CN113512552B - 与芥菜型油菜多室性状相关的两室基因BjMc2和三室基因Bjmc2及其应用 - Google Patents
与芥菜型油菜多室性状相关的两室基因BjMc2和三室基因Bjmc2及其应用 Download PDFInfo
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Abstract
本发明公开了与芥菜型油菜多室性状相关的两室基因BjMc2和三室基因Bjmc2及其应用，属于油菜育种技术领域，其中两室基因BjMc2的启动子序列如SEQ ID NO.1所示，编码区序列如SEQ ID NO.2所示；所述两室基因BjMc2的等位基因即三室基因Bjmc2的启动子相较于两室基因BjMc2缺失了914bp。本发明通过图位克隆技术，首次从芥菜型油菜中成功克隆得到了芥菜型油菜的两室基因BjMc2和三室基因Bjmc2，可将其用于油菜的基因工程改造，根据需求获得不同表型的油菜。同时本发明还提供了一个可分别特异性扩增两室基因BjMc2和三室基因Bjmc2的分子标记CM2，可将其应用于芥菜型多室油菜育种中，用于辅助选择具备三室性状的芥菜型油菜，从而减少育种工作量，缩短育种年限，加快芥菜型多室油菜育种进程。
Description
技术领域
本发明属于油菜育种技术领域，具体涉及与芥菜型油菜多室性状相关的两室基因BjMc2和三室基因Bjmc2及其应用。
背景技术
油菜作为我国最重要的大宗油料作物，其菜籽油产量占我国食用植物油总产量的半壁江山。近年来，由于多室油菜的每角果粒数显著多于普通两室油菜，且已有几种自然变异多室油菜种质资源在不同的杂交组合后代中，多室单株产量均高于两室单株，所以越来越多研究者认为这一性状具有潜在的增产效益。
研究人员在对油菜种质资源的收集、整理，发现几种特殊角果性状的种质资源，通过仔细观察发现其角果具有多个角果皮和果室，并称其为多室油菜。通常，普通油菜的成熟角果只有2个角果皮，角果内1个“I”型假隔膜将角果心房分为2个果室，每个果室都有1排沿果皮内则融合与假隔膜的连接处着生的种子，称为两室油菜；两室角果是由2个心皮先天性融合成柱状结构的雌蕊发育而来，而多室角果是由多个心皮的雌蕊发育而成，雌蕊的心皮数目不同导致发育成熟角果内的假隔膜类型不同，常见有“Y”型、“II”型和“+”型等，这些不同类型的假隔膜通常将角果分为3～4个果室，每个果室都含有1排以上种子。因此，多室油菜产量增加的主要原因是其每角果粒数要比普通两室油菜的显著增多。
以芥菜型三室油菜J163-4和两室油菜J268-2、J248-2作为亲本，通过对正反交F1、F1'以及F2、BC1F1群体单株的表型统计，证实J163-4的三室性状受2 对独立遗传的隐性核基因控制，并命名为Bjmc1和Bjmc2，其中控制三室性状的其中一个基因Bjmc1已被成功克隆。现有技术虽然已将另一个基因Bjmc2精细定位到一段共线于白菜A7 scaffold000019的946～1014kb区间之间，约68kb物理距离，然而现有技术并未成功克隆到控制三室性状的另一个基因Bjmc2及其基因序列。
发明内容
本发明针对现有技术的空白，提供了一种与芥菜型油菜多室性状相关的两室基因BjMc2和三室基因Bjmc2及其应用。
本发明的目的之一在于提供一种与芥菜型油菜多室性状相关的两室基因 BjMc2，所述两室基因BjMc2的启动子的核苷酸序列如SEQ ID NO.1所示，所述两室基因BjMc2的编码区的核苷酸序列如SEQ ID NO.2所示。
本发明的目的之二在于提供上述两室基因BjMc2编码的蛋白，所述蛋白的氨基酸序列如SEQ ID NO.3所示。
本发明的目的之三在于提供一种包含上述两室基因BjMc2的载体或遗传工程菌。
本发明的目的之四在于提供了所述两室基因BjMc2和/或包含上述两室基因BjMc2的载体或遗传工程菌在制备两室油菜中的应用。
本发明的目的之五在于提供了所述两室基因BjMc2的等位基因，所述等位基因为三室基因Bjmc2，所述三室基因Bjmc2的启动子核苷酸序列为：在如SEQ ID NO.1所示的两室基因BjMc2的启动子核苷酸序列中缺失如SEQ ID NO.5所示的914bp的核苷酸序列。
进一步地，所述三室基因Bjmc2的启动子核苷酸序列如SEQ ID NO.4所示。
进一步地，所述三室基因Bjmc2的编码区的核苷酸序列与两室基因BjMc2 的编码区的核苷酸序列相同，如SEQ ID NO.2所示。
本发明的目的之六在于提供了一种包含上述三室基因Bjmc2的载体或遗传工程菌。
本发明的目的之七在于提供了所述三室基因Bjmc2和/或包含上述三室基因Bjmc2的载体或遗传工程菌在制备多室油菜中的应用。
本发明的目的之八在于提供了一种用于区分所述两室基因BjMc2及其等位基因三室基因Bjmc2的分子标记，所述分子标记的核苷酸序列如SEQ ID NO.6-8 所示。
本发明的目的之九在于提供了所述分子标记在油菜多室品种选育中的应用。
本发明的目的之十在于提供了一种油菜多室品种的选育方法，所述方法包括：提取待测样品的总DNA，采用上述分子标记进行PCR扩增，扩增产物进行电泳检测：若电泳条带仅出现一条1299bp的特异性条带，则待测样品为仅包含两室基因BjMc2的纯合型两室油菜；若电泳条带只出现一条700bp的特异性条带，则待测样品为仅包含三室基因Bjmc2的纯合型三室油菜；若电泳条带同时包含一条1299bp和一条700bp的电泳条带，则待测样品为杂合型两室油菜。
与现有技术相比，本发明的有益效果是：本发明通过图位克隆技术，首次从芥菜型油菜中成功克隆得到了芥菜型油菜的两室基因BjMc2和三室基因 Bjmc2，可利用所述两室基因和三室基因对油菜进行基因工程改造，以根据需求获得不同表型的油菜，其中三室油菜的产量相对较高。同时本发明还提供了一个可分别特异性扩增两室基因BjMc2和三室基因Bjmc2的分子标记CM2，可将其应用于芥菜型多室油菜育种中，用于辅助选择具备三室性状的芥菜型油菜，从而克服了传统育种中依靠表型进行选择的缺陷，减少育种工作量，缩短育种年限，加快了芥菜型多室油菜育种的进程。
附图说明
图1为本发明实施例1中分离克隆候选基因区段的物理图谱，其中A为将 BjMc2基因登陆到芥菜型油菜A7染色体标记ZX17和BACsr96之间的35kb物理区间内，括号内的数字表示相应标记的交换单株数，下面的灰色物理图谱是标记ZX17和BACsr96在BAC测序序列contig1对应的物理区间；B为BjMc2 位点的35kb物理区间在芥菜型油菜参考基因组中包含6个注释基因，灰色箭头表示BjMc2的候选基因；C为候选基因的基因结构以及在双亲中的多态性，深灰色柱代表外显子，浅灰色柱代表启动子和3′UTR区，灰色虚线表示三室基因Bjmc2启动子区缺失914bp序列；
图2为本发明实施例2中利用分子标记CM2在芥菜型油菜不同群体中单株基因组DNA的扩增结果，其中A为以J163-4和J268-1为双亲构建的F2群体中单株检测结果；B为以J163-4和J268-2为双亲构建的BjMc2位点NIL-BC7F1群体中单株检测结果，图中M为DNAMarker；
图3为本发明实施例3中分2段在芥菜型油菜J163-4和J268-1的基因组DNA中分别扩增Bjmc2和BjMc2的检测结果，其中A为片段1(包含启动子) 扩增结果，左边为三室基因Bjmc2(3060bp)，右边为两室基因BjMc2(4014bp)，； B为片段2扩增结果，左边为三室基因Bjmc2，右边为两室基因BjMc2，片段大小均为3936bp，图中M为DNA Marker；
图4为本发明实施例3中互补载体的构建图，其中A为载体pMc2::Mc2；B 为载体pmc2::Mc2；
图5为本发明实施例3中阳性植株的PCR鉴定结果图，图中M为DNA Marker，DNAMarker左边为pmc2::Mc2转基因植株检测结果，DNA Marker右边为pMc2::Mc2转基因植株检测结果；
图6为本发明实施例3中qRT-PCR分析T1代转基因株系不同组织中BjMc2 的表达情况；
图7为本发明实施例3中BjMc2和Bjmc2基因在芥菜型油菜中互补验证的表型结果，其中图A为A-Bi株系植株的花发育第12期雌蕊横切、成熟果枝和角果表型；图B为A-Tri株系植株的花发育第12期雌蕊横切、成熟果枝和角果表型；图C为T1代pMc2::Mc2转基因植株的花发育第12期雌蕊横切、成熟果枝和角果表型；图D为T1代pmc2::Mc2转基因植株的花发育第12期雌蕊横切、成熟果枝和角果表型。
具体实施方式
下面将结合本发明中的实施例，对本发明的技术方案进行清楚、完整地描述，显然，所描述的实施例仅仅是本发明一部分实施例，而不是全部的实施例。基于本发明中的实施例，本领域普通技术人员在没有做出创造性劳动条件下所获得的所有其它实施例，都属于本发明保护的范围。
实施例1芥菜型油菜两室基因BjMc2和三室基因Bjmc2序列的获得
1、精细定位区间的染色体登陆
现有技术通过精细定位已经把BjMc2基因限定在分子标记ZX17和BACsr96 之间，遗传距离分别为0.048cM、0.34cM，并共线于白菜A7 scaffold000019的 946～1014kb之间，约68kb物理距离。随后，将挑到的两个BAC阳性单克隆 (002-O-21和009-M-2)的全长测序序列组装完成，一共获得3个contigs(分别命名为contige1、2、3)。经比对发现，标记ZX17和BACsr96在contige1的物理位置分别为89.5kb和54.6kb，覆盖34.9kb的物理区间。当芥菜型油菜的全基因组序列测序完成后，标记ZX17和BACsr96标记分别在芥菜型油菜参考基因组A7的物理位置为32935kb和32900kb，覆盖35kb的物理区间，说明之前BjMc2位点的定位结果是可靠的(如1-A所示，其中括号内的数字表示相应标记的交换单株数，下面的灰色物理图谱是标记ZX17和BACsr96在BAC测序序列contig1对应的物理区间)。如图1-B所示，这段基因组区域包含6个有注释的候选基因(BjuA029482、BjuA029483、BjuA029484、BjuA029485、BjuA029486 和BjuA029487)，其中BjuA029487只包含部分CDS。通过对6个候选基因的序列进行分析，预测其中BjuA029486可能是BjMc2的候选基因，并进一步对其进行验证。
2、两室基因BjMc2的候选基因BjuA029486在双亲中序列比较分析
为了确定BjMc2位点的候选基因，基于contig1测序序列，设计一系列特异引物，如表1所示：
表1候选基因比较测序的引物核苷酸序列
注：a在三室亲本中扩增无条带；b在三室亲本中扩增条带大小为1739bp
分别以两室亲本268-1和三室亲本163-4的基因组DNA为模板，利用高保真PCR方法进行基因扩增。TA克隆测序结果表明，在两室亲本268-1和三室亲本163-4中该基因的编码区(3043bp)和3'UTR(1248bp)序列没有差异，而在起始密码子ATG的上游启动子区域有差异，相较两室亲本，三室亲本在起始密码子ATG的上游-2865bp到-3778bp(914-bp)大片段缺失(如图1-C所示，其中深灰色柱代表外显子，浅灰色柱代表启动子和3′UTR区，灰色虚线表示三室基因Bjmc2启动子区缺失914-bp序列)。由此，得到了芥菜型油菜两室基因 BjMc2的候选基因完整序列，其启动子核苷酸序列(起始密码子ATG的上游序列，下同)如SEQ IDNO.1所示，编码区核苷酸序列如SEQ ID NO.2所示，编码的氨基酸序列如SEQ ID NO.3所示。同时，还得到了该基因在三室亲本J163-4 中的等位基因完整序列，即三室基因Bjmc2，其启动子核苷酸序列为SEQ ID NO. 4所示，相较于两室基因BjMc2的启动子序列，三室基因Bjmc2的启动子序列缺失了如SEQ ID NO.5所示的914bp序列。
实施例2与芥菜型油菜三室性状相关的分子标记及其应用
1、与芥菜型油菜三室性状相关的分子标记CM2的开发
本发明根据已获得的BjMc2基因和Bjmc2基因序列信息，在Bjmc2启动子缺失位点附近开发共分离分子标记CM2(包含CM2-F1、CM2-F2和CM2-R三条引物)。PCR扩增芥菜型两室油菜J268-1和芥菜型三室油菜J163-4的基因组 DNA，在J268-1中分子标记CM2预期扩增出1299bp的片段，而在J163-4中 CM2预期扩增出700bp的片段。
本实施例所开发的CM2共分离分子标记的序列如下：
CM2-F1：TACTACTTCCGTTGCCTTTTCG
CM2-F2：AAAATACTGGTGTACATTGGAA
CM2-R：AAATCCATACACTTGGGGTT
2、分子标记CM2在不同群体中的应用
采用本领域常规方法提取以J163-4和J268-2为双亲构建的BjMc2位点NIL -BC7F1群体和以J163-4和J268-1为双亲构建的F2及BC1F1群体的单株叶片总D NA，利用上述开发的分子标记CM2进行PCR扩增。
PCR的反应体系如下：1×PCR buffer，1.35mM MgCl2，0.08mM dNTP s，1.0U DNA聚合酶(均购自MBI Fermentas,Lithuanin公司)，100ng DNA，两条正向引物CM2-F1和CM2-F2各加0.5uM，反向引物CM2-R加1uM，ddH2 O补充至终体积20ul。
热循环参数为：94℃5min；94℃30sec，55℃30sec，72℃90sec，35个循环；72℃10min，1个循环。
扩增产物在水平电泳槽上1.0％琼脂糖凝胶电泳检测。检测结果如图2所示，其中图2-A为F2群体中单株检测结果，共检测出3种带型：同时包含1299bp 和700bp上下2条条带的BjMc2位点为杂合型，其基因型为Mc2/mc2；只含有 1条上条带(1299bp)的BjMc2位点为显性纯合型，其基因型为Mc2/Mc2；只含有1条下条带(700bp)的BjMc2位点为隐性纯合型，其基因型为mc2/mc2。图2-B为BjMc2位点NIL-BC7F1群体中单株检测结果，共检测出2种带型，其中同时包含上下2条条带的BjMc2位点为杂合型，其基因型为mc1mc1Mc2mc2 (田间表型为两室)，只含有1条下条带的BjMc2位点为隐性纯合型，其基因型为mc1mc1mc2mc2(田间表型为三室)。上述结果说明，本发明开发的CM2 分子标记可以作为芥菜型油菜三室性状的共显性分子标记。因此，CM2结合专利CN201610979823.2中公开的BjMc1位点的M1-1分子标记，可以在不同杂交组合的低世代(例如F2)就能对不同基因型的两室和三室单株进行区分，从而有效缩短了多室油菜的选育年限，提高选择效率。
实施例3两室基因BjMc2和三室基因Bjmc2在芥菜型油菜中的互补验证
1、遗传互补载体的构建
为了验证该候选基因的功能，根据比较测序得到的序列，设计特异引物在三室亲本J163-4和两室材料J268-1中分别扩增出该候选基因的基因组序列。本发明所用到的互补载体包括：pMc2::Mc2和pmc2::Mc2。其中，pMc2::Mc2的外源片段是通过两对引物G2EinfusionF/G2&G3MedinfuR(扩增条带大小为4014 bp)和G2&G3MedinfuF/G2&G3KinfusionR(扩增条带大小为3936bp)分2段扩增J268-1的基因组DNA，引物序列为：
G2EinfusionF： CTATGACATGATTACGAATTCGAATTGAAACCACTTTTTTTACACC
G2&G3MedinfuR： AGTGAGCTGAGGGGGAAGTGGAGTGAACCCAGTCGTGGAG
G2&G3MedinfuF： CACTTCCCCCTCAGCTCACTGTTCTTTCTCCGGCGTTTCA
G2&G3KinfusionR： TCTAGAGGATCCCCGGGTACCCAACATGGCTTTTATTTATTAATCTCCAC
PCR产物经过1.0％TAE琼脂糖凝胶电泳，检测结果如图3所示，其中3-A 为片段1(包含启动子)扩增结果，图3-B为片段2的扩增结果。将目的条带进行挖胶回收(琼脂糖凝胶DNA纯化回收试剂盒，Cat.No.DP219，天根)，再通过同源重组(ClonExpressⅡOne StepCloning Kit,Vazyme)的方法与经过EcoR Ⅰ和KpnⅠ双酶切处理的pCAMBIA 2300空载体连接，pMc2::Mc2载体的构建图如图4-A所示。插入片段为BjMc2全长7930bp的基因组序列，其中包括gDNA 序列3043bp、上游启动子序列3818bp和下游5'UTR序列1069bp。获得的阳性克隆经过PCR扩增、测序与比较测序序列比对无突变后，用于后续遗传转化。
同样，pmc2::Mc2的外源片段也是通过两对引物 G3EinfusionF/G2&G3MedinfuR(扩增条带大小为3060bp)和 G2&G3MedinfuF/G2&G3KinfusionR(扩增条带大小为3936bp)分2段扩增J163-4 的基因组DNA，引物序列为：
G3EinfusionF： CTATGACATGATTACGAATTCTGTACATTGGAAATGCAACTCCAGC
另外3条引物序列同上，PCR产物经过1.0％TAE琼脂糖凝胶电泳，检测结果如图3所示。将目的条带进行挖胶回收，扩增片段同样通过同源重组的方法与经过EcoRⅠ和KpnⅠ双酶切处理的pCAMBIA2300空载体连接，pmc2::Mc2 载体的构建图如图4-B所示。插入片段为Bjmc2全长6976bp的基因组序列，其中包括gDNA序列3043bp、上游启动子序列2864bp和下游5'UTR序列1069bp。阳性克隆经PCR扩增、测序与比较测序序列比对无突变后，用于后续遗传转化。
2、农杆菌介导的遗传转化
选择构建的BjMc2位点NIL-BC7F1群体中两室单株(基因型为 mc1mc1Mc2mc2)进行自交，接着在BC7F2株系中继续选择两室单株(基因型为 mc1mc1Mc2Mc2)和三室单株(基因型为mc1mc1mc2mc2)进行多代自交留种，并命名为A-Bi(BC7F2-Bi)和A-Tri(BC7F2-Tri)株系。正确的重组质粒通过常规冻融法导入农杆菌菌株GV3101，然后通过农杆菌介导的油菜下胚轴侵染法转入芥菜型油菜，受体材料均为A-Tri三室自交株系植株，具体操作方法为：
种子灭菌用75％酒精浸泡3min后，放入无菌水中冲洗1min。然后放入0.1％升汞中浸泡10min后，无菌水冲洗6-7次。将灭菌后的种子播种到种子萌发培养基上(所述的种子萌发培养基为：MS基本培养基+7g/L琼脂粉，pH值为 5.9)。于25℃按培养7天。在超净工作台内将获得的试管幼苗下胚轴切成1cm 左右的小段外植体，接种至含有农杆菌(悬浮过夜至对数生长期)的MS液体培养基(pH值为5.9)侵染10-15min后，吸干液体，于25℃黑暗条件下在共培养基上(所述的共培养基包括：MS培养基+30g/L蔗糖+18g/L甘露醇+1 mg/L 2,4-D+0.3mg/L Kinetin+100um/L乙酰丁香酮+6g/L琼脂粉，pH 值为5.9)共培养36h。将外植体转移到愈伤诱导培养基上(所述的愈伤诱导培养基包括：MS培养基+30g/L蔗糖+18g/L甘露醇+1mg/L 2,4-D+0.3mg/L Kinetin+300mg/L特美汀+50mg/L卡那霉素+0.03μM/L[Ag(NO3)2]3-+6g/L 琼脂粉，pH值为5.9)，25℃光照培养12-14d。将外植体转入不定芽诱导培养基上(所述的不定芽诱导培养基包括：MS培养基+30g/L蔗糖+1.5g/L TDZ+0.1 g/L NAA+300mg/L特美汀+50mg/L卡那霉素+5.0mg/L硝酸银+7g/L琼脂粉，pH值为5.9)，25℃光照培养12-14d。获得的带有不定芽点的愈伤组织转移到不定芽生长培养基中，进行生长培养。还未生长出不定芽点的外植体，则继续放到不定芽诱导培养基中进行诱导培养，待长出不定芽点后，转移到不定芽生长培养基中(所述的不定芽生长培养基包括：MS培养基+10g/L葡萄糖+ 0.25g/L木糖+0.6g/L MES+2.0mg/L反式-Zeatin+0.1mg/L IAA +300mg/L 特美汀+25mg/L卡那霉素+6g/L琼脂粉，pH值为5.9)。每15-20d继代一次，直至生长成幼苗。当幼苗长至2-3cm高时，将幼苗转入生根培养基中培养(所述生根培养基包括：MS培养基+10g/L蔗糖+10g/L琼脂粉，pH值为5.9)。对已生根的小苗进行炼苗，按常规方法移栽到温室里，进行土培生长。
3、阳性转化单株的分子及表型鉴定
从获得的阳性植株上按照常规方法提取叶片总DNA，用片段中间引物 CLV1-7.2F和通用引物M13-47(扩增条带大小约1.2kb)进行PCR鉴定，鉴定所用的引物序列如下：
CLV1-7.2F：ACTAACTTGATCGCCTTCTGA
M13-47：CGCCAGGGTTTTCCCAGTCACGAC
PCR扩增的反应体系是：30ng模板DNA，各35ng正向和反向特异引物(如上所示)，1×PCR缓冲液，1U Taq酶，0.15mmol/L dNTPs和2.0mmol/L Mg2+，加ddH2O到总体积为15μL。PCR循环参数为94℃(30s)—55℃(30s)—72℃ (90s)，共35个循环。
扩增结束后用1.0％琼脂凝胶检测扩增产物。检测结果如图5所示，图中M 为DNAMarker，片段大小从上至下依次为3000bp、2000bp、1000bp、750bp 和500bp，DNA Marker左边的条带为pmc2::Mc2转基因植株的检测结果，DNA Marker右边的条带为pMc2::Mc2转基因植株的检测结果，片段大小均约为1.2 kb，该结果说明阳性植株中包含候选基因。
对所有T0代阳性植株的表型进行鉴定，其中pMc2::Mc2载体18株有表型， pmc2::Mc2载体31株有表型。继续种植有表型的单株T1代，取苗期幼叶、花序和雌蕊子房提取总RNA(百泰克，RP3202，http://cn.bioteke.cn/)。每个样品吸取2毫克的总RNA进行反转录(Fermentas，RevertAid First Strand cDNA Synthes K1622)。将合成的第一链cDNA稀释20倍用作qRT-PCR的模板，在qRT-PCR 专用96孔板中先加入上下游引物(5.0μM)等比例混合液3.2μL，模板2.0μL 和
Green Realtime PCR Master Mix(TOYOBO)10μL，最后补ddH20 至20μL，每对引物3个技术重复，每个组织3次生物学重复，Actin2内参基因作为对照。qRT-PCR引物序列为：
RTA7Mc2F：TCAAGAATCTCCTCAACGGTAC
RTA7Mc2F：AGAAGAAGTTATCGGTGAGCTC
Actin2-F：GGAGCTGAGAGATTCCGTTG
Actin2-R：GAACCACCACTGAGGACGAT
反应程序如下：95℃预变性60s；95℃变性15s，55℃退火15s，72℃延伸 30s，39个cycles；熔解曲线程序：95℃10s，95℃-65℃5s，-0.5℃per cycle，收集数据。qRT-PCR在Bio-Red CFX96 TouchTM荧光定量PCR仪上进行，依照 2-ΔΔCt方法分析基因的相对表达量。
qRT-PCR分析结果如图6所示(Student’s t检验显著性差异，NS：不显著(P ≥0.05)；***：极显著(P<0.001))，其中pMc2::Mc2转基因植株的苗期幼叶、花序和雌蕊子房中BjMc2转录本的相对表达量相较于受体材料均有显著性提高。同样，pmc2::Mc2转基因植株的苗期幼叶和花序中BjMc2转录本的相对表达量相较于受体材料也有显著性提高，且和pMc2::Mc2转基因植株间没有显著性差异。然而，pmc2::Mc2转基因植株的雌蕊子房中BjMc2转录本的相对表达量相较于受体材料没有显著性差异，但显著性低于pMc2::Mc2转基因植株。这些结果表明Bjmc2启动子缺失914-bp不影响BjMc2基因在苗期幼叶和花序中的表达，但会影响BjMc2基因在雌蕊子房中表达。
同时取有表型T1代株系的花蕾做石蜡切片，不同植株的的花发育第12期雌蕊横切、成熟果枝和角果表型的切片结果如图7所示，其中图7-A为A-Bi株系植株；图7-B为A-Tri株系植株；图7-C为T1代pMc2::Mc2转基因植株；图7-D 为T1代pmc2::Mc2转基因植株。
根据图7-C，其中pMc2::Mc2转基因植株的雌蕊可以将三室株系A-Tri恢复成由两个心皮和一个“I”型假隔膜组成的两室表型，和图7-A中BC7F2-Bi(A-Bi) 两室株系的雌蕊表型类似。而图7-D中pmc2::Mc2转基因植株的雌蕊不可以恢复为两室表型，其雌蕊由4个心皮和1个“II”型假隔膜或3个心皮和1个“Y”型假隔膜组成的三室表型，和图7-B中BC7F2-Tri(A-Tri)三室株系的雌蕊表型类似。在成熟果枝和角果也能看出pMc2::Mc2转基因植株主要是两室角果组成，而pmc2::Mc2转基因植株仍然主要是3室角果组成，表明pMc2::Mc2转基因植株可恢复成两室角果表型，而pmc2::Mc2转基因植株不能恢复两室角果表型。
综上所述，本发明成功筛选克隆得到了两室基因BjMc2，其启动子核苷酸序列如SEQ ID NO.1所示，编码区的核苷酸序列如SEQ ID NO.2所示，编码的氨基酸序列如序列表SEQ ID NO.3所示；该两室基因的启动子突变等位基因即三室基因Bjmc2的启动子核苷酸序列如SEQ ID NO.4所示，三室基因Bjmc2启动子相较于两室基因BjMc2启动子缺失的914bp核苷酸序列如SEQ ID NO.5所示。将该两室基因BjMc2通过基因工程改造技术导入至以J163-4和J268-2为双亲构建的BjMc2位点NIL-BC7F2三室自交株系(A-Tri)植株中，可恢复两室角果表型，证明了这段914bp启动子核苷酸序列是两室基因BjMc2恢复两室角果表型所必须的。
以上所述，仅为本发明较佳的具体实施方式，但本发明的保护范围并不局限于此，任何熟悉本技术领域的技术人员在本发明揭露的技术范围内，可轻易想到的变化或替换，都应涵盖在本发明的保护范围之内。
序列表
<110> 华中农业大学
<120> 与芥菜型油菜多室性状相关的两室基因BjMc2和三室基因Bjmc2及其应用
<160> 8
<170> SIPOSequenceListing 1.0
<210> 1
<211> 3838
<212> DNA
<213> 芥菜型油菜(Brassica juncea)
<400> 1
atgtttgagt ttctggcaaa gaattgaaac cacttttttt acaccaaaaa aaaatactgg 60
tgttgcctct tcatgatgta aaataaatca aaattatttt tagatatctg ccgtgcaatg 120
gaacaaaaag tgaaaggtga ttcaagacct caagggtcat ggattgtgag aagtttcttc 180
gtccaatatc attgcatgtt ttgaatagat ttgggactaa agctgccaat agacagagag 240
aattcggtag tgactgaagc aaaagctcaa aatgccatta ggcaatcccc cgtttccaag 300
acagatagtg taagacacac ttgtgacaca actttgtctt cgtcttcatc atcatcttct 360
actactacta cttccgttgc cttttcgttt aattattaac ttgattgaca agacaatgac 420
ctatatttct tatatcatca ttgctctcaa tttgtttctt ttaataattg cttttgaaag 480
gtttatccca aaatgattag tgcgttgatt atatgtgaaa caaacaagct gatcacgaat 540
gaatcatgta tcattttcta gatatttact caaaaacgtt tactagattt tgtagtaaat 600
tttgaatgac aatacgccca ttttatttaa aatttgtatg ttttattttt ttgtatttaa 660
gagtttaata ttttatattt taactcatgt agtttaactg aaaactaaaa tatgtgatgg 720
gattatttta ttagaaaaag agaaataact aataaaataa agtatgtatg tatataattt 780
caaaaacgta tgatacaatt taaaacatca atctttcaaa tacataagaa atattatata 840
aacttgattt ttcttagccg ttttagattt gttctcccct caagttgcat atgattttca 900
aattgcttga gacaaacttt ttaaacagca attaactttc aagcaccaag tgaaaagtag 960
catatgatgc aaattgtaca ttggaaatgc aactccagct tctacagttc tactacctaa 1020
tcatgtttct ctttcttata attagtttca caaatttatt aattagcatt catctagaag 1080
ctagcctaaa gttcattcag acattcacca taaatcatta atgataaatg tcttacatta 1140
tgctcctaag tgtattgtca tgttatatca aaaaatgccc aaagttcata ttcctctcta 1200
cataaagaaa aatttctaat tactaagaaa agctcctttt ctgctgaaga gaacataaac 1260
ctttattccc caaagctcac aagtttgggc attttcagcg aatcagagaa aatatgcttc 1320
tcttcttcat cacatattga catctccctt ggttcaagac agtaccggtg gctctgattg 1380
tgtaaaccgt ccaatccagt tgaaccggtc aacagtatta attaaccaaa aataacacgt 1440
gaaagcatat atttatgttg ttaggtatgt atagcgttta ctttcttaaa tttaatgcat 1500
caagatcaaa tccaaaagag aatacacaac atcatataaa atgagaaatt cgctgcgcca 1560
tttaggaaag ggcctttata ccaatatgcc gcatatatgt agaacattgg tccccaagtg 1620
tatatgtgtg tgtgtgtcta tgtcaacccc aagtgtatgg atttctcaac aaatcaaagg 1680
ccctttgttg gtaaagctcc tcttgatatt taatcgtata aaatattgtt agcggggtga 1740
agatatattc gccaaaaacg gtagtgcttg tataaccgga aaccaataat ttacaaatag 1800
cattagcttt tatgttacaa atgttgagag aatgaataca cggaaaccaa taatttacaa 1860
atagccttag cttttatgtt acaaatgttg agagaataca ttttaattct ggcagaatcg 1920
attctgcata aaacttgtat aaactataac atggcttttg taactaaaaa ataaatattg 1980
gaaataaatt ggatatctcg cggatcaaac tcgtaaagtt cttaaaatct gtaaaagtta 2040
tgaaaaacaa atggaaacac ttgaaaatgt gtatcagcga tagtataata ttaagtaaga 2100
cacacatttg ctattcgatt tctctctagc aaaagctttg ttatgtagtt gtggacataa 2160
gcatacgatc agcgatatat atatttcgtt ctttgttttg ttatcgtcac acgattagca 2220
atcgcgatca aatagttgtt gttcatcatt tcatttcagt gtttggccat tgatagacaa 2280
gatatgtcaa agtcgctaga actttcagag tttctaaact aacgacagat tatttgttcc 2340
aataaagatc taagtgtgtg atattcgagc gtttccacta tgattttatt tatgattagt 2400
cattgttaac aactaaaact gtaagactat ccgaattttt tcacatattc cgaaattaat 2460
tcgagcttat tatcaaatca acgattttca taaggtatag tttaaaaaaa aaaattacaa 2520
gagagaagat aagggtaaag tcagagagat ctgcaaaacc aaaggtaaat aacacacaac 2580
gtggggaccc tgaagcacag cagaagaagg tctacaagaa gagtgagtgt cgacaaagac 2640
aagaaaagga gcgttgacta ttagattgaa gccaaaatag aaggggcaga tttgtctctt 2700
ttggaaagga cacagacaat cttttatacg ggccatttaa taactaggcc ctacttaata 2760
agcccattaa ctcccttctt tgtcctttta agtttttaaa atatatattc cattttcctt 2820
atcacgtgac aatgagaagg aaacgtcgtc ggattaaact ggttgagatg acaagacggc 2880
gataagaaga aagaaaccac agcagctata ggcagggaac gtaaagttaa gtctgtctat 2940
catctattaa cggtcagttc agatcataga cattgtaata gcgttttctt tgtcccgtta 3000
cattttaaac gccgtgaaga tatccattta ttttgatata tatagccaaa aagataaact 3060
ttctcgattt acatcgtgtc agaagattta acagaggaaa gaagacgatc acgacgtttc 3120
ccggttttcc atgtcttgtg tagtggcggc tactaaccaa ataactttgt ccgtaaaaac 3180
tgtaaaaaag actatggtaa aagtttcttt agtatgttta ctctgcacat atagcagtgg 3240
attatgggtt tgcattacaa tttaagcaca aatcgtttag ttacccacaa aaatatcata 3300
gattctctaa aaatcacatg attaaatgac atttacaaaa ttttgtgatt attactcgga 3360
ataattcaaa gaaaaatatt atttgagcac attaacaaaa aatggaagtt tgaagtatgt 3420
gacgtataaa gccaatgaaa gactgacact tcattgacca caagtcgtag ttcgtatcta 3480
caagctggat gtccgtttac ccgtttatat ccgtatgata atttcataaa tctaactcga 3540
taagtaaaca tcacaaaaat tacaagatac attggagaaa aaaaaggttt ttattcggcg 3600
cattatttct agtggtatat ttattctcat tcaaagtcat ataaaaaaag attttaaagt 3660
aaaaatattc tctctgaaag tctagcagta ttttagaaca gtaccactac gattctcttc 3720
ttcatcctct tcataatatg ccattgtggt gattgttcat atatctatat gtacctccta 3780
tcgcataaga ctcacgctaa cttcttattc tctctcaaaa gacagctttt aaataaaa 3838
<210> 2
<211> 3043
<212> DNA
<213> 芥菜型油菜(Brassica juncea)
<400> 2
atggagatga gacttctgaa aactcacctt ctgtttctcc atcttcatta cgttatctcg 60
atttcgcttc tatgtttctc actatgcctc gcttccactg acatggacca tctcctcaac 120
ctcaaatcct ccatggtcgg ccccaacggc cacggcctcc acgactgggt tcactccact 180
tccccctcag ctcactgttc tttctccggc gtttcatgcg acggcgacgc tcgtgtcatc 240
tccctcaacg tctctttcac tcctctcttc ggaaccatct cgccagagat tgggatgctg 300
aaccgtcttg tgaatctcac gttagctgct aataacttct ccggtatgtt gccgctggag 360
atgaagagtc tcacttctct aaaggttctc aacatctcca acaacgtgaa cctcaacggg 420
accttccccg gagagattct cactcccatg gtggacctcg aagtcctcga cgcgtacaac 480
aacaacttca caggcccatt acccccggag atccccgggc tcaagaagct gagacacctc 540
tctctcggag gaaacttctt aaccggagag atcccagaga gttacggaga tatccaaagc 600
ttggagtatc ttggcctcaa cggagccgga ctctccggtg aatctccggc gttcttgtct 660
cgcctcaaga atcttaaaga aatgtacgtc ggctacttca acagctacac cggcggcgtt 720
ccgccggagt tcggtgaatt gacaaaccta gaggtcctcg acatggcgag ctgtactctc 780
acaggagaga ttccgacaac actaagtaat ctaaaacatt tgcacacttt gtttctccac 840
atcaacaact taaccggaaa catcccacca gaactctccg gtttaatcag tttaaaatct 900
ctagacctct caataaacca gctaaccgga gagattcctc agagcttcat ctccctagga 960
aacatcactc tcatcaacct cttcagaaac aatctccacg ggccgatacc ggacttcatc 1020
ggagacatgc cgaacctcca agtcctccag gtgtgggaga acaacttcac gctagagcta 1080
ccggcgaatc tcggccggaa cgggaatctg aaaaagctcg acgtctctga taaccatctt 1140
accggactca tccccatgga tttgtgcaga ggcgggaagc tggagacgct cgtgctctcc 1200
aacaacttct tcttcggctc gatccctgag aagctaggtc aatgcaaatc gctaaacaag 1260
atcagaatcg tcaagaatct cctcaacggt acggttccgg agggattatt caatctaccg 1320
ctcgtaacga tcatcgagct caccgataac ttcttctccg gggagcttcc gggggagatg 1380
tcaggcgacg ttctcgatca tatctactta tctaacaatt ggtttaccgg tttaatcccc 1440
ccggctatcg gtaattttaa aaatctacaa gatttattct tagaccggaa ccggtttagc 1500
gggaatatcc cgagggaagt tttcgagtta aagcatctaa cgaagatcaa cacgagtgct 1560
aacaacctaa ccggcgacat ccctgactcg atctctcgct gcacttcctt aatctccgtc 1620
gatctcagcc gtaaccgaat cggcggagat attcctaaag acatccacga cgtgattaac 1680
ttaggaactc taaatctctc cgggaatcaa ctcaccggct cgatcccgat cggaatcggg 1740
aagatgacga gcttaaccac tctcgatctc tccttcaacg acctctccgg gagagtccca 1800
ctcggcggcc agttcctagt cttcaacgac acttccttcg ccggaaaccc ttacctctgc 1860
ctccctcacc acgtctcgtg ccttacgcgt ccggaacaaa cctccgatcg tatccacacg 1920
gctctcttct ctccgtcgag gatcgttatc acgatcgtcg cagcgatcac ggcgttgatc 1980
cttatcagcg tcgcgattcg tcagatgaac aagaagaagc acgagagatc tctctcctgg 2040
aagctaaccg ccttccaaag actcgatttc aaagcggaag acgtcctcga gtgtctccag 2100
gaagagaaca taatcggcaa aggcggagcg gggatcgtct accgcggatc catgccgaac 2160
aacgtcgacg tcgcgatcaa acgccttgta ggacgcggaa cagggaggag cgatcacgga 2220
ttcacggcgg agatacagac gctagggaga atccgccacc gtcatatagt gagactcctc 2280
ggatacgtgg cgaacaagga cacgaacctg cttctctacg agtacatgcc taacgggagc 2340
ctcggggagc ttttgcacgg atctaaagga ggtcatcttc agtgggagac gaggcacaga 2400
gtagccgtgg aagcggcgaa aggactgtgt tatcttcatc atgactgttc gccgttgatc 2460
ttgcatagag acgttaagtc caataacatt ttactggact ctgattttga ggcccatgtt 2520
gctgattttg ggcttgctaa gttcttagtg gacggtgctg cttctgagtg tatgtcttcg 2580
atagctggct cctatggata catcgctcca ggttagttta aacatgtttt aaataacaaa 2640
taatatgtat aaaactaact attgtttgtt ttggttttga attttgatag agtatgctta 2700
cactctcaaa gtggatgaga agagtgatgt gtatagtttc ggagtggtgt tattggagct 2760
gatagctggg aagaaaccag ttggtgagtt tggggaagga gtggatatag tgaggtgggt 2820
gaggaacacg gagggtgaga tacctcagcc gtcggatgca gctactgttg tggcgatcgt 2880
tgaccagagg ttgactggtt acccgttgac tagtgtgatt cacgtgttca agatagcgat 2940
gatgtgtgtg gaggatgagg cagcgacaag gccgacgatg agggaagttg tgcacatgct 3000
cactaaccct cccaagtccg tcactaactt gatcgccttc tga 3043
<210> 3
<211> 987
<212> PRT
<213> 芥菜型油菜(Brassica juncea)
<400> 3
Met Glu Met Arg Leu Leu Lys Thr His Leu Leu Phe Leu His Leu His
1 5 10 15
Tyr Val Ile Ser Ile Ser Leu Leu Cys Phe Ser Leu Cys Leu Ala Ser
20 25 30
Thr Asp Met Asp His Leu Leu Asn Leu Lys Ser Ser Met Val Gly Pro
35 40 45
Asn Gly His Gly Leu His Asp Trp Val His Ser Thr Ser Pro Ser Ala
50 55 60
His Cys Ser Phe Ser Gly Val Ser Cys Asp Gly Asp Ala Arg Val Ile
65 70 75 80
Ser Leu Asn Val Ser Phe Thr Pro Leu Phe Gly Thr Ile Ser Pro Glu
85 90 95
Ile Gly Met Leu Asn Arg Leu Val Asn Leu Thr Leu Ala Ala Asn Asn
100 105 110
Phe Ser Gly Met Leu Pro Leu Glu Met Lys Ser Leu Thr Ser Leu Lys
115 120 125
Val Leu Asn Ile Ser Asn Asn Val Asn Leu Asn Gly Thr Phe Pro Gly
130 135 140
Glu Ile Leu Thr Pro Met Val Asp Leu Glu Val Leu Asp Ala Tyr Asn
145 150 155 160
Asn Asn Phe Thr Gly Pro Leu Pro Pro Glu Ile Pro Gly Leu Lys Lys
165 170 175
Leu Arg His Leu Ser Leu Gly Gly Asn Phe Leu Thr Gly Glu Ile Pro
180 185 190
Glu Ser Tyr Gly Asp Ile Gln Ser Leu Glu Tyr Leu Gly Leu Asn Gly
195 200 205
Ala Gly Leu Ser Gly Glu Ser Pro Ala Phe Leu Ser Arg Leu Lys Asn
210 215 220
Leu Lys Glu Met Tyr Val Gly Tyr Phe Asn Ser Tyr Thr Gly Gly Val
225 230 235 240
Pro Pro Glu Phe Gly Glu Leu Thr Asn Leu Glu Val Leu Asp Met Ala
245 250 255
Ser Cys Thr Leu Thr Gly Glu Ile Pro Thr Thr Leu Ser Asn Leu Lys
260 265 270
His Leu His Thr Leu Phe Leu His Ile Asn Asn Leu Thr Gly Asn Ile
275 280 285
Pro Pro Glu Leu Ser Gly Leu Ile Ser Leu Lys Ser Leu Asp Leu Ser
290 295 300
Ile Asn Gln Leu Thr Gly Glu Ile Pro Gln Ser Phe Ile Ser Leu Gly
305 310 315 320
Asn Ile Thr Leu Ile Asn Leu Phe Arg Asn Asn Leu His Gly Pro Ile
325 330 335
Pro Asp Phe Ile Gly Asp Met Pro Asn Leu Gln Val Leu Gln Val Trp
340 345 350
Glu Asn Asn Phe Thr Leu Glu Leu Pro Ala Asn Leu Gly Arg Asn Gly
355 360 365
Asn Leu Lys Lys Leu Asp Val Ser Asp Asn His Leu Thr Gly Leu Ile
370 375 380
Pro Met Asp Leu Cys Arg Gly Gly Lys Leu Glu Thr Leu Val Leu Ser
385 390 395 400
Asn Asn Phe Phe Phe Gly Ser Ile Pro Glu Lys Leu Gly Gln Cys Lys
405 410 415
Ser Leu Asn Lys Ile Arg Ile Val Lys Asn Leu Leu Asn Gly Thr Val
420 425 430
Pro Glu Gly Leu Phe Asn Leu Pro Leu Val Thr Ile Ile Glu Leu Thr
435 440 445
Asp Asn Phe Phe Ser Gly Glu Leu Pro Gly Glu Met Ser Gly Asp Val
450 455 460
Leu Asp His Ile Tyr Leu Ser Asn Asn Trp Phe Thr Gly Leu Ile Pro
465 470 475 480
Pro Ala Ile Gly Asn Phe Lys Asn Leu Gln Asp Leu Phe Leu Asp Arg
485 490 495
Asn Arg Phe Ser Gly Asn Ile Pro Arg Glu Val Phe Glu Leu Lys His
500 505 510
Leu Thr Lys Ile Asn Thr Ser Ala Asn Asn Leu Thr Gly Asp Ile Pro
515 520 525
Asp Ser Ile Ser Arg Cys Thr Ser Leu Ile Ser Val Asp Leu Ser Arg
530 535 540
Asn Arg Ile Gly Gly Asp Ile Pro Lys Asp Ile His Asp Val Ile Asn
545 550 555 560
Leu Gly Thr Leu Asn Leu Ser Gly Asn Gln Leu Thr Gly Ser Ile Pro
565 570 575
Ile Gly Ile Gly Lys Met Thr Ser Leu Thr Thr Leu Asp Leu Ser Phe
580 585 590
Asn Asp Leu Ser Gly Arg Val Pro Leu Gly Gly Gln Phe Leu Val Phe
595 600 605
Asn Asp Thr Ser Phe Ala Gly Asn Pro Tyr Leu Cys Leu Pro His His
610 615 620
Val Ser Cys Leu Thr Arg Pro Glu Gln Thr Ser Asp Arg Ile His Thr
625 630 635 640
Ala Leu Phe Ser Pro Ser Arg Ile Val Ile Thr Ile Val Ala Ala Ile
645 650 655
Thr Ala Leu Ile Leu Ile Ser Val Ala Ile Arg Gln Met Asn Lys Lys
660 665 670
Lys His Glu Arg Ser Leu Ser Trp Lys Leu Thr Ala Phe Gln Arg Leu
675 680 685
Asp Phe Lys Ala Glu Asp Val Leu Glu Cys Leu Gln Glu Glu Asn Ile
690 695 700
Ile Gly Lys Gly Gly Ala Gly Ile Val Tyr Arg Gly Ser Met Pro Asn
705 710 715 720
Asn Val Asp Val Ala Ile Lys Arg Leu Val Gly Arg Gly Thr Gly Arg
725 730 735
Ser Asp His Gly Phe Thr Ala Glu Ile Gln Thr Leu Gly Arg Ile Arg
740 745 750
His Arg His Ile Val Arg Leu Leu Gly Tyr Val Ala Asn Lys Asp Thr
755 760 765
Asn Leu Leu Leu Tyr Glu Tyr Met Pro Asn Gly Ser Leu Gly Glu Leu
770 775 780
Leu His Gly Ser Lys Gly Gly His Leu Gln Trp Glu Thr Arg His Arg
785 790 795 800
Val Ala Val Glu Ala Ala Lys Gly Leu Cys Tyr Leu His His Asp Cys
805 810 815
Ser Pro Leu Ile Leu His Arg Asp Val Lys Ser Asn Asn Ile Leu Leu
820 825 830
Asp Ser Asp Phe Glu Ala His Val Ala Asp Phe Gly Leu Ala Lys Phe
835 840 845
Leu Val Asp Gly Ala Ala Ser Glu Cys Met Ser Ser Ile Ala Gly Ser
850 855 860
Tyr Gly Tyr Ile Ala Pro Glu Tyr Ala Tyr Thr Leu Lys Val Asp Glu
865 870 875 880
Lys Ser Asp Val Tyr Ser Phe Gly Val Val Leu Leu Glu Leu Ile Ala
885 890 895
Gly Lys Lys Pro Val Gly Glu Phe Gly Glu Gly Val Asp Ile Val Arg
900 905 910
Trp Val Arg Asn Thr Glu Gly Glu Ile Pro Gln Pro Ser Asp Ala Ala
915 920 925
Thr Val Val Ala Ile Val Asp Gln Arg Leu Thr Gly Tyr Pro Leu Thr
930 935 940
Ser Val Ile His Val Phe Lys Ile Ala Met Met Cys Val Glu Asp Glu
945 950 955 960
Ala Ala Thr Arg Pro Thr Met Arg Glu Val Val His Met Leu Thr Asn
965 970 975
Pro Pro Lys Ser Val Thr Asn Leu Ile Ala Phe
980 985
<210> 4
<211> 2924
<212> DNA
<213> 芥菜型油菜(Brassica juncea)
<400> 4
atgtttgagt ttctggcaaa gaattgaaac cacttttttt acaccaaaaa aaaatactgg 60
tgtacattgg aaatgcaact ccagcttcta cagttctact acctaatcat gtttctcttt 120
cttataatta gtttcacaaa tttattaatt agcattcatc tagaagctag cctaaagttc 180
attcagacat tcaccataaa tcattaatga taaatgtctt acattatgct cctaagtgta 240
ttgtcatgtt atatcaaaaa atgcccaaag ttcatattcc tctctacata aagaaaaatt 300
tctaattact aagaaaagct ccttttctgc tgaagagaac ataaaccttt attccccaaa 360
gctcacaagt ttgggcattt tcagcgaatc agagaaaata tgcttctctt cttcatcaca 420
tattgacatc tcccttggtt caagacagta ccggtggctc tgattgtgta aaccgtccaa 480
tccagttgaa ccggtcaaca gtattaatta accaaaaata acacgtgaaa gcatatattt 540
atgttgttag gtatgtatag cgtttacttt cttaaattta atgcatcaag atcaaatcca 600
aaagagaata cacaacatca tataaaatga gaaattcgct gcgccattta ggaaagggcc 660
tttataccaa tatgccgcat atatgtagaa cattggtccc caagtgtata tgtgtgtgtg 720
tgtctatgtc aaccccaagt gtatggattt ctcaacaaat caaaggccct ttgttggtaa 780
agctcctctt gatatttaat cgtataaaat attgttagcg gggtgaagat atattcgcca 840
aaaacggtag tgcttgtata accggaaacc aataatttac aaatagcatt agcttttatg 900
ttacaaatgt tgagagaatg aatacacgga aaccaataat ttacaaatag ccttagcttt 960
tatgttacaa atgttgagag aatacatttt aattctggca gaatcgattc tgcataaaac 1020
ttgtataaac tataacatgg cttttgtaac taaaaaataa atattggaaa taaattggat 1080
atctcgcgga tcaaactcgt aaagttctta aaatctgtaa aagttatgaa aaacaaatgg 1140
aaacacttga aaatgtgtat cagcgatagt ataatattaa gtaagacaca catttgctat 1200
tcgatttctc tctagcaaaa gctttgttat gtagttgtgg acataagcat acgatcagcg 1260
atatatatat ttcgttcttt gttttgttat cgtcacacga ttagcaatcg cgatcaaata 1320
gttgttgttc atcatttcat ttcagtgttt ggccattgat agacaagata tgtcaaagtc 1380
gctagaactt tcagagtttc taaactaacg acagattatt tgttccaata aagatctaag 1440
tgtgtgatat tcgagcgttt ccactatgat tttatttatg attagtcatt gttaacaact 1500
aaaactgtaa gactatccga attttttcac atattccgaa attaattcga gcttattatc 1560
aaatcaacga ttttcataag gtatagttta aaaaaaaaaa ttacaagaga gaagataagg 1620
gtaaagtcag agagatctgc aaaaccaaag gtaaataaca cacaacgtgg ggaccctgaa 1680
gcacagcaga agaaggtcta caagaagagt gagtgtcgac aaagacaaga aaaggagcgt 1740
tgactattag attgaagcca aaatagaagg ggcagatttg tctcttttgg aaaggacaca 1800
gacaatcttt tatacgggcc atttaataac taggccctac ttaataagcc cattaactcc 1860
cttctttgtc cttttaagtt tttaaaatat atattccatt ttccttatca cgtgacaatg 1920
agaaggaaac gtcgtcggat taaactggtt gagatgacaa gacggcgata agaagaaaga 1980
aaccacagca gctataggca gggaacgtaa agttaagtct gtctatcatc tattaacggt 2040
cagttcagat catagacatt gtaatagcgt tttctttgtc ccgttacatt ttaaacgccg 2100
tgaagatatc catttatttt gatatatata gccaaaaaga taaactttct cgatttacat 2160
cgtgtcagaa gatttaacag aggaaagaag acgatcacga cgtttcccgg ttttccatgt 2220
cttgtgtagt ggcggctact aaccaaataa ctttgtccgt aaaaactgta aaaaagacta 2280
tggtaaaagt ttctttagta tgtttactct gcacatatag cagtggatta tgggtttgca 2340
ttacaattta agcacaaatc gtttagttac ccacaaaaat atcatagatt ctctaaaaat 2400
cacatgatta aatgacattt acaaaatttt gtgattatta ctcggaataa ttcaaagaaa 2460
aatattattt gagcacatta acaaaaaatg gaagtttgaa gtatgtgacg tataaagcca 2520
atgaaagact gacacttcat tgaccacaag tcgtagttcg tatctacaag ctggatgtcc 2580
gtttacccgt ttatatccgt atgataattt cataaatcta actcgataag taaacatcac 2640
aaaaattaca agatacattg gagaaaaaaa aggtttttat tcggcgcatt atttctagtg 2700
gtatatttat tctcattcaa agtcatataa aaaaagattt taaagtaaaa atattctctc 2760
tgaaagtcta gcagtatttt agaacagtac cactacgatt ctcttcttca tcctcttcat 2820
aatatgccat tgtggtgatt gttcatatat ctatatgtac ctcctatcgc ataagactca 2880
cgctaacttc ttattctctc tcaaaagaca gcttttaaat aaaa 2924
<210> 5
<211> 914
<212> DNA
<213> 芥菜型油菜(Brassica juncea)
<400> 5
atttgcatca tatgctactt ttcacttggt gcttgaaagt taattgctgt ttaaaaagtt 60
tgtctcaagc aatttgaaaa tcatatgcaa cttgagggga gaacaaatct aaaacggcta 120
agaaaaatca agtttatata atatttctta tgtatttgaa agattgatgt tttaaattgt 180
atcatacgtt tttgaaatta tatacataca tactttattt tattagttat ttctcttttt 240
ctaataaaat aatcccatca catattttag ttttcagtta aactacatga gttaaaatat 300
aaaatattaa actcttaaat acaaaaaaat aaaacataca aattttaaat aaaatgggcg 360
tattgtcatt caaaatttac tacaaaatct agtaaacgtt tttgagtaaa tatctagaaa 420
atgatacatg attcattcgt gatcagcttg tttgtttcac atataatcaa cgcactaatc 480
attttgggat aaacctttca aaagcaatta ttaaaagaaa caaattgaga gcaatgatga 540
tataagaaat ataggtcatt gtcttgtcaa tcaagttaat aattaaacga aaaggcaacg 600
gaagtagtag tagtagaaga tgatgatgaa gacgaagaca aagttgtgtc acaagtgtgt 660
cttacactat ctgtcttgga aacgggggat tgcctaatgg cattttgagc ttttgcttca 720
gtcactaccg aattctctct gtctattggc agctttagtc ccaaatctat tcaaaacatg 780
caatgatatt ggacgaagaa acttctcaca atccatgacc cttgaggtct tgaatcacct 840
ttcacttttt gttccattgc acggcagata tctaaaaata attttgattt attttacatc 900
atgaagaggc aaca 914
<210> 6
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 6
tactacttcc gttgcctttt cg 22
<210> 7
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
aaaatactgg tgtacattgg aa 22
<210> 8
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 8
aaatccatac acttggggtt 20
Claims (10)
1.一种与芥菜型油菜多室性状相关的两室基因BjMc2，其特征在于，所述两室基因BjMc2的启动子的核苷酸序列如SEQ ID NO.1所示，所述两室基因BjMc2的编码区的核苷酸序列如SEQ ID NO.2所示。
2.如权利要求1所述的与芥菜型油菜多室性状相关的两室基因BjMc2编码的蛋白，其特征在于，所述蛋白的氨基酸序列如SEQ ID NO.3所示。
3.如权利要求1所述的与芥菜型油菜多室性状相关的两室基因BjMc2在制备两室油菜中的应用。
4.如权利要求1所述的两室基因BjMc2的等位基因，其特征在于，所述等位基因为三室基因Bjmc2，所述三室基因Bjmc2的启动子核苷酸序列为：在如SEQ ID NO.1所示的两室基因BjMc2的启动子核苷酸序列中缺失如SEQ ID NO.5所示的914bp的核苷酸序列。
5.根据权利要求4所述的两室基因BjMc2的等位基因，其特征在于，所述三室基因Bjmc2的启动子核苷酸序列如SEQ ID NO.4所示。
6.根据权利要求4所述的两室基因BjMc2的等位基因，其特征在于，所述三室基因Bjmc2的编码区的核苷酸序列如SEQ ID NO.2所示。
7.如权利要求4-6任一项所述的两室基因BjMc2的等位基因在制备多室油菜中的应用。
8.一种用于区分所述两室基因BjMc2及其等位基因三室基因Bjmc2的分子标记，其特征在于，所述分子标记的核苷酸序列如SEQ ID NO.6-8所示。
9.如权利要求8所述的分子标记在油菜多室品种选育中的应用。
10.一种油菜多室品种的选育方法，其特征在于，所述方法包括：提取待测样品的总DNA，采用如权利要求8所述的分子标记进行PCR扩增，扩增产物进行电泳检测：若电泳条带仅出现一条1299bp的特异性条带，则待测样品为仅包含两室基因BjMc2的纯合型两室油菜；若电泳条带只出现一条700bp的特异性条带，则待测样品为仅包含三室基因Bjmc2的纯合型三室油菜；若电泳条带同时包含一条1299bp和一条700bp的电泳条带，则待测样品为杂合型两室油菜。
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| Publication number | Priority date | Publication date | Assignee | Title |
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| CN106279388A (zh) * | 2016-11-08 | 2017-01-04 | 华中农业大学 | 与芥菜型油菜多室性状相关的两室基因BjMc1和三室基因Bjmc1及其应用 |
| CN108239647A (zh) * | 2017-11-29 | 2018-07-03 | 华中农业大学 | 一种控制油菜株型的基因、分子标记及应用 |
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Patent Citations (2)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN106279388A (zh) * | 2016-11-08 | 2017-01-04 | 华中农业大学 | 与芥菜型油菜多室性状相关的两室基因BjMc1和三室基因Bjmc1及其应用 |
| CN108239647A (zh) * | 2017-11-29 | 2018-07-03 | 华中农业大学 | 一种控制油菜株型的基因、分子标记及应用 |
Non-Patent Citations (2)
| Title |
|---|
| LOCUS XP_009106236;佚名;《NCBI》;20201207;参见序列 * |
| 芥菜型油菜多室基因 Bjmc2 的精细定位;王刚;《作物学报》;20161231;第1735-1742页 * |
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