CN112813075B - 芥蓝BoaWRKY4基因及其应用 - Google Patents
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Abstract
本发明涉及生物技术领域,尤其是涉及芥蓝BoaWRKY4基因在对核盘菌导致的菌核病抗性中的应用。本发明公开了提高芥蓝对核盘菌抗性的基因BoaWRKY4:为BoaWRKY4.1、BoaWRKY4.2、BoaWRKY4.3;本发明还公开了上述基因编码的蛋白质。基因BoaWRKY4沉默后,不仅能使芥子油苷合成相关基因的表达量降低,还能减弱芥蓝对核盘菌的抗性。
Description
技术领域
本发明涉及生物技术领域,尤其是涉及芥蓝BoaWRKY4基因在对核盘菌(Sclerotinia sclerotiorum(Lib.)de Bary)导致的菌核病抗性中的应用。
背景技术
芸薹属蔬菜不仅是我国栽培面积最大、产量最高和消费量最大的蔬菜,在世界上其他地区也具有不可估量的作用。核盘菌是一类寄主非常广泛的腐生真菌,其引发的菌核病对十字花科芸薹属蔬菜的危害十分严重。芥蓝是一种中国特产的芸薹属蔬菜,广泛种植于我国南部和东南亚地区。在生产中芥蓝易受到核盘菌的侵染,导致其产量、营养品质和安全性降低,挖掘芥蓝对抗核盘菌的抗性基因,并解析其功能和作用机制具有重要的生物学意义和经济价值。
核盘菌(Sclerotinia sclerotiorum(Lib.)de Bary)属真菌门、子囊菌亚门、盘菌纲、柔膜菌目、核盘菌属,其寄主范围和地理分布都十分广泛,是最具破坏力的植物病菌之一。核盘菌可以侵染400多种植物,主要包含十字花科和茄科等双子叶植物,也能侵染洋葱和郁金香等单子叶植物,导致60多种病害的发生,如菌核病、棉腐病、水霉病和茎腐病等。核盘菌引发的菌核病是十字花科蔬菜生产中最常见的真菌病害之一。以油菜为例,油菜菌核病在我国发病面积大,一般发病率为10%~30%,严重时高达80%以上,可导致产量减少10%~70%,严重影响油菜的生产和国家油料产量的稳定。
芥子油苷是芸薹属蔬菜中非常重要的次生代谢物质,在植物对抗生物胁迫中发挥了重要作用;芥蓝中含有丰富的芥子油苷。
发明内容
本发明要解决的技术问题是提供一种提高芥蓝对核盘菌抗性的基因、蛋白质及相应的用途。
为了解决上述技术问题,本发明提供一种提高芥蓝对核盘菌抗性的基因BoaWRKY4:为BoaWRKY4.1、BoaWRKY4.2、BoaWRKY4.3;
BoaWRKY4.1的核苷酸序列如SEQ ID NO:1所述;
BoaWRKY4.2的核苷酸序列如SEQ ID NO:2所述;
BoaWRKY4.3的核苷酸序列如SEQ ID NO:3所述。
本发明还同时提供了上述基因编码的蛋白质:
BoaWRKY4.1编码的蛋白质的氨基酸序列如SEQ ID NO:4所述;
BoaWRKY4.2编码的蛋白质的氨基酸序列如SEQ ID NO:5所述;
BoaWRKY4.3编码的蛋白质的氨基酸序列如SEQ ID NO:6所述。
本发明还同时提供了含有上述基因的质粒。
本发明还同时提供了含有上述基因的植物表达载体:为基因沉默VIGS载体PTY-S/BoaWRKY4;所述基因沉默VIGS载体PTY-S/BoaWRKY4同时沉默基因BoaWRKY4.1、BoaWRKY4.2、BoaWRKY4.3。
本发明还同时提供了含有上述基因的宿主细胞:该细胞为大肠杆菌细胞。
本发明还同时提供了基因或蛋白质在提高芥蓝对核盘菌菌核病抗性中的应用。
作为本发明应用的改进:基因BoaWRKY4沉默后,不仅能使芥子油苷合成相关基因的表达量降低,还能减弱芥蓝对核盘菌的抗性。芥子油苷合成相关基因包括吲哚族芥子油苷相关基因BoaMYB51.1、BoaMYB51.2以及BoaCYP83B1以及脂肪族芥子油苷相关基因BoaMYB28.1、BoaMYB29和BoaCYP79F1。
本发明经研究发现,来源于芥蓝的核盘菌抗性基因——BoaWRKY4不仅参与调控吲哚族芥子油苷和脂肪族芥子油苷相关基因的表达,还在对抗核盘菌中发挥着重要的作用。BoaWRKY4是芥蓝抗性相关的关键转录因子,属于第I类WRKY转录因子。本发明通过克隆BoaWRKY4基因,构建BoaWRKY4 VIGS基因沉默芥蓝材料和接种核盘菌实验,表明BoaWRKY4在提高芥蓝和其他芸薹属蔬菜对核盘菌的抗性方面具有很好的应用前景。
综上所述,本发明首次构建了芥蓝BoaWRKY4 VIGS基因沉默植株,并进行基因功能研究。通过测定芥子油苷合成相关基因表达量和核盘菌接种实验,发现BoaWRKY4不仅可以正调控吲哚族芥子油苷和脂肪族芥子油苷相关基因的表达,还在对抗核盘菌中发挥着重要的作用。
附图说明
为了使本发明的目的、技术方案和优点更加清楚,下面结合附图对本发明的具体实施方式作进一步详细说明。
图1是拟南芥(Arabidopsis thaliana)、甘蓝(Brassica oleracea)和芜箐(Brassica rapa)三种芸薹属蔬菜中同源的WRKY4的氨基酸序列的进化树分析。
图2是拟南芥AtWRKY4同芥蓝BoaWRKY4.1、BoaWRKY4.2和BoaWRKY4.3以及甘蓝Bol031460、Bol010361和Bol029198蛋白质序列同源性分析。
图3是BoaWRKY4.1、BoaWRKY4.2、BoaWRKY4.3在芥蓝不同组织中的表达量(相对表达水平)。
图4是BoaWRKY4 VIGS基因沉默芥蓝株系中BoaWRKY4.1、BoaWRKY4.2和BoaWRKY4.3基因的表达量,芥蓝叶片对核盘菌的抗性表型以及菌斑面积统计;其中,pTY-S为接种空载质粒的对照组,pTY-S/BoaWRKY4-1和pTY-S/BoaWRKY4-2代表不同的BoaWRKY4 VIGS基因沉默植株。
图5是芥子油苷合成相关基因在对照和BoaWRKY4 VIGS植株中的相对表达水平。
具体实施方式
下面结合具体实施例对本发明进行进一步描述,但本发明的保护范围并不仅限于此:
一、芥蓝BoaWRKY4的克隆
参考Brassica Database(BRAD,http://brassicadb.org)中甘蓝(Brassicaoleracea)的CDS序列,使用NCBI primer design tool设计特异性引物。PCR引物由北京擎科生物科技有限公司合成。以大笋迟花叶片的cDNA为模版,使用 HS DNAPolymerase(premix)(Takara,Japan)扩增BoaWRKY4.1、BoaWRKY4.2和BoaWRKY4.3片段。引物序列为:
BoaWRKY4.1F:ATGTCGGAGAAGGAAGAACTCC
BoaWRKY4.1R:AACTATGTGCTCTTCTTTGAGCC
BoaWRKY4.2F:ATGTCGGAGAAAGAAGAACGTCCG
BoaWRKY4.2R:AATTATTTGCTCTTCTTTCAGCCTT
BoaWRKY4.3F:ATGTCGGAGAAGGAAGATCTTCC
BoaWRKY4.3R:TTTCAGCCTTAGACACGCAAC
PCR反应体系:2xPrimerSTAR buffer 25μl、dNTP Mixture 5μl、PrimerSTAR HSDNA polymerase 1μl、ddH2O 16μl、cDNA 1μl、上下游引物(10μmol/L)各1μl,共50μl。PCR反应程序为:94℃90s;94℃30s,55℃45s,72℃1min30s,循环38次;72℃5min。扩增的PCR产物用1%琼脂糖凝胶电泳鉴定,然后将扩增条带割胶用GEL/PCR Purification Kit(toroivd,China)回收纯化,将回收产物构建到-Blunt Zero Cloning载体上,转化到Trans1-T1感受态细胞,PCR鉴定后将阳性克隆送到北京擎科生物科技有限公司测序。测序所得的BoaWRKY4的核苷酸序列如SEQ ID NO:1、3、5所示;该基因编码的蛋白质的氨基酸序列如SEQ ID NO:2、4、6所示。
具体为:
BoaWRKY4.1的核苷酸序列如SEQ ID NO:1所述;
BoaWRKY4.2的核苷酸序列如SEQ ID NO:2所述;
BoaWRKY4.3的核苷酸序列如SEQ ID NO:3所述。
二、BoWRKY4 VIGS基因沉默载体的构建
在BoaWRKY4.1、BoaWRKY4.2和BoaWRKY4.3的CDS序列中寻找5’端以NTAG、NTAA或者NTGA开头的共有40bp的寡核苷酸序列,然后在这段序列后加上其反向互补序列组成长度为80bp的自定义寡核苷酸序列作为目的片段,将目的片段与载体pTY-S进行酶切连接。
80bp的片段为:
TTGACCCGAGATTCAAGCAGAGCAGACCCACCGGTTTGATATCAAACCGGTGGGTCTGCTCTGCTTGAATCTCGGGTCAA;
具体步骤如下:
处理pTY-S质粒。首先,在37℃的金属浴里,用SnaBI(NEB,USA)酶切pTY-S质粒1-2h(Methods in Molecμlar Biology,2013,vol.975,197-210)。酶切后先用1%的琼脂糖对酶切产物进行凝胶电泳,再用GEL/PCR Purification Kit(toroivd,China)回收纯化约10000bp大小的线性片段。而后用T4 Polynucleotide Kinase(NEB,USA)处理回收的pTY-S质粒,再用GEL/PCR Purification Kit(toroivd,China)纯化回收去磷酸化的pTY-S质粒。
pTY-S质粒,在A Protocol for VIGS in Arabidopsis Thaliana Using a One-Step TYMV-Derived Vector中被告知。
处理寡核苷酸。首先,95℃水浴寡核苷酸5min,并让其自然冷却至室温形成双链。而后在37℃的金属浴中用Quick CIP(NEB,USA)处理寡核苷酸序列30min,使用GEL/PCRPurification Kit(toroivd,China)对磷酸化的寡核苷酸进行纯化回收。
参照DNA Ligation Kit Ver.2.1(Takara,Japan)使用说明,将处理过的pTY-S质粒和80bp寡核苷酸进行连接,并将连接产物转化大肠杆菌DH5α(唯地,中国),PCR鉴定后将阳性克隆送到北京擎科生物科技有限公司测序,测序正确的载体命名为pTY-S/BoaWRKY4,为BoaWRKY4.1、BoaWRKY4.2和BoaWRKY4.3这三个基因同源的序列构的载体。该载体序列为:GGTACTCTGGATCCTCTGGGTCACTATCCATCCCACCTTGCAAGCCCCAGCTTTCCCGACCACGGTCGGTGTCTGCTGGGTACCCGCCAATTCTCCAGTCACTCCCGCCCAAATCACCAAGACCTACGGTGGCCAGATCTTCTGCATTGGCGGCGCCATCAACACCCTCTCACCTCTCATCGTCAAGTGCCCACTTGAAATGATGAACCCCCGGGTCAAGGATTCGATTCAGTACCTTGACTCGCCCAAACTCCTCATCTCCATCACCGCTCAACCCACCGCTCCCCCCGCATCGACCTGCATAATAACTGTATCAGGAACTCTCTCGATGCACTCTCCGCTCATCACGGACACTTCTACTTGACCCGAGATTCAAGCAGAGCAGACCCACCGGTTTGATATCAAACCGGTGGGTCTGCTCTGCTTGAATCTCGGGTCAAGTAAATTCTCCATTTTAAAATCGTTAGCCTCCCCAAGAAAGCGAGAGCTGTCCCAAT
三、BoaWRKY4 VIGS基因沉默芥蓝的构建
将芥蓝播种于营养钵中,置于温度25℃,光照16h,黑暗8h的培养室,待芥蓝长到三叶一心时用于实验。侵染前浇足水并对植株进行16-24h的暗处理。侵染时,首先将少量碳化硅均匀撒在芥蓝的叶片上,轻轻摩擦3-4次造成机械损伤,用ddH2O将碳化硅冲洗掉,再用吸水纸将多余的水分吸干净。而后吸取约10μl(约1μg)的pTY-S/BoaWRKY4质粒滴在损伤处,1-2min钟后用ddH2O冲洗损伤处,再用吸水纸将多余的水分吸干净。接种后先暗处理12h,再将芥蓝植株放回光照16h,黑暗8h的培养室,待1-2周后长出新的叶片,再进行下述步骤。对照植株采取相同的处理方法,唯一不同的地方在于用pTY-S质粒滴在损伤处。
备注说明:选取BoaWRKY4.1、BoaWRKY4.2和BoaWRKY4.3表达量都显著低于对照株系pTY-S的两个基因沉默株系pTY-S/BoaWRKY4-1和pTY-S/BoaWRKY4-2作为下述步骤的研究对象。
四、RNA的提取和表达分析
称取约0.1g的新长出来的叶片,液氮冷冻3min,用研磨仪(60赫兹,120秒)将样品磨成粉末,加入1mL Trizol(Takara,Japan),涡旋振荡仪震荡3min,冰上静置3min。加入200μL氯仿,涡旋振荡仪震荡2min。4℃离心机12000rpm离心15分钟,吸取上清至新的去除RNA降解酶的离心管中,此步骤重复一次。而后加入等体积的异丙醇,放至于-20℃冰箱静置1h,4℃离心机12000rpm离心10min。弃上清,加入1mL的75%乙醇洗涤沉淀,4℃离心机12000rpm离心5min,此步骤重复一次,至于通风橱中10分钟,加入30μLDEPC水溶解RNA。使用NanoDrop(Thermo,USA)测量浓度。而后使用PrimeScriptTMRT Master Mix(Perfect Real Time)(Takara,Japan)将RNA反转录为cDNA。实时荧光定量PCR(Real-time qPCR)使用SYBR GreenPCR Master Mix(Takara,Japan),反应体系为:1μl稀释2倍的cDNA,引物各0.5μl(10μM)、10μl TB Premix Ex TaqTMII(Tli RNaseH Plus)(Takara,Japan)以及7μl的ddH2O。使用CFX96 Real-Time PCR Detection System(Bio-Rad)进行分析。qPCR程序为:95℃30s;95℃5s,60℃30s(39个循环);95℃10s,65℃5s,95℃5s。使用芥蓝β-BoaACTIN作为内参基因,基因相对表达水平采用2-△△CT方法计算。
β-BoaACTIN-F:CCAGAGGTCTTGTTCCAGCCATC
β-BoaACTIN-R:GTTCCACCACTGAGCACAATGTTAC
RT-BoaMYB51.1-F TTCGGAGTTAACGGTGACACTTG
RT-BoaMYB51.1-R GTTCTTAAACAGAGTAGCTAATAGGTTC
RT-BoaMYB51.2-F CCTTCACGGCAACAAATGGTCTGAT
RT-BoaMYB51.2-R CGGGTCGACACCTTTCTTGATCAA
RT-BoaCYP83B1-F TTAGAGAAGAAGAGTGCCAACG
RT-BoaCYP83B1-R CGAAATAAGGGAAAAGGTCGGA
RT-BoaMYB28.1-F GTTGCGGCTAAGGTCACTTCT
RT-BoaMYB28.1-R CAGAAGTAGCGTTGATCTCATGC
RT-BoaMYB29-F CGCCCAAGACTTCTGAGTT
RT-BoaMYB29-R TGATATTGCCCATGGAAGCTG
RT-BoaCYP79F1-F CCCATCCTAGGCAATTTACC
RT-BoaCYP79F1-R TGATTGCCTCCGATGGTTC
说明:MYB51.1和MYB51.2是调控吲哚族芥子油苷合成的转录因子,CYP83B1是参与吲哚族芥子油苷合成的结构基因;MYB28和MYB29是调控脂肪族芥子油苷合成的转录因子,CYP79F1是参与脂肪族芥子油苷合成的结构基因。
根据拟南芥、甘蓝和芜菁三种芸薹属蔬菜WRKY4蛋白质序列的进化树(图1),可以得知在芥蓝中有3个同源的WRKY4基因(BoaWRKY4.1、BoaWRKY4.2、BoaWRKY4.3)。
根据拟南芥AtWRKY4,芥蓝的BoaWRKY4.1、BoaWRKY4.2和BoaWRKY4.3和芥蓝的Bol031460、Bol010361和Bol029198蛋白质序列同源性分析(图2),可以得知芥蓝的BoaWRKY4.1与甘蓝Bol031460的氨基酸序列同源性高达99.3197%,BoaWRKY4.2与甘蓝Bol010361的氨基酸序列同源性高达95.6395%,BoaWRKY4.3与甘蓝Bol029198的氨基酸序列同源性高达99.2574%。其中芥蓝BoaWRKY4.1同拟南芥AtWRKY4氨基酸同源性最高,同源性大约为80%。
根据BoaWRKY4.1、BoaWRKY4.2和BoaWRKY4.3在芥蓝中的组织特异性表达(图3),可以得知BoaWRKY4.1、BoaWRKY4.2和BoaWRKY4.3在种子(Seed)中的表达量最高,在根(Root)中的表达量最低,在花(Flower)、叶(Leaf)和角果(Silque)中的表达量无显著性差异。
五、核盘菌菌核病病害研究
共试核盘菌[Sclerotinia sclerotiorum(Lib.)de Bary]为本实验室从芥蓝田间分离纯化得到。将核盘菌的菌核一分为二切开接种在马铃薯葡萄糖琼脂培养基(PDA)平板中央处,置于25℃生化培养箱暗培养4d左右,待菌丝体增殖接近覆盖整个培养基时,用于接种离体的芥蓝叶片,共试芥蓝品种为“大笋迟花”。用打孔器(孔径为5mm)在培养核盘菌的PDA培养基靠近边缘的同一半径处打孔,将菌丝面朝下放在离体的芥蓝叶片上。用湿润的棉球包裹芥蓝叶柄,保持湿润,放置于密封的玻璃培养皿中。接种后24h到48h后观察发病情况并统计菌斑面积大小,每组实验至少重复3次。菌斑面积统一拍照后,用ImageJ软件进行测量和统计。
根据图4可以得知,pTY-S/BoaWRKY4-1和pTY-S/BoaWRKY4-2两个株系中的BoaWRKY4.1、BoaWRKY4.2和BoaWRKY4.3基因表达量远低于对照pTY-S株系。核盘菌侵染后BoaWRKY4基因沉默株系pTY-S/BoaWRKY4-1和pTY-S/BoaWRKY4-2比对照pTY-S株系发病更为严重,病斑面积也显著高于对照pTY-S株系。
根据图5可以得知,pTY-S/BoaWRKY4-1和pTY-S/BoaWRKY4-2两个株系中吲哚族芥子油苷相关基因BoaMYB51.1、BoaMYB51.2以及BoaCYP83B1显著低于对照pTY-S株系,以及脂肪族芥子油苷相关基因BoaMYB28.1、BoaMYB29和BoaCYP79F1显著低于对照pTY-S株系。
根据图4和图5,可获得以下总结性结论:BoaWRKY4不仅可以正调控吲哚族芥子油苷和脂肪族芥子油苷相关基因的表达,还在对抗核盘菌中发挥着重要的作用。
综上所述,基因BoaWRKY4沉默后,不仅能使芥子油苷合成相关基因的表达量降低,还能减弱芥蓝对核盘菌的抗性。
最后,还需要注意的是,以上列举的仅是本发明的若干个具体实施例。显然,本发明不限于以上实施例,还可以有许多变形。本领域的普通技术人员能从本发明公开的内容直接导出或联想到的所有变形,均应认为是本发明的保护范围。
序列表
<110> 浙江大学
<120> 芥蓝BoaWRKY4基因及其应用
<160> 6
<170> SIPOSequenceListing 1.0
<210> 1
<211> 1458
<212> DNA
<213> 芥蓝(Brassica oleracea var. alboglabra Bailey)
<400> 1
atgtcggaga aggaagaact cccgtcgaca tctaagtcca ccggagctcc gtcacgaccg 60
actttatctc ttcctccacg gccgtttagt gagatgttct tcaacgctgg cgttggattc 120
agccctggtc cgatgactct ggtctcaaac atgttccccg agtccgatga gtttaggtct 180
ttctctcagc tcctcgccgg cgccatggct tctcccgcgg ctgcagctgc tgcatcagcc 240
gggacggcta gtgaagaagg gaataatagc tcgagcggtg gtgatgttga cccgagattc 300
aagcagagca gacccaccgg tttgatgatt tcgcagtcac cgtcgatgtt caccgtaccg 360
ccagggctaa gtccggcgat gcggctggat tcaccgagct tcttgggtct tttctctccc 420
gttcagggat catatggaat gacacatcag caagctctag cccaagtaac agcacaagca 480
gttcaagcca atgctaatat gcaaccacaa accgagtacc cttcttcctc tcaggttcaa 540
tcattttcat ctggaccgac cacagctaca gattcatcat taccagctcc aagagaaacc 600
tcaattacaa ccatcattga gcagaggtca caacaacagc ctctaaacgt tgacaaacca 660
gctgatgatg gctataactg gcgaaagtac ggacaaaagc aagttaaagg cagcgagttt 720
cctcgaagct attacaagtg caccaatcca ggatgtcctg tcaagaagaa ggtcgagaga 780
tcacttgatg gacaagtaac tgaaatcatc tacaagggcc agcacaatca tgagcctcct 840
cagaacacta agcgaggcaa caacaacaac aaagatagta atctaaacgg gagttcggtt 900
aataacaata agagtaagag ggaacaacat gaagcagcga gtcaagctac tactacagag 960
caaatgtctg aggctagtga cagcgaagaa gttggtaatg gagaaagtgg tgtcagagag 1020
aaagctgaag acgagcctga cgccaagaaa gttcgagttt cggaaccagc tgctgttgtt 1080
tctacttcac atagaactgt gacagagcct aagatcatcg tccaaacgac aagtgaagtt 1140
gatctgttag atgatgggta taggtggcgc aagtatggac agaaagtggt caaagggaat 1200
ccttatccga ggagttacta caagtgcaca acacagggat gtggagtgag gaaacatgta 1260
gagagagcag cagcagatcc aaaagctgta gtgacaacat atgaaggtaa acataaccat 1320
gaccttcctg cgtctaaatc aagaagccat gctgcagcag cagcagcagc acagttaagg 1380
ccagaaaatc gacatggcgg tttggctaat cagcagcctg ttgcgcgtct aaggctcaaa 1440
gaagagcaca tagtttaa 1458
<210> 2
<211> 1461
<212> DNA
<213> 芥蓝(Brassica oleracea var. alboglabra Bailey)
<400> 2
atgtcggaga aagaagaacg tccgtcgact tccaacaagt ccaccggacc tccgtcgcga 60
ccgaccttat ctctccctcc acggcccttt agtgagatgt tcttcagcgg cggagttgga 120
ttcagccctg gtccgatgac tctggtctcg aacatgttcc ccgattccga tgagtttagg 180
tctttctccc agctcctttc cggagtcatg ccttctccag cagctgcatc ttccgcagct 240
agtgaagaag ggggtaataa cagctcgagc ggtggtgatg ttgacccgag attcaagcag 300
agcagaccca ccggtttgat gatttcgcag tctccgtcga cgatgttcgc cgtaacgcca 360
gggctgagtc cggcgatgcg gctggattca ccgagcttct tgggtctttt ctctcccctt 420
cagggatcat atggaatgac acatcagcaa gctctagcac aagtcacagc acaagctgtt 480
caagccaatg ctaatatgca accacaaacc gagtaccctc cttcctctca ggtttcatct 540
gttcaacctc agaaccccac cacagctcct gattcttcag ctccaagaaa aacctcggat 600
acaaccatca tcgagcacag gtcacagcag cctctaaatg ttgacaaacc agctgatgat 660
ggctataact ggagaaagta cggacaaaag caagttaaag gcagcgagtt ccctcgaagc 720
tattacaagt gcacgaatcc agggtgtcct gtcaagaaaa aggttgagag atctctcgat 780
ggacaagtaa cggagatcat ctacaaaggt cagcacaatc atgaacctcc tcaaaacact 840
aggcgaggca acagagatag cacagctaat ctaaatggga gttcggttaa taacaaaact 900
acgagggaac aacatgaagc agcagcaagt caagctacga cagagcaaat gtctgaggca 960
agtgacagtg aagaagttgg taatggagaa actggtgtga ggaagaaagc tgaagacgag 1020
cctgatgcca agagaagaag tacagaagtt cgagtttcag aaccagctgc tgctgctgct 1080
gcttcacata gaactgtgac agagcctaga atcattgtcc aaacgacaag tgaagttgat 1140
ctattagatg atggatacag gtggcgtaaa tatggacaga aggttgtcaa agggaatcct 1200
tatccgagga gttactacaa gtgcacaaca cagggatgtg gagtgaggaa acatgtggag 1260
agagcagcaa cagatccaaa agctgtagta acaacatatg aagggaaaca taaccatgac 1320
cttcctgcat ctaaatcaag cagccatgcc gctgcagctg cagctgcggc acagttaagg 1380
ccagaaaatc gtttggctaa cttaaatcag cagcagcagc cggttgcgcg tctaaggctg 1440
aaagaagagc aaataattta a 1461
<210> 3
<211> 1386
<212> DNA
<213> 芥蓝(Brassica oleracea var. alboglabra Bailey)
<400> 3
atgtcggaga aggaagatct tccgtcgaca tcgaagtcca ccggagctgc tccgtcgcga 60
cccactttat atcttcctcc acggccattt actgagacgt tcttcaacgg tggcgtcgga 120
ttcagccctg gtccgatgac tctggtctca aacatgttct ccgattccga tgagtctagt 180
aggtctttct cccagctcct tgccggagtc gtcatgcctt ctcctgcaac gggtagtgaa 240
gggaactgta atagctctag tgttgatgta gttgacccga gattcaagca gagcagaccc 300
accggtttga tggtttctca gtcaccgtcg gtattcaccg tacctccagg gctaagtccg 360
gcgatgttgc tggattcacc gagcttcttg ggtcttttct ctcccgttca gggatcatat 420
ggaatgacac atcagcaagc tctagcacaa gtaacagcac aagctgttca agccaatgct 480
aatatgcaac ctcaaacaga gcaccctcct tcctctcagg ttcaatctca gaacccaacc 540
tcagctccag attcttcaca gctagctcaa agagaaacct cggatataaa catcatcgag 600
cacaacaggt cacaacaaca gcctctgaat gttgacaaac cagcaggtga tggctacaac 660
tggcgaaagt atggtcaaaa gcaagttaaa ggcagtgagt tccctcgaag ctattacaag 720
tgcaccaatc ctggatgtcc tgtcaagaag aaggttgaga ggtctcttga tggacaagta 780
actgaaatca tctacaaagg tcagcacaat cacgaacctc ctcaaaacaa taagcgtgga 840
aacaaagata gcaataacaa tcctaacaaa accacgaggg aacaacatga agcagcaagt 900
caagctacga cagagcaaat gtctgaggca agtgacagcg aagaagtcgg taacggagaa 960
gagcctgatc ctaagagaag aagtacagag gttcgagttc tggaaccagc tgctgctgtt 1020
gctgcttcac atagaacagt ggcagagcct agaattattg ttcagacaac aagtgaagtt 1080
gatcttctag acgatggata taggtggcgt aagtatggtc agaaaattgt taaagggaat 1140
ccttatccga ggagttatta caagtgtact acacagggat gtggagtgag gaaacacgtg 1200
gagagagcag caacagatcc aaaagctgta gtgacaacgt acgaaggaaa acataaccat 1260
gaccctccag cagctaaatc aagcagccat gctgctgttg ctgctgcggc acagttaagg 1320
catggcggtt tggctaactt aaatcagcag cagcagcagc ccgttgcgtg tctaaggctg 1380
aaataa 1386
<210> 4
<211> 485
<212> PRT
<213> 芥蓝(Brassica oleracea var. alboglabra Bailey)
<400> 4
Met Ser Glu Lys Glu Glu Leu Pro Ser Thr Ser Lys Ser Thr Gly Ala
1 5 10 15
Pro Ser Arg Pro Thr Leu Ser Leu Pro Pro Arg Pro Phe Ser Glu Met
20 25 30
Phe Phe Asn Ala Gly Val Gly Phe Ser Pro Gly Pro Met Thr Leu Val
35 40 45
Ser Asn Met Phe Pro Glu Ser Asp Glu Phe Arg Ser Phe Ser Gln Leu
50 55 60
Leu Ala Gly Ala Met Ala Ser Pro Ala Ala Ala Ala Ala Ala Ser Ala
65 70 75 80
Gly Thr Ala Ser Glu Glu Gly Asn Asn Ser Ser Ser Gly Gly Asp Val
85 90 95
Asp Pro Arg Phe Lys Gln Ser Arg Pro Thr Gly Leu Met Ile Ser Gln
100 105 110
Ser Pro Ser Met Phe Thr Val Pro Pro Gly Leu Ser Pro Ala Met Arg
115 120 125
Leu Asp Ser Pro Ser Phe Leu Gly Leu Phe Ser Pro Val Gln Gly Ser
130 135 140
Tyr Gly Met Thr His Gln Gln Ala Leu Ala Gln Val Thr Ala Gln Ala
145 150 155 160
Val Gln Ala Asn Ala Asn Met Gln Pro Gln Thr Glu Tyr Pro Ser Ser
165 170 175
Ser Gln Val Gln Ser Phe Ser Ser Gly Pro Thr Thr Ala Thr Asp Ser
180 185 190
Ser Leu Pro Ala Pro Arg Glu Thr Ser Ile Thr Thr Ile Ile Glu Gln
195 200 205
Arg Ser Gln Gln Gln Pro Leu Asn Val Asp Lys Pro Ala Asp Asp Gly
210 215 220
Tyr Asn Trp Arg Lys Tyr Gly Gln Lys Gln Val Lys Gly Ser Glu Phe
225 230 235 240
Pro Arg Ser Tyr Tyr Lys Cys Thr Asn Pro Gly Cys Pro Val Lys Lys
245 250 255
Lys Val Glu Arg Ser Leu Asp Gly Gln Val Thr Glu Ile Ile Tyr Lys
260 265 270
Gly Gln His Asn His Glu Pro Pro Gln Asn Thr Lys Arg Gly Asn Asn
275 280 285
Asn Asn Lys Asp Ser Asn Leu Asn Gly Ser Ser Val Asn Asn Asn Lys
290 295 300
Ser Lys Arg Glu Gln His Glu Ala Ala Ser Gln Ala Thr Thr Thr Glu
305 310 315 320
Gln Met Ser Glu Ala Ser Asp Ser Glu Glu Val Gly Asn Gly Glu Ser
325 330 335
Gly Val Arg Glu Lys Ala Glu Asp Glu Pro Asp Ala Lys Lys Val Arg
340 345 350
Val Ser Glu Pro Ala Ala Val Val Ser Thr Ser His Arg Thr Val Thr
355 360 365
Glu Pro Lys Ile Ile Val Gln Thr Thr Ser Glu Val Asp Leu Leu Asp
370 375 380
Asp Gly Tyr Arg Trp Arg Lys Tyr Gly Gln Lys Val Val Lys Gly Asn
385 390 395 400
Pro Tyr Pro Arg Ser Tyr Tyr Lys Cys Thr Thr Gln Gly Cys Gly Val
405 410 415
Arg Lys His Val Glu Arg Ala Ala Ala Asp Pro Lys Ala Val Val Thr
420 425 430
Thr Tyr Glu Gly Lys His Asn His Asp Leu Pro Ala Ser Lys Ser Arg
435 440 445
Ser His Ala Ala Ala Ala Ala Ala Ala Gln Leu Arg Pro Glu Asn Arg
450 455 460
His Gly Gly Leu Ala Asn Gln Gln Pro Val Ala Arg Leu Arg Leu Lys
465 470 475 480
Glu Glu His Ile Val
485
<210> 5
<211> 486
<212> PRT
<213> 芥蓝(Brassica oleracea var. alboglabra Bailey)
<400> 5
Met Ser Glu Lys Glu Glu Arg Pro Ser Thr Ser Asn Lys Ser Thr Gly
1 5 10 15
Pro Pro Ser Arg Pro Thr Leu Ser Leu Pro Pro Arg Pro Phe Ser Glu
20 25 30
Met Phe Phe Ser Gly Gly Val Gly Phe Ser Pro Gly Pro Met Thr Leu
35 40 45
Val Ser Asn Met Phe Pro Asp Ser Asp Glu Phe Arg Ser Phe Ser Gln
50 55 60
Leu Leu Ser Gly Val Met Pro Ser Pro Ala Ala Ala Ser Ser Ala Ala
65 70 75 80
Ser Glu Glu Gly Gly Asn Asn Ser Ser Ser Gly Gly Asp Val Asp Pro
85 90 95
Arg Phe Lys Gln Ser Arg Pro Thr Gly Leu Met Ile Ser Gln Ser Pro
100 105 110
Ser Thr Met Phe Ala Val Thr Pro Gly Leu Ser Pro Ala Met Arg Leu
115 120 125
Asp Ser Pro Ser Phe Leu Gly Leu Phe Ser Pro Leu Gln Gly Ser Tyr
130 135 140
Gly Met Thr His Gln Gln Ala Leu Ala Gln Val Thr Ala Gln Ala Val
145 150 155 160
Gln Ala Asn Ala Asn Met Gln Pro Gln Thr Glu Tyr Pro Pro Ser Ser
165 170 175
Gln Val Ser Ser Val Gln Pro Gln Asn Pro Thr Thr Ala Pro Asp Ser
180 185 190
Ser Ala Pro Arg Lys Thr Ser Asp Thr Thr Ile Ile Glu His Arg Ser
195 200 205
Gln Gln Pro Leu Asn Val Asp Lys Pro Ala Asp Asp Gly Tyr Asn Trp
210 215 220
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225 230 235 240
Tyr Tyr Lys Cys Thr Asn Pro Gly Cys Pro Val Lys Lys Lys Val Glu
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Arg Ser Leu Asp Gly Gln Val Thr Glu Ile Ile Tyr Lys Gly Gln His
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Asn His Glu Pro Pro Gln Asn Thr Arg Arg Gly Asn Arg Asp Ser Thr
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Ala Asn Leu Asn Gly Ser Ser Val Asn Asn Lys Thr Thr Arg Glu Gln
290 295 300
His Glu Ala Ala Ala Ser Gln Ala Thr Thr Glu Gln Met Ser Glu Ala
305 310 315 320
Ser Asp Ser Glu Glu Val Gly Asn Gly Glu Thr Gly Val Arg Lys Lys
325 330 335
Ala Glu Asp Glu Pro Asp Ala Lys Arg Arg Ser Thr Glu Val Arg Val
340 345 350
Ser Glu Pro Ala Ala Ala Ala Ala Ala Ser His Arg Thr Val Thr Glu
355 360 365
Pro Arg Ile Ile Val Gln Thr Thr Ser Glu Val Asp Leu Leu Asp Asp
370 375 380
Gly Tyr Arg Trp Arg Lys Tyr Gly Gln Lys Val Val Lys Gly Asn Pro
385 390 395 400
Tyr Pro Arg Ser Tyr Tyr Lys Cys Thr Thr Gln Gly Cys Gly Val Arg
405 410 415
Lys His Val Glu Arg Ala Ala Thr Asp Pro Lys Ala Val Val Thr Thr
420 425 430
Tyr Glu Gly Lys His Asn His Asp Leu Pro Ala Ser Lys Ser Ser Ser
435 440 445
His Ala Ala Ala Ala Ala Ala Ala Ala Gln Leu Arg Pro Glu Asn Arg
450 455 460
Leu Ala Asn Leu Asn Gln Gln Gln Gln Pro Val Ala Arg Leu Arg Leu
465 470 475 480
Lys Glu Glu Gln Ile Ile
485
<210> 6
<211> 461
<212> PRT
<213> 芥蓝(Brassica oleracea var. alboglabra Bailey)
<400> 6
Met Ser Glu Lys Glu Asp Leu Pro Ser Thr Ser Lys Ser Thr Gly Ala
1 5 10 15
Ala Pro Ser Arg Pro Thr Leu Tyr Leu Pro Pro Arg Pro Phe Thr Glu
20 25 30
Thr Phe Phe Asn Gly Gly Val Gly Phe Ser Pro Gly Pro Met Thr Leu
35 40 45
Val Ser Asn Met Phe Ser Asp Ser Asp Glu Ser Ser Arg Ser Phe Ser
50 55 60
Gln Leu Leu Ala Gly Val Val Met Pro Ser Pro Ala Thr Gly Ser Glu
65 70 75 80
Gly Asn Cys Asn Ser Ser Ser Val Asp Val Val Asp Pro Arg Phe Lys
85 90 95
Gln Ser Arg Pro Thr Gly Leu Met Val Ser Gln Ser Pro Ser Val Phe
100 105 110
Thr Val Pro Pro Gly Leu Ser Pro Ala Met Leu Leu Asp Ser Pro Ser
115 120 125
Phe Leu Gly Leu Phe Ser Pro Val Gln Gly Ser Tyr Gly Met Thr His
130 135 140
Gln Gln Ala Leu Ala Gln Val Thr Ala Gln Ala Val Gln Ala Asn Ala
145 150 155 160
Asn Met Gln Pro Gln Thr Glu His Pro Pro Ser Ser Gln Val Gln Ser
165 170 175
Gln Asn Pro Thr Ser Ala Pro Asp Ser Ser Gln Leu Ala Gln Arg Glu
180 185 190
Thr Ser Asp Ile Asn Ile Ile Glu His Asn Arg Ser Gln Gln Gln Pro
195 200 205
Leu Asn Val Asp Lys Pro Ala Gly Asp Gly Tyr Asn Trp Arg Lys Tyr
210 215 220
Gly Gln Lys Gln Val Lys Gly Ser Glu Phe Pro Arg Ser Tyr Tyr Lys
225 230 235 240
Cys Thr Asn Pro Gly Cys Pro Val Lys Lys Lys Val Glu Arg Ser Leu
245 250 255
Asp Gly Gln Val Thr Glu Ile Ile Tyr Lys Gly Gln His Asn His Glu
260 265 270
Pro Pro Gln Asn Asn Lys Arg Gly Asn Lys Asp Ser Asn Asn Asn Pro
275 280 285
Asn Lys Thr Thr Arg Glu Gln His Glu Ala Ala Ser Gln Ala Thr Thr
290 295 300
Glu Gln Met Ser Glu Ala Ser Asp Ser Glu Glu Val Gly Asn Gly Glu
305 310 315 320
Glu Pro Asp Pro Lys Arg Arg Ser Thr Glu Val Arg Val Leu Glu Pro
325 330 335
Ala Ala Ala Val Ala Ala Ser His Arg Thr Val Ala Glu Pro Arg Ile
340 345 350
Ile Val Gln Thr Thr Ser Glu Val Asp Leu Leu Asp Asp Gly Tyr Arg
355 360 365
Trp Arg Lys Tyr Gly Gln Lys Ile Val Lys Gly Asn Pro Tyr Pro Arg
370 375 380
Ser Tyr Tyr Lys Cys Thr Thr Gln Gly Cys Gly Val Arg Lys His Val
385 390 395 400
Glu Arg Ala Ala Thr Asp Pro Lys Ala Val Val Thr Thr Tyr Glu Gly
405 410 415
Lys His Asn His Asp Pro Pro Ala Ala Lys Ser Ser Ser His Ala Ala
420 425 430
Val Ala Ala Ala Ala Gln Leu Arg His Gly Gly Leu Ala Asn Leu Asn
435 440 445
Gln Gln Gln Gln Gln Pro Val Ala Cys Leu Arg Leu Lys
450 455 460
Claims (1)
1.芥蓝BoaWRKY4基因的用途,其特征在于:BoaWRKY4正调控吲哚族芥子油苷和脂肪族芥子油苷相关基因的表达;
吲哚族芥子油苷相关基因为BoaMYB51.1、BoaMYB51.2和BoaCYP83B1,脂肪族芥子油苷相关基因为BoaMYB28.1、BoaMYB29和BoaCYP79F1;
BoaWRKY4为BoaWRKY4.1、BoaWRKY4.2或BoaWRKY4.3;
BoaWRKY4.1的核苷酸序列如SEQ ID NO:1所述;
BoaWRKY4.2的核苷酸序列如SEQ ID NO:2所述;
BoaWRKY4.3的核苷酸序列如SEQ ID NO:3所述。
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