CN108823239B - 一种提高植物抗病性载体及其应用 - Google Patents
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Abstract
本发明公开了一种提高植物抗病性载体及其应用,所述载体由表达控制区与植物NPR1基因可操作连接构成,所述表达控制区包括免疫诱导型启动子和病原相关的上游开放阅读框。与非转基因相比,通过本发明获得的植株抗病性提高50%以上;与组成型启动子,如玉米泛素启动子pUBI相比,利用本发明中的表达控制区介导NPR1基因的表达,适应代价降低80%以上。同时,利用本发明中的表达控制区介导单子叶植物的NPR1基因表达,比介导双子叶植物的NPR1基因表达,更有利于提高单子叶植物玉米和水稻的抗病性,抗病性提供5%以上。
Description
(一)技术领域
本发明涉及一种免疫诱导型启动子和病原相关的上游开放阅读框(uORF)构成的表达控制区以及一种单子叶植物的NPR1基因的可操作连接构建成的载体,利用这种载体获得抗病性提高的转基因植物的方法。
(二)背景技术
植物经常受到包括病毒、细菌、真菌和线虫在内的各种致病生物体的攻击。然而,大部分植物具有抵抗致病生物体的先天机制。植物育种家和病理学家已经鉴定了对植物病原体具有抗性的天然变化形式并将其培育成许多农作物植物。这些天然病害抗性基因通常提供了抵抗病原体的高水平抗性或免疫性。
当植物受病虫害侵犯时,受伤部位会立刻启动细胞程序性死亡,即发生所谓超敏反应(hypersensitive response:HR)。HR通常会启动未受伤部位产生次级防御反应,从而对一般的病虫害产生普遍抗性,这种现象称为系统获得性抗性(systemic acquiredresistance,SAR)(Lamb,Cell 76:419-422,1994;Lamb等,Cell 56:215-224,1989)。发生在感染部位的一种防御反应叫作超敏反应("HR"),该反应涉及感染植物细胞或组织或该二者的快速局限性坏死。人们认为感染细胞的快速死亡使侵入病原体得不到足够营养供给,从而抑制病原体生长。进行HR的细胞表现出核DNA断裂(例如DNA梯化(DNA laddering)),DNA断裂是首先在动物系统中描述的编程性细胞死亡,表明HR涉及主动编程性细胞死亡(Mittler等,Plant Physlo1.108:489-493,1995;Greenberg等,Cell 77:551-5fi3,1994;Ryerson和Heath,Plant Cell 8:393-402,1996;Wang等,Plant Cell 8:375-391,1996)。HR也伴随有膜相关的氧化爆发,从而导致NADPH依赖性产生02和H202。这些活性氧类物质对侵入病原体可以有直接毒性,或者可以参与损伤周围的植物细胞壁的交联作用,以便对感染形成屏障(Bradly等,Ce11 70:21-30,1992;Levine等,Cell 79:583-593,1994)。
HR不仅阻止感染病原体的局部生长,还认为它引发该植物非感染部分的其它防御反应,使该植物对多种通常有毒的病原体产生抗性(Enyedi等,Cell 70:879-886,1992;Malamy和Klessig,Plant J.2:643-654,1992))。后一种现象称为系统获得性抗性(SAR),并且认为这是许多基因协同激活的结果,所述基因通常被称作病理相关(PR)基因。许多这类PR基因的生物学功能还不清楚;然而,大量生理、生化和分子证据提示,特定PR基因在赋予对病原体的抗性中起作直接作用。例如,某些PR因编码直接抑制病原体体外生长的壳多糖酶和β-1,3-葡聚糖酶(Mauch等,Plant Physlol.88:936-942,1988;Ponstein等,PlantPtrysio1.104:109-118,1994;Schlumbaum等,Nature 324:365-367,1986;Sela-Buurlage等,Plant Physiol.101:857-863,1993;Terms等,J.B1o1.Chew.267:15301-15309,1992;Woloshuk等,Plant Cell 3:619-628,1991)。此外,已表明在PR基因的转基因植物中的组成型表达,在少数情况下降低疾病易感性(Alexander等,Proc.Natl.Acad.Sci.USA 90:7327-7331,1993;Liu等,Proc.Natl.Acad.Sci.USA 91:1888-1892;1994;Terras等,Plant Cell7:573-588,1995;Zhu等,BioTechnology 12:807-812,1994)。
在理论上,SAR可以被分成两个阶段。在初始阶段中,识别病原体感染且释放通过韧皮部传输到远端组织的信号。这种系统性信号被因SAR基因和病害抗性的表达而起反应的靶细胞觉察。SAR的维持期是指从数周到植物的整个生命期的时间期限,在此过程中该植物是准稳态的且病害抗性得到控制(Ryals等,The plant cell,8(10):1809,1996)。
拟南芥NPR1(PR基因的非表达子)基因已被证明是SAR过程中的一个关键的调节基因。NPR1基因的突变体植株不能回应诱导SAR的信号,如水杨酸(SA),或SA诱导的致病相关(PR)基因(Cao等,The Plant Cell,6(11):1583-1592,1994)。Friedrich等和Zhang等(Friedrich等,Mol.Plant-Microbe Interact.9:1114-1124,2001)在拟南芥中过表达(At)NPR1后,转基因植株具有广谱抗病性,且没有发现转基因植株有显著的副作用。Chern(Chern等,Plant J.27:101-113,2001)等在水稻中过表达(At)NPR1基因,转基因植株对细菌病原菌X.oryzaepv.Oryzae的抗性增强,且植株没有明显副作用。Fitzgerald等(Fitzgerald,H.A.等,Molecular Plant-Microbe Interactions,17(2),140-151,2004)发现过表达(At)NPR1的水稻在特定的生长条件下(低光照和高湿度)产生有害的副作用-触发类病变/细胞死亡(LMD)表型。Xu等利用通过利用uORF(上游开放阅读框)在翻译水平上精准调控该抗病蛋白NPR1的表达-在没有病原菌入侵时候,NPR1蛋白处于极低表达水平;一旦有病原菌危害,NPR1蛋白快速表达,短时间内即可阻止病菌的入侵。在水稻中进一步研究发现精准调控NPR1的表达,不仅不影响水稻的农艺性状,并且对水稻生产上重要病害-稻瘟病、白叶枯病和细菌性条斑病均有很好的抗性。
目前已从动物、植物、病毒及微生物中分离到许多适用于植物的启动子。根据作用方式及功能可将启动子分为3类:组成型启动子、诱导型启动子和组织特异型启动子。诱导型启动子(inducible promoter)是指在某些特定的物理或化学信号的刺激下,该种类型的启动子可以大幅度地提高基因的转录水平。目前已经分离了光诱导表达基因启动子、热诱导表达基因启动子、创伤诱导表达基因启动子、真菌诱导表达基因启动子和共生细菌诱导表达基因启动子等。
烟草中SAR基因是一个至少包含十二个成员的家族,SAR也受一些诸如水杨酸(SA),INA(dichloroisonicotinic acid)和BTH(benzothiadiazole)等化学物诱导。烟草Sat8.2b基因启动子-205/-201是as-1元件(TGACG),-146/-141和-276/-271为两个GT-1结合序列(GGAAAT),-97/-94、-322/-318和-761/-758分别是Dof结合基序(AAAG),前两者被认为可以与SA应答的转录因子结合而起关键作用(Song,F.,&Goodman,R.M.,Gene,290(1):115-124,2002)。启动子缺失实验表明Sar8.2b启动子-927~-728和-351~-197分别包含有SAR高效诱导基因表达所需的顺式作用元件,缺少了这两个DNA片段,转基因烟草GUS表达活性明显降低(Song,F.,&Goodman,R.M.,Gene,290(1):115-124,2002)。拟南芥TBF1基因是负责生长-防御转换的关键基因。其本身的表达受到转录和翻译水平上的严格调控。TBF1基因翻译起始密码子5’端存在2个编码多芳香族氨基酸的上游开放阅读框在翻译水平上调控TBF1基因的表达。TBF1启动子上也含有多个上述病原体诱导表达相关元件(Karolina M等,Current Biology 22:103–112,2012)。
尽管作了大量研究且应用了包括植物的遗传转化在内的完善和深入细致的农作物保护措施,但是每年因病害导致的损失仍保持在10亿美元。因此,存在对开发基于植物中病害抗性的遗传基础不断增加的了解的新的农作物保护措施的持续需求。特别地,需要研究如果利用抗病基因,获得适用代价(fitness costs)最低的抗病植物的方法。目前,已经有很多报道在植物中利用各种调控元件调控拟南芥中的NPR1基因的表达来提供植物抗性,但是利用特异的表达调控元件来介导单子叶植物中NPR1基因表达,进而提高植物抗病性,减小其适用代价的研究还很少见。特别的,在长期的进化过程中,单子叶植物和双子叶植物中的NPR1基因对病原体的反应存在显著的差异。因此,研究如何利用适合的调控元件调控单子叶植物中NPR1基因在植物中表达来提高植物抗病性非常重要。
(三)发明内容
本发明目的是提供一种提高植物抗病性载体及应用,利用单子叶植物的NPR1基因提高植物抗病性,并降低植物适用代价。
本发明采用的技术方案是:
本发明提供一种提高植物抗病性载体,所述载体由表达控制区(uORF)与植物NPR1基因可操作连接构成,所述表达控制区包括免疫诱导型启动子和病原相关的上游开放阅读框。
进一步,所述上游开放阅读框编码富含芳香族氨基酸的多肽,所述免疫诱导型启动子为植物生长-防御过程转化的关键转录因子的启动子。
进一步,所述表达控制区由免疫诱导型启动子和病原相关的上游开放阅读框(uORF)构成,且病原相关的上游开放阅读框(uORF)受免疫诱导型启动子介导转录;表达控制区中免疫诱导型启动子和上游开放阅读框(uORF)可以来源于同一个物种,也可以来源于不同物种。
进一步,所述表达控制区核苷酸序列为SEQ ID NO.1或SEQ ID NO.2所示。
进一步,所述植物NPR1基因来自单子叶植物,所述单子叶植物为拟南芥、大豆和油菜等双子叶植物以外的被子植物,包括禾本科植物,优选水稻、玉米或小麦。
进一步,所述植物NPR1基因核苷酸序列为SEQ ID NO.3或SEQ ID NO.5所示。
本发明所述单子叶植物NPR1基因的终止子可以来源于生物基因组中,也可以通过人工合成的方式获得,优选终止子的核苷酸序列为SEQ ID NO:7所示。
本发明还提供一种所述提高植物抗病性载体在制备抗病植物中的应用,所述应用是将表达控制区导入植物内源NPR1基因编码区上游而实现的。
本发明利用所述的任意一种表达控制区序列和任意一种单子叶植物的NPR1基因构建T-DNA(启动子-植物NPR1-终止子-启动子-G10-终止子),利用T-DNA获得转基因作物,具体本发明精准控制NPR1基因表达来提高植物抗病性的方法为:通过基于基因编辑技术的DNA精准插入技术包括基于ZFN(zinc-finger nucleases)、TALEN(transcriptionactivator-like effector nucleases)、CRISPR/Cas9和Argonaute/gDNA等把本发明中的表达控制区导入植物内源NPR1基因编码区上游,实现利用本发明中的表达控制区调控NPR1基因表达。
所述T-DNA是将表达控制区,植物NPR1基因与双元载体连接构成,所述双元载体是将pCambia1300载体中的抗潮霉素基因置换为抗草甘膦基因G10构建成双元载体pCambia1300-p35S-G10。
与现有的发明相比,本发明有益效果主要体现在:本发明首次研究利用表达控制区介导单子叶植物的NPR1基因在植物中表达,利用免疫诱导型启动子和病原相关的上游开放阅读框(uORF)分别在转录水平和翻译水平对NPR1基因进行精细且严格的调控,进而使得获得的转基因植物具有更强的抗病性和更低的适应代价。与非转基因相比,通过本发明获得的植株抗病性提高50%以上;与组成型启动子,如玉米泛素启动子pUBI相比,利用本发明中的表达控制区介导NPR1基因的表达,适应代价降低80%以上。同时,利用本发明中的表达控制区介导单子叶植物的NPR1基因表达,比介导双子叶植物的NPR1基因表达,更有利于提高单子叶植物玉米和水稻的抗病性,抗病性提供5%以上。
(四)附图说明
图1:本发明中的表达控制区结构示意图。启动子为免疫诱导型启动子,uORF为病原相关的上游开放阅读框。在同一个表达控制区中可以有一个或多个uORFs。
图2:本发明中植物转化载体T-DNA结构示意图。该T-DNA中包括本发明中的表达控制区介导单子叶植物NPR1基因表达的表达框和其它靶标基因或标记基因表达框。
(五)具体实施方式
下面结合具体实施例对本发明进行进一步描述,但本发明的保护范围并不仅限于此:
本发明实施例所使用的分子生物学和生物化学方法均为已知的技术。在Ausubel编写的John Wiley and Sons公司出版的Current Protocols in Molecular Biology和J.Sambrook等编写Cold Spring Harbor Laboratory Press(2001)出版的MolecularCloning:A Labortory Manual,3rd ED.等文献均有详细的说明。
实施例1载体构建
1、通过PCR的方法获得表达控制免疫诱导型启动子和病原相关的上游开放阅读框(uORF)构成的表达控制区AtTBF1-D和OsTBF1-D(结构如图1所示)。
设计PCR引物AtTBF1-D-F(5’CAAGCTTCGACGACTAGTTTACAGAGAATTT)和AtTBF1-D-R(5’GGATCCCTTTTTTTATTTTACCACAGAAAAAT),以拟南芥的基因组DNA为模板,通过PCR扩增获得AtTBF1-D。PCR反应条件为:95℃3分钟;95℃15秒,68℃15秒,72℃4分钟,重复32个循环;然后72℃10分钟。将获得的大约3.6Kb的PCR产物克隆到pCambia1300载体中。然后,用HindIII和BamHI双酶切得到AtTBF1-D,并且DNA序列测定表明核苷酸序列正确(SEQ ID NO:1)。
设计PCR引物OsTBF1-D-F(5’GGTACCGATTTATAAATGCTGCTTTCACTGC)和OsTBF1-D-R(5’ATGGATCCCCTAACGCTATGATCTCTTTCTC),以商业玉米品种秀水134的基因组DNA为模板,通过PCR扩增获得pUBI。PCR反应条件为:95℃3分钟;95℃15秒,68℃15秒,72℃4分钟,重复32个循环;然后72℃10分钟。将获得的大约3.3Kb的PCR产物克隆到过渡载体pCambia1300中。然后,用KpnI和BamHI双酶切得到OsTBF1-D,并且DNA序列测定表明核苷酸序列正确(SEQ IDNO:2)。
2、人工合成玉米、水稻和拟南芥的NPR1基因的mRNA序列,其中玉米ZmNPR1基因的mRNA序列为SEQ ID NO:3(编码蛋白的氨基酸序列为SEQ ID NO:4),水稻OsNPR1基因的mRNA序列为SEQ ID NO:5(编码蛋白的氨基酸序列为SEQ ID NO:6),拟南芥AtNPR1基因的mRNA序列为SEQ ID NO:8(编码蛋白的氨基酸序列为SEQ ID NO:9)。
人工合成终止子序列(SEQ ID NO:7)。
3、合成引物:
ZmNPR1-F1:GGGATCCAACAATGGAGCCCATGGACAGCCAGCTCAC,
ZmNPR1-R1:GATCTTTTATCACCTCCGAGGCCGGACGGCCCCGATC,
ZmNPR1-F2:CGGCCTCGGAGGTGATAAAAGATCTGTTCTGCACAAAGT,
ZmNPR1-R2:GGGTACCCCTGGATTTTGGTTTTAGGAATTAGAAATTTT。以玉米ZmNPR1基因的mRNA(SEQ ID NO:3)为模板,通过引物ZmNPR1-F1和ZmNPR1-R1进行PCR扩增获得ZmNPR1-A。PCR反应条件为:95℃3分钟;95℃15秒,68℃15秒,72℃2分钟,重复32个循环;然后72℃10分钟。将获得的大约1.8Kb的PCR产物;以人工合成终止子(SEQ ID NO:7)为模板,通过引物ZmNPR1-F2和ZmNPR1-R2进行PCR扩增获得ZmNPR1-B。PCR反应条件为:95℃3分钟;95℃15秒,68℃15秒,72℃30秒,重复32个循环;然后72℃10分钟。将获得的大约0.2Kb的PCR产物;最终,以ZmNPR1-A和ZmNPR1-B为模板,通过引物ZmNPR1-F1和ZmNPR1-R2进行PCR扩增获得ZmNPR1-ter。PCR反应条件为:95℃3分钟;95℃15秒,68℃15秒,72℃2分钟,重复32个循环;然后72℃10分钟。将获得的大约2Kb的PCR产物。将获得的PCR产物克隆到载体pCambia1300中。然后,用KpnI和BamHI双酶切得到ZmNPR1-ter。
4、合成引物
OsNPR1-F1:GGATCCATCGATGGAGCCGCCGACCAGCCACGTC,
OsNPR1-R1:AACAGATCTTATCATCTCCTTGGTCGAATGGCCCCGATC,
OsNPR1-F2:TTCGACCAAGGAGATGATAAGATCTGTTCTGCACAAAGT,
ZmNPR1-R2:CGGTACCCCTGGATTTTGGTTTTAGGAATTAGAAATTTT。以水稻OsNPR1基因的mRNA(SEQ ID NO:5)为模板,通过引物OsNPR1-F1和OsNPR1-R1进行PCR扩增获得OsNPR1-A。PCR反应条件为:95℃3分钟;95℃15秒,68℃15秒,72℃2分钟,重复32个循环;然后72℃10分钟。将获得的大约1.8Kb的PCR产物;以人工合成终止子(SEQ ID NO:7)为模板,通过引物OsNPR1-F2和ZmNPR1-R2进行PCR扩增获得OsNPR1-B。PCR反应条件为:95℃3分钟;95℃15秒,68℃15秒,72℃30秒,重复32个循环;然后72℃10分钟。将获得的大约0.2Kb的PCR产物;最终,以OsNPR1-A和OsNPR1-B为模板,通过引物OsNPR1-F1和OsNPR1-R2进行PCR扩增获得ZmNPR1-ter。PCR反应条件为:95℃3分钟;95℃15秒,68℃15秒,72℃2分钟,重复32个循环;然后72℃10分钟。将获得的大约2Kb的PCR产物。将获得的PCR产物克隆到载体pCambia1300中。然后,用KpnI和BamHI双酶切得到OsNPR1-ter。
5、合成引物
AtNPR1-F1:TCGGATCCATGGACACCACCATCGACGGCTTC,
AtNPR1-R1:CTTGAGCTCTCATTACCTCCTCCTGTGGCTCAGC,
AtNPR1-F2:GGAGGAGGTAATGAGAGCTCAAGATCTGTTCTGCAC,
ZmNPR1-R2:CGGTACCCCTGGATTTTGGTTTTAGGAATTAGAAATTTT。以AtNPR1基因的mRNA(SEQ ID NO:8)为模板,通过引物AtNPR1-F1和AtNPR1-R1进行PCR扩增获得AtNPR1-A。PCR反应条件为:95℃3分钟;95℃15秒,68℃15秒,72℃2分钟,重复32个循环;然后72℃10分钟。将获得的大约1.8Kb的PCR产物;以人工合成终止子(SEQ ID NO:7)为模板,通过引物AtNPR1-F2和ZmNPR1-R2进行PCR扩增获得AtNPR1-B。PCR反应条件为:95℃3分钟;95℃15秒,68℃15秒,72℃30秒,重复32个循环;然后72℃10分钟。将获得的大约0.2Kb的PCR产物;最终,以AtNPR1-A和AtNPR1-B为模板,通过引物AtNPR1-F1和AtNPR1-R2进行PCR扩增获得AtNPR1-ter。PCR反应条件为:95℃3分钟;95℃15秒,68℃15秒,72℃2分钟,重复32个循环;然后72℃10分钟。将获得的大约2Kb的PCR产物。将获得的PCR产物克隆到载体pCambia1300中。然后,用KpnI和BamHI双酶切得到AtNPR1-ter。
6、G10为抗草甘膦基因(中国专利:201110009329.0)。通过人工合成G10基因。合成的基因5’端连接有玉米乙酰乳酸合成酶AHAS叶绿体转运信号肽且设置有XhoI酶切位点,3’端连接有终止子且设置有XhoI酶切位点。
7、农杆菌T-DNA载体的构建:双元载体pCambia1300-p35S-G10由载体pCambia1300修改而来,简单的说就是把pCambia1300载体中的抗潮霉素基因置换为抗草甘膦基因G10。具体把pCambia1300载体经过XhoI酶切以后,去磷酸化处理,然后与经过XhoI酶切后获得的人工合成的含有叶绿体转运信号肽和终止子的G10基因片段进行两端连接,转化,获得的载体即为pCambia1300-p35S-G10。
8、为了获得ZmNPR1基因表达载体,用HindIII和KpnI对之前构建好的pCambia1300-p35S-G10进行双酶切,回收获得载体;用HindIII和BamHI酶切含有AtTBF1-D表达控制区的质粒,获得AtTBF1-D片段;用BamHI和KpnI对含有ZmNPR1基因及其终止子的质粒,回收得到ZmNPR1-ter片段。然后,把上述酶切后的载体和两个片段进行三段连接,获得终载体。获得的T-DNA结构为:“启动子-ZmNPR1-终止子-启动子-G10-终止子”。这个载体命名为:pCambia1300-AtTBF1-D-ZmNPR1–p35S-G10(图2)。
9、为了获得OsNPR1基因表达载体,用KpnI对之前构建好的pCambia1300-p35S-G10进行单酶切,并且进行去磷酸化处理,回收获得载体;用KpnI和BamHI酶切含有OsTBF1-D表达控制区的质粒,获得OsTBF1-D片段;用BamHI和KpnI对含有OsNPR1基因及其终止子的质粒,回收得到OsNPR1-ter片段。然后,把上述酶切后的载体和两个片段进行三段连接,获得终载体。获得的T-DNA结构为:“启动子-OsNPR1-终止子-启动子-G10-终止子”。这个载体命名为:pCambia1300-OsTBF1-D-OsNPR1–p35S-G10(图2)。
10、为了获得AtNPR1基因表达载体,用KpnI对之前构建好的pCambia1300-p35S-G10进行单酶切,并且进行去磷酸化处理,回收获得载体;用KpnI和BamHI酶切含有AtTBF1-D表达控制区的质粒,获得AtTBF1-D片段;用BamHI和KpnI对含有AtNPR1基因及其终止子的质粒,回收得到AtNPR1-ter片段。然后,把上述酶切后的载体和两个片段进行三段连接,获得终载体。获得的T-DNA结构为:“启动子-AtNPR1-终止子-启动子-G10-终止子”。这个载体命名为:pCambia1300-AtTBF1-D-AtNPR1–p35S-G10(图2)。
最后,通过电转的方法把上述3个T-DNA质粒转入农杆菌LBA4404中,通过含有15μg/ml四环素和50μg/mL的卡那霉素的YEP固体培养基筛选出阳性克隆,并保菌,用于接下来的植物转化。
实施例2玉米转化
玉米的转化技术已经比较成熟。参考文献如:Vladimir Sidorov&David Duncan(in M.Paul Scott(ed.),Methods in MolecularBiology:TransgenicMaize,vol:526;Yuji Ishida,Yukoh Hiei&Toshihiko Komari(2007)Agrobacterium-mediatedtransformation of maize.Nature Protocols 2:1614-1622。基本方法如下:取授粉后8-10天的Hi-II玉米穗,收集所有的未成熟胚(大小为1.0-1.5mm)。将实施例1中制备的含有T-DNA载pCambia1300-AtTBF1-D-ZmNPR1–p35S-G10、pCambia1300-OsTBF1-D-OsNPR1–p35S-G10和pCambia1300-AtTBF1-D-AtNPR1–p35S-G10的农杆菌与未成熟胚在共培养培养基上(MS+2mg/L 2,4-D+30g/L蔗糖+3g/L琼脂(sigma 7921)+40mg/L乙酰丁香酮)共培养2-3天(22℃)。转移未成熟胚到愈伤诱导培养基上(MS+2mg/L 2,4-D+30g/L蔗糖+2.5g/L gelrite+5mg/L AgNO3+200mg/L乙酰丁香酮),28℃暗培养10-14天。将所有的愈伤转到带有2mM草甘膦的筛选培养基(与愈伤诱导培养基相同)上,28℃暗培养2-3周。转移所有的组织到新鲜含草甘膦的筛选培养基上,28℃暗培养2-3周。然后,转移所有筛选后成活的胚性组织到再生培养基(MS+30g/L蔗糖+0.5mg/Lkinetin+2.5g/L gelrite+200mg/L乙酰丁香酮)上,28℃暗培养10-14天,每皿一个株系。转移胚性组织到新鲜的再生培养基上,26℃光照培养10-14天。转移所有发育完全的植株到生根培养基(1/2MS+20g/L蔗糖+2.5g/L gelrite+200mg/L乙酰丁香酮)上,26℃光照培养直到根发育完全。分别获得含转化载体pCambia1300-AtTBF1-D-ZmNPR1–p35S-G10、pCambia1300-OsTBF1-D-OsNPR1–p35S-G10和pCambia1300-AtTBF1-D-AtNPR1–p35S-G10的转基因玉米植株。
实施例3、水稻的转化
转基因水稻的获得方法是采用现有技术(卢雄斌,龚祖埙(1998)生命科学10:125-131;刘凡等(2003)分子植物育种1:108-115)。选取成熟饱满的“秀水-134”种子去壳,诱导产生愈伤组织作为转化材料。分别取实施例1中构建的载体pCambia1300-AtTBF1-D-ZmNPR1–p35S-G10、pCambia1300-OsTBF1-D-OsNPR1–
p35S-G10和pCambia1300-AtTBF1-D-AtNPR1–p35S-G10的农杆菌划板。挑单菌落接种,准备转化用农杆菌。将待转化的愈伤组织放入OD为0.6左右的农杆菌菌液中(农杆菌菌液的制备:将农杆菌接种至培养基,培养至OD为0.6左右;培养基组成:3g/L K2HPO4、1g/LNaH2PO4、1g/LNH4Cl、0.3g/L MgSO4·7H2O、0.15g/L KCl、0.01g/L CaCl2、0.0025g/LFeSO4·7H2O、5g/L蔗糖、20mg/L乙酰丁香酮,溶剂为水,pH=5.8),让农杆菌结合到愈伤组织表面,然后把愈伤组织转移到共培养培养基(MS+2mg/L 2,4-D+30g/L葡萄糖+30g/L蔗糖+3g/L琼脂(sigma 7921)+20mg/L乙酰丁香酮)中,共培养2-3天。用无菌水冲洗转化后的愈伤,转移到筛选培养基(MS+2mg/L 2,4-D+30g/L蔗糖+3g/L琼脂(sigma 7921)+20mg/L乙酰丁香酮+2mM草甘膦(Sigma))上,筛选培养两个月(中间继代一次)。把筛选后,生长活力良好的愈伤转移到预分化培养基(MS+0.1g/L肌醇+5mg/L ABA+1mg/L NAA+5mg/L 6-BA+20g/L山梨醇+30g/L蔗糖+2.5g/L gelrite)上培养20天左右,然后将预分化好的愈伤组织移到分化培养基上,每天14小时光照分化发芽。2-3周后,把抗性再生植株转移到生根培养基(1/2MS+0.2mg/L NAA+20g/L蔗糖+2.5g/L gelrite)上壮苗生根,最后将再生植株洗去琼脂移植于温室,选择产量高、种子大或者生物量高等能够提高水稻产量的转基因株系,培育新品种。分别获得含上述转化载体和只含有筛选标记基因EPSPS的空载体的转基因水稻植株。
实施例4转基因玉米的抗病性增强
将实施例2制备的转基因玉米植株的T0代植株移栽到温室中,用商业化品种“郑丹958”的母本,“郑58”(Z58)的花粉进行授粉,收获T0代种子。然后将这些品系与商业化品种“郑丹958”的母本,“郑58”(Z58)进行回交转育,获得Z58近等位基因系。再对这些近等位基因系的抗病性进行比较分析。
我们对获得的87个转pCambia1300-AtTBF1-D-ZmNPR1–p35S-G10载体的转基因株系(命名为AZR)和56个转pCambia1300-AtTBF1-D-AtNPR1–p35S-G10载体的株系(命名为AAR)进行对不同病原菌的抗性效果鉴定,抗性效果如表1、表2所示:
表1:
注:玉米小斑病、灰斑病、纹枯病的抗病性根据病斑长度确定,分别对每个品系的30棵以上转基因植株进行接菌测试,统计病斑长度,与非转基因对照植株进行统计分析,确定转基因品系的抗病性。玉米茎腐病的抗性根据植株成活率确定,分别对每个品系的40棵以上转基因玉米植株进行接菌测试,统计成活株数,与非转基因对照植株进行统计分析,确定转基因品系的抗病性。
表2:
注:玉米小斑病、灰斑病的抗病等级根据病斑面积确定,与非转基因对照相比病斑面积减少50%以上的等级为优,与非转基因对照相比病斑面积减少20-50%以上的等级为良,与非转基因对照相比病斑面积减少5-20%以上的等级为合格。分别对测试品系的30棵以上转基因植株进行接菌测试,统计病斑面积,与非转基因对照植株进行统计分析,确定转基因品系的抗病性。
上述结果表明在玉米中,pCambia1300-AtTBF1-D-ZmNPR1–p35S-G10载体转基因植株与pCambia1300-AtTBF1-D-AtNPR1–p35S-G10载体的转基因植株相比,抗病性差异显著,ZmNPR1基因在玉米抗病上的可靠性远大于AtNPR1基因。
实施例5转基因水稻的抗病性增强
将实施例3制备的转基因水稻植株的T0代植株移栽到温室中,收获T0代种子。再对这些品系的T2代植株的抗病性进行比较分析。
我们获得143个转pCambia1300-OsTBF1-D-OsNPR1–p35S-G10载体的转基因株系(命名为OOR)和120个转pCambia1300-AtTBF1-D-AtNPR1–p35S-G10载体的株系(命名为AAR)进行不同病原菌的抗性效果鉴定,抗性效果如表3、表4所示:
表3:
注:表中抗病性根据病斑长度确定,分别对每个品系的30个以上转基因植株进行接菌测试,统计病斑长度,与非转基因对照植株进行统计分析,确定转基因品系的抗病性。
表4:
注:水稻条斑病菌、白叶枯病的抗病等级根据病斑面积确定,与非转基因对照相比病斑面积减少50%以上的等级为优,与非转基因对照相比病斑面积减少20-50%以上的等级为良,与非转基因对照相比病斑面积减少5-20%以上的等级为合格。分别对测试品系的30棵以上转基因植株进行接菌测试,统计病斑面积,与非转基因对照植株进行统计分析,确定转基因品系的抗病性。
上述结果表明在水稻中,pCambia1300-OsTBF1-D-OsNPR1–p35S-G10载体转基因植株与pCambia1300-AtTBF1-D-AtNPR1–p35S-G10载体的转基因植株相比,抗病性差异显著,OsNPR1基因在水稻抗病上的可靠性远大于AtNPR1基因。
实施例6转基因玉米的适应代价降低
作为对照,双元载体中的表达控制区AtTBF1-D被替换为玉米泛素pUBI启动子,替换后获得的载体为pCambia1300-pUBI-ZmNPR1–p35S-G10。通过转化上述载体,我们获得了81个独立转化体(命名为UZR)。我们把上述转化体与87个转pCambia1300-AtTBF1-D-ZmNPR1–p35S-G10载体的转基因株系(命名为AZR)的适应代价进行了调查和比较。
研究结果表明,使用由病毒免疫诱导型启动子和病原相关的上游开发阅读框构成的表达控制区AtTBF1-D介导玉米NPR1基因在玉米中表达比用组成型启动子pUBI介导玉米NPR1基因在玉米中表达,适应代价显著降低。具体结果如表5。
表5:
序列表
<110> 浙江大学
<120> 一种提高植物抗病性载体及其应用
<160> 9
<170> SIPOSequenceListing 1.0
<210> 1
<211> 3558
<212> DNA
<213> 未知(Unknown)
<400> 1
caagcttcga cgactagttt acagagaatt tggaccgtcc gatgtaaagc gaaaatagat 60
ctaggttttc cacgtgtccc ctattttaat gaaaccttct gattcatgta gaagttttac 120
tcaatttaat attttttagt atgtagtttt gtgtgtgtgt gtgtgtgtgt ttttatggct 180
ccacaccaac ttttaaaatg gtagaagcat gttgcatgtg atcgagtaaa aagccaataa 240
tgagattcag aaaaataaaa attacttata tagtttttta gagaaaaaat tgtattttgt 300
ttaaagcctt aatccggttg ttgaaagagc tgtgtcacga gttaaaaata ttttcttttc 360
attttttaag taattagttt ataatgcaaa aatggttttt atttatttgt cttcgcttat 420
agaactgcaa attgagagag aaaaaaatga attagtggtg gtgaccaaac attcaggaag 480
ctgtgattga tcatttgttt ttgaggtgag tgtagtggca acgtatgacg ttaacatatg 540
gcgtacataa taattacatg aacttaatca taataatcat attgcattta attcatatat 600
catatcccat tagttggacc acttgatttg aggtcatgag aagaacattt atgttttttt 660
tagtttgaat cggagtgatc actaaaaact agatactgaa aattttcaaa ctaaaatcat 720
attaatcttc aaaaaatgtg aaatctaaaa aaaaaaaaaa ttttaacgcg ttcattgtag 780
ccaagtagcc aagtattgtt aaagtagtag taaaagaagt ttagctttaa gtgatataat 840
ttgacacaaa tcctacttag atatggataa taggatatag cttcatgtat atttttatcg 900
ttgcttctgt aaccccaaaa tgtgttgata taagcatttg aatattcgta tgtataatgt 960
tttcttttca ccgtaaaaca tattacaatg ttagtttata ttggattttg aatgtgttta 1020
tgaacagttt ttgtcgactc aaaagttaag atgagaatat ggaagaaagt aaagtttaaa 1080
agtcatgatg ggaacaagga atggaactca aacattctaa tactcaacaa acgcaattat 1140
attattacca tgactcatct ttcaagttcc atcaaaaaga ttcgtggaaa ataatagact 1200
tacgtttcaa atccatgttt ctttctttat aacaaaaaaa atggatgttt cttgacgcgt 1260
gtcgagagta ctcaccatta ctctgacttc agtgagtttg gtcaagtggt cttttttttt 1320
ctcatgtcac caaaggtcca aaccctagaa attagttcga actttccata gaagaactga 1380
ataaatggtc caaaattgtt ttaaaaagga cctaagccat tagttcattg aattcgagtt 1440
aatgggtgaa gatttttatg ataacgaaag tcggagtaat tatgcttttg gtccgatagt 1500
tttctaattt gttttctttc catttttttt ttttcaaata ctacatacta tataagatag 1560
tggtttgtgt taatgtcatc gatgtgttac catccgcatt atattaatta tttatcccaa 1620
cataaagtca gaatctgtaa tttctttgtt ataaaataca gtaaatggtt ccgtttaagc 1680
tgttagatga tttttgagta aaaactaatg taaaaaaaac aaaaaaaaaa caatgtagtt 1740
cataatacat gcatgtttta aagaagtttc ttgtttacta tcaacttgaa tagtatttca 1800
cgaagtcaaa attgttcatt ccgacttttc tatgtggaga aaaaaaattc tatcattgtg 1860
cacaatttaa cagaatgtaa tttcttgtaa aagaagagga aacaattcgc tgttagtaaa 1920
tgtgaagtat agaagtctaa aatgagatac ctcaactagc ttgaattaag aaaaaaaaca 1980
aaaactctat cgacatgaaa aaggtcgcaa atatttatca tttatcaatg ccaaaggagt 2040
atttggttca caaaatactg aatcatttat atagatatat aattagctct aaattctact 2100
ataacttgca aaataagtat actgactcaa ttatatagcg tttaaaaata gacgatttgt 2160
atgatgaggt ccatatatat ggagatgtgc atgcaactat cgacattttc acacgttgat 2220
atcgtctttc tccaatggag acttgaattt gtgtaaacta tgaatactcg tctctctaag 2280
accttttttc ttcaaccatg ccaactattt aggtaagatt ttactgtctt tgattgatat 2340
taaatactta gccgtggcgt tatcaatgaa tgataataaa aatgcggata aaagccaaag 2400
gtgttggaaa taaatccaag aatgaagacg tagatgtcga tgggtatttt aagaacttga 2460
atttgtcacg actcacacgt taaaatatat tatccgaatt gtttagtcta aagacacaca 2520
tatattgaaa aagaaaaggt aaatgaagct cattggtgcc taaatgtgaa atgaagccga 2580
aatgtgttag gtgaacacat ttaaatatac aaaaagaaat ataatagaaa caaaactaat 2640
taacaaagtc gcaatttgta ttgtataaaa tatctttccg tctcccgtca tatttgaaaa 2700
aaaaaaaatt acaaatctgt taattttaaa actttctaga aaaacacaag tatataattt 2760
tctcttttcg tgcgtgtttg ttttaaaata acattgtttt gattggcgac tcaacatatt 2820
ttagcattta catatttctg catatattaa atgatttata aactcaacta tagattaaaa 2880
tataatttga catctaataa ttttaacaat aatataaaat atgagattta taaattacga 2940
atataaatat tcaagggaga gaaaaagtag aacataattc aaaagataag actttttaga 3000
cttttttaac aatatttttg atggataaaa attattcaaa agagaagaaa gtaagaagaa 3060
aagatgtttc tgagaatttc tagaaacagc atccgttttt ataatttaat tttcttacaa 3120
aggtaggacc aacatttgtg atctataaat cttcctacta cgttatatag agacccttcg 3180
acataacact taactcgttt atatatttgt tttacttgtt ttgcacatac acacaaaaat 3240
aaaaaagact ttatatttat ttacttttta atcacacgga ttagctccgg cgaagtatgg 3300
tcgtcgtctt catcttcttc ctccatcatc agatttttcc ttaaatggaa gaaaccaaac 3360
gaaactccga tcttctccgt tctcgtgttt tcctctctgg cttttattgc tgggattggg 3420
aatttctcac cgctctcttg ctttttagtt gctgattctt tttccttcga ctttctattt 3480
ccaatctttc ttcttctctt tgtgtattag attattttta gttttatttt tctgtggtaa 3540
aataaaaaaa gggatcca 3558
<210> 2
<211> 3314
<212> DNA
<213> 未知(Unknown)
<400> 2
ggtaccgatt tataaatgct gctttcactg caaatgtcct atgagccaat tttttgatta 60
tccttgcatt ataccttccc cttgatatga cttgttgagt acatcggttg tactcagtct 120
tgctttattt tttccaatcc cccagaagta gagaatttat tggatggtga gttctatgaa 180
gattaggctt ttgccagacg tcgaggtttt gcctgtggat tatggaggaa gcttcggttt 240
gctgagaaga ttaaatttag atggtcttta gtttttctgt tgcttttctg agtttatttt 300
atatttttgt aagacgtgga attgtatcag attatcgtct gtgtactctg gttgatgtct 360
ggacagaggt ttaatgcaca gatagccggt gattcgggtt gtcgaatccc tgggcgcgac 420
aggtcctgta tttatactgt agattatctt ggtctccaag cagaactcgg agatatctcc 480
ctatgtatga gatcacatat cccctatctc ctccgaatag gtctctaaca ccttccattt 540
gaggagacgg tttccttacg cattgacctt tttcttaatt tgtatgtaat gaccgtattc 600
gtatacttca caagtccaaa gatatgactt attcgtaaca ctgacagagc tatatatacc 660
tggttgccca cgttgataga tttctttttt gtttttagat aatacgttga tagatttcta 720
tccctatgtt atgtaaacta cattcttttc atattcagta tttctgaaag aggtgtattt 780
aaataagacc ttaattaatt gttcatgact gatccaaatt ctcgaatcta aaattcagca 840
aagccttaac ccagccacaa acacacacac aaaaagctat acgtgtctgt gtctctgtta 900
aaaaggaaga agctcagcac taatataata tgaactggtc agcaatcagc acaggcagga 960
tatggacgac atctttccag ctaaggtgtt agaccgtcag agagattaag caacaatcag 1020
aaccttctcc atcagaccat caccctttgt gtcgcatcgg cgtccgcgga tgaaatcagc 1080
aaagttgcag ctggctatgg catttattaa ggccctgctt agttccaaac aaaacttttc 1140
ccaaaaatat catattaaat ctttatatat atatatatgg agtataaaat atagataaaa 1200
attaaaacta attgcacagt ttgcatataa atcacgagac gaatcttttg agcctaatta 1260
cgccatgatt agtcataagt gctacagtaa tccacatgtg ataatgacgg attaattagg 1320
ctcaaaatat tcgtctcgcg gttttcaggc gagttatgta attagttttt tcattcgtgt 1380
ccgaaaatcc cttccaacat ccggtcaaac gtccgatgtg atacccaaaa attttcattt 1440
gagccctaac taagctccga tctttctctc cgcccagccc cggagccagt ttgccacaca 1500
atgcagacgc aaacatgtaa ctgaccggag aaatgaagtg gattttggca caaaagtcag 1560
gtttctacca cgcacccatc ggtcagcagg aactccctct tagacggcct catttttctc 1620
cagccttttg ttcaggttgg tgaggcgttt acttgcaaac accgagttgg acaagcaacc 1680
aaccccaaaa gccaaagggc agggaaaaaa aagaaagaaa gaaaaacgct agtccaaaat 1740
cacacggggt ccgggtctat cgttggattg acgattggct tgcgtcgttc gccgtctgct 1800
ccggggggtg tcacgttgag taggatgcag tgcaactagt tatactagaa aggggatcat 1860
gcagtgcagc cagtgatgtg tgagcatcgg gtggaacata tgtatccatg ggagatggga 1920
cagtgggatc aagaaaatga ggtaatgtac ttatgcctaa ttatgatgta gaatcatctt 1980
cttgttggtg tgattcgtgt gaagagcgat agaaacaact ccgcgggact atgggagttg 2040
gtgtaacagt acgtgctgta ccggagacat gctagctcca gaatgacatc aaggggaaaa 2100
accggtactt tgttatcaaa gctttcatat tggacgcagt gatccccggc ccgtttcagg 2160
agcagatatc tatatagtac tacacagtat acatgactac tacgactacg agtagtatac 2220
tacgagtagt cgttgaagta acatggttaa agtagcaatt acgagaaaaa agaaaaggtt 2280
gctgctactg tacgtgctgg gagccgggga ggccgtcctt gatagccaga tgattcccgt 2340
taataagatt gcagcagggc ttgtacatga atattcccaa gtacatatgg tctacgtacg 2400
ccgtcgagtt gggtgggaag aggaaggcgt atcatctcga ttagctcgtg ttggtcgatg 2460
tggacgactt tggacaaatc ggcaacggat ggaaacaagc gagccattaa agcaaaaccc 2520
gctttctcgc ccttcctttt tccacgcgcc tttggtttta ctttctttcg catctcgccc 2580
cggccccgcc cgggatgaca aggacgcatc gtagattacg gcgacgagag agaaaaattc 2640
cgctcccctc aagttttctg gaagcggcga agaagaagaa gaaccggaga gcggtgacgc 2700
ggacgtgcct gcccaacgca aattccgtcg gtcctcccta cgcccgttcc accgcacacc 2760
gtaacgggag ccccgtatta ttcctcgccg attcgtgcag tgatttggac ccggtgcacg 2820
gtgggagcgt ggtgcggcgc cacacgactc cctttgggga ggaagcttct tatttggacc 2880
ggacgggggc ctgacccacg cagacgtaaa cgccaggagg gccaacccgc cgctggctcg 2940
ctttatatcc cgcgtgagct ccacaccgag cgaagcggag gtccggagga ggaagcggcg 3000
agagaagctc agctaggcag ggcgacgggc agaaacgcga ccacggcaac aaaccccgcc 3060
gcgcgcgccc accgtgccgg ttacatggga gtagaggcgg gcggcggctg cggtgggagg 3120
gcggtagtca ccggattcta cgtctggggc tgggagttcc tcaccgccct cctgctcttc 3180
tcggccacca cctcctacta gctatacaca cccatctcac cataacacac atacatagat 3240
agatagatag atagatacat acacacaaac ataagtagct aggtagagaa agagatcata 3300
gcgttagggg atcc 3314
<210> 3
<211> 1780
<212> DNA
<213> 未知(Unknown)
<400> 3
atggagccca tggacagcca gctcacggct ctcgcgctct ccgactccga cagcgcctcc 60
gtcgagggcg ccgcagctga cgcggcggac gccgacctgc aggcgctccg ccgcctctcc 120
gacaacctcg cggcggcgtt ccgctcccca gacgacttcg ccttcctcgc cgacgcgcgc 180
atcgtggttc ccggcgcgcc cgacctccgc gtgcaccgct gcgtgctctg cgcgcgcagc 240
cccttcctgc gcgacgcctt tgcgcgccgc gccgccagcg ccggcgagga ggagaaggac 300
aaggacagtt acatgtgcaa ggtggagctc cgggatctcc tcggagacga ggtcgaggtc 360
gggtacgacg cgctgcggct ggtgctcgac tacctctaca gcggccgcgt cgccgccctg 420
cccaaggcgg cctgcctctg cgtcgacgag gacgcctgcg cgcacgtcgg ctgccgcccc 480
gccgtcgcct tcatggcgca ggtcctcttc gccgcgtcca ccttcgacgt cgccgagctc 540
accaatctct tccagcggcg gctccttgat gtccttgata aggttgaagt ggacaacctc 600
ccattggtct tatctgttgc caacttatgt agcaaatctt gtgtgaaact gcttgagagg 660
tgccttgatg tagtagttcg ttcaaatctt gatatgattg ctcttgaaaa aaagttgcct 720
ccagatgttg tcaaagaaat tgttgatgca agggtaagcc ttggattagt ttcaccggaa 780
gacaagggct tccctaacat acatgtaaga agaatacaca gagcactgga ttctgatgac 840
gtggagctag tccggatgct actcaaggaa ggaaaaacta atcttgatga tgcatatgca 900
ttacactatg ccgtcgaaca ttgcgactca aagatcacaa cagaacttct ggatctcgcg 960
cttgcagatg tcaatcatag gaacccaaga ggttatacgg tccttcacat tgctgctatg 1020
agaagggaac ctaaaatcat tgtctctctt ttgaccaagg gagctcggcc atcagacctc 1080
acatttgatg acagaaaagc agtgcagatc tctaaacgac ttacaaagca tggggattac 1140
tttgggccta ctgaggacgg aaagccttct cctaaagata gattatgtat tgaggtacta 1200
gagcaagctg aaagaaggga cccgcaactt ggagaagcat cagtttctct tgcgatagaa 1260
ggagactctg cgcgtgggag gctactctac cttgaaaacc gagttgctct agcaagaata 1320
ctgtttccta tggaggcaag ggtagcaatg gacattgctc aagtcgatgg aacactggaa 1380
ttcacccttg tttctagtgt gaatctgcct gccgagatac aacggacagt tgacctgaac 1440
gatactcctt tcacaatgaa ggaagaacac ctggctcgga tgagagccct atccaagaca 1500
gtggaagtcg ggaagcgttt cttcccgcgg tgttcgaagg tacttgacac gatcatggac 1560
gacgaagccg agatggcttc cctcgggcga gacacatccg cggagaagaa gaggaggttc 1620
catgacctgc aagacctggt ccagaaggca ttcagcgagg acaaggagga gaacgacagg 1680
tccgcggcgc ggtcaccttc ctcctcgtcg acgacgacga cgtcgatcgg ggccgtccgg 1740
cctcggaggt gaaagatctg ttctgcacaa agtggagtag 1780
<210> 4
<211> 583
<212> PRT
<213> 未知(Unknown)
<400> 4
Met Glu Pro Met Asp Ser Gln Leu Thr Ala Leu Ala Leu Ser Asp Ser
1 5 10 15
Asp Ser Ala Ser Val Glu Gly Ala Ala Ala Asp Ala Ala Asp Ala Asp
20 25 30
Leu Gln Ala Leu Arg Arg Leu Ser Asp Asn Leu Ala Ala Ala Phe Arg
35 40 45
Ser Pro Asp Asp Phe Ala Phe Leu Ala Asp Ala Arg Ile Val Val Pro
50 55 60
Gly Ala Pro Asp Leu Arg Val His Arg Cys Val Leu Cys Ala Arg Ser
65 70 75 80
Pro Phe Leu Arg Asp Ala Phe Ala Arg Arg Ala Ala Ser Ala Gly Glu
85 90 95
Glu Glu Lys Asp Lys Asp Ser Tyr Met Cys Lys Val Glu Leu Arg Asp
100 105 110
Leu Leu Gly Asp Glu Val Glu Val Gly Tyr Asp Ala Leu Arg Leu Val
115 120 125
Leu Asp Tyr Leu Tyr Ser Gly Arg Val Ala Ala Leu Pro Lys Ala Ala
130 135 140
Cys Leu Cys Val Asp Glu Asp Ala Cys Ala His Val Gly Cys Arg Pro
145 150 155 160
Ala Val Ala Phe Met Ala Gln Val Leu Phe Ala Ala Ser Thr Phe Asp
165 170 175
Val Ala Glu Leu Thr Asn Leu Phe Gln Arg Arg Leu Leu Asp Val Leu
180 185 190
Asp Lys Val Glu Val Asp Asn Leu Pro Leu Val Leu Ser Val Ala Asn
195 200 205
Leu Cys Ser Lys Ser Cys Val Lys Leu Leu Glu Arg Cys Leu Asp Val
210 215 220
Val Val Arg Ser Asn Leu Asp Met Ile Ala Leu Glu Lys Lys Leu Pro
225 230 235 240
Pro Asp Val Val Lys Glu Ile Val Asp Ala Arg Val Ser Leu Gly Leu
245 250 255
Val Ser Pro Glu Asp Lys Gly Phe Pro Asn Ile His Val Arg Arg Ile
260 265 270
His Arg Ala Leu Asp Ser Asp Asp Val Glu Leu Val Arg Met Leu Leu
275 280 285
Lys Glu Gly Lys Thr Asn Leu Asp Asp Ala Tyr Ala Leu His Tyr Ala
290 295 300
Val Glu His Cys Asp Ser Lys Ile Thr Thr Glu Leu Leu Asp Leu Ala
305 310 315 320
Leu Ala Asp Val Asn His Arg Asn Pro Arg Gly Tyr Thr Val Leu His
325 330 335
Ile Ala Ala Met Arg Arg Glu Pro Lys Ile Ile Val Ser Leu Leu Thr
340 345 350
Lys Gly Ala Arg Pro Ser Asp Leu Thr Phe Asp Asp Arg Lys Ala Val
355 360 365
Gln Ile Ser Lys Arg Leu Thr Lys His Gly Asp Tyr Phe Gly Pro Thr
370 375 380
Glu Asp Gly Lys Pro Ser Pro Lys Asp Arg Leu Cys Ile Glu Val Leu
385 390 395 400
Glu Gln Ala Glu Arg Arg Asp Pro Gln Leu Gly Glu Ala Ser Val Ser
405 410 415
Leu Ala Ile Glu Gly Asp Ser Ala Arg Gly Arg Leu Leu Tyr Leu Glu
420 425 430
Asn Arg Val Ala Leu Ala Arg Ile Leu Phe Pro Met Glu Ala Arg Val
435 440 445
Ala Met Asp Ile Ala Gln Val Asp Gly Thr Leu Glu Phe Thr Leu Val
450 455 460
Ser Ser Val Asn Leu Pro Ala Glu Ile Gln Arg Thr Val Asp Leu Asn
465 470 475 480
Asp Thr Pro Phe Thr Met Lys Glu Glu His Leu Ala Arg Met Arg Ala
485 490 495
Leu Ser Lys Thr Val Glu Val Gly Lys Arg Phe Phe Pro Arg Cys Ser
500 505 510
Lys Val Leu Asp Thr Ile Met Asp Asp Glu Ala Glu Met Ala Ser Leu
515 520 525
Gly Arg Asp Thr Ser Ala Glu Lys Lys Arg Arg Phe His Asp Leu Gln
530 535 540
Asp Leu Val Gln Lys Ala Phe Ser Glu Asp Lys Glu Glu Asn Asp Arg
545 550 555 560
Ser Ala Ala Arg Ser Pro Ser Ser Ser Ser Thr Thr Thr Thr Ser Ile
565 570 575
Gly Ala Val Arg Pro Arg Arg
580
<210> 5
<211> 1749
<212> DNA
<213> 未知(Unknown)
<400> 5
atggagccgc cgaccagcca cgtcaccaac gcgttctccg actcggacag cgcgtccgtg 60
gaggaggggg gcgccgacgc ggacgccgac gtggaggcgc tccgccgcct ctccgacaac 120
ctcgccgcgg cgttccgctc gcccgaggac ttcgcgttcc tcgccgacgc gcgcatcgcc 180
gtcccgggcg gcggcggcgg cggcggcgac ctgctggtgc accgctgcgt gctctccgcg 240
cggagcccct tcctgcgcgg cgtcttcgcg cgccgcgccg ccgccgccgc aggcggcggc 300
ggcgaggatg gcggcgagag gctggagctc cggggactcc tcggcggcgg cggcgaggag 360
gtggaggtcg ggtacgaggc gctgcggctg gtgctcgact acctctacag cggccgcgtc 420
ggcgacctgc ccaaggcggc gtgcctctgc gtcgacgagg actgcgccca cgtcgggtgc 480
caccccgccg tcgcgttcat ggcgcaggtc ctcttcgccg cctccacctt ccaggtcgcc 540
gagctcacca acctcttcca gcggcgtctc cttgatgtcc ttgataaggt tgaggtagat 600
aaccttctat tgatcttatc tgttgccaac ttatgcaaca aatcttgcat gaaactgctt 660
gaaagatgcc ttgatatggt agtccggtca aaccttgaca tgattactct tgagaagtca 720
ttgcctccag atgttatcaa gcagattatt gatgcacgcc taagcctcgg attaatttca 780
ccagaaaaca agggatttcc taacaaccat gtgaggagga tacacagagc ccttgactct 840
gacgatgtag agctagtcag gatgctgctc actgaaggac agacaaatct tgatgatgcg 900
tttgcactgc actacgccgt cgaacattgt gactcccaaa ttacaaccga gcttttggat 960
ctcgcacttg cagatgttaa tcatagaaac ccaagaggtt atactgttct tcacattgct 1020
gcgaggcgaa gagagcctaa aatcattgtc tcccttttaa ccaagggggc tcggccagca 1080
gatgttacat tcgatgggag aaaaggggtt caaatctcaa aaagactaac aaaacaaggg 1140
gattactttg gggttaccga agaaggaaaa ccttctccaa aagataggtt atgtattgaa 1200
atactggagc aagctgaaag aagggaccca caactcggag aagcatcagt ttctcttgca 1260
atggcaggtg agagtctacg aggaaggttg ctgtatcttg aaaaccgagt tgctttggcg 1320
aggattatgt ttccgatgga ggcaagagta gcaatggata ttgctcaagt ggatggaact 1380
ttggaattta acctgggttc tggtgcaaat ccacctcctg aaagacaacg gacaactgtt 1440
gatctaaatg aaagtccttt cataatgaaa gaagaacact tagctcggat gacggcactc 1500
tccaaaacag tggagctcgg gaaacgcttt ttcccgcgat gttcgaacgt gctcgacaag 1560
atcatggatg atgaaactga tccggtttcc ctcggaagag acacgtccgc ggagaagagg 1620
aagaggtttc atgacctgca ggatgttctt cagaaggcat tccacgagga caaggaggag 1680
aatgacaggt cggggctctc gtcgtcgtcg tcatcgacat cgatcggggc cattcgacca 1740
aggagatga 1749
<210> 6
<211> 582
<212> PRT
<213> 未知(Unknown)
<400> 6
Met Glu Pro Pro Thr Ser His Val Thr Asn Ala Phe Ser Asp Ser Asp
1 5 10 15
Ser Ala Ser Val Glu Glu Gly Gly Ala Asp Ala Asp Ala Asp Val Glu
20 25 30
Ala Leu Arg Arg Leu Ser Asp Asn Leu Ala Ala Ala Phe Arg Ser Pro
35 40 45
Glu Asp Phe Ala Phe Leu Ala Asp Ala Arg Ile Ala Val Pro Gly Gly
50 55 60
Gly Gly Gly Gly Gly Asp Leu Leu Val His Arg Cys Val Leu Ser Ala
65 70 75 80
Arg Ser Pro Phe Leu Arg Gly Val Phe Ala Arg Arg Ala Ala Ala Ala
85 90 95
Ala Gly Gly Gly Gly Glu Asp Gly Gly Glu Arg Leu Glu Leu Arg Gly
100 105 110
Leu Leu Gly Gly Gly Gly Glu Glu Val Glu Val Gly Tyr Glu Ala Leu
115 120 125
Arg Leu Val Leu Asp Tyr Leu Tyr Ser Gly Arg Val Gly Asp Leu Pro
130 135 140
Lys Ala Ala Cys Leu Cys Val Asp Glu Asp Cys Ala His Val Gly Cys
145 150 155 160
His Pro Ala Val Ala Phe Met Ala Gln Val Leu Phe Ala Ala Ser Thr
165 170 175
Phe Gln Val Ala Glu Leu Thr Asn Leu Phe Gln Arg Arg Leu Leu Asp
180 185 190
Val Leu Asp Lys Val Glu Val Asp Asn Leu Leu Leu Ile Leu Ser Val
195 200 205
Ala Asn Leu Cys Asn Lys Ser Cys Met Lys Leu Leu Glu Arg Cys Leu
210 215 220
Asp Met Val Val Arg Ser Asn Leu Asp Met Ile Thr Leu Glu Lys Ser
225 230 235 240
Leu Pro Pro Asp Val Ile Lys Gln Ile Ile Asp Ala Arg Leu Ser Leu
245 250 255
Gly Leu Ile Ser Pro Glu Asn Lys Gly Phe Pro Asn Asn His Val Arg
260 265 270
Arg Ile His Arg Ala Leu Asp Ser Asp Asp Val Glu Leu Val Arg Met
275 280 285
Leu Leu Thr Glu Gly Gln Thr Asn Leu Asp Asp Ala Phe Ala Leu His
290 295 300
Tyr Ala Val Glu His Cys Asp Ser Gln Ile Thr Thr Glu Leu Leu Asp
305 310 315 320
Leu Ala Leu Ala Asp Val Asn His Arg Asn Pro Arg Gly Tyr Thr Val
325 330 335
Leu His Ile Ala Ala Arg Arg Arg Glu Pro Lys Ile Ile Val Ser Leu
340 345 350
Leu Thr Lys Gly Ala Arg Pro Ala Asp Val Thr Phe Asp Gly Arg Lys
355 360 365
Gly Val Gln Ile Ser Lys Arg Leu Thr Lys Gln Gly Asp Tyr Phe Gly
370 375 380
Val Thr Glu Glu Gly Lys Pro Ser Pro Lys Asp Arg Leu Cys Ile Glu
385 390 395 400
Ile Leu Glu Gln Ala Glu Arg Arg Asp Pro Gln Leu Gly Glu Ala Ser
405 410 415
Val Ser Leu Ala Met Ala Gly Glu Ser Leu Arg Gly Arg Leu Leu Tyr
420 425 430
Leu Glu Asn Arg Val Ala Leu Ala Arg Ile Met Phe Pro Met Glu Ala
435 440 445
Arg Val Ala Met Asp Ile Ala Gln Val Asp Gly Thr Leu Glu Phe Asn
450 455 460
Leu Gly Ser Gly Ala Asn Pro Pro Pro Glu Arg Gln Arg Thr Thr Val
465 470 475 480
Asp Leu Asn Glu Ser Pro Phe Ile Met Lys Glu Glu His Leu Ala Arg
485 490 495
Met Thr Ala Leu Ser Lys Thr Val Glu Leu Gly Lys Arg Phe Phe Pro
500 505 510
Arg Cys Ser Asn Val Leu Asp Lys Ile Met Asp Asp Glu Thr Asp Pro
515 520 525
Val Ser Leu Gly Arg Asp Thr Ser Ala Glu Lys Arg Lys Arg Phe His
530 535 540
Asp Leu Gln Asp Val Leu Gln Lys Ala Phe His Glu Asp Lys Glu Glu
545 550 555 560
Asn Asp Arg Ser Gly Leu Ser Ser Ser Ser Ser Ser Thr Ser Ile Gly
565 570 575
Ala Ile Arg Pro Arg Arg
580
<210> 7
<211> 203
<212> DNA
<213> 未知(Unknown)
<400> 7
gagctcaaga tctgttctgc acaaagtgga gtagtcagtc atcgatcagg aaccagacac 60
cagactttta ttcatacagt gaagtgaagt gaagtgcagt gcagtgagtt gctggttttt 120
gtacaactta gtatgtattt gtatttgtaa aatacttcta tcaataaaat ttctaattcc 180
taaaaccaaa atccaggggt acc 203
<210> 8
<211> 1793
<212> DNA
<213> 未知(Unknown)
<400> 8
tggatccatg gacaccacca tcgacggctt cgccgacagc tacgagatca gcagcaccag 60
cttcgtggcc accgacaaca ccgacagcag catcgtgtac ctggccgccg agcaggtgct 120
gaccggcccg gacgtgagcg ccctgcagct gctgagcaac agcttcgaga gcgtgttcga 180
cagcccggac gacttctaca gcgacgccaa gctggtgctg agcgacggca gggaggtgag 240
cttccacagg tgcgtgctga gcgccaggag cagcttcttc aagagcgccc tggccgccgc 300
caagaaggag aaggacagca acaacaccgc cgccgtgaag ctggagctga aggagatcgc 360
caaggactac gaggtgggct tcgacagcgt ggtgaccgtg ctggcctacg tgtacagcag 420
cagggtgagg ccgccgccga agggcgtgag cgagtgcgcc gacgagaact gctgccacgt 480
ggcctgcagg ccggccgtgg acttcatgct ggaggtgctg tacctggcct tcatcttcaa 540
gatcccggag ctgatcaccc tgtaccagag gcacctgctg gacgtggtgg acaaggtggt 600
gatcgaggac accctggtga tcctgaagct ggccaacatc tgcggcaagg cctgcatgaa 660
gctgctggac aggtgcaagg agatcatcgt gaagagcaac gtggacatgg tgagcctgga 720
gaagagcctg ccggaggagc tggtgaagga gatcatcgac aggaggaagg agctgggcct 780
ggaggtgccg aaggtgaaga agcacgtgag caacgtgcac aaggccctgg acagcgacga 840
catcgagctg gtgaagctgc tgctgaagga ggaccacacc aacctggacg acgcctgcgc 900
cctgcacttc gccgtggcct actgcaacgt gaagaccgcc accgacctgc tgaagctgga 960
cctggccgac gtgaaccaca ggaacccgag gggctacacc gtgctgcacg tggccgccat 1020
gaggaaggag ccgcagctga tcctgagcct gctggagaag ggcgccagcg ccagcgaggc 1080
caccctggag ggcaggaccg ccctgatgat cgccaagcag gccaccatgg ccgtggagtg 1140
caacaacatc ccggagcagt gcaagcacag cctgaagggc aggctgtgcg tggagatcct 1200
ggagcaggag gacaagaggg agcagatccc gagggacgtg ccgccgagct tcgccgtggc 1260
cgccgacgag ctgaagatga ccctgctgga cctggagaac agggtggccc tggcccagag 1320
gctgttcccg accgaggccc aggccgccat ggagatcgcc gagatgaagg gcacctgcga 1380
gttcatcgtg accagcctgg agccggacag gctgaccggc accaagagga ccagcccggg 1440
cgtgaagatc gccccgttcc gcatcctgga ggagcaccag agcaggctga aggccctgag 1500
caagaccgtg gagctgggca agaggttctt cccgaggtgc agcgccgtgc tggaccagat 1560
catgaactgc gaggacctga cccagctggc ctgcggcgag gacgacaccg ccgagaagag 1620
gctgcagaag aagcagaggt acatggagat ccaggagacc ctgaagaagg ccttcagcga 1680
ggacaacctg gagctgggca acagcagcct gaccgacagc accagcagca ccagcaagag 1740
caccggcggc aagaggagca acaggaagct gagccacagg aggaggtaat gag 1793
<210> 9
<211> 593
<212> PRT
<213> 未知(Unknown)
<400> 9
Met Asp Thr Thr Ile Asp Gly Phe Ala Asp Ser Tyr Glu Ile Ser Ser
1 5 10 15
Thr Ser Phe Val Ala Thr Asp Asn Thr Asp Ser Ser Ile Val Tyr Leu
20 25 30
Ala Ala Glu Gln Val Leu Thr Gly Pro Asp Val Ser Ala Leu Gln Leu
35 40 45
Leu Ser Asn Ser Phe Glu Ser Val Phe Asp Ser Pro Asp Asp Phe Tyr
50 55 60
Ser Asp Ala Lys Leu Val Leu Ser Asp Gly Arg Glu Val Ser Phe His
65 70 75 80
Arg Cys Val Leu Ser Ala Arg Ser Ser Phe Phe Lys Ser Ala Leu Ala
85 90 95
Ala Ala Lys Lys Glu Lys Asp Ser Asn Asn Thr Ala Ala Val Lys Leu
100 105 110
Glu Leu Lys Glu Ile Ala Lys Asp Tyr Glu Val Gly Phe Asp Ser Val
115 120 125
Val Thr Val Leu Ala Tyr Val Tyr Ser Ser Arg Val Arg Pro Pro Pro
130 135 140
Lys Gly Val Ser Glu Cys Ala Asp Glu Asn Cys Cys His Val Ala Cys
145 150 155 160
Arg Pro Ala Val Asp Phe Met Leu Glu Val Leu Tyr Leu Ala Phe Ile
165 170 175
Phe Lys Ile Pro Glu Leu Ile Thr Leu Tyr Gln Arg His Leu Leu Asp
180 185 190
Val Val Asp Lys Val Val Ile Glu Asp Thr Leu Val Ile Leu Lys Leu
195 200 205
Ala Asn Ile Cys Gly Lys Ala Cys Met Lys Leu Leu Asp Arg Cys Lys
210 215 220
Glu Ile Ile Val Lys Ser Asn Val Asp Met Val Ser Leu Glu Lys Ser
225 230 235 240
Leu Pro Glu Glu Leu Val Lys Glu Ile Ile Asp Arg Arg Lys Glu Leu
245 250 255
Gly Leu Glu Val Pro Lys Val Lys Lys His Val Ser Asn Val His Lys
260 265 270
Ala Leu Asp Ser Asp Asp Ile Glu Leu Val Lys Leu Leu Leu Lys Glu
275 280 285
Asp His Thr Asn Leu Asp Asp Ala Cys Ala Leu His Phe Ala Val Ala
290 295 300
Tyr Cys Asn Val Lys Thr Ala Thr Asp Leu Leu Lys Leu Asp Leu Ala
305 310 315 320
Asp Val Asn His Arg Asn Pro Arg Gly Tyr Thr Val Leu His Val Ala
325 330 335
Ala Met Arg Lys Glu Pro Gln Leu Ile Leu Ser Leu Leu Glu Lys Gly
340 345 350
Ala Ser Ala Ser Glu Ala Thr Leu Glu Gly Arg Thr Ala Leu Met Ile
355 360 365
Ala Lys Gln Ala Thr Met Ala Val Glu Cys Asn Asn Ile Pro Glu Gln
370 375 380
Cys Lys His Ser Leu Lys Gly Arg Leu Cys Val Glu Ile Leu Glu Gln
385 390 395 400
Glu Asp Lys Arg Glu Gln Ile Pro Arg Asp Val Pro Pro Ser Phe Ala
405 410 415
Val Ala Ala Asp Glu Leu Lys Met Thr Leu Leu Asp Leu Glu Asn Arg
420 425 430
Val Ala Leu Ala Gln Arg Leu Phe Pro Thr Glu Ala Gln Ala Ala Met
435 440 445
Glu Ile Ala Glu Met Lys Gly Thr Cys Glu Phe Ile Val Thr Ser Leu
450 455 460
Glu Pro Asp Arg Leu Thr Gly Thr Lys Arg Thr Ser Pro Gly Val Lys
465 470 475 480
Ile Ala Pro Phe Arg Ile Leu Glu Glu His Gln Ser Arg Leu Lys Ala
485 490 495
Leu Ser Lys Thr Val Glu Leu Gly Lys Arg Phe Phe Pro Arg Cys Ser
500 505 510
Ala Val Leu Asp Gln Ile Met Asn Cys Glu Asp Leu Thr Gln Leu Ala
515 520 525
Cys Gly Glu Asp Asp Thr Ala Glu Lys Arg Leu Gln Lys Lys Gln Arg
530 535 540
Tyr Met Glu Ile Gln Glu Thr Leu Lys Lys Ala Phe Ser Glu Asp Asn
545 550 555 560
Leu Glu Leu Gly Asn Ser Ser Leu Thr Asp Ser Thr Ser Ser Thr Ser
565 570 575
Lys Ser Thr Gly Gly Lys Arg Ser Asn Arg Lys Leu Ser His Arg Arg
580 585 590
Arg
Claims (6)
1.一种提高植物抗病性载体,其特征在于所述载体由表达控制区与植物NPR1基因可操作连接构成,所述表达控制区核苷酸序列为SEQ ID NO.1或SEQ ID NO.2所示。
2.如权利要求1所述提高植物抗病性载体,其特征在于所述植物NPR1基因来自单子叶植物。
3.如权利要求1所述提高植物抗病性载体,其特征在于所述单子叶植物为水稻、玉米或小麦。
4.如权利要求1所述提高植物抗病性载体,其特征在于所述植物NPR1基因核苷酸序列为SEQ ID NO.3或SEQ ID NO.5所示。
5.一种权利要求1所述提高植物抗病性载体在制备抗病植物中的应用。
6.如权利要求5所述的应用,其特征在于所述应用是将表达控制区导入植物内源NPR1基因编码区上游而实现的。
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