二化螟诱导水稻表达的基因及二化螟特异诱导水稻启动子的克隆
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摘要
二化螟(Chilo suppressalis)是一种为害严重的世界性水稻害虫,目前还未见水稻对二化螟防御反应分子机制、水稻被二化螟取食特异诱导表达启动子研究的报道。本研究的目的在于:将扣除杂交技术和cDNA点阵技术相结合,得到水稻被二化螟诱导表达的基因,推测水稻对二化螟防御反应分子机制,克隆并鉴定水稻被二化螟特异诱导表达的启动子。
将水稻被二化螟取食后叶鞘mRNA及对照mRNA分别作为tester和driver构建cDNA扣除文库,文库Ⅰ以二化螟取食后叶鞘mRNA为tester,以对照mRNA为driver,文库Ⅱ刚好相反。理论上,来自于文库Ⅰ的EST为二化螟诱导上升表达的基因,来自于文库Ⅱ的EST为二化螟诱导下降表达的基因。从两个文库中共挑取864个克隆测序,序列拚接后得到271条unisequence。通过BLASTN和BLASTX分析这些序列的功能,结果表明这些基因涉及到光合作用、基础代谢、次生代谢、信号传导、蛋白降解、电子传递等多个代谢途径,这说明水稻对二化螟的防御反应是一个全基因组水平转录调整过程。
将这271个cDNA克隆、来自于本室明恢63全生育期平衡化cDNA文库的113个克隆、阳性对照及阴性对照cDNA克隆抽提质粒点在尼龙膜上,将二化螟取食后6h明恢63叶鞘及对照叶鞘总RNA反转录并用~(32)P标记成探针与尼龙膜杂交,杂交重复2次。结果得到在两次杂交中皆上升表达的基因12个,在两次杂交中皆下降表达的基因5个。尼龙膜杂交得到的差异表达克隆大大少于扣除杂交得到的差异表达克隆,但在这17个差异表达的克隆中,上升表达的基因基本上来自于文库Ⅰ(1个例外),下降表达的基因基本上来自于文库Ⅱ(1个例外),这说明扣除杂交技术与cDNA点阵技术之间的主要差异在于两者的灵敏度不同。
为了得到二化螟特异诱导表达的水稻启动子,从以上17个差异表达克隆中随机挑取4个上升表达克隆进行Northern blot分析。其中3个克隆同时被机械损伤和二化螟取食诱导。1个可能编码胰凝乳蛋白酶抑制剂的克隆—B1-A04在二化螟取食后急剧上升表达,但是机械损伤处理时其表达量几乎与对照一样,没有明显的变化。
通过PCR从基因组中克隆了B1-A04基因的启动子片段,如果将转录起始位点定位为1,则该启动子片段覆盖了-1569至+446之间的区域,包括B1-A04基因部分编码序列。将该片段(为了描述方便,命名为-1569)与GUS报告基因融合并通过农杆菌转化到水稻品种中花11中。在转基因植株中,二化螟未取食时,GUS基因在茎杆与幼穗中表达,在叶片与叶鞘中不表达;二化螟取食6h后,GUS基因在叶鞘与茎杆中的表达活性明显上升,但是叶片中仍然不表达。这说明该启动子具有器官特异表达、发育阶段特异表达的特点。GUS基因及内源B1-A04基因的表达水平在外源茉莉酸、脱落酸处理前后没有明显的变化。
为了了解-1569片段可能的调控区域,本研究从-1569的5'-end进行缺失,得到-1166至+446(命名为-1166)、-582至+446(命名为-582)、-371至+446(命名为-371)、-112至+446(命名为-112)4个截短的启动子片段。通过GUS活性测定及GUS组织染色可知在-1569位点至-1166位点、-1166位点至-582位点之间存在负调控顺式因子,在未被二化螟取食时抑制GUS基因在叶鞘中表达,被二化螟取食后开放。凝胶阻滞试验(EMSA)鉴定出7个与对照叶鞘核蛋白、二化螟取食后叶鞘核蛋白、机械损伤叶鞘核蛋白结合能力有差异的DNA探针,本文对这7段DNA对下游基因的可能调控作用进行了讨论。
本研究中得到的-1569启动子片段在水稻转基因育种中具有应用前景。
Rice striped stemborer,Chilo suppressalis(Walker),is one of the most damaging Lepidoptera pests of rice worldwide.There are few documents on molecular defensive mechanism of rice against C.suppressalis,and no reports about identification of rice promoter specifically responsive to C.suppressalis attack.The objective of this study was to identify rice genes responding to C.suppressalis and rice gene promoter that is specifically induced by feeding of C.suppressalis.
mRNA from rice sheath with and without chewing by C.suppressalis was used as tester and driver respectively to construct two subtractive cDNA libraries.LibraryⅠwas constructed using mRNA isolated from the feeding treatment as the tester with the undamaged control as the driver,and the reverse was done for libraryⅡ.Thus,libraryⅠandⅡwere expected to yield up-and down-regulated genes,respectively.A total of 864 clones randomly selected from the two libraries were sequenced,and 271 unique sequences longer than 100 bp were obtained.Functions of these ESTs were analyzed with BLASTN and BLASTX.These genes played roles in photosynthesis,primary metabolism, secondary metabolism,signal transduction,protein degration,electron transport and so on. This result indicated that the rice response triggered by C.suppressalis is likely to be involved comprehensive transcriptional reorganization.
These 271 cDNA clones together with other 113 ESTs from the normalized cDNA library and internal controls were arrayed on a nylon membrane.Total RNA of rice sheath harvested at 6 hours after C.suppressalis attack and control was reversely-transcribed and labeled with ~(32)P as the probes,and hybridized with the membranes.The hybridization was repeated twice,which identified 12 up-regulated and 5 down-regulated clones.The differentially expressed genes verified from the macroarray were much less than those from subtractive libraries,but only 2 differentially expressed genes were the exception which presented in the opposite library they should be.
Four of these up-regulated clones were randomely selected to be analyzed by Northern blot.Three of the selected clones were induced both by C.suppressalis infestation and wounding.One cDNA(B1-A04) encoding a putative subtilisin/chymotrypsin inhibitor was found to be rapidly and highly induced by C. suppressalis innfestation,compared with mechanical wounding.
The putative promoter region of B1-A04,spanning from-1569 to +446 relative to the transcriptional initiation site was isolated,fused to the GUS gene and introduced by Agrobacterium-mediated transformation to rice.In non-infested plants,the GUS activity driven by this promoter fragment was detected in culms and panicles,but not in leaves and sheaths.At 6 h after insect feeding,GUS activity was significantly induced in sheaths and culms,but not in leaves.GUS activity and native B1-A04 gene were not induced by JA and ABA treatment.These results showed that this promoter fragment had an organ-specific(i.e.culm and panicle) as well as developmental stage-specific(booting stage) expression pattern.
A serial deletion analysis revealed two regions(-1569 to-1166 and-1166 to-582) that negatively regulate the gene expression in sheaths of non-infested plants but not in insect-infested plants.An electrophoretic mobility shift assay(EMSA) identified 7 DNA fragments with various binding activities with nuclear proteins from mechanically wounded,insect-infested and untreated plants,and their possible roles in gene regulation were speculated.This promoter fragment should have utility in development of insect resistant transgenic crops.
将水稻被二化螟取食后叶鞘mRNA及对照mRNA分别作为tester和driver构建cDNA扣除文库,文库Ⅰ以二化螟取食后叶鞘mRNA为tester,以对照mRNA为driver,文库Ⅱ刚好相反。理论上,来自于文库Ⅰ的EST为二化螟诱导上升表达的基因,来自于文库Ⅱ的EST为二化螟诱导下降表达的基因。从两个文库中共挑取864个克隆测序,序列拚接后得到271条unisequence。通过BLASTN和BLASTX分析这些序列的功能,结果表明这些基因涉及到光合作用、基础代谢、次生代谢、信号传导、蛋白降解、电子传递等多个代谢途径,这说明水稻对二化螟的防御反应是一个全基因组水平转录调整过程。
将这271个cDNA克隆、来自于本室明恢63全生育期平衡化cDNA文库的113个克隆、阳性对照及阴性对照cDNA克隆抽提质粒点在尼龙膜上,将二化螟取食后6h明恢63叶鞘及对照叶鞘总RNA反转录并用~(32)P标记成探针与尼龙膜杂交,杂交重复2次。结果得到在两次杂交中皆上升表达的基因12个,在两次杂交中皆下降表达的基因5个。尼龙膜杂交得到的差异表达克隆大大少于扣除杂交得到的差异表达克隆,但在这17个差异表达的克隆中,上升表达的基因基本上来自于文库Ⅰ(1个例外),下降表达的基因基本上来自于文库Ⅱ(1个例外),这说明扣除杂交技术与cDNA点阵技术之间的主要差异在于两者的灵敏度不同。
为了得到二化螟特异诱导表达的水稻启动子,从以上17个差异表达克隆中随机挑取4个上升表达克隆进行Northern blot分析。其中3个克隆同时被机械损伤和二化螟取食诱导。1个可能编码胰凝乳蛋白酶抑制剂的克隆—B1-A04在二化螟取食后急剧上升表达,但是机械损伤处理时其表达量几乎与对照一样,没有明显的变化。
通过PCR从基因组中克隆了B1-A04基因的启动子片段,如果将转录起始位点定位为1,则该启动子片段覆盖了-1569至+446之间的区域,包括B1-A04基因部分编码序列。将该片段(为了描述方便,命名为-1569)与GUS报告基因融合并通过农杆菌转化到水稻品种中花11中。在转基因植株中,二化螟未取食时,GUS基因在茎杆与幼穗中表达,在叶片与叶鞘中不表达;二化螟取食6h后,GUS基因在叶鞘与茎杆中的表达活性明显上升,但是叶片中仍然不表达。这说明该启动子具有器官特异表达、发育阶段特异表达的特点。GUS基因及内源B1-A04基因的表达水平在外源茉莉酸、脱落酸处理前后没有明显的变化。
为了了解-1569片段可能的调控区域,本研究从-1569的5'-end进行缺失,得到-1166至+446(命名为-1166)、-582至+446(命名为-582)、-371至+446(命名为-371)、-112至+446(命名为-112)4个截短的启动子片段。通过GUS活性测定及GUS组织染色可知在-1569位点至-1166位点、-1166位点至-582位点之间存在负调控顺式因子,在未被二化螟取食时抑制GUS基因在叶鞘中表达,被二化螟取食后开放。凝胶阻滞试验(EMSA)鉴定出7个与对照叶鞘核蛋白、二化螟取食后叶鞘核蛋白、机械损伤叶鞘核蛋白结合能力有差异的DNA探针,本文对这7段DNA对下游基因的可能调控作用进行了讨论。
本研究中得到的-1569启动子片段在水稻转基因育种中具有应用前景。
Rice striped stemborer,Chilo suppressalis(Walker),is one of the most damaging Lepidoptera pests of rice worldwide.There are few documents on molecular defensive mechanism of rice against C.suppressalis,and no reports about identification of rice promoter specifically responsive to C.suppressalis attack.The objective of this study was to identify rice genes responding to C.suppressalis and rice gene promoter that is specifically induced by feeding of C.suppressalis.
mRNA from rice sheath with and without chewing by C.suppressalis was used as tester and driver respectively to construct two subtractive cDNA libraries.LibraryⅠwas constructed using mRNA isolated from the feeding treatment as the tester with the undamaged control as the driver,and the reverse was done for libraryⅡ.Thus,libraryⅠandⅡwere expected to yield up-and down-regulated genes,respectively.A total of 864 clones randomly selected from the two libraries were sequenced,and 271 unique sequences longer than 100 bp were obtained.Functions of these ESTs were analyzed with BLASTN and BLASTX.These genes played roles in photosynthesis,primary metabolism, secondary metabolism,signal transduction,protein degration,electron transport and so on. This result indicated that the rice response triggered by C.suppressalis is likely to be involved comprehensive transcriptional reorganization.
These 271 cDNA clones together with other 113 ESTs from the normalized cDNA library and internal controls were arrayed on a nylon membrane.Total RNA of rice sheath harvested at 6 hours after C.suppressalis attack and control was reversely-transcribed and labeled with ~(32)P as the probes,and hybridized with the membranes.The hybridization was repeated twice,which identified 12 up-regulated and 5 down-regulated clones.The differentially expressed genes verified from the macroarray were much less than those from subtractive libraries,but only 2 differentially expressed genes were the exception which presented in the opposite library they should be.
Four of these up-regulated clones were randomely selected to be analyzed by Northern blot.Three of the selected clones were induced both by C.suppressalis infestation and wounding.One cDNA(B1-A04) encoding a putative subtilisin/chymotrypsin inhibitor was found to be rapidly and highly induced by C. suppressalis innfestation,compared with mechanical wounding.
The putative promoter region of B1-A04,spanning from-1569 to +446 relative to the transcriptional initiation site was isolated,fused to the GUS gene and introduced by Agrobacterium-mediated transformation to rice.In non-infested plants,the GUS activity driven by this promoter fragment was detected in culms and panicles,but not in leaves and sheaths.At 6 h after insect feeding,GUS activity was significantly induced in sheaths and culms,but not in leaves.GUS activity and native B1-A04 gene were not induced by JA and ABA treatment.These results showed that this promoter fragment had an organ-specific(i.e.culm and panicle) as well as developmental stage-specific(booting stage) expression pattern.
A serial deletion analysis revealed two regions(-1569 to-1166 and-1166 to-582) that negatively regulate the gene expression in sheaths of non-infested plants but not in insect-infested plants.An electrophoretic mobility shift assay(EMSA) identified 7 DNA fragments with various binding activities with nuclear proteins from mechanically wounded,insect-infested and untreated plants,and their possible roles in gene regulation were speculated.This promoter fragment should have utility in development of insect resistant transgenic crops.
引文
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