利用大麦基因芯片筛选及克隆簇毛麦抗白粉病相关基因
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摘要
小麦(T．aestivum L)作为世界性的重要粮食作物，在农业生产中具有举足轻重的地位，但是其生产却受到很多生物胁迫和非生物胁迫的影响。由专性寄生真菌小麦白粉病菌(Erysiphegraminis DC)引起的小麦白粉病(Ergsiphe greminis f．sp．tritici)是中国和世界上许多国家小麦生产中危害日趋严重的病害之一。鉴别新的抗白粉病基因并研究它们的抗性机制变得越来越重要。簇毛麦中的抗白粉病基因Pm21被定位于染色体6V的短臂上，对白粉病抗性强、抗谱广。一方面，研究Pm21基因发挥抗病作用过程中所涉及的转录因子、信号传导途径和重要防卫反应基因，对于研究广谱抗性机理很有意义；另一方面，克隆抗病基因对今后利用基因工程手段提高小麦对白粉病的抗性具有重要意义。
小麦-簇毛麦易位系(6VS／6AL)已经在育种和生产上得到广泛应用，快速准确地鉴定6VS／6AL这条易位染色体的纯、杂合状态，可以提高育种过程中分子标记辅助选择的效率。由于以往开发的与Pm21连锁的分子标记都是显性标记，所以开发一个与Pm21基因连锁的共显性分子标记具有重要应用价值。另外，簇毛麦具有很多优良性状，通过细胞工程手段已经创造了很多异染色体系，所以开发一种能够在小麦背景中快速鉴定簇毛麦染色质的分子标记，可以大大提高鉴定效率。
1、利用大麦基因芯片筛选簇毛麦抗白粉病相关基因及Pm21基因抗性机理研究
为了研究Pm21基因对白粉病的抗性机理，利用经白粉菌诱导抗、感簇毛麦和非诱导的抗病簇毛麦为材料，借助于基因芯片分析手段进行分析。簇毛麦的cRNA与大麦基因芯片杂交，平均有5741个探针有杂交信号。通过比较抗病簇毛麦经白粉菌诱导前后的表达谱，筛选出簇毛麦经白粉菌诱导表达的基因，包括病程相关蛋白、防卫反应基因、转录因子、信号传导因子和抗病基因类似物等。通过比较诱导的抗病簇毛麦的表达谱与诱导的感病簇毛麦的表达谱，筛选出两者的差异表达基因。对这些差异表达基因的信息进行研究的结果表明，抗病簇毛麦中防卫反应受水杨酸和乙烯途径共同调控，而感病簇毛麦中茉莉酸信号途径被诱导，水杨酸途径受到了抑制。在簇毛麦的抗病过程中，水杨酸途径是一种最有效的信号传导途径。
2、抗病相关基因的克隆
根据基因芯片的杂交结果，选择了一些上调表达探针进行后续分析。Contig17515在白粉菌诱导的抗病簇毛麦中的表达量比非诱导的抗病簇毛麦和诱导的感病簇毛麦中的表达量都高。以该序列为模板设计引物，经RT-PCR克隆到一个定位于簇毛麦6VS上的丝氨酸／苏氨酸激酶基因Hv-S／TPK(cDNA)，进一步用RACE方法克隆到基因全长。该全长基因共有1376个碱基，编码的310个氨基酸中包含了丝氨酸／苏氨酸激酶的两个保守域(DaKXXN和GTAGYXAPCN／E)。用同一对引物把簇毛麦gDNA中的基因也克隆出来，并定名为Hv-S／TPK(gDNA)。经研究发现来自6VS染色体组的基因Hv-S／TPK(gDNA)与来自6AS、6BS、6DS染色体组的基因Ta-A-S／TPK(gDNA)、Ta-B-S／TPK(gDNA)、Ta-D-S／TPK(gDNA)是直系同源基因，Hv-S／TPK(gDNA)在簇毛麦和易位系6VS／6AL中均能表达，且该基因在B和D染色体组上的直系同源基因Ta-B-S／TPK(gDNA)和Ta-D-S／TPK(gDNA)在易位系6VS／6AL中也能表达。通过对Hv-S／TPK(cDNA)、Ta-B-S／TPK(cDNA)和Ta-D-S／TPK(cDNA)的核苷酸序列推导出的氨基酸序列进行分析，发现有4个氨基酸残基不同，其中的1处是疏水氨基酸和极性氨基酸之间的变化。
用根据Contig17515设计的引物进行PCR，可以从普通小麦的6AS、6BS、6DS和簇毛麦的6VS上扩增出大小不同的PCR条带。用该对引物在(扬麦5号X易位系)F_2群体中进行扩增，发现凡是田间鉴定抗病的单株都能扩增出6VS上的902bp的条带，感病单株则扩增不出这条特异带。另外抗病单株的扩增带型还分两种类型，一种类型的单株扩增不出6AS上的984bp条带，这种单株中的6VS／6AL染色体处于纯合状态；一种类型的单株可以扩增出6AS上的984bp条带，这种单株中的6VS／6AL染色体处于杂合状态。上述结果表明，本研究开发出一种基于PCR的与Pm21连锁的共显性分子标记，把该标记定名为NAU／xibao15_(902)。另外由于该引物只是从第六部分同源群染色体上扩增出大小不同的条带，所以可以用该引物进行PCR，鉴定涉及第六部分同源群染色体发生易位、缺失或添加的异染色体系。
基因芯片杂交结果表明，探针contig3156在抗、感簇毛麦中受白粉菌诱导的程度都最大，并且在抗病易位系中该探针的杂交信号比在感病扬麦5号中的杂交信号强。半定量RT-PCR结果表明：在抗、感簇毛麦和抗、感普通小麦中contig3156这个探针都受白粉菌诱导表达，但在抗病材料中比在感病材料中表达早或上调程度大。contig3156这个探针是一个推导的草酸盐氧化酶基因(Oxalate Oxydase Like Protein，OxOLP)。利用TAIL-PCR方法把簇毛麦中的Hv-OxOLP基因的全长序列克隆出，该基因包含一个内含子和两个外显子，ORF包含690个氨基酸，在启动子区包含两个W-box和一个TATA盒，在终止密码子后面还有一个polyA的加尾信号。由于Hv-OxOLP基因是一个受白粉菌诱导高度表达的基因，并且在抗、感植物材料中都上调表达，所以该基因可能是在植株识别白粉菌的非专化激发子后启动防卫反应时特异表达。鉴于Hv-OxOLP基因启动子的特点，该启动子可以在用基因工程手段改良作物抗病性中发挥作用。
3、建立在直系同源序列比较基础上的小麦进化研究
由于簇毛麦基因Hv-S／TPK(gDNA)和其在小麦中不同染色体组上的直系同源基因的序列之间有一定差异，所以比较小麦族不同物种中该直系同源基因中存在的SNPs可以进行物种进化的研究。将T.aestivum、T.turgidum、T.urartu、T.monococcum、Ae.speltoides、Ae.longissima、Ae.sharonensis、Ae.searsii、T.timopheevii、T.araraticum、A.tauschii中各个染色体组上Hv-S／TPK(gDNA)的直系同源基因片段分别克隆出来，比较了序列之间的相关性，进行小麦族几个不同物种的进化研究。T.timopheevii、T.araraticum中的A染色体组和T.aestivum、T.turgidum两者中的A染色体组之间存在一定的分化，可以分为两类；四倍体T.timopheevii中的A来自T.monococcum，而四倍体T.turgidum中的A染色体组来自T.urartu.B、G的序列与来自Ae.speltoides的序列之间的差异较小，与来自Ae.longissima、Ae.sharonensis、Ae.searsii的序列差异较大；T.timopheevii、T.araraticum两者中的两个G来源相同、T.turgidum、T.aestivum两者中的两个B来源相同，且B和G都可能来自Ae.speltoides。来自普通小麦D组的序列与来自Ae.tauschii的序列同源性非常高，为Ae.tauschii是普通小麦D组的供体提供了又一个证据。
Wheat (T.aestivum L) is one of the most important cereal crops in the world. However, the production of wheat is adversely affected by a number of biotic and abiotic stresses. Wheat powdery mildew disease, which is caused by Erysiphegraminis DC, is one of the most serious wheat diseases in China and many other countries of the world. Identifying more resistant genes to powdery mildew disease and studying the resistance mechanism become more and more important. Pm21 gene, which is located on the short arm of chromosome 6V of H. villosa, is a broad-spectrum and powerful resistant gene to the powdery mildew disease. So, on the one hand, studying the transcription factors, signal transduction pathway and important defense genes involved in the resistance progress will be valuable to investigate the mechanism of the broad-spectrum resistance; on the other hand, isolating the resistance gene will facilitate improving the level of resistance to powdery mildew in wheat through genetic engeneering.
Wheat-H.villosa translocation line (6VS/6AL) has been widely used in wheat breeding project as genetic material and some varieties containing the 6VS/6AL chromosome are now playing an important role in the wheat production. Identification the status of 6VS/6AL chromosome is tightly associated with the efficiency of the marker assisted selection. Because the marker published in the past years are all dominant markers, it is helpful to develop codominant PCR marker linked with Pm21 for the wheat breeding. Owing to the advantages of H.villosa, many genetic materials have been developed through molecular cytogenetic engineering. If a PCR marker is developed which can be used to identify the chromotin of H.villosa in the common wheat backgroud, the efficiency of screening will be improved dramaticly.
1. Identification of Genes Involved in the Resistance to the Powdery Mildew and Investigating the Resistance Mechanism of Pm21 in Haynaldia villosa Profiling Using Barley Genechip
The inoculated resistant and susceptible mutant of H.villosa and the uninoculated
resistant H. villosa were used to study the resistance mechanism of Pm21 to the powdery mildew by microarray analysis using the Barley1 genechip. About 5471 probes could hybridize with Haynaldia villosa genes in each split. Comparison of transcript profiling of inoculated sample with that of the uninoculated sample of resistant H.villosa, genes induced by Erysiphegraminis DC were identified, including pathogen related proteins, defense genes, transcription factors, signal transduction components and resistance gene analogs. Transcript profiling of the inoculated sample of the resistant H.villosa was also compared with the inoculated sample of the susceptible H.villosa mutant, the differentially expressed genes were screeded. The information of these genes indicated that salicytic acid and ethylene signal transduction pathway involved in the resistance defense pathway in resistant H.villosa, while jasmonic acid signal transduction pathway was induced instead of the salicytic acid signal transduction pathway in the susceptible H.villosa mutant. Salicytic acid signal transduction pathway may be the most effective in the powdery mildew resistance pathway of H. villosa.
2. Cloning of resistance related gene
According to the information of the upregulated genes, a few probes were analysed in detail in the following study. The expression level of the probe Contig17515 in the inoculated resistant H.villosa was higher than that in the uninoculated resistant H.villosa and than in the inoculated susceptible H.villosa. A pare of primers were designed based on the sequence of the Contig17515 and a putative serine/threonine kinase gene, named as Hv-S/TPK (cDNA), was cloned by RT-PCR. The complete sequence of Hv-S/TPK (cDNA), was obtained by RACE protocal and this gene was located on the short arm of the 6V chromosome. Hv-S/TPK (cDNA) contains 1376 nucleotides and the putative protein contains two conserved domain of serine/threonine kinase gene. The gene, named as Hv-S/TPK (gDNA), was cloned from the genomic DNA of H. villosa using the same primers. The results of southern blotting and the sequence comparison indicated that the Hv-S/TPK (gDNA) from the 6VS of H. villosa and Ta-A-S/TPK (gDNA), Ta-B-S/TPK (gDNA), Ta-D-S/TPK(gDNA) from the 6AS, 6BS and 6DS of common wheat were ortholog genes. Hv-S/TPK (gDNA) could express in both in the H. villosa and in the translocation line 6VS/6AL, more over the Ta-B-S/TPK (gDNA) and Ta-D-S/TPK(gDNA) could also espress in translocation line. According to the protein sequence of the gene Hv-S/TPK (cDNA)、Ta-B-S/TPK(cDNA) and Ta-D-S/TPK(cDNA), four differences were detected one of which involed the difference between the hydrophobic amino acid and the polar amino acid.
When the PCR was conducted using the genomic DNA of F_2 population derived from Yang5×92R137 as templates, it was found that the plants resistant to powdery mildew disease had the 902bp amplicon specific to H.villosa, while the susceptible plants didn't. Further analysis indicated that the resistant plants of F_2 population could be divided into two types: one type of plants containing one 6VS/6AL chromosome and these plant could amplify the 984bp fragment from the 6AS; another type of plants containing two 6VS/6AL chromosomes and these plants couldn't amplify the 984bp fragment. So, a codominant PCR marker Nau/xibao15_(902) was developed in this study. Since only fragments belonging to the group 6 chromosomes coule be amplified using 17515F and 17515R as primers, PCRs could be conducted using this pare of primers to detect whether the chromosomes added or translocatcd into other different species backgroud belong to the 6 homoeologous group.
The probe Contig3156 was mostly upregulated by the inoculation with Erysiphegraminis DC in the resistant and susceptible H.villosa, and its expression level in resistant 92R137 was also higher than that in the susceptible Yangmai5. Contig3156 was a putative OxOLP gene (Oxalate Oxydase Like Protein, OxOLP). The result of RT-PCR indicated that the OxOLP was induced by the Erysiphegraminis DC in all four plant matierials, but the expression level of OxOLP was higher or the OxOLP was induced earlier in the resistant plant than in the susceptible plant. The complete sequence of the Hv-OxOLP in the H.villosa was cloned by TAIL-PCR. The whole gene contained one intron and two extrons, and the ORF contained 690 nucleotides. In the promoter of Hv-OxOLP, there was a TATA box and two W-box. Following the stop code, there was a polyA signal motif. Because the gene OxOLP was upregulated by Erysiphegraminis DC both in the resistant and susceptible plant, the gene may express during the defense response after the plant recognized the general elicite of the Erysiphegraminis DC. Owing to the character of the promoter of Hv-OxOLP, it may be used in the improving the resistance to diseases of crop through genetic engineering.
3. Studying of the wheat evolution based on the sequence of the ortholog genes
Because there were differences among the sequences of the gene Hv-S/TPK (gDNA) of H.villosa and its orthologous genes in the different genomes of wheat, it was possible to study the evolution of Triticeae by analysing SNPs exsiting in the sequences from different species. The othologous genes of Hv-S/TPK (gDNA) from the different genomes of T.aestivum, T.turgidum, T.urartu, T.monococcum, Ae.speltoides, Ae.longissima, Ae.searsii, Ae.sharonensis, T.timopheevii, T.araraticum, A. tauschii were cloned and compared to study the relationship of these sequences and to investigate the evolution of the species in the Triticeae. The result indicated that the sequences of T. timopheevii and T. araraticum could be divided into one class, while the sequences of T.turgidum and T.aestivum could be divided into another. The A genome of tetraploid wheat T.timopheevii may originated from the T.monococcum, while the A genome of tetraploid wheat T.turgidum may originated from the T. urartu. The result of comparison among the sequences from the S, B, and G genomes indicated that the sequences from the Ae.searsii, Ae. longissima and Ae. sharonensis could be divided into one class, and the sequences from the G, B and the Ae.speltoides could be divided into another. T. timopheevii and T. araraticum came from a same donor and the B genome of the T. turgidum and T.aestivum came from another one. The G and B genome both may come from the S genome of Ae.speltoides. The sequence from the D genome of the T. aestivum shared high similarity with that from the D genome of the Ae. tauschii, so the above result supplied a new proof to the conclusion that the Ae. tauschii was the donor of the D genome of the T. aestivum.
小麦-簇毛麦易位系(6VS／6AL)已经在育种和生产上得到广泛应用，快速准确地鉴定6VS／6AL这条易位染色体的纯、杂合状态，可以提高育种过程中分子标记辅助选择的效率。由于以往开发的与Pm21连锁的分子标记都是显性标记，所以开发一个与Pm21基因连锁的共显性分子标记具有重要应用价值。另外，簇毛麦具有很多优良性状，通过细胞工程手段已经创造了很多异染色体系，所以开发一种能够在小麦背景中快速鉴定簇毛麦染色质的分子标记，可以大大提高鉴定效率。
1、利用大麦基因芯片筛选簇毛麦抗白粉病相关基因及Pm21基因抗性机理研究
为了研究Pm21基因对白粉病的抗性机理，利用经白粉菌诱导抗、感簇毛麦和非诱导的抗病簇毛麦为材料，借助于基因芯片分析手段进行分析。簇毛麦的cRNA与大麦基因芯片杂交，平均有5741个探针有杂交信号。通过比较抗病簇毛麦经白粉菌诱导前后的表达谱，筛选出簇毛麦经白粉菌诱导表达的基因，包括病程相关蛋白、防卫反应基因、转录因子、信号传导因子和抗病基因类似物等。通过比较诱导的抗病簇毛麦的表达谱与诱导的感病簇毛麦的表达谱，筛选出两者的差异表达基因。对这些差异表达基因的信息进行研究的结果表明，抗病簇毛麦中防卫反应受水杨酸和乙烯途径共同调控，而感病簇毛麦中茉莉酸信号途径被诱导，水杨酸途径受到了抑制。在簇毛麦的抗病过程中，水杨酸途径是一种最有效的信号传导途径。
2、抗病相关基因的克隆
根据基因芯片的杂交结果，选择了一些上调表达探针进行后续分析。Contig17515在白粉菌诱导的抗病簇毛麦中的表达量比非诱导的抗病簇毛麦和诱导的感病簇毛麦中的表达量都高。以该序列为模板设计引物，经RT-PCR克隆到一个定位于簇毛麦6VS上的丝氨酸／苏氨酸激酶基因Hv-S／TPK(cDNA)，进一步用RACE方法克隆到基因全长。该全长基因共有1376个碱基，编码的310个氨基酸中包含了丝氨酸／苏氨酸激酶的两个保守域(DaKXXN和GTAGYXAPCN／E)。用同一对引物把簇毛麦gDNA中的基因也克隆出来，并定名为Hv-S／TPK(gDNA)。经研究发现来自6VS染色体组的基因Hv-S／TPK(gDNA)与来自6AS、6BS、6DS染色体组的基因Ta-A-S／TPK(gDNA)、Ta-B-S／TPK(gDNA)、Ta-D-S／TPK(gDNA)是直系同源基因，Hv-S／TPK(gDNA)在簇毛麦和易位系6VS／6AL中均能表达，且该基因在B和D染色体组上的直系同源基因Ta-B-S／TPK(gDNA)和Ta-D-S／TPK(gDNA)在易位系6VS／6AL中也能表达。通过对Hv-S／TPK(cDNA)、Ta-B-S／TPK(cDNA)和Ta-D-S／TPK(cDNA)的核苷酸序列推导出的氨基酸序列进行分析，发现有4个氨基酸残基不同，其中的1处是疏水氨基酸和极性氨基酸之间的变化。
用根据Contig17515设计的引物进行PCR，可以从普通小麦的6AS、6BS、6DS和簇毛麦的6VS上扩增出大小不同的PCR条带。用该对引物在(扬麦5号X易位系)F_2群体中进行扩增，发现凡是田间鉴定抗病的单株都能扩增出6VS上的902bp的条带，感病单株则扩增不出这条特异带。另外抗病单株的扩增带型还分两种类型，一种类型的单株扩增不出6AS上的984bp条带，这种单株中的6VS／6AL染色体处于纯合状态；一种类型的单株可以扩增出6AS上的984bp条带，这种单株中的6VS／6AL染色体处于杂合状态。上述结果表明，本研究开发出一种基于PCR的与Pm21连锁的共显性分子标记，把该标记定名为NAU／xibao15_(902)。另外由于该引物只是从第六部分同源群染色体上扩增出大小不同的条带，所以可以用该引物进行PCR，鉴定涉及第六部分同源群染色体发生易位、缺失或添加的异染色体系。
基因芯片杂交结果表明，探针contig3156在抗、感簇毛麦中受白粉菌诱导的程度都最大，并且在抗病易位系中该探针的杂交信号比在感病扬麦5号中的杂交信号强。半定量RT-PCR结果表明：在抗、感簇毛麦和抗、感普通小麦中contig3156这个探针都受白粉菌诱导表达，但在抗病材料中比在感病材料中表达早或上调程度大。contig3156这个探针是一个推导的草酸盐氧化酶基因(Oxalate Oxydase Like Protein，OxOLP)。利用TAIL-PCR方法把簇毛麦中的Hv-OxOLP基因的全长序列克隆出，该基因包含一个内含子和两个外显子，ORF包含690个氨基酸，在启动子区包含两个W-box和一个TATA盒，在终止密码子后面还有一个polyA的加尾信号。由于Hv-OxOLP基因是一个受白粉菌诱导高度表达的基因，并且在抗、感植物材料中都上调表达，所以该基因可能是在植株识别白粉菌的非专化激发子后启动防卫反应时特异表达。鉴于Hv-OxOLP基因启动子的特点，该启动子可以在用基因工程手段改良作物抗病性中发挥作用。
3、建立在直系同源序列比较基础上的小麦进化研究
由于簇毛麦基因Hv-S／TPK(gDNA)和其在小麦中不同染色体组上的直系同源基因的序列之间有一定差异，所以比较小麦族不同物种中该直系同源基因中存在的SNPs可以进行物种进化的研究。将T.aestivum、T.turgidum、T.urartu、T.monococcum、Ae.speltoides、Ae.longissima、Ae.sharonensis、Ae.searsii、T.timopheevii、T.araraticum、A.tauschii中各个染色体组上Hv-S／TPK(gDNA)的直系同源基因片段分别克隆出来，比较了序列之间的相关性，进行小麦族几个不同物种的进化研究。T.timopheevii、T.araraticum中的A染色体组和T.aestivum、T.turgidum两者中的A染色体组之间存在一定的分化，可以分为两类；四倍体T.timopheevii中的A来自T.monococcum，而四倍体T.turgidum中的A染色体组来自T.urartu.B、G的序列与来自Ae.speltoides的序列之间的差异较小，与来自Ae.longissima、Ae.sharonensis、Ae.searsii的序列差异较大；T.timopheevii、T.araraticum两者中的两个G来源相同、T.turgidum、T.aestivum两者中的两个B来源相同，且B和G都可能来自Ae.speltoides。来自普通小麦D组的序列与来自Ae.tauschii的序列同源性非常高，为Ae.tauschii是普通小麦D组的供体提供了又一个证据。
Wheat (T.aestivum L) is one of the most important cereal crops in the world. However, the production of wheat is adversely affected by a number of biotic and abiotic stresses. Wheat powdery mildew disease, which is caused by Erysiphegraminis DC, is one of the most serious wheat diseases in China and many other countries of the world. Identifying more resistant genes to powdery mildew disease and studying the resistance mechanism become more and more important. Pm21 gene, which is located on the short arm of chromosome 6V of H. villosa, is a broad-spectrum and powerful resistant gene to the powdery mildew disease. So, on the one hand, studying the transcription factors, signal transduction pathway and important defense genes involved in the resistance progress will be valuable to investigate the mechanism of the broad-spectrum resistance; on the other hand, isolating the resistance gene will facilitate improving the level of resistance to powdery mildew in wheat through genetic engeneering.
Wheat-H.villosa translocation line (6VS/6AL) has been widely used in wheat breeding project as genetic material and some varieties containing the 6VS/6AL chromosome are now playing an important role in the wheat production. Identification the status of 6VS/6AL chromosome is tightly associated with the efficiency of the marker assisted selection. Because the marker published in the past years are all dominant markers, it is helpful to develop codominant PCR marker linked with Pm21 for the wheat breeding. Owing to the advantages of H.villosa, many genetic materials have been developed through molecular cytogenetic engineering. If a PCR marker is developed which can be used to identify the chromotin of H.villosa in the common wheat backgroud, the efficiency of screening will be improved dramaticly.
1. Identification of Genes Involved in the Resistance to the Powdery Mildew and Investigating the Resistance Mechanism of Pm21 in Haynaldia villosa Profiling Using Barley Genechip
The inoculated resistant and susceptible mutant of H.villosa and the uninoculated
resistant H. villosa were used to study the resistance mechanism of Pm21 to the powdery mildew by microarray analysis using the Barley1 genechip. About 5471 probes could hybridize with Haynaldia villosa genes in each split. Comparison of transcript profiling of inoculated sample with that of the uninoculated sample of resistant H.villosa, genes induced by Erysiphegraminis DC were identified, including pathogen related proteins, defense genes, transcription factors, signal transduction components and resistance gene analogs. Transcript profiling of the inoculated sample of the resistant H.villosa was also compared with the inoculated sample of the susceptible H.villosa mutant, the differentially expressed genes were screeded. The information of these genes indicated that salicytic acid and ethylene signal transduction pathway involved in the resistance defense pathway in resistant H.villosa, while jasmonic acid signal transduction pathway was induced instead of the salicytic acid signal transduction pathway in the susceptible H.villosa mutant. Salicytic acid signal transduction pathway may be the most effective in the powdery mildew resistance pathway of H. villosa.
2. Cloning of resistance related gene
According to the information of the upregulated genes, a few probes were analysed in detail in the following study. The expression level of the probe Contig17515 in the inoculated resistant H.villosa was higher than that in the uninoculated resistant H.villosa and than in the inoculated susceptible H.villosa. A pare of primers were designed based on the sequence of the Contig17515 and a putative serine/threonine kinase gene, named as Hv-S/TPK (cDNA), was cloned by RT-PCR. The complete sequence of Hv-S/TPK (cDNA), was obtained by RACE protocal and this gene was located on the short arm of the 6V chromosome. Hv-S/TPK (cDNA) contains 1376 nucleotides and the putative protein contains two conserved domain of serine/threonine kinase gene. The gene, named as Hv-S/TPK (gDNA), was cloned from the genomic DNA of H. villosa using the same primers. The results of southern blotting and the sequence comparison indicated that the Hv-S/TPK (gDNA) from the 6VS of H. villosa and Ta-A-S/TPK (gDNA), Ta-B-S/TPK (gDNA), Ta-D-S/TPK(gDNA) from the 6AS, 6BS and 6DS of common wheat were ortholog genes. Hv-S/TPK (gDNA) could express in both in the H. villosa and in the translocation line 6VS/6AL, more over the Ta-B-S/TPK (gDNA) and Ta-D-S/TPK(gDNA) could also espress in translocation line. According to the protein sequence of the gene Hv-S/TPK (cDNA)、Ta-B-S/TPK(cDNA) and Ta-D-S/TPK(cDNA), four differences were detected one of which involed the difference between the hydrophobic amino acid and the polar amino acid.
When the PCR was conducted using the genomic DNA of F_2 population derived from Yang5×92R137 as templates, it was found that the plants resistant to powdery mildew disease had the 902bp amplicon specific to H.villosa, while the susceptible plants didn't. Further analysis indicated that the resistant plants of F_2 population could be divided into two types: one type of plants containing one 6VS/6AL chromosome and these plant could amplify the 984bp fragment from the 6AS; another type of plants containing two 6VS/6AL chromosomes and these plants couldn't amplify the 984bp fragment. So, a codominant PCR marker Nau/xibao15_(902) was developed in this study. Since only fragments belonging to the group 6 chromosomes coule be amplified using 17515F and 17515R as primers, PCRs could be conducted using this pare of primers to detect whether the chromosomes added or translocatcd into other different species backgroud belong to the 6 homoeologous group.
The probe Contig3156 was mostly upregulated by the inoculation with Erysiphegraminis DC in the resistant and susceptible H.villosa, and its expression level in resistant 92R137 was also higher than that in the susceptible Yangmai5. Contig3156 was a putative OxOLP gene (Oxalate Oxydase Like Protein, OxOLP). The result of RT-PCR indicated that the OxOLP was induced by the Erysiphegraminis DC in all four plant matierials, but the expression level of OxOLP was higher or the OxOLP was induced earlier in the resistant plant than in the susceptible plant. The complete sequence of the Hv-OxOLP in the H.villosa was cloned by TAIL-PCR. The whole gene contained one intron and two extrons, and the ORF contained 690 nucleotides. In the promoter of Hv-OxOLP, there was a TATA box and two W-box. Following the stop code, there was a polyA signal motif. Because the gene OxOLP was upregulated by Erysiphegraminis DC both in the resistant and susceptible plant, the gene may express during the defense response after the plant recognized the general elicite of the Erysiphegraminis DC. Owing to the character of the promoter of Hv-OxOLP, it may be used in the improving the resistance to diseases of crop through genetic engineering.
3. Studying of the wheat evolution based on the sequence of the ortholog genes
Because there were differences among the sequences of the gene Hv-S/TPK (gDNA) of H.villosa and its orthologous genes in the different genomes of wheat, it was possible to study the evolution of Triticeae by analysing SNPs exsiting in the sequences from different species. The othologous genes of Hv-S/TPK (gDNA) from the different genomes of T.aestivum, T.turgidum, T.urartu, T.monococcum, Ae.speltoides, Ae.longissima, Ae.searsii, Ae.sharonensis, T.timopheevii, T.araraticum, A. tauschii were cloned and compared to study the relationship of these sequences and to investigate the evolution of the species in the Triticeae. The result indicated that the sequences of T. timopheevii and T. araraticum could be divided into one class, while the sequences of T.turgidum and T.aestivum could be divided into another. The A genome of tetraploid wheat T.timopheevii may originated from the T.monococcum, while the A genome of tetraploid wheat T.turgidum may originated from the T. urartu. The result of comparison among the sequences from the S, B, and G genomes indicated that the sequences from the Ae.searsii, Ae. longissima and Ae. sharonensis could be divided into one class, and the sequences from the G, B and the Ae.speltoides could be divided into another. T. timopheevii and T. araraticum came from a same donor and the B genome of the T. turgidum and T.aestivum came from another one. The G and B genome both may come from the S genome of Ae.speltoides. The sequence from the D genome of the T. aestivum shared high similarity with that from the D genome of the Ae. tauschii, so the above result supplied a new proof to the conclusion that the Ae. tauschii was the donor of the D genome of the T. aestivum.
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