CN102234327B - 植物耐盐相关蛋白AtST1及其编码基因与应用 - Google Patents
植物耐盐相关蛋白AtST1及其编码基因与应用 Download PDFInfo
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Abstract
本发明公开了植物耐盐相关蛋白AtST1及其编码基因与应用。本发明所提供的植物耐盐相关的蛋白,名称为AtST1,是如下1)或2)的蛋白质:1)由序列表中序列2所示的氨基酸序列组成的蛋白质;2)将序列表中序列2的氨基酸残基序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且与植物耐盐相关的由1)衍生的蛋白质。本发明提供的蛋白的编码基因AtST1受高盐诱导表达。而将AtST1基因导入野生型拟南芥中得到的转基因植物的耐高盐性、耐高渗性和耐ABA性均得到提高。本发明的耐盐基因对于植物耐高盐分子机制的研究,植物抗高盐品种的选育具有重要的理论及实际意义。
Description
技术领域
本发明涉及植物耐盐相关蛋白及其编码基因与应用。
背景技术
粮食问题是当今世界所面临的几大难题之一。土壤盐渍化是限制农业生产的一个重要限制因素。盐渍土是一种世界性的低产土壤,甚至是不毛之地。因此,认识植物对高盐胁迫的反应机制,提高植物自身对高盐胁迫的耐受能力,已成为如何进一步提高作物产量的重要研究工作,倍受世界各国政府和植物及农业科学家的关注。
在长期的进化过程中,高等植物逐渐演变产生了一套感受和传导高盐信号的系统,并形成一系列生理或发育的机制来响应环境中的高盐胁迫,最大限度地减轻高盐胁迫造成的伤害。利用现代分子生物学、植物生理学等技术手段研究植物抗高盐的分子机理,发现和挖掘抗盐基因,使通过基因工程技术改造植物,提高植物自身的抗盐能力成为可能。植物抗盐的分子机理研究及抗盐品种的培育已成为当前研究的热点。
拟南芥(Arabidopsis thaliana)是一种典型的双子叶模式植物,具有个体小、生长周期短、遗传背景简单清楚、易被诱变等特点,已成为植物科学研究的模式材料。广泛应用于植物遗传学、发育生物学和分子生物学的研究,拟南芥的大多数基因在其他植物中都能找到同源基因,在拟南芥研究中发现对植物生长发育具有重要作用的基因也可直接应用到其它植物研究中。
拟南芥基因组大小为125Mbp,约2.7万个基因,目前仍然还有许多基因的功能不十分清楚。对拟南芥中参与植物抗盐反应的基因的功能研究,对于进一步阐明植物抗盐的分子机理并通过基因工程的技术和手段培育优质、抗盐的作物新品种具有重要的理论意义和现实意义。
发明内容
本发明的目的是提供一种植物耐盐相关的蛋白及其编码基因。
本发明所提供的植物耐盐相关的蛋白,名称为AtST1,来源于拟南芥(Arabidopsis thaliana)Col-0生态型,是如下1)或2)的蛋白质:
1)由序列表中序列2所示的氨基酸序列组成的蛋白质;
2)将序列表中序列2的氨基酸残基序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且与植物耐盐相关的由1)衍生的蛋白质。
为了使1)中的AtST1便于纯化,可在由序列表中序列2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表1所示的标签。
表1.标签的序列
标签 | 残基 | 序列 |
Poly-Arg | 5-6(通常为5个) | RRRRR |
PoN-His | 2-10(通常为6个) | HHHHHH |
FLAG | 8 | DYKDDDDK |
Strep-tag II | 8 | WSHPQFEK |
c-myc | 10 | EQKLISEEDL |
上述2)中的AtST1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2)中的AtST1的编码基因可通过将序列表中序列1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。
上述植物耐盐相关蛋白的编码基因(命名为AtST1)也属于本发明的保护范围。
植物耐盐相关蛋白的编码基因具体可为如下1)-3)中任一所述的基因:
1)其编码序列是序列表中序列1;
2)在严格条件下与1)的基因杂交且编码所述蛋白的基因;
3)与1)的基因具有90%以上的同源性且编码所述蛋白的基因。
序列表中的序列1由1035个碱基组成,其开放阅读框架(ORF)为自5′末端第1-1035位碱基,编码具有序列表中序列2的氨基酸序列的AtST1蛋白。
上述严格条件可为用0.1×SSPE(或0.1×SSC),0.1%SDS的溶液,在DNA或者RNA杂交实验中65℃下杂交并洗膜。
扩增上述AtST1基因全长或其任一片段的引物对也属于本发明的保护范围。
含有上述植物耐盐相关蛋白的编码基因的表达盒、重组载体、转基因细胞系和重组菌也属于本发明的保护范围。
可用现有的植物表达载体构建含有AtST1基因的重组表达载体。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pBIB、pCAMBIA3301、pCAMBIA1300、pBI121、pBin19、pCAMBIA2301、pCAMBIA1301-UbiN或其它衍生植物表达载体。
使用AtST1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)、Super启动子等,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、萤光素酶基因等)、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。
所述重组载体具体为在pCAMBIA1300:Super的多克隆位点间插入上述基因得到的重组表达载体;所述pCAMBIA1300:Super是将核苷酸序列如序列表中序列3的自5’端第113-1112位所示的Super启动子插入pCAMBIA1300的HindIII和XbaI酶切位点间得到的载体。
本发明的另一个目的是提供一种培育转基因植物的方法。
本发明所提供的培育转基因植物的方法,是将上述基因AtST1导入目的植物中,得到转基因植物,所述转基因植物的耐盐性、耐渗透性和耐ABA性均强于所述目的植物。
上述耐盐可以是耐120mM NaCl的高盐。
上述耐渗透可以是耐300mM Sorbitol引起的高渗。
上述植物基因AtST1是通过上述重组表达载体导入目的植物中的。
携带有本发明的基因AtST1的植物表达载体可通过Ti质粒、Ri质粒、植物病毒载体、直接DNA转化、显微注射、电导、农杆菌介导等常规生物学方法转化到植物细胞或组织中。
上述目的植物可以是双子叶植物或单子叶植物,如拟南芥。
实验证明:在高盐条件下,AtST1基因表达明显升高,在高盐处理3天时,AtST1基因的表达达到最高,然后AtST1基因的表达逐步回到处理前水平,说明AtST1基因受高盐诱导表达。而将AtST1基因导入野生型拟南芥中得到的转基因植物的耐高盐性、耐高渗性和耐ABA性均得到提高。本发明的耐盐基因对于植物耐高盐分子机制的研究,植物抗高盐品种的选育具有重要的理论及实际意义。
附图说明
图1为AtST1过量表达材料Super:AtST1-1、Super:AtST1-2和突变体atst1的RP-PCR鉴定结果。
图2为高盐条件下AtST1过量表达材料(Super:AtST1-1和Super:AtST1-2)和突变体atst1的表型分析结果。
图3为高渗条件下AtST1过量表达材料(Super:AtST1-1和Super:AtST1-2)和突变体atst1的表型分析结果。
图4为外加ABA条件下AtST1过量表达材料(Super:AtST1-1和Super:AtST1-2)和突变体atst1的表型分析结果。
图5为高盐条件下AtST1基因的表达变化分析结果。
具体实施方式
下面结合具体实施例对本发明作进一步说明,但本发明并不限于以下实施例。
下述实施例中,如无特殊说明,均为常规方法。
实施例1、植物响应高盐胁迫的蛋白及其编码基因的获得
提取10天拟南芥(Columbia型又称Col-0生态型)(购自美国拟南芥生物资源中心(Arabidopsis Biological Resource Center,ABRC))幼苗的总RNA、用Takara公司的DNA酶消化RNA中的DNA,用消化后产物进行反转录得到cDNA。以此cDNA为模板、利用分别设有XbaI和KpnI酶切位点的一对引物进行PCR扩增,得到耐盐相关基因的cDNA全长。引物序列如下:
Primer1:5′-tctaga ATGGAATCGAGTAGCGTTGATGAG-3′(XbaI);
Primer2:5′-ggtacc TTACGAGGCGTGAAAGATGCGTTGC-3′(KpnI)。
采用Takara公司的高保真的Pyrobest DNA聚合酶进行扩增,扩增体系为50μL,含10×Pyrobest PCR缓冲液5.0μL,10mM dNTP mix 1.0μL,引物各1.0μL,Pyrobest酶0.5μL,模板3.0μL(0.1μg),补充灭菌超纯水至50μL,反应体系在冰上进行操作。在PE 9700仪器上进行扩增,预变性5min,95℃变性30s,58℃30s,72℃1.5min,循环数30个,72℃延伸5min。
将扩增片段用0.8%的琼脂糖凝胶进行电泳,回收扩增片段,连接到pMD18-T载体,酶切、扩增、测序鉴定,测序结果表明,上述PCR扩增得到的片段具有序列表中序列1的核苷酸序列,为本发明的调控植物响应高盐胁迫cDNA基因命名为AtST1,序列表中序列1的核苷酸序列由1035个碱基组成,其编码序列为自序列表中序列1的第1-1035位核苷酸,编码具有序列表中序列2的氨基酸残基序列。将上述获得的正确的含有AtST1的重组载体命名为pMD18-AtST1。
提取10天拟南芥(Columbia型)幼苗的总DNA,以此DNA为模板、利用一对引物进行PCR扩增,得到耐盐相关基因的基因组DNA全长。引物序列如下:
Primer1:5′-tctaga ATGGAATCGAGTAGCGTTGATGAG-3′(XbaI);
Primer2:5′-ggtacc TTACGAGGCGTGAAAGATGCGTTGC-3′(KpnI)。
采用Takara公司的高保真的Pyrobest DNA聚合酶进行扩增,扩增体系为50μL,含10×Pyrobest PCR缓冲液5.0μL,10mM dNTP mix 1.0μL,引物各1.0μL,Pyrobest酶0.5μL,模板3.0μL(0.1μg),补充灭菌超纯水至50μL,反应体系在冰上进行操作。在PE 9700仪器上进行扩增,预变性5min,95℃变性30s,56℃30s,72℃1.5min,循环数30个,72℃延伸5min。
对PCR产物进行测序,得到1035bp的核苷酸片段,将其命名为gAtST1。将gAtST1连接到pMD18-T载体得到的重组载体命名为pMD18-gAtST1。将gAtST1序列和序列1进行序列比对,发现gAtST1序列和序列1一致,说明gAtST1序列中没有内含子。
实施例2、植物响应高盐胁迫的蛋白及其编码基因的功能验证
一、AtST1过量表达植株(Super:AtST1-1和Super:AtST1-2)的获得
1、中间载体pCAMBIA1300:Super的构建
利用引物aagcttGTGGGCCTGTGGTCTCAAGAT和tctagaCTAGAGTCGATTTGGT从质粒pMSP-1(GenBank号为EU181145)中扩增出核苷酸序列如序列表中序列3的自5’端第113-1112位所示的Super启动子的Super启动子序列;将扩增得到的Super启动子用HindIII和XbaI双酶切,回收Super启动子片段,插入到质粒pCAMBIA1300(GenBank号为FJ362601)的HindIII和XbaI酶切位点之间中,得到pCAMBIA1300:Super质粒。
2、Super:AtST1的构建
将实施例1所获得的含有AtST1基因的重组载体pMD18-AtST1和上述步骤1获得的中间载体pCAMBIA1300:Super,同时用内切酶XbaI和KpnI进行大体系酶切。将所切得的AtST1基因片段和线性化的pCAMBIA1300:Super质粒片段进行回收,连接,并测序验证,即获得验证正确的含有AtST1基因的重组载体并将其命名为Super:AtST1。
将Super:AtST1利用农杆菌转化侵染Columbia野生型拟南芥植株。从而获得AtST1过量表达T 1代转Super:AtST1拟南芥植株。经过在含50μg/L潮霉素的MS培养基上筛选转Super:AtST1拟南芥阳性植株并进行分离比统计,在T 3代即得到转Super:AtST1拟南芥单拷贝纯合株系,即AtST1过量表达株系Super:AtST1-1和Super:AtST1-2。
将AtST1基因转化pBI121质粒,筛选得到反向插入的antisense-AtST1-pBI121质粒。将antisense-AtST1-pBI121质粒利用农杆菌转化侵染Columbia野生型拟南芥植株,获得AtST1的突变体atst1。
将野生型拟南芥(Columbia型)、AtST1过量表达植株(Super:AtST1拟南芥单拷贝纯合株系,名称为Super:AtST1-1和Super:AtST1-2)和AtST1突变体(atst1)拟南芥种子在MS培养基萌发生长7天,培养条件采用全日照光照培养,温度22℃,光强100μmol/(m2*s)。正常MS培养基组成为:每升溶液含1650mg NH4NO3,1900mgKNO3,370mg MgSO4·7H2O,170mg KH2PO4,440mg CaCl2·2H2O,22.3mgMnSO4·4H2O,0.83mg KI,0.025mg CuSO4·5H2O,6.25mg H3BO5,0.025mgCoCl·6H2O,8.65mg ZnSO4·7H2O,0.25mg Na2MoO4·2H2O,27.8mg FeSO4·7H2O,37.3mg Na2EDTA。植物生长7天后取样,提取总RNA,总RNA的提取按Invitrogen TrizolReagent的产品说明书进行。将总RNA用DNA酶进行消化,消化后的总RNA进行反转录,得到cDNA。以cDNA为模板,进行PCR扩增。引物序列如下:Primer1:5′-tctagaATGGAATCGAGTAGCGTTGATGAG-3′(XbaI);Primer2:5′-ggtaccTTACGAGGCGTGAAAGATGCGTTGC-3′(KpnI)。
结果如图1所示,在Super:AtST1拟南芥单拷贝纯合株系Super:AtST1-1和Super:AtST1-2中AtST1基因的表达水平明显高于野生型,而AtST1突变体atst1中AtST1基因的表达水平明显低于野生型。
二、野生型、AtST1过量表达植株(Super:AtST1-1和Super:AtST1-2)和AtST1突变体(atst1)在高盐、高渗和外加ABA条件下的表型观察
1、野生型、AtST1过量表达植株(Super:AtST1-1和Super:AtST1-2)和AtST1突变体(atst1)(Salk)在高盐条件下的表型观察结果
将野生型拟南芥(Columbia型)、AtST1过量表达植株(步骤一获得的转Super:AtST1拟南芥单拷贝纯合株系,名称为Super:AtST1-1和Super:AtST1-2)和AtST1突变体(atst1)拟南芥种子在含有120mM NaCl的高盐MS培养基(以正常MS培养基为基础,加入120mM NaCl,其它成分与正常MS培养基一致。正常MS培养基组成为:每升溶液含1650mgNH4NO3,1900mg KNO3,370mg MgSO4·7H2O,170mg KH2PO4,440mg CaCl2·2H2O,22.3mg MnSO4·4H2O,0.83mg KI,0.025mg CuSO4·5H2O,6.25mg H3BO5,0.025mgCoCl·6H2O,8.65mg ZnSO4·7H2O,0.25mg Na2MoO4·2H2O,278mg FeSO4·7H2O,37.3mg Na2EDTA)上萌发生长5天,观察其性状。以在MS培养基培养的各株系做为对照。
结果如图2所示,在高盐MS培养基上萌发生长,AtST1过量表达植株(Super:AtST1-1和Super:AtST1-2)表现出明显耐盐表型,较野生型萌发正常,子叶绿色;AtST1突变体(atst1)表现明显盐敏感表型,萌发明显差于野生型,子叶基本不能正常伸展。
2、野生型、AtST1过量表达植株(Super:AtST1-1和Super:AtST1-2)和AtST1突变体(atst1)在高渗条件下的表型观察结果
将野生型拟南芥(Columbia型)、AtST1过量表达植株(步骤一获得的转Super:AtST1拟南芥单拷贝纯合株系,名称为Super:AtST1-1和Super:AtST1-2)和AtST1敲除突变体(atst1)拟南芥种子在含有300mM山梨醇(Sorbitol)的高渗MS培养基(以正常MS培养基为基础,加入300mM Sorbitol,其它成分与正常MS培养基一致。)上萌发生长5天,观察其性状。以在MS培养基培养的各株系做为对照。
结果如图3所示,在高渗MS培养基上萌发生长,AtST1过量表达植株(Super:AtST1-1和Super:AtST1-2)表现出明显耐高渗表型,较野生型萌发正常,子叶绿色;AtST1突变体(atst1)表现明显高渗敏感表型,萌发明显差于野生型,子叶基本不能正常伸展。
3、野生型、AtST1过量表达植株(Super:AtST1-1和Super:AtST1-2)和AtST1突变体(atst1)在外加ABA条件下的表型观察结果
将野生型拟南芥(Columbia型)、AtST1过量表达植株(步骤一获得的转Super:AtST1拟南芥单拷贝纯合株系,名称为Super:AtST1-1和Super:AtST1-2)和AtST1敲除突变体(atst1)拟南芥种子在含有0.3μM ABA的外加ABA培养基(以正常MS培养基为基础,加入0.3μM ABA,其它成分与正常MS培养基一致。)上萌发生长5天,观察其性状。以在MS培养基培养的各株系做为对照。
结果如图4所示,在外加ABA的MS培养基上萌发生长,AtST1过量表达植株(Super:AtST1-1和Super:AtST1-2)表现出明显耐ABA表型,较野生型萌发正常,子叶完全展开,绿色;AtST1突变体(atst1)表现明显ABA敏感表型,萌发明显差于野生型,子叶基本不能正常伸展。
4、高盐条件下AtST1基因的表达变化结果
在MS培养基上萌发生长一周的野生型拟南芥(Columbia型)幼苗转移到高盐MS培养基(以正常MS培养基为基础,加入120mM NaCl,其它成分与正常MS培养基一致。)上培养。培养条件采用全日照光照培养,温度22℃,光强100μmol/(m2*s)。每隔一段时间对相关材料取样,提取总RNA,总RNA的提取按Invitrogen Trizol Reagent的产品说明书进行。将总RNA用Takara公司的DNA酶消化RNA中的DNA后进行反转录,得到cDNA。以cDNA为模板,按ABI SYBR Green的产品说明书进行Realtime PCR反应。
结果如图5所示,在高盐条件下,AtST1基因表达明显升高,在高盐处理3天时,AtST1基因的表达达到最高,然后AtST1基因的表达逐步回到处理前水平,说明AtST1基因受高盐诱导表达。
序列表
<110>中国农业大学
<120>植物耐盐相关蛋白AtST1及其编码基因与应用
<130>CGGNARL102267
<160>3
<210>1
<211>1035
<212>DNA
<213>拟南芥属拟南芥(Arabidopsis thaliana)
<400>1
atggaatcga gtagcgttga tgagagtact acaagtacag gttccatctg tgaaaccccg 60
gcgataactc cggcgaaaaa gtcgtcggta ggtaacttat acaggatggg aagcggatca 120
agcgttgtgt tagattcaga gaacggcgta gaagctgaat ctaggaagct tccgtcgtca 180
aaatacaaag gtgtggtgcc acaaccaaac ggaagatggg gagctcagat ttacgagaaa 240
caccagcgcg tgtggctcgg gacattcaac gaagaagacg aagccgctcg tgcctacgac 300
gtcgcggttc acaggttccg tcgccgtgac gccgtcacaa atttcaaaga cgtgaagatg 360
gacgaagacg aggtcgattt cttgaattct cattcgaaat ctgagatcgt tgatatgttg 420
aggaaacata cttataacga agagttagag cagagtaaac ggcgtcgtaa tggtaacggt 480
aacatgacta ggacgttgtt aacgtcgggg ttgagtaatg atggtgtttc tacgacgggg 540
tttagatcgg cggaggcact gtttgagaaa gcggtaacgc caagcgacgt tgggaagcta 600
aaccgtttgg ttataccgaa acatcacgca gagaaacatt ttccgttacc gtcaagtaac 660
gtttccgtga aaggagtgtt gttgaacttt gaggacgtta acgggaaagt gtggaggttc 720
cgttactcgt attggaacag tagtcagagt tatgttttga ctaaaggttg gagcaggttc 780
gttaaggaga agaatctacg tgctggtgac gtggttagtt tcagtagatc taacggtcag 840
gatcaacagt tgtacattgg gtggaagtcg agatccgggt cagatttaga tgcgggtcgg 900
gttttgagat tgttcggagt taacatttca ccggagagtt caagaaacga cgtcgtagga 960
aacaaaagag tgaacgatac tgagatgtta tcgttggtgt gtagcaagaa gcaacgcatc 1020
tttcacgcct cgtaa 1035
<210>2
<211>344
<212>PRT
<213>拟南芥属拟南芥(Arabidopsis thaliana)
<400>2
Met Glu Ser Ser Ser Val Asp Glu Ser Thr Thr Ser Thr Gly Ser Ile
1 5 10 15
Cys Glu Thr Pro Ala Ile Thr Pro Ala Lys Lys Ser Ser Val Gly Asn
20 25 30
Leu Tyr Arg Met Gly Ser Gly Ser Ser Val Val Leu Asp Ser Glu Asn
35 40 45
Gly Val Glu Ala Glu Ser Arg Lys Leu Pro Ser Ser Lys Tyr Lys Gly
50 55 60
Val Val Pro Gln Pro Asn Gly Arg Trp Gly Ala Gln Ile Tyr Glu Lys
65 70 75 80
His Gln Arg Val Trp Leu Gly Thr Phe Asn Glu Glu Asp Glu Ala Ala
85 90 95
Arg Ala Tyr Asp Val Ala Val His Arg Phe Arg Arg Arg Asp Ala Val
100 105 110
Thr Asn Phe Lys Asp Val Lys Met Asp Glu Asp Glu Val Asp Phe Leu
115 120 125
Asn Ser His Ser Lys Ser Glu Ile Val Asp Met Leu Arg Lys His Thr
130 135 140
Tyr Asn Glu Glu Leu Glu Gln Ser Lys Arg Arg Arg Asn Gly Asn Gly
145 150 155 160
Asn Met Thr Arg Thr Leu Leu Thr Ser Gly Leu Ser Asn Asp Gly Val
165 170 175
Ser Thr Thr Gly Phe Arg Ser Ala Glu Ala Leu Phe Glu Lys Ala Val
180 185 190
Thr Pro Ser Asp Val Gly Lys Leu Asn Arg Leu Val Ile Pro Lys His
195 200 205
His Ala Glu Lys His Phe Pro Leu Pro Ser Ser Asn Val Ser Val Lys
210 215 220
Gly Val Leu Leu Asn Phe Glu Asp Val Asn Gly Lys Val Trp Arg Phe
225 230 235 240
Arg Tyr Ser Tyr Trp Asn Ser Ser Gln Ser Tyr Val Leu Thr Lys Gly
245 250 255
Trp Ser Arg Phe Val Lys Glu Lys Asn Leu Arg Ala Gly Asp Val Val
260 265 270
Ser Phe Ser Arg Ser Asn Gly Gln Asp Gln Gln Leu Tyr Ile Gly Trp
275 280 285
Lys Ser Arg Ser Gly Ser Asp Leu Asp Ala Gly Arg Val Leu Arg Leu
290 295 300
Phe Gly Val Asn Ile Ser Pro Glu Ser Ser Arg Asn Asp Val Val Gly
305 310 315 320
Asn Lys Arg Val Asn Asp Thr Glu Met Leu Ser Leu Val Cys Ser Lys
325 330 335
Lys Gln Arg Ile Phe Hi s Ala Ser
340
<210>3
<211>1112
<212>DNA
<213>人工序列
<220>
<223>
<400>3
ggatccctga aagcgacgtt ggatgttaac atctacaaat tgccttttct tatcgaccat 60
gtacgtaagc gcttacgttt ttggtggacc cttgaggaaa ctggtagctg ttgtgggcct 120
gtggtctcaa gatggatcat taatttccac cttcacctac gatggggggc atcgcaccgg 180
tgagtaatat tgtacggcta agagcgaatt tggcctgtag gatccctgaa agcgacgttg 240
tgttaacatc tacaaattgc cttttcttat cgaccatgta cgtaagcgct tacgtttttg 300
gtggaccctt gaggaaactg gtagctgttg tgggcctgtg gtctcaagat ggatcattaa 360
tttccaccta cgatgggggg catcgcaccg gtgagtaata ttgtacggct aagagcgaat 420
ttggcctgta ggatccctga aagcgacgtt ggatgttaac atctacaaat tgccttttct 480
tatcgaccat gtacgtaagc gcttacgttt ttggtggacc cttgaggaaa ctggtagctg 540
ttgtgggcct gtggtctcaa gatggatcat taatttccac cttcacctac gatggggggc 600
atcgcaccgg tgagtaatat tgtacggcta agagcgaatt tggcctgtag gatccgcgag 660
ctgctcaatc ccattgcttt tgaagcagct caacattgat ctctttctcg atcgagggag 720
atttttcaaa tcagtgcgca agacgtgacg taagtatccg agtcagtttt tatttttcta 780
ctaatttggt cgtttatttc ggcgtgtagg acatggcaac cgggcctgaa tttcgcgggt 840
attctgtttc tattccaact ttttcttgat ccgcagccat taacgacttt tgaatagata 900
cgctgacacg ccaagcctcg ctagtcaaaa gtgtaccaaa caacgcttta cagcaagaac 960
ggaatgcgcg tgacgctcgc ggtgacgcca tttcgccttt tcagaaatgg ataaatagcc 1020
ttgcttccta ttatatcttc ccccaaatta ccaatacatt acactagcat ctgaatttca 1080
taaccaatct cgatacacca aatcgactct ag 1112
Claims (1)
1.一种培育耐逆性提高的转基因植物的方法,是将序列表中序列2所示的氨基酸序列组成的蛋白质的编码基因转入目的植物中,得到转基因植物,所述耐逆性提高为转基因植物的耐盐性、耐渗透性和耐ABA性均强于所述目的植物;所述目的植物为拟南芥;所述编码基因的序列是序列表中的序列1;序列表中序列2所示的氨基酸序列组成的蛋白质的编码基因是通过重组表达载体导入所述目的植物中;
所述重组载体为在pCAMBIA1300:Super的多克隆位点间插入所述编码基因得到的重组表达载体;
所述pCAMBIA1300:Super是将核苷酸序列如序列表中序列3的自5’端第113-1112位所示的Super启动子插入pCAMBIA1300的HindIII和XbaI酶切位点间得到的载体。
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Citations (2)
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US20040009476A9 (en) * | 2000-08-24 | 2004-01-15 | Harper Jeffrey F. | Stress-regulated genes of plants, transgenic plants containing same, and methods of use |
CN101386645A (zh) * | 2008-10-29 | 2009-03-18 | 中国农业科学院作物科学研究所 | 一种植物耐盐蛋白及其编码基因与应用 |
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US20040009476A9 (en) * | 2000-08-24 | 2004-01-15 | Harper Jeffrey F. | Stress-regulated genes of plants, transgenic plants containing same, and methods of use |
CN101386645A (zh) * | 2008-10-29 | 2009-03-18 | 中国农业科学院作物科学研究所 | 一种植物耐盐蛋白及其编码基因与应用 |
Non-Patent Citations (2)
Title |
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AY091291;Yamada,K et al.;《Genbank》;20020918;第2页CDS段、第2页最后1段-第3页第1段 * |
Yamada,K et al..AY091291.《Genbank》.2002,第1-3页. |
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