CN110791487B - 水稻受体激酶基因LOC_Os11g47290及其编码蛋白与应用 - Google Patents
水稻受体激酶基因LOC_Os11g47290及其编码蛋白与应用 Download PDFInfo
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Abstract
本发明公开了水稻受体激酶基因LOC_Os11g47290及其编码蛋白与应用。本发明发现水稻受体激酶基因LOC_Os11g47290及其编码蛋白在调控植物抗病性中的应用，通过破坏LOC_Os11g47290基因的编码蛋白的生物学功能能够显著提高水稻对白叶枯病的抗性。本发明利用CRISPR/Cas9技术实现了高效的LOC_Os11g47290基因定点敲除，定点敲除水稻接种白叶枯病菌GD1358和V后，病斑长度显著缩短。本发明提供的LOC_Os11g47290基因的新功能为植物抗病育种提供了新方法，在农业生产上具有十分重要的应用价值。
Description
技术领域
本发明涉及植物基因工程技术领域，具体涉及水稻受体激酶基因LOC_Os11g47290及其编码蛋白与应用。
背景技术
由黄单胞杆菌水稻致病变种(Xanthomonas oryzae pv.oryzae)引起的白叶枯病是制约水稻生产的一种重要细菌性病害，对于水稻种植产业危害严重，一般可致水稻减产20％-30％左右，严重可达50％。利用抗性基因培育和种植抗病品种是防治水稻白叶枯病最经济有效的措施。然而，目前已报道的44个水稻白叶枯病抗性基因/位点(http://www.shigen.nig.ac.jp/rice/oryzabase/gene/list)大部分表现出抗谱较窄或者难以利用的问题，仅Xa3、Xa4、Xa21和Xa23等基因在生产中得以广泛应用。
由于黄单胞杆菌水稻致病变种易于变异，水稻与白叶枯病菌(黄单胞杆菌水稻致病变种)的协同进化导致品种抗性极易丧失。因此，通过鉴定并敲除白叶枯病易感性基因，提高水稻品种的抗病性，对于水稻抗病育种具有重要的应用价值。
发明内容
为了解决现有技术存在的问题，本发明的一个目的是提供水稻受体激酶LOC_Os11g47290及其编码蛋白在调控植物抗病性中的应用。
本发明提供了如下A-C中任一种物质在调控植物抗病性中的应用；
A、LOC_Os11g47290蛋白；
B、所述LOC_Os11g47290蛋白的编码核酸；
C、含有所述核酸的表达盒、重组载体或重组微生物。
本发明还提供了如下b1或b2所示的物质在提高植物抗病性或改良植物抗病性中的应用：b1、抑制或降低植物中LOC_Os11g47290蛋白活性或者含量的物质；b2、抑制或降低植物中LOC_Os11g47290蛋白编码核酸表达的物质。
上述应用中，所述LOC_Os11g47290蛋白为如下(a1)-(a4)中任一种：
(a1)序列表中序列1所示的蛋白质；
(a2)在(a1)所述蛋白质的N端或/和C端连接标签得到的融合蛋白；
(a3)将(a1)经过一个或几个氨基酸残基的取代和/或缺失和/或添加得到的与植物抗病相关的蛋白质；
(a4)与(a1)具有98％以上同一性且与植物抗病性相关的蛋白质。
上述应用中，所述LOC_Os11g47290蛋白编码核酸为如下(b1)-(b3)中任一种：
(b1)编码区为序列表中序列2所示的DNA分子；
(b2)与(b1)具有95％以上同一性且编码所述蛋白质的DNA分子；
(b3)在严格条件下与(b1)或(b2)任一种限定的核苷酸序列杂交且编码所述蛋白的DNA分子。
上述如序列1所示的氨基酸序列为水稻LOC_Os11g47290基因的编码蛋白序列，本领域技术人员可根据本发明公开的氨基酸序列以及氨基酸的保守性替换等本领域常规技术手段，在不影响其活性的前提下，取代、缺失和/或增加一个或几个氨基酸，得到与本发明公开的水稻LOC_Os11g47290基因的编码蛋白具有相同活性的突变体。
上述如序列2所示的核苷酸序列为水稻LOC_Os11g47290基因的核苷酸序列。本发明所述的水稻LOC_Os11g47290基因可以为任意能够编码上述水稻LOC_Os11g47290基因的编码蛋白的核苷酸序列。考虑到密码子的简并性以及不同物种密码子的偏爱性，本领域技术人员可以根据需要使用适合特定物种表达的密码子。
上述应用中，所述抗病性为抗白叶枯病。
上述应用中，所述b1或b2所示的物质为LOC_Os11g47290蛋白或LOC_Os11g47290蛋白编码核酸的抑制因子，抑制因子可以为核酸，也可以含有所述核酸的表达盒、载体、重组菌或宿主细胞的形式存在；抑制因子具体如下：
1)干扰RNA；
2)CRISPR/Cas9***；
所述CRISPR/Cas9***中，所述sgRNA的靶序列为所述LOC_Os11g47290蛋白的编码核酸中的XXXNGG形式的核苷酸序列，其中，XXX为所述LOC_Os11g47290蛋白的编码核酸中任意19-20bp的核酸序列，N为A、T、G、C中的任意一个碱基。
利用CRISRP/Cas9***可使水稻LOC_Os11g47290基因中XXXNGG形式的核苷酸序列在NGG上游3-4bp处切割产生DNA双链断裂，从而引入核苷酸序列的***缺失，进而造成该基因的翻译提前终止或蛋白构象改变，最终破坏该基因的编码蛋白的生物学功能；其中XXXNGG形式的核苷酸序列，其中，XXX为19-20bp的核酸序列，N为A、T、G、C中的任意一个碱基。
优选为，XXX为LOC_Os11g47290蛋白的编码核酸(基因组)中第一外显子中任意19-20bp的核酸序列；在本发明的实施例中，更优选地，所述sgRNA为sgRNA1和/或sgRNA2；所述sgRNA1的靶序列为序列3(水稻LOC_Os11g47290基因的第367位至第386位)；所述sgRNA2的靶序列为序列4(水稻LOC_Os11g47290基因的第952位至第971位)。
在本发明的实施例中，pYLCRISPR/Cas9***包括如下任一种重组载体(CRISRP/Cas9基因编辑质粒)：
重组表达载体pYLCRISPR/Cas9Pubi-H-47290-T1含有U6a-47290-sgRNA1和cas9编码基因，U6a-47290-sgRNA1具有20个核苷酸组成的靶序列区，靶序列区在LOC_Os11g47290基因上对应的靶序列为序列3所示的47290-T1；该重组表达载体的具体构建方法见实施例；
重组表达载体pYLCRISPR/Cas9Pubi-H-47290-T2含有U6b-47290-sgRNA2和cas9编码基因，U6b-47290-sgRNA2具有20个核苷酸组成的靶序列区，靶序列区在LOC_Os11g47290基因上对应的靶序列为序列4所示的47290-T2；该重组表达载体的具体构建方法见实施例。
重组表达载体pYLCRISPR/Cas9Pubi-H-47290含有U6a-47290-sgRNA-1、U6b-47290-sgRNA-2和cas9编码基因，U6a-47290-sgRNA-1具有20个核苷酸组成的靶序列区，靶序列区在LOC_Os11g47290基因上对应的靶序列为序列3所示的47290-T1；U6b-47290-sgRNA-2具有20个核苷酸组成的靶序列结合区，靶序列区在LOC_Os11g47290基因上对应的靶序列为序列4所示的47290-T2；该重组表达载体的具体构建方法见实施例。
上述应用中，所述遗传育种为构建抗白叶枯病的转基因植物。
上述抗病性的提高可表现为植物白枯病的病斑长度减少。
本发明另一个目的是提供一种提高植物抗病性的方法，为如下1)-3)中任一种方法，实现提高植物抗病性；
1)包括如下步骤：抑制或降低目的植物中LOC_Os11g47290蛋白活性或者含量；
2)包括如下步骤：抑制或降低目的植物中LOC_Os11g47290蛋白编码核酸表达；
3)包括如下步骤：对目的植物中LOC_Os11g47290蛋白编码核酸进行基因编辑；
本发明还提供了一种制备高抗病性的转基因植物的方法，为如下1)-3)中任一种方法，
1)包括如下步骤：抑制或降低目的植物中LOC_Os11g47290蛋白活性或者含量，得到转基因植物；
2)包括如下步骤：抑制或降低目的植物中LOC_Os11g47290蛋白编码核酸表达，得到转基因植物；
3)包括如下步骤：对目的植物中LOC_Os11g47290蛋白编码核酸进行基因编辑，得到转基因植物；
所述转基因植物中的抗病性高于所述目的植物；
所述目的植物为野生型植物或受体植物。
上述方法还包括如下步骤：从转基因植物或基因编辑后植物中选取LOC_Os11g47290蛋白编码核酸被编辑的植株，为抗病性高的转基因植物。
所述从转基因植物或基因编辑后植物中选取LOC_Os11g47290蛋白编码核酸被编辑的植株的选取方法包括:1)直接扩增含有LOC_Os11g47290蛋白编码核酸抑制因子的载体片段；2)扩增并测序编辑后植株中含有LOC_Os11g47290蛋白编码核酸靶序列的基因组片段。
在本发明的实施例中，扩增并测序编辑后植株中含有LOC_Os11g47290蛋白编码核酸靶序列的基因组片段中采用的扩增引物具体为序列11所示的单链DNA分子和序列12所示的单链DNA分子组成的引物。
上述方法中，所述抗病性为抗白叶枯病。
上述方法中，所述抑制或降低植物中LOC_Os11g47290蛋白活性或者含量，或，抑制或降低植物中LOC_Os11g47290蛋白编码核酸表达是通过对LOC_Os11g47290蛋白编码核酸进行基因编辑实现。
上述方法中，所述基因编辑是借助CRISPR/Cas9***实现的；
所述CRISPR/Cas9***中，所述sgRNA的靶序列为所述LOC_Os11g47290蛋白的编码核酸中的XXXNGG形式的核苷酸序列，其中，XXX为所述LOC_Os11g47290蛋白的编码核酸中任意19-20bp的核酸序列，N为A、T、G、C中的任意一个碱基；
优选为，XXX为LOC_Os11g47290蛋白的编码核酸(基因组)中第一外显子中任意19-20bp的核酸序列；在本发明的实施例中，更优选地，所述sgRNA为sgRNA1和/或sgRNA2；所述sgRNA1的靶序列为序列3(水稻LOC_Os11g47290基因的第367位至第386位)；所述sgRNA2的靶序列为序列4(水稻LOC_Os11g47290基因的第952位至第971位)。
上述方法中，基因编辑具体为先构建含有序列3或/和序列4所示的sgRNA靶序列结合区的CRISRP/Cas9基因编辑质粒，再将该CRISRP/Cas9基因编辑质粒转入植物中，实现基因编辑。其中，CRISRP/Cas9基因编辑质粒为II型CRISPR***。在本发明的实施例中，pYLCRISPR/Cas9***包括如下任一种重组载体(pYLCRISRP/Cas9基因编辑质粒)：重组表达载体pYLCRISPR/Cas9Pubi-H-47290-T1、重组表达载体pYLCRISPR/Cas9Pubi-H-47290-T2或重组表达载体pYLCRISPR/Cas9Pubi-H-47290。
本发明还有一个目的是提供一种特异sgRNA或含有所述sgRNA编码基因的表达盒、载体、宿主细胞、工程菌或转基因植物细胞系。
本发明提供的特异sgRNA为sgRNA1和/或sgRNA2；
所述sgRNA1的靶序列为序列3；所述sgRNA2的靶序列为序列4。
在本发明中，所述植物可为单子叶植物或双子叶植物，优选为白叶枯病菌的寄主植物，包括但不限于水稻。
本发明通过实验证明，利用上述sgRNA进行CRISRP/Cas9介导的基因编辑能够高效获得LOC_Os11g47290基因突变的水稻植株，并且在如序列2所示序列的第367位至第386位或/和第952位至第971位发生***或缺失均会导致水稻LOC_Os11g47290基因的编码蛋白的生物学功能被破坏，使得水稻表现出白叶枯病抗性水平提高的性状，进而有效提高基于LOC_Os11g47290基因突变的抗病植物的选育效率。
本发明的有益效果在于：
(1)本发明发现水稻受体激酶基因LOC_Os11g47290及其编码蛋白参与调控水稻对白叶枯病菌的免疫响应，通过破坏LOC_Os11g47290基因编码蛋白的生物学功能能够显著提高水稻对白叶枯病的抗性。经实验验证：分别对LOC_Os11g47290定点敲除水稻接种白叶枯病菌，接种白叶枯病菌V叶片病斑长度缩短14.4％，接种白叶枯病菌GD1358叶片病斑长度缩短48.1％，证明通过突变LOC_Os11g47290基因的核苷酸序列制备水稻抗白叶枯病材料，在农业生产上具有十分重要的应用价值。
(2)本发明利用CRISRP/Cas9技术进行基因组靶向修饰LOC_Os11g47290基因，实现了高效的LOC_Os11g47290的定点敲除。本发明发现水稻LOC_Os11g47290基因中以第367位至第386位或/和第952位至第971位的核苷酸序列作为靶序列，能够高效实现LOC_Os11g47290的定点敲除，并且在如序列2所示序列的第367位至第386位或/和第952位至第971位发生***或缺失均会导致水稻LOC_Os11g47290基因的编码蛋白的生物学功能被破坏，使得水稻表现出白叶枯病抗性水平提高的性状，有效提高了基于LOC_Os11g47290基因突变的抗病植物的选育效率。
附图说明
图1为本发明实施例1提供的基于pYLCRISPR/Cas9技术水稻受体激酶LOC_Os11g47290定点敲除方法的载体活性检测测序峰图。
图2为本发明实施例2提供的水稻日本晴背景下Cas9-47290纯合突变植株中LOC_Os11g47290基因核苷酸序列的突变类型；下划线处核苷酸序列为靶序列，黑色框内为替换或***的核苷酸序列。
图3为本发明实施例2提供的水稻日本晴背景下Cas9-47290纯合突变植株中LOC_Os11g47290基因氨基酸序列的突变类型。
图4为本发明实施例2提供的水稻日本晴背景下Cas9-47290纯合突变植株分别接种白叶枯病菌GD1358和V后病斑表型统计图，**表示P<0.05。
具体实施方式
下述实施例中所使用的实验方法如无特殊说明，均为常规方法。
下述实施例中所用的材料、试剂等，如无特殊说明，均可从商业途径得到。
下述实施例中日本晴(Nipponbare,Oryza satiIVa ssp.japonica:记载过该材料的非专利文献是Yongqing Jiao，Yonghong Wang，Dawei Xue，Jing Wang，Meixian Yan,Guifu Liu，Guojun Dong，Dali Zeng，Zefu Lu，Xudong Zhu，Qian Qian and JiayangLi.Regulation of OsSPL14 by OsmiR156 defines ideal plant architecture inrice.Nature Genetics，2010,42，541-544；公众可从申请人处获得。
水稻白叶枯病菌菌株V：记载在“曾列先，黄少华，伍尚忠.IRBB21(Xa21)对广东稻白叶枯病菌5个小种的抗性反应.植物保护学报，2002,29(2):97-100”一文中，经广东省农业科学院曾列先老师同意后，公众可以从申请人处获得。
水稻白叶枯病菌菌株粤1358(GD1358)：记载在“方中达，许志刚，过崇俭，殷尚智，伍尚忠，徐羡明，章琦.中国水稻白叶枯病菌致病型的研究.植物病理学报，1990，20(2)：81-88”一文中，公众可以从申请人处获得。
实施例1、基于pYLCRISPR/Cas9***的水稻受体激酶基因LOC_Os11g47290定点敲除方法
一、水稻受体激酶基因LOC_Os11g47290的序列分析及靶序列筛选
水稻受体激酶基因LOC_Os11g47290的核苷酸序列为序列2所示，其编码的蛋白的氨基酸序列为序列表中序列1。序列分析显示，该基因共包括7个外显子，分别为序列2序列的第1-1696位(第一外显子)、第2042-2941位(第二外显子)、第3130-3278位(第三外显子)、第3677-3890位(第四外显子)、第6492-6611位(第五外显子)、第6976-7019位(第六外显子)、7163-7171位(第七外显子)。
以水稻受体激酶基因LOC_Os11g47290第一外显子上的序列为基于pYLCRISPR/Cas9***的水稻受体激酶基因LOC_Os11g47290定点敲除方法的47290-T1和47290-T2靶序列。
经大量筛选，确定利用pYLCRISPR/Cas9技术靶向水稻LOC_Os11g47290基因第一外显子的反义链第367位至第386位和正义链第952位至第971位分别作为靶序列47290-T1和47290-T2，靶序列如序列3和序列4所示。
pYLCRISPR/Cas9***载体(包括pYLsgRNA-OsU6a、pYLsgRNA-OsU6b和pYLCRISPR/Cas9Pubi-H等载体):记载过该材料的非专利文献是“Ma X.,Zhang Q.,Zhu Q.,Liu W.,Chen Y.,Qiu R.,Wang B.,Yang Z.,Li H.,Lin Y.,Xie Y.,Shen R.,Chen S.,Wang Z.,Chen Y.,Guo J.,Chen L.,Zhao X.,Dong Z.,and Liu Y.-G.(2015).A Robust CRISPR/Cas9 System for ConIVenient,High-Effificiency Multiplex Genome Editing inMonocot and Dicot Plants.Mol.Plant.8,1274–1284.”。经华南农业大学生命科学学院亚热带农业生物资源保护与利用国家重点实验室刘耀光老师同意后，公众可从申请人处获得该载体。
二、pYLCRISPR/Cas9***载体引物设计及其重组表达载体的构建
1、pYLCRISPR/Cas9技术靶序列引物的设计与合成
基于pYLCRISPR/Cas9技术设计靶向LOC_Os11g47290基因的靶序列引物，47290-T1靶序列引物47290-gRT1和47290-U6aT1序列分别如序列5和序列6所示；47290-T2靶序列引物47290-gRT2和47290-U6bT2序列分别如序列7和序列8所示；
分别合成基于pYLCRISPR/Cas9技术的47290-T1和47290-T2的相关引物。
序列5：47290-gRT1：5′-TGCAGCCTTCCTATAACACCgttttagagctagaaat-3′
序列6：47290-U6aT1：5′-GGTGTTATAGGAAGGCTGCACggcagccaagccagca-3′
序列7：47290-gRT2：5′-CATGTGCCGGAATGGTTAGCgttttagagctagaaat-3′
序列8：47290-U6bT2：5′-GCTAACCATTCCGGCACATGCaacacaagcggcagca-3′。
2、pYLCRISPR/Cas9技术重组表达载体的构建
1)单独含47290-T1或47290-T2的重组表达载体的构建
重组表达载体pYLCRISPR/Cas9Pubi-H-47290-T1为含有U6a-47290-sgRNA1和cas9编码基因，U6a-47290-sgRNA1具有20个核苷酸组成的靶序列区，靶序列区在LOC_Os11g47290基因上对应的靶序列为序列3所示的47290-T1；该重组表达载体的具体构建方法如下：
以pYLsgRNA-OsU6a载体为模板，使用引物UF(5′-CTCCGTTTTACCTGTGGAATCG-3′)和47290-U6aT1进行PCR扩增，将正确序列命名为U6aT1；以pYLsgRNA-OsU6a载体为模板，使用引物gR-R(5′-CGGAGGAAAATTCCATCCAC-3′)和47290-gRT1进行PCR扩增，将正确序列命名为gRT1；使用引物Pps-GGL(5′-TTCAGAGGTCTCTCTCGACTAGTATGGAATCGGCAGCAAAGG-3′)和Pgs-GGR(5′-AGCGTGGGTCTCGACCGACGCGTATCCATCCACTCCAAGCTC-3′)通过巢式PCR的方式将两个片段连接在一起，并将其命名为U6a-47290-sgRNA1；
再用BsaI酶同时对U6a-47290-sgRNA1和pYLCRISPR/Cas9Pubi-H进行酶切，以边酶切边连接的方式获得载体pYLCRISPR/Cas9Pubi-H-47290-T1。
重组表达载体pYLCRISPR/Cas9Pubi-H-47290-T2含有U6b-47290-sgRNA2和cas9编码基因，U6b-47290-sgRNA2具有20个核苷酸组成的靶序列区，靶序列区在LOC_Os11g47290基因上对应的靶序列为序列4所示的47290-T2；该重组表达载体的具体构建方法如下：
以pYLsgRNA-OsU6b载体为模板，使用引物UF(5′-CTCCGTTTTACCTGTGGAATCG-3′)和47290-U6bT2进行PCR扩增，将正确序列命名为U6bT2；以pYLsgRNA-OsU6b载体为模板，使用引物gR-R(5′-CGGAGGAAAATTCCATCCAC-3′)和47290-gRT2进行PCR扩增，将正确序列命名为gRT2；使用引物Pps-GGL(5′-TTCAGAGGTCTCTCTCGACTAGTATGGAATCGGCAGCAAAGG-3′)和Pgs-GGR(5′-AGCGTGGGTCTCGACCGACGCGTATCCATCCACTCCAAGCTC-3′)通过巢式PCR的方式将两个片段连接在一起，并将其命名为U6b-47290-sgRNA2；
再用BsaI酶同时对U6b-47290-sgRNA2和pYLCRISPR/Cas9Pubi-H载体进行酶切，以边酶切边连接的方式获得载体pYLCRISPR/Cas9Pubi-H-47290-T2。
2)含47290-T1和47290-T2的重组表达载体的构建
pYLCRISPR/Cas9Pubi-H-47290含有U6a-47290-sgRNA-1、U6b-47290-sgRNA-2和cas9编码基因，U6a-47290-sgRNA-1具有20个核苷酸组成的靶序列区，靶序列区在LOC_Os11g47290基因上对应的靶序列为序列3所示的47290-T1；U6b-47290-sgRNA-2具有20个核苷酸组成的靶序列区，靶序列区在LOC_Os11g47290基因上对应的靶序列为序列4所示的47290-T2。具体构建方法如下：1)使用引物Pps-GGL(5′-TTCAGAGGTCTCTCTCGACTAGTATGGAATCGGCAGCAAAGG-3′)和Pgs-GG2(5′-AGCGTGGGTCTCGTCAGGGTCCATCCACTCCAAGCTC-3)通过巢式PCR的方式将U6aT1、gRT1连接在一起，并将其命名为U6a-47290-sgRNA-1；2)使用引物Pps-GG2(5′-TTCAGAGGTCTCTCTGACACTGGAATCGGCAGCAAAGG-3′)和Pgs-GGR(5′-AGCGTGGGTCTCGACCGACGCGTATCCATCCACTCCAAGCTC-3′)通过巢式PCR的方式将U6bT2、gRT2连接在一起，并将其命名为U6b-47290-sgRNA-2；3)将BsaI酶同时对U6a-47290-sgRNA-1、U6b-47290-sgRNA-2和pYLCRISPR/Cas9Pubi-H进行酶切，以边酶切边连接的方式获得载体pYLCRISPR/Cas9Pubi-H-47290。
3、重组表达载体的活性检测
将上述2制备的重组表达载体pYLCRISPR/Cas9Pubi-H-47290-T1和pYLCRISPR/Cas9Pubi-H-47290-T2分别通过PEG介导导入(方法参见https://bio-protocol.org/bio101/e1010125)水稻日本晴原生质体，16小时后分别获得瞬时转导pYLCRISPR/Cas9Pubi-H-47290-T1和pYLCRISPR/Cas9Pubi-H-47290-T1质粒的原生质体。
分别提取瞬时转导pYLCRISPR/Cas9Pubi-H-47290-T1和pYLCRISPR/Cas9Pubi-H-47290-T2质粒的原生质体的基因组DNA，用序列表中序列11和序列12扩增LOC_Os11g47290基因的部分核苷酸序列，并进行测序验证。
结果如图1所示，图1为基于pYLCRISPR/Cas9技术水稻受体激酶基因LOC_Os11g47290定点敲除方法的载体活性检测测序峰图，可以看出，pYLCRISPR/Cas9技术可在靶序列47290-T1和47290-T2处诱导LOC_Os11g47290基因的突变。
4、重组根癌农杆菌的获得
将上述2获得的重组表达载体pYLCRISPR/Cas9Pubi-H-47290热激转化农杆菌EH105，获得含有重组表达载体pYLCRISPR/Cas9Pubi-H-47290的重组农杆菌，命名为EH105-Cas9-47290。
根癌农杆菌EHA105：BioIVector NTCC典型培养物保藏中心，可市售购得。
实施例2、基于pYLCRISPR/Cas9技术定点敲除方法在水稻品种的应用
一、pYLCRISPR/Cas9技术定点敲除LOC_Os11g47290基因
将重组农杆菌EH105-Cas9-47290侵染水稻品种日本晴(以下简称为野生型水稻)成熟胚诱导的愈伤组织，将获得的水稻转化植株分别命名为NIP-Cas9-47290；实验具体方法如下：
1、将实施例1获得的重组农杆菌接种于YEB液体培养基(含50μg/ml卡那霉素和20μg/ml利福平)中，28℃、200rpm条件下振荡培养至OD600为0.6-0.8；以5000rpm、4℃离心5min，用AAM液体培养基(乙酰丁香酮浓度为200μM/L，pH 5.2)重悬菌体沉淀浓度至OD600为0.6-0.8。
2、分别将水稻品种日本晴的成熟种子除去颖壳，在75％乙醇中浸泡1min，然后在NaClO溶液中(与水1:2混合，加1滴吐温20)振荡消毒20min，重复2次。经无菌水冲洗数次至无异味，将消毒后的水稻日本晴种子接种于NBD2培养基上诱导愈伤组织，26℃黑暗培养8-10天，切除根和残留胚乳，继代培养10天，获得成熟胚愈伤组织。
3、将步骤2获得的成熟胚愈伤组织分别浸于步骤1获得的重组农杆菌重悬液中，20-30min后移出水稻材料，接种于含有两层滤纸的共培养培养基(乙酰丁香酮浓度为100μM/L，pH 5.2)上，26℃黑暗条件下共培养3天。
4、将经过步骤3共培养的愈伤组织接种于筛选培养基(潮霉素浓度为50mg/L，pH5.8)中，28℃黑暗条件下筛选培养12天，转移抗性愈伤组织至含有50mg/L Hyg的选择培养基上继续筛选。
5、重复筛选2次后，转移抗性愈伤组织至分化培养基上(24小时光照/天)诱导分化；待新的无根幼苗生成，转移再生幼苗至1/2MS培养基上诱导生根；待小苗茁壮后移入人工气候室进行营养液栽培，得到再生植株NIP-Cas9-47290。
6、获得的再生植株移栽成活后，提取再生植株的叶片总DNA，基于重组表达载体pYLCRISPR/Cas9Pubi-H-47290的自身引物序列9、序列10所示序列进行PCR扩增筛选阳性转化植株，扩增出1225bp大小条带的植株即为阳性转化植株。
统计检测的再生植株数、阳性转化植株数及阳性转化植株数占检测的再生植株数的百分比即阳性率(％)，结果如表1所示。
表1.pYLCRISPR/Cas9Pubi-H-47290转化水稻品种的阳性率检测结果
7、以阳性转化植株数的基因组为模板，用水稻受体激酶基因LOC_Os11g47290特异性引物LOC_Os11g47290-TF如序列11所示序列和LOC_Os11g47290-TF如序列12所示序列(扩增基因LOC_Os11g47290中含有2个靶序列的片段)进行PCR扩增。以日本晴为对照。
将所获得1096bp的扩增产物进行测序验证，与日本晴相比，扩增产物有突变的记作发生突变转化植株数；测序验证结果如表2所示。统计检测的再生植株数、发生突变转化植株数及发生突变转化植株数占检测的再生植株数的百分比即突变效率(％)，结果如表2所示。
表2.pYLCRISPR/Cas9Pubi-H-47290诱导水稻受体激酶基因LOC_Os11g47290发生突变的检测结果
二、pYLCRISPR/Cas9技术定点敲除47290基因突变转化植株的表型
收集上述一获得的39株发生突变转化植株的种子，播种后，获得T1代植株。
提取200个T1代植株的基因组DNA，用序列11所示引物和序列12所示的引物进行扩增，得到PCR产物，送去测序，选取纯合突变株，获得1种纯合突变类型，该突变类型40株。
纯合突变类型的突变形式如下：两靶序列47290-T1和47290-T2均发生突变且在靶序列1的第371位碱基T替换为碱基A，第380位碱基G替换为碱基A；靶序列2的第968位与969位之间***碱基T，导致基因LOC_Os11g47290翻译提前终止。该种纯合突变类型的核苷酸序列如图2所示，氨基酸序列如图3所示。将具有该突变形式的植株命名为T1代Cas9-47290突变植株。
将T1代Cas9-47290突变植株(cas9-47290)接种白叶枯病菌GD1358、V。以野生型日本晴为对照。每个株系接种15株；实验重复3次，结果取平均值。
接种白叶枯病菌GD1358、V 14天后，观察叶片表型。
统计接菌叶片病斑长度，结果如图4所示，与野生型(条形图横坐标中的日本晴)相比，Cas9-47290突变体表现出白叶枯病斑变短的表型，接种白叶枯病菌GD1358的叶片病斑长度缩短48.1％，接种白叶枯病菌V的叶片病斑长度缩短14.4％。
上述结果说明利用CRISPR/Cas9技术创制的cas9-47290突变体增强了水稻对白叶枯病菌GD1358、V的抗病性。
最后应说明的是：以上实施例仅用以说明本发明的技术方案，而非对其限制；尽管参照前述实施例对本发明进行了详细的说明，本领域的普通技术人员应当理解：其依然可以对前述各实施例所记载的技术方案进行修改，或者对其中部分技术特征进行等同替换；而这些修改或者替换，并不使相应技术方案的本质脱离本发明各实施例技术方案的精神和范围。
SEQUENCE LISTING
<110> 中国科学院遗传与发育生物学研究所 中国农业科学院作物科学研究所
<120>水稻受体激酶基因LOC_Os11g47290及其编码蛋白与应用
<160> 12
<170> PatentIn version 3.5
<210> 1
<211> 1044
<212> PRT
<213> Artificial sequence
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His Arg Leu Glu Leu Leu Asp Leu Gly Tyr Asn Ala Leu Ser Gly Asn
130 135 140
Ile Pro Ala Thr Ile Gly Asn Leu Thr Lys Leu Glu Leu Leu Asn Leu
145 150 155 160
Glu Phe Asn Gln Leu Ser Gly Pro Ile Pro Ala Glu Leu Gln Gly Leu
165 170 175
Arg Ser Leu Gly Ser Met Asn Leu Arg Arg Asn Tyr Leu Ser Gly Leu
180 185 190
Ile Pro Asn Ser Leu Phe Asn Asn Thr Pro Leu Leu Gly Tyr Leu Ser
195 200 205
Ile Gly Asn Asn Ser Leu Ser Gly Pro Ile Pro His Val Ile Phe Ser
210 215 220
Leu His Val Leu Gln Val Leu Val Leu Glu His Asn Gln Leu Ser Gly
225 230 235 240
Ser Leu Pro Pro Ala Ile Phe Asn Met Ser Arg Leu Glu Lys Leu Tyr
245 250 255
Ala Thr Arg Asn Asn Leu Thr Gly Pro Ile Pro Tyr Pro Ala Glu Asn
260 265 270
Gln Thr Leu Met Asn Ile Pro Met Ile Arg Val Met Cys Leu Ser Phe
275 280 285
Asn Gly Phe Ile Gly Arg Ile Pro Pro Gly Leu Ala Ala Cys Arg Lys
290 295 300
Leu Gln Met Leu Glu Leu Gly Gly Asn Leu Leu Thr Asp His Val Pro
305 310 315 320
Glu Trp Leu Ala Gly Leu Ser Leu Leu Ser Thr Leu Val Ile Gly Gln
325 330 335
Asn Glu Leu Val Gly Ser Ile Pro Val Val Leu Ser Asn Leu Thr Lys
340 345 350
Leu Thr Val Leu Asp Leu Ser Ser Cys Lys Leu Ser Gly Ile Ile Pro
355 360 365
Leu Glu Leu Gly Lys Met Thr Gln Leu Asn Ile Leu His Leu Ser Phe
370 375 380
Asn Arg Leu Thr Gly Pro Phe Pro Thr Ser Leu Gly Asn Leu Thr Lys
385 390 395 400
Leu Ser Phe Leu Gly Leu Glu Ser Asn Leu Leu Thr Gly Gln Val Pro
405 410 415
Glu Thr Leu Gly Asn Leu Arg Ser Leu Tyr Ser Leu Gly Ile Gly Lys
420 425 430
Asn His Leu Gln Gly Lys Leu His Phe Phe Ala Leu Leu Ser Asn Cys
435 440 445
Arg Glu Leu Gln Phe Leu Asp Ile Gly Met Asn Ser Phe Ser Gly Ser
450 455 460
Ile Ser Ala Ser Leu Leu Ala Asn Leu Ser Asn Asn Leu Gln Tyr Phe
465 470 475 480
Tyr Ala Asn Asp Asn Asn Leu Thr Gly Ser Ile Pro Ala Thr Ile Ser
485 490 495
Asn Leu Ser Asn Leu Asn Val Ile Gly Leu Phe Asp Asn Gln Ile Ser
500 505 510
Gly Thr Ile Pro Asp Ser Ile Met Leu Met Asp Asn Leu Gln Ala Leu
515 520 525
Asp Leu Ser Ile Asn Asn Leu Phe Gly Pro Ile Pro Gly Gln Ile Gly
530 535 540
Thr Pro Lys Gly Met Val Ala Leu Ser Leu Ser Gly Asn Asn Leu Ser
545 550 555 560
Ser Tyr Ile Pro Asn Gly Gly Ile Pro Lys Tyr Phe Ser Asn Leu Thr
565 570 575
Tyr Leu Thr Ser Leu Asn Leu Ser Phe Asn Asn Leu Gln Gly Gln Ile
580 585 590
Pro Ser Gly Gly Ile Phe Ser Asn Ile Thr Met Gln Ser Leu Met Gly
595 600 605
Asn Ala Gly Leu Cys Gly Ala Pro Arg Leu Gly Phe Pro Ala Cys Leu
610 615 620
Glu Lys Ser Asp Ser Thr Arg Thr Lys His Leu Leu Lys Ile Val Leu
625 630 635 640
Pro Thr Val Ile Val Ala Phe Gly Ala Ile Val Val Phe Leu Tyr Leu
645 650 655
Met Ile Ala Lys Lys Met Lys Asn Pro Asp Ile Thr Ala Ser Phe Gly
660 665 670
Ile Ala Asp Ala Ile Cys His Arg Leu Val Ser Tyr Gln Glu Ile Val
675 680 685
Arg Ala Thr Glu Asn Phe Asn Glu Asp Asn Leu Leu Gly Val Gly Ser
690 695 700
Phe Gly Lys Val Phe Lys Gly Arg Leu Asp Asp Gly Leu Val Val Ala
705 710 715 720
Ile Lys Ile Leu Asn Met Gln Val Glu Arg Ala Ile Arg Ser Phe Asp
725 730 735
Ala Glu Cys His Val Leu Arg Met Ala Arg His Arg Asn Leu Ile Lys
740 745 750
Ile Leu Asn Thr Cys Ser Asn Leu Asp Phe Arg Ala Leu Phe Leu Gln
755 760 765
Phe Met Pro Asn Gly Asn Leu Glu Ser Tyr Leu His Ser Glu Ser Arg
770 775 780
Pro Cys Val Gly Ser Phe Leu Lys Arg Met Glu Ile Met Leu Asp Val
785 790 795 800
Ser Met Ala Met Glu Tyr Leu His His Glu His His Glu Val Val Leu
805 810 815
His Cys Asp Leu Lys Pro Ser Asn Val Leu Phe Asp Glu Glu Met Thr
820 825 830
Ala His Val Ala Asp Phe Gly Ile Ala Lys Met Leu Leu Gly Asp Asp
835 840 845
Asn Ser Ala Val Ser Ala Ser Met Leu Gly Thr Ile Gly Tyr Met Ala
850 855 860
Pro Val Phe Glu Leu Gly Leu Leu Cys Ser Ala Asp Ser Pro Glu Gln
865 870 875 880
Arg Thr Ala Met Ser Asp Val Val Val Thr Leu Lys Lys Ile Arg Lys
885 890 895
Asp Tyr Val Lys Leu Met Ala Thr Thr Arg Pro Gly Lys Lys Leu Met
900 905 910
Ala Thr Thr Ala Asn Arg Thr Ser Lys Gly Pro Gln Asp Asn Arg Val
915 920 925
Phe Arg Glu His Ile Phe Arg Leu Ser Cys Thr Gln Lys Arg Ser Asp
930 935 940
Ser Gln Met Arg Ala Gly Thr Ile Thr Gly Ser His Leu Ser Lys Thr
945 950 955 960
Asn Glu Ala Val Gly Gln Arg Arg Lys Lys Val Val Gly Thr Gly Asp
965 970 975
Asn Ser Arg Pro Ala Leu His Glu Leu Gln Gly Arg Glu Lys Gly Lys
980 985 990
Glu Glu Gly Gly Arg Gly Gly Ser Asp Trp Ile Gly Gly Ala Gly Asp
995 1000 1005
Arg Trp Glu Arg Ser Pro Ala His Gln Ala Tyr Pro Leu Met Gly
1010 1015 1020
Pro Glu Arg Lys Glu His Ser Lys Glu Arg Lys Gly Arg Lys Arg
1025 1030 1035
Asn Gly Glu Glu Asn Phe
1040
<210> 2
<211> 7171
<212> DNA
<213> Artificial sequence
<400> 2
atgggtgttg gtcctcattg tactactagt ctgctaataa tactggccgt cgtcatcacg 60
tcgtctttgc tcacgacgac gatcaaggca gatgagccga gcaatgatac cgacatcgcc 120
gcgctgcttg ccttcaaggc acagttctct gaccctctgg gtttcctccg tgacggctgg 180
agggaggaca atgcatcctg cttctgccag tggatcggcg tgtcgtgcag ccgccgccgg 240
cagcgcgtca ccgccctgga gctgccgggc attcccctgc aagggtcgat cacccctcac 300
ctcggtaacc tctctttcct ctacgtcctc aacctcgcca acaccagcct cacggggaca 360
ctcccgggtg ttataggaag gctgcatcgc ctggagctcc ttgatcttgg ctacaatgcc 420
ctgtcaggta acatcccagc caccatagga aacctcacca aacttgagct tcttaatctc 480
gagtttaacc agctatctgg tccaatccca gcagagctgc agggcctgcg aagccttggc 540
agtatgaatc tccgtaggaa ctatctcagt ggcttgattc ccaacagtct attcaacaac 600
accccattgt taggttatct cagcattggc aacaacagct tgtcagggcc aataccgcac 660
gtgatattct cgttgcacgt gctgcaggtc cttgttctag agcacaatca attgtccggc 720
tcactgcccc cagccatctt caacatgtcc agacttgaaa agctgtatgc cactcgaaac 780
aatctcactg gacctatccc atacccagct gaaaaccaga ccttgatgaa catccccatg 840
attcgggtga tgtgtctctc tttcaacgga ttcataggcc gaattccacc tgggcttgcg 900
gcatgccgga aactccagat gcttgagtta ggtgggaatc tcttgacgga tcatgtgccg 960
gaatggttag cgggcttgtc cctgctaagc accttggtta taggtcagaa tgagcttgtc 1020
ggttcgatcc cagttgtgct aagcaatctc accaagctca ccgtgcttga tctgtcatct 1080
tgcaagctaa gtggaatcat tccattggaa ctaggaaaga tgacacaact caacatcttg 1140
cacctctcat ttaatcgcct aactggtcct tttcctacct cccttggtaa cttgacaaaa 1200
ttatcttttc taggattaga atctaacctg ctgaccggac aagtacctga gacccttggg 1260
aacctcaggt ctctatactc ccttggtatt ggaaagaatc atctacaagg gaaacttcac 1320
ttctttgccc ttctctccaa ttgtagggaa ctccaattcc tcgacatagg aatgaattct 1380
ttctcaggga gcatttctgc gagtttacta gcaaacctct ctaacaactt acaatatttt 1440
tatgcaaatg ataacaactt aactggcagt attcctgcta ccatatcaaa tctgtctaac 1500
ctgaatgtaa taggcctttt tgacaaccaa ataagcggca caattccaga ttctataatg 1560
ctaatggata atctacaggc attggacctc tctataaaca atttgtttgg accaatccca 1620
ggacaaattg gtaccccaaa aggaatggtc gcattatctc tcagtggcaa caatctttct 1680
agttacatcc ctaacggtgt tggcaatcta agcacgttgc agtactgatt tctgtcatat 1740
aataggttgt catcagttat acccgcaagc ttagttaatc ttagtaatct tctccaacta 1800
gatatttcta ataataactt aactggttca ttgccttctg atctcagttc cttcaaagta 1860
ataggcctaa tggacatctc agcaaataat ttggtcggta gcctcccaac ttcgttggga 1920
tagctccaac tgtcaagcta cctgaattta tctcaaaaca cattcaatga ttcaattcca 1980
gactctttca aaggtctaat taatttagaa acattggatc tgtctcataa caatctttca 2040
ggaggcatac caaaatactt ttccaactta acctatctta cttctttgaa cctctccttt 2100
aacaatctac aaggtcagat accaagtgga ggtatttttt caaacatcac tatgcaatct 2160
ttaatgggaa atgctggact ctgtggtgct ccacgtctgg gatttcctgc atgtctggag 2220
aagtccgact cgactagaac aaaacacttg ctgaagattg tgctccctac tgtcattgtg 2280
gcgtttggtg ccattgttgt gttcctatac ctaatgattg caaagaaaat gaaaaatcca 2340
gatattacgg cttcttttgg catagcagat gcgatttgcc acaggctagt gtcctaccaa 2400
gaaatcgttc gcgctaccga aaatttcaat gaggacaacc tacttggagt tggaagtttt 2460
ggcaaagttt tcaagggtcg gctggatgac ggtttggtgg ttgcaatcaa aatcctcaac 2520
atgcaggttg aacgagctat taggagtttt gatgcagagt gccatgtctt gcggatggcc 2580
agacatcgca acctgataaa gatactaaat acatgttcca acttggattt cagagcactg 2640
tttcttcagt tcatgcccaa tggaaacttg gagtcatact tgcactctga aagcaggcct 2700
tgtgtgggat cattcctcaa aaggatggag attatgctag atgtgtcaat ggctatggaa 2760
tatctgcatc atgaacacca tgaggttgtc ctgcattgtg atttgaaacc tagcaacgtg 2820
ctatttgatg aagagatgac tgcacatgta gcagactttg gcatagcaaa gatgttgtta 2880
ggggatgaca attcagcagt ttcagcaagc atgctaggca caattgggta catggcacca 2940
ggtacttaca ctagccaatc taaagccaca tacatttttt gttggagttg ttacaaattc 3000
acttcaagat attggccggg cttctgacga agcggagcgg aggcccgtcg ccggaggctt 3060
gggccggagc cagtgggttg gtgtatcgtg tagccgccgc aggtaacatg catggcttcc 3120
ttgtgccagt gttcgagctg ggcttgctct gttcggctga ctcccccgag caaaggacgg 3180
cgatgagcga tgtggtcgtg acactgaaga agattagaaa ggactatgtc aaattgatgg 3240
caaccacccg acccggcaaa aaattgatgg caaccacagt aagcgttgtg cagcagtgat 3300
tcatcgctct atcgttgtat atgagcgaat gaaatacata tcatttgcat cattttcttc 3360
ttctgcatta ggaatagcat cagtgatcga ttaccctttg tttgtatttg tgtatggttg 3420
aattgaatat atatatatat atatatatat atatataaca atttcgttgg tgtaaatatg 3480
tgattgaacc gctggtcaat aaatttgcat cacgaaaatg ggagtagatg atgtgctact 3540
tatgttttct tatttctggc caaaataaat aaataaaaaa ggaatattat cggcacagca 3600
tcacaactcc ggctcgttca gccttaaaca accacaatta acagtcctaa gcagagaaac 3660
ttaacaagct tttcaggcta acagaacatc aaaaggtcca caagacaaca gggtcttcag 3720
agagcacatc ttcaggctgt catgcacgca aaaaagatca gacagccaga tgagagcggg 3780
tacaataacc ggtagccatc tcagtaaaac taacgaggct gtaggacaga gaagaaaaaa 3840
ggtagtagga acgggcgaca atagtcggcc agctctacat gagctccaag gtgaaaaagc 3900
ctacaagata ggaattttca atttttaaaa atggccatgg gcccacaagt agtgagatgc 3960
atgcaatgaa agatgctagt cagttgggtc ccacaagaaa aagaaataca tctcgggatt 4020
aactgctagc ttactctagt taatatggac atggatggag ccggcagcca gcaatactat 4080
tgaacctgct taagagcaag tgcaatagta ggctatatac cagctataaa catactttaa 4140
agagataaag gaagagagag aggaatagca gattacagca cggattacaa gacgtaatat 4200
gtgtataaca tgtgagacca gatattaata gtatagtaag caactattgt atgaactagg 4260
aaggaagccc gcgcagatgc gcaggcatct aatttattgt tttatgattt atgaaataat 4320
gctaaaagat gtcatgtctc atctttttta tgctgtgaga taaactgact ttaatatact 4380
tagtaaatga taagaatgtc tccaagatta aaagaaaatc agtctgttcc gattacttat 4440
aaattctcct attgtcacca caacttatta ttattgtttt aaaagttatg atctataaat 4500
agataaaact ggatatgtac caataaaata agcctaatac atcattgata cactaaaata 4560
ttcactactt atctaagtgt tcatgtttca cttatttccg aatcaatata taaatatttt 4620
aaaaatacca cttataccaa ttaatgaaat aacaccgtta aagataaaca tgaatgcccc 4680
tctaacgggt gaagcccatc gtagccacca agactttctt tctctgttga gttttctaaa 4740
aaaaacgaaa ataaaatttt tgataaatta atccaaaaaa ttaaaactaa attggattta 4800
cttaaaaaat agactaaatt gttttttatc attttttaca tttgtagatt taaattagtg 4860
ttgtatattt atcaagtgat attagaaatc ctaaaatata atgtgaaact atttgaaaat 4920
taccacaaca ttaattatta tatgttggtc ccatgtgtca tgatgtattt aggaccataa 4980
tatacctttt ataagacata agtatttcat ttaattaaag catgaaacta atcacaaatt 5040
gaatataatt gttgcttact atactattaa tatatggtcc cacctgtcat acgcatattg 5100
cgtcttgtag tccgtgctgc agctggctac aaatctgtag cccgctgctc ttctctctca 5160
tcgtttatct cattaaaata tatttgtagc tggctaatag cttgctattt tacttgctct 5220
aagataacag tcctaccaat ttaaataaaa attaatgatg agattagaca aagtcacagg 5280
gtggcctaca agagttggta gaaatggcac ataagataaa atatgatata gtagtaaaaa 5340
tatagctaat atagctcagg aattggtgat ttaattaaaa aaacctacag ctgctaaagt 5400
gtatgcgtag atcgtctata aaaataattg tttatttaca acaccagatc acttctattg 5460
taaattccta gcgtttgcgc attcacactc agattactct ctatgaaaaa actgatctaa 5520
taatttatga taataaatta tgatgtacat gtatcatgta atttgaaaaa agaagaaaaa 5580
aaggagaatt aggatctgga tggcatcaat cgagctcgac gaggcagatc agtctccatg 5640
gaaatgataa caaattatga tgcacatgta ttgtgtaatt agaaataaaa ggagagaggg 5700
agaatccaga tatggatggc atcaagctaa cagaagcata agtatgtacg tacgtactta 5760
tgggcggaaa atctaatggg cgatcaatcg ctaccgcgat ctggtagcga tcgcatcccc 5820
tcccccctcc ctccttccac gtttcctttt ctggcaccgc attacttttc tattttagta 5880
aaatttatgc acctaaagtt tatacaccta aagtttatag acccaaagtt tagagaccca 5940
aagtttataa atcaaaagtt tatatatccg attcaaattt aaatttgaat tcaaatattt 6000
tttatatata gtatttatat acatctaaag tttatacacc taaagtttat agacccaaag 6060
tttataaatc aaaagtttat ataccctatt caaatttgaa tttgaaatat attcgattca 6120
aaatttgaat ttgaattcaa atatttttta tatatagtat ttagatacat ctaaagttta 6180
tacacctaaa gtttatagac ccaaagttta gagacccaaa gtttacatac ccaattcaaa 6240
tttgaatttg aattgtatcc gattcaaatt taaatttgaa ttcaaatatt tttatatata 6300
gtatttctat acatctaaag tttatacacc taaagtttat agaccgaaag tttttagtaa 6360
aaagtttata tacccgattc aaatttgaat ttgaattcaa atattagttt atagatccaa 6420
agtttataag tcaaaaattt acataaccgt ttcaattctg aatttaaatt taaatattta 6480
tggtgtagta ggaagagaaa aaggaaagga ggaagggggg agaggaggga gcgactggat 6540
aggaggggcg ggtgatcgct gggagcgatc acccgcccat caggcatacc cacttatggg 6600
tccggagaga aagggtgaga aagatttagt tttgatccaa cggttaatat tattgggtcc 6660
accactttaa ataaaaactt acggtcagat atttttcttt ttctcagaat attatttaaa 6720
ttattagagc gccatgtggc gacttaggag cgtttatata taggagtctc acgtggtagc 6780
ttgagagcgt acgtagaaag tttaatgaac ttttagtata taaataatag atagatagat 6840
ttaggatttt ctctaatttg ctagaacgcc acgtggtggc ctaggatcgt tattggtcca 6900
aataatatgt aattacttat ttatataaat aatatgtaat ttagaagtat acttattcaa 6960
tatgtattgt tgcagaacat agtaaagaga ggaaggggag gaaaagaaac ggagaggagg 7020
tactcataca tgacgcacgg caggattcct tctttgtttt ttagaaagta agagaataaa 7080
ccaatctaat ctaacggctt ggagtaacgg gcccactaat tttaatgaaa attaatggct 7140
agatgttttg ctttttttat agaatttcta g 7171
<210> 3
<211> 20
<212> DNA
<213> Artificial sequence
<400> 3
tgcagccttc ctataacacc 20
<210> 4
<211> 20
<212> DNA
<213> Artificial sequence
<400> 4
catgtgccgg aatggttagc 20
<210> 5
<211> 37
<212> DNA
<213> Artificial sequence
<400> 5
tgcagccttc ctataacacc gttttagagc tagaaat 37
<210> 6
<211> 37
<212> DNA
<213> Artificial sequence
<400> 6
ggtgttatag gaaggctgca cggcagccaa gccagca 37
<210> 7
<211> 37
<212> DNA
<213> Artificial sequence
<400> 7
catgtgccgg aatggttagc gttttagagc tagaaat 37
<210> 8
<211> 37
<212> DNA
<213> Artificial sequence
<400> 8
gctaaccatt ccggcacatg caacacaagc ggcagca 37
<210> 9
<211> 22
<212> DNA
<213> Artificial sequence
<400> 9
gcggtgtcat ctatgttact ag 22
<210> 10
<211> 22
<212> DNA
<213> Artificial sequence
<400> 10
ccgacataga tgcaataact tc 22
<210> 11
<211> 20
<212> DNA
<213> Artificial sequence
<400> 11
atgagccgag caatgatacc 20
<210> 12
<211> 20
<212> DNA
<213> Artificial sequence
<400> 12
ccaagggagg taggaaaagg 20
Claims (5)
1.抑制或降低水稻中LOC_Os11g47290蛋白编码核酸表达的物质在提高水稻抗白叶枯病性或改良水稻抗白叶枯病性中的应用：
所述LOC_Os11g47290蛋白为如下（a1）-（a2）中任一种：
（a1）序列表中序列1所示的蛋白质；
（a2）在（a1）所述蛋白质的N端或/和C端连接标签得到的融合蛋白；
所述物质为如下：
1）干扰RNA；
2）CRISPR/Cas9***；
所述CRISPR/Cas9***中，sgRNA的靶序列为所述LOC_Os11g47290蛋白的编码核酸中的XXXNGG形式的核苷酸序列，其中，XXX为所述LOC_Os11g47290蛋白的编码核酸中任意19-20bp的核酸序列，N为A、T、G、C中的任意一个碱基。
2.根据权利要求1所述的应用，其特征在于：
所述LOC_Os11g47290蛋白编码核酸为编码区为序列表中序列2所示的DNA分子。
3.根据权利要求2所述的应用，其特征在于：
所述sgRNA为sgRNA1和/或sgRNA2；
所述sgRNA1的靶序列为序列3；所述sgRNA2的靶序列为序列4。
4.一种提高水稻抗白叶枯病性的方法，为如下1）-2）中任一种方法，实现提高水稻抗白叶枯病性；
1）包括如下步骤：抑制或降低目的水稻中LOC_Os11g47290蛋白编码核酸表达；
2）包括如下步骤：对目的水稻中LOC_Os11g47290蛋白编码核酸进行基因编辑；
或，一种制备高抗白叶枯病性的转基因水稻的方法，为如下1）-2）中任一种方法，
1）包括如下步骤：抑制或降低目的水稻中LOC_Os11g47290蛋白编码核酸表达，得到转基因水稻；
2）包括如下步骤：对目的水稻中LOC_Os11g47290蛋白编码核酸进行基因编辑，得到转基因水稻；
所述转基因水稻中的抗白叶枯病性高于所述目的水稻；
所述LOC_Os11g47290蛋白为如下（a1）-（a2）中任一种：
（a1）序列表中序列1所示的蛋白质；
（a2）在（a1）所述蛋白质的N端或/和C端连接标签得到的融合蛋白；
所述抑制或降低水稻中LOC_Os11g47290蛋白编码核酸表达是通过对LOC_Os11g47290蛋白编码核酸进行基因编辑实现的，所述基因编辑是借助CRISPR/Cas9***实现的；
所述CRISPR/Cas9***中， sgRNA的靶序列为所述LOC_Os11g47290蛋白的编码核酸中的XXXNGG形式的核苷酸序列，其中，XXX为所述LOC_Os11g47290蛋白的编码核酸中任意19-20 bp的核酸序列，N为A、T、G、C中的任意一个碱基。
5.根据权利要求4所述的方法，其特征在于：
所述sgRNA为sgRNA1和/或sgRNA2；
所述sgRNA1的靶序列为序列3；所述sgRNA2的靶序列为序列4。
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