CN114941008B - 菜心LRR受体蛋白激酶基因BraEFR在抗霜霉病中的应用 - Google Patents
菜心LRR受体蛋白激酶基因BraEFR在抗霜霉病中的应用 Download PDFInfo
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Abstract
本发明公开了菜心LRR受体蛋白激酶基因BraEFR在抗霜霉病中的应用，所述菜心BraEFR基因的核苷酸序列如SEQ ID NO：2所示。本发明首次发现了菜心LRR受体蛋白激酶基因BraEFR可以调控菜心抗霜霉病抗性。而且本发明中还提供了可用于调节菜心抗霜霉病抗性的产品，其能够通过上调菜心中BraEFR基因的表达从而达到提高菜心抗霜霉病抗性的作用，从而可以将其用于预防菜心霜霉病、培育抗菜心霜霉病转基因植物等相关方面，具有极高的实用价值。
Description
技术领域
本发明属于植物分子生物学与植物遗传工程领域，具体涉及菜心LRR受体蛋白激酶基因BraEFR在抗霜霉病中的应用。
背景技术
菜心霜霉病是由寄生霜霉菌(hyaloperonospora brassucae)所引发的菜心专性寄生性病害，在菜心苗期、成株期、开花期至结荚期均可发生，主要危害菜心叶片、种荚、花梗、茎顶等。发病一般由菜心的下部叶片开始，初呈水渍状褪绿病斑，逐渐扩大，受叶脉限制呈现黄褐色多角形斑。天气潮湿时病斑背面产生疏密不等的白色霉状物，严重时病斑融合成片。发病后期病斑破裂，病叶干枯变为暗褐色。当菜心采种株的茎顶及花梗染病时，多呈肥肿畸形，俗称“龙头拐”，种荚染病后会出现不同程度的变形。菜心霜霉病在20-24℃和高湿条件(如华南地区的冬季)下易发，降雨量比温度对病害的发展影响更为重要。生长发育较差、种植密度大或已感染其他病毒病的植株更易发生霜霉病，且往往发病更为严重。
因此，开发或培育抗病菜心品种是控制菜心霜霉病最经济、环保、有效的途径之一，而挖掘抗霜霉病基因(或蛋白)则是抗病育种的基础。
发明内容
本发明旨在至少解决上述现有技术中存在的技术问题之一。为此，本发明提出菜心LRR受体蛋白激酶基因BraEFR在抗霜霉病中的应用，发明人发现菜心LRR受体蛋白激酶基因BraEFR对菜心抗霜霉病具有重要作用，可用于预防菜心霜霉病、制备抗菜心霜霉病的产品、培育抗菜心霜霉病转基因植物，具有广泛的应用前景。
本发明的第一个方面，提供调控菜心BraEFR基因表达的产品在调控作物抗菜心霜霉病中的应用，所述菜心BraEFR基因的核苷酸序列如SEQ ID NO：2所示。
发明人首次发现了一种菜心LRR受体蛋白激酶基因BraEFR，并通过试验验证该基因对菜心霜霉病具有抗性调节作用，其在抗霜霉病菜心材料(R128)中受霜霉病菌诱导的表达量显著高于感霜霉病菜心材料(S002)受霜霉病菌诱导的表达量，且在过量表达BraEFR基因的菜心叶片上接种菜心霜霉病菌，叶片感染面积比对照显著减小，而在沉默表达BraEFR基因的菜心叶片上接种菜心霜霉病菌，叶片感染面积比对照显著增大。从而说明菜心LRR受体蛋白激酶基因BraEFR对菜心抗霜霉病具有重要作用。
根据本发明的第一个方面，在本发明的一些实施方式中，所述产品具有下述(1)～(5)中至少一项所述的功能；
(1)上调菜心BraEFR基因表达；
(2)下调或阻断菜心BraEFR基因表达；
(3)促进菜心BraEFR蛋白的分泌；
(4)抑制菜心BraEFR蛋白的分泌；
(5)靶向结合菜心BraEFR蛋白受体或靶向结合菜心BraEFR蛋白。
在本发明的一些实施方式中，所述菜心BraEFR蛋白的氨基酸序列如SEQ ID NO：3所示。
根据本发明的第一个方面，在本发明的一些实施方式中，所述产品包括下述(1)～(6)中任意一项：
(1)含有SEQ ID NO：1所示核酸分子的表达盒；
(2)含有SEQ ID NO：1所示核酸分子的微生物或细胞；
(3)含有(1)中所述表达盒的微生物或细胞；
(4)含有SEQ ID NO：4所示核酸分子的表达盒；
(5)含有SEQ ID NO：4所示核酸分子的微生物或细胞；
(6)含有(4)中所述表达盒的微生物或细胞。
在本发明的一些实施方式中，所述作物为菜心。
发明人发现，上述BraEFR基因能够正调控叶菜霜霉病的抗性，尤其是对于菜心而言，经试验验证发现其在过量表达BraEFR基因的菜心叶片上接种菜心霜霉病菌，叶片感染面积比对照显著减小，而在沉默表达BraEFR基因的菜心叶片上接种菜心霜霉病菌，叶片感染面积比对照显著增大。而且菜心中的寄生霜霉菌具有极强的专化性，不会发生跨种属传播的情况，因此，通过上述产品能够大大的提高菜心在抗霜霉病中的抗病力。
本发明的第二个方面，提供一种试剂，所述试剂中包括下述(1)～(3)中任意一项：
(1)含有SEQ ID NO：1所示核酸分子的表达盒；
(2)含有SEQ ID NO：1所示核酸分子的微生物或细胞；
(3)含有(1)中所述表达盒的微生物或细胞。
根据本发明的第二个方面，在本发明的一些实施方式中，所述试剂用于提高菜心的菜心霜霉病抗性。
发明人发现，通过构建过表达质粒上调菜心中的BraEFR基因表达量，能够显著增强菜心的霜霉病抗性。
本发明的第三个方面，提供一种培育抗菜心霜霉病作物的方法，包括如下步骤：上调菜心中BraEFR基因的表达。
根据本发明的第三个方面，在本发明的一些实施方式中，所述BraEFR基因的核苷酸序列如SEQ ID NO：2或SEQ ID NO：1所示。
在本发明的一些实施方式中，所述BraEFR基因的核苷酸序列如SEQ ID NO：2所示。
在本发明的一些实施方式中，所述作物为菜心。
在本发明的一些优选实施方式中，所述方法具体为：
(1)构建BraEFR基因过表达载体；
(2)将BraEFR基因过表达载体转化农杆菌，筛选阳性克隆，感染作物，即得。
在本发明的一些优选实施方式中，步骤(1)中具体的构建方法为：将BraEFR基因构建到pGWB5质粒中，具体插入位置为35S启动子后的attR1和attR2之间(具体为通过基因重组交换替代了pGWB5质粒中的ccdB基因)，从而得到了BraEFR基因过表达载体。
本发明的有益效果是：
1.本发明首次发现了菜心LRR受体蛋白激酶基因BraEFR在调控菜心抗霜霉病抗性中的应用，从而可以将其用于预防菜心霜霉病、制备抗菜心霜霉病的产品、培育抗菜心霜霉病转基因植物，具有广泛的应用前景。
2.本发明提供了可用于调节菜心抗霜霉病抗性的产品，其能够通过上调菜心中BraEFR基因的表达从而达到提高菜心抗霜霉病抗性的作用，从而可以将其用于预防菜心霜霉病、培育抗菜心霜霉病转基因植物等相关方面，具有极高的实用价值。
附图说明
图1是BraEFR基因在菜心抗病材料R128和感病材料S002中转录表达分析图。
图2是pGWB5质粒图谱。
图3是pK7GWIW质粒图谱。
图4是BraEFR基因瞬时过量表达和空白对照(CK)的菜心叶片中BraEFR基因的表达情况。
图5是BraEFR基因沉默和空白对照(CK)的菜心叶片中BraEFR基因的表达情况。
图6是过量表达BraEFR基因的菜心叶片与空白对照(CK)叶片的抗性鉴定分析。
图7是沉默BraEFR基因的菜心叶片与空白对照(CK)叶片的抗性鉴定分析。
具体实施方式
为了使本发明的发明目的、技术方案及其技术效果更加清晰，以下结合具体实施方式，对本发明进行进一步详细说明。应当理解的是，本说明书中描述的具体实施方式仅仅是为了解释本发明，并非为了限定本发明。
所使用的实验材料和试剂，若无特别说明，均为常规可从商业途径所获得的耗材和试剂。
下述实施例中，所用的菜心材料均由广东省农业科学院蔬菜研究所农业新技术研究室提供。其中，所用的菜心霜霉病菌来自广州白云区钟落潭实验地菜心霜霉病病株，且经鉴定确认无误。
BraEFR基因在抗、感菜心材料中转录表达测试
菜心BraEFR基因是一种LRR受体丝氨酸/苏氨酸蛋白激酶基因，发明人是通过对菜心抗、感材料的转录组分析中发现的。为了确定该基因在不同品种菜心中均可稳定表达，发明人在霜霉病菌接种前后，对菜心BraEFR基因在菜心抗病材料R128(于2018年在广州柯木塱种业博览会上收集，现保存于广东省农业科学院设施农业研究所)和感病材料S002(于2018年在广州柯木塱种业博览会上收集，现保存于广东省农业科学院设施农业研究所)中的转录表达水平进行检测(数据来自转录组测序，由广州基迪奥生物科技有限公司负责测序)。
结果如图1所示。
可以发现，菜心BraEFR基因在菜心抗病材料R128和感病材料S002中的转录表达水平相近，且表达量均较低。而在接种霜霉病菌24小时后，BraEFR基因在R128中的转录表达水平约是S002中的2倍，显著高于在S002中的转录表达水平，说明菜心BraEFR基因是潜在的抗霜霉病基因。
BraEFR基因过表达载体和基因沉默载体的构建
基于上述发现，发明人进一步构建BraEFR基因过表达载体和基因沉默载体以用于进一步验证BraEFR基因与菜心霜霉病抗性之间的关系。
(1)BraEFR基因过表达载体的构建：
采用本领域中基因重组常规方法，将BraEFR基因CDS全长序列构建到pGWB5质粒(来自中科院遗传与发育生物学研究所唐定中课题组，pGWB5质粒图谱如图2所示)中，具体插入位置为35S启动子后的attR1和attR2之间(具体为通过基因重组交换替代了pGWB5质粒中的ccdB基因)，从而得到了BraEFR基因过表达载体pGWB5-BraEFR。
其中，本实施例中所用的BraEFR基因CDS全长序列长度为2880bp，具体核苷酸序列为：
5’-ATGAGCCAAAACCCACAATCACTTTCATTCTCTTGGGACCCAAACACATCTCCTTGTAATTGGACTGGCGTGACTTGCGACACACGTATCAGAAGAGTGACTTCCATCAACCTCTCTGGTCATGGTCTCACCGGTTCCATAAGCCCAAGTATTGGTAATCTCTCTTTTCTTACATCTCTCCAACTTCAAAATAATCAACTTCGAGGTCCAATTCCAAAAGAGATCACAAATCTTTTTCGTCTGAGGGTTCTTAATTTAAGCTCTAACAGCTTAGACGGATCCTTACCTTCCAACCTAAGCAAGTTAATAGAGCTACGAGCCCTTGATCTAACCTCAAATATGATCACGGGTATCGTTCCGAACCAACTTGGTGATCTCAAGAACCTAACTATTTTAAACTTGGGAAAAAATCTGTTGCATGGACCCATTCCACCATCTCTTTCAAACATTTCATCTCTTACCGTTTTAAGCCTAGGCACAAACTCTCTTAGTGGTCCAGTTCCTAACGAGTTGGGTCGTCTTCAAAGGCTTCAAGTCCTTGATCTCACCATAAACAACCTCTCTGGAACAATTCCTCCTTCTATTTACAACATGTCGTCTCTAGAATCATTGGTTATTGCTTCAAACAACTTTTGGGGTCAGTTTCCTAGCAACATAGGATACACACTTCCAAAACTCTTGGTTTTCAACGTGTGTTTCAACAAGTTCAGCGGAGAGATTCCAGCTTCTTTATACAACCTCACAAATATTAAGGTTATCCGCGCTGCACACAACCTCTTAGAAGGGACCATACCATCAGGATTAGGAAATCTCCCATTCCTAGAAATGTACAACATTGGATTCAATAAATTAGTTTGGGGTAGAGATCAAAATCTTGATAGTTTCATCAAGTCTTTCAGTAATAGCTCCAAACTTAACTTCCTTGCATTTGATGGAAACCTCTTGGAGGGTGTTATTCCGGTATCTATTGGCAATCTTCCGAAGAATCTTTCCAAACTTTTCATGGGAGGAAACCGCTTCACGGGTATTATTCCTGAGTCCATCGGCGATCTGACAGGATTGACTCTGTTTAACATCAGTGACAACTCACTCACTGGTGAAATCCCTCAAGATATTGGGAAATTGAAAGGTTTGCAAGTGTTAGAACTTGCGAGAAATCAGCTTACTGGGAGAATACCAGATTCTATTGGTGATCTTGGCGGGTTAAATGAGATCAATCTATCACACAACAAACTACAAGGTAGGATCTCACTATCATTTGAAAACTTCAAGAATATGTTATCCATGGATCTATCCAGCAATATGCTAAATGGAAGCATACCAAATGGAGTTCTTAACCTACCTAGCTTGAGTGCCGTCTTGAATCTATCTAGCAACCTATTTTCTGGTCCAATTCCTCAAGATATAAGTCGCCTCGAAAGCCTTGTGAGTCTAGATCTCTCTAGTAACAACTTCTCAGGCCACATCCCTTCATCTATCAAAGATTGTCAAAGCTTGGAGAAGCTAAACATGGCTGGCAATAATCTAGACGGACCAATCCCGGATGCATTAGCAGAAGTAAAAGGTCTAGAGTTTCTTGATCTCTCTTCAAACCAGCTTTCCGGTGTTATTCCTCCAAGACTTCAAGATTTACAAGCAATGAAGTTCTTGAATATTTCATTCAACAATCTTGAAGGATGGGTCCCAAACCGTGGAGTTTTCAAAGATCATTCTAAAGCTTATATGGAGGGGAATCCAAAGCTCTGCATTCACACTTGTCGAAAAACTCGGACGCACAGAAAACTTTTGAAAGTTAGCGTTATTACATGTGCGGTTGGTCTGATTGCAATATGTGTGATAAGTTTCTTGATTTGGAAAAGAATGGAAAAGAGGTCTACCACATCTACCTCATCATCAAATTCACTCCTCAAGGAACCATTTATGAATGTTTCATATGATGAGCTCAGGAGAGCTACTGAAAATTTCAACCCTAGAAATATTCTTGGTGTTGGAAGCTTTGGATCGGTATTCAAAGGGGTGATACGAGGGGTTGATGTTGCGGTGAAAGTCATTGATCTCAAGGCAAACGGATACTACAAAGGCTTTATAGCAGAGTGTGAGGCTTTAAGGAATGTAAGGCACAGGAATCTTGTGAAACTAGTAACTTCTTGTTCAAGCATAGACTTCAAAAACAATGAGTTTTTGGCTTTGGTGTATGAATTTTTAAGCAATGGAAGCTTAGAAGAATGGATCAAAGGTAAACATAGAAAACCTGATGGGAGTGTTGGGTTAAATCTTGAAGAAAGAGTGAATGTAGCCATTGACATTGCATCTGCATTGGATTATTTGCACAATGATTGTGAAGTTCATGTGGTTCATTGTGATCTAAAACCTAGCAATATTCTTCTAACCGAAGACATGGTTACAAAGGTCGGCGATTTTGGGTTGGCTCGTGTGTTGTTTGACGCATCCGATGATTGTCATCACGCTTCTATAAGCTCCACACATGTCTTGAAAGGCTCCATTGGTTATATTCCTCCAGAGTATGGGCTTGGAGAAAAGCCATCACAAGCAGGAGATGTATATAGCTTTGGAGTGATGTTGTTAGAGCTCTTGAGTGGAAAAAGCCCAATGGACGAGCGCTTCGAAGGAGATCAAAATCTAATATCATGGATAAGTTATGGGTTTCAAAGTAACGCAATTATGGAAGTGATTGATCCTAAACTTAAGGGACTCATCGATGTTTCTGGTGCACAACTTCATGCAAAACTTGATTGCCTTAAGAAAACTATTGAAGTTGGCTTGGCTTGTACTGCGTATGCAGCTAGCGAACGGATGAAAATGAGAAACGTGCTACGTTTGTTGAAAGAGGCCAAGGGCATGCTTATCAAGGGAAACTAA-3’(SEQ ID NO：1)。
BraEFR基因的基因组DNA全长序列为：
5’-ATGAGCCAAAACCCACAATCACTTTCATTCTCTTGGGACCCAAACACATCTCCTT GTAATTGGACTGGCGTGACTTGCGACACACGTATCAGAAGAGTGACTTCCATCAACCTCTCTGGTCATGGTCTCACCGGTTCCATAAGCCCAAGTATTGGTAATCTCTCTTTTCTTACATCTCTCCAACTTCAAAATAATCAACTTCGAGGTCCAATTCCAAAAGAGATCACAAATCTTTTTCGTCTGAGGGTTCTTAATTTAAGCTCTAACAGCTTAGACGGATCCTTACCTTCCAACCTAAGCAAGTTAATAGAGCTACGAGCCCTTGATCTAACCTCAAATATGATCACGGGTATCGTTCCGAACCAACTTGGTGATCTCAAGAACCTAACTATTTTAAACTTGGGAAAAAATCTGTTGCATGGACCCATTCCACCATCTCTTTCAAACATTTCATCTCTTACCGTTTTAAGCCTAGGCACAAACTCTCTTAGTGGTCCAGTTCCTAACGAGTTGGGTCGTCTTCAAAGGCTTCAAGTCCTTGATCTCACCATAAACAACCTCTCTGGAACAATTCCTCCTTCTATTTACAACATGTCGTCTCTAGAATCATTGGTTATTGCTTCAAACAACTTTTGGGGTCAGTTTCCTAGCAACATAGGATACACACTTCCAAAACTCTTGGTTTTCAACGTGTGTTTCAACAAGTTCAGCGGAGAGATTCCAGCTTCTTTATACAACCTCACAAATATTAAGGTTATCCGCGCTGCACACAACCTCTTAGAAGGGACCATACCATCAGGATTAGGAAATCTCCCATTCCTAGAAATGTACAACATTGGATTCAATAAATTAGTTTGGGGTAGAGATCAAAATCTTGATAGTTTCATCAAGTCTTTCAGTAATAGCTCCAAACTTAACTTCCTTGCATTTGATGGAAACCTCTTGGAGGGTGTTATTCCGGTATCTATTGGCAATCTTCCGAAGAATCTTTCCAAACTTTTCATGGGAGGAAACCGCTTCACGGGTATTATTCCTGAGTCCATCGGCGATCTGACAGGATTGACTCTGTTTAACATCAGTGACAACTCACTCACTGGTGAAATCCCTCAAGATATTGGGAAATTGAAAGGTTTGCAAGTGTTAGAACTTGCGAGAAATCAGCTTACTGGGAGAATACCAGATTCTATTGGTGATCTTGGCGGGTTAAATGAGATCAATCTATCACACAACAAACTACAAGGTAGGATCTCACTATCATTTGAAAACTTCAAGAATATGTTATCCATGGATCTATCCAGCAATATGCTAAATGGAAGCATACCAAATGGAGTTCTTAACCTACCTAGCTTGAGTGCCGTCTTGAATCTATCTAGCAACCTATTTTCTGGTCCAATTCCTCAAGATATAAGTCGCCTCGAAAGCCTTGTGAGTCTAGATCTCTCTAGTAACAACTTCTCAGGCCACATCCCTTCATCTATCAAAGATTGTCAAAGCTTGGAGAAGCTAAACATGGCTGGCAATAATCTAGACGGACCAATCCCGGATGCATTAGCAGAAGTAAAAGGTCTAGAGTTTCTTGATCTCTCTTCAAACCAGCTTTCCGGTGTTATTCCTCCAAGACTTCAAGATTTACAAGCAATGAAGTTCTTGAATATTTCATTCAACAATCTTGAAGGATGGGTCCCAAACCGTGGAGTTTTCAAAGATCATTCTAAAGCTTATATGGAGGGGAATCCAAAGCTCTGCATTCACACTTGTCGAAAAACTCGGACGCACAGAAAACTTTTGAAAGTTAGCGTTATTACATGTGCGGTTGGTCTGATTGCAATATGTGTGATAAGTTTCTTGATTTGGAAAAGAATGGAAAAGAGGTCTACCACATCTACCTCATCATCAAATTCACTCCTCAAGGAACCATTTATGAATGTTTCATATGATGAGCTCAGGAGAGCTACTGAAAATTTCAACCCTAGAAATATTCTTGGTGTTGGAAGCTTTGGATCGGTATTCAAAGGGGTGATACGAGGGGTTGATGTTGCGGTGAAAGTCATTGATCTCAAGGCAAACGGATACTACAAAGGCTTTATAGCAGAGTGTGAGGCTTTAAGGAATGTAAGGCACAGGAATCTTGTGAAACTAGTAACTTCTTGTTCAAGCATAGACTTCAAAAACAATGAGTTTTTGGCTTTGGTGTATGAATTTTTAAGCAATGGAAGCTTAGAAGAATGGATCAAAGGTAAACATAGAAAACCTGATGGGAGTGTTGGGTTAAATCTTGAAGAAAGAGTGAATGTAGCCATTGACATTGCATCTGCATTGGATTATTTGCACAATGATTGTGAAGTTCATGTGGTTCATTGTGATCTAAAACCTAGCAATATTCTTCTAACCGAAGACATGGTTACAAAGGTCGGCGATTTTGGGTTGGCTCGTGTGTTGTTTGACGCATCCGATGATTGTCATCACGCTTCTATAAGCTCCACACATGTCTTGAAAGGCTCCATTGGTTATATTCCTCCAGGTTAGTTGCTCACCTTTAAATCTTATAAACAGTCTACATATGCTAAACTTGTTATTACTATTCAATATAGTTTTTTTACTACCTTGGTAACATATTTTATATTCTCACGCTGATTTGGGATTGTTATATATGCAGAGTATGGGCTTGGAGAAAAGCCATCACAAGCAGGAGATGTATATAGCTTTGGAGTGATGTTGTTAGAGCTCTTGAGTGGAAAAAGCCCAATGGACGAGCGCTTCGAAGGAGATCAAAATCTAATATCATGGATAAGTTATGGGTTTCAAAGTAACGCAATTATGGAAGTGATTGATCCTAAACTTAAGGGACTCATCGATGTTTCTGGTGCACAACTTCATGCAAAACTTGATTGCCTTAAGAAAACTATTGAAGTTGGCTTGGCTTGTACTGCGTATGCAGCTAGCGAACGGATGAAAATGAGAAACGTGCTACGTTTGTTGAAAGAGGCCAAGGGCATGCTTATCAAGGGAAACTAA-3’(SEQ ID NO：2)。
其编码的蛋白的氨基酸序列为：
MSQNPQSLSFSWDPNTSPCNWTGVTCDTRIRRVTSINLSGHGLTGSISPSIGNLSFLTSLQLQNNQLRGPIPKEITNLFRLRVLNLSSNSLDGSLPSNLSKLIELRALDLTSNMITGIVPNQLGDLKNLTILNLGKNLLHGPIPPSLSNISSLTVLSLGTNSLSGPVPNELGRLQRLQVLDLTINNLSGTIPPSIYNMSSLESLVIASNNFWGQFPSNIGYTLPKLLVFNVCFNKFSGEIPASLYNLTNIKVIRAAHNLLEGTIPSGLGNLPFLEMYNIGFNKLVWGRDQNLDSFIKSFSNSSKLNFLAFDGNLLEGVIPVSIGNLPKNLSKLFMGGNRFTGIIPESIGDLTGLTLFNISDNSLTGEIPQDIGKLKGLQVLELARNQLTGRIPDSIGDLGGLNEINLSHNKLQGRISLSFENFKNMLSMDLSSNMLNGSIPNGVLNLPSLSAVLNLSSNLFSGPIPQDISRLESLVSLDLSSNNFSGHIPSSIKDCQSLEKLNMAGNNLDGPIPDALAEVKGLEFLDLSSNQLSGVIPPRLQDLQAMKFLNISFNNLEGWVPNRGVFKDHSKAYMEGNPKLCIHTCRKTRTHRKLLKVSVITCAVGLIAICVISFLIWKRMEKRSTTSTSSSNSLLKEPFMNVSYDELRRATENFNPRNILGVGSFGSVFKGVIRGVDVAVKVIDLKANGYYKGFIAECEALRNVRHRNLVKLVTSCSSIDFKNNEFLALVYEFLSNGSLEEWIKGKHRKPDGSVGLNLEERVNVAIDIASALDYLHNDCEVHVVHCDLKPSNILLTEDMVTKVGDFGLARVLFDASDDCHHASISSTHVLKGSIGYIPPEYGLGEKPSQAGDVYSFGVMLLELLSGKSPMDERFEGDQNLISWISYGFQSNAIMEVIDPKLKGLIDVSGAQLHAKLDCLKKTIEVGLACTAYAASERMKMRNVLRLLKEAKGMLIKGN(SEQ ID NO：3)。
(2)BraEFR基因沉默载体的构建：
采用本领域中基因重组常规方法，将BraEFR基因CDS非保守区段的长度为350bp(CDS101-450bp)的片段及其互补片段构建到pK7GWIW质粒(购自上海远慕生物科技有限公司，pK7GWIW质粒图谱如图3所示)具体插入位置为35S启动子后的attR1和attR2之间(具体为通过基因重组交换替代了pK7GWIW质粒中的ccdB基因)，从而得到了BraEFR基因沉默载体pK7GWIW-BraEFR。
其中，本实施例中所用的BraEFR基因CDS非保守区段的长度为350bp(CDS 101-450bp)的片段具体核苷酸序列为：
5’-CTTCCATCAACCTCTCTGGTCATGGTCTCACCGGTTCCATAAGCCCAAGTATTGGT AATCTCTCTTTTCTTACATCTCTCCAACTTCAAAATAATCAACTTCGAGGTCCAATTCCAAAAGAGATCACAAATCTTTTTCGTCTGAGGGTTCTTAATTTAAGCTCTAACAGCTTAGACGGATCCTTACCTTCCAACCTAAGCAAGTTAATAGAGCTACGAGCCCTTGATCTAACCTCAAATATGATCACGGGTATCGTTCCGAACCAACTTGGTGATCTCAAGAACCTAACTATTTTAAACTTGGGAAAAAATCTGTTGCATGGACCCATTCCACCATCTCTTTCAAACATT-3’(SEQ ID NO：4)。
瞬时过量表达/基因沉默的菜心叶片基因表达模型的构建
具体构建方法为：
(1)取上述实施例中构建得到的BraEFR基因过表达载体pGWB5-BraEFR和BraEFR基因沉默载体pK7GWIW-BraEFR分别转化农杆菌GV3101，然后在对应抗性的培养基上倒置培养48h。
(2)分别挑取阳性单克隆加入到4mL含有相应抗生素及利福平的LB培养基中，28℃，180rpm摇菌24h。
(3)取菌液，按1:100的体积比加入至新鲜的含有相应抗生素及利福平的LB培养基中，28℃，180rpm摇菌到OD600值约为3.0。
(4)3000rpm离心5min，收集菌体，用悬浮液(含有终浓度为10mM MES缓冲液和10mMMgCl2的混合溶液)重悬菌体，并调整OD600值约为0.4。
(5)加入终浓度为200mM的乙酰丁香酮，室温静置3h。
(6)用注射器针头在试验用菜心幼苗的叶片主叶脉两侧各扎一个针孔。用1mL注射器吸取等量静置过后的菌液，对准菜心叶片背面的针孔注射进菌液(菜心叶片正面用手堵住)。
(7)将注射后的菜心幼苗(两叶一心)黑暗培养12h后22℃光照培养3天。
(8)取104个/mL浓度的霜霉菌孢子溶液，点滴接种于菜心幼苗叶片注射针眼处，按照白天12h(20℃)/夜晚12h(14℃)的条件，培养7天。
(9)收集菜心幼苗叶片，进行基因表达水平检测。
设置空白对照(CK)。
具体检测方法为：
a.将叶片组织在液氮中磨碎，每100mg组织加入1mL TRIzol(InvitrogenTRIzol)，用匀浆仪进行匀浆处理。
b.将匀浆样品在室温(15-30℃)放置5分钟，使核酸蛋白复合物完全分离。
c.每使用1mL TRIzol加入0.2ml氯仿，剧烈振荡15秒，室温放置3分钟。2-8℃10000×g离心15分钟。
d.把水相转移到新管中，用异丙醇沉淀水相中的RNA，每使用1mL TRIzol加入0.5ml异丙醇，室温放置10分钟。2-8℃10000×g离心10分钟，保留沉淀，移去上清。
e.用75％乙醇洗涤RNA沉淀，每使用1mL TRIzol加1mL 75％乙醇。2-8℃7000×g离心5分钟，弃上清。
f.室温放置干燥或真空抽干RNA沉淀，大约晾5-10分钟，加入25-200μL无RNase的水，用枪头吸打几次，55-60℃放置10分钟使RNA溶解，-70℃保存。
g.以EF1α基因为内参基因，目的基因表达量与内参基因表达量比值为基因的相对表达量。其中，BraEFR基因表达扩增引物为F:5’-CTTCCATCAACCTCTCTGGTC-3’(SEQ ID NO：5)，R:5’-TGAGGTTAGATCAAGGGCTCG-3’(SEQ ID NO：6)；EF1α基因表达扩增引物为：F:5’-ACTCCTCCCACATTGCTGTTA-3’(SEQ ID NO：7)，R:5’-CCTCAAGAAC TTGGGCTCCTT-3’(SEQ IDNO：8)。
结果如图4和图5所示。
可以发现，BraEFR基因瞬时过量表达的菜心叶片中BraEFR基因的表达量在霜霉病菌接种前后均显著高于未处理的对照(CK)(图4)。而BraEFR基因沉默的菜心叶片中BraEFR基因的表达量在接种霜霉病24h后，显著小于未处理的对照(CK)(图5)。上述结果说明BraEFR基因瞬时过量表达和基因沉默的菜心叶片模型均构建成功。
菜心叶片抗病实验
取上述实施例中构建得到的BraEFR基因瞬时过量表达和基因沉默的菜心叶片模型，按照上述实施例中的方法接种霜霉菌后培养7天，根据培养后的菜心叶片病斑大小来判断目的基因对霜霉病的抗性功能。以未处理的菜心作为空白对照。每个组设置30个样本。
对每个组中的30个叶片病斑面积进行统计，取平均数，并对不同组之间进行显著性分析和比较，检测方法同上述实施例，实验结果如图6和图7所示。
结果发现，在菜心叶片中瞬时过量表达BraEFR基因(通过pGWB5-BraEFR)能够使菜心叶片对霜霉病的抗性较对照(CK)明显增强，而在菜心叶片中瞬时沉默BraEFR基因(pK7GWIW-BraEFR)则使菜心叶片对霜霉病的抗性较对照(CK)显著减弱。上述结果说明通过调控BraEFR基因能够实现对于菜心霜霉病抗性的调控。
上述实施例为本发明较佳的实施方式，但本发明的实施方式并不受上述实施例的限制，其他的任何未背离本发明的精神实质与原理下所作的改变、修饰、替代、组合、简化，均应为等效的置换方式，都包含在本发明的保护范围之内。
SEQUENCE LISTING
<110> 广东省农业科学院设施农业研究所
<120> 菜心LRR受体蛋白激酶基因BraEFR在抗霜霉病中的应用
<130>
<160> 8
<170> PatentIn version 3.5
<210> 1
<211> 2880
<212> DNA
<213> BraEFR
<400> 1
atgagccaaa acccacaatc actttcattc tcttgggacc caaacacatc tccttgtaat 60
tggactggcg tgacttgcga cacacgtatc agaagagtga cttccatcaa cctctctggt 120
catggtctca ccggttccat aagcccaagt attggtaatc tctcttttct tacatctctc 180
caacttcaaa ataatcaact tcgaggtcca attccaaaag agatcacaaa tctttttcgt 240
ctgagggttc ttaatttaag ctctaacagc ttagacggat ccttaccttc caacctaagc 300
aagttaatag agctacgagc ccttgatcta acctcaaata tgatcacggg tatcgttccg 360
aaccaacttg gtgatctcaa gaacctaact attttaaact tgggaaaaaa tctgttgcat 420
ggacccattc caccatctct ttcaaacatt tcatctctta ccgttttaag cctaggcaca 480
aactctctta gtggtccagt tcctaacgag ttgggtcgtc ttcaaaggct tcaagtcctt 540
gatctcacca taaacaacct ctctggaaca attcctcctt ctatttacaa catgtcgtct 600
ctagaatcat tggttattgc ttcaaacaac ttttggggtc agtttcctag caacatagga 660
tacacacttc caaaactctt ggttttcaac gtgtgtttca acaagttcag cggagagatt 720
ccagcttctt tatacaacct cacaaatatt aaggttatcc gcgctgcaca caacctctta 780
gaagggacca taccatcagg attaggaaat ctcccattcc tagaaatgta caacattgga 840
ttcaataaat tagtttgggg tagagatcaa aatcttgata gtttcatcaa gtctttcagt 900
aatagctcca aacttaactt ccttgcattt gatggaaacc tcttggaggg tgttattccg 960
gtatctattg gcaatcttcc gaagaatctt tccaaacttt tcatgggagg aaaccgcttc 1020
acgggtatta ttcctgagtc catcggcgat ctgacaggat tgactctgtt taacatcagt 1080
gacaactcac tcactggtga aatccctcaa gatattggga aattgaaagg tttgcaagtg 1140
ttagaacttg cgagaaatca gcttactggg agaataccag attctattgg tgatcttggc 1200
gggttaaatg agatcaatct atcacacaac aaactacaag gtaggatctc actatcattt 1260
gaaaacttca agaatatgtt atccatggat ctatccagca atatgctaaa tggaagcata 1320
ccaaatggag ttcttaacct acctagcttg agtgccgtct tgaatctatc tagcaaccta 1380
ttttctggtc caattcctca agatataagt cgcctcgaaa gccttgtgag tctagatctc 1440
tctagtaaca acttctcagg ccacatccct tcatctatca aagattgtca aagcttggag 1500
aagctaaaca tggctggcaa taatctagac ggaccaatcc cggatgcatt agcagaagta 1560
aaaggtctag agtttcttga tctctcttca aaccagcttt ccggtgttat tcctccaaga 1620
cttcaagatt tacaagcaat gaagttcttg aatatttcat tcaacaatct tgaaggatgg 1680
gtcccaaacc gtggagtttt caaagatcat tctaaagctt atatggaggg gaatccaaag 1740
ctctgcattc acacttgtcg aaaaactcgg acgcacagaa aacttttgaa agttagcgtt 1800
attacatgtg cggttggtct gattgcaata tgtgtgataa gtttcttgat ttggaaaaga 1860
atggaaaaga ggtctaccac atctacctca tcatcaaatt cactcctcaa ggaaccattt 1920
atgaatgttt catatgatga gctcaggaga gctactgaaa atttcaaccc tagaaatatt 1980
cttggtgttg gaagctttgg atcggtattc aaaggggtga tacgaggggt tgatgttgcg 2040
gtgaaagtca ttgatctcaa ggcaaacgga tactacaaag gctttatagc agagtgtgag 2100
gctttaagga atgtaaggca caggaatctt gtgaaactag taacttcttg ttcaagcata 2160
gacttcaaaa acaatgagtt tttggctttg gtgtatgaat ttttaagcaa tggaagctta 2220
gaagaatgga tcaaaggtaa acatagaaaa cctgatggga gtgttgggtt aaatcttgaa 2280
gaaagagtga atgtagccat tgacattgca tctgcattgg attatttgca caatgattgt 2340
gaagttcatg tggttcattg tgatctaaaa cctagcaata ttcttctaac cgaagacatg 2400
gttacaaagg tcggcgattt tgggttggct cgtgtgttgt ttgacgcatc cgatgattgt 2460
catcacgctt ctataagctc cacacatgtc ttgaaaggct ccattggtta tattcctcca 2520
gagtatgggc ttggagaaaa gccatcacaa gcaggagatg tatatagctt tggagtgatg 2580
ttgttagagc tcttgagtgg aaaaagccca atggacgagc gcttcgaagg agatcaaaat 2640
ctaatatcat ggataagtta tgggtttcaa agtaacgcaa ttatggaagt gattgatcct 2700
aaacttaagg gactcatcga tgtttctggt gcacaacttc atgcaaaact tgattgcctt 2760
aagaaaacta ttgaagttgg cttggcttgt actgcgtatg cagctagcga acggatgaaa 2820
atgagaaacg tgctacgttt gttgaaagag gccaagggca tgcttatcaa gggaaactaa 2880
<210> 2
<211> 3015
<212> DNA
<213> BraEFR
<400> 2
atgagccaaa acccacaatc actttcattc tcttgggacc caaacacatc tccttgtaat 60
tggactggcg tgacttgcga cacacgtatc agaagagtga cttccatcaa cctctctggt 120
catggtctca ccggttccat aagcccaagt attggtaatc tctcttttct tacatctctc 180
caacttcaaa ataatcaact tcgaggtcca attccaaaag agatcacaaa tctttttcgt 240
ctgagggttc ttaatttaag ctctaacagc ttagacggat ccttaccttc caacctaagc 300
aagttaatag agctacgagc ccttgatcta acctcaaata tgatcacggg tatcgttccg 360
aaccaacttg gtgatctcaa gaacctaact attttaaact tgggaaaaaa tctgttgcat 420
ggacccattc caccatctct ttcaaacatt tcatctctta ccgttttaag cctaggcaca 480
aactctctta gtggtccagt tcctaacgag ttgggtcgtc ttcaaaggct tcaagtcctt 540
gatctcacca taaacaacct ctctggaaca attcctcctt ctatttacaa catgtcgtct 600
ctagaatcat tggttattgc ttcaaacaac ttttggggtc agtttcctag caacatagga 660
tacacacttc caaaactctt ggttttcaac gtgtgtttca acaagttcag cggagagatt 720
ccagcttctt tatacaacct cacaaatatt aaggttatcc gcgctgcaca caacctctta 780
gaagggacca taccatcagg attaggaaat ctcccattcc tagaaatgta caacattgga 840
ttcaataaat tagtttgggg tagagatcaa aatcttgata gtttcatcaa gtctttcagt 900
aatagctcca aacttaactt ccttgcattt gatggaaacc tcttggaggg tgttattccg 960
gtatctattg gcaatcttcc gaagaatctt tccaaacttt tcatgggagg aaaccgcttc 1020
acgggtatta ttcctgagtc catcggcgat ctgacaggat tgactctgtt taacatcagt 1080
gacaactcac tcactggtga aatccctcaa gatattggga aattgaaagg tttgcaagtg 1140
ttagaacttg cgagaaatca gcttactggg agaataccag attctattgg tgatcttggc 1200
gggttaaatg agatcaatct atcacacaac aaactacaag gtaggatctc actatcattt 1260
gaaaacttca agaatatgtt atccatggat ctatccagca atatgctaaa tggaagcata 1320
ccaaatggag ttcttaacct acctagcttg agtgccgtct tgaatctatc tagcaaccta 1380
ttttctggtc caattcctca agatataagt cgcctcgaaa gccttgtgag tctagatctc 1440
tctagtaaca acttctcagg ccacatccct tcatctatca aagattgtca aagcttggag 1500
aagctaaaca tggctggcaa taatctagac ggaccaatcc cggatgcatt agcagaagta 1560
aaaggtctag agtttcttga tctctcttca aaccagcttt ccggtgttat tcctccaaga 1620
cttcaagatt tacaagcaat gaagttcttg aatatttcat tcaacaatct tgaaggatgg 1680
gtcccaaacc gtggagtttt caaagatcat tctaaagctt atatggaggg gaatccaaag 1740
ctctgcattc acacttgtcg aaaaactcgg acgcacagaa aacttttgaa agttagcgtt 1800
attacatgtg cggttggtct gattgcaata tgtgtgataa gtttcttgat ttggaaaaga 1860
atggaaaaga ggtctaccac atctacctca tcatcaaatt cactcctcaa ggaaccattt 1920
atgaatgttt catatgatga gctcaggaga gctactgaaa atttcaaccc tagaaatatt 1980
cttggtgttg gaagctttgg atcggtattc aaaggggtga tacgaggggt tgatgttgcg 2040
gtgaaagtca ttgatctcaa ggcaaacgga tactacaaag gctttatagc agagtgtgag 2100
gctttaagga atgtaaggca caggaatctt gtgaaactag taacttcttg ttcaagcata 2160
gacttcaaaa acaatgagtt tttggctttg gtgtatgaat ttttaagcaa tggaagctta 2220
gaagaatgga tcaaaggtaa acatagaaaa cctgatggga gtgttgggtt aaatcttgaa 2280
gaaagagtga atgtagccat tgacattgca tctgcattgg attatttgca caatgattgt 2340
gaagttcatg tggttcattg tgatctaaaa cctagcaata ttcttctaac cgaagacatg 2400
gttacaaagg tcggcgattt tgggttggct cgtgtgttgt ttgacgcatc cgatgattgt 2460
catcacgctt ctataagctc cacacatgtc ttgaaaggct ccattggtta tattcctcca 2520
ggttagttgc tcacctttaa atcttataaa cagtctacat atgctaaact tgttattact 2580
attcaatata gtttttttac taccttggta acatatttta tattctcacg ctgatttggg 2640
attgttatat atgcagagta tgggcttgga gaaaagccat cacaagcagg agatgtatat 2700
agctttggag tgatgttgtt agagctcttg agtggaaaaa gcccaatgga cgagcgcttc 2760
gaaggagatc aaaatctaat atcatggata agttatgggt ttcaaagtaa cgcaattatg 2820
gaagtgattg atcctaaact taagggactc atcgatgttt ctggtgcaca acttcatgca 2880
aaacttgatt gccttaagaa aactattgaa gttggcttgg cttgtactgc gtatgcagct 2940
agcgaacgga tgaaaatgag aaacgtgcta cgtttgttga aagaggccaa gggcatgctt 3000
atcaagggaa actaa 3015
<210> 3
<211> 959
<212> PRT
<213> BraEFR
<400> 3
Met Ser Gln Asn Pro Gln Ser Leu Ser Phe Ser Trp Asp Pro Asn Thr
1 5 10 15
Ser Pro Cys Asn Trp Thr Gly Val Thr Cys Asp Thr Arg Ile Arg Arg
20 25 30
Val Thr Ser Ile Asn Leu Ser Gly His Gly Leu Thr Gly Ser Ile Ser
35 40 45
Pro Ser Ile Gly Asn Leu Ser Phe Leu Thr Ser Leu Gln Leu Gln Asn
50 55 60
Asn Gln Leu Arg Gly Pro Ile Pro Lys Glu Ile Thr Asn Leu Phe Arg
65 70 75 80
Leu Arg Val Leu Asn Leu Ser Ser Asn Ser Leu Asp Gly Ser Leu Pro
85 90 95
Ser Asn Leu Ser Lys Leu Ile Glu Leu Arg Ala Leu Asp Leu Thr Ser
100 105 110
Asn Met Ile Thr Gly Ile Val Pro Asn Gln Leu Gly Asp Leu Lys Asn
115 120 125
Leu Thr Ile Leu Asn Leu Gly Lys Asn Leu Leu His Gly Pro Ile Pro
130 135 140
Pro Ser Leu Ser Asn Ile Ser Ser Leu Thr Val Leu Ser Leu Gly Thr
145 150 155 160
Asn Ser Leu Ser Gly Pro Val Pro Asn Glu Leu Gly Arg Leu Gln Arg
165 170 175
Leu Gln Val Leu Asp Leu Thr Ile Asn Asn Leu Ser Gly Thr Ile Pro
180 185 190
Pro Ser Ile Tyr Asn Met Ser Ser Leu Glu Ser Leu Val Ile Ala Ser
195 200 205
Asn Asn Phe Trp Gly Gln Phe Pro Ser Asn Ile Gly Tyr Thr Leu Pro
210 215 220
Lys Leu Leu Val Phe Asn Val Cys Phe Asn Lys Phe Ser Gly Glu Ile
225 230 235 240
Pro Ala Ser Leu Tyr Asn Leu Thr Asn Ile Lys Val Ile Arg Ala Ala
245 250 255
His Asn Leu Leu Glu Gly Thr Ile Pro Ser Gly Leu Gly Asn Leu Pro
260 265 270
Phe Leu Glu Met Tyr Asn Ile Gly Phe Asn Lys Leu Val Trp Gly Arg
275 280 285
Asp Gln Asn Leu Asp Ser Phe Ile Lys Ser Phe Ser Asn Ser Ser Lys
290 295 300
Leu Asn Phe Leu Ala Phe Asp Gly Asn Leu Leu Glu Gly Val Ile Pro
305 310 315 320
Val Ser Ile Gly Asn Leu Pro Lys Asn Leu Ser Lys Leu Phe Met Gly
325 330 335
Gly Asn Arg Phe Thr Gly Ile Ile Pro Glu Ser Ile Gly Asp Leu Thr
340 345 350
Gly Leu Thr Leu Phe Asn Ile Ser Asp Asn Ser Leu Thr Gly Glu Ile
355 360 365
Pro Gln Asp Ile Gly Lys Leu Lys Gly Leu Gln Val Leu Glu Leu Ala
370 375 380
Arg Asn Gln Leu Thr Gly Arg Ile Pro Asp Ser Ile Gly Asp Leu Gly
385 390 395 400
Gly Leu Asn Glu Ile Asn Leu Ser His Asn Lys Leu Gln Gly Arg Ile
405 410 415
Ser Leu Ser Phe Glu Asn Phe Lys Asn Met Leu Ser Met Asp Leu Ser
420 425 430
Ser Asn Met Leu Asn Gly Ser Ile Pro Asn Gly Val Leu Asn Leu Pro
435 440 445
Ser Leu Ser Ala Val Leu Asn Leu Ser Ser Asn Leu Phe Ser Gly Pro
450 455 460
Ile Pro Gln Asp Ile Ser Arg Leu Glu Ser Leu Val Ser Leu Asp Leu
465 470 475 480
Ser Ser Asn Asn Phe Ser Gly His Ile Pro Ser Ser Ile Lys Asp Cys
485 490 495
Gln Ser Leu Glu Lys Leu Asn Met Ala Gly Asn Asn Leu Asp Gly Pro
500 505 510
Ile Pro Asp Ala Leu Ala Glu Val Lys Gly Leu Glu Phe Leu Asp Leu
515 520 525
Ser Ser Asn Gln Leu Ser Gly Val Ile Pro Pro Arg Leu Gln Asp Leu
530 535 540
Gln Ala Met Lys Phe Leu Asn Ile Ser Phe Asn Asn Leu Glu Gly Trp
545 550 555 560
Val Pro Asn Arg Gly Val Phe Lys Asp His Ser Lys Ala Tyr Met Glu
565 570 575
Gly Asn Pro Lys Leu Cys Ile His Thr Cys Arg Lys Thr Arg Thr His
580 585 590
Arg Lys Leu Leu Lys Val Ser Val Ile Thr Cys Ala Val Gly Leu Ile
595 600 605
Ala Ile Cys Val Ile Ser Phe Leu Ile Trp Lys Arg Met Glu Lys Arg
610 615 620
Ser Thr Thr Ser Thr Ser Ser Ser Asn Ser Leu Leu Lys Glu Pro Phe
625 630 635 640
Met Asn Val Ser Tyr Asp Glu Leu Arg Arg Ala Thr Glu Asn Phe Asn
645 650 655
Pro Arg Asn Ile Leu Gly Val Gly Ser Phe Gly Ser Val Phe Lys Gly
660 665 670
Val Ile Arg Gly Val Asp Val Ala Val Lys Val Ile Asp Leu Lys Ala
675 680 685
Asn Gly Tyr Tyr Lys Gly Phe Ile Ala Glu Cys Glu Ala Leu Arg Asn
690 695 700
Val Arg His Arg Asn Leu Val Lys Leu Val Thr Ser Cys Ser Ser Ile
705 710 715 720
Asp Phe Lys Asn Asn Glu Phe Leu Ala Leu Val Tyr Glu Phe Leu Ser
725 730 735
Asn Gly Ser Leu Glu Glu Trp Ile Lys Gly Lys His Arg Lys Pro Asp
740 745 750
Gly Ser Val Gly Leu Asn Leu Glu Glu Arg Val Asn Val Ala Ile Asp
755 760 765
Ile Ala Ser Ala Leu Asp Tyr Leu His Asn Asp Cys Glu Val His Val
770 775 780
Val His Cys Asp Leu Lys Pro Ser Asn Ile Leu Leu Thr Glu Asp Met
785 790 795 800
Val Thr Lys Val Gly Asp Phe Gly Leu Ala Arg Val Leu Phe Asp Ala
805 810 815
Ser Asp Asp Cys His His Ala Ser Ile Ser Ser Thr His Val Leu Lys
820 825 830
Gly Ser Ile Gly Tyr Ile Pro Pro Glu Tyr Gly Leu Gly Glu Lys Pro
835 840 845
Ser Gln Ala Gly Asp Val Tyr Ser Phe Gly Val Met Leu Leu Glu Leu
850 855 860
Leu Ser Gly Lys Ser Pro Met Asp Glu Arg Phe Glu Gly Asp Gln Asn
865 870 875 880
Leu Ile Ser Trp Ile Ser Tyr Gly Phe Gln Ser Asn Ala Ile Met Glu
885 890 895
Val Ile Asp Pro Lys Leu Lys Gly Leu Ile Asp Val Ser Gly Ala Gln
900 905 910
Leu His Ala Lys Leu Asp Cys Leu Lys Lys Thr Ile Glu Val Gly Leu
915 920 925
Ala Cys Thr Ala Tyr Ala Ala Ser Glu Arg Met Lys Met Arg Asn Val
930 935 940
Leu Arg Leu Leu Lys Glu Ala Lys Gly Met Leu Ile Lys Gly Asn
945 950 955
<210> 4
<211> 350
<212> DNA
<213> BraEFR
<400> 4
cttccatcaa cctctctggt catggtctca ccggttccat aagcccaagt attggtaatc 60
tctcttttct tacatctctc caacttcaaa ataatcaact tcgaggtcca attccaaaag 120
agatcacaaa tctttttcgt ctgagggttc ttaatttaag ctctaacagc ttagacggat 180
ccttaccttc caacctaagc aagttaatag agctacgagc ccttgatcta acctcaaata 240
tgatcacggg tatcgttccg aaccaacttg gtgatctcaa gaacctaact attttaaact 300
tgggaaaaaa tctgttgcat ggacccattc caccatctct ttcaaacatt 350
<210> 5
<211> 21
<212> DNA
<213> 人工序列
<400> 5
cttccatcaa cctctctggt c 21
<210> 6
<211> 21
<212> DNA
<213> 人工序列
<400> 6
tgaggttaga tcaagggctc g 21
<210> 7
<211> 21
<212> DNA
<213> 人工序列
<400> 7
actcctccca cattgctgtt a 21
<210> 8
<211> 21
<212> DNA
<213> 人工序列
<400> 8
cctcaagaac ttgggctcct t 21
Claims (3)
1.过量表达菜心BraEFR基因的载体在调控菜心抗菜心霜霉病中的应用，所述菜心BraEFR基因的核苷酸序列如SEQ ID NO：2或SEQ ID NO：1所示。
2.含有过量表达菜心BraEFR基因的载体的微生物或细胞在调控菜心抗菜心霜霉病中的应用，所述菜心BraEFR基因的核苷酸序列如SEQ ID NO：2或SEQ ID NO：1所示。
3.一种培育抗菜心霜霉病作物的方法，包括如下步骤：通过使用过量表达菜心BraEFR基因的载体正调菜心抗菜心霜霉病；所述BraEFR基因的核苷酸序列如SEQ ID NO：2或SEQID NO：1所示。
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