CN104119432A - 一种薄壳山核桃MADS-box类转录因子CiMAD9及其编码基因与应用 - Google Patents
一种薄壳山核桃MADS-box类转录因子CiMAD9及其编码基因与应用 Download PDFInfo
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Abstract
本发明公开了一种薄壳山核桃MADS-box类转录因子CiMADS9及其编码基因与应用。该薄壳山核桃MADS-box类转录因子CiMADS9,是具有序列表中的SEQIDNO.2所述氨基酸序列的蛋白质,其编码基因为CiMADS9基因SEQIDNO.1的DNA序列。薄壳山核桃MADS-box类转录因子CiMADS9基因可用于培育晚花植物品种。CiMADS9为特异表达基因,其表达模式与组织以及植株的发育阶段相关。过量表达CiMADS9的转基因拟南芥与对照组相比,开花比野生型植株晚,开花时间推迟8d以上,开花时基叶数较对照增加6片以上,说明CiMADS9可以控制植物的成花转变,影响植物的花期和营养生长,进而调控植物的生殖生长。
Description
技术领域
本发明涉及植物基因工程领域,具体涉及一种薄壳山核桃MADS-box类转录因子CiMAD9及其编码基因与应用。
背景技术
植物的开花时间直接控制植物营养生长期和生育期的长短,在相当程度上决定着繁育的成功与否,也与植物的产量和质量性状密切相关。MADS-box是一个保守性很强的DNA结合结构域,最早在酵母Mini Chromosome Maintenance1(MCM1)、拟南芥AGAMOUS(AG)、金鱼草DEFICIENS(DEF)和人类血清应答因子Serum ResponseFactor(SRF)中发现,因此具有此结构的功能基因被命名为MADS-box基因。MADS-box基因参与所有植物花发育过程的调节。最初认为MADS-box基因主要参与花器官的形态建成,目前发现MADS-bo基因功能具有多样性,参与几乎所有器官的形态建成及从胚到配子体的生长发育的每一个阶段。有些MADS-box基因的功能是多方面的,具有多效性,不仅仅参与某一方面的调节,在植物发育中的各个层次和方向发挥作用。
发明内容
发明目的:
本发明的目的是提供一种薄壳山核桃MADS-box类转录因子CiMAD9。
本发明的另一目的是提供该转录因子CiMAD9的编码基因。
本发明的又一目的是提供该转录因子的功能。
本发明的目的可通过如下技术方案实现:
本发明所提供的一种薄壳山核桃MADS-box类转录因子,命名为CiMAD9,来源于薄壳山核桃(Carya illinoensis(Wangench.)K.Koch)优良品种‘马罕’,氨基酸序列如SEQ ID NO.2所示。
本发明所述的的薄壳山核桃MADS-box类转录因子的编码基因,其cDNA序列如SEQ IDNO.1所示,含有768bp的最大开放阅读框,编码SEQ ID NO.2所示的255个氨基酸残基序列。
含有本发明CiMAD9基因SEQ ID NO.1的表达载体。
所述的表达载体优选将所述的薄壳山核桃MADS-box类转录因子CiMAD9的编码基因插入到pCAMBIA1301载体的KpnⅠ和SacⅠ酶切位点间所得。
宿主菌是指将pCAMBIA1301-CiMAD9转入的根癌农杆菌EHA105.
扩增CiMAD9cDNA全长的引物为:
CiMADS9-ORF sense:5'-ATGGGGAGAGGGAGGATAG-3'
CiMADS9-ORF antisense:5'-CCATCAGCCAAACCTTCT-3'
实时荧光定量RT-PCR分析中涉及的CiMAD9的qPCR引物为:
CiMADS9-qPCR sense:5'-GGTACAACGAGTGTGTAGATTCTCC-3'
CiMADS9-qPCR antisense:5'-TCCTCTCCTTCACAGACAATAACCC-3'
上述薄壳山核桃MADS-box类转录因子CiMAD9、其编码基因、含有编码基因的表达载体在培育晚花植物中的应用。
有益效果:
本发明在薄壳山核桃中克隆到一个MADS-box类转录因子CiMAD9基因,CiMAD9可能参与薄壳山核桃花发育、果实发育等生殖发育的大部分进程。CiMAD9编码基因在雄花、雌花和幼果中特异性表达。转基因拟南芥中过量表达CiMAD9编码基因,与对照组相比,转基因材料表现为晚花,开花时间推迟8d以上,开花时基叶数较WT增加6片以上。这一结果说明CiMAD9基因可以控制植物的成花转变过程,同时可以延长植株的营养生长。
本发明的CiMAD9对于培育晚花植物品种具有重要意义,在作物育种方面尤其是培育以营养器官为主的植物具有广泛的应用前景。
利用本发明的植物表达载体,将CiMAD9基因导入植物体内,可以获得开花延迟的转基因植株。
附图说明
图1CiMAD9基因在薄壳山核桃的表达特性
1,叶;2,当年生枝条;3,雌花;4,雄花;5,幼果
图2.CiMAD9过量表达表达载体的构建
图3.转CiMAD9基因拟南芥和对照组RT-PCR检测。
WT:野生型;1-5:转基因型
图4转CiMAD9基因拟南芥和对照的表型观察
WT:野生型;T:转基因株系
图5转基因与对照植株的基生叶数和开花时间比较
具体实施方式
下列实例中所有的方法无特别说明,均为常规方法
实施例1 薄壳山核桃CiMAD9基因的克隆与鉴定
实验材料为薄壳山核桃品种‘马罕’,根据本实验室已经克隆得到的保守片段(莫正海等,2013,南方农业学报,44(5):730-734.),设计两条特异引物,进行3'RACE PCR。基因特异引物序列分别为,GSP1:5'-ATGCAAATAGCAGGCAAGTCACATTCTC-3'(SEQ ID NO.3);GSP2:5'-CGCTGTTCTCTGTGAT GCTGAGGTCG-3'(SEQ ID NO.4),与其搭配的引物为接头引物均为AUAP:5'-GGCCACGCGTCGACTAGTAC-3'(SEQ ID NO.5)。取马罕雄花,进行RNA的提取,参照BioTeke通用植物总RNA提取试剂盒(离心柱型)进行。取总RNA1μg,cDNA合成采用PrimeScript RTase(Takara)反转录酶,以带接头的Oligo d(T)引物AP:5'-GGCCACGCGTCGACTAGTACTTTTTTTTTTTTTTTTT-3'(SEQ ID NO.6)进行反转录,获得第一链cDNA。以薄壳山核桃雄花第一链cDNA为模板,进行巢式PCR,第一轮PCR反应条件为:94℃5min热变性;94℃45s,65℃45s,72℃1min,共35个循环;72℃延伸10min。将第一轮PCR溶液稀释10倍,以此作为模板,进行第二轮PCR,反应条件同第一轮。将第二轮PCR产物用胶回收试剂盒(Genscript)回收后,与pMD19-T载体(Takara,Japan)连接,然后转化大肠菌DH5α,挑选阳性克隆,进行测序。将测序得到的片段与原来的保守片段进行比对、拼接,获得序列长度为1077bp(SEQ ID NO.7)。
为了获得其最大开放阅读框,在其两端设计基因特异引物,CiMADS9-ORF sense:5'-ATGGGGAGAGGGAGGATAG-3'(SEQ ID NO.8)和CiMADS9-ORF antisense:5'-CCATCAGCCAAACCTTCT-3'(SEQ ID NO.9),以薄壳山核桃雄花第一链cDNA为模板,进行PCR扩增,PCR反应条件为94℃5min热变性;94℃45s,48℃45s,72℃1min,35个循环;72℃延伸10min,4℃保存,将PCR产物回收、克隆、测序,经分析后获得SEQ IDNO.1所示序列。CiMAD9的ORF为768bp,编码255个氨基酸,氨基酸序列如SEQ ID NO.2所示。将所得到的序列SEQ ID NO.2在NCBI数据库中进行Blastp比对,结果表明,该序列与甜橙AGL15基因、川桑AGL15基因、鹰嘴豆AGL15基因的同源性分别为73%、71%、70%。
实施例2 薄壳山核桃CiMAD9基因在不同器官中的表达特征
以薄壳山核桃三个品种(绍兴、波尼、马罕)为材料,利用实时荧光定量RT-PCR技术对薄壳山核桃不同组织叶片、当年生枝条、雌花、雄花、幼果中的表达情况进行了研究。各组织总RNA的提取同实施例1,cDNA的合成用能消除RNA中残留DNA的试剂盒,PrimeScriopt RT reagent kit with gDNA Eraser(Takara,Japan),具体步骤参照试剂盒说明书进行。内参使用山核桃actin基因(周秦.山核桃果实成熟期间基因表达的初步分析[J].临安:浙江农林大学,2010),引物序列为ACTIN-F:5'-GCTGAACGGGAAATTGTC-3'(SEQ IDNO.10),ACTIN-R:5'-AGAGATGGCTGGAAGAGG-3'(SEQ ID NO.11)。根据CiMAD9基因序列设计表达检测引物CiMADS9-qPCR sense:5'-GGTACAACGAGTGTGTAGATTCTCC-3'(SEQ ID NO.12)和CiMADS9-qPCR antisense:5'-TCCTCTCCTTCACAGACAATAACCC-3'(SEQ ID NO.13)。以来自不同组织的cDNA为模板进行实时荧光定量RT-PCR分析。
结果分析表明(图1),CiMADS9基因在三个品种中均有表达,在雄花中的表达量最高。三个品种的表达分析均表明,CiMADS9基因在生殖组织(雄花、雌花、幼果)中的表达量显著高于营养组织(叶片、枝条)中的表达量。
实施例3 CiMADS9转录因子的功能鉴定
提取含有CiMADS9编码基因目的片段的T-载体及表达载体pCAMBIA1301质粒DNA,用限制性内切酶KpnⅠ和SacⅠ双酶切,切胶回收,然后用T4DNA连接酶连接,转化大肠杆菌。提取质粒,PCR和酶切鉴定。将大肠杆菌质粒用冻融法转入农杆菌(EHA105)感受态细胞,菌液PCR鉴定阳性克隆。获得CiMADS9过量表达表达载体pCAMBIA1301-CiMADS9(图2)。将过量表达载体通过用冻融法将pCAMBIA1301-CiMADS9转入根癌农杆菌株EHA105(Avsian-Kretchmer et al,2004,Plant Physiology,135:1685-1696)中。pCAMBIA1301-CiMADS9通过农杆菌株EHA105的介导,转化拟南芥,利用抗生素筛选和PCR技术筛选阳性转基因植株。以经抗生素筛选的抗性植株总RNA为模板,以未转基因拟南芥植株总RNA为阴性对照进行RT-PCR反应(图3),拟南芥18S作为RNA提取和cDNA合成成功的对照。将鉴定出的转基因T2代种子和野生型拟南芥种子同时播到栽培基质中,人工培养条件为:相对湿度80%,光照强度80-200μmol/M2/S,温光周期为16h光照,22℃/8h黑暗,17℃。对其表型进行分析,与WT相比,转基因材料表现为晚花(图4),开花时间推迟8d以上,开花时基叶数较WT增加6片以上(图5)。
Claims (10)
1.一种薄壳山核桃MADS‐box类转录因子CiMAD9,氨基酸序列如SEQ IDNO.2所示。
2.权利要求1所述的薄壳山核桃MADS‐box类转录因子CiMAD9的编码基因。
3.根据权利要求2所述的编码基因,其特征在于所述的薄壳山核桃MADS‐box类转录因子的cDNA基因核苷酸序列如SEQ ID NO.1所示。
4.含有权利要求2或3所述的薄壳山核桃MADS‐box类转录因子的编码基因的表达载体。
5.根据权利要求4所述的表达载体,其特征在于所述的表达载体是将所述的薄壳山核桃MADS‐box类转录因子CiMAD9的编码基因插入到pCAMBIA1301载体的KpnⅠ和SacⅠ酶切位点间所得。
6.含有权利要求2和3所述基因的宿主菌。
7.根据权利要求6所述的宿主菌,其特征在于所述的宿主菌是将pCAMBIA1301‐CiMAD9转入根癌农杆菌EHA105所得。
8.权利要求1所述的薄壳山核桃MADS‐box类转录因子CiMAD9在培育晚花植物或延长植株的营养生长中的应用。
9.权利要求2或3中所述的薄壳山核桃MADS‐box类转录因子CiMAD9的编码基因在培育晚花植物或延长植株的营养生长中的应用。
10.权利要求4或5所述的表达载体在培育晚花植物或延长植株的营养生长中的应用。
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CN113429465A (zh) * | 2021-05-24 | 2021-09-24 | 哈尔滨学院 | 一种桑黄MADS-box类转录因子PbMADS1及其编码基因与应用 |
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CN116003551A (zh) * | 2022-09-22 | 2023-04-25 | 四川农业大学 | 基因片段a在培育植物新材料中的应用 |
CN116003551B (zh) * | 2022-09-22 | 2024-03-01 | 四川农业大学 | 基因片段a在培育植物新材料中的应用 |
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