CN114381463A - 来源欧洲芜菁ECD04的BraA08g039212E基因在根肿菌抗性改良中的应用 - Google Patents
来源欧洲芜菁ECD04的BraA08g039212E基因在根肿菌抗性改良中的应用 Download PDFInfo
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Abstract
本发明公开了一种来源欧洲芜菁ECD04的BraA08g039212E基因在根肿菌抗性改良中的应用，本发明公开了来源于欧洲芜菁ECD04的BraA08g039212E基因对中国根肿菌4号生理小种的抗性作用，该基因核苷酸序列如SEQ ID No.1或SEQ ID No.2所示，氨基酸序列如SEQ ID No.3所示，包括利用图位克隆法精细定位该基因，明确其在A08染色体上的物理位置，然后利用该基因的gDNA或cDNA进行克隆并遗传转化不抗根肿病油菜，从而验证该基因对根肿菌的抗根肿病功能。本发明可以为通过基因工程手段或分子标记辅助选择技术选育芸薹属抗根肿病植物新品种提供科学和技术依据。
Description
技术领域
本发明涉及基因克隆技术领域，特别是涉及一种来源欧洲芜菁ECD04的BraA08g039212E基因在根肿菌抗性改良中的应用。
背景技术
根肿病是由芸苔根肿菌(Plasmodiophora brassicae Woronin)引起的一种土传病害，严重危害十字花科作物的生长发育。根肿菌在土壤中以休眠孢子的形式存在，萌发后在寄主体内以活体营养的方式进行繁殖，使寄主植物生长素和细胞分裂素代谢紊乱，导致植物的根部肿大。严重受到根肿菌侵染的植物，其主根系可能被完全破坏，从而影响地上部分的生长发育，造成植物的绝收。油菜是我国重要的油料作物，其产量深受根肿菌的影响。根肿菌在国内主要分布于四川、云南、安徽、湖北等省份。随着机械化程度不断提高，根肿菌扩散速度也不断加快，已经严重影响到我国油菜产业的安全。因此挖掘抗病基因，利用抗病位点去选育抗根肿病的优良油菜品种并解析根肿病的致病机理，对于病情的管控、稳定油菜产量、保证油菜产业安全具有重大意义。
目前，欧洲芜菁是抗根肿病QTL或者主效基因的主要来源，绝大多数抗病位点都分布在A基因组染色体上，分别为A01染色体上的Crr2和PbBa1.1、A02染色体上的CRc、A03染色体上的CRa和Crr3、A06染色体上的Crr4。在白菜品种“Debra”的A02和A03染色体上分别定位到了CRc和CRk。2013年，Chen等利用欧洲芜菁ECD04作为供体亲本，感病大白菜自交系C59-1作为受体亲本，用4种不同的根肿菌生理小种(Pb2、Pb4、Pb7、Pb10)对BC1F1世代部分材料进行接菌鉴定，同时结合已发表根肿病抗病基因相关特异性标记和SSR(Simple sequencerepeat)，利用QTL构建连锁遗传图谱，分别在A01、A03和A08染色体上一共定位出5个抗病位点：PbBa1.1、PbBa3.1、PbBa3.2、PbBa3.3和PbBa8.1(Chen J,Jing J,Zhan Z,Zhang T,Zhang C,Piao Z.Identification of novel QTLs for isolate-specific partialresistance to Plasmodiophora brassicae in Brassica rapa.PLoS One,2013,8:e85307.)。
测序技术的发展对根肿菌抗病位点研究起到了一定的推进作用。2019年，Huang等利用转录组测序结合的混合分组分析技术(BSR-Seq)在抗根肿病芜菁品种“Purple TopWhite Globe”的A03染色体上定位到Rcr5抗根肿病位点，并且将Rcr5的候选基因范围缩小在8个基因内(Huang Z,Peng G,Gossen BD,Yu F.Fine mapping of a clubrootresistance gene from turnip using SNP markers identified from bulkedsegregant RNA-Seq.Mol Breeding,2019,39:131)。
目前为止，CRa和Crr1是被克隆出来并且进行功能验证的两个根肿菌抗病基因。抗病基因CRa来自于ECD02中分离出一个抗病位点(Matsumoto E,Ueno H,Aruga D,SakamotoK,Hayashida N.Accumulation of three clubroot resistance genes through marker-assisted selection in Chinese cabbage(Brassica rapa ssp.pekinensis).J Jpn SocHortic Sci,2012,81:184–190)。Hatakeyama等人从“Siloga”中克隆出Crr1，而该位点存在两个抗病基因Crr1a和Crr1b。Crr1a为主效基因，单独存在时只能对生理小种起到微效抗性作用；与Crr2基因互作时，可以对根肿菌4号生理小种起到较强的抗性(Suwabe K,Tsukazaki H,Iketani H,Hatakeyama K,Fujimura M,Nunome T,Fukuoka H,Matsumoto S,Hirai M.Identification of two loci for resistance to clubroot(Plasmodiophorabrassicae Woronin)in Brassica rapa L.Theor Appl Genet,2003,107:997–1002)。
植物具备先天免疫系统，分别为病原相关分子模式激发的免疫反应(PTI)和效应蛋白激发的免疫反应(ETI)。抗病基因中最大的基因家族为NBS-LRR(Nucleotide-BlindingSite and Leucine Rich Repeat)基因家族。CRa和Crr1均具备典型的TIR-NBS-LRR结构域，属于NBS-LRR抗病基因家族。这一类抗病基因家族的功能通常具有专一性，针对某一或者一类生理小种具备强大的抗性。NBS-LRR基因家族发挥其功能往往需要LRR结构域识别病原效应因子从而激活植物的免疫反应(Porter B W，Paidi M，Ming R，et al.Genome-wideanalysis of Carica papaya reveals a small NBS resistance gene family[J].Molecular Genetics and Genomics，2009，281(6)：609-626)，使得N端的TIR/CC结构域去激活下游的免疫反应(Meyers B C，Kozik A，Griego A，et al.Genome-wide analysis ofNBS-LRR-encoding genes in Arabidopsis[J].The Plant Cell，2003，15(4)：809.)。
发明内容
本发明的目的是提供一种来源欧洲芜菁ECD04的BraA08g039212E基因在根肿菌抗性改良中的应用，以解决上述现有技术存在的问题，本发明以甘蓝型油菜J9707为转化受体，克隆并验证了一个对中国根肿菌4号优势生理小种具备有效抗性的基因BraA08g039212E，提供了一个依托该抗性基因BraA08g039212E，为通过分子标记辅助选择和基因工程方法改良油菜、白菜和甘蓝等芸薹属植物的根肿病抗性的育种方法。
为实现上述目的，本发明提供了如下方案：
本发明提供一种BraA08g039212E基因在赋予根肿菌抗性中的应用，所述BraA08g039212E基因的核苷酸序列如SEQ ID No.1或SEQ ID No.2所示，其编码蛋白的氨基酸序列如SEQ ID No.3所示。其中，SEQ ID No.1所示核苷酸序列为gDNA序列，包括启动子序列、基因全长序列、终止子序列，SEQ ID No.2所示核苷酸序列为cDNA序列。
进一步地，所述BraA08g039212E基因位于欧洲芜菁ECD04的A08染色体上，其物理位置为14863686-14873201。
本发明还提供一种BraA08g039212E基因在根肿病抗病育种中的应用，所述BraA08g039212E基因通过转基因或转育途径用于芸薹属植物的根肿病抗病育种中的应用，所述芸薹属植物包括油菜、白菜、甘蓝或萝卜，所述BraA08g039212E基因的核苷酸序列如SEQ ID No.1或SEQ ID No.2所示。
本发明还提供一种BraA08g039212E基因及根据其所开发的标记在芸薹属植物分子标记辅助选择中的应用，所述芸薹属植物包括油菜、白菜、甘蓝或萝卜，所述BraA08g039212E基因的核苷酸序列如SEQ ID No.1或SEQ ID No.2所示。
本发明还提供一种BraA08g039212E基因的启动子的用途，用于调控BraA08g039212E基因的表达，其序列如SEQ ID No.4所示。
本发明还提供一种用于扩增BraA08g039212E基因全长的引物，包括如SEQ IDNo.5所示的正向引物和SEQ ID No.6所示的反向引物。
本发明还提供一种用于构建BraA08g039212E基因遗传转化载体的引物，包括如SEQ ID No.7所示的正向引物和SEQ ID No.8所示的反向引物。
本发明还提供一种用于BraA08g039212E基因阳性检测的引物，包括如SEQ IDNo.9所示的正向引物和SEQ ID No.10所示的反向引物。
本发明还提供一种用于BraA08g039212E基因RT-qPCR的引物，包括如SEQ IDNo.11所示的正向引物和SEQ ID No.12所示的反向引物。
本发明公开了以下技术效果：
本发明公开了来源于欧洲芜菁ECD04的BraA08g039212E基因对中国根肿菌4号生理小种具有抗性功能，包括该BraA08g039212E基因抗病区间的精细定位和其在欧洲芜菁ECD04的A08染色体上的物理位置，并以甘蓝型油菜J9707为背景对其基因组序列进行克隆、遗传转化，从而验证BraA08g039212E基因对根肿菌的抗病功能。本发明可以提供一种BraA08g039212E基因对抗根肿菌病基础研究、抗病育种和分子标记辅助选择中的应用，通过该基因所在的定位区间去选育抗根肿菌的品种，从而缩短育种进程。
附图说明
为了更清楚地说明本发明实施例或现有技术中的技术方案，下面将对实施例中所需要使用的附图作简单地介绍，显而易见地，下面描述中的附图仅仅是本发明的一些实施例，对于本领域普通技术人员来讲，在不付出创造性劳动的前提下，还可以根据这些附图获得其他的附图。
图1：本发明实施例中BraA08g039212E基因在欧洲芜菁ECD04中的定位区间及其相应的物理位置。
图2：本发明实施例中用于定位左区间5778标记的20S029重组单株接中国根肿菌4号生理小种表型图，H5R为阳性对照，H5S为阴性对照，R为抗病材料，S为感病材料。
图3：本发明实施例中用于定位右区间JF07标记的18CZ3886重组单株接中国根肿菌4号生理小种表型图，H5R为阳性对照，H5S为阴性对照，R为抗病材料，S为感病材料。
图4：本发明实施例中BraA08g039212E基因结构域标注。
图5：ECD04和其他7个感病材料在BraA08g039212E位点附近100Kb内的LTR插入示意图。
图6-12：ECD04和其他7个感病材料的BraA08g039212E位点上游3000bp序列比对图。
图13：ECD04和其他7个感病材料的BraA08g039212E位点上游3000bp序列比对差异热图，a图为8个材料相互之间序列比对的覆盖程度；b图为8个材料相互之间在序列比对在a图覆盖程度上的差异性。
图14：本发明实施例中BraA08g039212E基因在改造过的pCAMBIA3301(将Bastar抗性改为Hygr抗性)植物表达载体上克隆模式图。
图15：本发明实施例中BraA08g039212E基因从启动子到终止子PCR扩增后的琼脂糖凝胶图，M为1Kb DNA Ladder，1-3为扩增目的条带，条带大小为8713bp。
图16：BraA08g039212E阳性克隆检测琼脂糖凝胶图，M为DL2000 DNA Marker，1-6为菌落PCR阳性检测条带，条带大小为711bp。
图17：本发明实施例中BraA08g039212E基因在甘蓝型油菜受体材料J9707中表达后接中国根肿菌4号优势生理小种表型图，Control以J9707受体材料作为感病对照，Grade0、Grade 1、Grade 2分别为株系接中国根肿菌4号优势生理小种发病的三个等级，Grade 0：主根与侧根完全不发病；Grade 1：侧根发生根肿现象；Grade 2：主根发生明显根肿现象。
图18：本发明实施例中BraA08g039212E转基因株系的两个株系检测目的基因琼脂糖凝胶图。a图为BraA08g039212E-L1株系基因型；b为BraA08g039212E-L2株系基因型。a，b图中，M为250bp DNA Ladder，N为阴性对照，P为阳性对照。
图19：本发明实施例中BraA08g039212E转基因株系的两个株系目的基因表达量RT-qPCR检测图。
具体实施方式
现详细说明本发明的多种示例性实施方式，该详细说明不应认为是对本发明的限制，而应理解为是对本发明的某些方面、特性和实施方案的更详细的描述。
应理解本发明中所述的术语仅仅是为描述特别的实施方式，并非用于限制本发明。另外，对于本发明中的数值范围，应理解为还具体公开了该范围的上限和下限之间的每个中间值。在任何陈述值或陈述范围内的中间值以及任何其他陈述值或在所述范围内的中间值之间的每个较小的范围也包括在本发明内。这些较小范围的上限和下限可独立地包括或排除在范围内。
除非另有说明，否则本文使用的所有技术和科学术语具有本发明所述领域的常规技术人员通常理解的相同含义。虽然本发明仅描述了优选的方法和材料，但是在本发明的实施或测试中也可以使用与本文所述相似或等同的任何方法和材料。本说明书中提到的所有文献通过引用并入，用以公开和描述与所述文献相关的方法和/或材料。在与任何并入的文献冲突时，以本说明书的内容为准。
在不背离本发明的范围或精神的情况下，可对本发明说明书的具体实施方式做多种改进和变化，这对本领域技术人员而言是显而易见的。由本发明的说明书得到的其他实施方式对技术人员而言是显而易见的。本发明说明书和实施例仅是示例性的。
关于本文中所使用的“包含”、“包括”、“具有”、“含有”等等，均为开放性的用语，即意指包含但不限于。
实施例1
1、精细定位群体构建
利用我国甘蓝型油菜常规品种华双5号(Brassica napus，AACC，2n＝38)作为受体亲本(又称H5S)，欧洲芜菁ECD04(Brassica rapa，AA，2n＝20)作为供体亲本，经过一次杂交和多次回交，并结合分子标记辅助筛选，将ECD04中抗根肿病位点PbBa8.1导入到华双5号中，获得了BC1、BC2和BC3材料。后续经过4次自交，并结合分子标记筛选和田间抗病鉴定，成功培育出中国根肿菌4号生理小种抗病品种华双5R(又称H5R)，并得到了BC3F4群体。该群体在病区田间接菌呈现出抗病表型分离，故选用作精细定位群体。
目前定位的区间位于欧洲芜菁ECD04的A08染色体的5778和JF07标记之间，物理位置为14765289和14965054，区间大小为199.77Kb。抗根肿病基因BraA08g039212E位于该区间中，在A08染色体上物理位置为14863686-14873201(图1)。
2、甘蓝型油菜株系接种体系
室内接种使用的根肿菌，是中国根肿菌4号生理小种(季海雯,任莉,陈坤荣,徐理,刘凡,孙超超,李俊,刘胜毅,方小平.油菜根肿病病原主要生理小种和品种抗病性鉴定[J].中国油料作物学报,2013,35(03):301-306.)。根肿病抗病性室内接菌鉴定采用的温室菌土法接种(战宗祥，江莹芬，朱紫媛等.与位点PbBa8.1紧密连锁分子标记的开发及甘蓝型油菜根肿病抗性育种[J].中国油料作物学报,2015,37(6):766-771.)。首先将冻存于-20℃的菌根置于室温下解冻后，用匀浆机将其磨碎，将菌根和风干的草炭土按1:20的比例混匀，混匀后的菌土放置于25℃黑暗环境下封存48h。播种时将种子播在菌土上并覆土，最后播种好的穴盘放在光照强度是200mmol·m-2·s-1，光照/黑暗时间16/8h，温度为25/20℃(昼夜)的条件下进行培养。培养6周后从土中拔出植株，用水清洗根部后调查抗病表型。
3、重组单株株系后代接菌表型验证
重组单株接种表型鉴定过程中采取了“自交系接菌鉴定表型原则”，即在非病区种植实验对象群体，通过分子标记鉴定筛选出目标单株，再对其进行套袋自交，收获自交系种后，分系进行室内接菌。20S029和18CZ3886是用于精细定位在5778标记到JF07标记区间的两个重组单株繁育的株系，将其后代接中国根肿菌4号生理小种后统计抗病表型，20S029后代144株全抗病，而18CZ3886后代抗、感表型出现分离，分离比为19：30(图2，图3)。
4、重组单株株系后代接菌基因型鉴定
对20S029后代接菌后的全部抗病病材料中进行随机取72个单株，对18CZ3886后代接菌从各表型中随机选取12棵单株，针对这些单株进行分子标记鉴定。
用CTAB法对待测样品进行基因组DNA提取。
利用聚丙烯酰胺凝胶电泳(PAGE)鉴定定位区间标记基因型，纯合抗病记为B，杂合基因型记为H，纯合感病记为A。
结果表明：20S029后代基因型在5778标记及其之前发生分离，而在5778标记之后为纯合抗病基因型(表1)，20S029后代接菌表型为144株全抗病，二者结合来看，该抗病位点定位区间在5778标记之后。18CZ3886后代基因型在JF07标记及其之前是分离的状况，而在JF07标记之后为纯合感病基因型(表2)，18CZ3886后代接菌表型分离比为19：30，二者相结合说明定位区间定在JF07标记之前。因此，抗病区间位于5778标记与JF07标记之间。
表1 20S029自交后代接中国根肿菌4号生理小种基因型鉴定结果
注：纯合抗病记为B，杂合基因型记为H，纯合感病记为A，R表示抗病单株，S表示感病单株。
表2 18CZ3886自交后代接中国根肿菌4号生理小种基因型鉴定结果
注：纯合抗病记为B，杂合基因型记为H，纯合感病记为A，R表示抗病单株，S表示感病单株。
5、通过欧洲芜菁ECD04供体材料全基因组注释，查询到5778标记和JF07标记在A08染色体上的物理位置分别为14765289和14965054，区间大小为199.77Kb，而抗根肿菌基因BraA08g039212E位于5778标记到JF07标记区间之间，其在A08染色体上的物理位置为14866737。BraA08g039212E基因的核苷酸序列如SEQ ID No.1和SEQ ID No.2所示，其编码蛋白氨基酸序列如SEQ ID No.3所示，具有典型的NBS-LRR结构域(图4)，因此该基因可能具有抗根肿菌的功能。
6、ECD04与其他7个感病材料的BraA08g039212E位点启动子分析比较。
ZS11、Quinta、Tapidor、Sheng Li、GanGan、Zhe You7和No2127为7个根肿病感病材料，通过生物信息学手段对BraA08g039212E所在ECD04的A08染色体100Kb区间进行分析，发现该区间中，ECD04与其他7个感病材料相比，BraA08g039212E上游17.59Kb在进化过程中有两个长末端重复序列(LTR)插入，物理位置分别为14856731和14857466，LTR片段大小分别为183bp和137bp。然而，感病材料在BraA08g039212E上游并不存在这两个LTR位点，因此可能会造成BraA08g039212E的启动子功能发生改变并影响基因表达水平(图5)。
将ECD04与其他7个感病材料的BraA08g039212E位点上游3000bp的DNA序列进行比较(图6-12)，结果显示ECD04与感病材料之间的BraA08g039212E位点上游3000bp分为了四类，第一类是ZS11、Tapidor、Quinta和Sheng Li；第二类是GanGan和Zhe You7；第三类是No2127；第四类是ECD04(图13)。从热图结果来看，ECD04的BraA08g039212E位点上游3000bp与其他7个感病材料差异显著，推测ECD04的BraA08g039212E启动子对BraA08g039212E的抗病功能具有一定的作用。由此将BraA08g039212E进行遗传转化时，使用的是其上游3000bp作为其启动子驱动该基因表达，其序列如SEQ ID No.4所示。
7、克隆BraA08g039212E基因，采用同源重组的方法，将目的基因PCR扩增出来，用于扩增BraA08g039212E基因组引物序列为：
正向引物(SEQ ID No.5)：5’-atggatccaatagagacagtgg-3’
反向引物(SEQ ID No.6)：5’-ttatggaaggctctcaagatttg-3’
然后连接至改造过的pCAMBIA3301(将Basta抗性改为Hyg抗性)植物表达载体为骨架，构建由自身启动子驱动的表达载体(图14)，最后转入GV3101农杆菌感受态细胞，详细说明如下：
以供体材料欧洲芜菁ECD04基因组为模板对BraA08g039212E基因进行扩增。
选取起始密码子ATG前3051bp序列作为启动子片段，终止密码TAA后924bp作为自身终止子元件，针对包含编码区，启动子和终止子全长为8713bp的gDNA区段设计引物进行扩增。PCR反应扩增引物由pCAMBIA3301载体同源臂、酶切位点、目的基因同源序列三部分组成，便于后续基因克隆操作。
正向引物序列(SEQ ID No.7)为5’-gctcggtacccggggatccgagggtactgcggagggta-3’
反向引物序列(SEQ ID No.8)为5’-gacggccagtgccaagcttttcctagtgactggcccatc-3’
同样采取CTAB法提取欧洲芜菁ECD04的DNA，使用Phanta Max Super-FidelityDNA Polymerase(南京诺唯赞生物科技公司)进行目的片段扩增。
50μl反应体系：2×Phanta Max Buffer 25μl、Forward Primer(10μM)2μl、Reverse Primer(10μM)2μl、dNTP Mix(10mM each)1μl、DNA模板2μl、ddH2O 19μl。
PCR反应条件：预变性95℃ 3min；变性95℃ 15s，复性55℃ 15s，延伸72℃ 8min，34个循环；72℃延伸5min；25℃ 5min。
扩增产物用1％的琼脂糖凝胶电泳检测，条带大小预期为8713bp(图15)。
对扩增出来的目的片段使用DNA Purification Kit(天根生化科技有限公司)试剂盒进行PCR原液回收并测量纯化回收后目的片段浓度。
酶切改造过的pCAMBIA3301(将Basta抗性改为Hyg抗性)植物表达载体。
酶切体系：
buffer 6μl，质粒DNA 50μl，NcoI FastDigest enzyme(s)1μl，Eco91I FastDigest enzyme(s)1μl，ddH2O 2μl。
将上述反应液离心混匀，置于37℃水浴锅中反应30min。反应结束后进行1％琼脂糖凝胶电泳检测，并切胶用DNA Purification Kit(天根生化科技有限公司)试剂盒回收并测量浓度。
使用ClonExpress II One Step Cloning Kit(南京诺唯赞生物科技公司)将目的基因片段和酶切好的植物表达载体进行连接。
将连接好的产物转入DH5α大肠感受态细胞：a.在冰上解冻克隆感受态细胞。b.取10μl重组产物加入到100μl感受态细胞中，轻弹管壁混匀(请勿振荡混匀)，冰上静置30min；c.42℃水浴热激45sec后，立即置于冰上冷却2-3min。d.加入900μl LB培养基(不添加抗生素)，37℃摇菌1h(转速200-250r/min)。e.将Kana抗性的LB平板固体培养基在37℃培养箱中预热。f.5000r/min离心5min，弃掉900μl上清。用剩余培养基将菌体重悬，用无菌涂布棒在含有Kana抗性的平板上轻轻涂匀。g.37℃培养箱中倒置培养12-16h。
单克隆阳性鉴定，在转化平板上挑取6个单克隆，于液体LB培养基中过夜培养。第二天直接吸取2μl菌液作为模板，进行阳性检测。
阳性检测正向引物序列(SEQ ID No.9)：5’-catggtctacaaggttccatg-3’
阳性检测反向引物序列(SEQ ID No.10)：5’-tcttcgctattacgccagc-3’
菌液PCR 12μl反应体系：2×Taq Master Mix(Dye Plus)6μl、Forward Primer(10μM)1μl、Reverse Primer(10μM)1μl、菌液2μl、ddH2O 2μl。
菌液PCR反应条件：预变性95℃ 3min；变性95℃ 15s，复性55℃ 15s，延伸72℃30s，34个循环；72℃延伸5min；25℃ 5min。
将PCR反应产物用1％琼脂糖凝胶电泳检测，条带在711bp为阳性单克隆(图16)，对于阳性克隆，直接将其转化好的DH5α菌液送往武汉擎科生物技术有限公司进行一代测序。
选取测序结果与基因组序列完全一致的克隆子扩大培养，使用TIANprep RapidMini Plasmid Kit(天根生化科技有限公司)提取质粒。将质粒体转化到农杆菌GV3101中，具体步骤如下：
取GV3101感受态细胞于冰上融化。向100μl感受态细胞中加入500ng重组质粒。冰上放置5min，转至液氮中放置5min，37℃水浴5min，冰上5min，加入400μl无抗LB，28℃摇床孵育1小时。取100μl转化产物涂布于带有Kana和Gent的抗性LB平板上。29℃培养36-48h。
8、将构建好的3301-hygr-212载体通过农杆菌介导的油菜下胚轴稳定遗传转化方法转化至甘蓝型油菜J9707。
9、将成功转入BraA08g039212E基因的株系培养至T1代，进行接中国根肿菌4号生理小种处理，观察其根部抗病表型。
把接菌后的株系表型分为三个等级，分别为Grade 0：Grade 1：Grade 2(图17)。Grade 0：主根与侧根完全不发病；Grade 1：侧根发生根肿现象；Grade 2：主根发生明显根肿现象。
统计BraA08g039212E的遗传转化所得株系L1和L2株系(2个独立转基因系)植株的抗病表型发现，L1株系接菌后表型分离比Grade 0：Grade 1：Grade 2＝12：16：7，L2株系接菌后表型分离比Grade 0：Grade 1：Grade 2＝9：16：7(表3)。
表3 BraA08g039212E转基因株系接“枝江”根肿菌表型分离比
对BraA08g039212E转基因株系接种表型抗感分离情况进行卡方(Chi-square)检验，L1株系X2＝1.67(X2<X2 0.05＝3.84),L2株系X2＝0.25(X2<X2 0.05＝3.84)，均符合3：1孟德尔分离定律。
计算接菌后株系的发病率和病情相关指数，公式如下：
株系发病率＝(感病株数/调查总株数)×100％
株系病情指数＝[∑(各级病株数×相应级数)/调查总株数×最高级别值]×100％
L1株系接菌后，发病率为65.70％，病情指数为42.90％；L2株系接菌后，发病率为71.80％，病情指数为46.90％(表4)。
表4 BraA08g039212E转基因株系接中国根肿菌4号生理小种的发病率以及病情指数
10、将接中国根肿菌4号生理小种的BraA08g039212E转基因株系进行基因型鉴定。
将L1和L2株系的样品用CTAB法提取DNA，用阳性检测引物。
12μl PCR扩增体系，2×Taq Master Mix(Dye Plus)6μl、Forward Primer(10μM)1μl、Reverse Primer(10μM)1μl、1μl、DNA模板1μl、ddH2O 3μl。
PCR反应条件：预变性95℃ 3min；变性95℃ 15s，复性55℃ 15s，延伸72℃ 30s，34个循环；72℃延伸5min；25℃ 5min，最后用1％琼脂糖凝胶电泳检测，711bp位置为目的条带(图18a，b)。
统计L1和L2株系基因型结果，L1株系基因型分离比为26：9，X2＝0.04(X2<X2 0.05＝3.84)，L2株系基因型分离比为23：9，X2＝0.17(X2<X2 0.05＝3.84)，均符合3：1孟德尔分离定律(图18a，b)。
11、BraA08g039212E基因在L1和L2株系接菌后的表达情况
使用
Super总RNA提取试剂盒(普洛麦格(北京)生物技术有限公司)对L1和L2株系接菌后根部样品的总RNA进行提取并进行RNA浓度测量。
根据样品RNA的浓度，以RNA含量1000ng为基准，用RNase-free ddH2O调整样品浓度使得RNA进样总体积为16μl，加入4×gDNA Wiper Mix，用移液器轻轻吹打混匀，42℃2min。
在上述反应管中直接加入5×HiScript II qRT SuperMix II，用移液器轻轻吹打混匀，然后进行逆转录反应：50℃ 15min，85℃5s。此时就得到了L1和L2株系接菌后的cDNA样品。
RT-PCR分析BraA08g039212E基因的表达量，利用CE Design设计BraA08g039212E基因的定量引物，定量引物序列如下：
正向定量引物(SEQ ID No.11)：5’-tggctgctcaaaccttc-3’
反向定量引物(SEQ ID No.12)：5’-acctcaacctgcgacaa-3’
qPCR反应体系：
qPCR SYBR Green Master Mix 11.8μl，F/R Primer 0.6μl，模板cDNA2μl，ddH2O to 15μl。
qPCR反应程序：95℃ 5min，95℃ 10s，52℃ 20s，72℃ 20s，40个循环，溶解曲线阶段默认设置1。
BraA08g039212E在转基因株系中的表达情况，该基因在J9707受体材料中稳定表达并且转基因材料的相对表达量均比9707受体材料高(图19)。
BraA08g039212E基因能够在受体材料J9707中稳定表达，并且能发挥其抗病功能，赋予感病材料9707抗根肿菌的能力。
BraA08g039212E基因在根肿病抗性育种和分子标记辅助选择中的应用
BraA08g039212E基因作为NBS-LRR基因，使得油菜转基因材料获得更好的根肿病抗性，可以抵御中国根肿菌4号生理小种的侵染，有效保护根肿菌污染田块中油菜的产量。BraA08g039212E除了对甘蓝型油菜有抗根肿效应，对其他农艺性状影响不大，表明该基因可以作为抗性基因通过转基因手段进行油菜、白菜和甘蓝等芸薹属植物的抗性育种。根据基因全长序列(包含上游序列和编码区)及与本基因连锁的其它序列所开发的标记进行的分子标记辅助育种，也可以大大加速根肿病抗性育种进程。
以上所述的实施例仅是对本发明的优选方式进行描述，并非对本发明的范围进行限定，在不脱离本发明设计精神的前提下，本领域普通技术人员对本发明的技术方案做出的各种变形和改进，均应落入本发明权利要求书确定的保护范围内。
序列表
<110> 华中农业大学
<120> 来源欧洲芜菁ECD04的BraA08g039212E基因在根肿菌抗性改良中的应用
<160> 12
<170> SIPOSequenceListing 1.0
<210> 1
<211> 8713
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 1
gagggtactg cggagggtac tgagccgtct caccatattg caagctcaac tcttcctggt 60
cacgacggat gaggtcttcc gtctcgctgt actcagatgt ctctccacca taacctccag 120
aatctgaagg ttggctgtaa ctatgtatcc ccattttata taaacctgca aaaacattta 180
tttcattagt acattataca taacgcaaga catcaccaaa caataattta aggaaacatt 240
ttcattactc cgaagcagaa atacattaga catggaagca gaaaaaacat aatttaaaaa 300
cagaattcaa gaaacagata cataataaaa tcgacaacat tttcagaaat acagagcgag 360
cagcttgttc ttcacaactt cctcaggctc ggttagtggt cctggcttgg ctaggagtgt 420
gtccagaatg gcaagctttg acaatctctc cttcagcagg agatcatcct tcttcaactc 480
cataacagtc gtaaaatcag ccatagattt tattccttga gtattattct ttctagcttt 540
agctgccttg ctaccctctg gacgcacctc ttcatcacca acagtagtgt taggtgtttg 600
cgaaccaaac tctccagcgt ttcgctttga actcacatta gttttaggag tgttaaggct 660
gagccacttc tgctcaaacc ttaacacaca ccacgcatgc tccagattga atttcctctt 720
gtgatcagcg aaataaatat catgagccac cttgaggaca tcagtgtcgc tctgaccaga 780
agtgatctgt ctctctgctg ccgcgtatgc accacaaaat ttgttggtgt agtcattaat 840
cttttgccac ctctgcttaa gatggagcca ctctataggt tcaccactct ctttagcatg 900
tggggttgat gcatagtact cagaaacacg tttccagaac gtcccggatt tttgttcgtt 960
tccaacaact gcatccttag atgtgttgag ccacgcactg attacgacct cgtcatcagc 1020
tggagtccat tgccttctct ccctagagcc aactggtggg tcttcacgtg aagctggagc 1080
gtcgggttgt tgtgaactaa atggaggtat ctcggatcct cttcctaagt tttgactcga 1140
aggaaaactt ccatactgga agtcttgact gtggagaaga tagctatagc taggggagtc 1200
actatatgga ttcactggat ccattatcga cctaactaag aggattatca aagaaagaga 1260
gagaaaccgg agagaaacaa gatagaaaca agagagaaac aagagggaga gataagagag 1320
aaagagataa ccacaacaag aagtaaagct tgttgcggat tatataaaga aataagagaa 1380
ctagatatca taaagacaaa gataacaacc atcaaagagc tttaaataaa aacctaacaa 1440
tcaactacta gggtctaatc aatagttatt acaagcataa caaccatcaa agcaatcaag 1500
taactaaata tgattttcat ttggttagaa accaaataca gtccactgac aaagagccat 1560
caaagcaatc aagttatact cgttacaagc ataacaacac tcactacaac gtcttccttt 1620
aatgtccaca ccaacttaaa caaatacaga cggcttttga gttcgaaact attaactaca 1680
aaaccaaact aacctcaaca atcactacaa cgtcttcctt aaatgtccac acctacttag 1740
tttaatttct tatttgtttt cattgtggaa ttcactaatc aatcaagaat acaatcaaat 1800
tgttaaacaa ccacagctca atcgagtata aaatcccgaa tttattctaa actacacaac 1860
aagcaaccac agatcagttc ataatcacat tcctcgccaa caaaaaccat caggaactat 1920
acaatcaaac attcagaaac tatacagtcg aactatacaa ccaaacagtc agtgacaaac 1980
caatcaatta gtttatatga ttcctctaaa tctttcgcta gaacacacat agaatattta 2040
cagagataga agacaaacct tgtagcgttt cgattcggag gacgacggcg agatccttct 2100
ggattgggtg agaatagaaa cgggcctccg aaatctaccg gagtgagaga gattagtgag 2160
actatgcaca ccgagatcaa tcaccacaca tgcaaaaagc catcaaacca aaaaaaagat 2220
tagatcgaca atctccctga ttataaaacc aaaaaaaaac catcttcgaa gaaggacgaa 2280
cctgctgaag attcgatgcg gggtttgagc cttcgatcgg tgtggtaagc gacgccgctt 2340
cactgctcga gaaatcgcct tctccggaga cagagagaga gagagaagag ggagacgaag 2400
gaaatgaatc gagacggaac gacctcttct ctctaccgct tccaacttcc tattaccgct 2460
tgccacgtga cacaagcact gcgaccacaa acttcttacc taaaaaacga ttctattaat 2520
attctccatt tttgatttaa attttaactt gaattgggct aagcatcgag gcttagaagc 2580
ccgataatgt tgctcttaag agcatgtgca aaggcaaaag actcttaggg gactttttct 2640
ttatttttta ttttgtctga tttaaaaaag aaaaattaaa aaaaaattat tgtgaatcgc 2700
tacatgtcag tgggagtcta tgaaaagtgg caaaaccctc tcagactcgc ctcttacgaa 2760
agaagcagaa gagacgtgat tttaaattat ggtaaggccc acataatttt aaattatttt 2820
ttttaaggag cagccttaag aggccccgtt gcacatgctc taacacaaga tggtgtggaa 2880
aaatttgcaa ggctttttag aaaagtcaaa gaagttgtgc tgaatgctga cgatgcaaag 2940
gaaccttcgt ggaaagattc cttgtccgaa cgactcatat attttaaagt cgtctcctct 3000
tcccccttca cttccgaaag agtccccgta tccagaaagc tgcatccgtt aatggcatct 3060
tcttcgtctt cctccaagcg ccagttcgac gtgtttgtga gtttcagagg cgccgacacg 3120
cgcaatacct tcaccgctca tctcctcaag tacctaagtg ggaaaggtat agacgctttc 3180
tctgacggga aactcctgag aggcgatgac ctgtcggttc tctttggccg gatcgagcaa 3240
tcgaagatgt cgatcgttgt cttctcggag aagtacgcca actccacctg gtgcttggag 3300
gaactctgga agatcatgca gtgcaggaag gaatccggtc atggggttat acccatcttc 3360
tacaaagtca agaaatctga tgtggaaatt cagaaaggga gtttcggagc tccatttcaa 3420
agccctaaag agagtttcaa gggagatggg cacaagattg aggaatggaa ggaagctctg 3480
aggactgctt ccaatgttct tggctttgta tatcctgagg acaggtagta atagctagtg 3540
ccttctgagc tcattattat ttggatttta ggaatattga aactgttctt ctatatattt 3600
ttgattctga aaagtgagtt agatctcatt catatatata acttttgtga ttaggaaaaa 3660
tgcttaagga agttttgctt cagggaaaat gaatagttct aagaggttgt aaattattta 3720
taaaaacgga tatttgcatc aatgtgtttt ctttctaagt gttctttggg gttaatttac 3780
cttttgttca caacatatat gcattcaatc atgttcgata ttgaacattt ggaccgactt 3840
tgcccaattt gcagaccgga gactgaattt ctcgatgaaa tcaccaagga taccttaagg 3900
atgatcaatg atttgtctcc atgtgaaacc agtggtctgc cggggatcga atcacgttca 3960
aaaaaactgg aggagttact catgtttggt aatgatgaat gtgtccgtac catcggagtt 4020
cttgggatga ctggaatcgg caagacactg gttgctgata tcgtctataa acgaaattgc 4080
aggcaatttg atggttacga tttcgtttat gacgttgaca gggagttgga attgcatcag 4140
ttgtgtcatt tgcgagagaa tctcctctgt aaagtattgg acgtagaaaa tttggatgac 4200
agagcgcacg gaagatcgga gaactatctt cggaacaaga agttgtttat cgttcttgat 4260
aatgtgactg ataaagaaca gatagatgtt ctcatcggag ataaagcagt gtaccggaaa 4320
ggaaccagga ttgttataat aaccagagac aagaagctgt tggagaacaa agctgacgca 4380
acatatgtag ttcccacact aaatgacacg gaagccatgg agcttttctg tcttagtgca 4440
ttccctagca acctctaccc gtcggaagaa tatattgatc tatcagaaaa attcgtatat 4500
tatgctaaag gtcatccttc agctttgaag tgtttaggtt cgggtctact taagaaggat 4560
aaatcatact ggagatggaa atgggagagc ttagaggtaa tgccagacaa ggatattcag 4620
agagtgctag aaaagagtta taagaaacta gatgatcaac agaagagcat gtttctggac 4680
atagcatgct ttttcagatc agaaaaagca gagttcatta caagcatcct gaaatcagac 4740
aacatcaatg ctgcagcagc tgtgatgcaa gagcttgtag ataaatgctt gttaactatt 4800
tcttatgata ggcttgagat gcatgatctc ttgcatataa tgggaaaaga tattggatat 4860
gaatcatcaa tcaaaaggct gtggaaacac aaagatattc gtcgtatcct ggagcggaac 4920
acggtcagct acacgtttcc attctttctc ccttaatttt ctctctctct atatatatat 4980
ccatttaaag tgattcttct cgtattgttt tctagctaac atttccatgt ccttaatctt 5040
ttatcttgct accagggtac tgaaaatgta agaggcatct tcttgaacat gtctgatgtc 5100
gaaaggatca agcttagtcc tgctgctttc atgaggatgt caaatctcaa gttcctgaaa 5160
ttccacaatt ctcattgttc tcagtggtgt gacaatgacc ataaatttca gttctgcaga 5220
gggctcgatc aatttccaga tgagcttgtg tatcttcact ggcaggggta cccttacaat 5280
caccttccat cagacttcta tccggatgaa cttgtggact taaatctgcg ttatagccgc 5340
atccaacaac tgtgggaaga ggaacaggta acgctacgtg ttttcttatc ttaaaaacat 5400
tactctttgg tttctctgct gaataattat ttgtcaatgt gccacagaat acagaaaatt 5460
tgagatgggt cgacctcagt cagtcaaaag acttgctcaa cttatcaggt ttgtccaagg 5520
ccaaaaatct cgaaagactg gatctcgaag gctgcactag tttggatacg ttaggcccat 5580
caatcgaaca gatgaacaag cttatttacc tgaacctccg agagtgcact agccttaaga 5640
gtctcccaga gggaatcaac ttagaatctc tgaagactct gatcctcagt ggctgctcaa 5700
accttcggta ctttcatatc atatcagaga gtattgaatc cctgtatttg gaaggctcag 5760
caatcgaaca agtcgttgaa cacatccaga gtcttcggag tctcatttta ctgagtctca 5820
agaattgtcg caggttgagg tgtcttccca acgatcttta caagctgaaa tctcttcaag 5880
aactgattct ctctggctgt tcagcgctgg agagtcttcc acccatcaga gaggaaatgg 5940
aatgcttaga gatcttgctt atggatggaa cgtccatcaa acagacacct gaaaccattt 6000
gtttgagtaa cctcaaggtt ttttcgctct gtgaatctag catcgacgat tccacagggt 6060
tggtactgcg gcctttctct ggcagctctc gtctattgga cctctatctc gcgaactgca 6120
atatcagcaa gttgccgaac aacttcagca gctccttaca ctcattgcgg tgtctatgct 6180
taagcagaaa caacattgag actctacccg aaagcatcga gaaactttat tctctgctgt 6240
tgcttgactt gaaacattgc cgcaggctca gttctctgcc ggtgcttcca tctaatctac 6300
agtgcttaga tgctcatggg tgtgtttctc tggaaaaggt tgctaaacaa gtaaaggttc 6360
ccctagtagc tgagaggatg catactaatt tcattttcac ggattgcttt atgctgaacc 6420
gtgctgagca agaagctatt gtagctcagg cccaactcaa gagtcagcta ctagcaagga 6480
catctcttca gtataatctt aaggtccgtt ttcaaaaccc taaaactcta aataaaaaca 6540
ctaaacccta aaattttcta aaccctaaat cttaaatttt aaactcttga gtgttttaga 6600
gtttaagatt tatcctaaag tttagggttt acccaaaggt ttagggttta ggatttaagg 6660
tttagagatt aagatttagg gtttaatgtt ttgctgacga cgttaaattt ttttttttgt 6720
aattactact attttttatt tatttctttt tatcttttaa ttttaaaaag ataatataat 6780
ttgacaatat tttgtttcat tttttaaaag ataacaaata tgaaataaca taatcttatt 6840
ggttgttgaa cctagaaccc aaaaatatat ctttcttata ttagtccatg gccggtcttg 6900
agcataggcc agtgaaatat tggtctatga ttcccaaatt ttgttttgaa gaaaatacat 6960
taaaataggt tttcaaactt ataaaaatta tttttattaa aattcttatt aaatcatata 7020
tggcccccaa aatctcaggg ttgacctaag agtcattcca actttttatg tgaaatttta 7080
tctctaatac acaccacgct tatgctttaa tgatttccgt tgtaactcat ttcagggact 7140
agttatggat cccctggcta ccgtttgctt tccaggaagt gacatacctt cgtggttatg 7200
tcaacagaga atgggatctt ccatagaaac cgacctggtt tcacactggt gtaatagtaa 7260
atttattgga gtttccctaa gtgttgttgt cagcttcaaa ggccatgaag attatcatgt 7320
caaccgttta tctgtaagat gcaagtgcaa cttcaaaaat caaaacggtc agtctatcag 7380
ctttagtttc tcgcttggag gatggaacga ctcgtgtggt tcatcttgcc atgaaccacg 7440
gaaacttgga tctgaccatg tgtttattag ctataacaaa tgtaatgtgc cagtcttccg 7500
atggagtgaa ggcagtgatg aagctaatag atgtcgtccc actagtgtcg catttgaatt 7560
ctaccttact gatgacactg aaaagaaact agaaagctgc aaggtgacaa ggtgtgggat 7620
gagtttgcta tatgctcctg atgagaatga ccgtggattc cagggaacac gggttacaga 7680
tatagttgag catacatcga gtgaagcttt tgtgcccata agaggttggt cacactcgca 7740
agttggagaa agaaggaatg gtataataag agatgaaatc cccttatgag tgtttattcc 7800
agagctggag gtcactggtc gtgaaatcat ttaaaggtac tagaatgctg cttaatagct 7860
ttcataagta tatgttatca ttttaaaaca caatttacca ttgaattagg gacttgacgc 7920
ctcattacta ttcagaccct ttggattttc aaagaaaagt tctatctact tatctggctg 7980
aagtttgacg tgaagcttat ccttggatct gattatcatt tcaggatgat cgatgaggtc 8040
gtcactttac tgcaactttg ttctgcgtcc caaatacaaa acagagtatt gttttctttc 8100
tctgtttctc cctttgcatg gtctacaagg ttccatgttt tgataaggca ttgtataaaa 8160
tgtatgtgcc caatgtcgaa gggaatttga tgaaagtcgc gggataaatc caaattattg 8220
ggccatgatt ttttgttatg tagttgatta aagaaacttg tatgtttgat gattgttggt 8280
atatgtaatt tgtggtatag agatgattgt tgtttgaagc ataaggaatg agtgtttgtt 8340
agctacgaag ttatctgtat tcattctatg acattagact catggatttg gcttgccacg 8400
aaagtgtgga tgtttagttg cagctgaaat ctccagctct ttctttgcaa actctatttc 8460
ttctcccaag ttccatacta aagcttttta gtccgattct tgaatcttga ttggttcagt 8520
attgacgtag atatgcttga ctgcgttgtt gagtggtcgt acttagaaag ctttaaagat 8580
tgctacgtta tggtttgact tttaggttgt ctaccagaca gaccaaaagt gtttacgaat 8640
ttgtcgacaa ggtatctttg aagagcttaa agctactaga agattcacag ctagatgggc 8700
cagtcactag gaa 8713
<210> 2
<211> 3564
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 2
atggcatctt cttcgtcttc ctccaagcgc cagttcgacg tgtttgtgag tttcagaggc 60
gccgacacgc gcaatacctt caccgctcat ctcctcaagt acctaagtgg gaaaggtata 120
gacgctttct ctgacgggaa actcctgaga ggcgatgacc tgtcggttct ctttggccgg 180
atcgagcaat cgaagatgtc gatcgttgtc ttctcggaga agtacgccaa ctccacctgg 240
tgcttggagg aactctggaa gatcatgcag tgcaggaagg aatccggtca tggggttata 300
cccatcttct acaaagtcaa gaaatctgat gtggaaattc agaaagggag tttcggagct 360
ccatttcaaa gccctaaaga gagtttcaag ggagatgggc acaagattga ggaatggaag 420
gaagctctga ggactgcttc caatgttctt ggctttgtat atcctgagga cagaccggag 480
actgaatttc tcgatgaaat caccaaggat accttaagga tgatcaatga tttgtctcca 540
tgtgaaacca gtggtctgcc ggggatcgaa tcacgttcaa aaaaactgga ggagttactc 600
atgtttggta atgatgaatg tgtccgtacc atcggagttc ttgggatgac tggaatcggc 660
aagacactgg ttgctgatat cgtctataaa cgaaattgca ggcaatttga tggttacgat 720
ttcgtttatg acgttgacag ggagttggaa ttgcatcagt tgtgtcattt gcgagagaat 780
ctcctctgta aagtattgga cgtagaaaat ttggatgaca gagcgcacgg aagatcggag 840
aactatcttc ggaacaagaa gttgtttatc gttcttgata atgtgactga taaagaacag 900
atagatgttc tcatcggaga taaagcagtg taccggaaag gaaccaggat tgttataata 960
accagagaca agaagctgtt ggagaacaaa gctgacgcaa catatgtagt tcccacacta 1020
aatgacacgg aagccatgga gcttttctgt cttagtgcat tccctagcaa cctctacccg 1080
tcggaagaat atattgatct atcagaaaaa ttcgtatatt atgctaaagg tcatccttca 1140
gctttgaagt gtttaggttc gggtctactt aagaaggata aatcatactg gagatggaaa 1200
tgggagagct tagaggtaat gccagacaag gatattcaga gagtgctaga aaagagttat 1260
aagaaactag atgatcaaca gaagagcatg tttctggaca tagcatgctt tttcagatca 1320
gaaaaagcag agttcattac aagcatcctg aaatcagaca acatcaatgc tgcagcagct 1380
gtgatgcaag agcttgtaga taaatgcttg ttaactattt cttatgatag gcttgagatg 1440
catgatctct tgcatataat gggaaaagat attggatatg aatcatcaat caaaaggctg 1500
tggaaacaca aagatattcg tcgtatcctg gagcggaaca cgggtactga aaatgtaaga 1560
ggcatcttct tgaacatgtc tgatgtcgaa aggatcaagc ttagtcctgc tgctttcatg 1620
aggatgtcaa atctcaagtt cctgaaattc cacaattctc attgttctca gtggtgtgac 1680
aatgaccata aatttcagtt ctgcagaggg ctcgatcaat ttccagatga gcttgtgtat 1740
cttcactggc aggggtaccc ttacaatcac cttccatcag acttctatcc ggatgaactt 1800
gtggacttaa atctgcgtta tagccgcatc caacaactgt gggaagagga acagaataca 1860
gaaaatttga gatgggtcga cctcagtcag tcaaaagact tgctcaactt atcaggtttg 1920
tccaaggcca aaaatctcga aagactggat ctcgaaggct gcactagttt ggatacgtta 1980
ggcccatcaa tcgaacagat gaacaagctt atttacctga acctccgaga gtgcactagc 2040
cttaagagtc tcccagaggg aatcaactta gaatctctga agactctgat cctcagtggc 2100
tgctcaaacc ttcggtactt tcatatcata tcagagagta ttgaatccct gtatttggaa 2160
ggctcagcaa tcgaacaagt cgttgaacac atccagagtc ttcggagtct cattttactg 2220
agtctcaaga attgtcgcag gttgaggtgt cttcccaacg atctttacaa gctgaaatct 2280
cttcaagaac tgattctctc tggctgttca gcgctggaga gtcttccacc catcagagag 2340
gaaatggaat gcttagagat cttgcttatg gatggaacgt ccatcaaaca gacacctgaa 2400
accatttgtt tgagtaacct caaggttttt tcgctctgtg aatctagcat cgacgattcc 2460
acagggttgg tactgcggcc tttctctggc agctctcgtc tattggacct ctatctcgcg 2520
aactgcaata tcagcaagtt gccgaacaac ttcagcagct ccttacactc attgcggtgt 2580
ctatgcttaa gcagaaacaa cattgagact ctacccgaaa gcatcgagaa actttattct 2640
ctgctgttgc ttgacttgaa acattgccgc aggctcagtt ctctgccggt gcttccatct 2700
aatctacagt gcttagatgc tcatgggtgt gtttctctgg aaaaggttgc taaacaagta 2760
aaggttcccc tagtagctga gaggatgcat actaatttca ttttcacgga ttgctttatg 2820
ctgaaccgtg ctgagcaaga agctattgta gctcaggccc aactcaagag tcagctacta 2880
gcaaggacat ctcttcagta taatcttaag ggactagtta tggatcccct ggctaccgtt 2940
tgctttccag gaagtgacat accttcgtgg ttatgtcaac agagaatggg atcttccata 3000
gaaaccgacc tggtttcaca ctggtgtaat agtaaattta ttggagtttc cctaagtgtt 3060
gttgtcagct tcaaaggcca tgaagattat catgtcaacc gtttatctgt aagatgcaag 3120
tgcaacttca aaaatcaaaa cggtcagtct atcagcttta gtttctcgct tggaggatgg 3180
aacgactcgt gtggttcatc ttgccatgaa ccacggaaac ttggatctga ccatgtgttt 3240
attagctata acaaatgtaa tgtgccagtc ttccgatgga gtgaaggcag tgatgaagct 3300
aatagatgtc gtcccactag tgtcgcattt gaattctacc ttactgatga cactgaaaag 3360
aaactagaaa gctgcaaggt gacaaggtgt gggatgagtt tgctatatgc tcctgatgag 3420
aatgaccgtg gattccaggg aacacgggtt acagatatag ttgagcatac atcgagtgaa 3480
gcttttgtgc ccataagagg ttggtcacac tcgcaagttg gagaaagaag gaatggtata 3540
ataagagatg aaatcccctt atga 3564
<210> 3
<211> 1187
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 3
Met Ala Ser Ser Ser Ser Ser Ser Lys Arg Gln Phe Asp Val Phe Val
1 5 10 15
Ser Phe Arg Gly Ala Asp Thr Arg Asn Thr Phe Thr Ala His Leu Leu
20 25 30
Lys Tyr Leu Ser Gly Lys Gly Ile Asp Ala Phe Ser Asp Gly Lys Leu
35 40 45
Leu Arg Gly Asp Asp Leu Ser Val Leu Phe Gly Arg Ile Glu Gln Ser
50 55 60
Lys Met Ser Ile Val Val Phe Ser Glu Lys Tyr Ala Asn Ser Thr Trp
65 70 75 80
Cys Leu Glu Glu Leu Trp Lys Ile Met Gln Cys Arg Lys Glu Ser Gly
85 90 95
His Gly Val Ile Pro Ile Phe Tyr Lys Val Lys Lys Ser Asp Val Glu
100 105 110
Ile Gln Lys Gly Ser Phe Gly Ala Pro Phe Gln Ser Pro Lys Glu Ser
115 120 125
Phe Lys Gly Asp Gly His Lys Ile Glu Glu Trp Lys Glu Ala Leu Arg
130 135 140
Thr Ala Ser Asn Val Leu Gly Phe Val Tyr Pro Glu Asp Arg Pro Glu
145 150 155 160
Thr Glu Phe Leu Asp Glu Ile Thr Lys Asp Thr Leu Arg Met Ile Asn
165 170 175
Asp Leu Ser Pro Cys Glu Thr Ser Gly Leu Pro Gly Ile Glu Ser Arg
180 185 190
Ser Lys Lys Leu Glu Glu Leu Leu Met Phe Gly Asn Asp Glu Cys Val
195 200 205
Arg Thr Ile Gly Val Leu Gly Met Thr Gly Ile Gly Lys Thr Leu Val
210 215 220
Ala Asp Ile Val Tyr Lys Arg Asn Cys Arg Gln Phe Asp Gly Tyr Asp
225 230 235 240
Phe Val Tyr Asp Val Asp Arg Glu Leu Glu Leu His Gln Leu Cys His
245 250 255
Leu Arg Glu Asn Leu Leu Cys Lys Val Leu Asp Val Glu Asn Leu Asp
260 265 270
Asp Arg Ala His Gly Arg Ser Glu Asn Tyr Leu Arg Asn Lys Lys Leu
275 280 285
Phe Ile Val Leu Asp Asn Val Thr Asp Lys Glu Gln Ile Asp Val Leu
290 295 300
Ile Gly Asp Lys Ala Val Tyr Arg Lys Gly Thr Arg Ile Val Ile Ile
305 310 315 320
Thr Arg Asp Lys Lys Leu Leu Glu Asn Lys Ala Asp Ala Thr Tyr Val
325 330 335
Val Pro Thr Leu Asn Asp Thr Glu Ala Met Glu Leu Phe Cys Leu Ser
340 345 350
Ala Phe Pro Ser Asn Leu Tyr Pro Ser Glu Glu Tyr Ile Asp Leu Ser
355 360 365
Glu Lys Phe Val Tyr Tyr Ala Lys Gly His Pro Ser Ala Leu Lys Cys
370 375 380
Leu Gly Ser Gly Leu Leu Lys Lys Asp Lys Ser Tyr Trp Arg Trp Lys
385 390 395 400
Trp Glu Ser Leu Glu Val Met Pro Asp Lys Asp Ile Gln Arg Val Leu
405 410 415
Glu Lys Ser Tyr Lys Lys Leu Asp Asp Gln Gln Lys Ser Met Phe Leu
420 425 430
Asp Ile Ala Cys Phe Phe Arg Ser Glu Lys Ala Glu Phe Ile Thr Ser
435 440 445
Ile Leu Lys Ser Asp Asn Ile Asn Ala Ala Ala Ala Val Met Gln Glu
450 455 460
Leu Val Asp Lys Cys Leu Leu Thr Ile Ser Tyr Asp Arg Leu Glu Met
465 470 475 480
His Asp Leu Leu His Ile Met Gly Lys Asp Ile Gly Tyr Glu Ser Ser
485 490 495
Ile Lys Arg Leu Trp Lys His Lys Asp Ile Arg Arg Ile Leu Glu Arg
500 505 510
Asn Thr Gly Thr Glu Asn Val Arg Gly Ile Phe Leu Asn Met Ser Asp
515 520 525
Val Glu Arg Ile Lys Leu Ser Pro Ala Ala Phe Met Arg Met Ser Asn
530 535 540
Leu Lys Phe Leu Lys Phe His Asn Ser His Cys Ser Gln Trp Cys Asp
545 550 555 560
Asn Asp His Lys Phe Gln Phe Cys Arg Gly Leu Asp Gln Phe Pro Asp
565 570 575
Glu Leu Val Tyr Leu His Trp Gln Gly Tyr Pro Tyr Asn His Leu Pro
580 585 590
Ser Asp Phe Tyr Pro Asp Glu Leu Val Asp Leu Asn Leu Arg Tyr Ser
595 600 605
Arg Ile Gln Gln Leu Trp Glu Glu Glu Gln Asn Thr Glu Asn Leu Arg
610 615 620
Trp Val Asp Leu Ser Gln Ser Lys Asp Leu Leu Asn Leu Ser Gly Leu
625 630 635 640
Ser Lys Ala Lys Asn Leu Glu Arg Leu Asp Leu Glu Gly Cys Thr Ser
645 650 655
Leu Asp Thr Leu Gly Pro Ser Ile Glu Gln Met Asn Lys Leu Ile Tyr
660 665 670
Leu Asn Leu Arg Glu Cys Thr Ser Leu Lys Ser Leu Pro Glu Gly Ile
675 680 685
Asn Leu Glu Ser Leu Lys Thr Leu Ile Leu Ser Gly Cys Ser Asn Leu
690 695 700
Arg Tyr Phe His Ile Ile Ser Glu Ser Ile Glu Ser Leu Tyr Leu Glu
705 710 715 720
Gly Ser Ala Ile Glu Gln Val Val Glu His Ile Gln Ser Leu Arg Ser
725 730 735
Leu Ile Leu Leu Ser Leu Lys Asn Cys Arg Arg Leu Arg Cys Leu Pro
740 745 750
Asn Asp Leu Tyr Lys Leu Lys Ser Leu Gln Glu Leu Ile Leu Ser Gly
755 760 765
Cys Ser Ala Leu Glu Ser Leu Pro Pro Ile Arg Glu Glu Met Glu Cys
770 775 780
Leu Glu Ile Leu Leu Met Asp Gly Thr Ser Ile Lys Gln Thr Pro Glu
785 790 795 800
Thr Ile Cys Leu Ser Asn Leu Lys Val Phe Ser Leu Cys Glu Ser Ser
805 810 815
Ile Asp Asp Ser Thr Gly Leu Val Leu Arg Pro Phe Ser Gly Ser Ser
820 825 830
Arg Leu Leu Asp Leu Tyr Leu Ala Asn Cys Asn Ile Ser Lys Leu Pro
835 840 845
Asn Asn Phe Ser Ser Ser Leu His Ser Leu Arg Cys Leu Cys Leu Ser
850 855 860
Arg Asn Asn Ile Glu Thr Leu Pro Glu Ser Ile Glu Lys Leu Tyr Ser
865 870 875 880
Leu Leu Leu Leu Asp Leu Lys His Cys Arg Arg Leu Ser Ser Leu Pro
885 890 895
Val Leu Pro Ser Asn Leu Gln Cys Leu Asp Ala His Gly Cys Val Ser
900 905 910
Leu Glu Lys Val Ala Lys Gln Val Lys Val Pro Leu Val Ala Glu Arg
915 920 925
Met His Thr Asn Phe Ile Phe Thr Asp Cys Phe Met Leu Asn Arg Ala
930 935 940
Glu Gln Glu Ala Ile Val Ala Gln Ala Gln Leu Lys Ser Gln Leu Leu
945 950 955 960
Ala Arg Thr Ser Leu Gln Tyr Asn Leu Lys Gly Leu Val Met Asp Pro
965 970 975
Leu Ala Thr Val Cys Phe Pro Gly Ser Asp Ile Pro Ser Trp Leu Cys
980 985 990
Gln Gln Arg Met Gly Ser Ser Ile Glu Thr Asp Leu Val Ser His Trp
995 1000 1005
Cys Asn Ser Lys Phe Ile Gly Val Ser Leu Ser Val Val Val Ser Phe
1010 1015 1020
Lys Gly His Glu Asp Tyr His Val Asn Arg Leu Ser Val Arg Cys Lys
1025 1030 1035 1040
Cys Asn Phe Lys Asn Gln Asn Gly Gln Ser Ile Ser Phe Ser Phe Ser
1045 1050 1055
Leu Gly Gly Trp Asn Asp Ser Cys Gly Ser Ser Cys His Glu Pro Arg
1060 1065 1070
Lys Leu Gly Ser Asp His Val Phe Ile Ser Tyr Asn Lys Cys Asn Val
1075 1080 1085
Pro Val Phe Arg Trp Ser Glu Gly Ser Asp Glu Ala Asn Arg Cys Arg
1090 1095 1100
Pro Thr Ser Val Ala Phe Glu Phe Tyr Leu Thr Asp Asp Thr Glu Lys
1105 1110 1115 1120
Lys Leu Glu Ser Cys Lys Val Thr Arg Cys Gly Met Ser Leu Leu Tyr
1125 1130 1135
Ala Pro Asp Glu Asn Asp Arg Gly Phe Gln Gly Thr Arg Val Thr Asp
1140 1145 1150
Ile Val Glu His Thr Ser Ser Glu Ala Phe Val Pro Ile Arg Gly Trp
1155 1160 1165
Ser His Ser Gln Val Gly Glu Arg Arg Asn Gly Ile Ile Arg Asp Glu
1170 1175 1180
Ile Pro Leu
1185
<210> 4
<211> 3051
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 4
gagggtactg cggagggtac tgagccgtct caccatattg caagctcaac tcttcctggt 60
cacgacggat gaggtcttcc gtctcgctgt actcagatgt ctctccacca taacctccag 120
aatctgaagg ttggctgtaa ctatgtatcc ccattttata taaacctgca aaaacattta 180
tttcattagt acattataca taacgcaaga catcaccaaa caataattta aggaaacatt 240
ttcattactc cgaagcagaa atacattaga catggaagca gaaaaaacat aatttaaaaa 300
cagaattcaa gaaacagata cataataaaa tcgacaacat tttcagaaat acagagcgag 360
cagcttgttc ttcacaactt cctcaggctc ggttagtggt cctggcttgg ctaggagtgt 420
gtccagaatg gcaagctttg acaatctctc cttcagcagg agatcatcct tcttcaactc 480
cataacagtc gtaaaatcag ccatagattt tattccttga gtattattct ttctagcttt 540
agctgccttg ctaccctctg gacgcacctc ttcatcacca acagtagtgt taggtgtttg 600
cgaaccaaac tctccagcgt ttcgctttga actcacatta gttttaggag tgttaaggct 660
gagccacttc tgctcaaacc ttaacacaca ccacgcatgc tccagattga atttcctctt 720
gtgatcagcg aaataaatat catgagccac cttgaggaca tcagtgtcgc tctgaccaga 780
agtgatctgt ctctctgctg ccgcgtatgc accacaaaat ttgttggtgt agtcattaat 840
cttttgccac ctctgcttaa gatggagcca ctctataggt tcaccactct ctttagcatg 900
tggggttgat gcatagtact cagaaacacg tttccagaac gtcccggatt tttgttcgtt 960
tccaacaact gcatccttag atgtgttgag ccacgcactg attacgacct cgtcatcagc 1020
tggagtccat tgccttctct ccctagagcc aactggtggg tcttcacgtg aagctggagc 1080
gtcgggttgt tgtgaactaa atggaggtat ctcggatcct cttcctaagt tttgactcga 1140
aggaaaactt ccatactgga agtcttgact gtggagaaga tagctatagc taggggagtc 1200
actatatgga ttcactggat ccattatcga cctaactaag aggattatca aagaaagaga 1260
gagaaaccgg agagaaacaa gatagaaaca agagagaaac aagagggaga gataagagag 1320
aaagagataa ccacaacaag aagtaaagct tgttgcggat tatataaaga aataagagaa 1380
ctagatatca taaagacaaa gataacaacc atcaaagagc tttaaataaa aacctaacaa 1440
tcaactacta gggtctaatc aatagttatt acaagcataa caaccatcaa agcaatcaag 1500
taactaaata tgattttcat ttggttagaa accaaataca gtccactgac aaagagccat 1560
caaagcaatc aagttatact cgttacaagc ataacaacac tcactacaac gtcttccttt 1620
aatgtccaca ccaacttaaa caaatacaga cggcttttga gttcgaaact attaactaca 1680
aaaccaaact aacctcaaca atcactacaa cgtcttcctt aaatgtccac acctacttag 1740
tttaatttct tatttgtttt cattgtggaa ttcactaatc aatcaagaat acaatcaaat 1800
tgttaaacaa ccacagctca atcgagtata aaatcccgaa tttattctaa actacacaac 1860
aagcaaccac agatcagttc ataatcacat tcctcgccaa caaaaaccat caggaactat 1920
acaatcaaac attcagaaac tatacagtcg aactatacaa ccaaacagtc agtgacaaac 1980
caatcaatta gtttatatga ttcctctaaa tctttcgcta gaacacacat agaatattta 2040
cagagataga agacaaacct tgtagcgttt cgattcggag gacgacggcg agatccttct 2100
ggattgggtg agaatagaaa cgggcctccg aaatctaccg gagtgagaga gattagtgag 2160
actatgcaca ccgagatcaa tcaccacaca tgcaaaaagc catcaaacca aaaaaaagat 2220
tagatcgaca atctccctga ttataaaacc aaaaaaaaac catcttcgaa gaaggacgaa 2280
cctgctgaag attcgatgcg gggtttgagc cttcgatcgg tgtggtaagc gacgccgctt 2340
cactgctcga gaaatcgcct tctccggaga cagagagaga gagagaagag ggagacgaag 2400
gaaatgaatc gagacggaac gacctcttct ctctaccgct tccaacttcc tattaccgct 2460
tgccacgtga cacaagcact gcgaccacaa acttcttacc taaaaaacga ttctattaat 2520
attctccatt tttgatttaa attttaactt gaattgggct aagcatcgag gcttagaagc 2580
ccgataatgt tgctcttaag agcatgtgca aaggcaaaag actcttaggg gactttttct 2640
ttatttttta ttttgtctga tttaaaaaag aaaaattaaa aaaaaattat tgtgaatcgc 2700
tacatgtcag tgggagtcta tgaaaagtgg caaaaccctc tcagactcgc ctcttacgaa 2760
agaagcagaa gagacgtgat tttaaattat ggtaaggccc acataatttt aaattatttt 2820
ttttaaggag cagccttaag aggccccgtt gcacatgctc taacacaaga tggtgtggaa 2880
aaatttgcaa ggctttttag aaaagtcaaa gaagttgtgc tgaatgctga cgatgcaaag 2940
gaaccttcgt ggaaagattc cttgtccgaa cgactcatat attttaaagt cgtctcctct 3000
tcccccttca cttccgaaag agtccccgta tccagaaagc tgcatccgtt a 3051
<210> 5
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
atggatccaa tagagacagt gg 22
<210> 6
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 6
ttatggaagg ctctcaagat ttg 23
<210> 7
<211> 38
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
gctcggtacc cggggatccg agggtactgc ggagggta 38
<210> 8
<211> 39
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 8
gacggccagt gccaagcttt tcctagtgac tggcccatc 39
<210> 9
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 9
catggtctac aaggttccat g 21
<210> 10
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 10
tcttcgctat tacgccagc 19
<210> 11
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 11
tggctgctca aaccttc 17
<210> 12
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 12
acctcaacct gcgacaa 17
Claims (9)
1.BraA08g039212E基因在赋予根肿菌抗性中的应用，其特征在于，所述BraA08g039212E基因的核苷酸序列如SEQ ID No.1或SEQ ID No.2所示，其编码蛋白的氨基酸序列如SEQ ID No.3所示。
2.根据权利要求1所述的应用，其特征在于，所述BraA08g039212E基因位于欧洲芜菁ECD04的A08染色体上，其物理位置为14863686-14873201。
3.BraA08g039212E基因在根肿病抗病育种中的应用，其特征在于，所述BraA08g039212E基因通过转基因或有性杂交转育途径用于芸薹属植物根肿病抗病育种中的应用，所述芸薹属植物包括油菜、白菜、甘蓝或萝卜，所述BraA08g039212E基因的核苷酸序列如SEQ ID No.1或SEQ ID No.2所示。
4.BraA08g039212E基因及根据其所开发的标记在芸薹属植物分子标记辅助选择中的应用，其特征在于，所述芸薹属植物包括油菜、白菜、甘蓝或萝卜，所述BraA08g039212E基因的核苷酸序列如SEQ ID No.1或SEQ ID No.2所示。
5.BraA08g039212E基因的启动子的用途，其特征在于，用于调控BraA08g039212E基因的表达，其序列如SEQ ID No.4所示。
6.一种用于扩增BraA08g039212E基因全长的引物，其特征在于，包括如SEQ ID No.5所示的正向引物和SEQ ID No.6所示的反向引物。
7.一种用于构建BraA08g039212E基因遗传转化载体的引物，其特征在于，包括如SEQID No.7所示的正向引物和SEQ ID No.8所示的反向引物。
8.一种用于BraA08g039212E基因阳性检测的引物，其特征在于，包括SEQ ID No.9所示的正向引物和SEQ ID No.10所示的反向引物。
9.一种用于检测BraA08g039212E基因RT-qPCR的引物，其特征在于，包括SEQ ID No.11所示的正向引物和SEQ ID No.12所示的反向引物。
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| CN116410282A (zh) * | 2023-03-17 | 2023-07-11 | 华中农业大学 | 来源欧洲芜菁ECD04的BraA03g008044E基因在根肿菌抗性改良中的应用 |
| EP4278891A1 (en) * | 2022-05-20 | 2023-11-22 | KWS SAAT SE & Co. KGaA | Clubroot resistance and markers in brassica |
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| CN116410282A (zh) * | 2023-03-17 | 2023-07-11 | 华中农业大学 | 来源欧洲芜菁ECD04的BraA03g008044E基因在根肿菌抗性改良中的应用 |
| CN116410282B (zh) * | 2023-03-17 | 2024-02-20 | 华中农业大学 | 来源欧洲芜菁ECD04的BraA03g008044E基因在根肿菌抗性改良中的应用 |
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