toxin treatment. Twenty-one transcript-derived fragments (TDFs), designatedH1 toH21, were specifically expressed inHZSHt2. BLAST analysis through the GenBank database indicated that the 21 TDFs identified can be separated into 4 groups. Basal energy metabolism gene and Signal transduction gene expressed at 3 h after inoculation. Transcription factor related gene and part of defiance gene expressed at 3 h after inoculation. Few genes expressed between 12 to 72 h. A gene cluster was found on bin 5.03 by MaizeGDB Blast analysis. The full length cDNA ofH8 (H8 cDNA, GenBank accession number FJ600319) was cloned and sequenced. RT-PCR analysis demonstrated thatH8 was up-regulated inHZSHt2 only at 6 h after inoculation withHT-toxin. The deduced protein encoded byH8 cDNA is 77% similar to Plus-3 domain containing protein which is found in yeast Rtf1.the the mechanism of geneHt2 regulate the resistance toHT-toxin is may similar to translation elongation factor.
5.eventy-six TDFs, designated Ex01 to Ex76, were specifically expressed inHZSHt2 by cDNA-AFLP analysis between near-isogenic lines (NILs)Huangzaosi (HZS) andHuangzaosiHt2 (HZSHt2) after inoculation with race 1 of Exserohilum turcicum. BLAST analysis showed that 52 of them were homologous to the genes in the GenBank database. The TDFs with significant protein homology were classified into eight functional categories: basal energy metabolism, transmembrane transport protein, defense/resistance protein, protein metabolism, protein of chromosome and proteins of DNA replication and transcription, signal transduction, growing development mediator and ranscription factor. Most of these genes were associated with basal energy metabolism and defence/resistance gene.Analyzed the space-time trait of 52 TDFs showed that those ranges from chromosome 1 to 9. Signal transduction genes expressed at 6 h after inoculation and defense/resistance genes star to express at 48~72 h.Ht2-related gene expression profiling in response to inoculation of E. turcicum were analyzed by means of cDNA-AFLP. The identification ofHt2-related genes in the present study are important to further study ofHt2 gene and the mechanism of resistance to E. turcicum in maize.
6. A full-length QM-like cDNA (designated ZmQM) was cloned from maize (Zea mays L.) leaf tissues using cDNA amplified fragment length polymorphism (cDNA-AFLP) and rapid amplification of cDNA ends (RACE) techniques. The expression of ZmQM was exa mined in leaves of theHt2 isogenic linesHuangzaosi andHuangzaosiHt2 carrying geneHt2 for resistance to northern corn leaf blight after inoculation with race 1 of Exserohilum turcicum. The nucleotide sequence of ZmQM shared 66–92% identity to QM genes isolated from other species. RT-PCR analysis showed that the expression of gene ZmQM was up-regulated inHuangzaosiHt2 at 12 h after inoculation with race 1 of E. turcicum and at 6 h after inoculation withHT-toxin compared withHuangzaosi. the mechanism of geneHt2 regulate the resistance toHT-toxin or E. turcicum by up-regulated QM-like protein (ZmQM) and then active the pathway of QM-like such as NIK-mediated antiviral signaling pathway.
7. For further function analysis on the ZmQM, the expression vector containing ZmQM were constructed.
鉴定了东北春玉米区的48个大斑病菌分离物,共鉴定出15个小种类型,1、0号小种较其它小种出现比例高,N和1N号小种有上升趋势,玉米大斑病病菌的优势小种已不明显,病菌呈现小种变异复杂化和小种类型多样化的群体格局。毒力分析表明,能克服1、2、3和4个抗性基因大斑病菌分别占41.7%、29.1%、10.4%和2.1%。
利用玉米离体叶片法对48个分离物的毒素粗提液进行了生物测定,其中43个的鉴定结果与生理小种鉴定结果基本一致,筛选出3个了对黄早四和黄早四Ht1严重致病,而对黄早四Ht2完全不致病的HT毒素,J15、J22和H7;2个对黄早四和黄早四Ht2严重致病,而对黄早四Ht1完全不致病的HT毒素,J13和L5。
建立一套适用于玉米的cDNA-AFLP技术体系,从1024对MseI+3和PstI+2的引物组合中筛选了70对指纹图谱清晰,多态性好,重复性好的引物组合。
对接种毒素后的黄早四和黄早四Ht2进行cDNA-AFLP分析,发现21个转录表达片段在Ht2背景下特异表达。按同源基因的功能分为3类,其中接种3 h表达的基因主要和基础能量代谢以及抗逆相关基因中的信号物质有关,6 h表达的基因主要是转录因子类基因和部分抗逆基因,接种12–72 h之间只有少数基因表达。利用MaizeGDB Blast对其进行了染色体定位分析,并发现了1个基因簇,位于bin5.03。克隆了H8的全长基因,命名为ZmHP,登录号FJ600319。ZmHP基因的表达量在接毒素后6 h达到高峰,其余时间与未接种对照植株的表达水平基本一致。
构建了Ht2基因背景下玉米对大斑病菌非亲和性1号小种的基因表达谱,识别了一批与Ht2相关的高表达基因。按同源基因的功能分为8类,在非亲和互作前期主要是一些信号相关基因的表达,接种后12 h表达的基因大多是各种抗病反应初期相关基因,与抗病有关的基因大多数在接种后48–72 h表达,生长发育类基因在非亲和互作的不同阶段均表现作用,后期主要是蛋白质代谢基因的表达。染色体定位分析发现了4个基因簇,并发现5个TDFs位于已报道的Ht2定位区域内bin8.04–8.05/8.06。
对病原菌处理和毒素处理后在Ht2背景下特异表达的片段进行了比较,发现:1)玉米受到大斑病菌胁迫后的抗性反应速度较受毒素胁迫时的慢,但抗性机制较复杂。2)两种互作模式中的抗病过程基本相同。3)bin 3.04是黄早四Ht2和病原菌非亲和小种互作的差异条带特有的基因簇,同时发现了两种互作模式的差异条带在bin 5.03处都含有基因簇。4)发现3对功能或序列相似的TDFs。
利用5' RACE技术克隆了ZmQM基因,登录号FJ600320。RT-PCR分析发现,与对照相比, ZmQM在接种病原菌12~24 h和接种毒素6~12 h的黄早四Ht2中表达量均上调。因此,推测ZmQM在Ht2基因介导的抗性反应过程中发挥重要作用。
构建了ZmQM基因的表达载体,为进一步研究ZmQM基因在抗玉米大斑病中的作用做好了准备。
Northern corn leaf blight (NCLB), caused by Exserohilum turcicum (Pass.) Leonard et Suggs, is a prevalent foliar disease occurred in most maize (Zea mays L.) production regions worldwide. Yield losses of susceptible cultivars due to occurrence of NCLB can be as high as 30% when the disease is severe. The most effective measure for controlling NCLB is application of host genetic resistance. Since 1961, six do minant genesHt1,Ht3 , HtN,HtM andHtP have been reported. Understanding of resistance to NCLB conferred byHt genes is important for improving maize against the disease.However, the information of the molecular characters ofHt genes orHt-relative genes is not available. This study was carried out to identify theHt2-related genes in RNA level and to provide the basis on studying geneHt2 and understanding the molecular mechanism of resistance to north corn leaf blight. The main results were as follow:
1. Using single spore isolating technique, 48 E.turcicum strains were purified from northeast provinces of China and the physiological races were analyzed by using differential host. Fifteen physiological races including 0、1、2、3、N、12、13、1N、23、2N、3N、12N、123N、123、23N were found. Race 0 and 1 were identified more frequently than others and the ratio of N and 1N are increasing, but there are no obvious do minant races in northeast provinces of China. About 41.7 percent E.turcicum can infect corn with one geneHt and 29.1percent, 10.4 percent and 2.1percent can infect corn with two, three and four geneHt respectively.
2. Based on the biological assay in vitro, the virulence of 48HT-toxins were identified. The result of virulence analysis fundamentally agreed with that of physiological races identification. At the same time J15, J22 andH7HT-toxins which induced necrotic spot on leaves ofHZS andHZSHt1, but no spot was present onHZSHt2 were selected. J13 and L5HT-toxins induced necrotic spot on leaves ofHZS andHZSHt2, but no spot was present onHZSHt1 were also selected.
3. cDNA-AFLP reaction system of corn was established. Seventy out of 1024 prime combinations with clear polymorphic AFLP fingerprints were screened
4. cDNA-AFLP was conducted to analyze differential expression ofHt2-related genes between near-isogenic lines (NILs) Huangzaosi (HZS) andHuangzaosiHt2 (HZSHt2) under crudeHT-toxin treatment. Twenty-one transcript-derived fragments (TDFs), designatedH1 toH21, were specifically expressed inHZSHt2. BLAST analysis through the GenBank database indicated that the 21 TDFs identified can be separated into 4 groups. Basal energy metabolism gene and Signal transduction gene expressed at 3 h after inoculation. Transcription factor related gene and part of defiance gene expressed at 3 h after inoculation. Few genes expressed between 12 to 72 h. A gene cluster was found on bin 5.03 by MaizeGDB Blast analysis. The full length cDNA ofH8 (H8 cDNA, GenBank accession number FJ600319) was cloned and sequenced. RT-PCR analysis demonstrated thatH8 was up-regulated inHZSHt2 only at 6 h after inoculation withHT-toxin. The deduced protein encoded byH8 cDNA is 77% similar to Plus-3 domain containing protein which is found in yeast Rtf1.the the mechanism of geneHt2 regulate the resistance toHT-toxin is may similar to translation elongation factor.
5.eventy-six TDFs, designated Ex01 to Ex76, were specifically expressed inHZSHt2 by cDNA-AFLP analysis between near-isogenic lines (NILs)Huangzaosi (HZS) andHuangzaosiHt2 (HZSHt2) after inoculation with race 1 of Exserohilum turcicum. BLAST analysis showed that 52 of them were homologous to the genes in the GenBank database. The TDFs with significant protein homology were classified into eight functional categories: basal energy metabolism, transmembrane transport protein, defense/resistance protein, protein metabolism, protein of chromosome and proteins of DNA replication and transcription, signal transduction, growing development mediator and ranscription factor. Most of these genes were associated with basal energy metabolism and defence/resistance gene.Analyzed the space-time trait of 52 TDFs showed that those ranges from chromosome 1 to 9. Signal transduction genes expressed at 6 h after inoculation and defense/resistance genes star to express at 48~72 h.Ht2-related gene expression profiling in response to inoculation of E. turcicum were analyzed by means of cDNA-AFLP. The identification ofHt2-related genes in the present study are important to further study ofHt2 gene and the mechanism of resistance to E. turcicum in maize.
6. A full-length QM-like cDNA (designated ZmQM) was cloned from maize (Zea mays L.) leaf tissues using cDNA amplified fragment length polymorphism (cDNA-AFLP) and rapid amplification of cDNA ends (RACE) techniques. The expression of ZmQM was exa mined in leaves of theHt2 isogenic linesHuangzaosi andHuangzaosiHt2 carrying geneHt2 for resistance to northern corn leaf blight after inoculation with race 1 of Exserohilum turcicum. The nucleotide sequence of ZmQM shared 66–92% identity to QM genes isolated from other species. RT-PCR analysis showed that the expression of gene ZmQM was up-regulated inHuangzaosiHt2 at 12 h after inoculation with race 1 of E. turcicum and at 6 h after inoculation withHT-toxin compared withHuangzaosi. the mechanism of geneHt2 regulate the resistance toHT-toxin or E. turcicum by up-regulated QM-like protein (ZmQM) and then active the pathway of QM-like such as NIK-mediated antiviral signaling pathway.
7. For further function analysis on the ZmQM, the expression vector containing ZmQM were constructed.
引文
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