CN110218247B - PwRBP1和PwNAC1两种蛋白互作协同提高植物耐逆性及其应用 - Google Patents
PwRBP1和PwNAC1两种蛋白互作协同提高植物耐逆性及其应用 Download PDFInfo
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- CN110218247B CN110218247B CN201910446974.5A CN201910446974A CN110218247B CN 110218247 B CN110218247 B CN 110218247B CN 201910446974 A CN201910446974 A CN 201910446974A CN 110218247 B CN110218247 B CN 110218247B
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Abstract
本发明首次公开了RNA结合蛋白PwRBP1和NAC转录因子PwNAC1蛋白能够互作形成异源二聚体，协同促进提高植物耐逆性。本发明将青杄中的PwRBP1和PwNAC1编码基因共同导入拟南芥中得到共转PwRBP1和PwNAC1拟南芥植株。实验证明，相比于受体植株以及单转PwRBP1或PwNAC1拟南芥植株，共转两种基因的拟南芥植株的耐旱性和耐盐性得到显著提高，表明PwRBP1和PwNAC1能够协同促进植物提高耐逆性，适于推广应用。
Description
技术领域
本发明属于生物技术领域，具体涉及青杄中的RNA结合蛋白PwRBP1以及NAC转录因子PwNAC1相互作用协同提高植物的耐逆性。
背景技术
干旱胁迫与盐胁迫是两种主要的非生物环境胁迫，其会导致植物产生诸如渗透伤害、离子伤害、活性氧和有毒物质的积累等，从而严重影响植物种子的萌发、光合作用、植物生长发育等各项生理和生长过程。而植物在长期进化过程中会形成一系列有效的作用机制，其中包括调控胁迫相关基因的表达以应对外界不良环境。
NAC转录因子是植物中特异的转录调控因子，RBP蛋白是一种RNA结合蛋白，其在真核生物中具有结合RNA能力的蛋白，能在转录后水平调控的过程中起关键作用，并协调相关mRNA的翻译，两者均能在生物体的生长发育、应对低温、水淹、干旱高盐、高温等胁迫中均能发挥重要的作用。目前在模式植物如拟南芥和水稻中的这两种基因与植物抗逆性相关研究已经取得了重要进展，但木本植物尤其针叶树种中有关NAC转录因子和RBP蛋白基因的功能及两者之间的关联的研究报道较少，基于此，提出本发明。
发明内容
本发明所要解决的技术问题是如何调控植物耐逆性。
为解决上述技术问题，本发明首先提供了两种可以相互作用且协同提高植物耐逆性相关的蛋白。
本发明所提供的可协同提高植物抗逆性相关蛋白的名称分别为PwNAC1及PwRBP1，来源于青杄(Picea wilsonii Mast.)，为如下a)或b)或c)或d)的蛋白：
a)包含SEQ ID NO.2或SEQ ID NO.4所示氨基酸序列的蛋白；在一些实施例中，是如SEQ ID NO.2或SEQ ID NO.4所示氨基酸序列的蛋白；
b)在SEQ ID NO.2或SEQ ID NO.4所示的蛋白质蛋白的N端和/或C端连接标签得到的融合蛋白；
c)将SEQ ID NO.2或SEQ ID NO.4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/或添加得到的具有相同功能的蛋白；
d)与SEQ ID NO.2或SEQ ID NO.4所示的氨基酸序列具有75％或75％以上的同源性且具有相同功能的蛋白。
其中，SEQ ID NO.2由430个氨基酸残基组成，SEQ ID NO.4由271个氨基酸残基组成。
上述b)中标签可以是表1所述序列标签，该标签为了使a)中的蛋白便于纯化，可连接在在序列表中SEQ ID NO.2或SEQ ID NO.4所示的蛋白的氨基末端或羧基末端。
表1、标签的序列
| 标签 | 残基 | 序列 |
| Poly-Arg | 5-6(通常为5个) | RRRRR |
| Poly-His | 2-10(通常为6个) | HHHHHH |
| FLAG | 8 | DYKDDDDK |
| Strep-tag II | 8 | WSHPQFEK |
| c-myc | 10 | EQKLISEEDL |
上述c)中的蛋白PwRBP1或PwNAC1，所述一个或几个氨基酸残基的取代和/或缺失和/或添加为不超过10个氨基酸残基的取代和/或缺失和/或添加。
上述c)中的蛋白PwRBP1或PwNAC1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c)中的蛋白PwRBP1或PwNAC1的编码基因可通过将SEQ ID NO.1或SEQ IDNO.3所示的DNA序列中缺失一个或几个氨基酸残基的密码子，和/或进行一个或几个碱基对的错义突变，和/或在其5′端和/或3′端连上表1所示的标签的编码序列得到。
为解决上述技术问题，本发明又提供了与PwRBP1和PwNAC1蛋白相关的生物材料。
本发明提供的与PwRBP1和PwNAC1蛋白相关的生物材料为下述A1)至A12)中的任一种：
A1)编码PwRBP1和PwNAC1蛋白的核酸分子；
A2)含有A1)所述核酸分子的表达盒；
A3)含有A1)所述核酸分子的重组载体；
A4)含有A2)所述表达盒的重组载体；
A5)含有A1)所述核酸分子的重组微生物；
A6)含有A2)所述表达盒的重组微生物；
A7)含有A3)所述重组载体的重组微生物；
A8)含有A4)所述重组载体的重组微生物；
A9)含有A1)所述核酸分子的转基因植物细胞系；
A10)含有A2)所述表达盒的转基因植物细胞系；
A11)含有A3)所述重组载体的转基因植物细胞系；
A12)含有A4)所述重组载体的转基因植物细胞系。
上述生物材料中，A1)所述核酸分子为如下1)或2)或3)所示的基因：
1)其编码序列是SEQ ID NO.1或SEQ ID NO.3所示的DNA分子，在一些实施例中所述DNA分子是cDNA分子或基因组DNA分子；
2)与1)限定的核苷酸序列具有75％或75％以上同一性，且编码PwRBP1或PwNAC1蛋白的cDNA分子或基因组DNA分子；
3)在严格条件下与1)或2)限定的核苷酸序列杂交，且编码PwRBP1或PwNAC1蛋白的cDNA分子或基因组DNA分子。
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。
其中，SEQ ID NO.1由1293个核苷酸组成，SEQ ID NO.3由816个核苷酸组成。
本领域相关技术人员能够通过使用已知的如定向进化和点突变等实验方法，对本发明的编码PwRBP1或PwNAC1的核苷酸序列进行突变。凡是经过人工修饰的，具有与本发明分离得到的PwRBP1或PwNAC1的核苷酸序列75％或者更高同一性的核苷酸，只要编码PwRBP1或PwNAC1且具有相同功能，均是衍生于本发明的核苷酸序列并且等同于本发明的序列。
这里使用的术语“同一性”指与天然核酸序列的序列相似性。“同一性”包括与本发明的编码SEQ ID NO.2所示的氨基酸序列组成的蛋白的核苷酸序列具有75％或更高，或85％或更高，或90％或更高，或95％或更高同一性的核苷酸序列。同一性可以用肉眼或计算机软件进行评价。使用计算机软件，两个或多个序列之间的同一性可以用百分比(％)表示，其可以用来评价相关序列之间的同一性。
上述75％或75％以上同一性，可为80％、85％、90％或95％以上的同一性。
上述生物材料中，A2)所述的含有编码PwRBP1或PwNAC1的核酸分子的表达盒(PwRBP1或PwNAC1基因表达盒)，是指能够在宿主细胞中表达PwRBP1或PwNAC1的DNA，该DNA不但可包括启动PwRBP1或PwNAC1转录的启动子，还可包括终止PwRBP1或PwNAC1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及诱导型启动子。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。
可用现有的表达载体构建含有所述PwRBP1或PwNAC1基因表达盒的重组载体。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、pBin438、pCAMBIA1205、pCAMBIA1302、pCAMBIA2300、pCAMBIA2301、pCAMBIA1301、pCAMBIA1300、pBI121、pCAMBIA1391-Xa或pCAMBIA1391-Xb(CAMBIA公司)等。使用PwRBP1或PwNAC1基因构建重组表达载体时，可在其转录起始核苷酸前加上任何一种增强型、组成型、组织特异型或诱导型启动子，如花椰菜花叶病毒(CAMV)35S启动子、泛素(Ubiquitin)基因启动子(pUbi)等，它们可单独使用或与其它的植物启动子结合使用；此外，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，如加入在植物中表达可产生颜色变化的酶或发光化合物的基因(GUS基因、GFP基因、荧光素酶基因等)、具有抗性的抗生素标记物(庆大霉素标记物、卡那霉素、氯霉素标记物等)或是抗化学试剂标记基因(如抗除莠剂基因)等。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，所述载体可为质粒、黏粒、噬菌体或病毒载体。在本发明的实施例中，所述重组载体具体可为在pCAMBIA1205载体上插入上述PwRBP1(SEQ ID NO.1)或PwNAC1(SEQ ID NO.3)基因得到的重组表达载体。
上述生物材料中，所述微生物可为酵母、细菌、藻或真菌，如农杆菌。在本发明的实施例中，采用的农杆菌为GV3101。
上述生物材料中，所述转基因植物细胞系均不包括繁殖材料。
为解决上述技术问题，本发明还提供PwRBP1和PwNAC1蛋白或上述生物材料的协同作用的新用途。
本发明提供了PwRBP1和PwNAC1蛋白或上述生物材料协同在调控植物耐逆性中的应用。
本发明还提供了PwRBP1或PwNAC1蛋白或上述生物材料协同在培育耐逆性提高或降低的转基因植物中的应用。
本发明还提供了PwRBP1或PwNAC1蛋白或上述生物材料协同在植物育种中的应用。
上述应用中，所述耐逆为耐盐性和/或耐旱性。
上述应用中，所述植物为单子叶植物或双子叶植物，所述双子叶植物具体可为豆科植物和/或十字花科植物和/或菊科植物；所述豆科植物可为大豆、百脉根、苜蓿或水黄皮；所述十字花科植物可为拟南芥或油菜；所述菊科植物可为向日葵；所述拟南芥可为拟南芥(哥伦比亚生态型col-0)。
为解决上述技术问题，本发明最后提供了一种通过共转能够协同互作的PwRBP1和PwNAC1蛋白培育抗逆性改善的转基因植物的方法；在一些实施例中，所述改善为提高；在另一些实施例中，所述改善为降低。
本发明提供的培育提高抗逆性的转基因植物的方法包括提高受体植物中PwRBP1和PwNAC1这两种协同互作蛋白的表达量和/或活性，得到转基因植物的步骤；所述转基因植物对于高盐和干旱的耐受能力均高于所述受体植物。
本发明所述的PwRBP1和PwNAC1两种蛋白经酵母双杂交和LCI实验验证能够形成异源二聚体，协同互作提高植物的耐逆性。
在本发明的一个试验方法中，为了验证PwRBP1和PwNAC1蛋白能够互作形成异源二聚体协同发挥提高植物耐逆性的作用，分别将PwRBP1和PwNAC1基因克隆到PDEST32和PDEST22载体中，构成PwRBP1-PDEST22、PwRBP1-PDEST32、PwNAC1-PDEST22以及PwNAC1-PDEST32重组载体，并分别将PDEST32+PDEST22、PwNAC1-PDEST32+PDEST22、PwRBP1-PDEST22+PDEST32、PwNAC1-PDEST32+PwRBP1-PDEST22以及PwRBP1-PDEST32+PwNAC1-PDEST22转化到MaV203酵母菌株中，以在酵母系统中验证PwRBP1和PwNAC1蛋白能够互作形成异源二聚体协同发挥作用。
在本发明的一个试验方法中，将分别将PwRBP1和PwNAC1克隆至pCAMBIA-NLuc(N端荧光素酶)和pCAMBIA-CLuc(C端荧光素酶)载体上分别构成PwRBP1-NLuc、PwRBP1-CLuc、PwNAC1-Nluc以及PwNAC1-CLuc重组载体。将混合载体PwNAC1-NLuc+PwRBP1-CLuc、PwNAC1-CLuc+PwRBP1-NLuc、PwRBP1-NLuc+CLuc、NLuc+PwRBP1-CLuc、NLuc+CLuc分别转入到GV3101农杆菌菌株中以验证本实验所述PwRBP1和PwNAC1能够形成异源二聚体协同发挥作用。
上述方法中，所述共同提高受体植物中PwRBP1和PwNAC1蛋白的表达量和/或活性的方法为在受体植物中均表达或过表达PwRBP1蛋白和PwNAC1。
上述方法中，所述共同表达或过表达的方法为将PwRBP1和PwNAC1蛋白质的编码基因共同导入受体植物；所述PwRBP1和PwNAC1蛋白的编码基因的核苷酸序列是SEQ ID NO.1和SEQ ID NO.3所示的DNA分子。
在本发明的一个试验方法中，分别将含有PwRBP1蛋白的编码基因(即SEQ ID NO.1所示的核苷酸)以及PwNAC1蛋白的编码基因(即SEQ ID NO.3所示的核苷酸)通过含有PwRBP1蛋白或PwNAC1蛋白的编码基因的表达盒的重组载体pCAMBIA1205-PwRBP1或pCAMBIA1205-PwNAC1分别导入农杆菌GV3101中。所述重组载体pCAMBIA1205-PwRBP1表达PwRBP1蛋白，重组载体pCAMBIA1205-PwNAC1表达PwNAC1蛋白。
上述方法中所述共转PwRBP1和PwNAC1两种蛋白的转基因植物的抗逆性高于所述野生型受体植物以及分别单转PwRBP1或PwNAC1蛋白转基因植物的具体表现为在逆境胁迫下产生如下方式：所述共转PwRBP1和PwNAC1两种蛋白的转基因植物在高浓度盐或高浓度甘露醇的胁迫下种子萌发率和/或幼苗根长和/或存活率大于所述所述受体野生型及单转PwRBP1或PwNAC1蛋白转基因植物。示例性的，上述高盐环境具体可以是100mM、200mM的NaCl水溶液引起的环境；上述干旱环境具体可以是200mM、300mM的甘露醇水溶液模拟得到的干旱环境，也可以是停止浇水11天的干旱处理环境。
上述方法中，所述转基因植物理解为不仅包含将所述PwRBP1和PwNAC1基因共同导入受体植物得到的第一代转基因植物同时包括其子代。对于转基因植物，可以在该物种中同时繁殖本发明中的两种基因，也可用常规育种技术将这两种基因共同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。
上述方法中，所述受体植物为单子叶植物或双子叶植物，所述双子叶植物具体可为豆科植物和/或十字花科植物和/或菊科植物；所述豆科植物可为大豆、百脉根、苜蓿或水黄皮；所述十字花科植物可为拟南芥或油菜；所述菊科植物可为向日葵；所述拟南芥可为拟南芥(哥伦比亚生态型Col-0)。
本发明首先发现了青杄中的PwRBP1或PwNAC1蛋白能够协同互作形成异源二聚体，并通过酵母双杂体系及烟草LCI体系中验证了该结果，然后将两者共同导入拟南芥中得到共转PwRBP1和PwNAC1拟南芥，并发现同时转入PwRBP1和PwNAC1拟南芥对于高盐和干旱的耐受能力均高于原受体野生型植株及单转PwRBP1或PwNAC1转基因植株。其具体表现如下：在100mM浓度的NaCl胁迫下，共转PwRBP1和PwNAC1基因植株的种子萌发率、根长均显著高于原野生型植株以及单转PwRBP1或PwNAC1转基因植株，以野生型为对照，其中萌发率在萌发初期提高了72.65％，单转PwRBP1和单转PwNAC1仅分别提高了45.82％和51.27％，共转植株的根长则比野生型对照组提高了46.83％，单转PwRBP1和单转PwNAC1分别提高了43.32％和43.67％。在浇灌200mM浓度的NaCl溶液胁迫下，处理11天后共转PwRBP1和PwNAC1基因幼苗的存活率比原野生型植株的存活率提高了65.18％，单转PwRBP1和单转PwNAC1仅分别提高了49.33％和53.39％；并且本发明获得的共转PwRBP1和PwNAC1基因拟南芥的种子在200mM浓度的甘露醇培养基胁迫下其种子萌发率、根长均显著高于原野生型植株以及单转PwRBP1或PwNAC1转基因植株，其中共转2种基因的植株萌发率为野生型的2.36倍，单转PwRBP1和单转PwNAC1仅为野生型的1.41和1.18倍，共转2种基因植株根长则比野生型对照组提高了44.23％，单转PwRBP1和单转PwNAC1仅分别提高了18.13％和30.50％。在培养基质中停止浇水11天，又复水3天后共转2种基因幼苗的存活率比原野生型植株提高了2倍，单转PwRBP1和单转PwNAC1仅为野生型的1.55和1.43倍。上述结果表明，青杄中的PwRBP1和PwNAC1蛋白能够协同互作形成异源二聚体共同提高植物耐旱性和耐盐性的功能。
附图说明
图1为酵母双杂和LCI实验验证PwRBP1与PwNAC1蛋白互作结果。
图2为转PwRBP1/NAC1拟南芥分子检测结果。
图3为转PwRBP1/NAC1拟南芥、转PwRBP1、转PwNAC1及野生型拟南芥种子在100mM的NaCl处理下萌发8天的观测结果。
图4为转PwRBP1/NAC1拟南芥、转PwRBP1、转PwNAC1及野生型拟南芥种子在100mM的NaCl处理下萌发率测定结果。
图5为转PwRBP1/NAC1拟南芥、转PwRBP1、转PwNAC1及野生型拟南芥种子在200mM的甘露醇处理下萌发8天的观测结果。
图6为转PwRBP1/NAC1拟南芥、转PwRBP1、转PwNAC1及野生型拟南芥种子在200mM的甘露醇处理下萌发率测定结果。
图7为转PwRBP1/NAC1拟南芥、转PwRBP1、转PwNAC1及野生型拟南芥种子萌发后转入含不同浓度NaCl和甘露醇的平板生长8天的根长观察结果。
图8为转PwRBP1/NAC1拟南芥、转PwRBP1、转PwNAC1及野生型拟南芥种子萌发后转入含不同浓度NaCl和甘露醇的平板生长8天的根长的数值统计。
图9为转PwRBP1/NAC1拟南芥、转PwRBP1、转PwNAC1及野生型拟南芥幼苗在200mM的NaCl处理11天后的观测结果。
图10为转PwRBP1/NAC1拟南芥、转PwRBP1、转PwNAC1及野生型拟南芥幼苗在干旱处理11天，复水3天后的观测结果。
图11为转PwRBP1/NAC1拟南芥、转PwRBP1、转PwNAC1及野生型拟南芥幼苗在盐处理和干旱处理后存活率测定结果。
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。
下述实施例中的pCAMBIA1205、PDEST32、PDEST22、pCAMBIA-NLuc(N端荧光素酶)和pCAMBIA-CLuc(C端荧光素酶)载体可从申请人(北京林业大学)处获得，该生物材料只为重复本发明的相关实验所用，不可作为其它用途使用。
实施例1、PwRBP1与PwNAC1蛋白互作
一、酵母双杂交体系验证PwRBP1与PwNAC1蛋白互作
为了验证PwRBP1与PwNAC1蛋白能够互作形成异源二聚体，通过Gateway技术(Invitrogen)分别将PwRBP1和PwNAC1基因克隆到PDEST32和PDEST22载体中，构成PwRBP1-PDEST22、PwRBP1-PDEST32、PwNAC1-PDEST22以及PwNAC1-PDEST32重组载体，并分别将PDEST32+PDEST22、PwNAC1-PDEST32+PDEST22、PwRBP1-PDEST22+PDEST32、PwNAC1-PDEST32+PwRBP1-PDEST22以及PwRBP1-PDEST32+PwNAC1-PDEST22并根据酵母方案手册(Invitrogen)转化到MaV203酵母菌株中并分别将含有重组质粒的酵母涂布代在SD-Trp-Leu-或SD-Trp-Leu-His-Ura+30mM 3AT+x-α-gal培养基平板上生长并观察酵母生长情况。其中PDEST32+PDEST22为阴性对照，PEXP32/Krev1+PEXP22/RalGDS-wt为阳性对照，每组实验重复3次。实验结果表明仅当PwRBP1和PwNAC1同时存在的情况下(PwNAC1-PDEST32+PwRBP1-PDEST22和PwRBP1-PDEST32+PwNAC1-PDEST22)酵母菌斑呈蓝色为阳性菌落(图1A)，表明PwRBP1能够与PwNAC1蛋白互作形成异源二聚体，共同发挥抗逆功能。
二、烟草LCI实验验证PwRBP1与PwNAC1蛋白互作
为了验证PwRBP1与PwNAC1蛋白能够互作形成异源二聚体，通过Gateway技术(Invitrogen)分别将PwRBP1和PwNAC1基因克隆到pCAMBIA-NLuc(N端荧光素酶)和pCAMBIA-CLuc(C端荧光素酶)载体上分别构成PwRBP1-NLuc和PwRBP1-CLuc重组载体。将混合载体PwNAC1-NLuc+PwRBP1-CLuc、PwNAC1-CLuc+PwRBP1-NLuc、PwRBP1-NLuc+CLuc、NLuc+PwRBP1-CLuc、NLuc+CLuc转入到GV3101农杆菌菌株，注射到烟草叶片细胞中。25℃黑暗处理72h后，通过荧光成像系统观察，在同时注射了PwNAC1和PwRBP1的烟草叶片细胞中可以检测到荧光素酶的表达，而在注射了PwRBP1-NLuc+CLuc、NLuc+PwRBP1-CLuc、NLuc+CLuc混合质粒的烟草叶片细胞没有检测到荧光素酶的表达(图1B)，说明PwNAC1与PwRBP1蛋白在烟草细胞可以形成异源二聚体结构。
实施例2、PwRBP1与PwNAC1蛋白协同提高植物耐逆性中的应用
一、共转PwRBP1与PwNAC1拟南芥的构建
1、PwRBP1和PwNAC1重组表达载体的获得
制备下述扩增引物对：
引物1：5’-ATGATGCAACCCACAGCAGG-3’；
引物2：5’-CTGATGAATATCGACTTTGGAAAGTC-3’。
引物3：5’-TGAGCAAAGACCCCAACGAGA-3’；
引物4：5’-CTTTATGCTTCCGGCTCGT-3’。
以青杄的cDNA为模板，用上述引物1和2进行PCR扩增，得到大小为1293bp的PCR产物1。以青杄的cDNA为模板，采用上述引物3和引物4进行PCR扩增，得到大小为816bp的PCR产物2。
将上述获得的PCR产物1和产物2分别和pCAMBIA1205载体经限制性内切酶酶切后进行连接，得到连接产物1和连接产物2。然后将连接产物1和2分别转化大肠杆菌DH5α感受态细胞，并涂布在含有35μg/ml氯霉素的LB平板上培养过夜。挑取白色单菌落于含有35μg/ml氯霉素的LB液体培养基中培养过夜并进行菌落PCR鉴定；同时碱法提取质粒DNA进行序列测定。测序结果表明：连接产物1所得的重组表达载体为将序列表中SEQ ID NO.1所示的PwRBP1插入表达载体pCAMBIA1205，且保持载体pCAMBIA1205其他序列不变后得到的载体，将其命名为pCAMBIA1205-PwRBP1。连接产物2所得的重组表达载体为将序列表中SEQ IDNO.3所示的PwNAC1插入表达载体pCAMBIA1205，且保持载体pCAMBIA1205其他序列不变后得到的载体，将其命名为pCAMBIA1205-PwNAC1。
2、共转PwRBP1与PwNAC1拟南芥的获得
1)转PwRBP1或PwNAC1拟南芥的获得
将上述实验中制备的重组载体pCAMBIA1205-PwRBP1或pCAMBIA1205-PwNAC1转化农杆菌GV3101的感受态细胞(购置于上海唯地生物技术公司)，得到重组菌GV3101/pCAMBIA1205-PwRBP1或GV3101/pCAMBIA1205-PwNAC1。
将重组菌GV3101/pCAMBIA1205-PwRBP1或GV3101/pCAMBIA1205-PwNAC1单克隆接种于含35mg/L氯霉素的YEB液体培养基中，28℃振荡培养两天。将培养液3000rpm/min离心5分钟，所得农杆菌沉淀用含5％蔗糖和0.03％SilwetL-77的侵染液悬浮。
采用花絮浸染法转化哥伦比亚生态型野生型拟南芥(Col-0)(购自ABRC)，收获当代转基因拟南芥植株所接的种子(T1代)，在含有40μg/ml潮霉素(Hygromycin B)和40μg/ml羧苄青霉素(Carbenicllin)的MS培养基筛选萌发的种子。将在上述培养基上萌发的T1代幼苗移到培养土中，收获种子(T2代)，然后经相同的筛选过程得到纯合的转PwRBP1或PwNAC1拟南芥植株(T4代)种子。最后将转PwRBP1或PwNAC1拟南芥植株(T4代)种子直接播种到培养土中，长出的PwRBP1或PwNAC1拟南芥植株(T4代)在长日照条件下生长两周左右开花。
采用同样的方法，将空载体pCAMBIA1205转入野生型拟南芥中，得到转空载体拟南芥，播种、传代，得到空载体过表达拟南芥植株(T4代)。
2)共转PwRBP1与PwNAC1拟南芥的获得
与上述1)中实验方法相同采用花絮浸染法转化，使用上述实验所述的GV3101/pCAMBIA1205-PwNAC1菌株侵染上述实验中获得的转PwRBP1基因的T3代拟南芥，或采用GV3101/pCAMBIA1205-PwRBP1菌株侵染上述实验中获得的转PwNAC1基因的T3代拟南芥以获得共转PwRBP1与PwNAC1两种基因的拟南芥转基因株系并同样筛选至T4代。最后将共转PwRBP1与PwNAC1拟南芥植株(T4代)种子直接播种到培养土中，长出的共转PwRBP1与PwNAC1拟南芥植株(T4代)在长日照条件下生长两周左右开花。
3)转基因拟南芥的分子检测
提取各转基因拟南芥植株(T4代)的RNA，反转录得到cDNA，以其为模板，进行实时荧光定量实验检测PwRBP1或PwNAC1的表达量，并以野生型拟南芥为对照。
结果如图2A所示，转PwNAC1、转PwRBP1以及共转两种基因的拟南芥植株(T4代)相应PwRBP1、PwNAC1的表达量显著高于野生型拟南芥(Col-0)以及转空载体PCM1205的株系，表明PwRBP1和PwNAC1分别在单转和共转拟南芥植株(T4代)中过表达。
此外，提取转基因拟南芥植株(T4代)的总蛋白，以其为模板，经SDS-PAGE跑胶、转膜、封闭后以GFP标签为抗体进行检测，并以野生型拟南芥为对照，检测结果如图2B所示，转基因拟南芥植株(T4代)经GFP抗体结合后有明显条带，而野生型拟南芥则没有检测到条带，证明该T4代株系中PwRBP1或PwNAC1的表达量明显提高。
二、共转PwRBP1与PwNAC1拟南芥功能研究
1、种子萌发试验
1)耐盐性研究
在含有100mM的NaCl的MS培养基上进行种子萌发实验，实验中取共转PwRBP1与PwNAC1拟南芥植株(T4代)、转PwRBP1拟南芥植株(T4代)、转PwNAC1拟南芥植株(T4代)、转空载体拟南芥植株(T4代)以及野生型拟南芥(Col-0)种子进行播种，其培养条件为光照16小时，黑暗8小时，光强为300-400μmol m-2s-1，光照下的室温为22-24℃，相对湿度为70-90％；黑暗条件下的室温为18-20℃，相对湿度大于90％。每个株系100粒种子，实验重复3次，结果取平均值。
实验进行第8天统计各类种子的萌发率结果如图3、4所示。在100mM的NaCl处理下，共转PwRBP1与PwNAC1、转PwRBP1、转PwNAC1、转空载体以及野生型拟南芥(Col-0)种子均能在第4天萌发，共转PwRBP1和PwNAC1基因植株的种子萌发率在萌发初期提高了72.65％，单转PwRBP1和单转PwNAC1仅分别提高了45.82％和51.27％，该表明结果表明在100mM NaCl胁迫处理下，共转PwRBP1与PwNAC1拟南芥的种子萌发率显著高于野生型及单转PwRBP1或PwNAC1植株，具有更好的耐盐能力。
野生型拟南芥和空载体过表达拟南芥植株(T4代)的结果无显著差异。
2)耐旱性研究
在含有200mM的甘露醇的MS培养基上进行种子萌发实验，实验中取共转PwRBP1与PwNAC1拟南芥植株(T4代)、转PwRBP1拟南芥植株(T4代)、转PwNAC1拟南芥植株(T4代)、转空载体拟南芥植株(T4代)以及野生型拟南芥(Col-0)种子进行播种，其培养条件为光照16小时，黑暗8小时，光强为300-400μmol m-2s-1，光照下的室温为22-24℃，相对湿度为70-90％；黑暗条件下的室温为18-20℃，相对湿度大于90％。每个株系100粒种子，实验重复3次，结果取平均值。
实验进行第8天统计各类种子的萌发率结果如图5、6所示。在200mM甘露醇胁迫处理下，共转PwRBP1与PwNAC1、转PwRBP1、转PwNAC1、转空载体以及野生型拟南芥(Col-0)种子在胁迫处理第1天时均未萌发，但均能在第5天时萌发，共转2种基因的植株萌发率为野生型的2.36倍，单转PwRBP1和单转PwNAC1仅为野生型的1.41和1.18倍，该结果表明在200mM甘露醇胁迫处理下，转PwRBP1与PwNAC1拟南芥的种子萌发率显著高于野生型及单转PwRBP1或PwNAC1植株，具有更好的抵御干旱胁迫的能力。
野生型拟南芥和空载体过表达拟南芥植株(T4代)的结果无显著差异。
2、根长测量耐盐性和耐旱性试验
实验中取共转PwRBP1与PwNAC1拟南芥植株(T4代)、转PwRBP1拟南芥植株(T4代)、转PwNAC1拟南芥植株(T4代)、转空载体拟南芥植株(T4代)以及野生型拟南芥(Col-0)种子进行播种，其培养条件为光照16小时，黑暗8小时，光强为300-400μmol m-2s-1，光照下的室温为22-24℃，相对湿度为70-90％；黑暗条件下的室温为18-20℃，相对湿度大于90％。待种子萌发后将萌发后的拟南芥幼苗，用镊子移到垂直平铺在含有100mM的NaCl或200mM甘露醇的MS培养基上，每个株系100粒种子，实验重复3次，结果取平均值。
实验进行第8天统计各类株系的根长，结果如图7、8所示。在100mM NaCl胁迫处理下，共转两种基因的植株的根长则比野生型对照组提高了46.83％，单转PwRBP1和单转PwNAC1分别提高了43.32％和43.67％；在200mM甘露醇胁迫处理下，共转PwRBP1与PwNAC1拟南芥根长则比野生型对照组提高了44.23％，单转PwRBP1和单转PwNAC1仅分别提高了18.13％和30.50％。该结果表明在100mM NaCl以及200mM甘露醇胁迫处理下，共转PwRBP1与PwNAC1拟南芥幼苗的根长显著长于野生型及单转PwRBP1或PwNAC1植株，具有更好的耐盐和耐旱性。
野生型拟南芥和空载体过表达拟南芥植株的结果无显著差异。
上述结果表明，PwRBP1和PwNAC1能够协同互作提高植物对盐和干旱的耐受能力。
3、幼苗耐逆实验
1)耐盐性研究
实验中取共转PwRBP1与PwNAC1拟南芥植株(T4代)、转PwRBP1拟南芥植株(T4代)、转PwNAC1拟南芥植株(T4代)、转空载体拟南芥植株(T4代)以及野生型拟南芥(Col-0)种子进行播种，其培养条件为光照16小时，黑暗8小时，光强为300-400μmol m-2s-1，光照下的室温为22-24℃，相对湿度为70-90％；黑暗条件下的室温为18-20℃，相对湿度大于90％。待种子萌发后将萌发后的拟南芥幼苗移至培养基质(营养土：蛭石为1:1)中，在培养基质中生长12天后，浇以200mM NaCl溶液，每两天浇一次，11天后统计存活率。每个株系100粒种子，实验重复3次，结果取平均值。
其实验结果如图9、11所示，共转PwRBP1和PwNAC1基因幼苗的存活率比原野生型植株的存活率提高了65.18％，单转PwRBP1和单转PwNAC1仅分别提高了49.33％和53.39％，具有更强的耐盐能力。
野生型拟南芥、和空载体过表达拟南芥植株的结果无显著差异。
2)耐旱性研究
实验中取共转PwRBP1与PwNAC1拟南芥植株(T4代)、转PwRBP1拟南芥植株(T4代)、转PwNAC1拟南芥植株(T4代)、转空载体拟南芥植株(T4代)以及野生型拟南芥(Col-0)种子进行播种，其培养条件为光照16小时，黑暗8小时，光强为300-400μmol m-2s-1，光照下的室温为22-24℃，相对湿度为70-90％；黑暗条件下的室温为18-20℃，相对湿度大于90％。待种子萌发后将萌发后的拟南芥幼苗移至培养基质(营养土：蛭石为1:1)中，在培养基质中生长12天后，停止浇水11天，复水3天后统计存活率。每个株系100粒种子，实验重复3次，结果取平均值。
其实验结果如图10、11所示，共转PwRBP1与PwNAC1基因幼苗的存活率比原野生型植株提高了2倍，单转PwRBP1和单转PwNAC1仅为野生型的1.55和1.43倍，表明共转PwRBP1与PwNAC1拟南芥植株具有更好的耐旱能力。
野生型拟南芥和空载体过表达拟南芥植株(T4代)的结果无显著差异。
上述结果表明，PwRBP1和PwNAC1能够协同互作提高植物对盐和干旱的耐受能力。
序列表
<110> 北京林业大学
<120> PwRBP1和PwNAC1两种蛋白互作协同提高植物耐逆性及其应用
<130> 20190525
<160> 4
<170> PatentIn version 3.3
<210> 1
<211> 1293
<212> DNA
<213> 人工序列
<400> 1
atgatgcaac ccacagcagg cgtcggccct ccttttgcaa accctaatca aaaccagcag 60
cggcagcaat ggcttcaaca gcagcagcag atggcgatgg cgatgcagca gcaacagcag 120
ccgcctcagc aacaggcgaa ccaggccatg gcgatgcagc aacaacaagc cccaatgatg 180
gctcagcagt actatgcaca gcagcctcaa tatcagcagc agcagctacc aatggtgatg 240
cagcaacatc agccgcagtc gagtgacgag gttaagactc tctgggtggg tgatttgcag 300
ttctggatgg acgagggcta tttgcacacc tgtttttccc acactggaga gcttgtttct 360
gccaagataa tccgtaataa gtatactgga cagtcagagg gttatggctt tatggagttc 420
ataacacgta cagctgctga gaagattatg caaacttata atgggacgct aatgcccaac 480
actgaacaag ttttcagaat gaattgggca acttttagca tgggagaaag gcgtctagat 540
ggaggcccag atttttctat ttttgtggga gatttggatt cagatgtctc agatttggtc 600
ttgcaggaga ctttccaaag tcgacattca tcagtgaaag ctgctaaggt tgtcatggat 660
gcaaacacag ggcgctcaaa aggttatggg tttgtgaggt ttggcgagga gagtgagagg 720
gcccgagcca tgacagaaat gaatggtgta tattgttcta ctagacctat gcgaatcagt 780
gcagccaccc caaggaagtc tgcaggggtt cagcaccagt attcaggaag agcaggcaat 840
ggcggatctc atgcccaagg attcccgtca gacaatgatt taaacaatac aactatattt 900
gtaggccggc tagacccaaa tgcgacagat gaagatctga gacaagtctt tggccagtat 960
ggagagcttg tgtctgtaaa aatacctgtt ggtaaaggtt gtggatttgt ccagtttggt 1020
aacagggctt ctgctgagga agccttgcaa aggcttcatg gtactgttat tcgtcagcaa 1080
actgtacgtc tttcttgggg tcgaagccct gcaaacaagc agcaacccca gccccagggg 1140
caacagcctc aatctgatcc aaatcaatgg aatggtgctt actatgggca aggatatgaa 1200
agctatggtt atgctccccc tcctcaagat cctgcaatgt atgcctatgg tggctaccct 1260
ggatatggga attataatca acaggtaagc tag 1293
<210> 2
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Met Met Gln Pro Thr Ala Gly Val Gly Pro Pro Phe Ala Asn Pro Asn
1 5 10 15
Gln Asn Gln Gln Arg Gln Gln Trp Leu Gln Gln Gln Gln Gln Met Ala
20 25 30
Met Ala Met Gln Gln Gln Gln Gln Pro Pro Gln Gln Gln Ala Asn Gln
35 40 45
Ala Met Ala Met Gln Gln Gln Gln Ala Pro Met Met Ala Gln Gln Tyr
50 55 60
Tyr Ala Gln Gln Pro Gln Tyr Gln Gln Gln Gln Leu Pro Met Val Met
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Gln Gln His Gln Pro Gln Ser Ser Asp Glu Val Lys Thr Leu Trp Val
85 90 95
Gly Asp Leu Gln Phe Trp Met Asp Glu Gly Tyr Leu His Thr Cys Phe
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Ser His Thr Gly Glu Leu Val Ser Ala Lys Ile Ile Arg Asn Lys Tyr
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Thr Gly Gln Ser Glu Gly Tyr Gly Phe Met Glu Phe Ile Thr Arg Thr
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Ala Ala Glu Lys Ile Met Gln Thr Tyr Asn Gly Thr Leu Met Pro Asn
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Thr Glu Gln Val Phe Arg Met Asn Trp Ala Thr Phe Ser Met Gly Glu
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Arg Arg Leu Asp Gly Gly Pro Asp Phe Ser Ile Phe Val Gly Asp Leu
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Asp Ser Asp Val Ser Asp Leu Val Leu Gln Glu Thr Phe Gln Ser Arg
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His Ser Ser Val Lys Ala Ala Lys Val Val Met Asp Ala Asn Thr Gly
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Arg Ser Lys Gly Tyr Gly Phe Val Arg Phe Gly Glu Glu Ser Glu Arg
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Ala Arg Ala Met Thr Glu Met Asn Gly Val Tyr Cys Ser Thr Arg Pro
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Met Arg Ile Ser Ala Ala Thr Pro Arg Lys Ser Ala Gly Val Gln His
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Gln Tyr Ser Gly Arg Ala Gly Asn Gly Gly Ser His Ala Gln Gly Phe
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Pro Ser Asp Asn Asp Leu Asn Asn Thr Thr Ile Phe Val Gly Arg Leu
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Asp Pro Asn Ala Thr Asp Glu Asp Leu Arg Gln Val Phe Gly Gln Tyr
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Gly Glu Leu Val Ser Val Lys Ile Pro Val Gly Lys Gly Cys Gly Phe
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Val Gln Phe Gly Asn Arg Ala Ser Ala Glu Glu Ala Leu Gln Arg Leu
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His Gly Thr Val Ile Arg Gln Gln Thr Val Arg Leu Ser Trp Gly Arg
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Ser Pro Ala Asn Lys Gln Gln Pro Gln Pro Gln Gly Gln Gln Pro Gln
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Ser Asp Pro Asn Gln Trp Asn Gly Ala Tyr Tyr Gly Gln Gly Tyr Glu
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Ser Tyr Gly Tyr Ala Pro Pro Pro Gln Asp Pro Ala Met Tyr Ala Tyr
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Gly Gly Tyr Pro Gly Tyr Gly Asn Tyr Asn Gln Gln Val Ser
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<210> 3
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<212> DNA
<213> 人工序列
<400> 1
atggagggat cggaattcca cccgcaggtc ctgcctccag gtttcagatt tcatcccaca 60
gatgaggagc tgctcatcca ttacctgaag aagaaggttt ctgcctctgc tctcccggcg 120
tcaattatcg cagagattga cctgtacaag catgacccct gggatttgcc cgcgaaggca 180
tgttacgggg agcgagaatg gtatttcttc agcccgaggg accggaaata tcccaacgga 240
gcgcggccaa acagatccgc cggctccggc tactggaaag ccactggagc agaaaagccc 300
atcgttgtca tgtcgggaac cacttcgtca cagaaagtgg gcgtgaagaa atccctggtg 360
ttttataaag gaatgccgct taagggcctc aagaccaact ggatcatgca cgaatactgc 420
ctcgctgaaa ccatgccaac gaaaagcaac cgatctctgc gcttggacga ttgggtactg 480
tgccggatat acaagaaagt gagtcattct gccacggcgg tctcgaaccc agaactggag 540
gcaccgtcac ctgctgacgt gcaacaagaa tccgtgatgc ccaaattcag ttccttctct 600
gggctgcttc agagcgacgg tccgtttatg gagagctttc taagccatga tttgtcagat 660
gcctacaaag ctgccctgtc cgatgcggag ccgtcatcca cggtagtccc ccagcagtta 720
aattcctccg agaccagagg ccacctgtca atgtccatgg ctctcaactg cgatgaatta 780
tcctccgtct accctgccga gtggcaaaca gtctga 816
<210> 4
<211> 271
<212> PRT
<213> 人工序列
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Met Glu Gly Ser Glu Phe His Pro Gln Val Leu Pro Pro Gly Phe Arg
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Val Ser Ala Ser Ala Leu Pro Ala Ser Ile Ile Ala Glu Ile Asp Leu
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Arg Glu Trp Tyr Phe Phe Ser Pro Arg Asp Arg Lys Tyr Pro Asn Gly
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Ala Arg Pro Asn Arg Ser Ala Gly Ser Gly Tyr Trp Lys Ala Thr Gly
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Ala Glu Lys Pro Ile Val Val Met Ser Gly Thr Thr Ser Ser Gln Lys
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Val Gly Val Lys Lys Ser Leu Val Phe Tyr Lys Gly Met Pro Leu Lys
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Gly Leu Lys Thr Asn Trp Ile Met His Glu Tyr Cys Leu Ala Glu Thr
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Met Pro Thr Lys Ser Asn Arg Ser Leu Arg Leu Asp Asp Trp Val Leu
145 150 155 160
Cys Arg Ile Tyr Lys Lys Val Ser His Ser Ala Thr Ala Val Ser Asn
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Pro Glu Leu Glu Ala Pro Ser Pro Ala Asp Val Gln Gln Glu Ser Val
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Phe Met Glu Ser Phe Leu Ser His Asp Leu Ser Asp Ala Tyr Lys Ala
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Ala Leu Ser Asp Ala Glu Pro Ser Ser Thr Val Val Pro Gln Gln Leu
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Asn Ser Ser Glu Thr Arg Gly His Leu Ser Met Ser Met Ala Leu Asn
245 250 255
Cys Asp Glu Leu Ser Ser Val Tyr Pro Ala Glu Trp Gln Thr Val
260 265 270
Claims (5)
1.两种能够互作形成同源二聚体的蛋白，是如下a)或b)的蛋白：
a)序列为SEQ ID NO .2和SEQ IDNO .4所示的蛋白；
b)在SEQ ID NO .2和SEQ ID NO .4所示的氨基酸的末端连接标签序列的融合蛋白。
2.与权利要求1所述的两种能够互作形成同源二聚体蛋白相关的生物材料，为下述A1)至A8)中的任一种：
A1)编码权利要求1所述的两种蛋白的核酸分子；
A2)含有A1)所述两种核酸分子的表达盒；
A3)含有A1)所述两种核酸分子的重组载体；
A4)含有A2)所述两种表达盒的重组载体；
A5)含有A1)所述两种核酸分子的重组微生物；
A6)含有A2)所述两种表达盒的重组微生物；
A7)含有A3)所述两种重组载体的重组微生物；
A8)含有A4)所述两种重组载体的重组微生物。
3.根据权利要求2所述的相关的生物材料，其特征在于：A1)所述核酸分子为如下的基因：序列如SEQ ID NO .1和SEQ ID NO .3所示的DNA分子。
4.权利要求1所述的蛋白或权利要求2或3所述的生物材料在共同调控植物耐逆性中的应用；
或，权利要求1所述的蛋白或权利要求2或3所述的生物材料共同在培育耐逆性提高的转基因植物中的应用；
所述耐逆性为耐盐性和/或耐旱性；所述调控为提高；所述植物为双子叶植物。
5.一种培育耐逆性改善的转基因植物的方法，包括同时调节受体植物中权利要求1所述的蛋白的表达量和/或活性，得到转基因植物的步骤；所述改善为提高；所述耐逆性为耐盐性和/或耐旱性，所述植物为双子叶植物。
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| CN107827964A (zh) * | 2017-12-11 | 2018-03-23 | 北京林业大学 | 一种与植物耐逆性相关的转录因子PwNAC2及其编码基因与应用 |
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| CN107827964A (zh) * | 2017-12-11 | 2018-03-23 | 北京林业大学 | 一种与植物耐逆性相关的转录因子PwNAC2及其编码基因与应用 |
Non-Patent Citations (1)
| Title |
|---|
| 青杄PwNAC42基因的克隆及表达模式分析;袁义杭等;《生物技术通报》;20171208(第03期);第113-120页 * |
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