CN114058627A - 基因PnMYB2及其在调控三七皂苷合成中的应用 - Google Patents
基因PnMYB2及其在调控三七皂苷合成中的应用 Download PDFInfo
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- CN114058627A CN114058627A CN202111181137.8A CN202111181137A CN114058627A CN 114058627 A CN114058627 A CN 114058627A CN 202111181137 A CN202111181137 A CN 202111181137A CN 114058627 A CN114058627 A CN 114058627A
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Abstract
本发明公开了一种基因PnMYB2及其在调控三七皂苷合成中的应用，该基因PnDCD的核苷酸序列如SEQ ID NO.1所示。本发明以基因PnSS和基因PnSE1上游启动子序列为研究对象，利用酵母单杂交方法从三七cDNA文库中筛选到与启动子片段相互作用的转录因子，为探究三七皂苷生物合成途径的转录调控奠定基础。
Description
技术领域
本发明涉及基因工程技术领域，主要涉及基因PnMYB2及其在调控三七皂苷合成中的应用。
背景技术
三七[Panax notoginseng(Burk.)F.H.Chen]为五加科(Araliaxeae)人参属(Panax)多年生直立草本植物，是我国传统名贵药材之一。三七皂苷(Panax notoginsengsaponins,PNS)是三七主要的药用活性成分，由多种四环三萜皂苷组成。已有研究表明三七总皂苷在中枢神经系统、心脑血管系统、血液系统、免疫系统以及抗纤维化、抗衰老、抗肿瘤等方面均具有较好的药理活性。三七对种植环境要求苛刻，生长周期长，轮作障碍严重，其产量难以满足市场的需求，严重制约了三七产业的可持续发展。三七皂苷主要由甲羟戊酸(MVA)途径合成，鲨烯合酶(Squalene synthase,SS)和鲨烯环氧酶(Squalene epoxidase,SE)是其中关键酶，对植物体内的三萜类和甾醇类化合物有着重要的调控作用。
近年来，药用植物次生代谢合成的基因调控已成为研究热点，且三七基因组测序的完成为三七皂苷生物合成调控机制的阐明以及传统中医药的发展提供了有力的研究与应用基础。转录因子能够作用于多个基因上游的启动子序列实现“多点调控”，影响与次生代谢产物合成相关的多个基因表达。转录因子(Transcription factor)对基因的转录激活是植物次生代谢过程重要的调控环节，而且具有“多点调控”的优势。
已有研究表明，基因PnSS和基因PnSE1是三七三萜皂苷生物合成途径的关键酶基因。PnSS是专属于三萜类和甾醇类物质合成分支代谢流上的一个关键酶，对植物体内的三萜类和甾醇类化合物的生物合成均起着重要的调控作用。Patel等抑制了人参SS基因的表达后，发现三萜皂苷的合成量降低，提高SS在人参不定根中的表达水平会同时使植物甾醇和人参皂苷的含量得以增加。当三七细胞系中的SS和DS共超表达时，皂苷含量明显提高，其中人参皂苷Re含量是对照的5-6倍。PnSE1是三萜皂苷生物合成途径中的另一个关键酶。研究发现三七中存在两种类型的PnSE基因，PnSE1编码537个氨基酸，在各个植物组织中均有表达，PnSE2编码545个氨基酸，但只在花中表达显著，其余组织较弱。推测PnSE1与PnSE2具有不同的表达模式，PnSE1参与三萜皂苷合成途径，而PnSE2参与甾醇合成途径。
发明内容
本发明提供了一种基因PnMYB2及其在调控三七皂苷合成中的应用，该基因PnMYB2为R2R3型MYB转录因子，与直接参与三七皂苷合成关键酶基因PnSS的启动子和基因PnSE1的启动子有相互作用，进而影响三七皂苷的合成。
具体技术方案如下：
本发明提供了基因PnMYB2，该基因的核苷酸序列如SEQ ID NO.1所示。
本发明提供了一种包含上文所述的基因PnMYB2的重组表达载体。
本发明提供了一种包含上文所述的基因PnMYB2的基因工程菌。
本发明提供了一种由权利要求1所述的基因PnMYB2编码的蛋白，其氨基酸序列如SEQ ID NO.2所示。
本发明还提供了如上文所述的基因PnMYB2或所述的蛋白在与基因PnSS的启动子和基因PnSE1的启动子相互作用中的应用。
本发明还提供了如上文所述的基因PnMYB2或所述的基因工程菌在调控三七皂苷合成中的应用。
进一步地，所述调控的途径为：通过基因PnMYB2编码的蛋白与基因PnSS的启动子或基因PnSE1的启动子结合来调控三七皂苷的合成。
本发明还提供了所述的蛋白在调控三七皂苷合成中的应用。
与现有技术相比，本发明具有以下有益效果：
本发明以基因PnSS和基因PnSE1上游启动子序列为研究对象，利用酵母单杂交方法从三七cDNA文库中筛选到与启动子片段相互作用的转录因子，为探究三七皂苷生物合成途径的转录调控奠定基础。
附图说明
图1为实施例3中PnMYB2蛋白二级结构的预测结果；
其中，α-螺旋：最长的竖直线；延伸链：第二长竖直线；β-转角：第三长竖直线；无规则卷曲：最短竖直线。
图2为实施例3中PnMYB2蛋白三级结构的预测结果。
图3为实施例3中PnMYB2蛋白的系统进化树。
图4为实施例4中X-α-gal显色反应验证PnSS(CK)和PnSE1(CK)与PnMYB2之间的互作结果。
图5为实施例5中PnMYB2蛋白的诱导表达结果。
图6为实施例6中PnMYB2蛋白亚细胞定位图(CK：35S-sGFP)。
具体实施方式
下面结合具体实施例对本发明作进一步描述，以下列举的仅是本发明的具体实施例，但本发明的保护范围不仅限于此。
实施例1酵母诱饵菌株的构建
随机选取PnSS与PnSE1启动子上的序列片段，每段上都包含主要的顺式作用元件至少一个，将它们通过PCR克隆的方式构建到pAbAi载体中。利用primer premier 5.0设计特异性引物，并利用SnapGene 3.2.1预测序列内可选的酶切位点，选取了SmaI和XhoI两个限制性内切酶位点。采用KOD高保真酶扩增目的片段，将PCR产物于1％的琼脂糖凝胶上进行电泳分离。然后将目的片段进行切胶回收。用SmaI和XhoI对目的片段和pAbAi载体进行双酶切，37℃酶切6h，然后加入5μL的10×DNA loading buffer终止酶切反应。将酶切产物于1％的琼脂糖凝胶上进行电泳分离，然后进行切胶回收处理。
采用T4连接酶将目的片段与载体连接过夜，连接产物全部转化进入大肠杆菌DH5α感受态(100μL)中。挑取单克隆到LB液体培养基(50mg/L Amp)中，扩大培养，然后吸取1μL菌液作为模板进行菌液PCR验证。验证得到的阳性克隆采用TIANGEN的TIANpure MiniPlasmid Kit II(Code No.DP107)进行质粒的提取。
将上述构建好的诱饵载体用BstBI限制性内切酶进行酶切线性化。纯化后的线性化质粒转入酵母Y1H株系中。酵母诱饵菌株鉴定采用Matchmaker Insert Check PCR Mix I试剂盒进行。鉴定成功的酵母菌落进行扩大培养，挑选到能被AbA抑制的合适的酵母诱饵菌株YS1262和YE697。
实施例2筛选与PnSS/SE1基因启动子互作的转录因子
参照TIANGEN RNAprep Pure植物总RNA提取试剂盒说明书提取三七总RNA。以三七总RNA为模板，经SMART逆转录合成cDNA第一链，利用长距离PCR(LD-PCR)扩增技术合成双链cDNA，并用1.2％琼脂糖凝胶电泳进行检测，最后用CHROMA SPIN+TE-400层析柱进行纯化，参照Matchmaker Gold Yeast One-Hybrid Library Screening System试剂盒使用说明构建三七cDNA文库。
将纯化后的三七cDNA文库与pGADT7载体共转化阳性诱饵酵母菌株感受态细胞，在SD/-Leu/AbA(500ng/mL)培养基上30℃培养3d，挑选阳性克隆至YPDA液体培养基扩大培养。用碧云天酵母质粒小量抽提试剂盒进行酵母质粒提取。以提取到的酵母质粒为模板进行PCR扩增，引物采用通用引物T7(5’-AATACGACTCACTATAGGGCG-3’)与3-AD(5’-AGATGGTGCACGATGCACAG-3’)。获得的PCR产物送样测序。将测序结果所得的数据在NCBI及三七基因组、转录组数据、拟南芥数据库中进行比对，获得与PnSS/SE1基因启动子互作的转录因子PnMYB2，碱基序列如SEQ ID NO.1所示，编码的蛋白的氨基酸序列如SEQ ID NO.2所示。
实施例3生物信息学分析
对上述转录因子PnMYB2的碱基和氨基酸序列提交至NCBI中进行分析。PnMYB2的开放阅读框(ORF)序列有864bp，编码287个氨基酸。使用ExPASy在线软件(https://web.expasy.org/compute_pi/)预测其分子量为32575.96Daltons，等电点(pI)为9.16，表明该蛋白为碱性蛋白。其中强碱性氨基酸(K,R)37个，强酸性氨基酸(D,E)26个，疏水性氨基酸(Hydrophobic Amino Acids)(A,I,L,F,W,V)78个，极性氨基酸(Polar Amino Acids)(N,C,Q,S,T,Y)105个。PnMYB2蛋白的不稳定指数(instability index,II)为55.23，亲水性总平均值(Grand average of hydropathicity,GRAVY)为-0.668，为不稳定亲水性蛋白。SMART在线软件(http://smart.embl-heidelberg.de/)预测结果显示，该蛋白没有跨膜结构(transmembranedomains)，但有两个SANT结构域，分别位于预测氨基酸序列的13～63aa和66～114aa。还含有三个低拷贝区域(low complexity)，分别位于预测氨基酸序列的131～142aa，183～193aa和262～272aa。
利用在线软件SOPMA(https://npsa-prabi.ibcp.fr/cgi-bin/npsa_automat.pl？page＝npsa_sopma.html)对转录因子PnMYB2蛋白的二级结构进行预测，结果如图1所示，该蛋白由23.34％的α-螺旋(Alpha helix)，10.45％的延伸链(Extended strand)，5.57％的β-转角(Beta turn)和60.63％的无规则卷曲(Random coil)构成，说明无规则卷曲结构是蛋白PnMYB2二级结构的骨架。利用在线软件SWISS-MODEL(http://swissmodel.expasy.org/)对PnMYB2蛋白的三维结构进行预测，使用方法为X-ray，分别率为
结果如图2所示。使用的模板编号为6kks.1.A，序列的一致性(Seq Identity)为61.82％，寡核苷酸的状态(Oligo-state)为Monomer(单体)，序列与模板序列的相似度(Seqsimilarity)为0.51，覆盖范围(Coverage)为0.38，对所预测序列的描述为R2R3型MYB转录因子。
转录因子PnMYB2在许多物种中被克隆和分析。通过软件Clustal X和MEGA6.0对PnMYB2的氨基酸序列和NCBI数据库中其他植物中该基因的氨基酸序列进行多序列比对并构建进化树，具体物种及蛋白序列号见表1。根据进化树结果表明(图3)，三七PnMYB2与人参PgMYB2归为一组，说明其在同属植物之间具有很高的同源性。
表1构建进化树的核苷酸序列
实施例4酵母体内验证
根据酵母单杂交的结果，挑选与PnSS和PnSE1启动子存在相互作用的基因序列作为酵母体内验证对象。根据三七转录组数据设计特异性引物(上游引物：5’-CGGAATTCATGGGCCGTTCACCTTGCT-3’，下游引物：5’-CGGGATCCTCAACAACATCTGTAAAACCCACTT-3’)，将EcoRI和BamHI酶切位点引入序列，以三七cDNA为模板，通过PCR扩增将基因构建到pGADT7载体中。将构建好的重组载体分别转入酵母株系YS1262和YE697中，用X-α-gal显色反应验证PnMYB2与PnSS和PnSE1启动子之间的相互作用(图4)。
实施例5体外验证
根据三七转录组数据设计特异性引物(上游引物：5’-CGGGATCCATGGGCCGTTCACCTTGCT-3’，下游引物：5’-CGGAATTCTCAACAACATCTGTAAAACCCACTT-3’)，将BamHI和EcoRI酶切位点引入序列，以三七cDNA为模板，通过PCR扩增将PnMYB2构建到原核表达载体pET-32a中。提取阳性重组质粒，转化大肠杆菌BL21(DE3)感受态细胞。用含有抗生素的LB培养基(50mg/L Amp)挑选阳性克隆。取200μL阳性克隆的菌液接种到5mL的LB液体培养基中扩大培养，当菌液达到对数增长期(OD600＝0.5)时，加入IPTG对重组蛋白进行诱导表达，IPTG的浓度为1mmol/L，适宜的诱导时间为6h，诱导温度为25℃。最后采用SDS-PAGE凝胶电泳技术对获得的蛋白进行凝胶电泳，结果如图5所示。PnMYB2融合蛋白的大小为45KD，除去His标签蛋白后，该条带大小与预测的PnMYB2蛋白分子量大小相吻合，约为32KD。
实施例6亚细胞定位
根据三七转录组数据设计特异性引物(上游引物：5’-TCCCCCGGGATGGGCCGTTCACCTTGCT-3’，下游引物：5’-CGGGATCCTCAACAACATCTGTAAAACCCACTT-3’)，将SmaI和BamHI酶切位点引入序列，以三七cDNA为模板，通过PCR扩增将基因构建到35S-sGFP载体中。将构建好的重组载体转化到大肠杆菌DH5α感受态中。挑取单克隆到LB液体培养基(50mg/L Kan)中，扩大培养，然后吸取1μL菌液作为模板进行菌液PCR验证。验证得到的阳性克隆用TIANGEN的TIANpure Mini Plasmid Kit II(Code No.DP107)进行质粒提取，然后转化根癌农杆菌株GV3101。将农杆菌H2B-RFP菌株与含有目的基因的GV3101菌株按照1：1的比例混匀后注射入同一片烟草叶片，H2B-RFP菌株会使细胞核呈现红色。注射完成后的烟草培养3天，然后取样放至激光共聚焦显微镜下进行观察。亚细胞定位结果显示，PnMYB2蛋白的荧光则与细胞核的红色荧光位置重合，表明其定位于细胞核中，符合转录因子的特性(图6)。
序列表
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Claims (8)
1.基因PnMYB2，其特征在于，该基因的核苷酸序列如SEQ ID NO.1所示。
2.一种包含权利要求1所述的基因PnMYB2的重组表达载体。
3.一种包含权利要求1所述的基因PnMYB2的基因工程菌。
4.一种由权利要求1所述的基因PnMYB2编码的蛋白，其氨基酸序列如SEQ ID NO.2所示。
5.如权利要求1所述的基因PnMYB2或权利要求4所述的蛋白在与基因PnSS的启动子和基因PnSE1的启动子相互作用中的应用。
6.如权利要求1所述的基因PnMYB2或如权利要求3所述的基因工程菌在调控三七皂苷合成中的应用。
7.如权利要求6所述的应用，其特征在于，所述调控的途径为：通过基因PnMYB2编码的蛋白与基因PnSS的启动子或基因PnSE1的启动子结合来调控三七皂苷的合成。
8.如权利要求4所述的蛋白在调控三七皂苷合成中的应用。
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