CN116410279B - 与调控水稻非生物胁迫耐受性相关的蛋白及其相关生物材料与应用 - Google Patents
与调控水稻非生物胁迫耐受性相关的蛋白及其相关生物材料与应用 Download PDFInfo
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Abstract
本发明涉及一种与调控水稻非生物胁迫耐受性相关的蛋白及其相关生物材料与应用,所述蛋白质,是如下A1)、A2)或A3)的蛋白质:A1)氨基酸序列是序列表中序列2的蛋白质;A2)将序列表中序列2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/或添加得到的与A1)所示的蛋白质具有90%以上的同一性且与植物非生物胁迫耐受性相关的蛋白质;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。本发明的OsCCA1在响应高盐、干旱以及渗透胁迫的过程中发挥正调控作用;OsCCA1可作为一个具有多重抗性的基因,在水稻抗盐、抗旱等方面发挥重要作用。
Description
技术领域
本发明涉及生物技术领域,具体的说,涉及一种与调控水稻非生物胁迫耐受性相关的蛋白OsCCA1及其相关生物材料与培育耐高盐且抗旱的植物方法。
背景技术
水稻生产在保障我国粮食安全问题上具有举足轻重的地位,也是我国人民主要粮食作物之一。然而,由于环境气候的变化,导致全球因干旱造成的农作物损失约数百亿美元,土壤的盐碱化问题也日益加重。因此,了解水稻在生长发育的过程中如何响应高盐、干旱以及渗透胁迫的环境,对于培育具有多重抗性的作物品种具有重要的意义。
干旱影响植物的生长发育、产量、细胞膜完整性、叶绿素含量以及光合作用效率等,是制约作物生产力直接因素之一(Praba et al.,2009)。伴随着产生的特点是含水量、叶水势和压势降低,叶表面气孔关闭、细胞增大等。另外,通过影响生理代谢过程如光合作用、呼吸作用、糖和营养代谢、离子吸收以及植物激素等导致植物生长减慢。干旱属于一种多维的压力胁迫,它会引发植物体从生理、生化到分子水平的各层级反应。其中不可避免的结果之一是,由线粒体和叶绿体产生的的活性氧(reactive oxygen species,ROS)量增加。在压力胁迫下产生的ROS可作为触发防御或警报的信号,H2O2作为二级信使向下游传递信号,藕连着脱落酸(abscisic acid,ABA)信号通路和Ca2+通道的打开或关闭。此外,一些干旱胁迫响应基因、转录因子、水孔道蛋白、以及脱水蛋白等在分子水平上发挥着不同功能,进而将干旱信号传递到下游,启动植物的防御系统(Kaur and Asthir,2017)。
植物感知水分缺失的信号并启动相应应对策略的能力称之为抗旱性,它是十分复杂的性状。当植物接收到干旱胁迫信号时,会启动不同的机制进行响应:(1)逃旱性:在干旱胁迫阻碍其生产力之前,加速植物的生长发育过程,帮助其完成生殖生长,便于繁衍后代;(2)避旱性:通过多重生理代谢活动,减少水分流失或增强水分吸收能力;(3)耐旱性:增强体内含水量低的耐力,维持基本代谢过程(Basu et al.,2016)。植物体内渗透势的调节是植物适应干旱的重要过程,它有助于维持组织的代谢活动。其调节过程主要依赖于一些化合物的合成,如氨基酸(天冬氨酸、脯氨酸、谷氨酸)、甘氨酸甜菜碱、蔗糖和果聚糖以及环醇类的合成等。其中,脯氨酸是最重要的一种,它参与稳定酶和蛋白质在内的大分子、保持膜的完整性以及活性氧的清除(Kaur and Asthir,2015)。除此之外,干旱环境也会触发一些激素信号,如乙烯(ethylene)、油菜素内酯(brassinosteroids,BR)以及ABA信号通路等(Janni et al.,2019)。其中ABA途径是调节植物的抗旱性,优化水分利用效率的重要方式,干旱环境会促进不同器官中ABA的产生和积累并激活下游信号通路(Chen et al.,2021)。
土壤盐渍化是一个世界性的问题,全球约1/3的灌溉土地受到盐碱化的影响,严重威胁着作物的生长和产量的提高,阻碍了现代农业的可持续发展。当土壤中盐浓度超过200mM时,主要粮食作物如水稻、小麦、玉米等均无法正常生长(Flowers et al.,2015)。因此提高作物的盐胁迫耐受性具有重要意义,在保障全球粮食安全上同样至关重要。了解高盐分环境如何影响植物形态发育、生理生化代谢以及基因表达的特性,是实现该目标的基础。模式植物拟南芥的基础性研究揭示了许多盐胁迫响应基因,其中一些基因可运用到作物中提高其耐盐性。SOS(salt overly sensitive)信号通路在植物响应盐胁迫的过程发挥重要作用,其功能在拟南芥和水稻中是保守的,主要将Na+排到细胞外(El Mahi et al.,2019;Shi et al.,2000)。近期研究表明,糖基肌醇磷酸神经酰胺鞘脂(glycosyl inositolphosphorylceramide,GIPC)在植物响应盐胁迫的过程中起到传感器的作用(Jiang etal.,2019)。高盐环境主要通过以下两个方面制约植物的生长:(1)渗透胁迫:降低外部水势,导致植物的吸水能力减弱;Na+与细胞壁成分结合导致细胞壁的发生改变;渗透压力还会导致气孔关闭,从而降低植物同化吸收CO2能力。(2)Na+毒害:在代谢活跃的细胞内隔室中Na+或Cl-过量积累导致。由于Na+、K+相似的化学属性,竞争性结合K+结合位点,影响酶活性(Benito et al.,2014);细胞内电荷和离子条件决定着局部水分与细胞骨架以及蛋白质的关系,进而决定着细胞的命运(Cheeseman,2013)。研究表明Cl-对植物的毒害可能不是毒性本身的结果,而是由于Cl-过量积累导致关键的大量元素N和S的缺乏,因为NO3 -、SO4 2-的吸收方式与Cl-相同,均是通过非选择性的阴离子通道吸收(Bazihizina et al.,2019)。
干旱和高盐胁迫等非生物逆境往往伴随着渗透胁迫产生,直接导致作物生产力降低,也是危害全球粮食安全的重要因素。固着生长的植物因难以躲避其所受到的渗透胁迫,进化出了感知和适应逆境的机制,包括信号接收与传递、ABA相关调控和后期应答等过程。渗透胁迫应答通过多重调控机制在不同时间段发挥作用,如瞬时产生的Ca2+信号(持续数秒)(Knight et al.,1997);SnRK2等蛋白激酶激活(数分钟)(Fujii and Zhu,2012);Ca2+依赖的应答以及ABA积累(Yuan et al.,2014)、气孔关闭和下游基因的表达等(持续几天)(Shkolnik et al.,2018),进而调控植物生长、叶片衰老、以及休眠过程等。
生物钟系统通过感知并整合外界环境信号,不仅调控植物对生长环境的适应性,还影响着作物的主要农艺性状(Müller et al.,2018)。深入了解昼夜节律的调节与植物反应之间的联系,对于预测植物对未来气候的适应性和农业可持续发展具有重要价值。然而,目前对水稻生物钟系统的解析仍相对较为局限,生物钟组分是否以及如何同时感知外界多重的胁迫压力如高盐、干旱,并调整水稻适应性的分子机理仍然未知。解析并挖掘具有多重抗性的基因资源,对于培育优质高产、环境广适的新型作物具有非同寻常的意义。
目前,虽然对高盐、干旱和渗透胁迫单独处理如何影响植物的耐受性以及生长发育了解相对较多,但对于植物如何协调并适应多重胁迫环境却知之甚少。
发明内容
本发明所要解决的技术问题在于提出一种与调控水稻非生物胁迫耐受性相关的蛋白OsCCA1及其相关生物材料,进而提供一种培育耐高盐且抗旱的植物方法。
为了解决以上技术问题,本发明提供了一种来源于水稻的非生物胁迫耐受性相关蛋白,名称为OsCCA1,来源于水稻,是如下A1)、A2)或A3)的蛋白质:
A1)氨基酸序列是序列表中序列2的蛋白质;
A2)将序列表中序列2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/或添加得到的与A1)所示的蛋白质具有90%以上的同一性且与植物非生物胁迫耐受性相关的蛋白质;
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。
所述非生物胁迫耐受性包括:高盐胁迫耐受性、渗透胁迫耐受性和干旱胁迫耐受性。
上述蛋白质中,序列表中的序列2由719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,Perresidue 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90%以上的同一性可为至少91%、92%、95%、96%、98%、99%或100%的同一性。
上述蛋白质中,所述OsCCA1可来源于水稻。
与OsCCA1相关的生物材料也属于本发明的保护范围。
本发明所提供的与OsCCA1相关的生物材料,为下述B1)至B7)中的任一种:
B1)编码OsCCA1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B1)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、转基因植物组织或转基因植物器官;
B6)降低OsCCA1表达的核酸分子;
B7)含有B6)所述核酸分子的表达盒、重组载体、重组微生物、转基因植物细胞系、转基因植物组织或转基因植物器官。
其中,所述核酸分子可以是DNA,如cDNA、基因组DNA或重组DNA;所述核酸分子也可以是RNA,如mRNA或hnRNA等。
上述生物材料中,B1)所述核酸分子具体可为如下1)或2)所示的基因:
1)编码序列(CDS)是序列表中序列1的第1-2160位核苷酸的DNA分子;
2)核苷酸序列是序列表中序列1的DNA分子。
上述生物材料中,B6)所述核酸分子具体可为与序列表中序列1的第1-2160位核苷酸所示的DNA分子中任一片段反向互补的DNA分子。
上述生物材料中,B7)所述重组载体可以为针对OsCCA1编码序列(CDS)设计的CRISPR/Cas9重组表达载体。
其中,序列表中的序列1由2160个核苷酸组成,其编码序列是序列表中序列1中的第1-2160位,编码序列表中的序列2所示的蛋白质。
上述生物材料中,B2)所述的含有编码OsCCA1的核酸分子的表达盒(OsCCA1基因表达盒),是指能够在宿主细胞中表达OsCCA1的DNA,该DNA不但可包括启动OsCCA1基因转录的启动子,还可包括终止OsCCA1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。启动子的例子包括但不限于:花椰菜花叶病毒的组成型启动子35S;来自西红柿的创伤诱导型启动子,亮氨酸氨基肽酶("LAP",Chao等人(1999)Plant Physiology 120:979-992);来自烟草的化学诱导型启动子,发病机理相关1(PR1)(由水杨酸和BTH(苯并噻二唑-7-硫代羟酸S-甲酯)诱导);西红柿蛋白酶抑制剂II启动子(PIN2)或LAP启动子(均可用茉莉酮酸曱酯诱导);热休克启动子(美国专利5,187,267);四环素诱导型启动子(美国专利5,057,422);种子特异性启动子,如谷子种子特异性启动子pF128(CN101063139B(中国专利2007 1 0099169.7)),种子贮存蛋白质特异的启动子(例如,菜豆球蛋白、napin,oleosin和大豆beta conglycin的启动子(Beachy等人(1985)EMBO J.4:3047-3053))。它们可单独使用或与其它的植物启动子结合使用。此处引用的所有参考文献均全文引用。合适的转录终止子包括但不限于:农杆菌胭脂碱合成酶终止子(NOS终止子)、花椰菜花叶病毒CaMV 35S终止子、tml终止子、豌豆rbcS E9终止子和胭脂氨酸和章鱼氨酸合酶终止子(参见,例如:Odell等人(I985)Nature 313:810;Rosenberg等人(1987)Gene,56:125;Guerineau等人(1991)Mol.Gen.Genet,262:141;Proudfoot(1991)Cell,64:671;Sanfacon等人GenesDev.,5:141;Mogen等人(1990)Plant Cell,2:1261;Munroe等人(1990)Gene,91:151;Ballad等人(1989)Nucleic Acids Res.17:7891;Joshi等人(1987)Nucleic Acid Res.,15:9627)。
可用现有的植物表达载体构建含有所述OsCCA1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。所述翻译控制信号和起始密码子的来源是广泛的,可以是天然的,也可以是合成的。翻译起始区域可以来自转录起始区域或结构基因。为了便于对转基因植物细胞或植物进行鉴定及筛选,可对所用植物表达载体进行加工,如加入可在植物中表达的编码可产生颜色变化的酶或发光化合物的基因(GUS基因、萤光素酶基因等)、抗生素的标记基因(如赋予对卡那霉素和相关抗生素抗性的nptII基因,赋予对除草剂膦丝菌素抗性的bar基因,赋予对抗生素潮霉素抗性的hph基因,和赋予对methatrexate抗性的dhfr基因,赋予对草甘磷抗性的EPSPS基因)或是抗化学试剂标记基因等(如抗除莠剂基因)、提供代谢甘露糖能力的甘露糖-6-磷酸异构酶基因。从转基因植物的安全性考虑,可不加任何选择性标记基因,直接以逆境筛选转化植株。
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。
为了解决上述技术问题,本发明还提供了植物非生物胁迫耐受性增强剂。
本发明所提供的植物非生物胁迫耐受性增强剂含有所述蛋白质或/和所述蛋白质相关的生物材料。
上述植物非生物胁迫耐受性增强剂的活性成分可为所述蛋白质或所述蛋白质相关的生物材料,上述植物非生物胁迫耐受性增强剂的活性成分还可含有其他生物成分或/和非生物成分,上述药剂的其他活性成分本领域技术人员可根据植物的非生物胁迫耐受效果确定。
上述植物非生物胁迫耐受性增强剂,可以为植物高盐胁迫耐受剂(用于提高植物的耐盐性)、植物渗透胁迫耐受剂(用于提高植物的渗透胁迫耐受性)和植物干旱胁迫耐受剂(用于提高植物的耐旱性)。
上述蛋白质、或上述生物材料的下述P1-P4中的任一种应用也属于本发明的保护范围:
P1、所述蛋白质、或所述生物材料在正调控植物非生物胁迫耐受性中的应用;
P2、所述蛋白质、或所述生物材料在高非生物胁迫耐受性植物中的应用;
P3、所述蛋白质、或所述生物材料在制备提高高非生物胁迫耐受性的产品中的应用;
P4、所述蛋白质、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耐盐植物的方法。
本发明所提供的培育高非生物胁迫耐受性植物的方法,包括提高目的植物中所述蛋白质或其编码基因的表达量,得到高非生物胁迫耐受性植物;所述高非生物胁迫耐受性植物的非生物胁迫耐受性高于所述目的种子植物的非生物胁迫耐受性。
上述方法中,所述提高目的植物中所述蛋白质或其编码基因的表达量可通过将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,其中所述蛋白质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)修饰邻近起始甲硫氨酸的基因序列,以使翻译有效起始;例如,利用在植物中已知的有效的序列进行修饰;
2)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;
3)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;
4)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)。
所述蛋白质的编码基因可通过使用Ti质粒,植物病毒栽体,直接DNA转化,微注射,电穿孔等常规生物技术方法导入植物细胞(Weissbach,1998,Method for PlantMolecular Biology VIII,Academy Press,New York,pp.411-463;Geiserson and Corey,1998,Plant Molecular Biology(2nd Edition)。
上述方法中,所述耐盐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。
为了解决上述技术问题,本发明还提供了一种培育非生物胁迫耐受性降低的转基因植物的方法。
本发明所提供的培育非生物胁迫耐受性降低的转基因植物的方法,包括降低目的植物中所述蛋白质的编码基因的表达,得到非生物胁迫耐受性低于所述目的植物的转基因植物。
上述方法中,所述降低目的植物中所述蛋白质的编码基因的表达是通过利用CRISPR/Cas9基因编辑系统抑制目的植物中所述的蛋白质的含量和/或活性实现的。
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。
上文中,所述植物和所述目的植物均为单子叶植物或双子叶植物。所述植物具体可以为水稻。
上文中,所述非生物胁迫耐受性包括:高盐胁迫耐受性、渗透胁迫耐受性和干旱胁迫耐受性。
本发明利用CRISPR/Cas9技术获得了OsCCA1功能缺失的突变体,分别进行180mMNaCl、180mM mannitol模拟高盐、渗透胁迫处理,以及干旱处理,观察表型并统计恢复生长后的存活率,最终确定OsCCA1在响应高盐、干旱以及渗透胁迫的过程中发挥正调控作用。本发明的OsCCA1可作为一个具有多重抗性的基因,在水稻抗盐、抗旱等方面发挥重要作用。
附图说明
图1为利用CRISPR/Cas9技术获得的Dongjin(DJ)背景下oscca1突变体,分别命名为oscca1-L1和oscca1-L2,二者均属于功能缺失型突变体。
图2为oscca1突变体盐胁迫耐受性分析,纯合突变体oscca1-L1和oscca1-L2以及野生型DJ四周龄幼苗材料进行盐胁迫处理的表型图以及存活率统计。
图3为oscca1突变体渗透胁迫耐受性分析,纯合突变体oscca1-L1和oscca1-L2以及野生型DJ四周龄幼苗材料进行180mM甘露醇模拟渗透胁迫处理的表型图以及存活率统计。
图4为oscca1突变体干旱胁迫耐受性分析,纯合突变体oscca1-L1和oscca1-L2以及野生型DJ三周龄土壤中生长的幼苗材料进行干旱胁迫处理的表型图以及存活率统计。
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。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。以下实施例中的定量试验,均设置三次重复实验,结果取平均值。
下述实施例中所用的实验材料为常用的粳稻品种Dongjin,公众可从中国科学院植物研究所获得,以重复本实验,以下简称DJ。
下述实施例中的CRISPR/Cas9载体系统记载于如下文献中(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,ChenY,Guo J,Chen L,Zhao X,Dong Z,Liu Y-G(2015)A Robust CRISPR/Cas9 System forConvenient,High-Efficiency Multiplex Genome Editing in Monocot and DicotPlants.Mol Plant8:1274-1284)由华南农业大学刘耀光实验室惠赠,公众可以从华南农业大学获得,以重复本申请实验,不可作为其它用途使用。
下述实施例中的感受态农杆菌为EHA105,购自北京博迈德公司。
培养水稻的营养液配方:由A母液、B母液、EDTA-Fe母液和微量元素母液,每升营养液中按照5:5:1:1的比例混合配制而成,再加入100~300mg硅酸钠,然后用浓盐酸调pH至5.8。母液配制:A母液由9.64g (NH4)2SO4、3.7g KNO3、4.96g KH2PO4、3.18g K2SO4和29.965gMgSO4.7H2O混合溶于1L蒸馏水中配制而成。B母液由17.235g Ca(NO3)2.4H2O溶于1L蒸馏水中配制而成。EDTA-Fe母液由7.45g Na2EDTA置于蒸馏水中加热溶解后与5.57g FeSO4.7H2O溶解液,不断搅拌混合,冷却后定容至1L。微量元素母液由2.86g H3BO4、0.08g CuSO4.H2O、0.22g ZnSO4.7H2O、1.81g MnCl2.4H2O、0.09g NaMO4.H2O溶于1L蒸馏水。
水稻愈伤诱导及转化的培养基所需试剂均可通过商业化途径获得,NB BasalMedium粉末,品牌:Phytotech,可从北京西美杰科技有限公司购买;MS培养基粉末,品牌:Caisson,可从北京兰博利德商贸有限公司购买。诱导愈伤培养基NB2:4.1g NB BasalMedium粉末溶于1L蒸馏水中,然后分别加入含300mg水解酪蛋白、500mg脯氨酸、500mg谷氨酰胺、30g蔗糖和2mg/L 2,4-D母液(1mg/L),蒸馏水定容至1L,调节pH至5.8,然后加入琼脂8g/L,高压120℃灭菌15min。
继代水稻愈伤培养基NB1:与NB2培养基配方基本一致,唯一不同的是加入0.5mg/L2,4-D母液。
愈伤转化后第一次筛选培养基NB1S1:NB1培养基高压灭菌后,待温度降至60℃左右时,加入600mg/L头孢霉素和25mg/L Hyg。
愈伤转化后第二次筛选培养基NB1S2:NB1培养基高压灭菌后,待温度降至60℃左右时,加入300mg/L头孢霉素和50mg/L Hyg。
抗性愈伤分化第一阶段培养基RE1:4.5gMS培养基粉末溶于蒸馏水1L中,然后分别加入300mg水解酪蛋白、30g蔗糖、30g山梨醇以及各种激素1mg/L 6-BA、0.5mg/L KT、0.2mg/L ZT、0.25mg/L NAA,蒸馏水定容至1L,调节pH至5.8,然后加入琼脂8g/L,高压120℃灭菌15min。
抗性愈伤分化第二阶段培养基RE2:与RE1培养基基本相同,区别在于RE2培养基中需要0.5mg/L NAA,其他成分均与RE1一致。
1/2MS培养基:2.25g MS培养基粉末溶于1L蒸馏水中,加入30g蔗糖,调节pH至5.8,加入8g琼脂粉,高压120℃灭菌15min。
下述实施例中使用的离心管规格均为1.5mL。
实施例1、OsCCA1功能缺失的突变体的构建
1、OsCCA1编码基因
OsCCA1为本发明的发明人从水稻Dongjin中分离克隆出一个基因,其核苷酸序列如序列表的序列1所示,将其编码蛋白命名为OsCCA1蛋白,如序列表的序列2所示。
2.pCRISPR/Cas9-OsCCA1载体
利用序列3所示的参考基因组DNA序列在E-CRISPR网站(http://www.e-crisp.org/E-CRISP/designcrispr.html),结合评分筛选合适的靶点,选择位于OsCCA1保守的MYB结构域,即第二个外显子上的GCAGAGGGAGCGTTGGACTG为CRISPR/Cas9靶点序列。按照要合成引物序列F:ggcAGCAGAGGGAGCGTTGGACTG和R:aaacCAGTCCAACGCTCCCTCTGC,用于构建pCRISPR/Cas9-OsCCA1载体。将重组质粒pCRISPR/Cas9-OsCCA1导入农杆菌感受态细胞EHA105中,得到重组农杆菌用于转化水稻愈伤。
pCRISPR/Cas9载体的构建方法,参考如下文献中(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,GuoJ,Chen L,Zhao X,Dong Z,Liu Y-G(2015)A Robust CRISPR/Cas9 System forConvenient,High-Efficiency Multiplex Genome Editing in Monocot and DicotPlants.Mol Plant8:1274-1284)实验步骤进行。具体可以包括如下步骤:
1)靶点接头制备:引物F和R分别用ddH2O溶解成100μM母液,F、R各取1μL加至98μLddH2O中混匀,置于PCR仪95℃5min,然后转移至室温冷却完成退火。
2)将靶点与pYLgRNA质粒进行边切边连反应,酶切体系如下:0.5μL的BsaⅠ-HF,BsaⅠ缓冲液1μL,10mM ATP 0.5μL,10ng/μL的pYLgRNA质粒1μL,T4 DNA连接酶0.1μL,接头1μL,ddH2O补齐至10μL,所有组分混匀离心,置于PCR仪中37℃5min,20℃5min,进行5个循环。
3)扩增gRNA表达盒,以上述连接产物为模板,第一轮PCR扩增,分两个反应进行,第一轮扩增反应一和反应二的体系如下表:
依次加入上表组分,轻轻混匀后离心,置于PCR仪,进行如下程序:98℃2min,98℃15s,60℃15s,68℃20s,进行28个循环。分别将反应一和反应二的产物取4μL,用1.5%琼脂糖凝胶进行检测,反应一产物目的条带约为425bp,反应二产物目的条带约为150bp。
第二轮PCR扩增:将通用引物B1’和BL混合成10×工作液,B1’3μL+BL 3μL+14μLddH2O。第一轮PCR产物各取1μL用ddH2O稀释10倍作为模板,分别为稀释后的反应一产物和稀释后的反应二产物,再按如下体系进行扩增:稀释后的反应一、反应二产物各1μL作为模板,KOD酶1μL,2×KOD buffer 25μL,dNTPs 10μL,通用引物B1’+BL混合液5μL,ddH2O 7μL,所有组分混合均匀,置于PCR仪中95℃2min,95℃10s,58℃15s,68℃20s,反应20个循环,取2μLPCR产物,进行1%的琼脂糖凝胶检测,目的条带约为500bp。
4)将上述第二轮PCR产物进行纯化,目的为了去除残留的KOD酶,测定浓度并记录。
5)边切边连至pYLCRISPR/Cas9Pubi-H载体,反应体系依次加入:纯化后产物总量至70ng,CRISPR/Cas9质粒总量为80ng,限制性内切酶BsaⅠ-HF 0.5μL,缓冲液1.5μL,ddH2O补齐至15μL。所有组分混合均匀,置于37℃反应15min。结束后取出,再加入1μL 10×T4连接酶缓冲液,以及1μL T4 DNA连接酶,吹打混匀,置于PCR中反应程序为37℃2min,10℃3min,20℃5min,37℃2min,进行25个循环。
6)将上述产物热击转化DH5α,37℃培养箱培养过夜。摇菌并提取质粒,电击转化农杆菌EHA105,得到农杆菌pCRISPR/Cas9-OsCCA1/EHA105。
3.pCRISPR/Cas9-OsCCA1/EHA105工程农杆菌转化水稻愈伤及抗性愈伤筛选
将步骤2中得到的农杆菌pCRISPR/Cas9-OsCCA1/EHA105培养至生长对数期,然后离心4000rpm,10min收集菌体,然后用无菌侵染液重悬。无菌操作挑选出分化良好呈淡黄色的水稻愈伤,放于无菌滤纸上吹20min左右,去除多余水分。然后将愈伤放于含有农杆菌的侵染液中浸泡15~20min,期间摇晃3次。倒掉侵染液,将愈伤放置于无菌滤纸上,吸干多余侵染液。
然后将愈伤转移至铺好滤纸的NB2C加乙酰丁香酮的培养基中,黑暗条件25℃培养3天。无菌水中加入300mg/L头孢霉素,洗涤共培的愈伤4~5次,直至无菌水变清澈,无浑浊物的状态。清洗后的水稻愈伤转移至无菌滤纸上吸干多余水分,再移放至含有600mg/L头孢霉素和25mg/L Hyg的NB1(NB1S1)培养基上,黑暗条件下培养两周。
然后无菌操作挑选淡黄色的抗性愈伤转移至含有300mg/L头孢霉素和50mg/L Hyg的NB1(NB1S2)培养基上进行第二次筛选,黑暗条件下培养两周。
筛选能继续分化的抗性愈伤,无菌操作转移至RE1培养基上开始分化培养,黑暗培养一周,再进行光照培养一周。将能分化出绿芽的抗性愈伤,无菌操作转移至含RE2培养基的组培瓶中,再继续分化,进行持续光照培养,直至分化出幼苗。期间每隔两周需更换一次RE2培养基。无菌操作将分化出的小苗,移接至1/2MS培养基上继续培养。然后打开组培瓶盖,炼苗3~4天,将T0代转基因苗移种到温室中生长,直至种子成熟,得到OsCCA1功能缺失的突变体T0代种子。
实施例二、获得OsCCA1功能缺失的突变体
通过CRISPR/Cas9技术获得oscca1相关突变体,对T0代转基因材料的突变类型进行鉴定。先将T0代苗,收取叶片,提取DNA,具体实验流程如下:收取水稻叶片放入1.5mL离心管,并研磨成粉末;然后加入500μL DNA提取液,震荡混匀;再加入等体积的氯仿和苯酚(1:1)混合液,震荡混匀,13000rpm离心5min;缓慢吸取上清300μL,再加入等体积的氯仿,剧烈震荡混匀,13000rpm离心5min;收集上清150μL,加入两倍体积预冷的无水乙醇,置于-20℃沉淀10min;4℃,13000rpm,离心10min,收集DNA;倒掉上清,加入800μL的75%乙醇洗两遍,吹干后加入50μL ddH2O溶解。用OsCCA1的特异性引物,F的序列为TCTAGGAGGAGGTATGAGGTA、R的序列为GCACAAAAGCCAGCCGTTTTT,以提取的DNA为模板,进行PCR,扩增目的条带。然后将产物进行一代测序,与目的片段进行比对。通过分析我们鉴定到OsCCA1基因发生突变的两个独立转基因植株(分别为oscca1-L1和oscca1-L2),其突变类型如图1所示。oscca1-L1缺失了2bp,导致OsCCA1的mRNA发生移码突变,导使mRNA编码提前终止,进而丧失OsCCA1正常功能。oscca1-L2在第二个外显子上插入一个碱基A,同样导致移码突变,编码仅有27个氨基酸的短肽,如图1所示。
实施例三、oscca1突变体材料对高盐分胁迫耐受性分析
本发明下述实施例中使用的水稻材料培养方法为:挑选饱满野生型DJ和oscca1-L1、oscca1-L2纯合突变体的种子,浸泡于蒸馏水中37℃培养箱生长48小时,然后倒掉多余蒸馏水,冲洗种子,密封继续催芽24小时。挑选萌发状态一致的种子点放至无底的96孔板中,置于营养液中培养。生长条件为12小时光照/12小时黑暗,温度为30±2℃。下述实施例中存活率统计方法:存活率=恢复生长后存活的水稻幼苗数/总数×100%。
将生长至四周的材料,进行拍照,作为处理前的对照,然后将材料转移至含有180mMNaCl的营养液中,开始模拟高盐胁迫处理。盐处理24天后进行拍照,然后再转换至新鲜不含盐的营养液中恢复生长,恢复10天后,进行拍照如图2A所示。恢复生长10天后,统计DJ、oscca1-L1和oscca1-L2的存活率,实验重复四次取平均值,如图2B所示。图2中(A)DJ和oscca1突变体材料进行180mM NaCl模拟盐胁迫处理,处理前、处理24天以及恢复生长10天后的材料状态。比例尺=5厘米;(B)DJ和oscca1-L1、oscca1-L2突变体盐处理后并恢复生长10天之后的存活率统计。显示的值为平均值±标准误,四次生物学重复。***,P<0.001,差异极显著,统计分析方法为单因素方差分析。结果表明,DJ的存活率高达70%左右,但是oscca1-L1和oscca1-L2的存活率仅有20%左右,统计说明OsCCA1的功能缺失会导致植株对盐胁迫高度敏感。由此可见,OsCCA1在植物响应盐胁迫的过程中起正调控的作用,对于增强水稻的耐盐性是必不可少的。
实施例四、oscca1突变体材料对渗透胁迫耐受性分析
按照上述水稻材料培养方法,培养一批DJ、oscca1-L1和oscca1-L2材料,在生长四周时,进行拍照,作为处理前对照。然后将材料转移至含180mM甘露醇的营养液中,模拟渗透胁迫环境处理24天,然后拍照。在将处理后的材料放于不含甘露醇的正常营养液中,开始进行恢复生长24天,然后拍照记录,如图3A所示。最后对恢复生长后的材料,统计它们的存活率,实验进行了三次生物学重复,统计结果取三次的平均值,如图3B所示。图3中(A)DJ和oscca1突变体材料进行180mM甘露醇模拟渗透胁迫处理,处理前、处理24天、恢复生长24天后的材料状态,比例尺=5厘米;(B)DJ和oscca1-L1、oscca1-L2突变体渗透胁迫处理后并恢复生长24天之后的存活率统计。显示的值为平均值±标准误,三次生物学重复。**,P<0.01,差异极显著,统计分析方法为单因素方差分析。结果表明,在渗透胁迫的环境中,野生型DJ的存活率达50%左右,而oscca1-L1和oscca-L2突变体的存活率仅有15%左右。渗透胁迫中的处理结果说明,OsCCA1在调控水稻对渗透胁迫环境中的适应性中同样发挥着关键作用,对于提高水稻对渗透胁迫的耐受性所发挥的作用是不容忽视的。
实施例五、oscca1突变体材料干旱胁迫耐受性分析
将萌发状态一致的DJ和oscca1-L1、oscca1-L2的种子种植于土壤中,正常浇水灌溉生长至三周时,拍照记录其生长状态。然后将材料开始模拟干旱处理,即停止浇水10天,然后再恢复正常灌溉,生长5天后,拍照记录所有材料的状态,如图4A所示。然后统计DJ和oscca1-L1、oscca1-L2材料的存活率。干旱胁迫处理进行了三次生物学重复,三次存活率统计的平均值如图4B所示。图4中,(A)DJ和oscca1突变体材料进行干旱胁迫处理,处理前、处理10天并恢复生长5天后的材料状态,比例尺=5厘米;(B)DJ和oscca1-L1、oscca1-L2突变体干旱胁迫处理并恢复生长5天之后的存活率统计。显示的值为平均值±标准误,三次生物学重复。*,P<0.05,差异显著,统计分析方法为单因素方差分析。干旱环境中DJ的存活率达60%左右,而oscca1-L1和oscca1-L2仅有30%左右。由此看见,oscca1突变体材料对干旱环境也更加敏感。因此,OsCCA1在调控水稻耐旱性中所发挥的功能同样至关重要。
通过上述的实施例中,研究结果表明,OsCCA1在调控水稻对高盐、干旱以及渗透胁迫抗性中发挥正调节作用,对提高水稻对多重胁迫耐受性具有重要价值。
序列表
<110> 中国科学院植物研究所
<120> 与调控水稻非生物胁迫耐受性相关的蛋白及其相关生物材料与应用
<160> 3
<170> SIPOSequenceListing 1.0
<210> 1
<211> 2160
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 1
atggagatta attcctctgg tgaggaagcg gtggtaaagg tgaggaagcc atacacaatc 60
acaaagcaga gggagcgttg gactgaggca gagcacaaca ggttccttga agccttgaaa 120
ctgtatggga gagcctggca gcgcatagaa gagcatgttg ggacaaagac agctgtgcag 180
atcagaagtc atgctcaaaa gttcttcacc aagttggaaa aggaagctat caacaatggc 240
acttctccag gacaagctca tgacatcgac atacctccac cacgaccaaa aagaaaacct 300
aacagtccat atcctcgaaa aagttgtctc agctctgaga catccaccag ggaagttcaa 360
aatgataagg caacaatatc aaatatgacg aacaatagca ctgcacaaat ggcaggtgat 420
gcagctcttg agaaacttca aagaaaggag atatctgaaa aaggaagttg ctccgaagtt 480
cttaatctct ttcgagaagt cccatcggca tcattttctt cagttaacaa aagctcttca 540
aatcatggtg catccagggg gctggaaccg actaaaacag aagtcaaaga tgtggtcatc 600
ttggaaaggg attctatttc caatggtgca gggaaggatg caaaagatat caatgatcaa 660
gaaatggaaa ggctcaatgg gatacacatc agctcgaagc ctgatcattc tcatgaaaac 720
tgtttggata cctcaagcca acaatttaag ccaaaatcaa actctgtgga gacaacatat 780
gtggattggt ctgctgcaaa agcttcacac taccaaatgg acagaaatgg ggttactggc 840
tttcaagcca ctggaactga aggaagccat cctgatcaaa caagtgatca aatgggagga 900
gccagcggaa ctatgaatca atgcatccat ccaacacttc ctgtggatcc aaaattcgac 960
ggcaatgccg cagcacagcc ctttcctcac aactatgcag cctttgcacc aatgatgcaa 1020
tgccactgca accaagatgc ctacagatct tttgccaata tgtcatccac cttctccagc 1080
atgcttgtct ccacattgtt gtcaaaccct gcaatccatg cagctgccag gcttgcagca 1140
tcgtactggc ctacagtaga cggcaatact cctgatccaa atcaagaaaa tctttctgag 1200
agtgctcaag gaagccacgc tggctctcct cccaacatgg catctattgt cacagctaca 1260
gttgctgcag catcagcatg gtgggcaaca caaggtcttc tccctctttt tcctccacct 1320
atagcttttc catttgttcc agctcctagt gctccctttt ccacagcaga tgttcagcga 1380
gctcaagaga aagatataga ctgcccaatg gataatgcac agaaggaatt gcaagaaact 1440
cggaaacaag ataattttga agctatgaag gtcatagtgt cttcagagac tgatgagagt 1500
ggaaaaggag aagtgtcgct ccacactgag ttaaagatat ctccagcaga taaggccgac 1560
accaaacctg ccgcaggagc tgaaacaagt gacgtttttg gaaataagaa aaagcaggat 1620
cgctcttcat gtggttccaa cacaccgtca agtagtgata tagaagcaga taatgctcct 1680
gagaatcaag aaaaggctaa cgacaaggca aagcaagcat cttgcagtaa ctcttcagcc 1740
ggtgacaata accaccgtag atttaggagc agtgcaagca caagtgattc atggaaggaa 1800
gtttctgaag agggtcgtct ggcttttgat gcactgttca gtagagaaag gcttccccaa 1860
agcttttctc ctccgcaagt agaaggatca aaggagatta gcaaggagga agaagatgaa 1920
gtaaccacgg tgacggttga cctcaacaag aatgccgcta ttattgatca agaactcgac 1980
acagcggatg agccaagagc ttcctttcct aatgaattgt caaacctgaa gctgaaatct 2040
cgcaggaccg gtttcaaacc atacaagagg tgctcagtgg aagcgaagga gaacagggta 2100
ccggctagcg atgaggttgg taccaagagg attcgtcttg agagcgaagc atcgacatga 2160
<210> 2
<211> 719
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 2
Met Glu Ile Asn Ser Ser Gly Glu Glu Ala Val Val Lys Val Arg Lys
1 5 10 15
Pro Tyr Thr Ile Thr Lys Gln Arg Glu Arg Trp Thr Glu Ala Glu His
20 25 30
Asn Arg Phe Leu Glu Ala Leu Lys Leu Tyr Gly Arg Ala Trp Gln Arg
35 40 45
Ile Glu Glu His Val Gly Thr Lys Thr Ala Val Gln Ile Arg Ser His
50 55 60
Ala Gln Lys Phe Phe Thr Lys Leu Glu Lys Glu Ala Ile Asn Asn Gly
65 70 75 80
Thr Ser Pro Gly Gln Ala His Asp Ile Asp Ile Pro Pro Pro Arg Pro
85 90 95
Lys Arg Lys Pro Asn Ser Pro Tyr Pro Arg Lys Ser Cys Leu Ser Ser
100 105 110
Glu Thr Ser Thr Arg Glu Val Gln Asn Asp Lys Ala Thr Ile Ser Asn
115 120 125
Met Thr Asn Asn Ser Thr Ala Gln Met Ala Gly Asp Ala Ala Leu Glu
130 135 140
Lys Leu Gln Arg Lys Glu Ile Ser Glu Lys Gly Ser Cys Ser Glu Val
145 150 155 160
Leu Asn Leu Phe Arg Glu Val Pro Ser Ala Ser Phe Ser Ser Val Asn
165 170 175
Lys Ser Ser Ser Asn His Gly Ala Ser Arg Gly Leu Glu Pro Thr Lys
180 185 190
Thr Glu Val Lys Asp Val Val Ile Leu Glu Arg Asp Ser Ile Ser Asn
195 200 205
Gly Ala Gly Lys Asp Ala Lys Asp Ile Asn Asp Gln Glu Met Glu Arg
210 215 220
Leu Asn Gly Ile His Ile Ser Ser Lys Pro Asp His Ser His Glu Asn
225 230 235 240
Cys Leu Asp Thr Ser Ser Gln Gln Phe Lys Pro Lys Ser Asn Ser Val
245 250 255
Glu Thr Thr Tyr Val Asp Trp Ser Ala Ala Lys Ala Ser His Tyr Gln
260 265 270
Met Asp Arg Asn Gly Val Thr Gly Phe Gln Ala Thr Gly Thr Glu Gly
275 280 285
Ser His Pro Asp Gln Thr Ser Asp Gln Met Gly Gly Ala Ser Gly Thr
290 295 300
Met Asn Gln Cys Ile His Pro Thr Leu Pro Val Asp Pro Lys Phe Asp
305 310 315 320
Gly Asn Ala Ala Ala Gln Pro Phe Pro His Asn Tyr Ala Ala Phe Ala
325 330 335
Pro Met Met Gln Cys His Cys Asn Gln Asp Ala Tyr Arg Ser Phe Ala
340 345 350
Asn Met Ser Ser Thr Phe Ser Ser Met Leu Val Ser Thr Leu Leu Ser
355 360 365
Asn Pro Ala Ile His Ala Ala Ala Arg Leu Ala Ala Ser Tyr Trp Pro
370 375 380
Thr Val Asp Gly Asn Thr Pro Asp Pro Asn Gln Glu Asn Leu Ser Glu
385 390 395 400
Ser Ala Gln Gly Ser His Ala Gly Ser Pro Pro Asn Met Ala Ser Ile
405 410 415
Val Thr Ala Thr Val Ala Ala Ala Ser Ala Trp Trp Ala Thr Gln Gly
420 425 430
Leu Leu Pro Leu Phe Pro Pro Pro Ile Ala Phe Pro Phe Val Pro Ala
435 440 445
Pro Ser Ala Pro Phe Ser Thr Ala Asp Val Gln Arg Ala Gln Glu Lys
450 455 460
Asp Ile Asp Cys Pro Met Asp Asn Ala Gln Lys Glu Leu Gln Glu Thr
465 470 475 480
Arg Lys Gln Asp Asn Phe Glu Ala Met Lys Val Ile Val Ser Ser Glu
485 490 495
Thr Asp Glu Ser Gly Lys Gly Glu Val Ser Leu His Thr Glu Leu Lys
500 505 510
Ile Ser Pro Ala Asp Lys Ala Asp Thr Lys Pro Ala Ala Gly Ala Glu
515 520 525
Thr Ser Asp Val Phe Gly Asn Lys Lys Lys Gln Asp Arg Ser Ser Cys
530 535 540
Gly Ser Asn Thr Pro Ser Ser Ser Asp Ile Glu Ala Asp Asn Ala Pro
545 550 555 560
Glu Asn Gln Glu Lys Ala Asn Asp Lys Ala Lys Gln Ala Ser Cys Ser
565 570 575
Asn Ser Ser Ala Gly Asp Asn Asn His Arg Arg Phe Arg Ser Ser Ala
580 585 590
Ser Thr Ser Asp Ser Trp Lys Glu Val Ser Glu Glu Gly Arg Leu Ala
595 600 605
Phe Asp Ala Leu Phe Ser Arg Glu Arg Leu Pro Gln Ser Phe Ser Pro
610 615 620
Pro Gln Val Glu Gly Ser Lys Glu Ile Ser Lys Glu Glu Glu Asp Glu
625 630 635 640
Val Thr Thr Val Thr Val Asp Leu Asn Lys Asn Ala Ala Ile Ile Asp
645 650 655
Gln Glu Leu Asp Thr Ala Asp Glu Pro Arg Ala Ser Phe Pro Asn Glu
660 665 670
Leu Ser Asn Leu Lys Leu Lys Ser Arg Arg Thr Gly Phe Lys Pro Tyr
675 680 685
Lys Arg Cys Ser Val Glu Ala Lys Glu Asn Arg Val Pro Ala Ser Asp
690 695 700
Glu Val Gly Thr Lys Arg Ile Arg Leu Glu Ser Glu Ala Ser Thr
705 710 715
<210> 3
<211> 7705
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 3
ttgtgtattt tgttgttctg gcaggatcaa ggaagtgttt ggccttcgta ggtgagcctg 60
attactgttt gcattgctgc ttcatatatt tttcatactt ctaattggtt ttatgtttct 120
aattaattct tcttttttta atggttgcat aatttgagaa ctgcgagttt taatctgatt 180
ctagagttga agataacgaa aggagcaagg gtgaactctg ctgacttggt ttttgtgtgc 240
acagaccaac tgcagctgag gatttagtag gtgagtggag tggtgctgta ctgcgctgtc 300
cctggatttt tgtctcgagt aattgttgta gttgtggaac cctttagttt tatcaaatta 360
agtgatgcct tcttttgggg aaataaatgt ttttttcttg gtggagttca gtttgaccat 420
ttggagatgc ttatgaattt aaaatgatgt ggccccttat tgcatttttt ttttcagatt 480
gactgcaatt ggatcaattc aagtaggtga gtggaattga tcctcttgaa gattttgacc 540
caagtccaat tcacaagtag aattgtgaat tctttaggtt ctaccttggg tttgtaacac 600
taatttggct gctcttttgt ggttggccgc acaaatttga aatactggga gtgttggcgt 660
gatatagagg tctaggagga ggtatgaggt actacttgcg tcaggtccag cgtttggttg 720
gagatatgga aggaggagtg cttttgtttg tggaagacct caactctaac tgacaaagcg 780
accaattgcc aacacttctg ttatatttct tctttttctt gaattgggaa tggagattaa 840
ttcctctggt gaggaagcgg tggtaaaggt gagttgttct ttgatgtgca tgttgattcg 900
tgttaaaagg gagcagcatt ttattagggg ttatttgtag gtgaggaagc catacacaat 960
cacaaagcag agggagcgtt ggactgaggc agagcacaac aggttccttg aagccttgaa 1020
actgtatggg agagcctggc agcgcataga aggtgaattc tccacatata tatgtcatat 1080
cacaaatagt tctatgctat tactttctga atttactttt tgtactcact gtgattatta 1140
gctggttaat aaatacggct ttaattgagt tattgatatg gatcttgcat cttttttttt 1200
tcgttttctt gtttgccgtc tgatttattc tatagagcat gttgggacaa agacagctgt 1260
gcagatcaga agtcatgctc aaaagttctt caccaaggtt cttcctctat cttattggcc 1320
agtgctcctt tttttggttt agcgctatct ccttagcttt ctggtttcct tgtctttacc 1380
tttgggataa aaacggctgg cttttgtgct tactttctta ttattttttc ttactttctg 1440
cagtattatt tctttctttc cttcttttat tttacatatg agtacagcaa ttatgtacat 1500
agagtctgaa ctgtgtgtga tcagttcctt tcgattatta tgagaaatac acttatggct 1560
tttgtgctct gttagttgga tgaaacgtta actaaacagt gctgagatgg aaaagaagca 1620
ctaggaagtg tttgtctcgt agtaaaaaaa aaataatgtt gcaaaattga tgctattaaa 1680
cattctgttt gcatgttcct ttatttacat atccatccta tcactgagat tttgagcatg 1740
cacagatgca cttatgacaa aaaaaaaagc attttggtac tgttatattt acatactaag 1800
tcaagtcaaa ttgtatatct ggactatcat caagattagt aataaagatt tagttaaaca 1860
tgttccagtg cctacgtggt gttcataact aataaggatt cttaaaaaaa aaaagaacta 1920
gggcagccat ttgagacaat ttattaaaat aagaaaaatg taaaatgatt tgcaagttga 1980
ccaaggtatc accttccatt agaaatagcc actcaatcgc gcatttgcgc gactttgact 2040
gacagcatat aatatgtagg agtagttata acagtcacga gtggattaag tttcttttat 2100
ttatacatta ggcttatttt ttaatgaaat tctaatttac gcggtagaat tgtatcttta 2160
tttttgaaaa gggttgtcta ttattaatga tttttttaat caaggtgtaa tattggaatt 2220
cacagttatt tgttcggcaa acattgtaat aattgtctgt agattcacta aagtaaaggt 2280
cggaataatt tttgccatgc tggctttttc tttagaaaaa aaaagaagaa aactgttgtc 2340
cggtttcctc caattttttt tctacgtggg ctgcctgcag gttctggagg tttctctttg 2400
gaactctcct tgtttgatta gggctaatcc aacggttgtt gttgcttatt tctttgcatt 2460
ggacggtcct acatttcttt cttatgtggc tcaaccagat tgtgtgaaag tagttctttc 2520
actactttat attgtgtgaa aggtcggaaa acatcaaaaa taatactgta cacctattgt 2580
ttatcagtca aaacagaagg acctagtcca ttcccttctt ttaaaaaatt tactcccaat 2640
ctaagtcttg ttttttttcc tgttgcgcaa tctacccaaa ctgtgccttt taggggcttt 2700
acactgccac ctgatggagg tcacgttgcc atcactgtgc aacagccact ggactgttgc 2760
attgttgatt gcatcatcat cctctcaaag gtctcattgc aagatctatg aatggggagg 2820
aacacaaggt tagtaattta agaaacatga tggctttctg gtttctcaac tctatatcca 2880
gccaacatcc atgagcatct gaccgtccat tgttgtggtg ggttggattt ttggttgttg 2940
attaaagagg tgaataggtc tgccctaatc tccataaaat atgtaataca gctaataggt 3000
tttcgtcgct gcaagtactt gccagtaact gttctccatg tctaggctca taaaaaatat 3060
agagtgacaa gtaacaacgt ttaacctgac caatacttat actgcttaaa caccttctgt 3120
tttctaacaa gtatgcagta aatgtaatca gtgctacaat aacttcttga acattaacca 3180
tgcctagaac ttgggaccat acgctcataa cattggtttg agaagggtag tggactggga 3240
ggatatactc tagtcacgta tttcctagta atcccttggt tttgccaaaa aaaaggcaac 3300
caaggcaaat taccactgtt cattgaaccc tgatagtcaa tagttcgaaa agtggtaact 3360
aataactgtg aaatgtccat tgtggttgat gtcaattaaa ttagtcttta ctttgataag 3420
atgtccaaga gctttttgga gccccgaagg gttatgagtg atggccattc caggcttcag 3480
atgcacagtc gcttttcttt tgcaaagaca tgcaacaatg tttggttcta catcagcact 3540
ataccagtaa ttgcagtagt atagttttgg cttgtcgtta ctggtttctg ggctgaagtc 3600
caacatgact aggctccatt tgtggtctta tctagtgact aggctccatt tggttcttgt 3660
cataattatc cacagagatc tggttcattg atcacttttt ctactgtatc agcgaaaagt 3720
ttgatgatat taccgcctat acaaatgtta tagcatgctg ctaatcatac ccaccaatgc 3780
atactctccc ctgtttggtc aatgatgatt gatgccatgg ttgatgttca aactaaacct 3840
cggcaggtca ttctgagatc agaaggataa tataggcata aaactaaatc tatgacaacc 3900
tgtaatgtgg aaatggcctt tttcttttat gcaggtgaaa aaaaataaat gcgtactaaa 3960
ttaataataa cccaagggga gagacatccc aaaggcattg catataaggt tgagcaaagg 4020
ccccaaagcc agctcataac cgtatgggta ctttagggga acatatagcg aggggtcttt 4080
ttttttgtga gaactaggaa gcctcatgac tggaggactt accagttacc accgtgctac 4140
aagcacgttc tctctactaa atttataata gcaaagtact tggcttcaat gtgacttttt 4200
aggtccagga aaaatgcttt caaatttctt atcaacttat catcacaata gctagacgat 4260
gattttgatc aataaaagag caggggaatg tctagttgac tgaaagtaga agaggaacaa 4320
ttttctttct gcaaatttgg cttcaaaata atctgttgaa atattcatca ttacatgttt 4380
tcttttggac aaaggaagtg taacttccag cctctgcacc aactaaggat gcaaacaacc 4440
atatttgtgg ataccaggat ttgaaccctg gtggctggag tcattacatg ttttctatat 4500
tccaacttcc atgtcaaacc atgttcctga agtataatgc tcattaatta tctaatatgg 4560
cagcaaggca gacataatcc ggtaagtaaa gggtagggct gaccgccctc tggtcaagca 4620
ggctctccct gcatagcagg aggctctgct cggtgaaata caaaatcctt ctctgatatt 4680
attgcttatt tttccttaat acacatacta gtccaccttt atagcacaag tgctgagata 4740
tgtacttgga agtactaaat aaactaggaa gctctagcta gaaagtaccg gttcccgaac 4800
cgatggcttc atgaaacttt ttgccaaagg tttttagtaa ccttctacat gacactccct 4860
ctactccaca atatttgtcc aagactacca tctgcataga ccaaggaggt gcaagatata 4920
agaaatagtt aatgataatg gactatatat accatctgca taacttttat acctctttta 4980
agcattatat aaagtttagt agagagttgg gacatcgcaa taagtccaaa ctcctaattt 5040
tctcgacaaa cagacattgt caaaggggac agccccgaga aaatgaacaa atcttgtgga 5100
gcagacggag tagcattatg taaaatatca atatcatgta gtatggcact ttcatatatg 5160
tcaaaattac tgttagatat gcctgatttg ttttctgttt ctttatgtaa tttaactgta 5220
tctaactaat tgaaatatta cagttggaaa aggaagctat caacaatggc acttctccag 5280
gacaagctca tgacatcgac atacctccac cacgaccaaa aagaaaacct aacagtccat 5340
atcctcgaaa aagttgtctc agctctgaga catccaccag ggaagttcaa aatgataagg 5400
caacaatatc aaatatgacg aacaatagca ctgcacaaat ggcaggtgat gcagctcttg 5460
aggtacatac ctttgtagta tttcgtttta gttaggtttc tatttttgat ttatcctttt 5520
ctttaatgtg ctatcctgtt tctttaacaa gaaactcact tacattcaga aacttcaaag 5580
aaaggagata tctgaaaaag gaagttgctc cgaagttctt aatctctttc gagaagtccc 5640
atcggcatca ttttcttcag ttaacaaaag ctcttcaaat catggtgcat ccagggggct 5700
ggaaccgact aaaacagaag tcaaagatgt ggtcatcttg gaaagggatt ctatttccaa 5760
tggtgcaggg aaggatgcaa aagatatcaa tgatcaagaa atggaaaggc tcaatgggat 5820
acacatcagc tcgaagcctg atcattctca tgaaaactgt ttggatacct caagccaaca 5880
atttaagcca aaatcaaact ctgtggagac aacatatgtg gattggtctg ctgcaaaagc 5940
ttcacactac caaatggaca gaaatggggt tactggcttt caagccactg gaactgaagg 6000
aagccatcct gatcaaacaa gtgatcaaat gggaggagcc agcggaacta tgaatcaatg 6060
catccatcca acacttcctg tggatccaaa attcgacggc aatgccgcag cacagccctt 6120
tcctcacaac tatgcagcct ttgcaccaat gatgcaatgc cactgcaacc aagatgccta 6180
cagatctttt gccaatatgt catccacctt ctccagcatg cttgtctcca cattgttgtc 6240
aaaccctgca atccatgcag ctgccaggct tgcagcatcg tactggccta cagtagacgg 6300
caatactcct gatccaaatc aagaaaatct ttctgagagt gctcaaggaa gccacgctgg 6360
ctctcctccc aacatggcat ctattgtcac agctacagtt gctgcagcat cagcatggtg 6420
ggcaacacaa ggtcttctcc ctctttttcc tccacctata gcttttccat ttgttccagc 6480
tcctagtgct cccttttcca cagcagatgt tcagcgagct caagagaaag atatagactg 6540
cccaatggat aatgcacaga aggaattgca agaaactcgg aaacaagata attttgaagc 6600
tatgaaggtc atagtgtctt cagagactga tgagagtgga aaaggagaag tgtcgctcca 6660
cactgagtta aagatatctc cagcagataa ggccgacacc aaacctgccg caggagctga 6720
aacaagtgac gtttttggaa ataagaaaaa gcaggatcgc tcttcatgtg gttccaacac 6780
accgtcaagt agtgatatag aagcagataa tgctcctgag aatcaagaaa aggctaacga 6840
caaggcaaag caagcatctt gcagtaactc ttcagccggt gacaataacc accgtagatt 6900
taggagcagt gcaagcacaa gtgattcatg gaaggaagtt tctgaagagg tggtcgtcta 6960
ccagcattcc gcatattcac atttatctta ctcgctgaac atccatcact gcttttctaa 7020
ttcacatttc tgctgtcagg gtcgtctggc ttttgatgca ctgttcagta gagaaaggct 7080
tccccaaagc ttttctcctc cgcaagtaga aggatcaaag gagattagca aggaggaaga 7140
agatgaagta accacggtga cggttgacct caacaagaat gccgctatta ttgatcaaga 7200
actcgacaca gcggatgagc caagagcttc ctttcctaat gaattgtcaa acctgaagct 7260
gaaatctcgc aggaccggtt tcaaaccata caagaggtgc tcagtggaag cgaaggagaa 7320
cagggtaccg gctagcgatg aggttggtac caagaggatt cgtcttgaga gcgaagcatc 7380
gacatgattt gctttccacc tggttgctgg cctctaccaa gtcagaagtt aaatttacat 7440
ccgagctacc ataggacttc agaccttcca atgcattacc tcacaaactc aatttattgt 7500
gtgttgtatc ttaatgcttg ccaagcagct cctatagact gcttttaaac cgtatctcat 7560
agacttttga ttagcattta agcgactgct aaactttctt cataaaaggg tattatcctt 7620
attatgcatt atagtctggg gaaggtaata agtggaattt tggttcattt ttctgggtca 7680
tattttacca actgcatttt tatcc 7705
Claims (4)
1.蛋白质、或生物材料的下述P1-P2中的任一种应用:
P1、所述蛋白质、或生物材料在调控植物非生物胁迫耐受性中的应用,所示调控为降低植物非生物胁迫耐受性;
P2、所述蛋白质、或生物材料在培育非生物胁迫耐受性降低植物中的应用;
其中,所述蛋白质如下A1)或A2)所示:
A1)氨基酸序列是序列表中序列2的蛋白质;
A2)在A1)的N末端或/和C末端连接蛋白标签得到的融合蛋白质,所述蛋白标签为Flag标签、His标签、MBP标签、HA标签、myc标签、GST标签和/或SUMO标签;
所述生物材料如下述B1)至B4)中的任一种所示:
B1)编码A1)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;
所述非生物胁迫耐受性为高盐胁迫耐受性、渗透胁迫耐受性或干旱胁迫耐受性;
所述植物为水稻。
2.根据权利要求1所述的应用,其特征在于:B1)所述核酸分子为序列表中序列1的cDNA分子或DNA分子。
3.一种培育非生物胁迫耐受性降低的转基因植物的方法,包括降低目的植物中权利要求1中所述蛋白质的编码基因的表达,得到非生物胁迫耐受性低于所述目的植物的转基因植物;所述非生物胁迫耐受性为高盐胁迫耐受性、渗透胁迫耐受性或干旱胁迫耐受性,所述植物为水稻。
4.根据权利要求3所述的方法,其特征在于:所述降低目的植物中权利要求1中所述蛋白质的编码基因的表达是通过利用CRISPR/Cas9基因编辑系统抑制目的植物中权利要求1中所述的蛋白质的含量和/或活性实现的。
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