CN109134632A - 调控植物根发育的蛋白及其编码基因和应用 - Google Patents
调控植物根发育的蛋白及其编码基因和应用 Download PDFInfo
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Abstract
本发明涉及生物技术领域,特别涉及一种植物发育相关的蛋白及其编码基因在控制植物根生长与发育中的应用。具体而言,本发明公开了一种蛋白质,其氨基酸序列如SEQ ID No:1所示;本发明还同时公开了编码上述蛋白质的基因,该基因的编码区的核苷酸序列如SEQ ID NO:2所示。本发明还同时公开了该基因的用途:用于构建转基因植物,所述转基因植物的根长度变长。
Description
技术领域
本发明涉及生物技术领域,特别涉及一种植物发育相关的蛋白及其编码基因在控制植物根生长与发育中的应用。
背景技术
根是植物重要的地下器官,其不仅可以固定植株,而且还可以从土壤中吸收营养、水分等物质供植物的生长发育。此外,植物的根系还可以监测土壤环境中的各种信号分子,比如土壤中的水分含量,营养物质的水平以及一些植物激素及信号等,以提高植物对其生长环境的适应性。植物根系主要分为两大类,分别为直根系和须根系。水稻是单子叶模式植物,其根系为须根系,主要由种子根和不定根及种子根和不定根上的侧根和根毛组成(Rebouillat et al.2009;Coudert et al.2010)。研究表明冠根比高,发达的根系有助于提高水稻的抗旱能力及产量(Fukai et al.1995;Gowda et al.2011)。
之前的研究表明植物激素生长素对水稻根的发生与发育有着重要的作用。生长素转运蛋白基因OsAUX1的T-DNA插入突变体Osaux1,在水培条件下表现为根及根毛短的表型(Yu et al.2015)。降低生长素响应基因OsARF12的表达可使水稻根变短(Qi et al.2012)。水稻生长素受体相关基因OsTIR1和OsAFB2的表达受到miR393调控进而调控水稻主根、冠根、根毛的生长发育(Bian et al.2012;Xia et al.2012)。此外Oscyp2突变可以降低OsTIR1与OsIAA11间的相互作用,从而导致OsIAA11积累并抑制生长素信号,从而调控侧根的发育(Kang et al.,2013)。类似地,基因OsIAA11或OsIAA13功能获型突变体表现为无侧根的表型(Kitomi et al.2012;Zhu et al.2012)。最近研究表明,OsCYP2也可以与C2HC型锌指蛋白OsZFP相互作用形成复合体共同调节侧根发育(Cui et al.2017)。
除了生长素外,植物激素乙烯也参与了水稻根发育。乙烯信号转导途径的转录因子OsEIL1参与调控水稻根伸长(Mao et al.2006)。最近的研究表明与野生型相比,乙烯响应因子的功能获得型突变体Oserf2植株为短根表型,而沉默OsERF2的植株表现为长根表型。OsERF2参与协调糖代谢和激素信号传导的一系列基因的表达(Xiao et al.2016)。另一个编码AP2/ERF转录因子的基因SHB影响赤霉素的生物合成,进而导致根分生组织细胞的伸长和增殖(Li et al.2015)。此外,最近的研究表明,独脚金内酯与水杨酸生物合成相关基因也在水稻根生长中起着重要作用(Sun et al.2016;Xu et al.2017)。
尽管许多调控水稻根生长于发育的关键基因被克隆研究,但是我们对详细的根的发生及伸长的机制还知之甚少。因此,分离鉴定水稻根长发育的新基因对进一步全面了解其发生与发育的机理,获得优良基因以培育新的水稻品种是十分重要的和必要的。
参考文献:
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发明内容
本发明要解决的技术问题是提供一种调控植物根发育相关蛋白及其编码基因和应用。
为了解决上述技术问题,本发明提供如下技术方案:
本发明提供的蛋白(OsLR1),来自粳稻秀水134(Oryza sativa L.ssp.Japonicacv.Xiushui134),是如下(a)或(b)的蛋白质:
(a)由序列表中序列1所示的氨基酸序列组成的蛋白质;
(b)将序列1的蛋白质经过一个或几个氨基酸残基的取代和/或缺失和/或添加且与植物根发育相关的由其衍生的蛋白质;
所述植物发育体现在植株根长或者特定器官的大小等性状上。
为了使(a)中的OsLR1便于纯化,可在由序列表中序列1所示的氨基酸基酸序列组成的蛋白质的氨基末端或羧基末端连接上包括但不限于如表1所示的标签。
表1.标签的序列。
上述(b)中的OsLR1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(b)中的OsLR1的编码基因可通过将序列表中序列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1所示的标签的编码序列得到。
所述蛋白的编码基因也属于本发明的保护范围。
所述蛋白的编码基因(OsLR1)可为如下1)或2)或3)或4)的DNA分子:
1)、其编码序列是序列表中序列2所示的DNA分子,即,基因的编码区的核苷酸序列如SEQ ID NO:2所示;
2)、序列表中序列3所示的DNA分子,即,基因的基因组的核苷酸序列如SEQ ID NO:3所示;
3)、在严格条件下与1)或2)限定的DNA序列杂交且编码相同功能蛋白质的DNA分子;
4)、与1)或2)或3)限定的DNA序列具有90%以上同源性,且编码相同功能蛋白质的DNA分子。
含有所述基因或其反义基因的重组表达载体、表达盒、转基因细胞系或重组菌均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OsLR1基因的重组表达载体。
所述植物表达载体包括但不限于如双元农杆菌载体和可用于植物微弹轰击的载体等。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,如包括但不限于农杆菌冠瘿瘤诱导(Ti)质粒基因(如胭脂合成酶Nos基因)、植物基因(如大豆贮藏蛋白基因)3'端转录的非翻译区均具有类似功能。
使用OsLR1构建重组植物表达载体时,在其转录起始核苷酸前可加上任何一种增强型启动子或组成型启动子,包括但不限于如花椰菜花叶病毒(CAMV)35S启动子、玉米的泛素启动子(ubiquitin),它们可单独使用或与其它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白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,如加入包括但不限于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、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、卡那霉素标记物等)或是抗化学试剂标记基因(如抗除草剂基因)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。
所述重组表达载体具体可为如下(I)或(II):
(I)将序列表的序列2所示DNA分子插入质粒pCAMBIA1300-35S-EGFP的多克隆位点得到的含有上述基因的重组表达载体;
(II)将序列表的序列2所示基因DNA片段通过正向与反向连接到过渡载体pBSSK-in后插入质粒pCAMBIA1300-35S-EGFP得到的重组表达载体;
(I)或(II)中的所述质粒pCAMBIA1300-35S-EGFP与pBSSK-in都是本实验室改造得到的(已公开)。
本发明还保护一种培育转基因植物的方法,是将所述基因或基因的片段通过正反向连接到过渡载体pBSSK-in后导入目的植物中,所述转基因植物为与目的植物相比根长度改变的转基因植物。
利用任何一种可以引导外源基因在植物中表达的载体,将本发明所提供的OsLR1基因或基因的片段通过正反向连接到过渡载体pBSSK-in后导入植物细胞,可获得根长改变的转基因细胞系及转基因植株。携带有OsLR1基因或其部分基因正反向连接的表达载体可通过使用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植物细胞或组织,并将转化的植物组织培育成植株。被转化的植物宿主可为禾本科植物,如水稻(如秀水134)。
当培育根长大于目的植物的转基因植物时;所述方法是将(I)所述重组表达载体导入目的植物中,得到根变长的转基因植物。当培育根长小于目的植物的转基因植物时;所述方法是将(II)所述重组表达载体导入目的植物中,得到根变短的转基因植物。
本发明发现了一个新蛋白OsLR1及其编码基因OsLR1,并获得了含有该编码基因或该基因的部分片段通过正向与反向连接到过渡载体pBSSK-in的重组表达载体,用重组载体转化目的植物,可以得到根长改变的转基因植物。因此OsLR1可以作为一种潜在的分子育种工具,通过改良植物的根长来提高植物产量。
附图说明
下面结合附图对本发明的具体实施方式作进一步详细说明。
图1为实施例1中突变体Oslr1表型;
A为突变体Oslr1和野生型WT 7天的表型(标尺为5厘米),B为突变体Oslr1和野生型WT 7天的株高统计图,C为突变体Oslr1和野生型WT 7天的根长统计图;误差用SD用表示(统计的个体数大于30),***P<0.001(T检验)。
图2为实施例1中突变体Oslr1和野生型WT的根尖细胞观察及数据统计;
A为扫描电子显微镜观察到的主根根尖伸长区细胞(标尺为100微米),B为主根伸长区细胞长、宽的统计数据,***P<0.001(T检验)。
图3为实施例1中不同浓度IAA处理下Oslr1的表型;
A为浓度为1μM的IAA处理WT与Oslr1 7天的表型(标尺为5厘米),B为不同浓度IAA培养7天苗的根长度统计数据。
图4为实施例1中不同浓度NAA处理下Oslr1的表型;
A为浓度为0.1μM的NAA处理WT与Oslr1 7天的表型(标尺为5厘米),B为不同浓度NAA培养7天苗根长度的统计数据。
图5为实施例1中OsLR1的染色体位置及编码蛋白结构示意图;
黑色的方框表示外显子,黑色线条表示内含子。其编码的蛋白胞外域有一个LRR功能域,胞内域包含有KD激酶功能域,JM功能域及CT结构域。
图6为实施例1中7天苗龄野生型主根中OsLR1的表达对不同植物激素的响应;
H2O,水;Ethanol,乙醇;2,4D,2,4-二氯苯氧乙酸;6-BA,6-苄氨基腺嘌呤;ABA,脱落酸;e-BL,表油菜素内酯;GA3,赤霉素;GR24,独脚金内酯类似物;IAA,吲哚-3-乙酸。
图7为实施例1中不同组织器官OsLR1的表达;
根、茎、叶、叶鞘样品取自7天苗龄水稻苗,YP为幼穗(数字代表幼穗的长度,单位为厘米)。
图8为实施例2中回复表型分析;
A为WT、Oslr1及不同OsLR1转基因回复互补株系C4、C7植株7天苗龄的表型(标尺为5厘米);B为相应株系的株高统计;C为相应株系的根长统计;误差用SD用表示(统计的个体数目大于30),字母a、b、c代表具有显著差异,P<0.01(单因素方差分析SLD检验)。
图9为实施例2中过表达表型分析;
A为WT与不同OsLR1转基因过表达株系OE5、OE10和OE14植株7天苗龄的表型(标尺为5厘米);B为相应株系的株高统计;C为相应株系的根长统计;误差用SD用表示(统计的个体数目大于30),***P<0.001(T检验)。
图10为实施例2中OsLR1过表达转基因植株的根长度与OsLR1以及蛋白OsLR1的表达量呈正相关;
A为qRT-PCR检测7天苗龄WT与OsLR1过表达转基因株系OE5、OE10和OE14根中OsLR1的表达量;B为检测7天苗龄的WT与OsLR1过表达转基因株系OE5、OE10和OE14根中OsLR1蛋白的表达量。
图11为实施例2中RNAi干涉表型分析;
A为WT与不同OsLR1转基因沉默株系Ri5、Ri7和Ri49植株7天苗龄的表型(标尺为5厘米);B为相应株系的株高统计;C为相应株系的根长统计;误差用SD用表示(统计的个体数目大于30),***P<0.001(T检验)。
图12为实施例2中OsLR1抑制表达转基因株系的根长度与OsLR1以及蛋白OsLR1的表达量呈正相关;
A为qRT-PCR检测7天苗龄WT与OsLR1抑制表达转基因株系Ri5、Ri7和Ri49根中OsLR1的表达量;B为检测7天苗龄的WT与OsLR1抑制表达转基因株系Ri5、Ri7和Ri49根中OsLR1蛋白的表达量。
具体实施方式
下面结合具体实施例对本发明进行进一步描述,但本发明的保护范围并不仅限于此:
水培培养基的成分如下
以下实施例中的基因表达量检测结果,如无特殊说明,均是以野生型植株秀水134的目的基因表达量为1,其它植株的目的基因表达量与野生型植株的目的基因表达量进行比较。
实施例1、突变体植株Oslr1的发现
一、突变体植株Oslr1的获得和形态观察
1、突变体植株Oslr1的获得
构建秀水134水稻甲基磺酸乙酯(Ethyl Methylsulfonate,EMS)突变体库,从中发现了一株根变长的突变体植株Oslr1(Oryza sativa longer root1)。
突变体库的构建方式为:用浓度为0.8%的甲基磺酸乙酯对秀水134进行12小时的浸泡处理,用清水洗净后田间播种,成熟后单株收种构成突变体库。
2、突变体植株Oslr1的形态观察
(1)根长
Oslr1的表型是在幼苗期,突变体Oslr1萌发后生长7天,与野生型(WT)相比其株高没有显著变化,但其根长显著增长(见图1)。
(2)根尖的伸长区细胞观察
用扫描电镜的方法对幼苗期的野生型秀水134(WT)及突变体Oslr1根尖的伸长区细胞进行观察,统计结果表明,变体Oslr1该区域的细胞比野生型秀水134相同部位细胞的长度显著减小,但是宽度没有改变(见图2)。
3、突变体植株Oslr1的子代的形态观察
将Oslr1种子繁殖后,较野生型其后代持续出现根变长的表型。这些结果表明突变体Oslr1长根的表型是可稳定遗传的。
4、突变体植株Oslr1对生长素敏感性的分析
以野生型秀水134(WT)及突变体Oslr1为材料,将种子经破休眠催芽处理后,生长素IAA处理7天后对根长进行统计。结果表明,突变体Oslr1在IAA处理下根长的变短的程度要比野生型大,尤其在高浓度IAA反应更为明显(见图3),用不同浓度的生长素类似物NAA也得到了类似的结果(见图4)。这两项生理实验均表明突变体Oslr1对生长素的敏感性增强。
二、基因OsLR1的发现
以突变体Oslr1纯合体为母本,XS134为父本杂交获得F1,发现F1植株根长介于野生型和突变体之间,说明该Oslr1为半显性突变。统计F2代根性状分离结果,发现在131株F2群体中,33株为长根表型(Oslr1表型),68株的根长度介于野生型和突变体表型之间,30株为短根表型(XS134表型),三种根长表型植株的比例大约为1:2:1(χ2=0.169<χ2 0.05(2)=5.99),该分离规律符合孟德尔的单基因半显性遗传分离规律,说明Oslr1为单基因突变导致的。接着我们通过全基因组二代测序,分析得到了该突变基因的侯选区域。对区域内的侯选基因进行进一步分析,得到候选突变基因为01g0816600,我们将其命名为OsLR1。根据注释,该基因编码一个含LRR结构域、激酶域、JM功能域及CT结构域的蛋白。扩增该基因测序,测序结果表明该基因在突变体中存在三个突变位点,分别位于该基因的第3、5、10外显子中。
第一个突变位点位于该基因基因组序列(从启始密码子开始计数)的406位,突变方式为将胞嘧啶(cytimidine)突变为鸟嘌呤(guanine),记为C406G,并且此突变造成了该基因所编码蛋白质的第82位的苏氨酸(threonine)变为丝氨酸(serine),记为T82S。第二个突变位点位于该基因基因组序列的790位,突变方式为将鸟嘌呤(guanine)突变为腺嘌呤(adenine),记为G790A,并且此突变造成该基因所编码蛋白质的第144位的缬氨酸(valine)突变为异亮氨酸(isoleucine),记为V144I。第三个突变位点位于该基因基因组序列的1610位,突变方式为将鸟嘌呤(guanine)突变为胞嘧啶(cytimidine),记为G1610C,此突变为同义突变,没有造成该基因所编码蛋白质氨基酸位点的突变(见图5)。
突变基因01g0816600的编码区序列如SEQ ID NO:2所示;基因组序列如SEQ IDNO:3所示。该基因编码的蛋白质的氨基酸序列如SEQ ID No:1所示。
三、基因OsLR1的功能鉴定
1植物激素对基因OsLR1表达的调节
以野生型秀水134为材料,将种子经破休眠催芽处理后,在正常营养液中培养7天,使用不同的植物激素如2,4D、6-BA、ABA、BL、GA3、GR24、IAA对水稻幼苗处理1小时后取样,提取根部RNA并进行逆转录后获得相应的cDNA,用qRT-PCR方法确定OsLR1在不同激素处理下的表达差异。结果表明,基因OsLR1可被所检测的激素所诱导,但是对于生长素IAA的响应最为明显(见图6)。
2基因OsLR1的表达模式分析
以野生型秀水134水稻为材料,在正常营养液中培养7天,取其根、茎、叶及叶鞘,然后待植株将要抽穗时候,取其不同发育水平的幼穗。所有样品于液氮中速冻并研磨,分别抽提总RNA并进行逆转录获得相应的cDNA。用实时定量PCR方法检测OsLR1在各组织中的表达。结果表明,OsLR1基因在我们所检测的不同组织中都有表达,在根中表达量较高(见图7)。
实施例2、转OsLR1基因水稻材料的获得
一、互补重组载体的构建
提取秀水134水稻的gDNA,以gDNA为模板进行PCR扩增制备OsLR1的基因组序列(SEQ ID NO:3所示DNA)。PCR扩增的引物为:
F 5'-ACGCGTCGACATCCGATGGAGCGAGCGGTACA-3';
R 5'-CGGGGTACCGTACGCACGCAAGATTACGG-3'。
将获得PCR产物连入载体pCAMBIA1300NH-sGFP(购自Invitrogen)的SalI和KpnI酶切位点之问,得到含有OsLR1的互补表达载体。OsLR1的互补表达载体经测序检验序列正确。
二、OsLR1互补表达水稻的获得
将步骤一构建的OsLR1的互补表达载体转入农杆菌EHA105,用于转化突变体OsLR1,具体步骤如下:
(1)取培养好的菌液500μl于1.5ml离心管中,室温,4000rmp,离心2分钟,去上清。用含200μmol/L乙酰丁香酮的30ml AAM感菌液制成悬浮液,菌液终浓度OD600为0.01;将长到一定大小的80~120个水稻愈伤组织挑出,放入农杆菌悬浮液,在水平摇床上摇晃5分钟;
(2)将愈伤组织取出,置于无菌的滤纸上沥干30到40分钟;
(3)将愈伤组织置于有一张无菌滤纸的共培养基上,25℃暗培养3天;
(4)上述的愈伤组织取出,再用无菌水清洗5到6次,其间需不停的振荡。用含300mg/L羧苄青霉素钠(Carb)的无菌水清洗2遍,每次在水平摇床上摇晃30分钟。最后置于无菌滤纸上沥干2小时;
(5)将晾干的愈伤转入含300mg/L羧苄青霉素钠和相应筛选压力的选择培养基上进行第一轮选择,28℃,光照培养14天;
(6)将长有抗性愈伤的初始愈伤转到含300mg/L羧苄青霉素钠和相应筛选压力的培养基上进行第二轮选择,28℃,光照培养,直到颗粒性的抗性愈伤组织长出(约14天左右);
(7)挑取从不同愈伤来的颜色鲜黄的抗性愈伤3到5颗,移入装有分化培养基的塑料广口瓶中,用封口膜封好,放入恒温(25℃)培养室中(光周期:16小时光照),等待分化成苗(约40天);
(8)待苗长至3厘米左右,用剪刀从苗基部剪去老根和愈伤组织,放入生根培养基中壮苗(约1周)。将根部和茎叶分化得较完好的苗从试管挑出(苗长至试管顶部,就要及时开盖),打开封口膜,加入适量蒸馏水或无菌水(防止培养基长菌),炼苗2到3天,然后洗去琼脂,移栽到温室的水培条件中生长。利用抗赤霉素基因的引物检测转基因植物,引物序列如下:
5'-ATGAAAAAGCCTGAACTCACC-3';
5'-CTATTCCTTTGCCCTCGGACG-3',
得到的12个T1代转移基因阳性水稻株系,选择其中两个代表性株系(C4,C7)进行详细的表型分析,结果表明与突变体Oslr1相比,两个互补转基因株系的株高没有显著变化,其根长显著缩短(见图8)。这说明确实是基因OsLR1的突变导致了突变体Oslr1的长根表型。
三、OsLR1过表达重组载体的构建
提取秀水134水稻的mRNA并逆转录为cDNA,以此cDNA为模板进行PCR扩增制备OsLR1序列(SEQ ID NO:2所示DNA)。PCR扩增的引物如下:
5'-ACGGGGGACGAGCTCATGAAGAATTGGCAGAATGAA-3';
5'-GACTCTAGAGGATCCCTTTGGTTCAACAACTGA-3',
将获得PCR产物利用重组克隆的方法插入到本实验室改造的载体pCAMBIA1300-35S-EGFP(Chen et al.2015)的SacI和BamHI酶切位点之间,得到OsLR1过表达载体。OsLR1过表达载体经过测序检验正确。
四、OsLR1过表达水稻的获得
将步骤三构建的OsLR1过表达载体转入农杆菌EHA105,用于转化水稻秀水134,得到32株阳性转基因植株,具体步骤同实施例2的步骤二。
挑选三株代表性的过表达植株(OE5,OE10,OE14)。与野生型相比,转基因株系的株高没有显著变化,其根长显著增加(见图9)。
用实时定量PCR的方法对水稻秀水134与OsLR1过表达转基因植株中(OE5,OE10,OE14)中内源OsLR1相对表达量进行分析(引物为:AATCTGAAGGGTAAAGGAG;GATCCAAATAACCCAGTGTG),结果表明,OsLR1的过表达转基因株系的主根长度与OsLR1表达量正相关(见图10A);此外,对这些转基因株系中OsLR1蛋白的检测结果也表明,植株中OsLR1蛋白的表达量与其主根的长度正相关(见图10B)。
五、OsLR1抑制表达重组载体的构建
提取水稻秀水134的mRNA并逆转录为cDNA,以此长cDNA为模板进行PCR扩增制备OsLR1的部分DNA序列:
CACCGCGATGGTTTTCAACATTAACATCCTTGACTACCTTATTTATGGATAGTGATCATCTTACTGGAACAATCCCCAGTGCCCTGTTCAGTTTCCCGCAATTGCAGCAAATATCATTAGCTAAGAATTCATTCAGTGGGGAACTTAATATGAGCAGTAACATCAGTTCACTACTGCGGGTTGTTAATTTGACAAACAATCAGATCTTCAATGCTGAAGTTGACCCAAGCT。
PCR扩增的引物如下:
5'-CACCGCGATGGTTTTCAACA-3';
5'-AGCTTGGGTCAACTTCAGCA-3',
将克隆到的PCR产物与T载体(购自TAKARA公司)连接,连接好的质粒分别用PstI、BamH I以及Pst I、Sal I酶切得到两片段;两片段一同连入本实验室改造的pBSSK-in载体(Chen et al.2015),分两步进行。pBSSK-in先用Pst I、BamH I酶切,连上一个片段后,再Nsi I、Sal I酶切,连另一片段;最后用Sac I、Sal I将两片段以及intron切下,连入相同酶切的植物双元载体pCAMBIA1300-35S-EGFP中,得到OsLR1抑制表达载体。OsLR1抑制表达载体经过测序检验正确。
六、OsLR1抑制表达水稻的获得
将步骤五构建的OsLR1抑制表达载体转入农杆菌EHA105,用于转化水稻秀水134,得到53株阳性转基因植株。具体步骤同实施例2的步骤二。
在得到的T1代转移基因水稻中,挑选三株代表性过表达植株(Ri5,Ri7,Ri49)进行表型分析,与野生型相比,与野生型相比,转基因株系的株高没有显著变化,其根长显著缩短(见图11)。
用实时定量PCR的方法对秀水134与OsLR1抑制表达转基因植株(Ri5,Ri7,Ri49)中内源OsLR1相对表达量进行分析(引物为:AATCTGAAGGGTAAAGGAG;GATCCAAATAACCCAGTGTG),结果表明,基因OsLR1的过表达转基因株系的主根长度与OsLR1表达量正相关(见图12A);此外,对这些转基因株系中OsLR1蛋白的检测结果也表明,植株中OsLR1蛋白的表达量与其主根的长度正相关(见图12B)。
综合以上的结果,在Oslr1突变体中恢复表达基因OsLR1可以抑制其长根表型,OsLR1的过量表达促使水稻根伸长,而抑制OsLR1的表达量后,会使抑制水稻根伸长,这表明可以通过调控OsLR1控制水稻根的生长发育。
最后,还需要注意的是,以上列举的仅是本发明的若干个具体实施例。显然,本发明不限于以上实施例,还可以有许多变形。本领域的普通技术人员能从本发明公开的内容直接导出或联想到的所有变形,均应认为是本发明的保护范围。
序列表
<110> 浙江大学
<120> 调控植物根发育的蛋白及其编码基因和应用
<160> 3
<170> SIPOSequenceListing 1.0
<210> 1
<211> 926
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 1
Met Lys Asn Trp Gln Asn Glu Pro Gln Ser Trp Met Gly Ser Thr Asp
1 5 10 15
Pro Cys Thr Ser Trp Asp Gly Ile Ser Cys Ser Asn Gly Arg Val Thr
20 25 30
Glu Met Arg Leu Ser Gly Ile Asn Leu Gln Gly Thr Leu Ser Asn Ala
35 40 45
Ile Asp Gln Leu Ser Ser Leu Thr Tyr Leu Asp Leu Ser Asn Asn Leu
50 55 60
Asn Leu Gly Gly Pro Leu Pro Pro Ser Ile Val Asn Leu Lys Gln Leu
65 70 75 80
Thr Thr Leu Ile Leu Leu Gly Cys Ser Phe Thr Gly Asp Ile Pro Glu
85 90 95
Gln Ile Gly Ala Leu Arg Gln Leu Thr Phe Leu Ala Leu Asn Ser Asn
100 105 110
Lys Phe Thr Gly Gly Ile Pro Pro Thr Leu Gly Leu Leu Ser Lys Leu
115 120 125
Phe Trp Leu Asp Leu Ser Asp Asn Gln Leu Ser Gly Lys Ile Pro Val
130 135 140
Ser Ser Gly Ser Asn Pro Gly Leu Asp Gln Leu Val Asn Ala Glu His
145 150 155 160
Phe His Phe Ser Glu Asn Gln Leu Thr Gly Pro Ile Asp Glu Lys Leu
165 170 175
Phe Ser Glu Lys Met Asn Leu Ile His Val Ile Phe Asp Asn Asn Asn
180 185 190
Phe Thr Gly Pro Ile Pro Gly Ser Leu Gly Arg Val Ser Ser Ile Gln
195 200 205
Ile Ile Arg Leu Asp His Asn Gln Phe Ser Gly Pro Val Pro Gly Ser
210 215 220
Ile Ala Asn Leu Ser Arg Leu Met Glu Leu Ser Leu Ala Ser Asn Gln
225 230 235 240
Leu Asn Gly Thr Val Pro Asp Leu Thr Ser Ala Asn Ala Leu Thr Tyr
245 250 255
Val Asp Leu Ser Asn Asn Asn Phe Met Ser Ser Pro Ala Pro Arg Trp
260 265 270
Phe Ser Thr Leu Thr Ser Leu Thr Thr Leu Phe Met Asp Ser Asp His
275 280 285
Leu Thr Gly Thr Ile Pro Ser Ala Leu Phe Ser Phe Pro Gln Leu Gln
290 295 300
Gln Ile Ser Leu Ala Lys Asn Ser Phe Ser Gly Glu Leu Asn Met Ser
305 310 315 320
Ser Asn Ile Ser Ser Leu Leu Arg Val Val Asn Leu Thr Asn Asn Gln
325 330 335
Ile Phe Asn Ala Glu Val Asp Pro Ser Tyr Thr Gly Ser Leu Ile Leu
340 345 350
Ser Gly Asn Leu Ile Cys Phe Asn Asn Ile Ser Phe Cys Thr Leu Lys
355 360 365
Gln Lys Gln Gln Val Pro Tyr Ser Thr Asn Leu Gly Pro Cys Gly Ala
370 375 380
Ile Ser Cys Pro Thr Asp Gln Ser Ala Asn Pro Val Ala Ser Gln Asn
385 390 395 400
Cys Ala Cys Ala Ser Pro Phe Gln Gly Leu Met Ile Phe Arg Ala Pro
405 410 415
Ala Phe Ser Asp Val Thr Asn Pro Lys Ser Phe Gln Pro Leu Glu Phe
420 425 430
Thr Leu Val Gln Asn Leu Ser Leu Ala Pro Gly Ser Val Ala Ile Ser
435 440 445
Asn Val Glu Phe Ser Pro Gly Glu Pro Leu Thr Phe Thr Val Lys Val
450 455 460
Phe Pro Glu Ser Gly Thr Ser Phe Asn His Ser Glu Val Ile Arg Ile
465 470 475 480
Ser Ser Ser Leu Val Asn Gln Thr Tyr Lys Ala Pro Ala Tyr Phe Gly
485 490 495
Pro Tyr Ser Phe Ile Ala Ser Thr Tyr Phe Ala Ser Pro Ser Gly Lys
500 505 510
Arg Ser Ser Met Gly Lys Gly Ala Ile Ile Gly Ile Ala Val Ala Gly
515 520 525
Phe Leu Leu Leu Val Gly Leu Ile Leu Val Ala Met Tyr Ala Leu Arg
530 535 540
Gln Lys Lys Ile Ala Lys Glu Ala Val Glu Arg Thr Thr Asn Pro Phe
545 550 555 560
Ala Ser Trp Gly Gln Gly Gly Lys Asp Asn Gly Asp Val Pro Gln Leu
565 570 575
Lys Gly Ala Arg Tyr Phe Ala Phe Glu Glu Leu Lys Arg Cys Thr Asn
580 585 590
Asn Phe Ser Glu Thr Gln Glu Ile Gly Ser Gly Gly Tyr Gly Lys Val
595 600 605
Tyr Lys Gly Met Leu Ala Asn Gly Gln Met Ala Ala Ile Lys Arg Ala
610 615 620
Gln Gln Gly Ser Met Gln Gly Ala Ala Glu Phe Lys Asn Glu Ile Glu
625 630 635 640
Leu Leu Ser Arg Val His His Lys Asn Leu Val Ser Leu Val Gly Phe
645 650 655
Cys Tyr Glu Gln Gly Glu Gln Met Leu Val Tyr Glu Tyr Ile Pro Asn
660 665 670
Gly Thr Leu Arg Glu Asn Leu Lys Gly Lys Gly Gly Met His Leu Asp
675 680 685
Trp Lys Lys Arg Leu Gln Ile Ala Val Gly Ser Ala Lys Gly Leu Ala
690 695 700
Tyr Leu His Glu Leu Ala Asp Pro Pro Ile Ile His Arg Asp Ile Lys
705 710 715 720
Ser Thr Asn Ile Leu Leu Asp Glu Ser Leu Asn Ala Lys Val Ala Asp
725 730 735
Phe Gly Leu Ser Lys Leu Val Ser Asp Thr Lys Lys Gly His Val Ser
740 745 750
Thr Gln Val Lys Gly Thr Leu Gly Tyr Leu Asp Pro Glu Tyr Tyr Met
755 760 765
Thr Gln Gln Leu Ser Glu Lys Ser Asp Val Tyr Ser Phe Gly Val Val
770 775 780
Met Leu Glu Leu Ile Thr Ser Arg Gln Pro Ile Glu Lys Gly Thr Tyr
785 790 795 800
Ile Val Arg Glu Ile Arg Thr Ala Ile Asp Gln Tyr Asp Gln Glu Tyr
805 810 815
Tyr Gly Leu Lys Ser Leu Ile Asp Pro Thr Ile Arg Asp Ser Ala Lys
820 825 830
Met Val Gly Phe Arg Arg Phe Val Gln Leu Ala Met Glu Cys Val Glu
835 840 845
Glu Ser Ala Ala Asp Arg Pro Thr Met Asn Asp Val Val Lys Glu Leu
850 855 860
Glu Ile Ile Ile Gln Asn Glu Gly Ala Gln Leu Leu Asn Ser Ala Ser
865 870 875 880
Leu Ser Ala Gln Gln Phe Gly Tyr Ala Lys Gly Arg Asp Pro Asp Pro
885 890 895
Tyr Gly Asp His Val Pro Ile Asn Asp Asp Ser Ser Ser Ser Ala Phe
900 905 910
Asp Tyr Asn Ser Val Tyr Ser Tyr Ser Val Val Glu Pro Lys
915 920 925
<210> 2
<211> 2781
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 2
atgaagaatt ggcagaatga accgcaaagc tggatgggat cgactgatcc ctgcacctcc 60
tgggatggaa tttcctgttc caatgggagg gtgacagaaa tgagattatc aggcattaat 120
ctgcaaggca cattaagcaa tgcaatagac caactttctt ctttgacata tctggatctg 180
tctaacaacc taaatcttgg aggtccactt cctccaagca ttgtgaatct gaagcagctc 240
acaactctga ttttacttgg atgcagtttc actggtgata ttccagagca gatcggagca 300
ttaaggcaac tcacattcct ggccctgaac tcaaataagt tcactggtgg aatcccccca 360
acacttggcc ttctctctaa gcttttttgg ttggacttgt cagacaatca gctgtcgggg 420
aaaataccgg tttcatcagg ttcaaaccca gggctggatc agcttgttaa tgcagaacat 480
ttccatttca gtgagaacca gttgacaggc ccaattgacg aaaagctttt cagtgaaaaa 540
atgaacctta tacacgtgat atttgacaac aataacttta ctggaccaat cccaggatct 600
ctcgggagag tctcatcaat tcaaattatc cgactagatc ataaccaatt cagtggtcca 660
gttccaggaa gtatagcaaa cctatctcgt ctgatggaac tgagcttagc aagcaaccaa 720
ctgaatggga cagtgccaga cctcaccagt gcaaatgctc tcacttatgt ggacctaagc 780
aacaataact tcatgagctc accagcaccg cgatggtttt caacattaac atccttgact 840
accttattta tggatagtga tcatcttact ggaacaatcc ccagtgccct gttcagtttc 900
ccgcaattgc agcaaatatc attagctaag aattcattca gtggggaact taatatgagc 960
agtaacatca gttcactact gcgggttgtt aatttgacaa acaatcagat cttcaatgct 1020
gaagttgacc caagctacac cggcagcctc atactatcag gcaatcttat atgcttcaac 1080
aatatcagtt tctgcacact caagcaaaag cagcaagtgc catactcaac aaacctaggt 1140
ccatgtggtg ccatttcatg ccccactgac cagtcagcaa atccggtggc ttcacagaac 1200
tgtgcttgcg ccagcccctt ccagggtttg atgatcttcc gagcaccagc cttctctgat 1260
gtgaccaatc ccaagtcatt ccaacctttg gaatttactc ttgttcagaa ccttagccta 1320
gctccaggat cagttgccat ttccaatgtt gagttcagtc caggggagcc actaacattc 1380
acagtgaagg tttttccaga gagcggaaca agcttcaatc actcagaggt tatcagaatc 1440
agttcttctt tggtcaacca aacctacaaa gctccagcat attttggacc atatagcttc 1500
atagcaagca catattttgc aagccccagt ggtaaaaggt cgtcaatggg caaaggtgca 1560
ataatcggaa tagcggttgc tggttttctc cttcttgttg gccttatcct agtagcaatg 1620
tatgctctaa ggcagaaaaa aatagctaag gaggcagtag aacgaactac taatcctttt 1680
gcatcatggg gacaaggtgg taaagataat ggagatgtgc cacagctgaa gggagcaaga 1740
tactttgcat ttgaggaact gaagaggtgc acaaacaatt tctcagaaac ccaagaaata 1800
ggatcaggag gatatgggaa ggtgtacaaa ggaatgctcg caaatgggca aatggctgca 1860
ataaaacgcg cacagcaagg atctatgcaa ggtgcagctg aatttaagaa tgagatagaa 1920
ctactttcca gggttcatca caagaacctg gtgagcttag taggtttctg ctatgaacaa 1980
ggggagcaga tgttggttta tgagtacatt cctaatggga ccttaaggga gaatctgaag 2040
ggtaaaggag gaatgcactt ggattggaag aaacgccttc aaattgcagt tggctctgct 2100
aaaggtctag cgtatcttca tgaacttgct gatccaccaa ttatccatag agatatcaaa 2160
tcaaccaata tccttctgga tgaaagtcta aatgcaaagg ttgctgattt tggtctttca 2220
aagctggtat ctgacacaaa aaagggccat gtttctaccc aagtgaaggg cacactgggt 2280
tatttggatc ctgaatacta catgactcag cagctatctg agaagagtga tgtatatagc 2340
tttggagttg tcatgctaga actgataacg tccaggcagc ccatagagaa aggcacatat 2400
attgtccgtg agatcaggac agcaatagat cagtatgacc aagagtatta cggattgaag 2460
agcctaattg atccaacaat ccgggattca gcaaaaatgg ttggctttag gagatttgtg 2520
cagttggcta tggaatgtgt agaggagtct gctgctgacc gcccgacaat gaatgatgtg 2580
gtgaaggaac ttgagattat aatacaaaat gaaggggcac agttgctaaa ctcagcatct 2640
ttatcggctc agcagtttgg atatgcaaag ggtagagatc cagatcctta tggagatcat 2700
gtgcctatca atgatgacag cagcagcagt gcttttgatt acaacagtgt atattcatac 2760
tcagttgttg aaccaaagta g 2781
<210> 3
<211> 8632
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 3
atccgatgga gcgagcggta caccacagtg ctacaccact ccggatcgat gcggcggcgt 60
tcgaattctt ttttcctttt ttttcttttt ggaaagattt tgaagttccc gggaagaccg 120
gggtgactgg gatcactgta aatcaggaat ttcctttctc ccacacgggc cattcgggaa 180
ttgggcccac tggtcctggg ccagtttctc ttcatcttct cgtctttgag gaataagtcc 240
aatttgactc cctcaaatat agttctaatc taatttgtgt ccctcaaccg taatattgga 300
tacctcgatc tctaaactat taaaactgat gcaaaatgac acccttggtg gttttgaaga 360
ctgttttcct gatgtggtgc gttgacttga tccaaccgac tgggtcagca tgtggggccc 420
atatgccaac ttctctttat tcttctccat ttctttctcc ctctcctttc ccacacacgc 480
acacaacgac tcgagggtgg accgcagtag cgcggcccaa ggcggagcgg cgtctaggtg 540
ggcccaatgc atagtgatgg cacagactgg aggcggagga gtggcggcat ggcctcgagg 600
ctagaggcgg agcgatggtg tggcctcatc cttcttgcag cccttccatg ccaagctcga 660
cagtggccat gtgagccaac cggtggcggt ggtgctagag aagaccggcg gggagatggg 720
agagccggtg gaggccagcc gctagagacg gtggtggcct gtgggcgccg ctcccttcca 780
ccggctggag ggggtcagga tttgaatttt gtatgttaga ttatagggat tttaaaattt 840
aaatttcatt taaaatatga acaaaatgat ttagctaagt gttatcaaca taatcgtaga 900
tttatatatc attggatttc tgaagctagg aatatccggc ttcaagaaat cttgaaaagg 960
tttagtatct actccatcca ttttatatta taattcgttt tgattttttt cctagtcaaa 1020
ctttttttaa gtttgaccga gtttatagaa aaatatatta gtatttttaa tacaaaacag 1080
acatattatc aaaatatatt caatgctcga tttgatgaaa ctaatttagt gttttagatg 1140
ttactacatt agtttcatca aatcgaacgt tgaatatatt ttgataatat atctgttttg 1200
tattgaaaat actaatatat ttttctataa actcggtcaa acttaataaa gtttgactag 1260
gaaaaaagtc aaaacaaatt ataatataaa acggatggag tagatactaa acctgttcaa 1320
gatttcttga agctggatat tcctaacttc ggaaatccaa tgatatataa atttacgatt 1380
acgttgataa cacttagcta aatcattttg ttcatatttt aaataaaatt aaaaatttaa 1440
gatccctata atctaacata caatcttcaa aatccactat ggatctgagt tcgggagccc 1500
tgccaaacat gttctaagtt gccaacacga aacgttcctt cagttttgga tgcaagattc 1560
tgaattgatc ttacgcaatc tccagttgtc aaattagacc gagttggagg cggcgagcaa 1620
gcgcagctac cgtgctgttg cttacgttga agccacaacc aacgtgatca agagcatatt 1680
acagttatat attttccaaa caaatggcaa tttgaagcca catccaacgt gatctaaggg 1740
catattacag tatagtattc tccaaacaaa ttgcaagcaa aggatcctca ccagaagagc 1800
ctgaactgtt tgttcgcaat cactttcgta ggatcataca ttcacactcc gtttgtcaaa 1860
agctttcgag gccaatgatt ggcgcatata aaaataagaa gcaagcaact gctctggaca 1920
tactaataat ctgaaattat ttcagtaatt ctatctcagt ttgtcctctc ggggaagatg 1980
atatcatgca tgtagtaact tgtgaactcc atattccttt cagtaaatac tttcagctgc 2040
agaatatcaa ccagcttatc attagactaa actccgaaga acatttcagc agttaagttg 2100
ccaagtaaac ggcacatgtg acatgttgtt cgctttgacc gattactccc acagaagaaa 2160
aaagaagaca tttttctcac accagtacat ttcatacgag cacgattgtc accaagccat 2220
ctaactcaac tgcgtcaggc ttatattttc aattagaaca atggacgaaa caaatgcgaa 2280
tgcattatat cagctggcca gtggaaggaa gcaagaggca actgggtgcc attgagtttt 2340
ggcctcatgc agaaggatct ttttcgtgtg tcgccggctc cgccatttgc tgtgtatata 2400
gtgcacctaa tgttacctga tgaagctgtt aacaatacct acaaaaggaa gaagccggcc 2460
ggccatgtat taacagaaat tggccttcct gccccactga atgtgaccgt gaattgcata 2520
atttgcgttg catgatgtca caccgaatca tttttcggtg tttactttgt tggatccagc 2580
ggagaacata atcctctaca aaatcttggc ttcctgctga cataacccaa agctttcact 2640
gtccttgaga agcctgcaaa acaagatttt atattcatgt tctattgctt acaatcagat 2700
ttctagagct ttttctgaat caaatttctg gaagatgaaa atttgaaaca agaccaacca 2760
tggaagcatt cacaggcagt tacttaggat aatatgtcga tccattttct gatgccatat 2820
cccatcagaa aggttggcat tggcaggcac tttgacttag cctgaaaact gttgtttaca 2880
gctccaacgg tcccaactat acaagggaaa acagatatga taagaacagc ctgagcgatt 2940
aacacagtgt ctgacatgca ggtacaggac aaaaatatcc aatcaatggc cgtatcaaaa 3000
aaataaatta agtaaaagca cagaatatac tgaccaaaaa cgtataaaag ccaagaacgt 3060
ccaaaccaag aagttctgac tggatcaaaa cccagaaggt gttcccagat aaggctccat 3120
agataaaaca ggcaagaagt aactgcaagt aaacggttta ttaaatcatt tcacattttg 3180
atcagcaact gagctcttca gaggtttctt ttaattaagc ctcccactac aacgcataca 3240
tgcattttct atgtgcatgt tgcttggacc tggtaagcat tctcataggc aatgtttgat 3300
aagaattccc ttgaatcaag ttatataagg ttgtcttgtt tgattagcta ttttctttgc 3360
ctggttgcca gcctcaagga gggagacgcc atttggcatc tactcgagct tgttcactgc 3420
acagtgctct tcgacaagaa caatcagcag attttgctat cctccggtgt gaatttgatt 3480
tgcgtggagg gtgacaccgt gtggtacgat ggtggagatg cagtggctgc tacttctgtt 3540
catgctcttg gtgagcctac ggctaagctt cagtcagaca aatcctcaag atggtaagtc 3600
tttgatcttt ctgcttttca ataggttaca ggaagaaagg tgacatcttt tccaggattt 3660
gtcactcaga ctggccattg acctgaaaac aaggtgagta ggattaggtc tgccagaagt 3720
acagcaagag aagaatctat ttgtggagca tgaatgacta gatgctacag aaaaacaaaa 3780
tccttaaaga tagccttctg ccatcaatac tatgttctcc ttgggtagtt atgcaacaaa 3840
ttgtgatcat ttggttattt ccatctaata acctaggaag atttttttta tgaccaaatt 3900
catagtgtca tgtacaaaga atagtttccc atgatgaaaa ggggagctgc aacagctata 3960
tctcatgcac gcacgatcaa taattttaga gagatgactc cttcctttcc tgcaggagcg 4020
aatccatgaa aaagccatgt gatgcacagg acaggcggaa cttttgaatc ccattaattt 4080
tagctaatgt tagaatgtaa acatagtatt atatccatga ttcatacagg acacctggta 4140
atttttgttt ttagattcgc ccctgctttc ctggaccata ctttcctgtg aaattttcca 4200
tgatgatgat aaactaatgg atgatattga cctaataggc caagatgact tggatcaacc 4260
actggagttg ttgacacatt ttgcttgact aacatttgtt ccatttattc tacagtttct 4320
gcactccagg cattgatgaa gaattggcag aatgaaccgc aaagctggat gggatcgact 4380
gatccctgca cctcctggga tggaatttcc tgttccaatg ggagggtgac agaaatgtga 4440
gtgatcaact tcgattcatc atatttgtac ttatttttga gacaagaact acaccgaaac 4500
ctggtattta caggagatta tcaggcatta atctgcaagg cacattaagc aatgcaatag 4560
accaactttc ttctttgaca tatctgtaag tccctgctta ctgagttttg aacaaactac 4620
cacactgcaa aatgctttat gctcctgatg taaaataaac tacatccagg gatctgtcta 4680
acaacctaaa tcttggaggt ccacttcctc caagcattgt gaatctgaag cagctcacaa 4740
ctctgtaaga acagcatgtc actataagaa aacatttctt cctttttggt ctgtggagag 4800
gactaagaaa aataggggaa aacactgatt tatatcttgg ttttgtcagg attttacttg 4860
gatgcagttt cactggtgat attccagagc agatcggagc attaaggcaa ctcacattcc 4920
tgtaagaaac aactcaaacg tgggcagata tattgaatgg acatcaatct cttcacgata 4980
tgatttactt tagcctaaga aatgctctca tgcagggccc tgaactcaaa taagttcact 5040
ggtggaatcc ccccaacact tggccttctc tctaagcttt tttggttgga cttgtcagac 5100
aatcagctgt cggggaaaat accggtttca tcaggttcaa acccagggct ggatcagctt 5160
gttaatgcag aacatttgta agttctcaaa tggtgtacca gttctagaac aaaagaatca 5220
agatatactg ctgcttaaat gtatggtatc ctcatatgac tcatacattt ccaatcaaca 5280
gccatttcag tgagaaccag ttgacaggcc caattgacga aaagcttttc agtgaaaaaa 5340
tgaaccttat acacgtgtaa gctgcacact gtttgtttgt gcttggtagt atcatcttgc 5400
catcacctca ctgccttcag taataattgt caggatattt gacaacaata actttactgg 5460
accaatccca ggatctctcg ggagagtctc atcaattcaa attatgtaag tacctcgtaa 5520
tatgatttac cctatgtttt gacccccttt gaacagtttc atgttttttt tttcctgcct 5580
attgcaacag ccgactagat cataaccaat tcagtggtcc agttccagga agtatagcaa 5640
acctatctcg tctgatggaa ctgtaagtga ttaccatcaa ggaacatcaa caaatatttc 5700
ttattcctgg aaacaacatt gcaaatattt gtttttgttt ttgcaggagc ttagcaagca 5760
accaactgaa tgggacagtg ccagacctca ccagtgcaaa tgctctcact tatgtgtaag 5820
gaagaaagta tatgagcatt taaggttaat gattctttga gttatgggaa attttcttcc 5880
tttttacagg gacctaagca acaataactt catgagctca ccagcaccgc gatggttttc 5940
aacattaaca tccttgacta ccttgtgagt tgtctgatga cgaaaaaaca gtgagtatca 6000
aatggactta gtaagataca tcacaattaa tgtttctatt cctgttgaca acttacagat 6060
ttatggatag tgatcatctt actggaacaa tccccagtgc cctgttcagt ttcccgcaat 6120
tgcagcaaat gtaatttcct catggatcat gctacagctc tctgatagca tctctagcta 6180
aaagtttaac ttcagtgaac ttatgccttc tgatgttata aattggaatg cagatcatta 6240
gctaagaatt cattcagtgg ggaacttaat atgagcagta acatcagttc actactgcgg 6300
gttgttaatt tgacaaacaa tcagatcttc aatgctgaag ttgacccaag ctacaccggc 6360
agcctcatgt aagcatcatc ctgtactgta taagggtgta caagtgagtg catgcattct 6420
cacccaacgt attctttcct tgtccagact atcaggcaat cttatatgct tcaacaatat 6480
cagtttctgc acactcaagc aaaagcagca agtgccatac tcaacaaacc taggtccatg 6540
tggtgccatt tcatgcccca ctgaccagtc agcaaatccg gtggcttcac agaactgtgc 6600
ttgcgccagc cccttccagg gtttgatgat cttccgagca ccagccttct ctgatgtgac 6660
caatcccaag tcattccaac ctttggaatt tactcttgtt cagaacctta gcctagctcc 6720
aggatcagtt gccatttcca atgttgagtt cagtccaggg gagccactaa cattcacagt 6780
gaaggttttt ccagagagcg gaacaagctt caatcactca gaggttatca gaatcagttc 6840
ttctttggtc aaccaaacct acaaagctcc agcatatttt ggaccatata gcttcatagc 6900
aagcacatat tttgcaagta tggcgttgct taggtctgca atttctttag tttgctttat 6960
tttcaaacat tacataactc attcatttca ggccccagtg gtaaaaggtc gtcaatgggc 7020
aaaggtgcaa taatcggaat agcggttgct ggttttctcc ttcttgttgg ccttatccta 7080
gtagcaatgt atgctctaag gcagaaaaaa atagctaagg aggcagtaga acgaactact 7140
aatccttttg gtaatgtgtt atttcatcat tcagtatcca tccaataggg aagtgttaag 7200
catgattgct aatctgtgct agtgaatgct ctggtgccac agcatcatgg ggacaaggtg 7260
gtaaagataa tggagatgtg ccacagctga agggagcaag atactttgca tttgaggaac 7320
tgaagaggtg cacaaacaat ttctcagaaa cccaagaaat aggatcagga ggatatggga 7380
aggtatattg gtgaatgaat ttccatgcta gtatagtcac aaaaagaaat aaggatttcc 7440
aaaagcaatt aatctattat cttttctgtt aaaaggtgta caaaggaatg ctcgcaaatg 7500
ggcaaatggc tgcaataaaa cgcgcacagc aaggatctat gcaaggtgca gctgaattta 7560
agaatgagat agaactactt tccagggttc atcacaagaa cctggtgagc ttagtaggtt 7620
tctgctatga acaaggggag cagatgttgg tttatgagta cattcctaat gggaccttaa 7680
gggagaatct gaagggtata cttttcaaac ttcatcacca ctactacccc catttataac 7740
ctatttacat aattaataca tgctacaaca aaaggacatg ggggttcatc cacatcttaa 7800
ctgcaggtaa aggaggaatg cacttggatt ggaagaaacg ccttcaaatt gcagttggct 7860
ctgctaaagg tctagcgtat cttcatgaac ttgctgatcc accaattatc catagagata 7920
tcaaatcaac caatatcctt ctggatgaaa gtctaaatgc aaaggttgct gattttggtc 7980
tttcaaagct ggtatctgac acaaaaaagg gccatgtttc tacccaagtg aagggcacac 8040
tggtatgtac ttctggaact tcaaaatgca taacttttat tgacaaaaaa atatttctag 8100
ttgattactc ttctgtgttg atatctaggg ttatttggat cctgaatact acatgactca 8160
gcagctatct gagaagagtg atgtatatag ctttggagtt gtcatgctag aactgataac 8220
gtccaggcag cccatagaga aaggcacata tattgtccgt gagatcagga cagcaataga 8280
tcagtatgac caagagtatt acggattgaa gagcctaatt gatccaacaa tccgggattc 8340
agcaaaaatg gttggcttta ggagatttgt gcagttggct atggaatgtg tagaggagtc 8400
tgctgctgac cgcccgacaa tgaatgatgt ggtgaaggaa cttgagatta taatacaaaa 8460
tgaaggggca cagttgctaa actcagcatc tttatcggct cagcagtttg gatatgcaaa 8520
gggtagagat ccagatcctt atggagatca tgtgcctatc aatgatgaca gcagcagcag 8580
tgcttttgat tacaacagtg tatattcata ctcagttgtt gaaccaaagt ag 8632
Claims (8)
1.一种蛋白,其特征是:为如下(a)或(b)所述的蛋白质:
(a)、蛋白质的氨基酸序列如SEQ ID No:1所示;
(b)、将SEQ ID No:1所示的蛋白质经过一个或几个氨基酸残基的取代和/或缺失和/或添加且与植物发育相关的由其衍生的蛋白质。
2.编码如权利要求1所述蛋白质的基因,其特征是:所述基因为如下1)~4)中的任一:
1)、基因的编码区的核苷酸序列如SEQ ID NO:2所示;
2)、基因的基因组的核苷酸序列如SEQ ID NO:3所示;
3)、为在严格条件下与1)或2)限定的DNA序列杂交且编码相同功能蛋白质的DNA分子;
4)、为与1)或2)或3)限定的DNA序列具有90%以上同源性,且编码相同功能蛋白质的DNA分子。
3.含有权利要求2所述基因或其反义基因的重组表达载体、表达盒、转基因细胞系或重组菌。
4.根据权利要求3所述的重组表达载体,其特征是:所述重组表达载体为如下(I)或(II):
(I)、将SEQ ID NO:2所示DNA分子插入质粒pCAMBIA1300-35S-EGFP的多克隆位点得到的重组表达载体Ⅰ--OsLR1过表达载体;
(II)、将SEQ ID NO:2所示基因DNA片段通过正向与反向连接到过渡载体pBSSK-in后插入质粒pCAMBIA1300-35S-EGFP得到的重组表达载体Ⅱ---OsLR1抑制表达载体。
5.如权利要求2所述基因的用途,其特征是用于构建转基因植物,所述转基因植物的根长度发生变长的改变。
6.根据权利要求5所述基因的用途,其特征是:
将重组表达载体Ⅰ导入目的植物中,得到长度变长的转基因植物。
7.根据权利要求5所述基因的用途,其特征是:
将重组表达载体Ⅱ导入目的植物中,得到长度变短的转基因植物。
8.根据权利要求5~7任一所述基因的用途,其特征是:所述植物为水稻。
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CN109628462A (zh) * | 2018-12-13 | 2019-04-16 | 宁波大学 | 一种水稻根系伸长控制基因OsKSR7及编码的蛋白质 |
CN111793119A (zh) * | 2019-04-04 | 2020-10-20 | 中国科学院遗传与发育生物学研究所 | 调控植物抗旱性的蛋白质及其编码基因与应用 |
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CN112280784B (zh) * | 2020-10-30 | 2022-12-02 | 宁波大学科学技术学院 | 一种水稻侧根发育控制基因OsLRD2、编码蛋白质及其应用 |
CN116622761A (zh) * | 2023-05-05 | 2023-08-22 | 青岛农业大学 | 玉米生长素响应蛋白iaa15的用途 |
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CN109628462A (zh) * | 2018-12-13 | 2019-04-16 | 宁波大学 | 一种水稻根系伸长控制基因OsKSR7及编码的蛋白质 |
CN109628462B (zh) * | 2018-12-13 | 2021-02-19 | 宁波大学 | 一种水稻根系伸长控制基因OsKSR7及编码的蛋白质 |
CN111793119A (zh) * | 2019-04-04 | 2020-10-20 | 中国科学院遗传与发育生物学研究所 | 调控植物抗旱性的蛋白质及其编码基因与应用 |
CN112280784B (zh) * | 2020-10-30 | 2022-12-02 | 宁波大学科学技术学院 | 一种水稻侧根发育控制基因OsLRD2、编码蛋白质及其应用 |
CN112779268A (zh) * | 2021-01-15 | 2021-05-11 | 南京农业大学 | 大豆GmCRF4a基因及其应用 |
CN112779268B (zh) * | 2021-01-15 | 2022-07-01 | 南京农业大学 | 大豆GmCRF4a基因及其应用 |
CN112899300A (zh) * | 2021-02-20 | 2021-06-04 | 浙江大学 | 水稻根分泌多肽pep1及其编码基因和应用 |
CN116622761A (zh) * | 2023-05-05 | 2023-08-22 | 青岛农业大学 | 玉米生长素响应蛋白iaa15的用途 |
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