CN109082428B - 水稻气孔开放型钾离子通道基因OsK2-1的应用及其表达载体 - Google Patents
水稻气孔开放型钾离子通道基因OsK2-1的应用及其表达载体 Download PDFInfo
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Abstract
本申请通过克隆获得了水稻气孔开放型钾离子吸收通道Osk2‑1基因,该基因主要在水稻地上部叶肉及气孔保卫细胞中表达,并且该基因蛋白具有介导钾离子内向吸收的转运活性;本发明首次发现该水稻气孔开放型钾离子通道基因OsK2‑1在提高农作物光合速率、气孔导度、蒸腾速率以及提高农作物高氮环境中氮素利用效率中的应用;本发明同时提供了含有上述水稻气孔开放型钾离子吸收通道基因OsK2‑1的动物表达载体和植物超表达载体,使得转Osk2‑1基因水稻生物量和籽粒产量显著增加,尤其是在高氮条件下展现出良好的增产效果和应用前景。
Description
技术领域
本发明涉及植物基因工程技术领域,特别是一种水稻气孔开放型钾离子通道蛋白基因Osk2-1的应用。
背景技术
钾是作物生长发育所必需的关键矿质营养元素,它不仅具有提高植物体内酶系统的活性、促进植物代谢、体内物质运输和蛋白质的合成等功能,而且能够增强植物的光合作用及植株的抗逆性。植物对钾的吸收主要由各种类别的钾转运体和钾通道介导(Epstein等,1963)。转运体,其主要特点是和被转运离子相结合,通过一系列构象变化实现跨膜转运,迄今为止,在植物中已发现存在三大钾转运体家族,分别为KUP/HAK/KT家族,TRK/HKT家族,CPA(cation proton antiporter)家族,这些转运体都具有转运钾的功能(Gierth and2007),但其转运钾的具体机制和特征还不完全清楚。钾离子通道是细胞膜上具有选择性转运钾离子功能的孔道蛋白,主要负责植物体内钾离子的动态平衡及分配。根据蛋白序列和结构特征将钾通道划分为三个类型:Shaker型、TPK型和Kir-like 型,其中研究最深入的是Shaker型钾通道。同样,根据蛋白质序列、结构及功能,将Shaker型钾通道细分为GroupⅠ、GroupⅡ、GroupⅢ、GroupⅣ和Group Ⅴ五组,这些钾通道在植物中的分布呈现多样性。
气孔是由一对特化的表皮细胞(保卫细胞)围成的孔状结构,归属于Group Ⅱ中的KAT型钾通道,主要在叶片和保卫细胞中表达,介导K+进入保卫细胞调控气孔的开放而与植物的光合作用和水分蒸腾密切相关(Lebaudy等,2008)。因为气孔是植物水分蒸腾和获取光合CO2的主要门户,促进气孔的高效开放不仅利于生物量的积累,更有利于养分的高效运输,从而有助于提高养分利用率和产量。
水稻是重要的粮食作物,生长在淹水条件下以吸收铵态氮为主要的氮源,当前的种植模式下,为保证水稻产量不得不施用远超过水稻所需要的氮素化肥“堆砌”充足的氮素养分供应以发挥现有高产水稻品种的增产潜力,这不仅导致氮肥利用率(NUE)走低,同时过量的氮也对环境造成巨大压力。解决上述问题的关键是充分挖掘水稻自身吸收/利用氮素的基因资源并通过转基因/育种技术,定向培育水稻的优良氮素营养形状,充分开发水稻自身吸收和利用氮素的潜力。
大量化肥的施用及集约化施肥方式使得现有农田的铵态氮水平可高达 20mM(Britto等,2002)。现有通过分子调控手段超表达铵吸收同化或响应基因改良水稻氮素营养形状的研究,超表达水稻铵吸收系统OsAMT1;1的水稻仅在小于等于0.3mM NH4 +(远低于现实农田铵水平)条件下展现出一定的正面效果,在较高铵(3mM NH4 +)条件下水稻生长反而出现生长抑制现象(Ranathunge等, 2014);调控另一个水稻铵吸收系统OsAMT1;3后,水稻体内出现C;N代谢失衡进而导致产量降低(Bao等,2015),而超表达铵同化基因(如OsGS1;1,1;2及 glnA)的水稻产量均降低(Cai等,2009)。超表达氮代谢调控基因来自玉米的转录因子Dof1能够提高水稻的氮素利用效率,但是这种正面效果也仅在低氮条件下出现(Kurai等,2011)。近年来发现的一些其他的转录因子如HAP3,14-3-3 蛋白,qNGR9,MiRNA,以及激酶CIPK8和CIPK23都参与氮素吸收代谢的调控,但是尚未见这些基因在提高水稻氮素利用效率中的应用。并且在应用中发现的有作用的基因都在氮素供应较低的条件下有效,这与我国当前高度集约化的生产模式不匹配。因此,充分发掘水稻自身具有应用前景尤其是在高氮条件下具有应用前景的基因资源具有重要的理论和现实意义。
发明内容
针对上述问题,本发明的目的之一是提供一种水稻气孔开放型钾离子吸收通道基因Osk2-1,其编码的Osk2-1蛋白在高氮条件下可以通过调控气孔开度增强光合速率和蒸腾速率,进而提供该蛋白在提高水稻氮素利用效率及水稻产量方面的应用。
为实现上述发明目的,本申请采用如下技术方案:
一种水稻气孔开放型钾离子吸收通道基因Osk2-1在提高农作物光合速率、气孔导度及蒸腾速率中的应用,该Osk2-1基因DNA的核苷酸序列如SEQ ID No.1所示;所述农作物优选水稻。
一种水稻气孔开放型钾离子通道基因OsK2-1在提高农作物高氮环境中氮素利用效率中的应用;所述钾离子通道基因OsK2-1的核苷酸序列如SEQ ID NO.1 所示;所述农作物优选水稻。
含有核苷酸序列如SEQ ID NO.1所示的水稻气孔开放型钾离子吸收通道基因Osk2-1的动物表达载体,如将水稻OsK2-1蛋白编码基因构建到动物表达载体pTracer-CMV3中,所得到的重组质粒pTracer-CMV3-OsK2-1。
含有核苷酸序列如SEQ ID NO.1所示的水稻气孔开放型钾离子吸收通道基因Osk2-1的植物超表达载体,如将水稻OsK2-1蛋白编码基因构建到单子叶植物超表达载体pCambia1301-ubiquitin-NOS(简称pUN1301)中,所得到的重组质粒pUN1301-OsK2-1。
本申请中,技术术语“低氮环境”是指:外界氮含量不高于66.7kg纯N/hm2;“高氮环境”是指:外界氮含量不低于300kg纯N/hm2。
本发明首次发现并应用了Osk2-1基因编码的了Osk2-1蛋白在高氮条件下,在调控水稻气孔开度、速率及蒸腾效率方面的应用,通过将该基因导入水稻并表达,实现在高氮条件下能够提高水稻的气孔开度、光合速率及蒸腾效率,从而提高农作物(特别是水稻)对高氮环境中氮的有效利用能力。
附图说明
图1为水稻Osk2-1基因克隆及生物信息学分析结果示意图;
其中图1A:M1为250bp DNAladder marker,lane 1为Osk2-1(NotI/NotI)的 PCR结果,lane 2为水稻Osk2-1(BamHI/SacI的PCR结果;图1B:为Osk2-1的生物信息学-进化树分析。
图2重组质粒图pTracer-CMV3-OsK2-1及pUN1301-OsK2-1的双酶切鉴定结果电泳图;图2A:M1为250bp DNAladder Marker,lane1-lane4为重组质粒 pTracer-CMV3-OsK2-1的NotI/NotI双酶切鉴定结果;图2B:M2为DL15,000 DNA Marker,lane1-6为pUN1301-OsK2-1的BamHI/SacI双酶切鉴定结果,M1 为250bp DNAladder Marker。
图3 OsK2-1的时空表达模式和精确组织定位结果示意图;
其中,图3A:OsK2-1的时空表达模式;图3B:OsK2-1的精确组织定位结果。
图4是OsK2-1的钾离子转运活性验证结果示意图;
其中图4A:转染pTracer-CMV3-OsK2-1的HEK-293T细胞转运钾离子特征电流曲线;图4B:转染空载体pTracer-CMV3的HEK-293T细胞转运钾离子特征电流曲线;图4C:OsK2-1介导的钾离子吸收所产生的净电流曲线示意图;
图5是小区实验低氮和高氮条件下转OsK2-1水稻株系各项指标实验结果示意图;
其中,图5A为气孔开度检测结果示意图,图5B为光合速率检测结果示意图,图5C为蒸腾速率检测结果示意图,其中LN表示小区实验中的低氮条件, HN表示小区实验中的高氮条件。
图6是小区实验低氮和高氮条件下条件下转OsK2-1水稻株系的产量示意图;
其中图6A为总生物量检测结果示意图;图6B为稻草重量检测结果示意图;图 6C为籽粒产量结果示意图。
具体实施方式
实施例中,日本晴和中花11水稻均购自于北京未名凯拓农业生物技术有限公司、pMD19-T Simple购自TAKARA公司。
以下实施例所涉及的原料和试剂,除非特殊说明,均为市售渠道购买。
实施例1水稻OsK2-1基因的克隆及生物信息学分析
提取水培生长10天的日本晴水稻的总RNA并以其为模板,以Oligo(dT)18 为引物反转录,得到反转录产物cDNA。根据水稻OsK2-1的全长编码序列,设计扩增出完整编码阅读框的引物,根据需要设计两对引物,一对为上游引物F1 和下游引物R1(forpTracer-CMV3),另一对为上游引物F2和下游引物R2(for pUN1301),并在各引物上引入不同的酶切位点,如下:
F1(NotI,SEQ ID NO.3):
GTCGCGGCCGCATGACCCAAGCTCACTCAAAATCTTGCTTC(for pTracer-CMV3);
R1(NotI,SEQ ID NO.4):
GTCGCGGCCGCTACATCTCAAGAAGGAATAGATGGTCGCCATC(for pTracer-CMV3);
F2(BamHI,SEQ ID NO.5):
GTCGGATCCATGACCCAAGCTCACTCAAAATCTTGCTTC(forpUN1301);
R2(SacI,SEQ ID NO.6):GTCGAGCTCCTACATCTCAAGAAGGAATAGATGGTCGCCATC(forpUN1301);
使用高保真酶进行PCR扩增,PCR反应体系为50μL,5PrimeScriptbuffer(Mg2+)为10μL,dNTPs(2.5μmol/L)为4μL,上、下游引物(10μmol/L)各为1μL,(2.5u/μL)为0.5μL,DMSO为2.5μL,加水补齐至50μL。
PCR反应程序:95℃预变性5min;95℃变性30sec,60℃退火30sec,72℃延伸2min30sec,30个循环;72℃延伸10min;4℃保存。
电泳检测,结果得到两条约2.2kb左右的单一产物(图1A)。通过凝胶电泳回收扩增片段,与pMD19-T Simple载体相连接获得TA-OsK2-1质粒,委托北京六合华大基因科技有限公司上海分公司进行测序,确定获得水稻OsK2-1的 cDNA序列,其cDNA序列(大小为2157bp)及其编码的蛋白质的氨基酸序列 (大小为719aa)分别如SEQ ID No.1和SEQ IDNo.2所示。进化树分析表明,水稻OsK2-1与已经报道的拟南芥气孔钾离子吸收通道AtKAT1具有高度的同源性(图1B),暗示了其在功能上的相似性。
实施例2:水稻OsK2-1基因表达的组织特异性
1.水稻OsK2-1基因的时空表达模式
为了解OsK2-1基因在水稻不同组织器官中的表达情况,将萌发生长一致的水稻苗在木村营养液(参见文献:Li SM等,2006)中培养7d后再进行不同浓度的钾处理,这些处理分别为:
对照组:,完全木村营养液(2mM KCl);
缺K组:,0mM KCl(2mM NaCl代替);
高K组:,20mM KCl;
每天更换营养液,处理两天后分根和地上部取样,冻于液氮中,分别提取了水稻根和地上部的总RNA,以Oligo(dT)18为引物反转录(操作步骤按照逆转录试剂盒说明(AMV)进行),得到反转录产物cDNA。以此cDNA为模板,以特异引物Osk2-1F1和Osk2-1R1进行半定量PCR分析。
PCR反应体系为25μl:2μl MgCl2,2μl dNTP,2.5μl ExTaqbuffer,0.125μl ExTaq酶,上下游特异引物各1.25μl,模板1μl,用dH2O补齐至25μl;
PCR反应程序为:94℃,3min;94℃,30sec,60℃,30sec,72℃,1min,35个循环;72℃,10min。
内参OsActin2为28个循环,上游引物和下游引物分别为OsActin-F1和 OsActin-R1。
OsK2-1F1(SEQ ID NO.7):5'TTCTTTCCGATCATCCGGCAAGCCT 3';
OsK2-1R1(SEQ ID NO.8):5'GTGGAGACCCTGTCTGCACTGTTGC 3';
OsActin-F1(SEQ ID NO.9):5'CTCCATCATGAAGTGCGACGTGGAT 3';
OsActin-R1(SEQ ID NO.10):5'TGGTACCCGCATCAGGCATCTGATT 3';
结果显示,在不同钾浓度条件下,OsK2-1基因都主要在水稻地上部的表达,而在根中几乎检测不到其表达(图2A),表明OsK2-1基因在水稻中有明显的组织特异性,其在地上部的表达较为活跃。
2.水稻OsK2-1基因的精确组织定位
为进一步了解OsK2-1的功能特征,通过原位杂交研究了OsK2-1在水稻地上部的精确组织定位特征,根据Osk2-1的mRNA序列,设计特异性寡核苷酸序列正义(sense)链(SEQID NO.11):5’CATGGGACCAGCCGCACCAG 3’和反义(anti-sense)链(SEQ ID NO.12):5’TGGTCGCCATCGCGGATAACA 3’,并在5’分别进行地高辛(Digoxigenin)标记,在3’分别进行磷酸化修饰,从而得到sense和anti-sense的探针并由上海Invitrogen公司合成探针。
通过制备水稻叶片组织切片并进行杂交(该方法为本领域常规技术,具体可参见文献:Zhao等,2009),结果显示OsK2-1主要定位于叶片的叶肉和保卫细胞(图2B)中,暗示其在气孔开度调控方面的作用。
实施例3:重组质粒pTracer-CMV3-OsK2-1和pUN1301-OsK2-1的构建
将水稻OsK2-1基因经由酶切位点经酶切位点NotI/NotI构建到 pTracer-CMV3上,转化大肠杆菌感受态细胞DH5α后,提取质粒并酶切(图 3A)鉴定(Kanwar,等,1980),结果表明已经成功获得重组质粒 pTracer-CMV3-OsK2-1。
将水稻OsK2-1基因经由酶切位点BamHI/SacI构建到pUN1301,转化大肠杆菌感受态细胞DH5α后,提取质粒并酶切(图3B)鉴定。结果表明已经成功获得重组质粒pUN1301-OsK2-1。
实施例4:水稻OsK2-1基因的电生理功能分析
1.溶液配制
电极内液成分为:120mM K-Gluconate,5mM NaCl,5mM KCl,2mM MgCl2, 2mMCaCl2,4mM EGTA,10mM HEPES,14mM Tris-creatine phosphate,4mM Tris-ATP,0.3mMTris-GTP,用1M KOH将pH调至7.3,K+终浓度约为140 mM,渗透压约为300mOsM。
细胞外液成分为:1.0mM MgCl2,1.2mM CaCl2,10.0mM HEPES,20mM Glucose,150mMKCl(低于150mM时用NaCl代替KCl补充至150mM),用 1M NaOH调节pH至7.4,渗透压用Sucrose调至约320mOsm。
2.质粒转染
将质粒pTracer-CMV3-Osk2-1用Lipofectamine转染到HEK293-T细胞中,对照转染空载体pTracer-CMV3,再放回细胞培养箱中表达一天,第二天挑取带绿色荧光的细胞检测。
3.检测程序设置:
在显微镜下,将电极(已灌注电极内液)通过电动微操接触HEK293-T细胞,施加负压使电极尖端与HEK293-T质膜之间形成高阻封接(GΩ),继续施加负压,吸破HEK293-T细胞膜,使电极与细胞之间形成全细胞记录模式。稳定5min后,开始记录电流大小。记录过程中,吸收型钾离子通道电流电压施加方案为钳制电压-40mV,阶越电压为20mV,共11个刺激电压,从-180mV到+20mV,刺激间隔5.5s,采样频率10kHz。数据采集用软件为pCLAMP 10.0。
记录到表达pTracer-CMV3-Osk2-1的细胞电流特征曲线如图4A所示,记录到表达pTracer-CMV3空载体的细胞电流特征曲线如图4B所示,获得Osk2-1通道净电流特征曲线如图4C所示,由图可知Osk2-1介导K+吸收产生了电流,是典型的内向吸收型钾离子通道。
实施例5利用pUN1301-Osk2-1转基因水稻植株研究Osk2-1的应用
1.转Osk2-1转基因水稻材料的制备
采用根癌农杆菌介导法(该方法为本领域常规技术,具体可参见文献: Valvekens等,1988)将实施例3中构建获得的重组质粒pUN1301-Osk2-1转入水稻中花11品种。诱导水稻成熟愈伤,将长到适度大小的愈伤组织挑出,放入农杆菌悬浮液侵染5-10分钟,将愈伤组织取出,置于无菌的滤纸上沥干30-40 分钟,愈伤组织置于共培养基上,28℃暗培养3天,然后将愈伤转入含50mg/L 潮霉素的选择培养基上进行筛选,挑取颜色亮黄的抗性愈伤移入装有分化培养基的培养皿中,放入恒温培养箱分化成苗。再放入生根培养基中壮苗一到两周。通过鉴定获得Osk2-1超表达水稻株系OE-26。
2.小区实验检验转Osk2-1水稻的气孔导度、光合速率及蒸腾速率检测
将转Osk2-1的水稻(水稻株系OE-26)播种至低氮(LN,66.7kg纯N/hm2)、高氮(HN,300kg纯N/hm2)两个氮素水平小区中,在结实灌浆期采用LiCor-6400 光合仪测定剑叶的光合速率、气孔导度及蒸腾速率(该测定技术为本领域常规技术,具体可参见文献:Reguera,等,2013),结果显示,转Osk2-1水稻的的气孔开度(图5A)、光合速率(图5B)及蒸腾速率(图5C)显著高于野生型(中花11),增加趋势在高氮条件下尤为显著(图5),表明Osk2-1基因能够调控气孔的开度并响应环境中不同的氮水平。
3.转Osk2-1水稻的氮素利用效率
在成熟期将地上部齐地面割下,单株收获装至独立的网袋中,晾干后进行总生物量、稻草重量及籽粒产量的鉴定,结果显示,高氮条件(外界氮含量不低于 300kg纯N/hm2)下(转Osk2-1株系的总生物量(图6A)包括稻草重量(图 6B)及籽粒产量(图6C)显著高于野生型。也就是说,Osk2-1通过调控水稻的气孔开度,增强了水稻的光合速率及蒸腾速率,光合效率的增强促进了光合产物的积累,蒸腾速率的提高带动了氮钾养分向地上部的转移,促进了结实灌浆期关键功能叶剑叶氮的积累,利于其光合作用时间的延长及碳水化合物的积累,并且这些优势最终体现在转Osk2-1水稻的产量上。
本技术领域技术人员可以理解的是,除非另外定义,这里使用的所有术语(包括技术术语和科学术语)具有与本发明所属领域中的普通技术人员的一般理解相同的意义。还应该理解的是,诸如通用工具书中定义的那些术语应该被理解为具有与现有技术的上下文中的意义一致的意义,并且除非像这里一样定义,不会用理想化或过于正式的含义来解释。
以上所述的具体实施方式,对本发明的目的、技术方案和有益效果进行了进一步详细说明,所应理解的是,以上所述仅为本发明的具体实施方式而已,并不用于限制本发明,凡在本发明的精神和原则之内,所做的任何修改、等同替换、改进等,均应包含在本发明的保护范围之内。
序列表
<110> 中国科学院南京土壤研究所
<120> 水稻气孔开放型钾离子通道基因OsK2-1的应用及其表达载体
<141> 2018-08-22
<160> 12
<170> SIPOSequenceListing 1.0
<210> 1
<211> 2157
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 1
atgacccaag ctcactcaaa atcttgcttc caccagttct gggatggact ccagatcaag 60
aggagcagcg atagcttcac tgtagagcta ctgccatccc ttggtgcaac tataaaccac 120
tccaacaagt tgcagaagtt catcatatcg ccttatgatc cccggtacag atcctgggag 180
ctgttcctta tagtcctagt tgtttactct gcctggattt gtccgtttga actagcattc 240
ctgagggact taccatctaa gcttttgcta gttgagaaca ttgtggatat attctttgcc 300
attgatattg ttttgacgtt cttcgttgct tatgtcgata gcaaaacaca tcttcttgtg 360
gatgaccgaa agagaattgc aatgaggtac ctatccactt ggtttatctt tgatgtatgt 420
tcaacagcac catttcaacc aatcatcctt ctatttacac acaaggggaa tgacattgct 480
ttcaaggtac tcaatttgct caggctgtgg cgtcttcacc gagtcagttc actatttgcc 540
aggctggaga aagacatccg attcaactat ttctggacta ggtgctcaaa acttatttct 600
gtgaccctct tcgcagtaca ttgtgcagga tgcttcaatt atatgattgc tgacagatat 660
cccaacccag agaaaacatg gataggagct gtgatgtcaa ctttcagatc agagagctta 720
tggactagat atattactgc tctttactgg tccattacaa cactgacaac aacaggatat 780
ggggacctac atgctgaaaa tccaacagag atgctatttg atattgtcta tatgatgttt 840
aatctgggct tgacagctta tctcattggc aacatgacca acctagttgt ccatgggacc 900
agccgcacca ggaaatttag ggactcaatc caagcagcat cagagtttgc agcacgcaat 960
cagctacctg agaacataaa gcagcaagtg ctatctcact tctgcctaca attcaagaca 1020
gagggactca accagcaggt catgttagac tgtctaccaa aaggaatccg ctcaagcata 1080
gcatacagct tattctttcc gatcatccgg caagcctatc tcttcaatgg agtatctggc 1140
aatttcattg cagaactggt catggaagtg caggctgagt acttcccacc aaaggaagat 1200
ataatattgc agaatgaagg agaagcagat gtttacatag tagtttcagg agcagtgaat 1260
ataataacaa caatacatgg gaatgagcag gtatatgaga aaattgcaga gggagaaatg 1320
tttggagagg ttggttcttt atgtaacata ccccagccat ttacttgccg cacagcggag 1380
ctatcacagc ttttaagaat aagcaaaaca aggcttagag agatcattga agaaaacagg 1440
gaagacagta acattctaat gaacaaccta gttcagaagc tgaagctacg agaaagttta 1500
cctgacatga atcaaccaga ccgaagattc ttgagcaagt atgagctatt ccacgttcct 1560
cgagaagcat ggctgctgaa aaagtcacaa ctacattaca cagagcacac aagtagggat 1620
tctagtaaca acactccagt atttggaggt gacagatatt ccagacagtt acttggagaa 1680
gcaacccgat cttcggcatc agaaaacgaa aatagcagta tgacagataa agaagagaac 1740
catgatgaag ttcacacaaa ctgtgagatc aaaaaaagaa cggaagagca ctgtatccag 1800
ataaactctg aagatagcag ctccacatac agtcagcgaa caatgaatgc aacagtgcag 1860
acagggtctc cacataaaac agaagaaaac ataacaagaa gaattccaga tgagtactac 1920
ataaaagaag caaacaaaag agttacaatt cacaaatatc gtcataactc aacagtctct 1980
gctgcgcaaa atggaaagct tatcaagctg cctacatcat tggaggaact gttcaaaatt 2040
ggcagtcaga agtttcaagg ttttcatcct agaaaggtgg tcagtagaga ttatgctgaa 2100
atagatgatg tcagtgttat ccgcgatggc gaccatctat tccttcttga gatgtag 2157
<210> 2
<211> 718
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 2
Met Thr Gln Ala His Ser Lys Ser Cys Phe His Gln Phe Trp Asp Gly
1 5 10 15
Leu Gln Ile Lys Arg Ser Ser Asp Ser Phe Thr Val Glu Leu Leu Pro
20 25 30
Ser Leu Gly Ala Thr Ile Asn His Ser Asn Lys Leu Gln Lys Phe Ile
35 40 45
Ile Ser Pro Tyr Asp Pro Arg Tyr Arg Ser Trp Glu Leu Phe Leu Ile
50 55 60
Val Leu Val Val Tyr Ser Ala Trp Ile Cys Pro Phe Glu Leu Ala Phe
65 70 75 80
Leu Arg Asp Leu Pro Ser Lys Leu Leu Leu Val Glu Asn Ile Val Asp
85 90 95
Ile Phe Phe Ala Ile Asp Ile Val Leu Thr Phe Phe Val Ala Tyr Val
100 105 110
Asp Ser Lys Thr His Leu Leu Val Asp Asp Arg Lys Arg Ile Ala Met
115 120 125
Arg Tyr Leu Ser Thr Trp Phe Ile Phe Asp Val Cys Ser Thr Ala Pro
130 135 140
Phe Gln Pro Ile Ile Leu Leu Phe Thr His Lys Gly Asn Asp Ile Ala
145 150 155 160
Phe Lys Val Leu Asn Leu Leu Arg Leu Trp Arg Leu His Arg Val Ser
165 170 175
Ser Leu Phe Ala Arg Leu Glu Lys Asp Ile Arg Phe Asn Tyr Phe Trp
180 185 190
Thr Arg Cys Ser Lys Leu Ile Ser Val Thr Leu Phe Ala Val His Cys
195 200 205
Ala Gly Cys Phe Asn Tyr Met Ile Ala Asp Arg Tyr Pro Asn Pro Glu
210 215 220
Lys Thr Trp Ile Gly Ala Val Met Ser Thr Phe Arg Ser Glu Ser Leu
225 230 235 240
Trp Thr Arg Tyr Ile Thr Ala Leu Tyr Trp Ser Ile Thr Thr Leu Thr
245 250 255
Thr Thr Gly Tyr Gly Asp Leu His Ala Glu Asn Pro Thr Glu Met Leu
260 265 270
Phe Asp Ile Val Tyr Met Met Phe Asn Leu Gly Leu Thr Ala Tyr Leu
275 280 285
Ile Gly Asn Met Thr Asn Leu Val Val His Gly Thr Ser Arg Thr Arg
290 295 300
Lys Phe Arg Asp Ser Ile Gln Ala Ala Ser Glu Phe Ala Ala Arg Asn
305 310 315 320
Gln Leu Pro Glu Asn Ile Lys Gln Gln Val Leu Ser His Phe Cys Leu
325 330 335
Gln Phe Lys Thr Glu Gly Leu Asn Gln Gln Val Met Leu Asp Cys Leu
340 345 350
Pro Lys Gly Ile Arg Ser Ser Ile Ala Tyr Ser Leu Phe Phe Pro Ile
355 360 365
Ile Arg Gln Ala Tyr Leu Phe Asn Gly Val Ser Gly Asn Phe Ile Ala
370 375 380
Glu Leu Val Met Glu Val Gln Ala Glu Tyr Phe Pro Pro Lys Glu Asp
385 390 395 400
Ile Ile Leu Gln Asn Glu Gly Glu Ala Asp Val Tyr Ile Val Val Ser
405 410 415
Gly Ala Val Asn Ile Ile Thr Thr Ile His Gly Asn Glu Gln Val Tyr
420 425 430
Glu Lys Ile Ala Glu Gly Glu Met Phe Gly Glu Val Gly Ser Leu Cys
435 440 445
Asn Ile Pro Gln Pro Phe Thr Cys Arg Thr Ala Glu Leu Ser Gln Leu
450 455 460
Leu Arg Ile Ser Lys Thr Arg Leu Arg Glu Ile Ile Glu Glu Asn Arg
465 470 475 480
Glu Asp Ser Asn Ile Leu Met Asn Asn Leu Val Gln Lys Leu Lys Leu
485 490 495
Arg Glu Ser Leu Pro Asp Met Asn Gln Pro Asp Arg Arg Phe Leu Ser
500 505 510
Lys Tyr Glu Leu Phe His Val Pro Arg Glu Ala Trp Leu Leu Lys Lys
515 520 525
Ser Gln Leu His Tyr Thr Glu His Thr Ser Arg Asp Ser Ser Asn Asn
530 535 540
Thr Pro Val Phe Gly Gly Asp Arg Tyr Ser Arg Gln Leu Leu Gly Glu
545 550 555 560
Ala Thr Arg Ser Ser Ala Ser Glu Asn Glu Asn Ser Ser Met Thr Asp
565 570 575
Lys Glu Glu Asn His Asp Glu Val His Thr Asn Cys Glu Ile Lys Lys
580 585 590
Arg Thr Glu Glu His Cys Ile Gln Ile Asn Ser Glu Asp Ser Ser Ser
595 600 605
Thr Tyr Ser Gln Arg Thr Met Asn Ala Thr Val Gln Thr Gly Ser Pro
610 615 620
His Lys Thr Glu Glu Asn Ile Thr Arg Arg Ile Pro Asp Glu Tyr Tyr
625 630 635 640
Ile Lys Glu Ala Asn Lys Arg Val Thr Ile His Lys Tyr Arg His Asn
645 650 655
Ser Thr Val Ser Ala Ala Gln Asn Gly Lys Leu Ile Lys Leu Pro Thr
660 665 670
Ser Leu Glu Glu Leu Phe Lys Ile Gly Ser Gln Lys Phe Gln Gly Phe
675 680 685
His Pro Arg Lys Val Val Ser Arg Asp Tyr Ala Glu Ile Asp Asp Val
690 695 700
Ser Val Ile Arg Asp Gly Asp His Leu Phe Leu Leu Glu Met
705 710 715
<210> 3
<211> 41
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 3
gtcgcggccg catgacccaa gctcactcaa aatcttgctt c 41
<210> 4
<211> 43
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 4
gtcgcggccg ctacatctca agaaggaata gatggtcgcc atc 43
<210> 5
<211> 39
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
gtcggatcca tgacccaagc tcactcaaaa tcttgcttc 39
<210> 6
<211> 42
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 6
gtcgagctcc tacatctcaa gaaggaatag atggtcgcca tc 42
<210> 7
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
ttctttccga tcatccggca agcct 25
<210> 8
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 8
gtggagaccc tgtctgcact gttgc 25
<210> 9
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 9
ctccatcatg aagtgcgacg tggat 25
<210> 10
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 10
tggtacccgc atcaggcatc tgatt 25
<210> 11
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 11
catgggacca gccgcaccag 20
<210> 12
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 12
tggtcgccat cgcggataac a 21
Claims (1)
1.水稻气孔开放型钾离子通道基因OsK2-1在提高农作物在高氮环境中对氮素利用效率中的应用;所述钾离子通道基因OsK2-1的核苷酸序列如SEQ ID NO.1所示;
所述高氮环境是指:外界氮含量不低于300 kg 纯N /hm2。
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Citations (3)
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WO2003000898A1 (en) * | 2001-06-22 | 2003-01-03 | Syngenta Participations Ag | Plant genes involved in defense against pathogens |
CN102492028A (zh) * | 2011-11-29 | 2012-06-13 | 中国科学院南京土壤研究所 | 水稻钾双向整流型离子通道OsAKT2/3分子及应用 |
CN103215279A (zh) * | 2013-04-26 | 2013-07-24 | 大连理工大学 | 一种钾离子通道蛋白基因、其编码的蛋白及其应用 |
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Patent Citations (3)
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WO2003000898A1 (en) * | 2001-06-22 | 2003-01-03 | Syngenta Participations Ag | Plant genes involved in defense against pathogens |
CN102492028A (zh) * | 2011-11-29 | 2012-06-13 | 中国科学院南京土壤研究所 | 水稻钾双向整流型离子通道OsAKT2/3分子及应用 |
CN103215279A (zh) * | 2013-04-26 | 2013-07-24 | 大连理工大学 | 一种钾离子通道蛋白基因、其编码的蛋白及其应用 |
Non-Patent Citations (5)
Title |
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"Optimization of ammonium acquisition and metabolism by potassium in rice ( Oryza sativa L.cv.IR-72)";KONSTANTINE D. BALKOS et al.;《Plant, Cell and Environment》;20101231;第33卷;第23-34页 * |
"Predicted:Oryza sativa Japonica Group potassium channel KAT1-like(LOC4328864),mRNA",Accession Number:XM_015767872.2;genbank;《GenBank》;20180807;第1-2页 * |
"Unique Features of Two Potassium Channels, OsKAT2 and OsKAT3, Expressed in Rice Guard Cells";Hyunsik Hwang et al.;《PLOS ONE》;20130813;第8卷(第8期);第1-14页 * |
"应用钾通道基因对水稻和棉花钾素营养的研究";王义霞;《中国优秀硕士学位论文全文数据库 农业科技辑》;20141215(第12期);第12页第1.3节、第13页第2.1.1.3节、第32页第3.3.1.1节-第35页第3.3.1.3节、第41页第3.3.4.1节-第43页第3.3.4.2节,表3-9和表3-10 * |
"水稻钾离子通道基因OsKAT1.1的克隆、表达载体的构建及其电生理功能";李俊林 等;《基因组学与应用生物学》;20111231;第30卷(第4期);第346页摘要和第347页第1.3节,第348-349页第1.4节 * |
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