CN110878303B - 水稻Os11g0681100基因及其编码蛋白的功能和应用 - Google Patents
水稻Os11g0681100基因及其编码蛋白的功能和应用 Download PDFInfo
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Abstract
本发明涉及植物基因工程技术领域,具体涉及水稻Os11g0681100基因及其编码蛋白的功能和应用。本发明发现水稻Os11g0681100基因及其编码蛋白参与调控水稻对白叶枯病的免疫响应,通过破坏Os11g0681100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。本发明利用CRISPR/Cas9技术实现了高效的Os11g0681100基因的定点敲除,定点敲除水稻植株接种白枯病菌后,病斑长度显著缩短。本发明提供的Os11g0681100基因的新功能为植物抗病育种提供了新方法,在农业生产中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Os11g0681100基因及其编码蛋白在调控植物抗病性中的应用。
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(ht tp://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,本发明提供水稻基因Os11g0681100及其编码的蛋白在调控植物抗病性中的应用。
为实现上述目的,本发明的技术方案如下:
本发明通过对1440份国内外水稻材料接种白叶枯病菌IV后病斑长度的全基因组关联分析(Genome-wide associated study,GWAS)鉴定到白叶枯病抗性相关候选基因Os11g0681100,该基因编码水稻表达蛋白,其核苷酸序列如SEQ ID NO.2所示,编码蛋白的氨基酸序列如SEQ ID NO.1所示。进一步地,本发明利用CRISPR/Cas9技术创制了定点敲除Os11g0681100基因的水稻植株,并对该水稻植株进行白叶枯病抗性评价,发现该基因的敲除能够显著提高水稻对白叶枯病的抗性。
具体地,第一方面,本发明提供水稻Os11g0681100基因、其编码蛋白或水稻Os11g0681100基因的抑制因子在调控植物抗病性中的应用。
第二方面,本发明提供水稻Os11g0681100基因、其编码蛋白或水稻Os11g0681100基因的抑制因子在改良植物抗病性的遗传育种中的应用。
作为优选,本发明所述抗病性为植物对白叶枯病的抗病性。
作为优选,所述遗传育种为构建抗白叶枯病的转基因植物或通过杂交等育种方式培育抗白叶枯病植物。
作为优选,上述应用中,通过破坏水稻Os11g0681100基因的编码蛋白的生物学功能,提高植物的抗病性。
上述抗病性的提高可表现为水稻白枯病的病斑长度缩短。
本发明中,所述水稻Os11g0681100基因的编码蛋白具有如下任一种氨基酸序列:
(1)如SEQ ID NO.1所示的氨基酸序列;
(2)如SEQ ID NO.1所示的氨基酸序列经一个或多个氨基酸的替换、插入或缺失得到的具有相同功能蛋白的氨基酸序列;
(3)与如SEQ ID NO.1所示的氨基酸序列具有至少80%同源性的氨基酸序列;优选地,所述同源性为至少90%;更优选为95%。
上述如SEQ ID NO.1所示的氨基酸序列为水稻Os11g0681100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Os11g0681100基因的编码蛋白具有相同活性的突变体。
本发明中,所述水稻Os11g0681100基因具有如下任一种核苷酸序列:
(1)如SEQ ID NO.2所示的核苷酸序列;
(2)如SEQ ID NO.2所示的核苷酸序列经一个或多个核苷酸的替换、插入或缺失得到的编码相同功能蛋白的核苷酸序列。
上述如SEQ ID NO.2所示的核苷酸序列为水稻Os11g0681100基因的核苷酸序列。本发明所述的水稻Os11g0681100基因可以为任意能够编码上述水稻Os11g0681100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。
本发明中,所述水稻Os11g0681100基因的抑制因子为能够破坏水稻Os11g0681100基因的编码蛋白的生物学功能的核酸。
作为优选,所述核酸为sgRNA或干扰RNA。
作为优选,所述sgRNA的靶序列为所述水稻Os11g0681100基因中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。
更优选地,所述sgRNA的靶序列为如SEQ ID NO.2所示序列的第569位至第588位。
上述水稻Os11g0681100基因、其编码蛋白或水稻Os11g0681100基因的抑制因子的应用可以水稻Os11g0681100基因、其编码蛋白或水稻Os11g0681100基因的抑制因子本身的形式应用,或者以含有水稻Os11g0681100基因或其抑制因子的表达盒、载体、含有所述表达盒或所述载体的宿主细胞的形式应用。
第三方面,本发明提供一种用于突变水稻Os11g0681100基因的sgRNA,所述sgRNA的靶序列为如SEQ ID NO.2所示序列的第569位至第588位。
作为优选,所述sgRNA包含如SEQ ID NO.3所示的序列。
上述sgRNA可与CRISRP/Cas9基因编辑工具配合作用,实现水稻Os11g0681100基因的高效定点敲除,破坏水稻Os11g0681100基因的编码蛋白的生物学功能。
上述sgRNA为本发明通过大量筛选获得,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Os11g0681100基因突变的水稻植株,并且在如SEQ ID NO.2所示序列的第569位至第588位发生插入或缺失均会导致水稻Os11g0681100基因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Os11g0681100基因突变的抗病植物的选育效率。
第四方面,本发明还提供包含所述用于突变水稻Os11g0681100基因的sgRNA的生物材料,所述生物材料包括表达盒、载体、宿主细胞、工程菌或转基因植物细胞系。
第五方面,本发明提供一种调控植物白叶枯病抗性或培育转基因植物的方法,包括:调控植物中水稻Os11g0681100基因的编码蛋白的生物学功能;所述水稻Os11g0681100基因的编码蛋白具有如下任一种氨基酸序列:
(1)如SEQ ID NO.1所示的氨基酸序列;
(2)如SEQ ID NO.1所示的氨基酸序列经一个或多个氨基酸的替换、插入或缺失得到的具有相同功能蛋白的氨基酸序列;
(3)与如SEQ ID NO.1所示的氨基酸序列具有至少80%同源性的氨基酸序列;优选地,所述同源性为至少90%;更优选为95%。
作为优选,通过破坏所述水稻Os11g0681100基因的编码蛋白的生物学功能,提高植物的白叶枯病抗性。
上述破坏水稻Os11g0681100基因编码蛋白的生物学功能可通过本领域常规技术手段实现。
作为优选,利用CRISRP/Cas9技术破坏所述水稻Os11g0681100基因编码蛋白的生物学功能;所述CRISRP/Cas9技术使用的sgRNA的靶序列为如SEQ ID NO.2所示序列的第569位至第588位。
更优选地,所述sgRNA包含如SEQ ID NO.3所示的序列。
利用CRISRP/Cas9技术可使水稻Os11g0681100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插入缺失,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。
作为本发明的一种优选方案,所述调控植物白叶枯病抗性或培育转基因植物的方法包括如下步骤:
(1)构建含有如SEQ ID NO.3所示的sgRNA的CRISRP/Cas9基因编辑质粒;
(2)将步骤(1)构建的CRISRP/Cas9基因编辑质粒转入水稻中;
(3)经筛选和鉴定获得抗水稻白叶枯病材料。
作为优选,上述步骤(1)中,CRISRP/Cas9基因编辑质粒为II型CRISPR系统;
作为优选,上述步骤(2)中,将步骤(1)构建的CRISRP/Cas9基因编辑质粒采用根癌农杆菌介导的方式转入水稻品种中。
作为优选,上述步骤(3)中,所述筛选的方法具体为:利用针对CRISRP/Cas9基因编辑质粒的自身引物对进行PCR扩增,若扩增出744bp,则表明含有SEQ ID NO.3的CRISRP/Cas9基因编辑质粒已成功转化植株。
作为优选,上述步骤(3)中,所述鉴定的方法具体为:利用Os11g0681100基因的特异性引物扩增转化植株基因组的Os11g0681100基因片段并进行测序,若如SEQ ID NO.2所示序列的第569位至第588位的核酸序列发生插入或缺失突变,则说明植株Os11g0681100基因的生物学功能受到破坏。
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。
本发明的有益效果在于:
(1)本发明发现水稻基因Os11g0681100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Os11g0681100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,Os11g0681100定点敲除水稻接种白枯病菌IV病斑长度缩短41.2%,证明通过敲除Os11g0681100基因可制备水稻抗白叶枯病材料,在农业生产中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Os11g0681100基因,实现了高效的Os11g0681100的定点敲除。本发明发现水稻Os11g0681100基因中以第569位至第588位的核苷酸序列作为靶序列,能够高效实现Os11g0681100的定点敲除,并且在如SEQ IDNO.2所示序列的第569位至第588位发生插入或缺失均会导致水稻Os11g0681100基因的编码蛋白被破坏,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Os11g0681100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中基于CRISPR/Cas9技术的水稻表达蛋白Os11g0681100定点敲除方法的载体活性检测的测序峰图。
图2为本发明实施例2中水稻日本晴背景下Cas9-Os11g0681100纯合突变植株中Os11g0681100基因核苷酸序列的突变类型。
图3为本发明实施例2中水稻日本晴背景下Cas9-Os11g0681100纯合突变植株中Os11g0681100基因氨基酸序列的突变类型。
图4本发明实施例2中水稻日本晴背景下Cas9-Os11g0681100纯合突变植株分别接种白叶枯病菌IV后病斑表型统计图,***表示P<0.01。
具体实施方式
下面将结合实施例对本发明的优选实施方式进行详细说明。需要理解的是以下实施例的给出仅是为了起到说明的目的,并不是用于对本发明的范围进行限制。本领域的技术人员在不背离本发明的宗旨和精神的情况下,可以对本发明进行各种修改和替换。
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。
实施例1基于pYLCRISPR/Cas9系统的水稻表达蛋白基因Os11g0681100的定点敲除
1、水稻表达蛋白Os11g0681100的序列分析及靶序列筛选
水稻表达蛋白Os11g0681100的编码基因序列如SEQ ID NO.2所示。序列分析显示,该基因共包括1个外显子,位于如SEQ ID NO.2所示序列的第1-1971位(第一外显子)。
本发明以水稻表达蛋白Os11g0681100第一外显子上的序列为基于CRISPR/Cas9技术的水稻表达蛋白Os11g0681100定点敲除方法的Os11g0681100-T靶序列。
本发明经大量筛选,确定利用pTLCRISPR/Cas9技术靶向水稻Os11g0681100基因第一外显子的反义链第569位至588位作为靶序列(Os11g0681100-T),靶序列如SEQ ID NO.3所示。
pYLCRISPR/Cas9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 RobustCRISPR/Cas9 System for Convenient,High-Effificiency Multiplex Genome Editingin Monocot and Dicot Plants.Mol.Plant.8,1274-1284.”。
2、pYLCRISPR/Cas9Pubi-H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Os11g0681100基因的靶序列Os11g0681100-T的引物Os11g0681100-gRT和Os11g0681100-U6aT,其序列分别如SEQ ID NO.4和SEQ IDNO.5所示
分别合成基于pYLCRISPR/Cas9技术的Os11g0681100-T的相关引物。
SEQ ID NO.4:Os11g0681100-gRT:5′-TGGTACCCTTGCATGTCGAgttttagagctagaaat-3′
SEQ ID NO.5:Os11g0681100-U6aT:5′-TCGACATGCAAGGGTACCACggcagccaagccagca-3′
(2)pYLCRISPR/Cas9技术重组表达载体的构建
以pYLsgRNA-OsU6a载体为模板,使用引物UF(SEQ ID NO.6:5′-CTCCGTTTTACCTGTGGAATCG-3′)和Os11g0681100-U6aT进行PCR扩增,将正确序列命名为U6aT;以pYLsgRNA-OsU6a载体为模板,使用引物gR-R(SEQ ID NO.7:5′-CGGAGGAAAATTCCATCCAC-3′)和Os11g0681100-gRT进行PCR扩增,将正确序列命名为gRT;使用引物Pps-GGL(SEQ ID NO.8:5′-TTCAGAGGTCTCTCTCGACTAGTATGGAATCGGCAGCAAAGG-3′)和Pgs-GGR(SEQ ID NO.9:5′-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-Os11g0681100-sgRNA1;U6a-Os11g0681100-sgRNA1和经BsaI酶处理的pYLCRISPR/Cas9Pubi-H以边酶切边连接的方式获得载体pYLCRISPR/Cas9Pubi-H-Os11g0681100。
3、pYLCRISPR/Cas9Pubi-H-Os11g0681100-T表达载体的活性检测
将重组表达载体pYLCRISPR/Cas9Pubi-H-Os11g0681100-T通过PEG介导导入水稻原生质体,获得重组表达载体pYLCRISPR/Cas9Pubi-H-Os11g0681100-T的瞬时表达结果。基于pYLCRISPR/Cas9技术的水稻表达蛋白Os11g0681100定点敲除方法的载体活性检测测序峰图如图1所示,结果表明所构建的重组表达载体可以对基因Os11g0681100进行编辑。
4、重组根癌农杆菌的获得
将重组表达载体pYLCRISPR/Cas9Pubi-H-Os11g0681100热激转化农杆菌EH105,获得含有重组表达载体pYLCRISPR/Cas9Pubi-H-Os11g0681100的重组农杆菌,命名为EH105-Cas9-Os11g0681100。
根癌农杆菌EHA105:BioVector NTCC典型培养物保藏中心,可市售购得。
实施例2基于pYLCRISPR/Cas9技术的水稻Os11g0681100基因敲除突变体的构建及其白叶枯病抗性检测
采用实施例1构建的重组农杆菌EH105-pYLCRISPR/Cas9Pubi-H-Os11g0681100侵染水稻品种日本晴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-Os11g0681100,具体方法如下:
1、将实施例1获得的重组农杆菌EH105-pYLCRISPR/Cas9Pubi-H-Os11g0681100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。
6、获得的再生植株移栽成活后,提取再生植株的叶片总DNA,利用重组表达载体pYLCRISPR/Cas9Pubi-H-Os11g0681100的自身引物(序列如SEQ ID NO.10和SEQ ID NO.11所示)进行PCR扩增筛选阳性转化植株。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-Os11g0681100转化水稻品种的阳性率检测结果
再生植株 | 再生植株数 | 阳性转化植株数 | 阳性率(%) |
Nip-Cas9-Os11g0681100 | 49 | 42 | 85.7 |
7、以阳性转化植株数的基因组为模板,用水稻表达蛋白Os11g0681100特异性引物Os11g0681100-TF(序列如SEQ ID NO.12所示序列)和Os11g0681100-TR(序列如SEQ IDNO.13所示)进行PCR扩增,将所获得954bp扩增产物进行测序验证。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-Os11g0681100诱导水稻表达蛋白Os11g0681100发生突变的检测结果
再生植株 | 再生植株数 | 发生突变转化植株数 | 突变效率(%) |
Nip-Cas9-Os11g0681100 | 49 | 42 | 85.7 |
8、收集水稻品种日本晴中不同突变植株的种子,分别以自主分离的方式筛选纯合突变植株,共获得1种纯合突变类型的植株(Cas9-Os11g0681100),其中,在如SEQ ID NO.2所示序列的第572位添加碱基A,导致基因Os11g0681100翻译提前终止,其核苷酸序列如图2所示,氨基酸序列如图3所示。
对上述纯合突变类型植株接种白叶枯病菌IV,检测该突变植株的白叶枯病抗性,结果如图4所示,结果显示,与野生型(日本晴)相比,Cas9-Os11g0681100突变体表现出白叶枯病斑变短的表型,接种白叶枯病菌IV的病斑长度缩短41.2%,这说明利用CRISPR/Cas9技术创制的Cas9-Os11g0681100突变体对白叶枯病菌IV的抗病性明显增强,表明使Os11g0681100基因丧失生物学功能能够显著提高水稻的白叶枯病抗性。
日本晴(Nipponbare,Oryza sativa 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 Jiayang Li.Regulationof OsSPL14 by OsmiR156 defines ideal plant architecture in rice.NatureGenetics,2010,42,541-544。
水稻白叶枯病菌菌株IV:记载过该材料的非专利文献是“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”。
虽然,上文中已经用一般性说明、具体实施方式及试验,对本发明作了详尽的描述,但在本发明基础上,可以对之作一些修改或改进,这对本领域技术人员而言是显而易见的。因此,在不偏离本发明精神的基础上所做的这些修改或改进,均属于本发明要求保护的范围。
序列表
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Claims (8)
1.水稻Os11g0681100基因、其编码蛋白或水稻Os11g0681100基因的抑制因子在调控水稻对白叶枯病的抗病性中的应用;
所述应用为通过破坏水稻Os11g0681100基因的编码蛋白的生物学功能,提高水稻对白叶枯病的抗病性;
所述水稻Os11g0681100基因的编码蛋白的氨基酸序列如SEQ ID NO.1所示。
2.水稻Os11g0681100基因、其编码蛋白或水稻Os11g0681100基因的抑制因子在改良水稻对白叶枯病的抗病性的遗传育种中的应用;
所述应用为通过破坏水稻Os11g0681100基因的编码蛋白的生物学功能,提高水稻对白叶枯病的抗病性;
所述水稻Os11g0681100基因的编码蛋白的氨基酸序列如SEQ ID NO.1所示。
3.根据权利要求1或2所述的应用,其特征在于,所述水稻Os11g0681100基因的抑制因子为能够破坏水稻Os11g0681100基因的编码蛋白的生物学功能的核酸。
4.根据权利要求3所述的应用,其特征在于,所述核酸为sgRNA或干扰RNA。
5.根据权利要求4所述的应用,其特征在于,所述sgRNA的靶序列为所述水稻Os11g0681100基因中的XXXNGG形式的核苷酸序列,其中,XXX为19-20 bp的核酸序列,N为A、T、G、C中的任意一个碱基。
6.根据权利要求5所述的应用,其特征在于,所述sgRNA的靶序列为如SEQ ID NO.2所示序列的第569位至第588位。
7.一种提高水稻对白叶枯病抗性或培育抗白叶枯病转基因水稻的方法,其特征在于,包括:通过破坏水稻Os11g0681100基因的编码蛋白的生物学功能,提高水稻对白叶枯病的抗病性;
所述水稻Os11g0681100基因的编码蛋白的氨基酸序列如SEQ ID NO.1所示。
8.根据权利要求7所述的方法,其特征在于,利用CRISPR/Cas9技术破坏所述水稻Os11g0681100基因的编码蛋白的生物学功能;所述CRISPR/Cas9技术使用的sgRNA的靶序列为如SEQ ID NO.2所示序列的第569位至第588位。
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US5977434A (en) * | 1995-01-17 | 1999-11-02 | The Regents Of The University Of California | Procedures and materials for conferring disease resistance in plants |
CN109369790A (zh) * | 2018-12-04 | 2019-02-22 | 中国农业科学院作物科学研究所 | 水稻白枯病抗性相关蛋白OsBBR1及其编码基因与应用 |
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US5977434A (en) * | 1995-01-17 | 1999-11-02 | The Regents Of The University Of California | Procedures and materials for conferring disease resistance in plants |
CN109369790A (zh) * | 2018-12-04 | 2019-02-22 | 中国农业科学院作物科学研究所 | 水稻白枯病抗性相关蛋白OsBBR1及其编码基因与应用 |
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