CN116732000A - 一种构树低温胁迫相关的蛋白及其编码基因与应用 - Google Patents
一种构树低温胁迫相关的蛋白及其编码基因与应用 Download PDFInfo
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Abstract
本发明公开了一种构树低温胁迫相关的蛋白及其编码基因与应用。本发明公开的构树低温胁迫相关的蛋白为序列表中SEQ ID No.1所示的蛋白质。实验证明,本发明的构树低温胁迫相关的蛋白及其编码基因可以调控植物的耐冷性:在正常生长条件下,BpPP2C1过表达株系与野生型生长状态没有明显差异,当低温处理后,二者同时出现叶片萎蔫的状态,但重新放回正常生长条件恢复培养3天后,野生型死亡,而BpPP2C1过表达株系则恢复为正常生长状态。本发明的BpPP2C1及其编码基因可以用于提高植物的耐冷性,具有很好的应用前景。
Description
技术领域
本发明涉及生物技术领域中,一种构树低温胁迫相关的蛋白及其编码基因与应用。
背景技术
温度是影响植物生长和农作物产量的最主要的环境因子。利用基因工程手段改良作物的耐低温能力,提高农作物和经济作物对逆境的适应能力是新品种培育亟需解决的关键问题和重大问题。人们从生理、生化、代谢、生态、以及遗传、进化等角度对植物响应干旱等逆境胁迫的机制进行了大量研究,积累了丰富的资料,特别是随着分子生物学的发展,人们能够在基因组成、表达调控及信号传导等分子水平上认识植物对胁迫的耐逆性机理,为利用基因工程手段改良植物的抗胁迫性能开拓了新的途径。由于植物抗逆性状的复杂性,采用传统的育种方法提高植物的抗逆性十分困难,随着分子生物学的发展,基因工程手段开辟了植物抗逆育种的新途径,但高效抗逆基因的分离成为限制植物抗逆基因工程的主要因素。
PP2C蛋白家族是植物磷酸酶家族的重要成员,占比达到60-65%。PP2C是一种多功能单体酶,其活性依赖于Mg2+或Mn2+。脱落酸(ABA)受体(ABARs)将PP2C蛋白置于调节植物应激反应和植物发育的主要信号通路的中心阶段;此外,PP2C蛋白在植物非生物和生物胁迫信号和反应中也发挥重要作用。
发明内容
本发明所要解决的技术问题是如何提高植物的耐冷性。
为解决上述技术问题,本发明首先提供了耐冷相关蛋白或调控所述耐冷相关蛋白含量或活性的物质的下述任一应用:
D1)调控植物耐冷性;
D2)制备调控植物耐冷性产品;
D3)提高植物耐冷性;
D4)制备提高植物耐冷性产品;
D5)培育耐冷植物;
D6)制备培育耐冷植物产品;
所述耐冷相关蛋白来源于构树(Broussonetia papyrifera),其名称为BpPP2C1,BpPP2C1为如下A1)、A2)或A3):
A1)氨基酸序列是SEQ ID No.1的蛋白质;
A2)将序列表中SEQ ID No.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/或添加且具有相同功能的蛋白质;
A3)在A1)或A2)的N端或/和C端连接标签得到的融合蛋白质。
为了使A1)中的蛋白质便于纯化,可在由序列表中SEQ ID No.1所示的氨基酸序列组成的蛋白质的氨基末端或羧基末端连接上如下表所示的标签。
表: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A2)中的蛋白质,为与SEQ ID No.1所示蛋白质的氨基酸序列具有75%或75%以上同一性且具有相同功能的蛋白质。所述具有75%或75%以上同一性为具有75%、具有80%、具有85%、具有90%、具有95%、具有96%、具有97%、具有98%或具有99%的同一性。
上述A2)中的蛋白质可人工合成,也可先合成其编码基因,再进行生物表达得到。
上述A2)中的蛋白质的编码基因可通过将SEQ ID No.2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′端连上表所示的标签的编码序列得到。其中,SEQ ID No.2所示的DNA分子编码SEQ IDNo.1所示的蛋白质。
本发明还提供了与BpPP2C1相关的生物材料的下述任一应用:
D1)调控植物耐冷性;
D2)制备调控植物耐冷性产品;
D3)提高植物耐冷性;
D4)制备提高植物耐冷性产品;
D5)培育耐冷植物;
D6)制备培育耐冷植物产品;
所述生物材料为下述B1)至B7)中的任一种:
B1)编码BpPP2C1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。
上述应用中,B1)所述核酸分子可为如下b11)或b12)或b13)或b14):
b11)编码序列是序列表中SEQ ID No.2的cDNA分子或DNA分子;
b12)序列表中SEQ ID No.2所示的DNA分子;
b13)与b11)或b12)限定的核苷酸序列具有75%或75%以上同一性,且编码BpPP2C1的cDNA分子或基因组DNA分子;
b14)在严格条件下与b11)或b12)或b13)限定的核苷酸序列杂交,且编码BpPP2C1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。
本领域普通技术人员可以很容易地采用已知的方法,例如定向进化和点突变的方法,对本发明的编码BpPP2C1蛋白质的核苷酸序列进行突变。那些经过人工修饰的,具有与本发明分离得到的BpPP2C1蛋白质的核苷酸序列75%或者更高同一性的核苷酸,只要编码BpPP2C1蛋白质且具有BpPP2C1蛋白质功能,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序列1所示的氨基酸序列组成的蛋白质的核苷酸序列具有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。
上述应用中,所述严格条件可为如下:50℃,在7%十二烷基硫酸钠(SDS)、0.5MNaPO4和1mM EDTA的混合溶液中杂交,在50℃,2×SSC,0.1%SDS中漂洗;还可为:50℃,在7%SDS、0.5M NaPO4和1mM EDTA的混合溶液中杂交,在50℃,1×SSC,0.1%SDS中漂洗;还可为:50℃,在7%SDS、0.5M NaPO4和1mM EDTA的混合溶液中杂交,在50℃,0.5×SSC,0.1%SDS中漂洗;还可为:50℃,在7%SDS、0.5M NaPO4和1mM EDTA的混合溶液中杂交,在50℃,0.1×SSC,0.1%SDS中漂洗;还可为:50℃,在7%SDS、0.5M NaPO4和1mM EDTA的混合溶液中杂交,在65℃,0.1×SSC,0.1%SDS中漂洗;也可为:在6×SSC,0.5%SDS的溶液中,在65℃下杂交,然后用2×SSC,0.1%SDS和1×SSC,0.1%SDS各洗膜一次;也可为:2×SSC,0.1%SDS的溶液中,在68℃下杂交并洗膜2次,每次5min,又于0.5×SSC,0.1%SDS的溶液中,在68℃下杂交并洗膜2次,每次15min;也可为:0.1×SSPE(或0.1×SSC)、0.1%SDS的溶液中,65℃条件下杂交并洗膜。
上述75%或75%以上同一性,可为80%、85%、90%或95%以上的同一性。
上述应用中,B2)所述的含有编码BpPP2C1蛋白质的核酸分子的表达盒(BpPP2C1基因表达盒),是指能够在宿主细胞中表达BpPP2C1蛋白质的DNA,该DNA不但可包括启动BpPP2C1基因转录的启动子,还可包括终止BpPP2C1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。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 Physiol 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(均可用茉莉酮酸甲酯诱导);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200710099169.7)),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Genes Dev.,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 AcidRes.,15:9627)。
可用现有的表达载体构建含有所述BpPP2C1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、pCAMBIA1302、pCAMBIA2301、pCAMBIA1301、pCAMBIA1300、pBI121、pCAMBIA1391-Xa、PSN1301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,和赋予对氨甲喋呤抗性的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。
上述应用中,所述载体可为质粒、黏粒、噬菌体或病毒载体。所述质粒具体可为pCAMBIA1300-GFP载体。
B3)所述重组载体具体可为p1300-BpPP2C1。所述p1300-BpPP2C1为在pCAMBIA1300-GFP中插入序列表中SEQ ID No.2所示的DNA分子得到的能够表达BpPP2C1融合蛋白的重组载体。
上述应用中,所述微生物可为酵母、细菌、藻或真菌。其中,细菌可为农杆菌,如农杆菌EHA105。
上述应用中,所述转基因植物细胞系、转基因植物组织和转基因植物器官均不包括繁殖材料。
本发明还提供了下述任一方法:
X1)培育耐冷植物的方法,包括使受体植物中表达BpPP2C1,或提高受体植物中BpPP2C1的含量,或提高受体植物中BpPP2C1的活性,得到与所述受体植物相比耐冷性增强的目的植物;
X2)提高植物耐冷性的方法,包括使受体植物中表达BpPP2C1,或提高受体植物中BpPP2C1的含量,或提高受体植物中BpPP2C1的活性,得到与所述受体植物相比耐冷性增强的目的植物,实现植物耐冷性的提高。
上述方法可通过向所述受体植物中导入BpPP2C1的编码基因并使所述编码基因得到表达实现。
上述方法中,所述编码基因可为B1)所述核酸分子。
上述方法中,其中所述BpPP2C1的编码基因可先进行如下修饰,再导入受体植物中,以达到更好的表达效果:
1)根据实际需要进行修饰和优化,以使基因高效表达;例如,可根据受体植物所偏爱的密码子,在保持本发明所述BpPP2C1的编码基因的氨基酸序列的同时改变其密码子以符合植物偏爱性;优化过程中,最好能使优化后的编码序列中保持一定的GC含量,以最好地实现植物中导入基因的高水平表达,其中GC含量可为35%、多于45%、多于50%或多于约60%;
2)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;
3)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;
4)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;
5)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BpPP2C1的编码基因可利用含有所述BpPP2C1的编码基因的重组表达载体导入受体植物。所述重组表达载体具体可为所述p1300-BpPP2C1。
所述重组表达载体可通过使用Ti质粒,植物病毒载体,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 Plant MolecularBiology VIII,Academy Press,New York,pp.411-463;Geiserson and Corey,1998,PlantMolecular Biology(2nd Edition).)。
所述目的植物理解为不仅包含BpPP2C1蛋白或其编码基因被改变的第一代植物,也包括其子代。对于所述目的植物,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。所述目的植物包括种子、愈伤组织、完整植株和细胞。
本发明还提供了具有耐冷性的产品,所述产品含有BpPP2C1或所述生物材料。
所述产品可以BpPP2C1或所述生物材料作为其活性成分,还可将BpPP2C1或所述生物材料与其他具有相同功能的物质一起作为其活性成分。
上文中,所述植物可为M1)或M2)或M3)或M4)或M5):
M1)双子叶植物或单子叶植物;
M2)荨麻目植物;
M3)桑科植物;
M4)构属植物;
M5)构树。
BpPP2C1或所述生物材料,也属于本发明的保护范围。
实验证明,本发明的BpPP2C1及其编码基因可以调控植物的耐冷性:在正常生长条件下,BpPP2C1过表达株系与野生型生长状态没有明显差异,当低温处理后,二者同时出现叶片萎蔫的状态,但重新放回正常生长条件恢复培养3天后,野生型死亡,而BpPP2C1过表达株系则恢复为正常生长状态;低温处理前,BpPP2C1过表达植株与野生型构树植株各生理指标差别变化不大,但低温处理后,POD、SOD活性以及Fv/Fm的比值在BpPP2C1过表达植株中均明显高于野生型构树,而超氧阴离子含量、相对电导率以及MDA含量则明显低于野生型构树。本发明的BpPP2C1及其编码基因可以用于提高植物的耐冷性,具有很好的应用前景。
附图说明
图1为PCR扩增BpPP2C1全长cDNA的琼脂糖凝胶电泳检测结果。
图2为BpPP2C1基因表达模式分析结果。
图3为BpPP2C1基因的亚细胞定位结果。A中上图为p1300-BpPP2C1的结构特征,下图为pCAMBIA1300-GFP的结构特征。BpPP2C1-GFP表示p1300-BpPP2C1,35S::GFP表示pCAMBIA1300-GFP;Chloroplast:叶绿体;Bright:明场;Merged:合并。
图4为BpPP2C1基因过表达载体转化构树阳性植株结果。B中,M为DNA分子量标准,L1、L2、L3为3个BpPP2C1过表达株系,(-)表示去离子水,P表示p1300-BpPP2C1,WT-1、WT-2表示野生型构树。
图5为BpPP2C1基因过表达植株在低温处理后与野生型构树之间的表型差异。A:不同状态下的植株照片;B:植株叶绿素荧光变化图。
图6为BpPP2C1基因过表达植株在低温处理后与野生型构树相比生理指标测定的结果。A-E中每图从左至右依次为WT、L1、L2、L3,25℃为冷处理前的结果,4℃为冷处理后恢复培养3天后的结果。*表示差异达到显著水平,p<0.05。F中0、2、4、6、8、10d分别为低温处理后第0、2、4、6、8、10天的结果。
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,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、仪器等,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,如无特殊说明,序列表中各核苷酸序列的第1位均为相应DNA/RNA的5′末端核苷酸,末位均为相应DNA/RNA的3′末端核苷酸。
下述实施例中的pCAMBIA1300-GFP(Tang F,Zhang D,Chen N,Peng X,ShenS.Genome-Wide Analysis of BpYABs and Function Identification Involving in theLeaf and Silique Development in Transgenic Arabidopsis.International Journalof Molecular Sciences.2022;23(3):1670.),公众可从申请人处获得该生物材料,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、构树低温胁迫特异表达基因BpPP2C1全长cDNA序列的获得
一、构树低温胁迫特异表达基因BpPP2C1序列的克隆
1、植物材料处理及总RNA的提取
以构树(Broussonetia papyrifera)幼苗为材料提取其总RNA,进行1%琼脂糖凝胶电泳检测,并利用Scan Drop测定RNA浓度及完整性,结果表明所提RNA质量完好,浓度适宜。
2、构树低温胁迫特异表达基因BpPP2C1全长cDNA克隆
以上述步骤1提取的构树幼苗的总RNA为模板,用PrimeScriptTM 1st Strand cDNASynthesized Kit试剂盒(Takara公司)并参照试剂盒说明书的要求,反转合成其第一链cDNA。将合成的第一链cDNA贮存于-20℃备用。
再以获得的第一链cDNA为模板,参照构树已测序基因组序列中P1959同源基因Bp01g0320的CDS序列设计引物,引物1959-F:5′-ATGGGTTGTGTGTACTCGAGGGTTT-3′与引物1959-R:5′-CTACCAGTCATCTAGCCATTCGGGA-3′配对进行PCR扩增。
PCR反应结束后,对PCR扩增产物进行1%琼脂糖凝胶电泳检测,结果如图1所示。其中,泳道M为TRANS2000 DNA分子量标准(北京全式金生物技术有限公司)的DNA分子量标准,泳道BpPP2C1为PCR扩增产物,结果表明,经PCR扩增获得了长度约为3300bp的目的片段。回收并纯化扩增产物,将其连接到Blunt Zero载体上(CB501-01,北京全式金科技有限公司),连接产物转化大肠杆菌DH5α感受态细胞,筛选阳性克隆进行菌液PCR鉴定,提取阳性克隆的质粒进行测序,对测序结果进行比对分析。结果表明,P1959该片段的长度为3234bp,其脱氧核糖核苷酸序列如序列表中SEQ ID No.2所示,编码由1077个氨基酸残基组成的蛋白质(序列为序列表中SEQ ID No.1),与构树中Bp01g0320基因具有较高的同源性,表明该片段为构树P1959基因序列,将该基因记为BpPP2C1基因。
实施例2、BpPP2C1表达模式分析
一、BpPP2C1在构树不同组织中的表达模式分析
取正常生长8周的构树幼苗的根、茎和叶三个组织的样品,分别提取总RNA,利用荧光定量PCR的方法分析BpPP2C1基因在不同组织中表达模式的变化。
用BpGADPH基因作为反应的内参,引物如下:
BpGADPH-F:5′-CCGTGCTCAATGGGATACTTC-3′,
BpGADPH-R:5′-CCCTCGTCTGTGACAATGGTAC-3′。
BpPP2C1基因引物如下:
QF:5′-CACGGAGAAGATGCAAAGAA-3′;QR:5′-GATTAGCAGCAGCAAGGGAG-3′。
所得数据及Ct值的分析用Mx3000p软件进行。结果如图2中A所示。结果表明,BpPP2C1基因在叶中表达量高于茎,而在茎中的表达量又明显高于根。
二、构树BpPP2C1基因在低温处理后表达模式分析
将正常生长8周的构树幼苗分别进行不同时间(0、1、3、6和12小时)的低温(4℃)胁迫处理后,分别提取上述不同处理的构树幼苗的总RNA,利用荧光定量PCR方法分析BpPP2C1基因在低温胁迫条件下的表达模式变化。所用引物同步骤一。
结果如图2中B所示。结果表明,BpPP2C1在低温处理1小时后,表达量开始缓慢上升,在处理6小时达到最高为未处理的2.5倍,在12小时的时候开始下降。说明,BpPP2C1基因在低温胁迫后表达量有所升高,但并不明显。
实施例3、BpPP2C1蛋白体内功能分析
一、BpPP2C1的亚细胞定位
将上述实施例1扩增得到的BpPP2C1基因以及植物双元表达载体pCAMBIA1300-GFP利用碱基合成的方法获得重组表达载体命名为p1300-BpPP2C1。p1300-BpPP2C1能表达序列表中SEQ ID No.2所示的BpPP2C1与GFP的融合蛋白,BpPP2C1基因的表达由35S驱动,SEQ IDNo.1所示的BpPP2C1基因位于p1300-BpPP2C1的Xba Ⅰ和Sal Ⅰ识别序列间。载体构建情况如图3中A。
将1μg上述重组表达载体导入到农杆菌EHA105菌株中,转化烟草叶片表皮细胞瞬时表达,同时以转入空载体pCAMBIA1300-GFP的烟草表皮细胞作为对照。基因转化后的烟草叶片表皮细胞培养24-48小时后,在激光共聚焦扫描显微镜(Bio-Rad MRC 1024)下观察并照相,结果如图3中B所示。从中可以看出,转入空载体pCAMBIA1300-GFP的转基因细胞中表达的蛋白分布在整个细胞内,而转入重组表达载体p1300-BpPP2C1的转基因细胞中表达的蛋白则定位于细胞质膜上。同时转入定位于质膜的标记基因PIP2-mCherry载体与p1300-BpPP2C1载体共表达,结果如图3中C。结果表明,BpPP2C1定位于质膜上,与PIP2共表达。
二、BpPP2C1基因的过表达植株构建
将步骤二中构建好的p1300-BpPP2C1导入农杆菌LBA4404菌株中,得到重组菌LBA4404/p1300-BpPP2C1。将LBA4404/p1300-BpPP2C1利用圆盘法侵染构树叶片,经过抗性筛选等过程,获得阳性芽。阳性芽经过DNA、RNA和蛋白质水平鉴定后,确认有BpPP2C1-GFP融合蛋白表达的株系被认定为BpPP2C1过表达植株,结果如图4。结果表明,经DNA、RNA和蛋白质水平的鉴定,共得到3个BpPP2C1过表达株系,记为L1、L2、L3。
DNA水平鉴定:所用引物为P1(5’-CCAAGATAGCAAAGGGACAG-3’)与P2(5’-GGTGCAGATGAACTTCAGGGT-3’)、P3(5’-TCGTTATGTTTATCGGCACTTTG-3’)与P4(5’-TGTTGGCGACCTCGTATTGG-3’),其在载体中的位置如图4中A所示。利用p1300-BpPP2C1作为阳性对照,利用去离子水和野生型构树作为阴性对照。结果见图4中B,L1、L2、L3均含有目的DNA片段。
RNA水平鉴定:所用引物为F:5’-ATGGGTTGTGTGTACTCGAGGG-3’,R:5’-CTACCAGTCATCTAGCCATTCG-3’。利用Actin基因作为内参。结果见图4中C,L1、L2、L3中BpPP2C1基因的表达量均显著高于野生型构树(WT)。
蛋白质水平鉴定:利用Actin基因作为内参,利用野生型构树(WT)作为对照,所用抗体为GFP抗体(Anti-GFP)(柏奥易杰(北京)科技有限公司)、Actin抗体(Anti-Actin)(柏奥易杰(北京)科技有限公司。结果见图4中D,L1、L2、L3中含有目的融合蛋白,野生型构树中不含有目的融合蛋白。
三、BpPP2C1过表达植株低温表型分析
将步骤二所得BpPP2C1过表达株系L1、L2、L3与野生型构树(WT)的生根苗在正常生长状态(25℃)下,培养13天后,移入-4℃冷处理24小时后,恢复正常培养3天,观察记录表型数据,同时测定其Fv/Fm变化情况,结果如图5。
结果表明,在正常生长条件下,BpPP2C1过表达株系与野生型构树生长状态没有明显差异,当低温处理后,二者同时出现叶片萎蔫的状态,但重新放回正常生长条件恢复培养3天后,野生型构树死亡,而BpPP2C1过表达株系则恢复为正常生长状态,其Fv/Fm值也印证了这一点(图6中F)。这说明,过表达BpPP2C1可以提高植株的耐低温能力。
为了检测BpPP2C1过表达植株的耐低稳能力,发明人还测定了一系列的生理生化指标,包括过氧化物酶(POD)活性,超氧化物歧化酶(Superoxide dismutase,SOD)活性,超氧阴离子含量(O2 -含量),相对电导率(REL),丙二醛(MDA)含量,结果如图6所示。
结果表明,低温处理前,BpPP2C1过表达植株与野生型构树植株各生理指标差别变化不大,但低温处理后,POD、SOD活性以及Fv/Fm的比值在BpPP2C1过表达植株中均明显高于野生型构树,而超氧阴离子含量、相对电导率以及MDA含量则明显低于野生型构树,证明BpPP2C1过表达株系确实对低温有较高的耐受力。
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> 一种构树低温胁迫相关的蛋白及其编码基因与应用
<160> 2
<170> PatentIn version 3.5
<210> 1
<211> 1077
<212> PRT
<213> 构树(Broussonetia papyrifera)
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atgggttgtg tgtactcgag ggtttgtatc ggagaggtct gtactccgag ggacacgaga 60
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acaatttacg tcgcgaattc gggcgattcg agggcggtga tcgccgagaa gagaggggat 660
gaatttgttg ctgtcgacct ctccattgac cagactccgt ttagggaaga tgagctcgaa 720
agggtcaagc tttgcggcgc cagagttctc acgttggatc agattgaagg gctgaagaac 780
ccggatgtgc agtgttgggg cactgaggaa ggagatgatg gtgacccacc tagattgtgg 840
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atttctaaga accaacgtgt tagacatgat ttatcaaggg cacgtattcg tgctattgaa 1320
agttctcttg agaatggaca agtttgggtt cctccatctc cagcccatag aaaaacttgg 1380
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acggctgaaa agctatcatc ctttggagaa ctggcactga tgtataacaa accactccag 1680
gcctccgttc gtgccgtaac aaatggaact ctatgggctt taagaagaga agactttcgt 1740
ggaatactca tgtcagagtt ttctaatttg tcatccttga agttactccg atcggtagat 1800
cttctttcaa ggttgacaat cttacagctg agtcacattg cagattcttt atctgaggtt 1860
tccttctcag atgggcagac aatagtgaaa aagaatgaag ccctctttgc attatacatt 1920
attcagaagg gacgagtgag gattacttat aaagcagatt tagttggtcc aaatgtttcc 1980
agtcttaatt ctgacaatca aaaagaaggt gacaatcctc caggcagtaa cgagcttaca 2040
gtggagaaga cagagggaag ctattttggt gaatggacgc ttcttggtga acatattgat 2100
tccataagtg cagttgctgt aggtgacgtc gtttgttcct ttttaacaaa ggaaaaattt 2160
gagtcagttg ttggcccttt gcgaaagctt tctcaggatg atcagaagtc aagagatcat 2220
tctttagatt tctccaaaga atctgctaaa aatattgata tttcaacact ttccgaagtt 2280
cagttctctg atatggaatg gaagaaatgt ttatattcta ccgattgcag tgaaattggg 2340
ctcgtacttt tgagagattc agaaaatttg cttagtttga aaaggttttc aaagcagaag 2400
gtcaagaggc tgggaaagga ggcacaggtc ttgaaagaaa agaacttgat gaaaagtatc 2460
agccattcag cccatgtgcc aaaggtcctg tctacttgtg ttgatcagtc acatgccggc 2520
atattgcttg acacatgtct tgcttgtcct ttggcctcta tacttcacac cccactggat 2580
gagctgtctg cacaattttg tgcagcttgt gttgttaatg ccttggaaca tttacacaag 2640
aacgatatcc tatacagagg cgtgtcccat gatgttctga tgttgaacca gacgggatat 2700
ttacaggtgg tggacttcag atttggaaaa agattatctg gtgagaggac atatacaatt 2760
tgtggaatgg cagacttttt agctccagaa atagtccagg gaaaaggcca cggcttcaca 2820
gctgactggt gggcattggg agtcttgatc tatttcatgc taaaaggtga aatgccattt 2880
ggttcatgga gagaaagtga actcgataca tttgccaaga tagcaaaggg acagctaaat 2940
cttcctgcaa atttcagccc tgaagttgtg gatctcatca ccaagttact tgacgttgac 3000
gaacaaacta gactgggaaa cacgggtccc aattctatca gaagtcaccc ttggtttgat 3060
ggtattgatt ggaaagcgat agagaaccat agttttcccg ttcctaatga gatcacttcc 3120
cggattgctc aacacttgga ggtgaactcc gaggactcta cttttccgcg cctctctctc 3180
tcccaagacg tcgaagaaga tgacgacgtt cccgaatggc tagatgactg gtag 3234
Claims (9)
1.耐冷相关蛋白或调控所述耐冷相关蛋白含量或活性的物质的下述任一应用:
D1)调控植物耐冷性;
D2)制备调控植物耐冷性产品;
D3)提高植物耐冷性;
D4)制备提高植物耐冷性产品;
D5)培育耐冷植物;
D6)制备培育耐冷植物产品;
所述耐冷相关蛋白为如下A1)、A2)或A3):
A1)氨基酸序列是SEQ ID No.1的蛋白质;
A2)将序列表中SEQ ID No.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/或添加且具有相同功能的蛋白质;
A3)在A1)或A2)的N端或/和C端连接标签得到的融合蛋白质。
2.与权利要求1中所述耐冷相关蛋白相关的生物材料的下述任一应用:
D1)调控植物耐冷性;
D2)制备调控植物耐冷性产品;
D3)提高植物耐冷性;
D4)制备提高植物耐冷性产品;
D5)培育耐冷植物;
D6)制备培育耐冷植物产品;
所述生物材料为下述B1)至B7)中的任一种:
B1)编码权利要求1中所述耐冷相关蛋白的核酸分子;
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。
3.根据权利要求2所述的应用,其特征在于:B1)所述核酸分子为如下b11)或b12)或b13)或b14):
b11)编码序列是序列表中SEQ ID No.2的cDNA分子或DNA分子;
b12)序列表中SEQ ID No.2所示的DNA分子;
b13)与b11)或b12)限定的核苷酸序列具有75%或75%以上同一性,且编码权利要求1中所述耐冷相关蛋白的cDNA分子或基因组DNA分子;
b14)在严格条件下与b11)或b12)或b13)限定的核苷酸序列杂交,且编码权利要求1中所述耐冷相关蛋白的cDNA分子或基因组DNA分子。
4.下述任一方法:
X1)培育耐冷植物的方法,包括使受体植物中表达权利要求1中所述耐冷相关蛋白,或提高受体植物中权利要求1中所述耐冷相关蛋白的含量,或提高受体植物中权利要求1中所述耐冷相关蛋白的活性,得到与所述受体植物相比耐冷性增强的目的植物;
X2)提高植物耐冷性的方法,包括使受体植物中表达权利要求1中所述耐冷相关蛋白,或提高受体植物中权利要求1中所述耐冷相关蛋白的含量,或提高受体植物中权利要求1中所述耐冷相关蛋白的活性,得到与所述受体植物相比耐冷性增强的目的植物,实现植物耐冷性的提高。
5.根据权利要求4所述的方法,其特征在于:所述方法通过向所述受体植物中导入权利要求1中所述耐冷相关蛋白的编码基因并使所述编码基因得到表达实现。
6.根据权利要求5所述的方法,其特征在于:所述编码基因为权利要求2或3中B1)所述核酸分子。
7.具有耐冷性的产品,含有权利要求1中所述耐冷相关蛋白或权利要求2或3中所述生物材料。
8.根据权利要求1-3中任一所述的应用,或权利要求4-6中任一所述的方法,或权利要求7所述的产品,其特征在于:所述植物为M1)或M2)或M3)或M4)或M5):
M1)双子叶植物或单子叶植物;
M2)荨麻目植物;
M3)桑科植物;
M4)构属植物;
M5)构树。
9.权利要求1中所述耐冷相关蛋白或权利要求2或3中所述生物材料。
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