CN111662928B - 调控植物耐盐性的方法及耐盐相关蛋白 - Google Patents
调控植物耐盐性的方法及耐盐相关蛋白 Download PDFInfo
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Abstract
本发明公开了一种调控植物耐盐性的方法及耐盐相关蛋白，本发明提供了一种培调控植物耐盐性的方法，包括调控目的植物中耐盐相关蛋白质的编码基因的表达，其中，所述耐盐相关蛋白质是来源于水稻的耐盐性相关蛋白，名称为OsPRR73，来源于水稻日本晴。本发明利用CRISPR/Cas9技术获得了OsPRR73功能缺失的突变体，进行180mM NaCl模拟盐胁迫处理，观察表型并统计恢复处理后的存活率，最终确定OsPRR73在响应水稻盐胁迫的过程中发挥正调控作用。本发明的OsPRR73可作为一个耐盐基因，在水稻抗盐性方面发挥重要作用。
Description
技术领域
本发明涉及生物技术领域中耐盐相关蛋白OsPRR73及其相关生物材料与培育耐盐植物的方法。
背景技术
水稻(Oryza sativa L.)是我国最主要的粮食作物之一，随着现代农业发展，水稻产量已经大大提高。但是环境恶化带来的负面效应，已经严重阻碍了现代农业的发展。其中，土壤盐渍化是植物面临的主要胁迫之一，已成为威胁全球农业发展的严重问题。据估计，全球陆地约6％的面积受到盐渍化的影响，其中包含近4500万公顷的灌溉用地(Munnsand Tester,2008)。
禾本科作物对非生物胁迫极为敏感，尤其是水稻和小麦，在干旱和盐渍环境下产量下降最为明显(Lobell et al.,2014；Landi et al.,2017)。盐胁迫是一种常见的非生物胁迫，高浓度的盐主要会导致植物体内渗透胁迫和离子毒害。植物在响应盐胁迫初期，主要是渗透胁迫抑制植物幼叶的生长，而后期是离子失衡会加速叶片的衰老(Urano et al.,2014)。盐胁迫产生的影响主要有以下两个方面：一方面会影响植物体的Na+/K+离子稳态平衡，进而导致离子毒害的产生；另一方面，盐胁迫会诱导植物体内活性氧(ROS)的积累，进而产生氧化胁迫(Mittler,2017)。在植物响应盐胁迫的过程中，少量的ROS会增强植物体的适应性；而过量的ROS累积会直接引起细胞程序性死亡和组织坏死。Na+是造成离子毒害的最关键的因素，主要是因为Na+在叶片中的积累会加速叶片的坏死，导致作物产量的降低(Munns,1993)。在植物体内K+参与许多重要的代谢过程，而过量Na+能够竞争性结合K+位点，如50多种生物酶均需要K+来激活，而这是Na+所无法代替的，所以高浓度的Na+会破坏细胞中的多种酶活过程，进而导致植物体内的代谢紊乱(Bhadal and Malik,1988)。
盐胁迫还会对植物光合作用产生影响。盐胁迫下，植株叶片的类胡卜素与叶绿素含量普遍降低。随着盐胁迫时间的延长，叶片开始枯萎凋零(Hernandez et al.,1999；Ramanjulu et al.,1998)。研究发现在盐胁迫条件下番茄叶片中的总叶绿素、叶绿素α和β-胡萝卜素的含量因盐胁迫时间延长而逐渐降低(Kurth et al.,1986)。不同基因型的水稻研究中同样发现随着盐胁迫时间的延长，光合色素普遍降低(Yu et al.,2012)。盐胁迫对光合作用的影响又分为短期与长期效应，短期效应发生在盐胁迫开始后几小时或1天内，这种效应对植物影响很大，直接会导致碳同化的停止；长期影响发生在盐胁迫环境下的几天之后，由于发育的叶片中盐的积累导致碳同化减少，进而导致植物生长缓慢或死亡(Ramanjulu et al.,1998)。
盐胁迫会影响着植物生长发育的各个阶段，盐胁迫导致的离子不平衡直接影响种子萌发和幼苗的生长；在营养生长阶段其敏感性降低；但在开花阶段盐胁迫将直接影响植物种子灌浆。目前对于这方面的研究较多集中于模式植物拟南芥中，比如研究较为透彻的SOS(Salt Over Sensitive)途径，其重要生理作用就是在拟南芥面临盐胁迫时进行的稳态调节。EF-hand结构的钙结合蛋白SOS3(Salt Overly Sensitive 3)和SCaBP8(SOS3-LikeCalcium Binding Protein 8)/CBL10(Calcineurin B-Like Protein 10)能够感受并解码盐胁迫刺激产生的钙信号(Zhu,2016)，进而激活丝氨酸/苏氨酸蛋白激酶SOS2(SaltOverly Sensitive 2)(Lin et al.,2009)。被激活后的SOS2可以与细胞质膜上SOS1相互作用并磷酸化SOS1，然后激活Na+/H+交换通道，将胞质内积累的Na+排至胞外(Shi et al.,2000)。除此之外，大量研究表明植物响应盐胁迫等外界压力是通过多种复杂的信号通路相互作用，如介导着Ca2 +离子的信号转运(Yuan et al.,2014)、ABA的积累(Ma et al.,2009)、离子转运和转录因子活性(Vahisalu et al.,2008；Fujita et al.,2011)以及活性氧稳态平衡等。
然而，目前在农作物水稻中的盐胁迫调控网络研究相对较少，虽然已经初步积累了一些耐盐的种质资源，但是对控制耐盐的主要关键基因或者主效QTL的了解还相对较少。前期研究通过正向遗传学方法图为克隆定位到SKC1即OsHKT8，属于HKT家族的转运蛋白，位于1号染色体上，通过维持根中K+离子含量，从而保持着植物体内Na+/K+的稳态平衡，使植物在盐胁迫的压力下具有更强的耐受性(Ren et al.,2005)。后续又利用EMS诱变的群体，进行筛选鉴定到Drought Salt Tolerance(DST)，属于锌指结构的转录因子，发现其主要通过控制气孔密度和开度，进而调控水稻对不利环境的适应性(Huang et al.,2009)。随后研究人员又利用酵母双杂交技术筛选到与DST互作的蛋白DCA1，属于CHY型锌指结构的转录因子，发现它能够与DST互作形成异源四聚体的转录复合体，结合在靶基因过氧化物酶体peroxidase24 precursor(Prx 24)的启动子调控其表达。Prx 24属于H2O2的清除者，主要在保卫细胞中表达，通过控制保卫细胞中过氧化氢的量，来调控气孔的开闭，控制水稻对盐胁迫的响应(Cui et al.,2015)。
发明内容
本发明所要解决的技术问题是如何调控植物的耐盐性能。
为了解决以上技术问题，本发明提供了一种培调控植物耐盐性的方法，包括调控目的植物中耐盐相关蛋白质的编码基因的表达，其中，所述耐盐相关蛋白质，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耐盐性相关的蛋白质；
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。
本发明提供了一种来源于水稻的耐盐性相关蛋白，名称为OsPRR73，来源于水稻日本晴，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耐盐性相关的蛋白质；
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。
上述蛋白质中，序列表中的序列2由767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OsPRR73可来源于水稻。
与OsPRR73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OsPRR73相关的生物材料，为下述B1)至B7)中的任一种：
B1)编码OsPRR73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OsPRR73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1-2304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1-2304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OsPRR73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2304个核苷酸组成，其编码序列是序列表中序列1，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OsPRR73的核酸分子的表达盒(OsPRR73基因表达盒)，是指能够在宿主细胞中表达OsPRR73的DNA，该DNA不但可包括启动OsPRR73基因转录的启动子，还可包括终止OsPRR73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 Physiology120:979-992)；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)EMBOJ.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rbcSE9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OsPRR73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耐盐剂。
本发明所提供的植物耐盐剂含有所述蛋白质或/和所述蛋白质相关的生物材料。
上述植物耐盐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，上述植物耐盐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耐盐效果确定。
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、权利要求1所述蛋白质、或权利要求2或3所述生物材料在调控植物耐盐性中的应用；
P2、权利要求1所述蛋白质、或权利要求2或3所述生物材料在培育耐盐/不耐盐植物中的应用；
P3、权利要求1所述蛋白质、或权利要求2或3所述生物材料在制备提高/降低植物耐盐性的产品中的应用；
P4、权利要求1所述蛋白质、或权利要求2或3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耐盐物的方法，包括提高目的植物中所述蛋白质或其编码基因的表达量，得到耐盐植物；所述耐盐植物的耐盐性高于所述目的种子植物的耐盐性。
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耐盐性降低的转基因植物的方法。
本发明所提供的培育耐盐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，得到耐盐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。
本发明利用CRISPR/Cas9技术获得了OsPRR73功能缺失的突变体，进行180mM NaCl模拟盐胁迫处理，观察表型并统计恢复处理后的存活率，最终确定OsPRR73在响应水稻盐胁迫的过程中发挥正调控作用。本发明的OsPRR73可作为一个耐盐基因，在水稻抗盐性方面发挥重要作用。
附图说明
图1为180mM NaCl模拟盐胁迫处理，检测OsPRR73响应盐胁迫表达变化。材料为野生型两周龄幼苗，进行180mM NaCl处理。处理时为液体木村B营养液，将材料分别放入加盐和不加盐的营养液中，在处理过程中取材(即地上部分)，每个时间点三个生物学重复。图1中柱状图在正常和盐胁迫4小时条件下OsPRR73相对表达量。
图2为利用CRISPR/Cas9技术获得的NIP背景下osprr73突变体，分别命名为osprr73-C1和osprr73-C2，二者均属于功能缺失型突变体。
图3为osprr73-C1和osprr73-C2盐胁迫耐受性分析，纯合突变体osprr73-C1、osprr73-C2和野生型NIP四周龄幼苗材料进行盐胁迫处理的情况图。在处理前进行拍照，处理时营养液中加入180mM NaCl，处理21天后拍照，然后恢复时加入不含盐的正常营养液，恢复9天后拍照，并统计存活率(A图是未进行NaCl处理、处理21天和恢复处理9天的材料状态；B图是恢复生长之后存活率统计)。
图4为正常和盐胁迫处理7天后，NIP和osprr73-C1、osprr73-C2叶片中叶绿素含量和离子泄露率测定结果。(A图叶绿素含量测定结果；B图是离子泄露率结果)。
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。
本研究所用的实验材料为常用的粳稻品种日本晴(Nipponbare，简称NIP)。
下述实施例中的CRISPR/Cas9载体体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 Plant 8:1274-1284)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。
下述实施例中的农杆菌EHA105感受态细胞，北京博迈德公司。
营养液木村B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。其中，1L(200X)的A母液中包含(NH4)2SO49.64g、KNO33.7g、KH2PO4 4.96g、K2SO4 3.18g和MgSO4.7H2O 29.965g；1L(200X)的B母液中包含Ca(NO3)2.4H2O 17.235g；1L(1000X)的EDTA-Fe母液中：首先溶解5.57g FeSO4.7H2O于200mL蒸馏水中，然后加热溶解7.45g Na2EDTA于200mL蒸馏水中，将FeSO4.7H2O溶液与Na2EDTA溶液不断搅拌混合，冷却后定容至1L。1L(1000X)的微量元素母液中包含：H3BO4 2.86g、CuSO4.H2O 0.08g、ZnSO4.7H2O 0.22g、MnCl2.4H2O 1.81g和NaMO4.H2O 0.09g。每升营养液加入100～300ng硅酸钠，然后用浓盐酸调pH至5.8。
NB2培养基：NB Basal Medium粉末(品牌：Phytotech，从北京西美杰科技有限公司购买)(4.1g/L)溶于蒸馏水中，然后分别加入含300mg/L水解酪蛋白、500mg/L脯氨酸、500mg/L谷氨酰胺、30g/L蔗糖和2mg/L 2,4-D。
NB1培养基：在NB2培养基配方的基础上加入0.5mg/L 2,4-D。
NB1S1培养基：NB1培养基灭菌后，加入600mg/L头孢霉素和25mg/L Hyg。
NB1S2培养基：NB1培养基灭菌后，加入300mg/L头孢霉素和50mg/L Hyg。
RE1分化培养基：MS培养基粉末(4.5g/L)溶于蒸馏水中，然后分别加入300mg/L水解酪蛋白、1mg/L 6-BA、0.5mg/L KT、0.2mg/L ZT、0.25mg/L NAA、30g/L蔗糖和30g/L山梨醇。
RE2分化培养基：MS培养基粉末(4.5g/L)溶于蒸馏水中，然后分别加入300mg/L水解酪蛋白、50mg/L潮霉素、1mg/L 6-BA、0.5mg/L KT、0.2mg/L ZT、0.5mg/L NAA、30g/L蔗糖和20g/L山梨醇。
1/2MS培养基：MS培养基粉末(品牌：Caisson，从北京兰博利德商贸有限公司购买)(2.25g/L)溶于蒸馏水中，加入30g/L蔗糖。所有培养基均调节pH至5.8，培养基中加入琼脂8g/L，高压120℃灭菌15min
本发明下述实施例中所用的水稻材料培养方法为：挑选发育良好且饱满的种子，用蒸馏水在37℃培养箱中浸泡48小时，然后倒掉蒸馏水，在潮湿环境下继续催芽24小时。挑选萌发状态一致的种子点放至无底的96孔板中，放于盛有木村B营养液的器皿生长。温室生长条件为12小时光照/12小时黑暗，温度为30±2℃。在培养的过程中每隔3天更换一次新鲜营养液。
实施例1、耐盐相关的水稻OsPRR73编码基因的克隆
本发明的发明人从水稻日本晴中分离克隆出一个耐盐相关的水稻基因OsPRR73，如序列表的序列1所示，将其编码蛋白命名为OsPRR73蛋白，如序列表的序列2所示。
提取水稻日本晴总RNA并反转录成cDNA，以F:ATGGGTAGCGCCTGCGAAGC和R:TTATCTGTCTTCGTCTTGGC为引物进行PCR扩增。PCR反应程序为：94℃预变性2min；94℃变性30s，56℃退火30s，72℃延伸30s，35个循环；72℃延伸5min。将PCR产物进行Sanger测序。测序结果表明，该PCR扩增产物的核苷酸序列是序列表中序列1，其编码序列是序列表中序列1的第1-2304位核苷酸，编码序列2所示的蛋白质OsPRR73(序列表中序列2的第1-767位氨基酸残基)；将序列表中序列1所示的DNA命名为OsPRR73基因。
实施例2.OsPRR73缺失的突变体水稻构建
1.载体及重组菌的构建
利用序列1所示的DNA序列在E-CRISPR网站(http://www.e-crisp.org/E-CRISP/designcrispr.html)上筛选合适的靶点。结合评分及靶点位置，选择位于OsPRR73第一个外显子上的ATGGGTAGCGCCTGCGAAGCTGG为CRISPR/Cas9靶点之一。合成引物序列F:ggcATGGGTAGCGCCTGCGAAGC和R:aaacGCTTCGCAGGCGCTACCCA，按照文献中(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,Guo J,Chen L,Zhao X,Dong Z,Liu Y-G(2015)A Robust CRISPR/Cas9 Systemfor Convenient,High-Efficiency Multiplex Genome Editing in Monocot and DicotPlants.Mol Plant 8:1274-1284)进行pCRISPR/Cas9载体系统的构建，到载体pCRISPR/Cas9-OsPRR73。将载体pCRISPR/Cas9-OsPRR73导入农杆菌EHA105感受态细胞，得到重组农杆菌-pCRISPR/Cas9-OsPRR73。
2.pCRISPR/Cas9-OsPRR73转化水稻愈伤及阳性愈伤筛选
选取成熟饱满的日本晴水稻种子，去除外壳；用75％酒精消毒3～5min，倒去酒精；用灭菌后的蒸馏水冲洗3次；加入10％的NaClO(含有0.1％的吐温20)浸泡15～30min；倒掉NaClO，用灭菌后的蒸馏水冲洗5次。将消毒后的种子接入诱导愈伤培养基，26摄氏度培养20天，得到NIP水稻愈伤。
收集处于生长对数期的重组农杆菌-pCRISPR/Cas9-OsPRR73，用侵染液重悬后，挑选淡黄色的NIP水稻愈伤浸泡15-20min。倒掉侵染液，放于无菌滤纸上吹干。无菌操作放入含有10g/L的葡萄糖NB2(NB2C)培养基上，黑暗共培养两天。用含有300mg/L头孢霉素灭菌蒸馏水洗涤4-5次，直至变成清澈。放入无菌滤纸吸干多余水分，放入含有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。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。将能继续分化出的抗性愈伤转移至RE1培养基上进行分化，黑暗培养一周，再进行光照培养一周。挑选能分化出绿芽的愈伤转移至RE2培养基上继续分化，直至长出小苗。将分化出的小苗转移至1/2MS培养基上进行生根培养，然后练苗3-4天，移种到温室中。
根据上述步骤，获得了NIP背景下CRISPR/Cas9-OsPRR73的T0代转基因株系，即水稻OsPRR73突变体T0代。
实施例3转基因水稻CRISPR/Cas9-OsPRR73 T0代植株鉴定
取水稻OsPRR73突变体T0代的叶片提取DNA，以提取的DNA为模板，用OsPRR73的特异性引物，进行PCR扩增，引物序列分别为F:ACGTTGGGCGTATGATGCA；R:TCACCATGCTATGAAGCT。PCR反应程序为：94℃预变性2min；94℃变性30s，56℃退火30s，72℃延伸30s，35个循环；72℃延伸5min。将PCR产物进行Sanger测序，检查靶点位置是否发生突变。
其中，叶片DNA的提取按照如下方法进行：将水稻叶片放入1.5mL离心管，加入钢珠用研磨器研磨成粉末。离心管中加入500μL提取缓冲液，震荡混匀。加入等体积的氯仿和苯酚(1：1)混合液，震荡混匀，13500rpm离心5min。取300μL上清，加入等体积的氯仿，剧烈震荡混匀，13500rpm离心5min。取上清150μL，加入两倍体积的无水乙醇在-20℃沉淀30min。4℃，13500rpm，离心10min。倒掉上清，加入800μL的75％乙醇洗两遍，吹干后加入50μL ddH2O溶解。
鉴定结果表明，最终鉴定得到OsPRR73基因发生突变的2个独立转基因植株(分别为osprr73-C1和osprr73-C2)，比对DNA序列，其突变位点如图2所示。其中osprr73-C1缺失了5bp，使OsPRR73移码突变，导致其功能完全缺失。osprr73-C2在ATG下游17bp位置插入一个碱基A，同样导致移码突变，功能完全丧失。
实施例4OsPRR73在植物盐胁迫响应能力分析
为了检测OsPRR73如何参与盐胁迫响应，首先检测了其表达水平在正常和盐胁迫的条件下是否会发生改变。按照上述培养条件，将野生型生长14天后，开始进行180mM NaCl处理。将同一批NIP分别放入含180mM NaCl的木村B营养液和不含盐的正常营养液中，分别在0小时、4小时取地上部分材料，用锡纸包裹后迅速放入液氮中，然后-80℃保存备用，每个时间点取3个生物学重复。提取RNA用于检测OsPRR73的表达水平变化。植物RNA提取过程如下：将研磨好的水稻材料转移到干冰上速冻的1.5mL离心管中。往离心管中加0.5mL的Trizol，剧烈涡旋震荡混匀，室温放置5min。加入100μL的氯仿，混匀后室温静置2min，13000rpm，4℃离心15min。取300μL上清，加入等体积的氯仿，13000rpm，4℃离心12min。取150μL的上清，加入两倍体积的无水乙醇，-20℃放置30min，13000rpm，4℃离心15min。倒掉上清，瞬时离心后，去掉残留的乙醇，在超净台吹1min，加入35μL的DEPC水溶解RNA。测量RNA浓度，取1μg进行反转录，剩余RNA在-80℃保存。
按照上述方法提取RNA后，用Takara的试剂盒进行反转录，然后将cDNA稀释10倍，进行qPCR检测OsPRR73的表达水平变化。OsPRR73的qPCR引物F:AGGAGCGGAAAGAAACATAA，引物R:AACCTTGGAGGAGCAATCAG。内参基因Ubiquitin的qPCR引物F:AACCAGCTGAGGCCCAAGA，引物R:ACGATTGATTTAACCAGTCCATGA。qPCR的体系为7.5μL2×SYBR Green Master Mix，3μL稀释10倍的cDNA模板，10μM qPCR引物F和R各0.3μL，ddH2O定容至15μL。qPCR反应程序95℃预变性2min，95℃变性15s，55℃退火15s，72℃延伸15s，进行40个循环。程序结束后导出数据进行分析，采用2-ΔΔCT相对定量的方法对目的基因表达量进行分析。qPCR分析结果如图1所示：OsPRR73在盐胁迫处理4小时，表达水平明显上升。因此，OsPRR73对水稻抗盐胁迫可能起正向调控作用。
实施例5osprr73-Cs突变体植物耐盐性分析
同样按照上述培养条件，将野生型NIP和osprr73-C1、osprr73-C2突变体材料生长四周后，开始进行180mM NaCl胁迫处理，处理21天和未进行处理的材料进行拍照后，开始进行恢复处理，恢复9天后，进行拍照如图3A所示，其中图3A中的osprr73-C/L1表示osprr73-C1，osprr73-C/L2表示osprr73-C2，NIP表示日本晴野生型。恢复处理9天后，统计NIP和osprr73-C1、osprr73-C2的存活率，如图3B所示。NIP的存活率高达80％左右，而osprr73-C1和osprr73-C2的存活率仅有30％左右，该结果说明OsPRR73的功能缺失导致植株对盐胁迫更加敏感。由此可见，OsPRR73在植物响应盐胁迫的过程中起正调节的作用。
为了验证OsPRR73在植物盐胁迫耐受性中发挥的作用，我们又进一步检测了在正常和盐胁迫处理7天后的NIP和osprr73-C1、osprr73-C2突变体叶片中叶绿素含量和离子泄露率情况。二者的含量均与植物受胁迫的程度息息相关，具体测定方法如下，测定结果如图4所示。
其中，叶绿素含量测定方法为：取180mM NaCl处理7天和未经过处理的14天幼苗的叶片，记录离体的重量作为鲜重(W)。将叶片浸泡于3ml浓度为80％丙酮(v/v)中，在黑暗条件下于室温孵育48小时，并将记录其体积为V。取上清，用分光光度计在波长为OD645和OD663nm时测量吸光值，并记录。然后根据公式(8.02 A663+20.21 A645)*V/W计算叶绿素含量。
其中，离子泄露率测定方法为：取180mM NaCl处理7天和未处理的叶片在无菌的去离子水中轻轻摇晃过夜。用电导率仪测量电导率，记录为C1。然后将样品煮沸10-20分钟之后，冷却至室温，用电导率仪测量电导率，记录为C2。C1∶C2的比值为离子泄漏率。
根据上述方法，对测定结果进行分析，在正常条件下NIP和osprr73-Cs突变体叶片中叶绿素含量基本没有明显变化，但是在盐胁迫处理7天后，osprr73-C1和osprr73-C2突变体叶片的叶绿素含量明显低于野生型NIP。该结果说明，osprr73对盐胁迫更加敏感，其受盐胁迫危害的程度明显强于NIP。测定的离子泄露率结果表明，在正常条件下NIP和osprr73-C1、osprr73-C2突变体的离子泄露率均很低，且二者没有明显变化；然而NaCl胁迫导致osprr73-C1、osprr73-C2突变体中离子泄露率明显高于NIP，如图4B所示。由此可见，OsPRR73在盐胁迫响应中的确发挥正调控作用。
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> 中国科学院植物研究所
<120> 调控植物耐盐性的方法及耐盐相关蛋白
<160> 2
<170> SIPOSequenceListing 1.0
<210> 1
<211> 2304
<212> DNA
<213> Oryza sativa L.
<400> 1
atgggtagcg cctgcgaagc tggtacggac gagccttccc gagacgatgt taaggggaca 60
gggaatggca tcctggagaa tggtcatagt cacaagccag aggaggagga atggaggaat 120
ggcatgggag aggacttacc caatgggcac agtacaccac cagagcccca gcaaacagat 180
gaacagaagg agcaccaagt gcagattgtc cggtgggaga ggttcctccc tgtgaagaca 240
ctgagggtct tgctggtgga gaatgatgac tctacccgtc aggtggtcag cgcactgctt 300
cgtaagtgtt gttatgaagt tatccctgct gaaaatgggc tacatgcatg gcaatgtctt 360
gaagatctgc aaaaccacat tgaccttgta ttgaccgagg tcgtaatgcc acgtctgtct 420
ggcattggtc tgcttagtaa gatcacaagc cacaaaattt gcaaggatat tcccgtgatt 480
atgatgtctt cgaatgactc aatgggtaca gtctttaagt gtttgtcaaa aggagcagtt 540
gactttctag tgaagcctat acgtaagaat gaacttaaga acctttggca gcatgtttgg 600
agacgatgcc acagttccag tggcagtgga agcgaaagtg gcatccgaac acaaaagtgt 660
accaaaccaa aggttgatga tgaatatgag aataacagcg gtagcaataa tgacaacgag 720
gatgatgatg acaatgatga agatgatgac gacttaagtg ttggacacaa cgctagggat 780
ggcagtgata atggcagtgg cactcaaagt tcatggacaa agcgtgcagt ggagattgac 840
agcccacaac aaatgtctcc tgatcaacca tccgatctac cagatagtac ttgtgcgcaa 900
gtaattcacc ccacatcaga gatatgcagc aacaggtggt taccgactgc aaataaaagg 960
agcggaaaga aacataaaga aaataacgat gactccatgg ggaagtactt agaaatagga 1020
gctcctagaa attctagtat ggagtaccaa tcttctccaa gagagatgtc cgttaatcca 1080
acagaaaaac agcatgaaac tctcatgccc caaagtaaaa caacaagaga aacagatagt 1140
aggaacacac agaatgaacc aactactcaa actgttgatt taattagttc aatagccaga 1200
agcacagatg acaaacaagt agttagaatc aataatgctc ctgattgctc ctccaaggtt 1260
ccagatggaa atgataaaaa tcgtgattct ctcattgata tgacatctga agagttgggt 1320
ttgaagagat tgaaaacaac tggatctgca actgaaatcc atgatgaacg aaatattctg 1380
aaaagatcag atctctcagc tttcaccagg taccatacaa ctgtggcttc taatcaaggt 1440
ggagctggat ttgggggaag ctgttcacct caagataaca gttcagaggc tctgaaaaca 1500
gactccaact gcaaggtgaa gtcaaattca gatgctgctg aaataaagca aggctccaat 1560
ggtagtagca acaacaatga catgggctcc agtactaaga atgccatcac aaaaccttct 1620
tcaaacaggg gaaaagtgat atcaccatca gctgtcaaag ctacccaaca tacatcagca 1680
ttccatcctg tgcagcgtca aacgtcacct gctaatgttg tagggaaaga caaagttgat 1740
gaaggaattg ctaatggagt taatgtgggc caccctgtag atgtacaaaa tagctttatg 1800
cagcaccatc atcatgttca ttactacgtc catgttatga cacagcagca gcagcagcca 1860
tccattgagc gaggatcatc agatgctcag tgtggttcat ccaatgtatt tgatcctccc 1920
attgaaggtc atgcggcaaa ctatagtgtg aacgggagct tttcaggtgg ccataatgga 1980
aacaatgggc aaagaggacc tagtactgct cccaatgttg ggaggccaaa catggagact 2040
gttaatggta tcgtggatga aaatggggct ggaggtggca atggaagtgg gagcggtagt 2100
ggtaatgact tgtatcagaa tggggtctgt taccgagaag ctgcattgaa caaattcaga 2160
cagaaacgga aagtgaggaa ctttggaaaa aaggtgcgct atcagagcag aaagaggttg 2220
gctgagcagc gccctcggat ccgcgggcaa ttcgtgcgac aatctggaca ggaagatcag 2280
gcaggccaag acgaagacag ataa 2304
<210> 2
<211> 767
<212> PRT
<213> Oryza sativa L.
<400> 2
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Thr Lys Arg Ala Val Glu Ile Asp Ser Pro Gln Gln Met Ser Pro Asp
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Gln Pro Ser Asp Leu Pro Asp Ser Thr Cys Ala Gln Val Ile His Pro
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Ser Gly Lys Lys His Lys Glu Asn Asn Asp Asp Ser Met Gly Lys Tyr
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Leu Glu Ile Gly Ala Pro Arg Asn Ser Ser Met Glu Tyr Gln Ser Ser
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Met Pro Gln Ser Lys Thr Thr Arg Glu Thr Asp Ser Arg Asn Thr Gln
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Asn Glu Pro Thr Thr Gln Thr Val Asp Leu Ile Ser Ser Ile Ala Arg
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Ser Thr Asp Asp Lys Gln Val Val Arg Ile Asn Asn Ala Pro Asp Cys
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Ser Ser Lys Val Pro Asp Gly Asn Asp Lys Asn Arg Asp Ser Leu Ile
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Asp Met Thr Ser Glu Glu Leu Gly Leu Lys Arg Leu Lys Thr Thr Gly
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Gly Ala Gly Phe Gly Gly Ser Cys Ser Pro Gln Asp Asn Ser Ser Glu
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Gln Lys Arg Lys Val Arg Asn Phe Gly Lys Lys Val Arg Tyr Gln Ser
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Arg Lys Arg Leu Ala Glu Gln Arg Pro Arg Ile Arg Gly Gln Phe Val
740 745 750
Arg Gln Ser Gly Gln Glu Asp Gln Ala Gly Gln Asp Glu Asp Arg
755 760 765
Claims (4)
1.一种正调控水稻耐盐性的方法，其特征在于，正调控目的水稻中耐盐相关蛋白质的编码基因的表达，其中，所述耐盐相关蛋白质，是氨基酸序列是序列表中序列2的蛋白质。
2.权利要求1中所述耐盐相关蛋白质在培育耐盐/不耐盐水稻中的应用。
3.一种培育耐盐性降低的转基因水稻的方法，包括降低目的水稻中权利要求1中所述耐盐相关蛋白质的编码基因的表达，得到耐盐性低于所述目的水稻的转基因水稻。
4.根据权利要求3所述的方法，其特征在于：所述降低目的水稻中权利要求1中所述耐盐相关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水稻中权利要求1中所述耐盐相关蛋白质的含量和/或活性实现的。
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