CN114891773B - 一种提高大白菜叶绿素含量的蛋白dBrFC2与编码基因及其应用 - Google Patents
一种提高大白菜叶绿素含量的蛋白dBrFC2与编码基因及其应用 Download PDFInfo
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- CN114891773B CN114891773B CN202210342867.XA CN202210342867A CN114891773B CN 114891773 B CN114891773 B CN 114891773B CN 202210342867 A CN202210342867 A CN 202210342867A CN 114891773 B CN114891773 B CN 114891773B
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Abstract
本发明公开了一种提高大白菜叶绿素含量的蛋白dBrFC2及编码基因在培育提高大白菜叶绿素含量的大白菜中的应用,本发明设计特异性引物,利用PCR技术,从大白菜叶绿素含量增加的EMS突变体中克隆出基因,命名为dBrFC2;利用qPCR分析发现该基因在大白菜的根茎叶中均表达,在叶中为优势表达,在根中的表达量较低;含有dBrFC2基因的大白菜光合效率显著提高;通过转基因技术获得dBrFC2过表达转基因株系,测定过表达转基因株系叶绿素含量,其叶绿素含量显著增加。因此dBrFC2是一个可提高叶绿素含量的功能获得型基因,可以作为植物改良叶绿素合成靶点的基因资源,可实现植物基因编辑精准设计育种,在创制新种质及改良产量和品质方面具有重要的理论意义和应用价值。
Description
技术领域
本发明涉及生物技术领域,特别是一种提高大白菜叶绿素含量的蛋白dBrFC2与编码基因及其应用。
背景技术
大白菜(Brassica rapa L.ssp.pekinensis)属十字花科芸薹属(Brassica),是我国的重要蔬菜作物,其种植面积和消费量皆居蔬菜作物前列。近年来,人们对大白菜的品种特性提出了进一步的需求,叶色多样性特色大白菜日益受到关注,其中叶片深绿色的大白菜品种更容易受到消费者的青睐,叶色深绿可促进光合作用,有效提高产量和品质,具有重要的应用价值。提高叶绿素(Chlorophyll,Chl)含量可作为提高作物光合效率的有效措施,进而显著提高产量和品质。近年来,科学家们已鉴定出大量的叶绿素相关基因,不仅涉及到叶绿素的合成、积累和降解途径,更与叶绿体的发育、光形态建成及叶绿体与细胞核信号转导途径紧密联系在一起。但这些基因突变体多为叶绿素缺失突变体(如黄化/白化突变体),影响植物光合作用正常行使功能,而提高叶绿素含量的突变体及基因尚未有报道。目前缺乏可作为改良靶点的、加强叶绿素合成途径的基因资源,不仅难以满足大白菜分子育种的需求,也限制了复杂的叶绿素生物合成调控机制的研究进程。
针对以上问题,研究一种可提高植物叶绿素含量的基因尤为重要。
发明内容
本发明的目的是要解决现有技术中存在的不足,提供一种提高大白菜叶绿素含量的蛋白dBrFC2与编码基因及其应用。
为达到上述目的,本发明是按照以下技术方案实施的:
本发明的第一个目的是要提供一种提高大白菜叶绿素含量的蛋白dBrFC2,为如(a)或(b)或(c)所示的蛋白:
(a)氨基酸序列是大白菜中BrFC2氨基酸序列所示的蛋白质,所述蛋白BrFC2的氨基酸序列如SEQ ID NO.1所示;
(b)将(a)所示的蛋白质通过一个氨基酸的替换而衍生得到的蛋白dBrFC2,所述蛋白dBrFC2的氨基酸序列如SEQ ID NO.2所示;
(c)在SEQ ID NO.2所示的蛋白质的N端或/和C端连接标签得到的融合蛋白质。
进一步地,在SEQ ID NO.2所示的蛋白质的N端或/和C端连接的标签如下:
标签 | 残基 | 序列 |
Poly-Arg | 5-6(通常为5个) | RRRRR |
Poly-His | 2-10(通常为6个) | HHHHHH |
FLAG | 8 | DYKDDDDK |
Strep-tagII | 8 | WSHPQFEK |
c-myc | 10 | EQKLISEEDL |
。
本发明的第二个目的是要提供一种提高大白菜叶绿素含量的蛋白dBrFC2的编码基因,所述蛋白dBrFC2的编码基因为编码如权利要求1所述的蛋白dBrFC2的DNA分子得到,所述DNA分子为:
(1)核苷酸序列是BrFC2基因的DNA分子,所述BrFC2基因的核苷酸序列如SEQ IDNO.3所示;
(2)核苷酸序列是在BrFC2基因的核苷酸序列基础上进行一个或多个碱基的替换、缺失或/和插入而衍生得到的dBrFC2基因,所示dBrFC2基因的核苷酸序列如SEQ ID NO.4所示。
进一步地,所述dBrFC2基因的开放阅读框为1482bp,编码由494个氨基酸残基组成。
本发明的第三个目的是要提供一种提高大白菜叶绿素含量的蛋白dBrFC2或编码基因在培育提高大白菜叶绿素含量的大白菜中的应用。
与现有技术相比,本发明设计特异性引物,利用PCR技术,从大白菜叶绿素含量增加的EMS突变体中克隆出基因,命名为dBrFC2;利用qPCR分析发现该基因在大白菜的根茎叶中均表达,在叶中为优势表达,在根中的表达量较低;含有dBrFC2基因的大白菜光合效率显著提高;通过转基因技术获得dBrFC2过表达转基因株系,测定过表达转基因株系叶绿素含量,其叶绿素含量显著增加。因此dBrFC2是一个可提高叶绿素含量的功能获得型基因,可以作为植物改良叶绿素合成靶点的基因资源,可实现植物基因编辑精准设计育种,在创制新种质及改良产量和品质方面具有重要的理论意义和应用价值。
附图说明
图1为大白菜dBrFC2其他植物FC2系统发育树分析对比图,图中大白菜BrFC2(Brassica rapa WT)、大白菜具有提高叶绿素含量功能的基因dBrFC2(Brassica rapadg)、甘蓝型油菜(Brassica oleracea)、欧洲油菜(Brassica napus)十字花科模式植物拟南芥(Arabidopsis thaliana)、花生(Arachis hypogaea)、单子叶模式植物水稻(Oryzasativa)、小立碗藓(Physcomitrellapatens)、番茄(Solanum lycopersicum)、黄瓜(Cucumis sativus)、玉米(Zea mays);
图2为BrFC2和dBrFC2在大白菜不同组织器官的特异性表达分析图;
图3为含有BrFC2和dBrFC2基因的大白菜叶绿素含量分析图;
图4为含有BrFC2和dBrFC2基因的大白菜光合效率分析图;
图5为pGWB-BrFC2和pGWB-dBrFC2过表达载5体构建示意图;
图6为BrFC2和dBrFC2在大白菜转基因过表达株系的表型图;
图7为BrFC2和dBrFC2在大白菜转基因过表达株系的叶绿素含量分析图。
具体实施方式
为使本发明的目的、技术方案及优点更加清楚明白,以下结合实施例,对本发明进行进一步的详细说明。此处所描述的具体实施例仅用于解释本发明,并不用于限定发明。
实施例1
1.实验材料及试剂
大白菜野生型WT,用于创制EMS突变体的纯合基因型,含有BrFC2基因;叶色深绿突变体dg,由EMS诱变野生型种子后筛选获得,M6世代,含有dBrFC2基因;大肠杆菌‘DH5α’感受态细胞和根癌农杆菌菌株‘GV3101’由河北农业大学华北作物改良与调控国家重点实验室保存;各种限制性内切酶及Buffer购自TaKaRa公司,柱式植物总RNA分离提取纯化试剂盒购自上海生工生物工程股份有限公司,质粒小量制备试剂盒(升级版离心柱型)购自上海捷瑞生物工程有限公司;Taq DNA聚合酶、dNTPs、TaqBuffer、T4 DNA连接酶、PCR纯化回收试剂盒、PCR切胶回收试剂盒、克隆载体pEASY-T1试剂盒、EasyScript One-Step gDNA Removeland cDNA Synthesis SuperMix试剂盒、TransStart Top Green qPCR SuperMix试剂盒均购自北京全式金生物技术有限公司。
2.实验方法及结果分析:大白菜BrFC2和dBrFC2基因的克隆和表达分析
(1)总RNA的提取和cDNA第一链的合成
大白菜幼苗长出真叶后分别摘取幼苗的根、茎、花和叶样品,使用柱式植物总RNA分离提取纯化试剂盒提取RNA。cDNA第一链的合成参照EasyScript One-Step gDNARemovaland cDNA Synthesis SuperMix试剂盒说明进行。
(2)大白菜BrFC2和dBrFC2基因克隆
设计引物,引物为GhWRKY40-F:5’-atgaactgcccagccatgagtg-3’;GhWRKY40-R:5’-agatgaaggcagaatgcccca-3’。以大白菜叶的cDNA为模板进行PCR扩增,将PCR产物连接到T载体上进行测序,测序由上海生工生物工程股份有限公司完成。结果表明获得的片段大小与预期的大小一致,该片段全长1482bp,包含了从ATG开始TGA结束完整的ORF阅读框,编码了494个氨基酸残基,分子量为51.9KDa(参见序列表中序列3和序列4)。
(3)BrFC2和dBrFC2基因序列系统进化树构建
在NCBI中选取不同植物的FC2的氨基酸序列,它们分别是:大白菜BrFC2(Brassicarapa WT)、大白菜具有提高叶绿素含量功能的基因dBrFC2(Brassica rapa dg)、甘蓝型油菜(Brassica oleracea)、欧洲油菜(Brassica napus)十字花科模式植物拟南芥(Arabidopsis thaliana)、花生(Arachis hypogaea)、单子叶模式植物水稻(Oryzasativa)、小立碗藓(Physcomitrellapatens)、番茄(Solanum lycopersicum)、黄瓜(Cucumis sativus)、玉米(Zea mays)。MEGA 7软件的neighbor-joining方法进行1000次bootstrap重复,从而建立了系统发育树。结果见图1,从图中可以看出,大白菜FC2蛋白与同为芸薹属的甘蓝FC2蛋白最为相近,其次为同为十字花科的拟南芥FC2蛋白。
(4)BrFC2和dBrFC2基因在不同组织的表达分析
抽取大白菜的根、茎、花和叶器官的总RNA,反转录成cDNA,利用该cDNA作为模板进行qRT-PCR分析,利用qRT-PCR方法,检测BrFC2和dBrFC2基因在大白菜的根、茎、花和叶器官中的表达情况,扩增BrFC2和dBrFC2的引物为:F:5’-CGTTATTGGCATCCATTCACT-3’;R:5’-CGACTGTCGAGCTTCAAGGTTT-3’。以大白菜Actin作为内参基因,扩增引物为:F:5’-CGAAACAACTTACAACTCCA-3’;R:5’-CTCTTTGCTCATACGGTCA-3’。qRT-PCR所用的反应体系为20μL,以cDNA为模板,具体反应体系按照试剂盒使用说明进行配置。扩增条件为:预变性95℃2min;95℃30s,退火58℃10s,延伸68℃10s,40个循环;然后65-95℃进行反应熔解曲线分析。每个反应设置3个技术重复。目标基因表达差异采用2-△△CT分析组织特异表达分析。结果表明,BrFC2和dBrFC2基因在这些组织器官中均表达,参见图2,但是其在成熟叶里的表达量较高,幼叶次之,根中的表达量最低,暗示该基因的功能可能存在组织特异性。
(5)含有BrFC2和dBrFC2基因的大白菜叶绿素含量及光合效率分析
大白菜野生型WT,含有BrFC2基因;叶色深绿突变体dg,含有dBrFC2基因。在苗期,WT和突变体dg各从3株中选取由内向外数第五片真叶,采用乙醇-丙酮浸提法提取叶绿素:取0.5g新鲜叶片,擦净叶片表面污物并用吸水纸吸干表面附着的水,尽量避开叶脉部位,剪成适当大小放入50mL的离心管中,加入10mL的提取液(乙醇:丙酮=1:1),黑暗条件下浸提24h,并不时摇动至叶片变白,待其叶片颜色完全褪去。使用UV1800紫外可见分光光度计测定分光光度值,提取液为空白对照,分别测定在波长646nm和663nm下的吸光度,每个样品重复测定3次。按照Inskeep公式计算光合色素含量:
叶绿素a(mg·g-1)=(12.21D663-2.81D646)V/1000W
叶绿素b(mg·g-1)=(20.13D646-5.03D663)V/1000W
叶绿素总含量(mg·g-1)=chl a+chl b
式中:V为提取液体积(mL),W为材料重(g)。
并选择晴朗天气,采用翼鬃麒科技(北京)有限公司生产的YZQ-100E多叶室动态光合仪测定其光合速率。
叶色深绿突变体dg含有dBrFC2基因,其叶片叶绿素b含量显著高于野生型,见图3;光合效率显著高于野生型,见图4。
实施例2
1.实验材料及试剂
同实施例1。
2.实验方法及结果分析:BrFC2和dBrFC2在大白菜转基因过表达株系的表型及叶绿素含量分析
(1)BrFC2和dBrFC2在大白菜转基因过表达株系植株的培育
将连有att位点的纯化后BrFC2和dBrFC2序列,利用BP反应连入pDONR221入门载体:PCR产物3μL,BP ClonaseTMII酶0.5μL,pDONR221入门载体2μL,反应体系25℃孵育过夜。取出冷冻贮藏于-80℃低温冰箱中的感受态细胞悬浮液DH5α,置于冰上解冻,上步连接产物每管加入50μL融解的细胞感受态悬浮液,置于冰中30min,42℃热击90s,迅速置于冰上冷却5min以上。每管混合物中加入400μL LB液体培养基,37℃180rmp震荡培养90min。将混合物均匀的涂抹于LB固体培养基(Kan+)中,吹干后,37℃培养箱倒置暗培养12-16h。用灭菌牙签各挑取平板中3个白色菌落,菌落PCR扩增检测目的片段。将扩增正确的单克隆,加入LB液体培养基(Kan+)中,37℃条件下200rmp震荡培养过夜。利用Gateway系统的LR反应,将测序准确的重组入门载体连入过表达载体pGWB505中,反应体系为:重组载体3μL,LR ClonaseTMII酶0.5μL,pGWB505载体2μL,25℃孵育过夜。载体pGWB505由荷兰瓦赫宁根植物育种组提供。取50μL GV3101农杆菌感受态细胞,加入10μL载体质粒DNA,放置于冰上30min,液氮速冻1min,37℃水浴5min,然后加入800μL LB液体培养基。28℃150rmp培养3个小时。将菌液涂布于LB(Spec+,Rif+)的平板上,28℃培养48小时。挑取平板上的单菌落,接种于3mL LB液体培养基中,28℃200rmp震荡培养48小时。28℃200rmp扩大培养。大白菜遗传转化步骤由河北农业大学华北作物改良与调控国家重点实验室完成,见图5。
(2)BrFC2和dBrFC2在大白菜转基因过表达株系的表型及叶绿素含量分析
各选取3个过表达转基因株系观察表型,并利用实例1(5)中的方法测定叶绿素含量。过表达野生型基因BrFC2,转基因植株与含有BrFC2野生型WT表型相同;过表达突变基因dBrFC2,转基因植株与含有dBrFC2突变体表型相同,为深绿色,见图6。叶绿素含量结果显示,过表达dBrFC2转基因植株叶绿素b含量显著高于过表达BrFC2转基因植株,见图7。
综述,本发明的dBrFC2是一个可提高叶绿素含量的功能获得型基因,含有dBrFC2基因的大白菜光合效率显著提高;通过转基因技术获得dBrFC2过表达转基因株系,测定过表达转基因株系叶绿素含量,其叶绿素含量显著增加。
本发明的技术方案不限于上述具体实施例的限制,凡是根据本发明的技术方案做出的技术变形,均落入本发明的保护范围之内。
序列表
<110> 河北农业大学
<120> 一种提高大白菜叶绿素含量的蛋白dBrFC2与编码基因及其应用
<130> 2022-4-2
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tggaacggaa aagcagcaat gttggcagtg ctcgcactct tggtactaga agtaaccacc 1440
ggaaaagggt ttctgcatca gtggggcatt ctgccttcat ct 1482
Claims (5)
1.一种提高大白菜叶绿素含量的蛋白dBrFC2,其特征在于:
所述蛋白dBrFC2的氨基酸序列如SEQ ID NO.2所示;
在SEQ ID NO.2所示的蛋白质的N端或/和C端连接标签得到的融合蛋白质。
2.根据权利要求1所述的提高大白菜叶绿素含量的蛋白dBrFC2,其特征在于:在SEQ IDNO.2所示的蛋白质的N端或/和C端连接的标签如下:
。
3.一种提高大白菜叶绿素含量的蛋白dBrFC2的编码基因,其特征在于,所述蛋白dBrFC2的编码基因为编码如权利要求1所述的蛋白dBrFC2的DNA分子得到,所述DNA分子为:
所示dBrFC2基因的核苷酸序列如SEQ ID NO.4所示。
4.根据权利要求3所述的提高大白菜叶绿素含量的蛋白dBrFC2的编码基因,其特征在于:所述dBrFC2基因的开放阅读框为1482bp,编码由494个氨基酸残基组成。
5.一种如权利要求1所述的蛋白dBrFC2及其编码基因在培育提高大白菜叶绿素含量的大白菜中的应用。
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Citations (4)
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