小麦应答农杆菌转化相关基因的筛选及TaUGT在小麦与农杆菌互作过程中的功能研究
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摘要
小麦是目前世界上主要的粮食作物之一，小麦的产量与品质直接关系到粮食安全。转基因技术是目前小麦基因改良的重要方法之，其效率高低直接影响小麦遗传改良进程。备受关注的小麦基因枪转化法容易引发转基因高嵌合度、多位点整合及诱发转基因沉默等问题，农杆菌介导转化法则可很好地解决或缓解上述问题，但农杆菌介导转化小麦的效率亟待提高。本研究从小麦与农杆菌互作的分子机制出发，探索影响农杆菌介导转化小麦转化效率的功能基因，取得了如下主要结果：
1、利用抑制性差减杂交、蛋白质组学、数字化表达谱测序来筛选小麦对农杆菌侵染的应答基因，分别得到20、90和4,889个差异表达基因。其中，数字化表达谱测序灵敏度最高。差别基因聚类分析表明，参与物质代谢途径的基因所占比例最大，其次是次生代谢物质合成通路。其中，超过3%的差异表达基因参与到目前已经明确的植物-病原体互作通路，而在生物素合成通路、花生四烯酸代谢通路、光合作用生物通路差异表达基因定位最少。
2、KEGG通路分析表明，小麦UDP葡萄糖基转移酶（TaUGT）编码基因主要参与抗坏血酸代谢通路和药物代谢-细胞色素P450通路。而这两条通路都与植物抗病相关。Real-time PCR分析显示，TaUGT在小麦受到农杆菌侵染12小时后，表达量大幅上调，并随着共培养时间延伸而逐渐下降。表明该基因是在农杆菌侵染小麦细胞的初期起了较重要的调控作用。
3、基因结构分析表明，TaUGT为真核生物中少见的不存在内含子的核编码功能基因之一，与二倍体祖先种的同源性极高，进化上十分保守。经Southern杂交分析及将TaUGT基因序列向GrainGenes数据库的已定位小麦EST数据库进行blastn比对，推测TaUGT在小麦基因组中存在9个拷贝，在A、B、D基因组上各存在3个拷贝。亚细胞定位显示，TaUGT定位在细胞膜上，可跨膜发挥功能。
4、TaUGT功能分析发现，过表达TaUGT的拟南芥及小麦幼胚，农杆菌介导的GUS基因瞬时表达能力明显降低。电镜观察结果表明，过表达TaUGT后减少了农杆菌在小麦幼胚细胞表面附着的数量。利用农杆菌介导再次转化过表达TaUGT的烟草植株，转化效率明显降低。利用基因枪介导抑制小麦成熟胚愈伤组织中TaUGT的表达，增加了农杆菌侵染后小麦成熟胚愈伤组织的分化能力，但没有提高最终转化率。此外，体外真核表达的TaUGT并不影响农杆菌的生长。
综上所述，很多参与小麦正常代谢途径的基因都与小麦防御农杆菌侵染有关。TaUGT在农杆菌对小麦的侵染阶段，从农杆菌吸附小麦细胞表面到T-DNA转入小麦细胞的过程中调节小麦对农杆菌的防御反应。
Wheat is one of most important food crops worldwide, which yield and qulity affects the foodsecurity directly. Genetically modified (GM) technology is of great significance to wheat geneticalimprovement. A common limitation in wheat biotechnological breeding is the low transformationefficiency. Remarkably, gene gun-mediated transformation in wheat often triggers many seriousproblems like high degree of chimeric, multiple integration sites for foreign genes, and transgenesilencing. Howerver, Agrobacterium-mediated transformation can effectively avoid or alleviate thesedrawbacks. Still, Agrobacterium-mediated transformation efficiency in wheat needs to be improved.This study intends to screen and characterize the genes responsible for the transformation efficiencyfrom the view of molecular mechanisms underlying the interactions between wheat andAgrobacterium cells during their co-cultivation. The main results achieved in this study were asfollows:
1. According to the results of suppression subtractive hybridization (SSH), proteomics andRNA-seq,20,90, and4,889differential expressed genes were got in response to the Agrobacteriuminfection in wheat, respectively. Of the three methods above, RNA-seq is the most effective, accurate,and suitable for screening differential transcripts on the whole. Based on clustering analysis, most ofthe differential transcripts were involved in metabolic pathways; the second is in secondarymetabolism. Of those,3%participated in plant-pathogenic interaction pathway known so far.However, few were found in biotin synthesis pathway, arachidonic acid metabolism pathway, andphotosynthesis pathway.
2. KEGG pathway analysis indicated that wheat UDP-glucosyltransferase encoded gene (TaUGT)was mainly involved in ascorbic acid metabolic pathway and drug metabolism-cytochrome P450pathway. And these two pathways were related to the process of plant disease resistance. qRT-PCRshowed the expression of TaUGT upregulated at12h after Agrobacterium infection and decreasedgradually following the time span of co-culture in wheat. It inferred that TaUGT played a regulatingfactor in the initial stage of infection by Agrobacterium.
3. Structural analysis revealed that TaUGT in wheat was representative of those nuclear encodedwithout intron available in eukaryote, which showed high identity with its wild relative-orginatedspecies in evolution. Inferiring that there are nine copies were present in wheat genome based onSouthern blot analysis and the wheat mapped expression sequence tags (EST) in GrainGenes database.And each three copies are located on genome A, B, and D of wheat, respectively. Furthermore,according to the result of subcellular localization, TaUGT located on cytomembrane and function wellthrough transmembrane.
4. Functional identification of TaUGT showed that overexpression of TaUGT inhibited transientexpression of GUS in Arabidopsis and wheat after Agrobacterium infection, reduced theAgrobacterium population attaching to the surface of wheat cells, and decreased the Agrobacterium mediated transformation efficiency in tobacco. On ther other hand, inhibition of TaUGT expressioncan increase the amount of regeneration seedlings but not the transformation efficiency in theAgrobacterium mediated transformation using wheat mature embryo derived callus as target tissues.Besides, in vitro eukaryotic expression of TaUGT had no effect on the growth of Agrobacterium.
In conclusion, many genes involved in wheat normal metabolism pathway all showed highrelevance to the resistance to Agrobacterium infection in wheat. TaUGT played a defensive role in theprocess of Agrobacterium infection before T-DNA integrating into genomic DNA of wheat.
1、利用抑制性差减杂交、蛋白质组学、数字化表达谱测序来筛选小麦对农杆菌侵染的应答基因，分别得到20、90和4,889个差异表达基因。其中，数字化表达谱测序灵敏度最高。差别基因聚类分析表明，参与物质代谢途径的基因所占比例最大，其次是次生代谢物质合成通路。其中，超过3%的差异表达基因参与到目前已经明确的植物-病原体互作通路，而在生物素合成通路、花生四烯酸代谢通路、光合作用生物通路差异表达基因定位最少。
2、KEGG通路分析表明，小麦UDP葡萄糖基转移酶（TaUGT）编码基因主要参与抗坏血酸代谢通路和药物代谢-细胞色素P450通路。而这两条通路都与植物抗病相关。Real-time PCR分析显示，TaUGT在小麦受到农杆菌侵染12小时后，表达量大幅上调，并随着共培养时间延伸而逐渐下降。表明该基因是在农杆菌侵染小麦细胞的初期起了较重要的调控作用。
3、基因结构分析表明，TaUGT为真核生物中少见的不存在内含子的核编码功能基因之一，与二倍体祖先种的同源性极高，进化上十分保守。经Southern杂交分析及将TaUGT基因序列向GrainGenes数据库的已定位小麦EST数据库进行blastn比对，推测TaUGT在小麦基因组中存在9个拷贝，在A、B、D基因组上各存在3个拷贝。亚细胞定位显示，TaUGT定位在细胞膜上，可跨膜发挥功能。
4、TaUGT功能分析发现，过表达TaUGT的拟南芥及小麦幼胚，农杆菌介导的GUS基因瞬时表达能力明显降低。电镜观察结果表明，过表达TaUGT后减少了农杆菌在小麦幼胚细胞表面附着的数量。利用农杆菌介导再次转化过表达TaUGT的烟草植株，转化效率明显降低。利用基因枪介导抑制小麦成熟胚愈伤组织中TaUGT的表达，增加了农杆菌侵染后小麦成熟胚愈伤组织的分化能力，但没有提高最终转化率。此外，体外真核表达的TaUGT并不影响农杆菌的生长。
综上所述，很多参与小麦正常代谢途径的基因都与小麦防御农杆菌侵染有关。TaUGT在农杆菌对小麦的侵染阶段，从农杆菌吸附小麦细胞表面到T-DNA转入小麦细胞的过程中调节小麦对农杆菌的防御反应。
Wheat is one of most important food crops worldwide, which yield and qulity affects the foodsecurity directly. Genetically modified (GM) technology is of great significance to wheat geneticalimprovement. A common limitation in wheat biotechnological breeding is the low transformationefficiency. Remarkably, gene gun-mediated transformation in wheat often triggers many seriousproblems like high degree of chimeric, multiple integration sites for foreign genes, and transgenesilencing. Howerver, Agrobacterium-mediated transformation can effectively avoid or alleviate thesedrawbacks. Still, Agrobacterium-mediated transformation efficiency in wheat needs to be improved.This study intends to screen and characterize the genes responsible for the transformation efficiencyfrom the view of molecular mechanisms underlying the interactions between wheat andAgrobacterium cells during their co-cultivation. The main results achieved in this study were asfollows:
1. According to the results of suppression subtractive hybridization (SSH), proteomics andRNA-seq,20,90, and4,889differential expressed genes were got in response to the Agrobacteriuminfection in wheat, respectively. Of the three methods above, RNA-seq is the most effective, accurate,and suitable for screening differential transcripts on the whole. Based on clustering analysis, most ofthe differential transcripts were involved in metabolic pathways; the second is in secondarymetabolism. Of those,3%participated in plant-pathogenic interaction pathway known so far.However, few were found in biotin synthesis pathway, arachidonic acid metabolism pathway, andphotosynthesis pathway.
2. KEGG pathway analysis indicated that wheat UDP-glucosyltransferase encoded gene (TaUGT)was mainly involved in ascorbic acid metabolic pathway and drug metabolism-cytochrome P450pathway. And these two pathways were related to the process of plant disease resistance. qRT-PCRshowed the expression of TaUGT upregulated at12h after Agrobacterium infection and decreasedgradually following the time span of co-culture in wheat. It inferred that TaUGT played a regulatingfactor in the initial stage of infection by Agrobacterium.
3. Structural analysis revealed that TaUGT in wheat was representative of those nuclear encodedwithout intron available in eukaryote, which showed high identity with its wild relative-orginatedspecies in evolution. Inferiring that there are nine copies were present in wheat genome based onSouthern blot analysis and the wheat mapped expression sequence tags (EST) in GrainGenes database.And each three copies are located on genome A, B, and D of wheat, respectively. Furthermore,according to the result of subcellular localization, TaUGT located on cytomembrane and function wellthrough transmembrane.
4. Functional identification of TaUGT showed that overexpression of TaUGT inhibited transientexpression of GUS in Arabidopsis and wheat after Agrobacterium infection, reduced theAgrobacterium population attaching to the surface of wheat cells, and decreased the Agrobacterium mediated transformation efficiency in tobacco. On ther other hand, inhibition of TaUGT expressioncan increase the amount of regeneration seedlings but not the transformation efficiency in theAgrobacterium mediated transformation using wheat mature embryo derived callus as target tissues.Besides, in vitro eukaryotic expression of TaUGT had no effect on the growth of Agrobacterium.
In conclusion, many genes involved in wheat normal metabolism pathway all showed highrelevance to the resistance to Agrobacterium infection in wheat. TaUGT played a defensive role in theprocess of Agrobacterium infection before T-DNA integrating into genomic DNA of wheat.
引文
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