CN113122547B - CsMYB110基因在调控类胡萝卜素合成中的应用 - Google Patents
CsMYB110基因在调控类胡萝卜素合成中的应用 Download PDFInfo
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Abstract
本发明属于基因工程技术领域，公开了CsMYB110基因在调控类胡萝卜素合成中的应用。其中CsMYB110基因的核苷酸序列如序列表中SEQ ID NO.1所示；CsMYB110基因编码的蛋白的氨基酸序列如序列表中SEQ ID NO.2所示。CsMYB110的表达模式与茶树叶片中类胡萝卜素合成高度相关，该基因能够显著促进植物的类胡萝卜素合成积累。该基因的克隆将不仅有利于解析茶树叶片类胡萝卜素的调控机制，而且有助于培育类胡萝卜素含量较高的茶树品种，提高茶叶品质，具有很大的应用价值。
Description
技术领域
本发明涉及基因工程技术领域，具体为CsMYB110基因在调控类胡萝卜素合成中的应用。
背景技术
茶树(Camellia sinensis(L.)O.Kuntze)是我国重要的经济作物，其叶片中的色素主要分为脂溶性色素和水溶性色素两大类，其中黄酮类、花青素类以及茶黄素等属于水溶性色素，而叶绿素，类胡萝卜素，原花色素等属于脂溶性色素。这些色素都与茶叶品质质量密切相关，包括茶叶的干茶、茶汤及叶底的色泽以及茶叶香气的形成。
类胡萝卜素广泛存在于高等植物的叶、花和果实中，是一种天然的黄色、橙红色或红色的亲脂分子，属于萜类色素。茶叶中类胡萝卜素在每100g干重36～73mg之间，主要由β-胡萝卜素、叶黄素和玉米黄质组成，在植物的生理、发育、生态和进化中起到重要作用。在高等植物中，类胡萝卜素在质体中合成，首先从甲羟戊酸途径(MEP)途径开始，由丙酮酸和3-磷酸甘油醛合成异戊二烯焦磷酸(IPP)及其异构体二甲烯丙基焦磷酸(DMAPP)，然后三个IPP分子与一个DMAPP分子通过牻牛儿基焦磷酸合成酶(GGPPS)缩合成牻牛儿基牻牛儿基焦磷酸(GGPP)，这个途径是二萜类化合物、类胡萝卜素、生育酚和叶绿素侧链合成过程中共有的。八氢番茄红素合成酶(PSY)催化两分子的GGPP缩合生成C40八氢番茄红素，这是生成的第一个类胡萝卜素化合物，然后再经过脱氢、环化、羟基化等转变为其他的类胡萝卜素，如β-胡萝卜素、叶黄素等。近年来，人们对类胡萝卜素的生物合成途径进行了广泛研究，并且已经在许多植物中鉴定了大部分催化生物合成步骤的酶基因，但是对于调控机制的研究大多集中在花与果实方面，对于叶片、种子以及根中研究较少。
通常来说，消费者对茶类饮品的青睐很大一部分是取决于其香气与味道，而大多数香气化合物是在茶类的加工过程中由类胡萝卜素、氨基酸、不饱和脂肪酸等转化形成的。类胡萝卜素在茶树叶片中含量较高，不仅是植物光合作用和光保护所必需的色素，参与光合作用以及多种生理逆境调节，而且是形成茶叶色泽和香气的重要化合物，其氧化降解产物如β-紫罗酮、二氢海葵内脂等对增进茶叶香气有重要作用。有研究表明，类胡萝卜素在生产过程中的降解导致了红茶与乌龙茶加工过程中萜类风味化合物的形成，由此鉴定类胡萝卜素是生产高品质茶类香气的重要前体物质。近年来茶树基因组数据的公布，虽然为我们从分子水平解析茶树叶片中类胡萝卜素合成机理提供了理论参考，但是在基因层次研究茶树类胡萝卜素合成的研究较少，不能够为基因工程育种提供优秀的基因资源。
发明内容
本发明的目的在于：提供CsMYB110基因在调控类胡萝卜素合成中的应用，实现调控茶树叶片中类胡萝卜素合成和茶叶品质形成的应用研究，为实现茶树选择性农艺性状育种提供了理论和实际参考基础。
为了实现上述目的，本发明提供如下技术方案：
一种CsMYB110基因，所述CsMYB110基因为茶树MYB类转录因子基因，所述基因的核苷酸序列，如序列表SEQ ID NO.1所示。
优选地，所述CsMYB110基因编码的蛋白序列，如序列表SEQ ID NO.2所示。
优选地，CsMYB110基因在调控类胡萝卜素合成中的应用，所述CsMYB110基因应用于调控茶树中类胡萝卜素合成和茶叶品质的形成。
本发明的有益效果在于：
本发明中，首次克隆并验证了调控茶树类胡萝卜素合成的关键转录因子CsMYB110，该转录因子在茶树中促进类胡萝卜素类化合物合成，影响茶叶品质形成，本发明还提供了含有CsMYB110基因的重组质粒、转基因工程菌和转基因植株。本发明丰富了茶树次生代谢的认知，以及茶叶品质形成特别是香气形成的遗传机制。本发明为实现茶树选择性农艺性状育种提供了理论和实际参考基础。
附图说明
图1：为茶树不同组织中类胡萝卜素化合物的含量差异和CsMYB110以及类胡萝卜素合成基因的表达差异图；
图2：为CsMYB110基因在烟草叶片中的亚细胞定位分析图；
图3：为CsMYB110基因促进烟草叶片中类胡萝卜素化合物的合成比较图；
图4：为CsMYB110基因调控茶树叶片类胡萝卜素化合物合成积累图。
具体实施方式
在本发明中所使用的术语，除非有另外说明，一般具有本领域普通技术人员通常理解的含义。
下面结合具体的制备实施例和应用实施例，并参照数据进一步详细地描述本发明。应理解，这些实施例只是为了举例说明本发明，而非以任何方式限制本发明的范围。
在以下的实施例中，未详细描述的各种过程和方法是本领域中公知的常规方法。所用到的引物，均在首次出现时标明，其后所用相同引物，均以首次标明的内容相同。
实施例1：
1、CsMYB110基因的克隆与序列结构分析
CsMYB110基因，为茶树MYB类转录因子基因，其克隆与序列结构分析，具体如下：
茶树国家级良种舒茶早种植于安徽省庐阳区合肥安徽农业大学农业产业园，取幼嫩叶片用于RNA的提取。总RNA的抽提采用Trizol试剂(Invitrogen，USA)按照说明操作，用分光度计检测其RNA含量和质量。
反转录生成第一链：取1μg RNA作模板，根据Promega公司M-MLV1反转录酶配套试剂使用说明，分别加入下游引物0.5μg，定容至15μl，70℃变性5min，立即在冰上放置5min；然后分别加入M-MLV 5×buffer 5μl，dNTP(25mM)5μl，rRNasin RibonucleaseInhibitor25U，M-MLV反转录酶200U，DEPC水补足25μl，42℃温育1h，95℃5min灭活反转录酶。经过优化后，取适量的反转录产物用于随后的PCR。以cDNA第一链作为RT-PCR模板，常规方法做PCR，扩增CsMYB110基因。其中上游引物：(5’-AGGTGCATGGACTGAGGAAG-3’)，下游引物：(5’-CAAAGATCCCAAAGGTCCATTTT-3’)。25μl PCR反应体系为：10×Ex taq buffer 2.5μl，dNTP 2.0μl，Mg2+1.5μl，上、下游引物各1μl，Ex taq 0.2μl，模板1μl，ddH20 15.8μl。反应程序如下为95℃5min，95℃50sec，58℃50sec，72℃1min，72℃10min，35个循环。PCR产物CsMYB110基因经纯化回收后，连接到pMDTM19-T Simple Vector载体(Takara，Japan)上得到pMDTM19-T-CsMYB110质粒，转化大肠杆菌感受态细胞DH5α，送上海生工公司测序，得到的CsMYB110基因的核苷酸序列如序列表SEQ ID NO.1所示，具体如下：
ATGGAAGGTGTTCCTTTAGGAGTGAGAAAAGGTGCATGGACTGAGGAAGAAGACAACCTCCTTAAGAAGTGCATTGAAACTAATGGAGAAGGAAAGTGGCACCAAGTTCCTTTCAAAGCAGGATTGAACAGATGCAGGAAGAGTTGTAGATTGAGATGGTTGAACTATCTGAGGCCCAATATTAAGAGAGGAAGCTTTGGGGTAGATGAAGTTGATCTCATTGTTAGGCTTCATAAGCTTCTAGGAAACAGATGGTCGCTAATTGCGGGCAGACTTCCAGGAAGGACAGCAAACGATGTAAAAAACTACTGGAATACCCACTTGCAGAAGAAGCTGATACCTCAAATTGAAGTGGTGAAAGTTAAGACTCCGAGGATGATGGAGACCAAAGCTATACGACCTCGACCTCGAACCTTCTCAAAAAACCTAATTTGGTTAAAATCCAAAACAACCGCCATAGCTAATATTGAAACAAGAAACAATCTCTTCAAGCAACTATCACCACCACTATCGCCGCCAAGGGACGATGGAATATCGTGGTGGGAAAACATGTTTGTTGACTTGGAAATTAACAAAGAAATCACATTGTCAATTGATGGATCAAACGAGGAGAAATGGCATGAAAAGGAAACACAAGGTATTCTGGCAATTGGCGACAGTTCTGTTCAAGGAGAGAGTGATTGGAATGACATTTTTATTGATAAAATGGACCTTTGGGATCTTTGA。
CsMYB110基因编码的蛋白序列，如序列表SEQ ID NO.2所示具体如下：
MEGVPLGVRKGAWTEEEDNLLKKCIETNGEGKWHQVPFKAGLNRCRKSCRLRWLNYLRPNIKRGSFGVDEVDLIVRLHKLLGNRWSLIAGRLPGRTANDVKNYWNTHLQKKLIPQIEVVKVKTPRMMETKAIRPRPRTFSKNLIWLKSKTTAIANIETRNNLFKQLSPPLSPPRDDGISWWENMFVDLEINKEITLSIDGSNEEKWHEKETQGILAIGDSSVQGESDWNDIFIDKMDLWDL。
2、茶树不同组织类胡萝卜素含量和CsMYB110基因表达分析
(1)茶树不同组织类胡萝卜素含量分析
茶树国家级良种舒茶早种植于安徽省庐阳区合肥安徽农业大学农业产业园，通过8个组织器官用来分析类胡萝卜素含量分布和基因表达。这8个组织器官包括顶芽、嫩叶、成熟叶、老叶、嫩茎、花、果实、根。类胡萝卜素化合物测定采用安捷伦高效液相色谱仪(HPLC)进行分析。
取新鲜研磨的样品粉末约0.1g，加入1ml氯仿匀混，室温超声仪处理1h；再加入800μl水混匀，2500r/min离心10min，分层后取下部有色液体至新管中，真空干燥至液体挥发；加入500μl异丙醇和100μl饱和的KOH-甲醇溶液混匀避光放置于-20℃过夜；第二天加入500μl水和400μl氯仿混匀，2500r/min离心10min，分层后取下部有色液体至新管中，真空干燥至液体挥发；加入少量乙酸乙酯混匀，用0.22μm滤膜过滤，然后分装到进样瓶中准备进样分析。
HPLC参数设置如下：反相Spherisorb ODS2色谱柱(5μm，4.6×250mm)(Waters)，柱温30℃，进样量为10μl，检测波长为440nm，采用二元流动相，流速为1.0mL/min，梯度洗脱(A80％甲醇+，B 100％乙酸乙酯：0.1-5min 95％A 5％B；5min-8min 75％A 25％B；8min-15min 70％A 30％B；15min-25min 60％A 40％B；25min-45min55％A 45％B；45min-60min45％A 55％B；60min-65min 30％A 70％B；65min-70min 100％B；70min-75min 95％A 5％B)。所检测物质均采用内标法进行定性和定量。
(2)CsMYB110和相关基因在茶树不同组织中表达分析
上文中茶树8个不同组织用于总RNA的提取，然后交由中国华大公司进行二代转录组测序，每个样品三个生物学重复。基因表达量计算由标准的转录组测序分析流程产生，基因表达量用FPKM(Fragments Per Kilobase of exon model per Million mappedfragments)值表示。
图1为茶树不同组织中类胡萝卜素化合物的含量差异和CsMYB110以及类胡萝卜素合成基因的表达差异图。如图1所示，通过HPLC分析可以看到茶树8个不同组织的叶黄素与β-胡萝卜素含量分布不同，在老叶(ML)中叶黄素含量最高，并且随叶片成熟度不断升高而升高，β-胡萝卜素也呈现类似趋势。在茶树嫩茎(ST)、嫩芽(AB)、花(FL)中类胡萝卜素含量也比较高，在其他组织中含量比较低。其中茶树花中只特异性地含有β-胡萝卜素。通过基因表达聚类分析表明，CsMYB110与一些胡萝卜素代谢途径结构基因具有相似的表达趋势，都高表达在茶树花中。
3、CsMYB110基因亚细胞定位和烟草叶片瞬时过量表达分析
(1)CsMYB110-pK7GW7载体构建
以pMDTM 19-T Simple:CsMYB110质粒为模板，利用引物(其中上游引物为
5’-GGGGACAAGTTTGTACAAAAAAGCAGGCTTCAGGTGCATGGACTGAGGAAG-3’，下游引物为5’-GGGGACCACTTTGTACAAGAAAGCTGGGTCAAAGATCCCAAAGGTCCATTTT-3’)进行PCR扩增。PCR产物用1％的琼脂糖胶电泳条带进行回收。利用Gateway克隆技术，加入1μl PCR回收产物，与其质量相当的pDONR221中间载体，最后加入1μl BP Clonase Mix，室温过夜后转化DH5α，送上海生工公司测序。测序正确的阳性克隆的质粒，取1μl质粒并加入等量pK7GW7过表达载体，最后加入1μl LR Clonase Mix，室温过夜后转化DH5α，送上海生工公司测序验证。
(2)CsMYB110亚细胞定位分析
取适量的本式烟草种子加去离子水，置于4℃冰箱春化处理72h之后播种。播种完覆盖保鲜膜，置于合适条件(湿度60％，温度23℃，16小时光照/8小时黑暗)下等待发芽。种子出芽之后，尽量选取大小一致的苗进行移栽，正常培养。把CsMYB110-pK7GW7载体电激法转化到GV3101农杆菌当中，并通过常规PCR方法鉴定阳性克隆。挑取含有目的基因的阳性菌落，在5mL含有相应抗生素的LB液体培养基中，28℃，200r/min培养约24h；吸取培养过的菌液2ml，加入到50mL新鲜的含有相应抗生素的LB液体培养基，继续振荡培养至OD600约为1.0，离心收集菌体，用适量转化液(MS+0.3mg/L6-BA+150g/L蔗糖+15g/LEMS+0.06％silwetL-77，PH＝5.7)重悬菌体，终浓度OD600约为0.8。将准备好的转化液装在注射器中，轻轻地注射到烟草叶片背部，然后黑暗放置24H。三天之后，取注射的叶片进行激光共聚焦显微镜观察。以不含有目的基因的空载体作为对照。
图2为CsMYB110基因在烟草叶片中的亚细胞定位分析图。如图2所示，CsMYB110在烟草叶片中具有明显的转录因子核定位信号，这说明CsMYB110可能在细胞核中调控相关基因的转录表达。
(3)烟草叶片瞬时表达代谢物分析
通过激光共聚焦显微镜对烟草叶片进行荧光分析，选择带有核定位荧光信号的烟草叶片作为CsMYB110瞬时过表达遗传材料，空载体作为对照。然后用HPLC分别分析烟草叶片中类胡萝卜素化合物含量的差异。
图3为CsMYB110基因促进烟草叶片中类胡萝卜素化合物的合成比较图。图3所示，图3(A)中，在烟草叶片中瞬时表达CsMYB110，能显著促进β胡萝卜素的合成。图3(B)中，在烟草叶片中瞬时表达CsMYB110，能显著促进黄体黄质的合成。在图3中，把CsMYB110瞬时过表达在烟草叶片中，能显著提高烟草叶片的类胡萝卜素化合物的合成积累，其中黄体黄质和β-胡萝卜素升高最显著。
结果表明，瞬时过表达CsMYB110能显著提高烟草叶片中类胡萝卜素化合物的含量。
4、CsMYB110基因在茶树体内功能验证
(1)体外寡核苷酸反义抑制实验
根据CsMYB110预测序列设计合成寡核苷酸反义的引物，设计在网站http://sfold.wadsworth.org/cgi-bin/soligo.pl上完成，引物序列如所示：
P1，(5’-GTTGTCTTCTTCCTCAGTCC-3’)；
P2，(5’-GGTCGTATAGCTTTGGTCTC-3’)；
P3，(5’-CATCGTTTGCTGTCCTTCCT-3’)；
用80mM蔗糖溶液溶解，配制并获得体外寡核苷酸反义抑制缓冲液，空白为蔗糖溶液；用剪刀剪下大小基本一致，颜色鲜艳，色泽健康，无虫无病的一芽一叶，插入装有缓冲液的96孔板中，要确保一芽一叶尾部没入缓冲液中。将96孔板放入光照培养箱中按光照16h/黑暗8h进行光照培养，培养箱温度为28℃。处理后3d引物处理样和空白样分别取样进行代谢和基因表达分析。
(2)体外寡核苷酸反义抑制样品的代谢物和相关基因表达分析
对处理样品和对照样品分别用于总RNA的提取以及cDNA第一链合成。反转录产物(cDNA第一条链)稀释80倍作为模板，使用SYBR Realtime Mix(TOYOBO,Osaka,Japan)，配制20μl反应体系：8.4μl稀释80倍的逆转录产物，上下游引物各0.8μl(10pmol/μl)，10μl2×SYBR Green PCR Master Mix，每个反应配3个重复。然后在bio-radCFX-96上以程序：①95℃3min；②95℃10s，60℃15s，72℃30s运行45个循环；③从65℃到95℃，以0.1℃/s绘制熔解曲线。
上游引物：(5’-ACCTCGACCTCGAACCTTCT-3’)，下游引物：(5’-TATTCCATCGTCCCTTGGCG-3’)，以茶树ACTIN基因为内参，上游引物：(5’-GCCATATTTGATTGGAATGG-3’)，下游引物：(5’-GGTGCCACAACCTTGATCTT-3’)通过仪器自带分析软件，计算出CsMYB110的相对表达数值。同理，计算PSY(上游引物5’-GTGGCGAAGTTTGTGCAGAG-3’，下游引物：5’-GTGACGCATTAGGCCCATCA-3’)、PDS(上游引物5’-TTTGGCAGATGCAGGTCACA-3’，下游引物：5’-TCAAATCGGCTGAACTCCCC-3’)、ZDS(上游引物5’-GCCATGAGGTTGGTGCTGAT-3’，下游引物：5’-GATCAACAAGAGCCCGCACA-3’)、CRTISO(上游引物5’-TTAGGGTTTAATGGGGCGGG-3’，下游引物：5’-AAACCCAGAACTCCCACCAG-3’)、LYCe(上游引物5’-GGCGCTTATTCTGCAACTGG-3’，下游引物：5’-TCGGCGGAGGAAAGAGTAGA-3’)、BCH(上游引物5’-ACGACGGTCTCGTTCACAAA-3’，下游引物：5’-CAACTCATCAAGCCCTCCCA-3’)的相对表达量。结果显示CsMYB110体外寡核苷酸反义抑制能显著干扰目的基因表达水平。类胡萝卜素合成关键基因表达分析显示：PSY、PDS、ZDS、CRTISO、LYCe、BCH等基因表达都显著下调。代谢物含量分析表明叶黄素及β-类胡萝卜素化合物含量显著降低。
图4为CsMYB110基因调控茶树叶片类胡萝卜素化合物合成积累图。其中图4(A)为寡核苷酸反义抑制实验示意图，一芽一叶材料取自于舒茶早品种，Control treatment，空白对照；AsODN-CsMYB110，CsMYB110寡核苷酸反义链抑制处理。图4(B)为CsMYB110基因在处理样品中的基因表达量图。图4(C)为茶树叶片中类胡萝卜素化合物含量变化图。图4(D)为类胡萝卜素代谢途径关键基因表达量变化图。
如图4所示，可以看到通过体外寡核苷酸反义抑制实验可以使得CsMYB110的表达量和对照相比显著被抑制，通过体外寡核苷酸反义抑制CsMYB110基因表达3d后代谢物质测定结果可以看到，抑制CsMYB110的表达可以显著抑制茶树叶片类胡萝卜素化合物的合成积累，类胡萝卜素合成途径中的关键结构基因的表达量也显著降低。
综上所述，CsMYB110蛋白及其编码基因与茶树叶片类胡萝卜素化合物合成代谢调控相关，能显著提高茶叶品质形成，CsMYB110基因能够应用于调控茶树中类胡萝卜素合成和茶叶品质的形成。
CsMYB110的表达模式与茶树叶片中类胡萝卜素合成高度相关，将该基因瞬时过量表达在植物叶片中，能够显著促进植物的类胡萝卜素合成积累，通过反义寡核苷酸技术抑制CsMYB110的表达，使茶树叶片中类胡萝卜素相关化合物含量显著下降，类胡萝卜素合成相关基因表达也显著下降。该基因的克隆将不仅有利于解析茶树叶片类胡萝卜素的调控机制，而且有助于培育类胡萝卜素含量较高的茶树品种，提高茶叶品质，具有很大的应用价值。
本发明中，首次克隆并验证了调控茶树类胡萝卜素合成的关键转录因子CsMYB110，该转录因子在茶树中促进类胡萝卜素类化合物合成，影响茶叶品质形成，本发明还提供了含有CsMYB110基因的重组质粒、转基因工程菌和转基因植株。本发明丰富了茶树次生代谢的认知，以及茶叶品质形成特别是香气形成的遗传机制。本发明为实现茶树选择性农艺性状育种提供了理论和实际参考基础。
上述是对发明进行了示例性描述，显然本发明具体实现并不受上述方式的限制，只要采用了本发明的方法构思和技术方案进行的这种非实质改进，或未经改进将发明的构思和技术方案直接应用于其他场合的，均在本发明的保护范围之内。
序列表
<110> 安徽农业大学
<120> CsMYB110基因及在调控类胡萝卜素合成中的应用
<130> NO
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ctaggaaaca gatggtcgct aattgcgggc agacttccag gaaggacagc aaacgatgta 300
aaaaactact ggaataccca cttgcagaag aagctgatac ctcaaattga agtggtgaaa 360
gttaagactc cgaggatgat ggagaccaaa gctatacgac ctcgacctcg aaccttctca 420
aaaaacctaa tttggttaaa atccaaaaca accgccatag ctaatattga aacaagaaac 480
aatctcttca agcaactatc accaccacta tcgccgccaa gggacgatgg aatatcgtgg 540
tgggaaaaca tgtttgttga cttggaaatt aacaaagaaa tcacattgtc aattgatgga 600
tcaaacgagg agaaatggca tgaaaaggaa acacaaggta ttctggcaat tggcgacagt 660
tctgttcaag gagagagtga ttggaatgac atttttattg ataaaatgga cctttgggat 720
ctttga 726
<210> 2
<211> 241
<212> PRT
<213> 茶树(Camellia sinensis L. O. Kuntze)
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Claims (1)
1.一种CsMYB110基因在调控类胡萝卜素合成中的应用，其特征在于，CsMYB110基因为茶树MYB类转录因子基因，所述CsMYB110基因的核苷酸序列，如序列表SEQ ID NO.1所示；所述CsMYB110基因应用于调控茶树中类胡萝卜素合成和茶叶品质的形成。
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