CN111118028A - 一种白花虎眼万年青矮化多分蘖OtDWARF53基因及应用 - Google Patents
一种白花虎眼万年青矮化多分蘖OtDWARF53基因及应用 Download PDFInfo
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Abstract
本发明涉及植物基因工程技术领域，尤其涉及一种白花虎眼万年青矮化多分蘖OtDWARF53基因及应用。其核苷酸序列如SEO ID NO.1所示，该基因编码的蛋白的氨基酸序列如SEO ID NO.2所示；本发明在已经获得的白花虎眼万年青叶上珠芽的诱导和组织培养的发明专利的基础上，以及根据白花虎眼万年青三代和二代转录组联合分析的结果（3+2），从中分离出来了调控独脚金内酯的关键基因DWARF53（DWF53），并构建了组成型植物表达载体pHB+ OtDWARF53，将其转入烟草中，发现转基因烟草出现了植株矮化、茎尖产生多个顶芽、叶腋形成腋芽侧芽、外形更加丰满等表型，表明该基因能够使得植株矮化、及促进多分蘖多分枝的功能，具体是该基因在调节植株高度和促进腋芽侧芽发育等调控株型中的应用。
Description
技术领域
本发明涉及植物基因工程技术领域，尤其涉及一种白花虎眼万年青矮化多分蘖OtDWARF53基因及应用。
背景技术
高等植物株型形成是指在植物整个生长发育过程中植株形态相关器官的发生，尤其是指分枝、叶片和花器官的形成、形状与着生位置等。植物株型的形成过程主要受遗传与植物激素等内在因素的调控，同时还受光周期、温度、水肥等外界环境因素的影响。高等植物株型形成的分子机理是植物生长发育研究的基本科学问题，对于科研和生产均具有重要的价值和意义。
在农作物中，株型直接影响植株分蘖、以及有效穗数和穗粒数，是控制产量的核心要素，决定收获、产量和经济效益，同时也与抗逆性、生态适应能力和竞争能力密切相关；
不但农作物株型的改良具有重要的生产应用价值，而且在园林园艺植物中，植物的茎秆矮化、分蘖分枝的角度和数量及侧生器官的数量等株型方面的研究；对于植物的形态建成、光能的利用以及经济器官的产量和品质有重要影响，并且丰满而紧凑的叶片和株型也可以提高整株植物的观赏价值及经济效益，同时也决定其抗逆性能、生态适应能力和生态效益；
所以园林园艺植物的茎秆矮化和分蘖分枝等株型问题的研究也具有重要的价值意义，仅在苹果等园艺植物中有一定的研究，但是在鳞茎球根花卉中却鲜见报道。
鳞茎球根花卉是指植株地下部分的茎或根变态膨大并贮藏大量养分的一类多年生草本植物，基生叶片较多，且多既可观花又可赏叶，多可作为地被植物栽培，由于其种类丰富、抗逆性强、适应范围广、栽培简易、管理方便等特点，在园林绿化中应用广泛，在药用价值开发、植物生态修复和生态文明建设中也有广阔的发展潜力。
白花虎眼万年青（Ornithogalum thyrsoides）是百合科虎眼万年青属（Ornithogalum spp）多年生鳞茎球根花卉，其植株低矮、簇生的基生叶覆盖地面、且花序花朵繁多、开花时间长、观赏价值高、适应范围广和栽培简易等特点，可作为切花、盆花和地被等，在园林绿化中应用广泛，也可用于废弃土地和重金属污染土壤的植物生态修复中，在生态文明建设中具有广阔的发展潜力。
白花虎眼万年青不但具备多数鳞茎球根花卉所具有的观赏价值高、抗逆强和适应范围广等优点，而且具有再生能力强、繁殖速度快的特点，其叶片可产生多个珠芽（叶上珠芽）来进行繁育，是鳞茎球根花卉中的一朵奇葩；
尤其是白花虎眼万年青具有极其缩短矮化的主茎，多个近基生的叶片簇生覆盖地面，以及多个珠芽着生在叶片上以及数十枚小花产生于无限花序上；其表现出来的株高极度矮化、分蘖分枝众多（基生叶片、花朵和珠芽等侧生器官多）和株型丰满紧凑等特点，不但能够更充分利用光照和营养等外部条件，而且使其抗逆性和竞争力更强，以及繁殖更多的个体并提高产量，获得更加理想的观赏效果、景观效益、经济效益及生态效益，是非常理想的株型和园艺性状，但是其分子机制尚需深入挖掘。
以前的研究中，人们一直认为只有细胞分裂素和生长素是影响植物的分枝两大激素，自2008年以来发现独脚金内酯（Strigolactones, SLs）是一类新的植物激素，调控侧芽伸长、株高、叶片形状、衰老、种子萌发、侧根生长等发育过程，在单子叶植物和双子叶植物中具有功能保守性。
并且人们发现独脚金内酯是受到以DWARF53（DWF53）蛋白为主导的信号转导途径所调控的，其最初是从水稻的矮化多分蘖突变体中分离出来的，是一类新发现的高等植物所特有的蛋白家族，作为独脚金内酯信号途径的抑制因子和关键开关，通过泛素化介导降解特定的靶蛋白来发挥作用，通过影响其下游目标基因的转录水平来调控独脚金内酯的信号传递，参与调控植物分枝分蘖，促进水稻分蘖生长发育；可通过调节DWARF53基因在植物体内的表达水平来实现对水稻分蘖数量和株型的定向调控，从而适当增加分蘖以显著提高水稻产量，并且提高水稻的抗逆性能、生态适应能力和竞争能力等，同时也为水稻的杂种优势的利用等研究开发提供了重要功能基因和有价值的植物材料；是实际生产中可用于调控植物分蘖分枝和株型的重要功能基因；
可见DWARF53蛋白作为独脚金内酯信号的抑制因子和关键开关来调控分蘖和株型；而且DWARF53蛋白可以为深入研究在独脚金内酯信号传递中的调控机制以及其参与的分枝、次生生长、根系发育和光形态建成等发育途径的分子机制提供了突破口和奠定了基础；
这表明，DWARF53基因对于科研和生产具有重要的价值和意义，但是仅仅限于在水稻等极少数农作物中，在园林园艺植物中却鲜见报道；
综上所述，白花虎眼万年青表现出的极度矮化的主茎、多分蘖多分枝（叶片、珠芽和花朵等侧生器官较多）及株型丰满等特点，可能是由其同源的DWARF53蛋白及其所主导的独脚金内酯信号途径所决定的，因此克隆并分离白花虎眼万年青的DWARF53蛋白，即OtDWARF53基因(OtDWF53)，是掌控其独脚金内酯信号途径进而调控植物多分蘖多分枝和茎节等器官的发育进而获得理想株型的关键基因。
发明内容
本发明要解决的技术问题是克隆并分离白花虎眼万年青的DWARF53蛋白，即OtDWARF53基因(OtDWF53)，是掌控其独脚金内酯信号途径进而调控植物多分蘖多分枝和茎节等器官的发育进而获得理想株型的关键基因。
为解决上述问题，本发明在稳定成熟的白花虎眼万年青叶上珠芽高效再生体系及其转录组分析的基础上的，成功克隆了DWARF53在白花虎眼万年青中的同源基因OtDWARF53，为掌控其独脚金内酯信号途径进而调控植物多分蘖多分枝和茎节等器官的发育进而获得理想株型提供了基础。
为达到上述目的，本发明具体通过以下技术方案实现，一种白花虎眼万年青矮化多分蘖OtDWARF53基因，包括SEO ID NO.1所示的核苷酸序列和SEO ID NO.1所示的核苷酸序列经替换，缺失或添加碱基形成的具有同等功能的核苷酸序列。
一种上述白花虎眼万年青矮化多分蘖Ot DWARF53基因编码的蛋白，其氨基酸如SEO ID NO.2所示。
一种含有白花虎眼万年青矮化多分蘖OtDWARF53基因的核苷酸序列的白花虎眼万年青Ot DWARF53基因表达载体，及表达载体的宿主细胞。
一种上述表达载体的构建方法，该方法中用到的无缝连接引物包括HindIIIOtDWARF53和BamHIOtDWARF53R；其核苷酸序列分别如SEO ID NO.5和SEO ID NO.6所示。
一种Ot DWARF53基因的制备方法，其特征在于，包括以下步骤：
步骤1、在无菌的超净工作台上，剪取白花虎眼万年青的离体叶片，放到含有细胞分裂素6BA的培养基上进行离体培养和珠芽的诱导；
步骤2、待白花虎眼万年青的离体叶片上产生珠芽后，剪取珠芽并提取其RNA，并反转录获得cDNA模板；
步骤3、对cDNA模板进行PCR扩增，获得OtDWARF53基因编码区PCR产物；所述PCR扩增的引物包括OtDWARF53F和OtDWARF53R，其核苷酸序列分别如SEO ID NO.3和SEO ID NO.4所示；对OtDWARF53基因编码区PCR产物连接到T载体中，将其转入大肠杆菌感受态中，涂布平板，37℃倒置培养后挑取单克隆进行PCR鉴定及测序，得到核苷酸序列如SEO ID NO.1所示的OtDWARF53基因，其所编码的氨基酸序列如SEO ID NO.2所示；
步骤4、从pHB植物表达载体上多克隆酶切位点筛选出HindIII和BamHI两个酶切位点用于目的基因植物表达载体的构建，同时根据目的基因的完整开发阅读框架的起始密码子和终止密码子来设计目的基因的上下游特异引物，并分别在其5，端添加根据选取的这两个酶切位点及其相连的碱基序列，从而完成用于植物表达载体构建的两条无缝连接引物包括HindIII OtDWARF53和BamHIOtDWARF53R，其核苷酸序列分别如SEO ID NO.5和SEO ID NO.6所示；用上述两条引物对已经获得的cDNA模板进行PCR扩增，随后对获得的PCR产物进行回收和纯化处理；
步骤5、对pHB植物表达载体上进行HindIII和BamHI双酶切使其线性化，按照无缝融合试剂盒的要求和步骤，对线性化后的pHB植物表达载体和纯化后的PCR产物在PCR上进行连接操作， 并将连接后的产物转化大肠杆菌及涂板和进行抗性筛选，随后挑取单菌落并摇菌，最后进行测序和酶切鉴定分析以确定植物表达载体构建成功。
上述白花虎眼万年青OtDWARF53基因调控植物侧芽腋芽以及株型发育中的应用。
进一步的，所述植物为鳞茎球根花卉。
进一步的，所述鳞茎球根花卉为白花虎眼万年青。
本发明的有益效果在于：
本发明是建立在稳定成熟的白花虎眼万年青叶上珠芽高效再生体系及其转录组分析的基础上的，克隆了DWARF53在白花虎眼万年青中的同源基因OtDWARF53，并将其置于双35S启动子下，构建了组成型的植物表达载体pHB+ OtDWARF53，将其导入模式植物烟草中，发现转基因烟草与野生型相比，原有野生型植株的顶端优势基本被打破，出现了节间明显缩短、植株高度显著矮化、茎尖出现多个顶芽、叶腋长出数个腋芽等根本性的变化，转基因烟草的株型更加丰满，更符合人们在生产上对矮化的株型和多分蘖多分枝等理想株型的需求。
这表明OtDWARF53基因能够调控植物节间和植株高度，促进植物多分蘖多分枝，是可应用于研究植物叶片、珠芽及节间等器官和株型的分子机理，以及培育植株矮化并多分蘖多分枝等理想株型的花卉新品种，并可用于深入研究独脚金内酯在虎眼万年青和鳞茎球根花卉中的信号途径和分子机制，是非常有价值的重要功能基因，无论对于科研和生产，均具有很好的应用前景。
当然，实施本发明的任一产品并不一定需要同时达到以上所述的所有技术效果。
附图说明
图1是本发明白花虎眼万年青叶上珠芽总RNA的1%琼脂糖凝胶电泳图；其中，1和2均为总RNA的1%琼脂糖凝胶电泳条带，可以条带清晰明亮；
图2是本发明白花虎眼万年青OtDWARF53基因完整编码区所编码的氨基酸与海枣（Phoenix dactylifera，登录号： XM_008806797）、野蕉(Musa acuminata subsp.malaccensis，登录号：XM_009412288)、油棕（Elaeis guineensis，登录号：XM_010910100）等进行蛋白质保守序列分析，表明与它们的相似度分别为68.09%、68.33%和67.79%，并包括同源基因的保守区域，为DWARF53的同源基因；
图3是本发明pHB+ OtDWARF53植物表达载体构建过程示意图；
图4是本发明所获得的转DWF53基因的烟草幼苗，其茎尖出现多个顶芽、叶腋产生数个侧芽，株型也更加矮化丰满；
图5是与野生型烟草幼苗对比图，其中，A．野生型烟草，仅有一个顶芽，叶腋无腋芽；B.转基因烟草茎尖多顶芽（从侧面观察）；C. 转基因烟草茎尖多顶芽（从正上方观察）；D. 转基因烟草形成多个小腋芽；E. 转基因烟草产生的腋芽分化形成叶片；
图6是与野生型烟草对比图， 其中， A.为野生型烟草，叶片阔卵形，叶缘全缘； B.为部分转基因烟草，叶片呈狭长的条形，叶缘波浪状；
图7是与野生型烟草成年植株对比图，其中，A.为野生型烟草，植株较高大，节间较长，茎尖仅有一个顶芽，叶腋无腋芽侧芽，叶片多阔卵形，叶缘多全缘；B.转DWF基因烟草，植株严重矮化，节间明显缩短，茎尖出现两个以上的顶芽，叶腋也出现腋芽侧芽，叶片呈条形，边缘表现出波浪状。
具体实施方式
为使本发明实施例的目的、技术方案和优点更加清楚，下面将结合本发明的实施例，对本发明实施例中的技术方案进行清楚、完整地描述。显然，所描述的实施例是本发明一部分实施例，而不是全部的实施例。基于本发明中的实施例，本领域普通技术人员在没有做出创造性劳动前提下所获得的所有其他实施例，都属于本发明保护的范围。
实施例1 制备Ot DWARF53基因：
在经过灭菌的超净工作台上剪切白花虎眼万年青无菌组培苗的叶片并转接到含有细胞分裂素的培养基上，经过20~30天后，叶片表面逐渐长出多个绿色的小球，叶上珠芽开始形成，以白花虎眼万年青叶上珠芽为材料，提取RNA，并反转录成cDNA，设计相应引物进行PCR，琼脂糖凝胶电泳后，回收目的条带，与pMD19-T载体连接，转入大肠杆菌，测序并分析。挑取阳性克隆进行质粒抽提，根据pHB植物表达载体的序列信息设计两条无缝融合的引物并对阳性质粒采用高保真酶进行PCR扩增，与进过线性化处理的pHB植物表达载体进行无缝融合来进行植物表达载体构建，将构建好的植物表达载体用冻融法导入根瘤农杆菌LBA4404中，用叶盘法对烟草叶片进行侵染，在含有潮霉素的培养基上进行筛选，获得的抗性植株进行PCR鉴定后，观察其表型，并拍照记录。
（1）总RNA 提取
在经过灭菌的超净工作台上剪切白花虎眼万年青无菌组培苗的叶片并转接到含有细胞分裂素的培养基上，经过20~30天后，叶片表面逐渐长出多个绿色的小球，叶上珠芽开始形成，以白花虎眼万年青叶上珠芽为材料，采用大连宝生物的总RNA提取试剂盒（RNAisoPlus）的操作步骤进行RNA的提取，所使用的试剂盒耗材均处理使其无RNA酶。白花虎眼万年青总RNA经过1%琼脂糖电泳检测结果如图1所示，a和b两个电泳道中均出现了两条清晰明亮的谱带，18S和28S均完整良好，证明获得了较高质量的RNA；经过紫外分光光度计检测，其OD260/OD280值为1.90，OD260OD230值为2.00，表明RNA质量较好，可用于下一步试验。
RNA提取的具体步骤为：用剪刀将组培玻璃瓶的白花虎眼万年青叶上珠芽剪取下来放到研钵内，加入液氮后研磨，用小勺小心地将匀浆装入已经加入RNAiPlus提取液的1.5ml的EP管中，并震荡使得植物匀浆与提取液充分接触混匀，室温静置5分钟，然后在冷冻离心机上12000g4℃离心5分钟，将上清液转移至新的1.5ml的EP管中，加入等体积的氯仿，震荡混匀，室温静置5分钟，12000g4℃离心15分钟，用移液枪吸取上清液转移至新的离心管中，加入等体积的异丙醇，室温静置10分钟，12000g4℃离心10分钟，吸去上清液，用1ml的75%乙醇漂洗两次，在超净台上干燥，然后用适量的DEPC水溶解。
（2）cDNA模板的制备
以所得的RNA为模板，反转录获得cDNA，所使用的是诺唯赞公司的HiScriptII 1stStrand Cdna Synthesis Kit试剂盒，1）反应混合液的制备：在无RNA酶的离心管内依次加入下列试剂：1uLPrimer(OligodT23VN)，总RNA6ul，并加入RNase free ddH2O至12 ul，在PCR仪上加热5分钟，迅速置于冰上骤冷，并在冰上静置2分钟，然后加入10×RT Mix 2ul和HiScript II Enzyme Mix 2ul轻轻混匀后在PCR仪上，程序为50℃50分钟，85℃5分钟，2）然后在上述反应液中加入1ul的RNaseH,37℃20分钟以去除DNA污染。
（3）基因完整编码区CDS的克隆
根据白花虎眼万年青叶上珠芽转录组获得的OtDWARF53基因完整编码区序列，设计引物进行PCR扩增，引物如下：
OtDWARF53F： ATGCCGACACCGGTCAGTAGCGCGCGGC
其核苷酸序列如SEO ID NO.3所示；
OtDWARF53R：ATCCAAAATAATCCTGGAAGGAAGAAG，
其核苷酸序列如SEO ID NO.4所示；
使用诺唯赞公司Phanta Super –Fidelity DNA Polymerase高保真酶进行基因克隆，25ul反应体系，5×buffer(with 10mMMgSO4)5ul，2.5mMdNTP2ul，正向引物2ul，反向引物2ul，cDNA模板2ul，Super –Fidelity DNA Polymerase0.5ul，余量加ddH2O补足。
PCR反应条件：95℃，4min;95℃ 30s，66℃ 30s， 72℃ 2min； 35℃ cycle； 72℃10 min； 4℃，Forever。
OtDWARF53基因编码区PCR产物在1%琼脂糖凝胶电泳后，在紫外灯下，用解剖刀将目的条带的DNA从琼脂糖凝胶上小心切割下来，进行纯化回收，采用宝生物Takara pMD-TVector 载体系统进行连接反应，反应体系如下：5ul Ligation Buffer，1ul pMD-TVector，4ul PCR纯化产物，ddH2O补足至10ul，混匀，16度过夜后，将其转入大肠杆菌感受态中，涂布平板，37℃倒置培养后挑取单克隆进行PCR鉴定及测序；
对获得的阳性克隆进行测序分析，发现OtDWARF53基因编码区长度为3369bp，序列为SEO ID NO.1所示，所编码的氨基酸序列如SEO ID NO.2所示，包括1123个氨基酸的开发阅读框（ORF）。将测序获得的序列在NCBI上进行Blast比对，查找到其同源序列，采用DANMAN等软件进行其同源序列比对，发现与海枣、野蕉和油棕等物种的DWARF53基因同源性较高。
本发明在对白花虎眼万年青多年的组织培养和栽培生产的基础上，根据其植株低矮及侧生器官多（基生叶片多、珠芽多、花朵多）的生态习性和生物学特点，结合分子生物学和生物信息学等先进技术及独脚金内酯调控株型的研究，将白花虎眼万年青的植物生物学特性、植物表型学和三代和二代联合转录组（3+2）进行系统分析，认为可能是具有较高生物活性的独角金内酯所主导的信号代谢途径在其中发挥了重要作用，从而进行筛选出调控其株型的独角金内酯的主导基因OtDWARF53，并且研究表明该基因表现的功能与其矮化及多分蘖等株型特点相吻合，仅仅在拟南芥和水稻中进行过初步研究，在虎眼万年青属植物、鳞茎球根花卉等园艺植物中尚未见到报道，并且OtDWARF53基因功能强大，表明其是控制植株高度、调控顶芽和侧芽的发育、促使植物多分枝多分蘖、从而获得矮化丰满的理想株型的有价值的重要功能基因，并且为深入研究独角金内酯的信号传导途径、分子机制及其在虎眼万年青等鳞茎球根花卉中的应用提供了突破口，在科研和生产上均具有广阔的应用前景。
实施例2 基因功能验证
首先构建白花虎眼万年青OtDWARF53基因植物表达载体，转入野生型烟草中，进行真核细胞表达及转基因烟草植株的表型观察、鉴定与分析
（一）OtDWARF53基因的组成型植物表达载体的构建
1.将OtDWARF53基因的ORF克隆至pHB载体并置于泛素启动子的控制下；
采用HindIII和BamHI对pHB植物表达载体进行双酶切并纯化，使其线性化
2.根据上述白花虎眼万年青OtDWARF53基因cDNA序列和pHBYFP载体的HindIII和BamHI两个个酶切位点旁15~20个左右的碱基设计无缝连接引物，序列如下：
HindIIIOtDWARF53F:accactctctgtctcAAGCTTATGCCGACACCGGTCAGTAGC，其核苷酸序列如SEO ID NO.5所示；小写部分表示植物表达载体上酶切位点旁的碱基序列斜体且划线部分表示酶切位点，大写部分表示根据目的基因的起始密码子及其后面的碱基序列；
BamHIOtDWARF53R:ggtcaggatactagtGGATCCATCCAAAATAATCCTGGAAG，其核苷酸序列如SEO ID NO.6所示；小写部分表示植物表达载体上酶切位点旁的碱基序列斜体且划线部分表示酶切位点，大写部分表示根据目的基因的终止密码子后面的碱基序列；
3.采用HindIIIOtDWARF53F 和BamHIOtDWARF53R引物对已经获得的白花虎眼万年青cDNA模板进行扩增后回收目的条带，与经过HindIII和BamHI双酶切处理的线性化pHBYFP载体进行无缝融合连接，并将反应产物转化大肠杆菌，涂布平板，挑取单克隆后摇菌，提取质粒后用根据pHB植物表达载体上的序列所设计的测序引物pHBR（GTAGCGGGCGAAGCACT）和目的基因的序列所设计的特异引物进行测序，测序结果表明该植物表达载体构建成功
本发明采用了无缝融合植物表达载体构建技术与T4DNA连接的植物表达载体构建方法相比，只需一次扩增和转化即可完成，无需连接到T载体上及两次转化，节约了时间和试剂；并采用了快切酶和双酶切的方法，节约了时间，效率也明显提高。
（二）叶盘法转化烟草
1.将pHB+OtDWARF53植物表达载体采用冻融法转入根瘤农杆菌LBA4404感受态中，涂布平板；
2.在无菌条件下挑取平板上的阳性菌落，接种于YEB液体培养基中（Sm+，Kan+），28度，200rpm振荡培养24~36小时；
3.在离心机上4000g离心5min后，丢弃上清，用1/2MS液体培养基悬浮菌丝体并稀释，使得菌液的浓度OD600值为0.6左右；
4.选取生长健壮的烟草组培苗的无菌叶片，去其主脉，剪成1平方厘米左右的小叶片；
5.将上述小叶片放入制备好的菌液中，浸泡5分钟，在无菌滤纸上吸干菌液；
6.把经侵染的叶片接种于MS培养基中，28度黑暗条件下共培养2~3天；
7.将经过共培养的烟草叶片转接到抑菌培养基（MS+6BA2.0mg/L+Carb250mg/L）上，在25度左右光照下培养，15~20天左右，以去除根瘤农杆菌；
8.将上述烟草接种于筛选培养基上（MS+6BA2.0mg/L+Hyg50mg/L）上，进行抗性不定芽的筛选，
9.将筛选出的不定芽切下转接到生根培养基（1/2MS+NAA1.0mg/L）上进行生根培养，一周后逐渐开始生根，并进行观察、记录和拍照，如图5-图7所示。
（三）转OtDWARF53基因烟草表型观察
1.如图4所示，说明经过潮霉素筛选已经获得转基因烟草的幼苗，茎尖和叶腋均产生了多个不定芽，株型也更加矮化丰满。
2.如图5所示，本发明转基因烟草幼苗在茎尖和叶腋等方面所发生的具体变化，主要表现为：原有的顶端优势从根本上被打破，茎尖产生多个顶芽并且叶腋也产生了多个腋芽，部分植株叶片形状发生了明显变化，从阔卵形变成了狭长的条形或披针形，叶片边缘从全缘变成了浅裂或锯齿；进一步表明，与野生型相比，转基因烟草的顶端优势被打破，茎尖产生多个顶芽、叶腋也产生了侧芽。
3.如图6所示，本发明所获得转基因烟草在叶片形状上的明显变化，从阔卵形变成了狭长的条形，叶缘也从全缘到波浪状。即与野生型相比，转基因烟草叶片从阔卵形变为狭长的条形，叶缘也从全缘变成波浪状，表明OtDWARF53基因能够改变叶片内激素和影响叶脉的发育从而对转基因烟草的叶片产生影响。
4图7所示，本发明转基因烟草成年植株节间明显缩短，及植株显著矮化，并且其植株顶端产生数个顶芽，叶腋也产生了多个不定芽，并且叶腋的不定芽可以发育成为叶片等器官。即野生型烟草顶端优势明显，茎尖仅有一个独立的顶芽，且叶腋无侧芽腋芽；而转基因烟草茎尖产生多个顶芽，叶腋产生多个腋芽侧芽，且腋芽可以继续发育形成叶片；野生型烟草顶端优势明显，节间较长，茎尖仅有一个顶芽，叶腋无腋芽，叶片阔卵形，较大，且叶缘全缘；转基因烟草株型矮小节间极度缩短，茎尖产生多个顶芽，叶腋也有多个腋芽，叶片变得狭长成条形，且叶片边缘浅裂或成锯齿状
上述说明示出并描述了发明的若干优选实施例，但如前所述，应当理解发明并非局限于本文所披露的形式，不应看作是对其他实施例的排除，而可用于各种其他组合、修改和环境，并能够在本文所述发明构想范围内，通过上述教导或相关领域的技术或知识进行改动。而本领域人员所进行的改动和变化不脱离发明的精神和范围，则都应在发明所附权利要求的保护范围内。
序列表
<110> 四川天艺生态园林集团股份有限公司
四川农业大学
四川天艺优境环境科技有限公司
<120> 一种白花虎眼万年青矮化多分蘖基因OtDWARF53及应用
<130> 2020
<141> 2020-01-17
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<213> 白花虎眼万年青(Ornithogalum thyrsoides)
<400> 1
<210> 2
<211> 1122
<212> PRT
<213> 白花虎眼万年青(Ornithogalum thyrsoides)
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Glu Thr Thr Arg Gly Leu Gln Cys Phe Lys Glu Lys Leu Trp Val Met
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Claims (10)
1.一种白花虎眼万年青矮化多分蘖Ot DWARF53基因，其特征在于包括：
a)SEO ID NO.1所示的核苷酸序列；
b)SEO ID NO.1所示的核苷酸序列经替换，缺失或添加碱基形成的具有同等功能的核苷酸序列。
2.一种权利要求1所述的白花虎眼万年青矮化多分蘖Ot DWARF53基因编码的蛋白。
3.如权利要求2所述的蛋白，其特征在于：其氨基酸如SEO ID NO.2所示。
4.一种含有权利要求1所述的核苷酸序列的白花虎眼万年青Ot DWARF53基因表达载体。
5.一种含有权利要求4所述的表达载体的宿主细胞。
6.一种权利要求4所述的表达载体的构建方法，其特征在于，该方法中用到的无缝连接引物包括HindIII OtDWARF53和BamHIOtDWARF53R；其核苷酸序列分别如SEO ID NO.5和SEOID NO.6所示。
7.一种Ot DWARF53基因的制备方法，其特征在于，包括以下步骤：
步骤1、在无菌的超净工作台上，剪取白花虎眼万年青的离体叶片，放到含有细胞分裂素6BA的培养基上进行离体培养和珠芽的诱导；
步骤2、待白花虎眼万年青的离体叶片上产生珠芽后，剪取珠芽并提取其RNA，并反转录获得cDNA模板；
步骤3、对cDNA模板进行PCR扩增，获得OtDWARF53基因编码区PCR产物；所述PCR扩增的引物包括OtDWARF53F和OtDWARF53R，其核苷酸序列分别如SEO ID NO.3和SEO ID NO.4所示；对OtDWARF53基因编码区PCR产物连接到T载体中，将其转入大肠杆菌感受态中，涂布平板，37℃倒置培养后挑取单克隆进行PCR鉴定及测序，得到核苷酸序列如SEO ID NO.1所示的OtDWARF53基因，其所编码的氨基酸序列如SEO ID NO.2所示；
步骤4、从pHB植物表达载体上多克隆酶切位点筛选出HindIII和BamHI两个酶切位点用于目的基因植物表达载体的构建，同时根据目的基因的完整开发阅读框架的起始密码子和终止密码子来设计目的基因的上下游特异引物，并分别在其5，端添加根据选取的这两个酶切位点及其相连的碱基序列，从而完成用于植物表达载体构建的两条无缝连接引物包括HindIII OtDWARF53和BamHIOtDWARF53R，其核苷酸序列分别如SEO ID NO.5和SEO ID NO.6所示；用上述两条引物对已经获得的cDNA模板进行PCR扩增，随后对获得的PCR产物进行回收和纯化处理；
步骤5、对pHB植物表达载体上进行HindIII和BamHI双酶切使其线性化，按照无缝融合试剂盒的要求和步骤，对线性化后的pHB植物表达载体和纯化后的PCR产物在PCR上进行连接操作， 并将连接后的产物转化大肠杆菌及涂板和进行抗性筛选，随后挑取单菌落并摇菌，最后进行测序和酶切鉴定分析以确定植物表达载体构建成功。
8.权利要求1所述的白花虎眼万年青OtDWARF53基因调控植物侧芽腋芽以及株型发育中的应用。
9.如权利要求8所述的应用，其特征在于：所述植物为鳞茎球根花卉。
10.如权利要求9所述的应用，其特征在于：所述鳞茎球根花卉为白花虎眼万年青。
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