CN110607310B - 一种调控茶叶茶毫形成的基因及应用 - Google Patents
一种调控茶叶茶毫形成的基因及应用 Download PDFInfo
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Abstract
本发明公开了一种调控茶叶茶毫形成的基因及应用。其中CsMYB184基因的核苷酸序列如序列表中SEQ ID NO.1所示；CsMYB184基因编码的蛋白的氨基酸序列如序列表中SEQ ID NO.2所示。CsMYB184的表达模式与茶毫分布高度相关，将该基因超量表达转化植物，能够完全使转基因植物的表皮毛发育缺陷表型恢复正常，通过反义寡核苷酸技术抑制CsMYB184的表达，使茶树叶片中特征性化合物含量显著下降。该基因的克隆不仅将有利于理解茶树表皮毛形态建成的分子机制，而且将有助于认识茶毫这个重要外观和品质农艺性状形成的分子机理，为茶树品质育种提供理论依据，加快特定茶树品种育种进程，本发明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，具体涉及一种茶树MYB类转录因子基因及其编码蛋白和应用。
背景技术
茶树(Camellia sinensis(L.)O.Kuntze)是我国重要的经济作物。茶叶富含氨基酸、生物碱、茶多糖、多酚类和挥发类等物质，这些内含物含量和组分的不同，共同赋予了茶叶色、香、味等品质特征。茶毫是茶树重要的农艺性状，其丰富多样的次生代谢物不仅是茶叶品质形成的关键，而且对茶树逆境胁迫响应具有重要作用。茶毫是一种茶叶单细胞类型表皮毛，主要分布在幼嫩的芽叶背部，长度依叶序而异，并随着幼叶成熟而自行脱落，至成熟叶基本全部脱落。茶毫中含有18种人体所需要的氨基酸、多酚类物质和咖啡碱，以及含量丰富的挥发类物质，对成茶的外观、滋味和香气都具有重要贡献。茶毫对多种名优茶的外形有着画龙点睛的作用，如工夫红茶、烘青、白茶等都要求显毫。因此，芽叶茶毫的多少，已经成为茶叶原材料幼嫩和优质的标志，而成茶毫的多少及隐显亦是评定茶叶品质优劣的重要因素之一。
表皮毛除了作为植物抵御外界病虫害、天敌取食等第一道物理屏障外，也能通过合成、积累或分泌大量的次生代谢产物来抵御外界胁迫。螨类和假眼小绿叶蝉是危害茶树生长的两类重要害虫，大量的研究表明，茶毫的富集对这两种害虫的取食具有阻碍作用。也有研究表明，茶毫由于木质素含量高，阻碍某些鳞翅目害虫的取食和产卵。前人的研究表明，茶毫在茶树抵御外界病虫害侵害以及非生物逆境胁迫中，具有重要作用。茶毫对茶树生理生长和成茶品质都具有重要影响，但是其形成的分子机理尚不清楚，严重阻碍了我们对茶树发育生物学和茶叶品质形成机理的深入认识。
在茶树叶片的发育和成茶的品质形成以及茶树对外界逆境胁迫响应，茶毫都起着非常重要的作用，随着茶树基因组数据的公布，为我们从分子水平解析茶毫发育机理提供了理论参考。对茶毫发育、生物学功能的研究，将能更好地丰富植物表皮毛的发育机理，以及为优质、抗逆基因工程育种提供优秀的基因资源。
发明内容
本发明要解决的技术问题是提供一种来源于茶树的调控茶毫形成的基因(下称CsMYB184基因或转录因子)。
本发明要解决的另外一个技术问题是提供上述CsMYB184基因在茶毫形成方面的应用。
对于CsMYB184基因，本发明提供的技术方案是，一种具有调控茶毫形成的基因，其核苷酸序列如SEQ ID NO.1所示，其编码氨基酸序列如SEQ ID NO.2所示。
本发明还包括上述CsMYB184基因在茶毫形成方面的应用。
本发明的有益效果是：
首次克隆并验证了控制茶毫发育的关键转录因子，同时该转录因子还调控茶叶特征性化合物合成，影响茶叶品质形成，本发明还提供了含有CsMYB184基因的重组质粒、转基因工程菌和转基因植株。本发明丰富了茶树发育生物学的认知，以及茶树次生代谢调控新的机制。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
下面结合附图和具体实施方式对本发明作进一步详细的说明。
图1是本发明实施例2的茶树不同组织茶毫分布差异和CsMYB184编码蛋白的表达差异。以富毫茶树品种福鼎大白为研究材料，不同组织中茶毫分布和CsMYB184表达模式高度正相关。
图2是本发明实施例2的茶毫分布差异显著的茶树品种中CsMYB184编码蛋白的表达差异。茶毫显著富集品种：FDDB，福鼎大白；DJBH，短节白毫；TYDY，桃源大叶。茶毫显著少品种：HJG，黄金桂；LJ，龙井；HK，黄魁。A，六种茶树品种的一叶的茶毫表型。B，不同茶树品种中茶毫密度定量分析。C，不同茶树品种中茶毫长度定量分析。D，不同茶树品种中CsMYB184及相关基因表达模式。
图3是本发明实施例3的CsMYB184基因能完全互补拟南芥表皮毛发育缺陷突变体gl1的表型。A，半定量分析CsMYB184在突变体gl1、野生型WS和转基因CsMYB184/gl1中表达分析。B，突变体gl1、野生型WS和转基因CsMYB184/gl1表皮毛表型观测。C，拟南芥中GL1下游靶基因AtGL2和AtGL3表达分析。
图4是本发明实施例4的CsMYB184基因调控茶树叶片特征性化合物合成积累。A，寡核苷酸反义抑制实验示意图，一芽一叶材料取自于福鼎大白品种，Control，空白对照；asODN-CsMYB184，CsMYB184寡核苷酸反义链抑制处理。B，空白对照和CsMYB184寡核苷酸反义链抑制处理材料中咖啡碱、儿茶素类和没食子酸化合物含量变化。表没食子儿茶素没食子酸酯(EGCG)，表儿茶素没食子酸酯(ECG)，表没食子儿茶素(EGC)，表儿茶素(EC)，儿茶素(C)，没食子酸(GA)空白对照和CsMYB184寡核苷酸反义链抑制处理材料中主要挥发类物质含量检测。
图5本发明实施例4的体外寡核苷酸反义抑制CsMYB184，对儿茶素、咖啡碱、没食子酸、萜类代谢途径关键基因变化。A，CsMYB184基因在处理样品中表达显著受到抑制。B，儿茶素代谢途径关键基因表达变化，Leucoanthocyanidin reductase(LAR)，Anthocyanidinreductase(ANR)，Anthocyanidinsynthase(ANS)，Serine carboxypeptidases like(SCPL)。C，咖啡碱代谢途径关键基因表达变化，Tea caffeine synthase(TCS)，7-Methyxanthine methyl transferase(MXMT)。D，没食子酸代谢途径关键基因表达变化，3-Dehydroquinic acid dehydrase/shikimate dehydrogenase(DHD/SDH)。E，萜类代谢途径关键基因表达变化，Terpenoid synthase(TPS)。
具体实施方式
实施例1.CsMYB184基因的克隆与序列结构分析
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，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-MLV反转录酶配套试剂使用说明合成cDNA。经过优化后，取适量的反转录产物用于随后的PCR。以cDNA作为RT-PCR模板，常规方法做PCR，扩增CsMYB184基因。上游引物：(5’-ATGGCTCCGAAGAGCAGTGA-3’)，下游引物：(5’-TTACCATTTATCGGTAAGTGCC-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 taq0.2μL，模板1μL，ddH2015.8μL。反应程序如下为95℃5min，95℃50sec，58℃50sec，72℃1min，72℃10min，35个循环。PCR产物CsMYB184基因经纯化回收后，连接到pMDTM19-TSimple Vector载体(Takara，Japan)上得到pMDTM19-T-CsMYB184质粒，转化大肠杆菌感受态细胞DH5α，送去测序，得到如下的核苷酸序列：
上述CsMYB184基因编码的蛋白序列如下：
实施例2.CsMYB184基因的表达差异与茶毫分布差异相关性：
1，茶树不同组织茶毫分布和CsMYB184基因表达
茶树国家级良种福鼎大白品种种植于安徽省庐阳区合肥安徽农业大学农业产业园，13个组织器官用来分析茶毫分布和基因表达。这13个组织器官包括嫩芽、一叶、二叶、三叶、四叶、老叶、第一茎段、第二茎段、第三茎段、第四茎段、花、果实、根。通过体式显微镜或者数码相机对13个品种的茶毫分布进行拍照记录。同时这些样品用于总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)，10μl 2×SYBR Green PCR Master Mix，每个反应配3个重复。然后在bio-rad CFX-96上以程序：①95℃3min②95℃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’-CCGAATATCAAGCGAGGCAACA-3’)，下游引物：(5’-ATCGGTTCGTCCAGGAAGTCTC-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CsMYB184在不同组织中的相对表达数值。
2，茶树不同种质资源茶毫分布和CsMYB184基因表达
茶树品种种质资源丰富，表皮毛数量和长度等都有显著差异。通过茶树资源资料结合实际考察，选取了茶毫显著富集的茶树品种福鼎大白、桃源大叶、短节白毫以及茶毫显著少的茶树品种黄金桂、龙井和黄魁，这些品种均种植于安徽省庐阳区合肥安徽农业大学农业产业园。不同品种均选择第一叶进行茶毫性状的考察以及基因表达分析，利用体式显微镜记录茶毫分布情况，选取不少于30个叶片进行定量分析，结果如图1。并采用高通量二代测序的方法(参考illumina公司具体流程)检测各个样本中CsMYB184以及相关基因的表达量。
在图1中，可以看到福鼎大白13个不同组织中茶毫分布差异显著，其中幼嫩叶片中茶毫分布较多，随着叶片成熟茶毫逐渐减少。茎部也有少量茶毫分布，但整体表现短小稀疏。而花、果实和根部表皮毛几乎不可见。通过qRT-PCR检测结果显示CsMYB184表达量与茶毫分布高度正相关。CsMYB184高表达在茶毫丰富的幼嫩叶片当中，其它组织表达较低。结合蛋白序列基因注释，暗示CsMYB184可能与茶树表皮毛的起始发育有关系。
在图2中，可以看到茶毫作为一种重要的农艺性状，在不同茶树品种资源中差异显著。在三个茶毫富集的品种中CsMYB184的表达显著高于三个茶毫少的品种，而与茶毫发育相关的其它转录因子的表达没有变现出明显的相关性。可见CsMYB184可能是茶树资源中茶毫分布显著差异的重要遗传因素。
实施例3.CsMYB184基因在拟南芥体内功能验证
1，CsMYB184-pB2GW7载体构建
以pMDTM 19-T Simple::CsMYB184质粒为模板，利用以下引物进行PCR扩增：
上游引物：(5’-GGGGACAAGTTTGTACAAAAAAGCAGGCTTCATGGCTCCGAAGAGCAGTGA-3’)，
下游引物：(5’-GGGGACCACTTTGTACAAGAAAGCTGGGTTTACCATTTATCGGTAAGTGCC-3’)，
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1μlPCR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，送去测序。测序正确的阳性克隆的质粒，取1μL质粒并加入等量pB2GW7过表达载体，最后加入1μL LR Clonase Mix，室温过夜后转化DH5α，送去测序验证。
2，拟南芥遗传转化
各取适量的拟南芥野生型和gl1突变体种子加去离子水，置于4度冰箱春化春化处理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℃；光周期16h光/8h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CsMYB184-pB2GW7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，用适量拟南芥转化液(MS+0.3mg/L6-BA+150g/L蔗糖+15g/LEMS+0.06％silwet L-77，PH＝5.7)重悬菌体，终浓度OD600约为0.8。拟南芥种植约一个月，植株开始陆续开花，挑选生长健壮的植株为待转化植株，转化前不断去除顶端花序，以使植株产生更多的花蕾。转化前一天需将待转化植株充分浇水。将准备好的转化液装在容器内，轻轻把拟南芥花序浸泡在转化液当中约30S，然后黑暗放置24H，然后正常培养，收获种子。将收获的拟南芥种子至于离心管中，先用1ml0.1％的升汞灭菌2min,然后用无菌水冲洗5-6遍，用枪头吸取种子，播于潮霉素浓度为25mg/L的MS固体培养基上。4℃黑暗条件下春化72h，转移至培养室，温度23℃；光周期16h光/8h暗条件下培养。约两周后，选取叶片绿色、根系发育正常的抗性植株移植到栽培基质中继续培养。移植前栽培基质充分吸水，移植后覆盖保鲜膜，约3d去除，以后管理同上，收获T2代种子，做好标记，继续筛选，至得到纯合转化植株。提取拟南芥苗期DNA和RNA，利用基因特异引物进行PCR检测目的基因表达。对转基因拟南芥在生长不同阶段进行观察，并用显微镜、相机等工具记录表皮毛发育表型。
在图3中，在拟南芥中gl1突变体表皮毛发育缺陷是由于拟南芥R2R3-MYB转录因子GL1基因的功能丧失，导致无法激活下游GL2，GL3等靶基因从而导致表皮毛细胞发育无法启动，形成地上组织表面光滑表型。CsMYB184可以完全互补拟南芥gl1突变体表皮毛发育缺陷的表型，进一步qRT-PCR分析显示CsMYB184可能是通过激活下游GL2，GL3靶基因，进而决定表皮毛发育。
实施例4.CsMYB184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CsMYB184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’-TTGTCGTCATTTTTCCCTTC-3’)；
P2，(5’-AGTTGTCGTCATTTTTCCCT-3’)；
P3，(5’-GAAGTTGTCGTCATTTTTCC-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的测定
儿茶素、咖啡碱和没食子酸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0.05g，加入1ml提取液(80％甲醇)匀混，室温超声仪处理1h。4度120r/min震荡提取过夜。2500r/min离心10min，取上清液再用0.2μm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：C18色谱柱(5um 4.6×250mm)，柱温39℃，进样量为10μl，检测波长为280nm，采用二元流动相，流速为1.0mL/min，梯度洗脱(A 0.1％冰醋酸，B 100％甲醇：0.1-5min 95％A 5％B；5min-8min 75％A 25％B；8min-15min 70％A30％B；15min-25min 60％A 40％B；25min-45min 55％A 45％B；45min-60min 45％A55％B；60min-65min 30％A 70％B；65min-70min 100％B；70min-75min 95％A5％B)。所检测物质均采用外标法进行定性和定量。挥发物的测定采用顶空固相微萃取萃取(headspacesolid-phase microextraction，HS-SPME)结合气相色谱-质谱联用(GC-MS)(Agilent7890A)的方法来测，用含有50/30um DVB/CAR/PDMS(Supelco)的萃取头萃取体外寡核苷酸反义抑制样品和空白样品的挥发物，60℃萃取1h，然后取出萃取头插入GC-MS进样口解吸附5min，同时启动仪器开始收集数据。色谱条件：进样口温度为240℃，载气为高纯氮气，纯度大于99.99％,流速0.8ml/min,不分流进样；色谱柱型号为HP-5MS石英毛细管柱(60m×0.32mm×0.25um)；柱温起始为40℃，保持3min，以2℃/min升至90℃,保持5min，再以3℃/min升至160℃，最后以10℃/min升至250℃，保持5min；质谱条件：离子源为EI源，离子源温度230℃，电子能量70eV，四极杆温度150℃,接口温度280℃，电子倍增器电压1680V，扫描范围m/z为35～350amu。
3、体外寡核苷酸反义抑制影响茶树基因表达分析
对处理样品和对照样品分别提取总mRNA，然后进行反转录，合成第一链cDNA，用定量PCR检测相关基因表达。首先对对照和处理样品中CsMYB184的基因表达水平进行了检测，结果显示CsMYB184体外寡核苷酸反义抑制能显著干扰目的基因表达水平，下降幅度约为8倍。儿茶素合成关键基因表达分析显示：LAR，ANR，ANS以及SCPL基因表达都显著下调。咖啡碱合成关键基因表达分析显示：TCS和MXMT基因表达显著下调。没食子酸合成关键基因DHD/SDH基因表达分析显示：3个DHD/SDH拷贝基因表达也都显著下调。
在图4中，可以看到体外寡核苷酸反义抑制CsMYB184基因表达3d后代谢物质测定结果，可以看到抑制CsMYB184的表达可以显著抑制咖啡碱、儿茶素类、没食子酸以及部分挥发类物质合成积累。
在图5中，可以看到体外寡核苷酸反义抑制实验中CsMYB184的表达量和对照相比显著被抑制，抑制率接近80％。同时儿茶素类、咖啡碱、没食子酸以及萜类物质合成途径中关键基因的表达也在寡核苷酸反义抑制处理样品中显著被抑制。
综上所述，CsMYB184蛋白及其编码基因与植物表皮毛发育和特征性化合物合成代谢调控相关，能显著提高茶叶外观和内在品质形成。
以上所述的本发明实施方式，并不构成对本发明保护范围的限定。任何在本发明的精神和原则之内所作的修改、等同替换和改进等，均应包含在本发明的权利要求保护范围之内。
SEQUENCE LISTING
<110> 安徽农业大学
<120> 一种调控茶叶茶毫形成的基因及应用
<130> NO
<160> 8
<170> PatentIn version 3.5
<210> 1
<211> 582
<212> DNA
<213> 茶（Camellia sinensis）
<400> 1
atggctccga agagcagtga gggactggct aaaaaagttt acaataaagg agcatggaca 60
tctgaggagg atagaaagct ggctcagtat gttgaagttc atggagcaaa gaagtggaag 120
actatcgcta ccaagtcagg tttgaaccga tgcgggaaga gttgtagatt gagatggttg 180
aattatctta gaccgaatat caagcgaggc aacattactg atgaagaaga ggacttgata 240
cttaggcttc ataagctatt agggaacagg tggtccttga ttgctgggag acttcctgga 300
cgaaccgata atgagattaa gaactattgg aattctcatt tgagcaggaa aataaatcag 360
aagggaaaaa tgacgacaac ttcgccggaa caagaaagca cgcctgagaa aactgcagac 420
tctgacgtca aaagagaagg caccaaagga agtggagacg gagagtttat gcttgatgtg 480
aatgaattct tcgatttctc taccggaggt acctacgggt tagattgggt taataaattt 540
cttgaactcg atgatgatca ggcacttacc gataaatggt aa 582
<210> 2
<211> 193
<212> PRT
<213> 茶（Camellia sinensis）
<400> 2
Met Ala Pro Lys Ser Ser Glu Gly Leu Ala Lys Lys Val Tyr Asn
1 5 10 15
Lys Gly Ala Trp Thr Ser Glu Glu Asp Arg Lys Leu Ala Gln Tyr
20 25 30
Val Glu Val His Gly Ala Lys Lys Trp Lys Thr Ile Ala Thr Lys
35 40 45
Ser Gly Leu Asn Arg Cys Gly Lys Ser Cys Arg Leu Arg Trp Leu
50 55 60
Asn Tyr Leu Arg Pro Asn Ile Lys Arg Gly Asn Ile Thr Asp Glu
65 70 75
Glu Glu Asp Leu Ile Leu Arg Leu His Lys Leu Leu Gly Asn Arg
80 85 90
Trp Ser Leu Ile Ala Gly Arg Leu Pro Gly Arg Thr Asp Asn Glu
95 100 105
Ile Lys Asn Tyr Trp Asn Ser His Leu Ser Arg Lys Ile Asn Gln
110 115 120
Lys Gly Lys Met Thr Thr Thr Ser Pro Glu Gln Glu Ser Thr Pro
125 130 135
Glu Lys Thr Ala Asp Ser Asp Val Lys Arg Glu Gly Thr Lys Gly
140 145 150
Ser Gly Asp Gly Glu Phe Met Leu Asp Val Asn Glu Phe Phe Asp
155 160 165
Phe Ser Thr Gly Gly Thr Tyr Gly Leu Asp Trp Val Asn Lys Phe
170 175 180
Leu Glu Leu Asp Asp Asp Gln Ala Leu Thr Asp Lys Trp
185 190
<210> 3
<211> 22
<212> DNA
<213> 人工序列(Artificial sequence)
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CCGAATATCA AGCGAGGCAA CA 22
<210> 4
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<212> DNA
<213> 人工序列(Artificial sequence)
<400> 4
ATCGGTTCGT CCAGGAAGTC TC 22
<210> 5
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 5
GCCATATTTG ATTGGAATGG 20
<210> 6
<211> 20
<212> DNA
<213> 人工序列(Artificial sequence)
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GGTGCCACAA CCTTGATCTT 20
<210> 7
<211> 51
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<213> 人工序列(Artificial sequence)
<400> 7
GGGGACAAGT TTGTACAAAA AAGCAGGCTT CATGGCTCCG AAGAGCAGTG A 51
<210> 8
<211> 51
<212> DNA
<213> 人工序列(Artificial sequence)
<400> 8
GGGGACCACT TTGTACAAGA AAGCTGGGTT TACCATTTAT CGGTAAGTGC C 51
Claims (1)
1.如SEQ ID NO.1所示核苷酸序列的基因在调控茶叶茶毫形成方面的应用，其特征在于，调控茶毫的分布与数量。
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| CN113234737A (zh) * | 2021-06-30 | 2021-08-10 | 安徽农业大学 | 茶树myb类转录因子基因在调控茶树咖啡碱生物合成方面的应用 |
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