CN114805507B - 水稻OsREIN1T219I蛋白及其编码基因与应用 - Google Patents
水稻OsREIN1T219I蛋白及其编码基因与应用 Download PDFInfo
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- CN114805507B CN114805507B CN202110118542.9A CN202110118542A CN114805507B CN 114805507 B CN114805507 B CN 114805507B CN 202110118542 A CN202110118542 A CN 202110118542A CN 114805507 B CN114805507 B CN 114805507B
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Abstract
本发明公开了水稻OsREIN1T219I蛋白及其编码基因与应用。本发明所要保护的OsREIN1T219I蛋白为序列表中序列1所示的蛋白或由序列1衍生的或与序列1中所示蛋白具有80%以上的同一性且具有相同功能的蛋白质。本发明的实验证明,含有OsREIN1T219I蛋白纯合编码基因的水稻材料MC41与含有野生型OsREIN1蛋白编码基因的水稻材料Nip相比,对稻瘟菌具有更高的抗性,OsREIN1T219I可应用于分子设计育种培育高抗病水稻。
Description
技术领域
本发明涉及生物技术领域,具体涉及水稻OsREIN1T219I蛋白及其编码基因与应用。
背景技术
水稻是最主要的粮食作物之一,水稻的侵染性病害有300余种,常年发生的水稻病害有29种,例如,稻瘟病、白叶枯病、稻纹枯病及近年来逐渐蔓延的稻曲柄、稻黑粉病等,这些病害的发生和流行是影响水稻高产稳产的主要因素,因此水稻抗病相关基因的发掘得到极大关注。
等位基因是指在一对同源染色体上,占有相同座位的一对基因,它控制一对相对性状。不同的等位基因会导致一些遗传特征的变化。等位变异(allelic variation) 使人们有可能利用优异等位,培育优良品种。目前已经发展了许多有效的分子生物学技术用于优异等位基因的发掘,例如,SSR标记、自然群体的关联分析等等。利用上述技术已经鉴定了众多与耐逆、抗病、优质和高产等性状相关的优异等位及其载体,广泛应用于培育品种最常用的有效方法杂交育种中。
细胞编程性死亡(Programmed cell death,PCD)是指由基因控制的,细胞主动、有序的死亡,是一种生命现象。PCD普遍存在于植物组织器官分化、生长和发育过程中。其典型的生化特征为细胞收缩、核浓缩、染色质边缘化、DNA被剪切后由膜包成凋亡小体等等。在植物和病原菌互作中,寄主受病原菌侵染的部位经常出现一种形态多样的坏死斑,可以是单细胞死亡,也会有一个较大的坏死区域,成班或条状等。这种坏死现象1915年Stakman命名为超敏反应(hypersensitive response,HR),是植物抗病性的常见反应。目前,HR反应定义为:植物细胞为限制病原菌生长的一种快速死亡。
发明内容
本发明所要解决的技术问题是如何提高水稻的抗病性或如何培育高抗病性水稻。
为了解决上述技术问题,本发明首先提供了一种蛋白质。所述蛋白质为OsREIN1T219I。所述OsREIN1T219I可为如下A1)、A2)或A3)的蛋白质:
A1)氨基酸序列是序列表中序列1的蛋白质;
A2)将序列表中序列1所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加得到的且具有相同功能的由A1)衍生的或与A1)所示的蛋白质具有80%以上的同一性且具有相同功能的蛋白质;
A3)在A1)或A2)的N末端或/和C末端连接蛋白标签得到的融合蛋白质。
上述A2)中所述的蛋白质也就是将序列表中序列1所示的氨基酸序列除第219位氨基酸为异亮氨酸不变外,其他的氨基酸序列可经过一个或几个氨基酸残基的取代和/ 或缺失和/或添加得到的且具有相同功能的由A1)衍生的或与A1)所示的蛋白质具有 80%以上的同一性且具有相同功能的蛋白质。
上述蛋白质中,序列表中序列1由981个氨基酸残基组成。
上述蛋白质可人工合成,也可先合成其编码基因,再进行生物表达得到。
上述蛋白质中,所述蛋白标签(protein-tag)是指利用DNA体外重组技术,与目的蛋白一起融合表达的一种多肽或者蛋白,以便于目的蛋白的表达、检测、示踪和/ 或纯化。所述蛋白标签可为Flag标签、His标签、MBP标签、HA标签、myc标签、GST 标签和/或SUMO标签等。
上述蛋白质中,同一性是指氨基酸序列的同一性。可使用国际互联网上的同源性检索站点测定氨基酸序列的同一性,如NCBI主页网站的BLAST网页。例如,可在高级BLAST2.1中,通过使用blastp作为程序,将Expect值设置为10,将所有Filter设置为OFF,使用BLOSUM62作为Matrix,将Gap existence cost,Per residue gap cost 和Lambda ratio分别设置为11,1和0.85(缺省值)并进行检索一对氨基酸序列的同一性进行计算,然后即可获得同一性的值(%)。
上述蛋白质中,所述80%以上的同一性可为至少81%、82%、85%、86%、88%、90%、 91%、92%、95%、96%、98%、99%或100%的同一性。
为了解决上述技术问题,本发明还提供了与上述蛋白质相关的生物材料。所述生物材料可为下述B1)至B9)中的任一种:
B1)编码上述蛋白质的核酸分子;
B2)含有B1)所述核酸分子的表达盒;
B3)含有B1)所述核酸分子的重组载体、或含有B2)所述表达盒的重组载体;
B4)含有B1)所述核酸分子的重组微生物、或含有B2)所述表达盒的重组微生物、或含有B3)所述重组载体的重组微生物;
B5)含有B1)所述核酸分子的转基因植物细胞系、或含有B2)所述表达盒的转基因植物细胞系;
B6)含有B1)所述核酸分子的转基因植物组织、或含有B2)所述表达盒的转基因植物组织;
B7)含有B1)所述核酸分子的转基因植物器官、或含有B2)所述表达盒的转基因植物器官;
B8)提高或促进权利要求1中所述蛋白质表达的核酸分子;
B9)含有B8)所述核酸分子的表达盒、重组载体、重组微生物或转基因植物细胞系。
上文所述的生物材料中,B1)所述核酸分子可为如下b1)、b2)或b3)所示的所述蛋白质的编码基因:
b1)编码序列是序列表中序列2的核苷酸的cDNA分子或DNA分子;
b2)核苷酸是序列表中序列2的DNA分子;
b3)与b2)限定的cDNA或DNA分子杂交且编码具有相同功能的蛋白质的cDNA分子或DNA分子。
上述生物材料中,B2)所述的含有核酸分子的表达盒,是指能够在宿主细胞中表达上述应用中所述蛋白质的DNA,该DNA不但可包括启动蛋白编码基因转录的启动子,还可包括终止蛋白编码基因转录的终止子。进一步,所述表达盒还可包括增强子序列。可用于本发明的启动子包括但不限于:组成型启动子,组织、器官和发育特异的启动子,和诱导型启动子。
可用现有的植物表达载体构建含有所述蛋白编码基因表达盒的重组表达载体。所述植物表达载体包括双元农杆菌载体和可用于植物微弹轰击的载体等。如pAHC25、pWMB123、pBin438、pCAMBIA1302、pCAMBIA2301、pCAMBIA1301、pCAMBIA1300、pBI121、pCAMBIA1391-Xa或pCAMBIA1391-Xb(CAMBIA公司)等。所述植物表达载体还可包含外源基因的3’端非翻译区域,即包含聚腺苷酸信号和任何其它参与mRNA加工或基因表达的DNA片段。所述聚腺苷酸信号可引导聚腺苷酸加入到mRNA前体的3’端,如农杆菌冠瘿瘤诱导(Ti)质粒基因(如胭脂碱合成酶基因Nos)、植物基因(如大豆贮存蛋白基因)3’端转录的非翻译区均具有类似功能。使用本发明的基因构建植物表达载体时,还可使用增强子,包括翻译增强子或转录增强子,这些增强子区域可以是ATG起始密码子或邻接区域起始密码子等,但必需与编码序列的阅读框相同,以保证整个序列的正确翻译。
上述生物材料中,所述重组微生物具体可为酵母,细菌,藻和真菌。
上文所述蛋白质和/或上文所述生物材料的下述任一种应用也属于本发明的保护范围:
Q1、所述蛋白质和/或所述生物材料在调控植物抗病性中的应用,
Q2、所述蛋白质和/或所述生物材料在制备提高植物抗病性的产品中的应用,
Q3、所述蛋白质和/或所述生物材料在培育抗病性植物中的应用,
Q4、所述蛋白质和/或所述生物材料在制备植物抗病性产品中的应用,
Q5、所述蛋白质和/或所述生物材料在植物育种中的应用。
为了解决上述技术问题,本发明还提供了一种培育抗病植物的方法。所述方法包括提高目的植物中上文所述蛋白质的活性或/和上文所述蛋白质的编码基因的表达量,得到抗病植物。所述抗病植物的抗病性高于所述目的植物的抗病性。
所述提高目的植物中上文所述蛋白质的活性或/和上文所述蛋白质的编码基因的表达量是通过将上文所述蛋白质的编码基因导入所述目的植物实现的。
上述方法中,所述蛋白的编码基因可先进行如下修饰,再导入目的植物中,以达到更好的表达效果:
1)与各种植物表达的启动子连接,以利于其在植物中的表达;所述启动子可包括组成型、诱导型、时序调节、发育调节、化学调节、组织优选和组织特异性启动子;启动子的选择将随着表达时间和空间需要而变化,而且也取决于靶物种;例如组织或器官的特异性表达启动子,根据需要受体在发育的什么时期而定;尽管证明了来源于双子叶植物的许多启动子在单子叶植物中是可起作用的,反之亦然,但是理想地,选择双子叶植物启动子用于双子叶植物中的表达,单子叶植物的启动子用于单子叶植物中的表达;
2)与适合的转录终止子连接,也可以提高本发明基因的表达效率;例如来源于CaMV的tml,来源于rbcS的E9;任何已知在植物中起作用的可得到的终止子都可以与本发明基因进行连接;
3)引入增强子序列,如内含子序列(例如来源于Adhl和bronzel)和病毒前导序列(例如来源于TMV,MCMV和AMV)。
上述方法中,所述对逆境胁迫敏感植物可为转基因植物,也可为通过杂交等常规育种技术获得的植物。
上述方法中,所述转基因植物理解为不仅包含第一代到第二代转基因植物,也包括其子代。对于转基因植物,可以在该物种中繁殖该基因,也可用常规育种技术将该基因转移进入相同物种的其它品种,特别包括商业品种中。所述转基因植物包括种子、愈伤组织、完整植株和细胞。
上文所述植物或/和目的植物可为下述任一种:
C1)双子叶植物;
D1)单子叶植物;
D3)禾本目植物;
D4)禾本科植物;
D5)稻属植物;
D6)水稻。
上文所述抗病可为抗稻瘟病。
检测水稻基因组中SNP1的多态性或基因型的物质的下述任一种应用也属于本发明的保护范围:
F1、在鉴定或辅助鉴定水稻抗病性中的应用;
F2、在制备鉴定或辅助鉴定水稻抗病性产品中的应用;
F3、在检测或辅助检测水稻抗病性中的应用;
F4、在制备检测或辅助检测水稻抗病性产品中的应用;
F6、在水稻抗病育种中的应用;
所述SNP1是水稻基因组的一个SNP位点,对应于序列表中序列2的第656位核苷酸,其可为T或C。SNP 1位点有三种基因型,即CC、TT或CT。基因型TT是SNP1位点为T的纯合型,基因型CC是位点为C的纯合型,基因型CT是SNP1位点为C和T的杂合型。
所述SNP分子标记位点的基因型为基因型TT的水稻的抗病性高于或候选高于所述SNP分子标记位点的基因型为基因型CC的水稻。
所述水稻抗病可为水稻抗稻瘟病。
本申请从1500余份水稻资源中筛选、获得乙烯突变体,其中来自越南的Tang10 编号为MC41的材料对稻瘟病有明显抗性。应用图位克隆获得目的基因,为NBS-LRR 家族成员REIN1,该基因的突变基因OsREIN1T219I可激发超敏反应,增加了水稻的抗病性。本发明的实验证明,OsREIN1T219I蛋白具有提高水稻抗病能力的功能。导入 OsREIN1T219I基因的烟草可引起超敏反应,而同样条件下导入OsREIN1基因和GFP基因的烟草并不能引起超敏反应。OsREIN1T219I蛋白及其编码基因对培育抗病水稻品种具有重要意义。
附图说明
图1为MC41的乙烯不敏感表型由隐性突变所致。Air表示未进行乙烯处理,Ethylene表示进行乙烯处理。
图2为水稻REIN1T219I的图位克隆。
图3为OsREIN1T219I可激发超敏反应。
图4为纯合OsREIN1T219I增加水稻对稻瘟病的抗性。
图5为pCAMBIA2300载体经改造后得到的pCAMBIA2300-35S-OCS载体示意图。
具体实施方式
下面结合具体实施方式对本发明进行进一步的详细描述,给出的实施例仅为了阐明本发明,而不是为了限制本发明的范围。以下提供的实施例可作为本技术领域普通技术人员进行进一步改进的指南,并不以任何方式构成对本发明的限制。
下述实施例中的实验方法,如无特殊说明,均为常规方法,按照本领域内的文献所描述的技术或条件或者按照产品说明书进行。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。
水稻品种日本晴(O.Sativa L.spp.japonica,var nipponbare,AA genome,Nip)属粳亚种,记载于如下文献:傅秀林等,水稻品种日本晴,农业科技通讯,1973,2,11-15;由中国科学院遗传与发育生物学研究所朱立煌提供。水稻品种Tang 10(Oryza sativaL.Indica,Tang 10),编号为MC41(来自越南)和明恢63(MH63)(Oryza sativa L.MH63)由中国科学院遗传与发育生物学研究所储成才研究员提供。
植物双元表达载体pCAMBIA2300:购自酶研生物,货号MY1383。Julie Leclercq etal.,Development of a new pCAMBIA binary vector using technology,Plasmid.2015Sep;81:50-4。
同源重组试剂盒:美国Clone Smarter品牌的Seamless Assembly Cloning Kit,货号为C5891-50,国内由中美泰和生物技术有限公司代理。
稻瘟菌Guy11:由中国科学院微生物研究所刘俊提供。S Fukiya et al.,Establishment of a new cross of the rice blast fungus derived from Japanesedifferential strain Ina168 and hermaphroditic rice pathogen Guy11,BiosciBiotechnol Biochem.2001Jul;65(7):1464-73.
实施例1、水稻乙烯反应缺陷材料的筛选和遗传分析
在筛选1500份水稻资源中乙烯反应缺陷材料的过程中,发现Tang10,编号为MC41的材料,其黄化苗在乙烯处理时与对照日本晴(Oryza sativa L.cv.Nipponbare,Nip) 比较,其根长没有变短的乙烯反应,并且将MC41与Nip杂交,获得的F1代,其乙烯反应与Nip相似,即,在乙烯处理下,其根长明显变短(图1)。因此推测MC41乙烯反应异常的性状可能由隐性单基因决定。
对MC41的乙烯反应异常变异的显隐性做进一步分析。以MC41为母本与MH63作为父本杂交,MH63与Nip有同样正常的乙烯反应。F1代黄化苗为MH63相同表型,表明 MC41的乙烯异常反应为隐性突变。F2代黄化苗表型分析发现为3:1(MH63表型:MC41 表型)分离,且符合χ2检验(表1)。以上结果表明,MC41的变异是由单基因控制的隐性突变。
表1 MC41乙烯反应异常表型的显隐性分析
注:“+”为MC41乙烯不敏感表型;“-”为MH63正常乙烯敏感表型;critical value(0.05, 1)=3.84.
实施例2、MC41变异等位基因的鉴定
通过图位克隆将MC41变异位点确定在7号染色体长臂上的一个73k的区间里,并在其中的一个基因上鉴定到了一个位于水稻基因LOC_Os07g40810的特异SNP(命名为SNP1)可导致氨基酸的非同义替换,将该基因列为候选基因。该基因编码蛋白为 NBS-LRR家族成员REIN1,该基因命名为OsREIN1基因。按照水稻基因组参考基因序列,设计引物:
REIN1F:5’-ATGGAACATGCTGTTGTTAGTGC-3’
REIN1R:5’-TCAGTCTTCGTCGAAAGATGAAGATAGT-3’
以MC41总mRNA为模板,由上述引物扩增获得约3Kb DNA条带。经测序后确认其为属于NBS-LRR家族的REIN1基因,但与水稻基因组参考序列中LOC_Os07g40810的序列有差异,MC41中OsREIN1发生了单个碱基的变化,即位于NB-ARC结构域 P-loop(ATP/ADP binding)中第656位碱基由C突变为T(C656T),使其编码蛋白 OsREIN1中219位单个氨基酸发生改变,由苏氨酸突变为异亮氨酸(T219I)(图2)。因此,将水稻MC41中该突变后的蛋白命名为OsREIN1T219I,OsREIN1T219I蛋白的氨基酸序列为序列表中的序列1的所示。编码OsREIN1T219I蛋白的基因(OsREIN1T219I)的编码链的CDS序列为序列表中的序列2所示的DNA分子。在MC41的OsREIN1T219I基因中SNP1 特异突变位点(序列表中序列2的第656位)的核苷酸为T,对应OsREIN1T219I蛋白的该位点的氨基酸(序列表中序列1的第219位)为异亮氨酸(I)。
在水稻MH63与Nip中,该SNP1位点的核苷酸为C(对应于序列表中序列2的第 656位),对应的氨基酸为苏氨酸(T)(对应于序列表中序列1的第219位)。因此,在水稻基因组中,该SNP1位点的核苷酸为C或T,对应的氨基酸为苏氨酸(T)或异亮氨酸(I);该位点有三种基因型,即CC、TT或CT,基因型TT是SNP1位点为T的纯合型,基因型CC是位点为C的纯合型,基因型CT是SNP1位点为C和T的杂合型。
接着,在5518份水稻材料中检测了OsREIN1的等位基因,发现只有MC41中OsREIN1基因有该SNP1位点的点突变,因此这一突变为稀有突变。
实施例3、OsREIN1T219I参与抗病性调控
HR(超敏反应)细胞死亡对病原菌获取养分起阻隔作用,限制病原菌生长和繁衍,同时可激发邻近组织的特异性防御反应和植株的***获得抗性(systematic acquiredresistance,SAR)。因此超敏反应对植物的抗病性起重要作用。
在烟草***中检测OsREIN1T219I是否能启动植物的超敏反应。具体实验步骤如下:
1.超敏反应载体构建:
1)使用下列引物,分别以Nip和MC41的cDNA为模板,通过PCR扩增出目的片段:
F:5’-TGGAGAGGACAGGGTACCCGGATGGAACATGCTGTTGTTAGTGC-3’
(划线部分序列为同源重组臂序列,粗体部分序列为3xMYC标签序列)
2)将植物表达载体pCAMBIA2300进行改造:在NcoI和KpnI酶切位点之间*** 35S启动子序列(序列表中序列3),在PstI和HindⅢ之间***了OCS终止子序列 (序列表中序列4)得到改造后的载体pCAMBIA2300-35S-OCS(图5);然后用BamHI 酶切pCAMBIA2300-35S-OCS载体,回收得到线性化载体。
3)将步骤1)中的以Nip为模板的PCR回收产物和以MC41为模板的PCR回收产物,分别与步骤2)中得到的线性化载体通过同源重组试剂盒进行混合连接,分别构建得到重组载体pCAMBIA2300-REIN1和pCAMBIA2300-REIN1T219I。其中, pCAMBIA2300-REIN1是来源于Nip的OsREIN1基因与线性化载体连接的重组载体,表达OsREIN1蛋白(OsREIN1蛋白是将序列表中的序列1的第219位I替换为T,保持序列1的其它氨基酸残基不变得到的蛋白质);pCAMBIA2300-REIN1T219I是来源于MC41的 OsREIN1T219I基因与线性化载体连接的重组载体,表达氨基酸序列为序列表中的序列1 的OsREIN1T219I蛋白。
4)将步骤3)中构建得到的两种重组载体pCAMBIA2300-REIN1和 pCAMBIA2300-REIN1T219I分别单独转化至E.coli.DH5α感受态细胞,铺至含50μl/ml 卡那霉素的固体LB培养基中,过夜培养。
5)挑取固体培养基上生长出的单克隆进行菌落PCR鉴定,阳性重组菌菌落提取质粒并测序验证得到pCAMBIA2300-REIN1重组质粒和pCAMBIA2300-REIN1219I重组质粒后进行下一步实验。
参照上述载体构建方法构建GFP基因的重组载体pCAMBIA2300-GFP,用作对照进行后续实验。pCAMBIA2300-GFP与pCAMBIA2300-REIN1T219I的区别仅在于将 pCAMBIA2300-REIN1T219I中的OsREIN1T219I基因替换为GFP基因,其它核苷酸相同。
2.HR反应检测:
1)本氏烟草(Nicotiana benthamiana)在14h光照/10h黑暗,温度25℃,相对湿度70%条件下培养约4周。
2)将上述1中得到的pCAMBIA2300-REIN1、pCAMBIA2300-REIN1T219I及GFP对照重组载体pCAMBIA2300-GFP分别转入农杆菌EHA105中,分别得到重组农杆菌 EHA105/pCAMBIA2300-REIN1、EHA105/pCAMBIA2300-REIN1219I和 EHA105/pCAMBIA2300-GFP。
3)分别挑取重组农杆菌EHA105/pCAMBIA2300-REIN1、 EHA105/pCAMBIA2300-REIN1219I和EHA105/pCAMBIA2300-GFP这三种重组农杆菌的单克隆培养、扩增。收集菌体,用侵染液(含10mM MgCl2,10mM MES,150μM乙酰丁香酮,其余为水,pH=5.6)悬浮重组农杆菌菌体得到重组农杆菌 EHA105/pCAMBIA2300-REIN1侵染液、重组农杆菌EHA105/pCAMBIA2300-REIN1219I侵染液和重组农杆菌EHA105/pCAMBIA2300-GFP侵染液这三种重组农杆菌侵染液,每种重组农杆菌侵染液的OD600nm=1.0(以侵染液为空白对照)。室温静止2~3h(至少0.5 h,至多不超过3h。)
4)将各种重组农杆菌侵染液单独或按不同的比例混合,并用1mL注射器在烟草背面注射,用记号笔标记烟草叶片水渍状区域。实验重复三次,每次重复设置以下9 种处理,每种处理注射10个烟草叶片。
重组农杆菌REIN1-3Xmyc单独侵染烟草处理:用1mL注射器在烟草背面注射3) 的重组农杆菌EHA105/pCAMBIA2300-REIN1侵染液,用记号笔标记烟草叶片水渍状区域。
重组农杆菌REIN1219I-3Xmyc单独侵染烟草处理:用1mL注射器在烟草背面注射3)的重组农杆菌EHA105/pCAMBIA2300-REIN1219I侵染液,用记号笔标记烟草叶片水渍状区域。
重组农杆菌GFP-3Xmyc单独侵染烟草处理:用1mL注射器在烟草背面注射3)的重组农杆菌EHA105/pCAMBIA2300-GFP侵染液,用记号笔标记烟草叶片水渍状区域。
重组农杆菌REIN1219I-3Xmyc和重组农杆菌REIN1-3Xmyc的1:1处理 (REIN1219I-3Xmyc/REIN1-3Xmyc 1:1):将3)的重组农杆菌 EHA105/pCAMBIA2300-REIN1219I侵染液和重组农杆菌EHA105/pCAMBIA2300-REIN1侵染液按照1:1的体积比混合得到重组农杆菌混合侵染液。用1mL注射器在烟草背面注射该重组农杆菌混合侵染液,用记号笔标记烟草叶片水渍状区域。
重组农杆菌REIN1219I-3Xmyc和重组农杆菌REIN1-3Xmyc的1:0.5处理 (REIN1219I-3Xmyc/REIN1-3Xmyc 1:0.5):将3)的重组农杆菌 EHA105/pCAMBIA2300-REIN1219I侵染液和重组农杆菌EHA105/pCAMBIA2300-REIN1侵染液按照1:0.5的体积比混合得到重组农杆菌混合侵染液。用1mL注射器在烟草背面注射该重组农杆菌混合侵染液,用记号笔标记烟草叶片水渍状区域。
重组农杆菌REIN1219I-3Xmyc和重组农杆菌REIN1-3Xmyc的1:0.1处理 (REIN1219I-3Xmyc/REIN1-3Xmyc 1:0.1):将3)的重组农杆菌 EHA105/pCAMBIA2300-REIN1219I侵染液和重组农杆菌EHA105/pCAMBIA2300-REIN1侵染液按照1:0.1的体积比混合得到重组农杆菌混合侵染液。用1mL注射器在烟草背面注射该重组农杆菌混合侵染液,用记号笔标记烟草叶片水渍状区域。
重组农杆菌REIN1219I-3Xmyc和重组农杆菌GFP-3Xmyc的1:1处理(REIN1219I-3Xmyc/GFP-3Xmyc 1:1):将3)的重组农杆菌EHA105/pCAMBIA2300-REIN1219I侵染液和重组农杆菌EHA105/pCAMBIA2300-GFP侵染液按照1:1的体积比混合得到重组农杆菌混合侵染液。用1mL注射器在烟草背面注射该重组农杆菌混合侵染液,用记号笔标记烟草叶片水渍状区域。
重组农杆菌REIN1219I-3Xmyc和重组农杆菌GFP-3Xmyc的1:0.5处理(REIN1219I-3Xmyc/GFP-3Xmyc 1:0.5):将3)的重组农杆菌EHA105/pCAMBIA2300-REIN1219I侵染液和重组农杆菌EHA105/pCAMBIA2300-GFP侵染液按照1:0.5的体积比混合得到重组农杆菌混合侵染液。用1mL注射器在烟草背面注射该重组农杆菌混合侵染液,用记号笔标记烟草叶片水渍状区域。
重组农杆菌REIN1219I-3Xmyc和重组农杆菌GFP-3Xmyc的1:0.1处理(REIN1219I-3Xmyc/GFP-3Xmyc 1:0.1):将3)的重组农杆菌EHA105/pCAMBIA2300-REIN1219I侵染液和重组农杆菌EHA105/pCAMBIA2300-GFP侵染液按照1:0.1的体积比混合得到重组农杆菌混合侵染液。用1mL注射器在烟草背面注射该重组农杆菌混合侵染液,用记号笔标记烟草叶片水渍状区域。
5)注射后40h左右即可观察到HR反应。
图3显示了HR检测结果。在烟草中瞬时表达实验表明,注射分别含对照GFP-3XMyc和REIN1-3xMyc的重组农杆菌时,叶片无明显变化,表明均不能启动HR,而注射含REIN1T219I-3XMyc农杆菌,叶片变色,则可激发强烈的HR。含REIN1T219I-3XMyc和 GFP-3Xmyc农杆菌无论以1:1、1:0.5和1:0.1混合注射,均能激发HR(图3中左图所示)。图3中右图显示,当将含REIN1T219I-3XMyc农杆菌与含REIN1-3xMyc农杆菌混合注射1:0.1时,HR不受影响,1:0.5时HR下降,而1:1时则不发生HR,上述结果说明REIN1T219I-3Xmyc重组农杆菌可引起植物HR,而REIN1-3Xmyc重组农杆菌不能引起植物HR。REIN1-3xMyc的增加抑制了HR反应,OsREIN1可抑制OsREIN1T219I1引起的HR。进一步说明sREIN1T219I为隐性变异,只有隐性纯合时才引发HR反应,赋予植株抗病性。 OsREIN1T219I能引起植物HR,可用于调控植物的抗病性,纯合的OsREIN1T219I可提高水稻的抗病性。
3.水稻MC41和Nip的稻瘟病抗性检测
3.1稻瘟菌准备:
将供试的稻瘟菌Guy11接种至酵母淀粉斜面培养基上(水溶性淀粉10g,酵母浸膏2g,琼脂20g加水至1000ml),待菌丝长满后,用1.5ml无菌水,轻轻刮取菌丝,将菌丝移至燕麦片培养基上(燕麦片30g,CaCl2 0.6g,琼脂15g加水至1000ml),涂布均匀后置于28℃,约10天后培养皿中长满菌丝,25-28℃光照培养约2天,待其充分产孢,用无菌蒸馏水洗下稻瘟病菌孢子,经三层擦镜纸过滤后,在100倍显微镜下计算孢子浓度,调节孢子浓度至5×105个/ml(含0.02%Tween-20)得到孢子悬浮液备用。
3.2受试植物的抗病性检测:
将生长四周的日本晴和MC41水稻幼苗置于接种箱内,用喷雾器对其进行喷雾接种(每个接种箱接种30mL3.1中得到的孢子悬浮液),于28℃黑暗保湿培养24小时后,将接种箱移至正常光照(16h光照/8h黑暗)培养箱生长,5天后拍照并记录水稻的发病情况。
图4显示,Nip在接种6天后病斑明显,新叶和老叶均开始萎蔫,叶尖明显枯萎,受害面积约为40-70%,而MC41老叶上虽有针尖状斑点,叶尖略有枯黄,但仍呈绿色。结果表明MC41抗病性明显高于对照Nip。
上述实验表明,来自MC41(Tang 10)的OsREIN1突变基因OsREIN1T219I与调控水稻抗病性相关,其纯合水稻品种MC41(Tang 10)与含未突变OsREIN1基因的对照Nip相比增加了抗病性。因此OsREIN1T219I可应用于分子设计育种,培育高抗病水稻。
以上对本发明进行了详述。对于本领域技术人员来说,在不脱离本发明的宗旨和范围,以及无需进行不必要的实验情况下,可在等同参数、浓度和条件下,在较宽范围内实施本发明。虽然本发明给出了特殊的实施例,应该理解为,可以对本发明作进一步的改进。总之,按本发明的原理,本申请欲包括任何变更、用途或对本发明的改进,包括脱离了本申请中已公开范围,而用本领域已知的常规技术进行的改变。按以下附带的权利要求的范围,可以进行一些基本特征的应用。
序列表
<110> 中国科学院遗传与发育生物学研究所
<120> 水稻OsREIN1T219I蛋白及其编码基因与应用
<130> GNCSQ210207
<160> 4
<170> PatentIn version 3.5
<210> 1
<211> 981
<212> PRT
<213> 水稻(Oryza sativa)
<400> 1
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Thr Ala Phe Leu Gln Asp Leu Ser Gly Arg Asp Glu Cys Gly Lys Gln
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Val Lys Ile Trp Lys Lys His Val Arg Glu Ile Ala Tyr Asp Ile Glu
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Asp Cys Ile Asp Glu Phe Lys His Gln Leu Gly Asp Ser Ser Ser Ala
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Asn His Leu Ile Ser Gly Thr Ala Gly Asn Ser Met Ala Ala Tyr Asp
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Val His Trp Leu Leu Glu Ala His Val Gln Gln Leu Arg Val Val Ser
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Ile Phe Gly Phe Gly Gly Leu Gly Lys Thr Ile Leu Ala Met Thr Thr
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Tyr Gln Ser Leu Ser Gly Arg Asn Gly Pro Phe Gln Cys Gln Ala Phe
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<210> 2
<211> 2946
<212> DNA
<213> 水稻(Oryza sativa)
<400> 2
atggaacatg ctgttgttag tgccgcggaa ggtgcgatcc acactctctt gggaaagctc 60
ggcacaatcg tcctccaaga agcccagctt ctgggaggta ttcggggtga actgcaacac 120
ttgaaggatg aactggagag catgacagcc tttcttcagg acctttctgg gagagatgaa 180
tgcggcaagc aagttaagat ttggaagaag catgtgcgtg agattgcata tgatatcgag 240
gattgtattg atgagttcaa acatcaactt ggtgacagca gtagtgccgg tggcagcggc 300
cctgtagtgt ttttccgcaa ggccacccac atattgcaga ccaccagagt gaggcatcag 360
attgccaaac aaatccaaga actaaaaagg cgtactatga atatcagtgc ccgaaattca 420
aggtatagtg ccaatcatct catctctgga actgctggga atagcatggc tgcatatgat 480
agccaagcta atcttttaaa tgttgatact cgcattactg cactctttcc agagagaagg 540
cagcttgttg gcattgaacc acgtcaggga aatcttgtgc actggttatt ggaggcacat 600
gtgcaacaat tacgggtggt gtctatattt ggttttggtg gtttgggcaa gacaatactt 660
gctatgacaa catatcaaag tctatctgga agaaatggac cttttcaatg tcaagctttt 720
gtaactgtgt cccagagttt tgatgtcaag gttctgatga gagatattct tctccaaatc 780
actcaaccag ttaatcaacc gagttctcca tcaactggag ctggcaaagg tcccatggaa 840
ggcctactca agggcatgga agcatggaat gtggtacaac ttgcgagcat cctcaggcag 900
caattggaaa ataagagata tctgattgtt cttgatgata tctggagcat gactgcatgg 960
gaaggtattc ggttttcttt gccggactca aataatggta gcagaatagt ggttactaca 1020
cgaatcagag ctgtagcaca cacctgttgt ttccatgagt acgaccgagc ttacgaaatc 1080
aaacctctca ctgattgtga atccagagac ttattcttca aaagaatatt tggcagctca 1140
atttgtcctg agcacttaga agatatttca gctaagattc tgggaaaatg tggtggcaca 1200
cctttatcca tagtcagtat agcaggcctc ttggctagca agcctgtgca cagtaaagat 1260
ctttgggaga agatttatag ctctcttggt tcagagattg aaaccaatcc atcacttgat 1320
agattgaaga aaatacttga gcttagctat aatgatcttc cttaccactt gaagacttgc 1380
ttcttatatt taagcattta ccctgaggac cataatatca gaaggaaaac tatactcagg 1440
cgatgggtag cagaacgctt tgtgactggc aaacgtggac tgagtgtttt tgaggtggca 1500
gaaagctact ttgatgaatt cattaaccgg agcattattc agccagtgac cactagcttt 1560
acagggaagg ttaaaacatt ccgagttcat gatgtgatgt tggagatcat tgtgtcaaag 1620
tcaattgaag ataattttat cactctagta ggagaacaga acactttgtt tccgcaagag 1680
aagataaggc ggctaactgt tcatagcaga ggtgtgaaat atattgccac aagagaaata 1740
ttatgccatg tccggtcctt gagcatattt gccgatggag aaacattgca gtttggttgg 1800
atgaaattaa tgagaatatt ggacttagaa ggctacgaat tcctgagaaa tagagatctc 1860
aaagacttgt gcaggctgtt tcaattggaa taccttaatc tgagaagaac acatattacg 1920
gaactccctg cacaaatagg aaacctgaag aagttggaga ctctggatat aagggacaca 1980
gccataaagc atttgcctcc tggcataacc aatcttccac atttagcaaa cttacttgga 2040
ggaagaagat cctataacca cactgggcgg tggcctattt ctgagttctg gggtttacat 2100
atccctaacg aacttcgaaa aatggattca cttacaacac ttgcacaggt agaaatcacg 2160
acctctacat cacattatat cagcgagttg tcgaagctat cccggttgag gaaacttgga 2220
gtactgatgt ttgttgatga cgacagcacc tgggcatcct tgatctctgc tcttgagaag 2280
ctcagcggca gccttcgctc gttattactt tggcgacctg atggcgcgat gaacttcaac 2340
attgtcaatt cattatccag tccaccaata tttacgaaaa gtatgaacct tcgaggtcag 2400
ttaacacaac taccatgttg gtttcctttg ctttccaaca tcacagaatt aacccttcgc 2460
gcaaccgaac taagcgccga ggaagacctg aaggtactag gaagtttacc tagcctgcta 2520
tacctcaggt tgcatcacaa tgcatacata ggaacagaat tttctgcttc agcaggggaa 2580
ttcccatctc ttagactgct tgtcattcat ctcgacatgt ctgaagattg ggaggcaaga 2640
tttgaagaag gggcactacc aaagctagca aggctagagc tgtctctgtt cgaagaagcc 2700
tccatccaag agatcactgg catagagttc cttccgagtc tgaaggaagt ctcgattcgt 2760
gcttgtcaca gcaatatcgt gaatgtggaa gagatagcta cctctctgag ggctgatgcc 2820
gagaagaaca tcaacaaacc catcgtcact ttcgaggaga aacagtgggt gccgatgagg 2880
tcaaggacgg atcctccttt agatcacatg ggaaacctac tatcttcatc tttcgacgaa 2940
gactga 2946
<210> 3
<211> 531
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 3
ccatggagtc aaagattcaa atagaggacc taacagaact cgccgtaaag actggcgaac 60
agttcataca gagtctctta cgactcaatg acaagaagaa aatcttcgtc aacatggtgg 120
agcacgacac gcttgtctac tccaaaaata tcaaagatac agtctcagaa gaccaaaggg 180
caattgagac ttttcaacaa agggtaatat ccggaaacct cctcggattc cattgcccag 240
ctatctgtca ctttattgtg aagatagtgg aaaaggaagg tggctcctac aaatgccatc 300
attgcgataa aggaaaggcc atcgttgaag atgcctctgc cgacagtggt cccaaagatg 360
gacccccacc cacgaggagc atcgtggaaa aagaagacgt tccaaccacg tcttcaaagc 420
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<210> 4
<211> 233
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 4
gcatgccctg ctttaatgag atatgcgaga cgcctatgat cgcatgatat ttgctttcaa 60
ttctgttgtg cacgttgtaa aaaacctgag catgtgtagc tcagatcctt accgccggtt 120
tcggttcatt ctaatgaata tatcacccgt tactatcgta tttttatgaa taatattctc 180
cgttcaattt actgattgta ccctactact tatatgtaca atattaaaat gaa 233
Claims (10)
1.一种蛋白质,其特征在于:所述蛋白质为OsREIN1T219I;所述OsREIN1T219I为氨基酸序列是序列表中序列1的蛋白质。
2.与权利要求1中所述蛋白质相关的生物材料,所述生物材料为下述B1)至B4)中的任一种:
B1)编码权利要求1中所述蛋白质的核酸分子;
B2)含有B1)所述核酸分子的表达盒;
B3)含有B1)所述核酸分子的重组载体、或含有B2)所述表达盒的重组载体;
B4)含有B1)所述核酸分子的重组微生物、或含有B2)所述表达盒的重组微生物、或含有B3)所述重组载体的重组微生物。
3.根据权利要求2所述的生物材料,其特征在于:B1)所述核酸分子的编码序列是序列表中序列2。
4.权利要求1中所述蛋白质和/或权利要求2或3中所述生物材料的下述任一种应用:
Q1、权利要求1中所述蛋白质和/或权利要求2或3中所述生物材料在调控植物抗病性中的应用;
Q2、权利要求1中所述蛋白质和/或权利要求2或3中所述生物材料在制备提高植物抗病性的产品中的应用;
Q3、权利要求1中所述蛋白质和/或权利要求2或3中所述生物材料在培育抗病性植物中的应用;
Q4、权利要求1中所述蛋白质和/或权利要求2或3中所述生物材料在制备植物抗病性产品中的应用;
Q5、权利要求1中所述蛋白质和/或权利要求2或3中所述生物材料在植物抗病育种中的应用。
5.一种培育抗病植物的方法,包括提高目的植物中权利要求1中所述蛋白质的活性或/和权利要求1中所述蛋白质的编码基因的表达量,得到抗病植物;所述抗病植物的抗病性高于所述目的植物的抗病性。
6.根据权利要求5所述的方法,其特征在于:所述提高目的植物中权利要求1中所述蛋白质的活性或/和权利要求1中所述蛋白质的编码基因的表达量是通过将权利要求1中所述蛋白质的编码基因导入所述目的植物实现的。
7.根据权利要求4所述的应用和/或权利要求5或6所述的方法,其特征在于:所述植物或/和目的植物为单子叶植物。
8.根据权利要求4所述的应用和/或权利要求5或6所述的方法,其特征在于:所述植物或/和目的植物为水稻;所述抗病为抗稻瘟病。
9.检测水稻基因组中SNP1的多态性或基因型的物质的下述任一种应用:
F1、在鉴定或辅助鉴定水稻抗病性中的应用;
F2、在制备鉴定或辅助鉴定水稻抗病性产品中的应用;
F3、在检测或辅助检测水稻抗病性中的应用;
F4、在制备检测或辅助检测水稻抗病性产品中的应用;
F5、在水稻抗病育种中的应用;
所述SNP1是水稻基因组的一个SNP位点,对应于序列表中序列2的第656位核苷酸,其为T或C,所述SNP分子标记位点的基因型为基因型TT的水稻的抗病性高于或候选高于所述SNP分子标记位点的基因型为基因型CC的水稻。
10.根据权利要求9所述的应用,其特征在于:所述水稻抗病为水稻抗稻瘟病。
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Citations (7)
Publication number | Priority date | Publication date | Assignee | Title |
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