CN110241125B - 大豆百粒重增效基因及其分子标记和应用 - Google Patents
大豆百粒重增效基因及其分子标记和应用 Download PDFInfo
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Abstract
本发明公开了一个大豆百粒重增效基因及其分子标记和应用，该基因碱基序列如SEQ ID No.2所示，该基因对大豆百粒重具有增效作用；本发明鉴定了一个位于大豆Gm16号染色体上调控大豆百粒重的基因GmSLK，该基因的在拟南芥中过表达可以显著提高其籽粒大小。同时提供了一个鉴定该基因的标记InDelSLK。该标记为在种质资源和杂交后代中鉴定其GmSLK是增效基因还是减效基因提供了方便，为大豆育种选材提供了重要的判断价值。
Description
技术领域
本发明属于育种领域，具体涉及调控大豆籽粒百粒重的基因GmSLK的挖掘及其应用。
背景技术
大豆是具有重要经济意义的作物，为人类和动物提供食用油和蛋白质等。现随着人类生活质量的不断提高，对大豆的需求量也越来越大，所以提高大豆的产量一直以来都是国内外大豆育种的主要目标。然而，百粒重作为大豆产量的重要决定因素，也被育种家列为主要的育种性状之一。但是大豆百粒重性状是由多基因控制的数量性状，易受环境的影响，这给研究者带来了很大的阻碍。
针对大豆百粒重性状，国内外许多学者利用不同的初级作图群体和基于不同标记图谱定位了大量的QTL(Quantitative trait loci)。但初级作图群体会使得QTL定位区间过大，大量基因位点同时分离，造成遗传背景效应，增加了主效QTL精细定位的难度。利用次级作图群体，如染色体片段代换系(chromosome segment substitution line,CSSL)的应用，为解决这一难题找到了理想途径。理想状态下，染色体片段代换系个体基因组带有供体亲本染色体的少量片段，其遗传背景高度一致，无疑是进行QTL定位的理想材料。理论上讲，染色体片段代换系的性状值可与轮回亲本的性状值比较，两个株系之间性状差异都因为导入株系的渗入片段存在差异QTL而造成的，因此，利用这一同源的染色体片段代换系进行QTL分析可以提高定位的精确度。
为了更好地实现QTL精细定位及分子辅助育种，可根据染色体片段代换系的定位结果，对检测到的具有重要目标性状的代换系有目的地进行标记辅助回交，构建次级分离群体，将这些重要位点分散在不同的个体中，在高度一致的遗传背景下逐一评价每个候选位点的效应，可实现候选基因的定位和分子辅助育种及基因功能研究的有效手段。
目前虽然定位了大量的关于大豆百粒重的QTL，但是确定直接调控大豆百粒重的基因确很少，因此挖掘调控大豆百粒重的候选基因及基因的功能研究，可为大豆百粒重分子辅助育种提供有用的信息和重要的育种材料。
发明内容
本发明所要解决的技术问题为：提供一个调控大豆百粒重的关键候选基因GmSLK及应用，为大豆百粒重分子辅助育种提供有用的信息和重要的育种材料。
本发明的技术方案为：
大豆百粒重增效基因，其碱基序列如SEQ ID No.2所示。
大豆百粒重增效基因编码的蛋白，其氨基酸序列如SEQ ID No.4所示。
编码SEQ ID No.4所示的氨基酸序列的多核苷酸片段。
重组载体，含有上述所述的多核苷酸片段；优选地，所述多核苷酸片段的序列如SEQ ID No.2所示。
扩增上述所述的大豆百粒重增效基因的特异引物，该特异引物的上游引物序列为SEQ ID No.5，特异引物的下游引物序列为SEQ ID No.6。
大豆百粒重增效基因的分子标记，该分子标记位于大豆染色体第16号染色体Chr16:35923082-35923257bp的位置，该分子标记的引物如SEQ ID No.7和SEQ ID No.8所示。
鉴定大豆是否含有上述所述的大豆百粒重增效基因的方法，以待鉴定大豆的基因组DNA为模板，以SEQ ID No.7和SEQ ID No.8所示的引物进行PCR扩增，若能得到175bp的扩增产物，则该大豆含有所述的大豆百粒重增效基因。
上述所述的大豆百粒重增效基因或蛋白或重组载体或特异引物或分子标记在大豆育种中的应用。
本发明发现了一个调控大豆百粒重基因GmSLK，所述基因位于大豆染色体第16号染色体Chr16:35920962-35928190bp，全基因组长7229bp，完整编码序列长2640bp，分析比较发现来自野生豆N24852的GmSLK基因序列相比较栽培豆NN1138-2的GmSLK基因序列在第一个外显子的谷氨酸串联重复区域多了一个谷氨酰胺。且来自野生豆的GmSLK基因对大豆百粒重具有减效作用，来自栽培豆的GmSLK基因对大豆百粒重具有增效作用。通过进一步定位发现一个分子标记InDelSLK位于GmSLK基因在野生豆N24852与栽培豆NN1138-2的差异区间，因此可以通过这标记来判断不同大豆品种中的GmSLK基因是增效基因还是减效基因，在大豆育种选材中具有重要的价值。
与现有技术相比，本发明具有以下有益效果：
本发明鉴定了一个位于大豆Gm16号染色体上调控大豆百粒重的基因GmSLK，该基因的过表达可以显著提高大豆百粒重性状。同时提供了一个鉴定该基因的标记InDelSLK。该标记为在大豆种子资源和杂交后代中鉴定其GmSLK是增效基因还是减效基因提供了方便，为大豆育种选材提供了重要的判断价值。
附图说明
图1：群体构建示意图；以野生豆N24852为父本，栽培豆NN1138-2为轮回亲本，构建染色体片段代换系(王吴彬等人2013年构建)，CSSL3068为代换系中的一个家系。次级群体以CSSL3068作为母本，NN1138-2作为父本，杂交、自交得到294个后代，单株种植并收获得到294个家系的F2群体，然后每个家系种植一行，自交，并按株行收获得到294个F2:3家系，同理得到F2:4和F2:5家系。
图2：亲本及定位群体百粒重表型；(A)为次级群体亲本NN1138-2和CSSL3068的种子大小示意图；(B)为次级群体亲本NN1138-2和CSSL3068的百粒重比较(**P<0.01)；(C)定位群体F2:3于2015年种植江苏南京(2015NJ)，F2:4群体于2016年种植于江苏南京(2016NJ)和F2:5群体于2017年分别种植江苏南京(2017NJ)和安徽当涂(2017AH)的百粒重表型的次数分布图。
图3：精细定位过程；(A)为16号染色体的初级物理图谱；(B)为16号染色体的局部高密度物理图谱；(C)利用单标记分析法(IciMapping2.0)结合高密度图谱分析次级群体F2:3、F2:4及F2:5在四个环境中的表型数据；(D)利用置换作图法分析11个重组个体的表型和基因型定位候选基因；(E)GmSLK基因结构示意图。
图4：GmSLK扩增条带图；GmSLK-N24852代表GmSLK来自野生豆N24852，GmSLK-NN1138-2代表GmSLK来自栽培豆NN1138-2，GmSLK-CSSL3068代表GmSLK来自代换系CSSL3068，Marker为5000bp，红色箭头指示的条带为3000bp条带。
图5：比较亲本籽粒发育不同时期的籽粒大小和鲜重。(A)比较亲本NN1138-2和CSSL3068在籽粒发育不同时期籽粒大小；(B)比较亲本NN1138-2和CSSL3068在籽粒发育不同时期籽粒鲜重。每个样品统计分析3个生物学重复和3个技术重复(*p<0.05,**p<0.01)。
图6：GmSLK在亲本NN1138-2和CSSL3068籽粒发育不同时期的相对表达量。内参基因为UBI3基因，每个样品统计分析3个生物学重复和3个技术重复(*p<0.05,**p<0.01)。
图7：转基因拟南芥分析GmSLK-NN1138-2(左边)和GmSLK-CSSL3068(右边)基因功能。(A)纯合转基因株系的相对表达量分析，内参基因为Actin基因；(B)转基因植株和野生型的3周植株形态大小比较；(C)比较转基因植株和野生型植株的1000粒重，每个样品统计分析3个生物学重复和3个技术重复(*p<0.05,**p<0.001)。
具体实施方式
实施例1.大豆百粒重主效位点qSW16.1的获得
(1)、定位群体构建及种植
2013年王吴彬等人利用野生豆N24852和栽培豆NN1138-2构建了一套151个家系组成的染色体片段代换系群体，根据群体家系的百粒重表型信息发现其中一个家系CSSL3068的百粒重为14.68g，相比较轮回亲本NN1138-2百粒重(18.7g)显著变小，根据群体的标记信息发现CSSL3068含有一个百粒重的野生位点Sat_224。2014年利用家系CSSL3068与轮回亲本NN1138-2杂交构建次级群体，获得294个株系的F2代次级定位群体。2015年于江苏南京(2015NJ)将F2次级群体种植294个株行，3个重复，然后每行单独收获，获得294个家系的F2:3群体，测定F2:3群体家系的百粒重表型。2016年于江苏南京(2016NJ)将F2:3群体种植294个株行，3个重复，然后每行单独收获，获得294个家系的F2:4群体，测定F2:4群体家系的百粒重表型。2017年将F2:4群体294个株行分别种植于江苏南京(2017NJ)和安徽当涂(2017AH)，3个重复，然后每行单独收获，获得294个家系的F2:5群体，测定F2:5群体家系的百粒重表型(图1)。
(2)、亲本及定位群体百粒重表型的测定及分析
株行种植，株行收获，然后在30℃恒温烘箱至种子恒重，每个亲本及家系挑选发育良好、饱满的100粒种子称重。
首先鉴定分析了两个亲本之间的种子大小(图2A和B)，结果发现亲本NN1138-2的百粒重在四个不同的环境下都显著大于亲本CSSL3068的百粒重(如表1)。此外利用SPSS2.0统计分析了次级群体在四个环境下百粒重表型的频率分布图(图2C)，结果显示都呈连续分布，且在2017AH的环境下F2:5群体百粒重平均值最大为16.94g，在2015NJ环境下群体群体百粒重平均值最小为13.88g(如表1)，这些结果都显示大豆百粒重表型为数量性状，为多基因控制。另外，定位群体在不同的环境下都具有较低的表型变异系数(CV)和较高的遗传率(h2)，这说明了大豆百粒重表型主要变异都是由基因控制的。
表1亲本NN1138-2和CSSL3068不同环境下的百粒重
CV，变异系数；h2，遗传率。
(3)、群体基因型鉴定
采用CTAB法提取次级群体F2的294个株系的大豆叶片的基因组DNA，所用引物由英潍捷基(上海)贸易公司合成。PCR反应体系10ul，其中含有3ulDNA(15ul)、上下游引物(0.2mmol/L)各1.5μL、1.2μL Mgcl2(2.5mmol/L)、0.24μL dNTP(10mmol/L，N＝A，C，G，T),0.12μL Taq酶(5U/ul)和1.4μL ddH2O。PCR反应程序：94℃变性5min；随后进行33个循环的94℃变性30s，55℃退火30s，72℃延伸40s；再经过72℃彻底延伸10min；最后4℃保存。PCR扩增产物用8％聚丙烯酰胺凝胶电泳，银染显色鉴定条带的多态性。
(4)、遗传连锁图谱的构建
利用第16号染色体上的49个SSR分子标记分析两个亲本，统计分析亲本间具有多态性的位点共7个，在次级群体F2株系中鉴定亲本间具有多态性标记的遗传类型，然后对其进行连锁分析，构建次级群体的遗传连锁图谱(图3A)。
(5)、分子标记开发
利用Song等公布的大豆基因组SSR标记信息及野生豆N24852和栽培豆NN1138-2在Gm16号染色体的InDel信息，设计了新的分子标记，统计分析亲本之间具有多态性的SSR标记3个，16个InDel标记(表2)，在次级群体F2中鉴定亲本间具有多态性标记的遗传类型，然后对其进行连锁分析，构建次级群体的高密度图谱(图3B)。
表2本发明涉及的分子标记信息
(6)、结果分析
利用SAS9.4结合次级群体表型数据和分子数据初定位结果显示大豆百粒重主效位点qSW16.1位于第16号染色体Sat_224和Satt431之间。
结合高密度图谱及次级群体F2:3、F2:4及F2:5在四个环境中的表型数据，利用单标记分析法(IciMapping2.0)进行精细定位，结果将大豆百粒重主效位点qSW16.1定位于InDel16_01与BARCSOYSSR_16_1215两个标记之间(图3C)。
实施例2 调控大豆百粒重基因GmSLK的获得
基因GmSLK的确定
通过精细定位将大豆百粒重主效位点qSW16.1定位于Indel16_01与BARCSOYSSR_16_1215两个标记之间，通过比对大豆参考基因序列发现这两个标记之间共有两个候选基因，为了进一步确定候选基因，从F2群体中挑选了11个在Gm16号染色体35889420bp-35952811bp区域有重组的家系，通过置换作图法比较分析这11个重组家系与两个亲本的基因型和百粒重表型数据进一步进行精细定位，发现InDel16_03与BARCSOYSSR_16_1215两个标记之间没有重组位点，与基因Glyma.16g198300共分离(图3D和E)。通过基因注释分析发现基因Glyma.16g198300属于LIM结构域，其功能注释为转录调控因子(TranscriptionalRegulator,SLK2)，因此命名为GmSLK。
实施例3 调控大豆百粒重基因GmSLK在大豆育种中的应用
本发明所述的调控大豆百粒重性状基因的特异扩增上游引物序列为SEQ IDNo.5，特异扩增下游引物序列为SEQ ID No.6(表3)。
表3扩增GmSLK基因的特异引物序列
利用引物序列SEQ ID No.5和SEQ ID No.6，分别以野生豆N24852和两个亲本(NN1138-2和CSSL3068)的RNA为模板扩增GmSLK的编码区(CDS)序列。扩增使用的是Takara公司的高保真酶(Phanta Super-Fidelity DNA Poymerase,Vazyme)，PCR反应体系及程序如表4和表5，扩增条带大小如图4：
表4.PCR反应体系
表5.PCR反应程序
通过测序扩增产物分析比较两个亲本及野生豆N24852之间该基因的CDS序列差异，发现基因GmSLK在亲本CSSL3068和野生豆N24852的序列是一致的，核苷酸序列如SEQ IDNo.1所示，编码的氨基酸序列如SEQ ID No.3所示。说明亲本CSSL3068的GmSLK来自野生豆。栽培豆NN1138-2的GmSLK基因核苷酸序列如SEQ ID No.2所示，编码的氨基酸序列如SEQ IDNo.4所示。野生豆的GmSLK基因在第一个外显子的谷氨酸串联重复区域比栽培豆NN1138-2的GmSLK基因多了一个谷氨酰胺(图3E)。且来自野生豆的GmSLK基因对大豆百粒重具有减效作用，来自栽培豆(NN1138-2)的GmSLK基因对大豆百粒重具有增效作用。
通过序列比对发现InDel16_03位于GmSLK基因在野生豆N24852与栽培豆NN1138-2的差异区间，故将Indel16_03命名为InDelSLK。因此可以通过这标记来判断不同大豆品种中的GmSLK基因是增效基因还是减效基因，在大豆育种选材中具有重要的价值。
鉴定大豆是否含有上述所述的大豆百粒重增效基因的方法，以待鉴定大豆的基因组DNA为模板，以SEQ ID No.7和SEQ ID No.8所示的引物进行PCR扩增，若能得到175bp的扩增产物，则该大豆含有所述的大豆百粒重增效基因。
实施例4 GmSLK基因功能验证
为了能够快速鉴定GmSLK基因对籽粒粒重的影响，本实验选择模式作物拟南芥(Arabidopsis thaliana)Columbia-0野生型作为转基因受体材料，分别过表达亲本CSSL3068的GmSLK(GmSLK-CSSL3068)和栽培豆NN1138-2的GmSLK(GmSLK-NN1138-2)基因，分析比较单拷贝纯合体的1000粒重，从而验证GmSLK基因对籽粒粒重的影响。
(1)GmSLK基因表达水平分析
为了更好的理解GmSLK基因对大豆籽粒发育时期的影响，本实验分析比较GmSLK基因在亲本(NN1138-2和CSSL3068)的籽粒发育不同时期的表达水平。分别取8棵单株NN1138-2和CSSL3068开花后7_DAF、14_DAF、21_DAF,28_DAF、35_DAF及42_DAF(DAF：day afterflower)的籽粒，每个样3个生物学重复，分别测定表型(图5)并提取RNA做(Real-time RT-PCR)RT-PCR分析(图6)。
观察比较亲本NN1138-2和CSSL3068在籽粒发育不同时期的籽粒大小，发现NN1138-2的籽粒在不同的发育时期籽粒相比较CSSL3068籽粒都较大(图5A)，通过测定籽粒的鲜重，结果表明NN1138-2的籽粒在不同的发育时期籽粒鲜重都显著大于CSSL3068籽粒显著(图5B)。
分析比较GmSLK基因在亲本的籽粒发育不同时期的相对表达水平，本实验选取大豆的UBI3(ribosomal-ubiquitin fusion protein)基因作为内参。结果分析表明GmSLK基因在双亲中的表达丰富度随着籽粒的发育而增大，且在42_DAF时期表达量达到最大。然而，GmSLK基因在双亲籽粒发育前期和中期(14_DAF-35_DAF)表达量水平很相似，但是在42_DAF时期，GmSLK基因在亲本NN1138-2的相对表达量水平显著高于在亲本CSSL3068的相对表达量水平。
(2)GmSLK基因转基因拟南芥验证
本研究拟南芥转化所选用的是pCAMBIA 3301载体，酶切位点为(Bst EII和NcoI)，GmSLK基因扩增上下游引物分别是SEQ ID No.3和SEQ ID No.4(表6)。使用克隆重组试剂盒(ClonExperss II)进行重组反应。然后将构建好的过表达载体pCAMBIA 3301-GmSLK-NN1138-2和pCAMBIA 3301-GmSLK-CSSL30068分别转化入DH5a感受态中进行质粒扩繁，然后通过克隆鉴定，提取质粒后转入农杆菌EHA105中。通过花粉侵泡法，分别将构建好的过表达载体转入拟南芥Columbia-0野生型中。
表6GmSLK基因构建载体引物设计
过表达载体pCAMBIA 3301具有草丁膦抗性，阳性苗通过1/2MSA+10mg/l的草丁膦培养基进行筛选。收获侵染的拟南芥植株的T1代种子，首先再抗性培养基上筛选T2代阳性苗，然后种植收获T2种子，抗性培养基上对T2种子种植并统计分析存活/死亡比例，通过卡方检测选择存活/死亡符合3:1，说明该转基因植株属于单拷贝植株，然后种植并收获T3代种子，再种植于抗性培养基选择存活率100％的纯合植株，种植并收获T4种子并统计1000粒重表型。对于GmSLK-NN1138-2和GmSLK-CSSL30068过表达植株分别选择3个单拷贝纯合株系，每个株系分别种植3棵，然后取叶片进行RT-PCR分析，选择拟南芥的Actin基因作为内参基因，检测转基因植株的外源基因的相对表达量(图7A)。本实验也观察分析了转基因植株3周大小的植株大小，发现过表达的GmSLK-NN1138-2和GmSLK-CSSL30068植株相比较野生型植株大小没有差异(图7B)。收获T5种子统计分析1000粒重表型(图7C)，结果分析表明，相比较野生型，过表达的GmSLK-NN1138-2可以显著增大粒重，然而过表达GmSLK-CSSL30068显著降低粒重。因此可以证明来自野生豆或者亲本CSSL3068的GmSLK基因型属于减效基因，可以降低籽粒粒重，而来自栽培豆NN1138-2的GmSLK基因型属于增效基因，可以增加籽粒粒重，这为大豆的产量育种提供了有力的证据。
序列表
<110> 南京农业大学
<120> 大豆百粒重增效基因及其分子标记和应用
<160> 8
<170> SIPOSequenceListing 1.0
<210> 1
<211> 2640
<212> DNA
<213> 大豆(Glycine max)
<400> 1
atgacacctt tgcgagtggc aggtggatta acccaatcat catcaaattc tggaattttc 60
tatcaaggag atgggcagtc acagaatgta gttaactctc acttaagctc atcttttgtt 120
aactcgtcta gcacggtttc tggagcttct cgttcaaacc tgggtccggt ttctggggat 180
atgaataatg cagttttgaa cagtgtggca aactcagcac caagtgtagg agccagctct 240
ttagttacag atgcaaattc tgcactctct ggtggcccac atttgcagag aagtgccagc 300
gttaacacag actcatactt acgattacct gcctcaccta tgtcatttac atcaaataat 360
atcagtattt ctggatcatc agtgatggat gtttcctctg tagtacaaca gagctctcat 420
caagatcaga atgttcaaca attgcagcag aatcagcagc agccacaggg tgcttcaagt 480
gctatgtctt tgtctgcatc tcaaactggc ccttctatgc tccaaatggg tgcacaaatc 540
ccaggatctt tcattcaaga tccaaataat atgtctcatc tgtcaaagaa acctagaatg 600
gatatcaaac aggaagatat gatgcagcag caggttatac aacagattct tcagagacaa 660
gattccatgc aattccaggg tcgtaatccc cagttacagg ctttccttca gcagcagcag 720
cagagactga gacaacaaca aatgtttcag caaatgccac aattacaccg agcacacttg 780
cagcagcagc aacaacaaca acaaatgcaa ttgaggcagc agcagcagca gcagcaacaa 840
caacaacaac aacaacaaca acaacaacaa caacaacaac aacaacaaca acaacaacag 900
caacagcaag tgatgcagcc ctcttctgtg gtcaagcgtc catatgacag tagtgttagt 960
ggggtatgtg cccgtcgatt gatgcagtat ctctatcatc aaaggcaacg accaaatgat 1020
aatagtattg cctattggag aaaatttgtg gctgaatatt actctcctcg tgcaaagaaa 1080
cggtggtgct tgtcattata tagtaatgtt gggcatcacg cacttggtgt ttttccccaa 1140
gcatctatgg atgcatggca ctgtgacata tgtggttcta aatctggaag gggatttgag 1200
gcaacttatg aagtattacc tagacttaat gaaatcaaat ttggcagtgg agtaattgat 1260
gaactattgt ttctggacat gccacgtgaa atgagattcg cttctggtgc gatgatgtta 1320
gaatatggaa aagcagttca agagagtgta tatgagcagc ttcgtgttgt tcgtgaaggt 1380
caacttcgta tcatattcac tcaagacttg aagatattat cttgggagtt ctgtgcaagg 1440
tgccatgagg aacttcttcc tcgaaggttg gttgcaccac aggtcaacca gttagttcag 1500
gtagctaaaa aatgtcaaag tacaattgct gaaagtggtt ctgatggggt ttctcaacaa 1560
gacatccaaa caaacagcaa catgttgttg acagctgggg gtcagcttgc gaagattttg 1620
gagatgcaat cgctaaatga gttgggcttt tctaaaagat acgtgagatg tttgcaaatt 1680
tcggaggttg tcaatagcat gaaagaccta atagatatct gtgcagatca caaaattggt 1740
gccattgaga gtttgaaaaa ttttcctcgt ctagcaacag cctcaaaggt ccagatgcag 1800
aagatgcagg aaatggaaca gctagcaaat gttcaaggtc tgccaactga tcgaaacaca 1860
ctcaataagc taatggcact gaatcctgga ttgaacaacc atataaacaa ccctcataat 1920
atggttaatc gtggtgcttt gagtgggtca gcccaagctg ctttagcact gaacaactac 1980
caaaatcttc tcatgaggca aaattcaatg aactctagcc ctggctcact tcagcgcgaa 2040
gggtcctctt tcaataattc aaaccagagt ccatcttcag ctttgcaagg agctggtcct 2100
gctttaattc caggcccaat gcagaattca tctgttagtg gtttcccaag cccccgtcta 2160
cccccacagc agcagcaaca ccacctacaa cagccctcat taagtgcaaa tgctttactg 2220
caacaaaatc attcacaggg ttcccaagga aatcaagctc tgcagcagca gatgatccat 2280
caactactgc aggagatgtc aaataacaac gggggagtgc aaccacagtc tcttggtgga 2340
cccagtgcaa atatggcaaa gaatgcactg gggtttgggg gccattatcc atccttaagt 2400
ggaggttctg ccaatgttac aggaaacaat ggacctatgt caaggaataa tagcttcaaa 2460
acaactgcaa atagtgattc ttctgctgct ggtggcaaca atggattaaa ccagagaaca 2520
tctgagatgc cacaaaatct acatttgcaa gatgtggttc aggatattgg caatgaattc 2580
acggataatc ccttccttaa cagtgatctt gatgataaca tgggttttgg ctggaaggca 2640
<210> 2
<211> 2637
<212> DNA
<213> 大豆(Glycine max)
<400> 2
atgacacctt tgcgagtggc aggtggatta acccaatcat catcaaattc tggaattttc 60
tatcaaggag atgggcagtc acagaatgta gttaactctc acttaagctc atcttttgtt 120
aactcgtcta gcacggtttc tggagcttct cgttcaaacc tgggtccggt ttctggggat 180
atgaataatg cagttttgaa cagtgtggca aactcagcac caagtgtagg agccagctct 240
ttagttacag atgcaaattc tgcactctct ggtggcccac atttgcagag aagtgccagc 300
gttaacacag actcatactt acgattacct gcctcaccta tgtcatttac atcaaataat 360
atcagtattt ctggatcatc agtgatggat gtttcctctg tagtacaaca gagctctcat 420
caagatcaga atgttcaaca attgcagcag aatcagcagc agccacaggg tgcttcaagt 480
gctatgtctt tgtctgcatc tcaaactggc ccttctatgc tccaaatggg tgcacaaatc 540
ccaggatctt tcattcaaga tccaaataat atgtctcatc tgtcaaagaa acctagaatg 600
gatatcaaac aggaagatat gatgcagcag caggttatac aacagattct tcagagacaa 660
gattccatgc aattccaggg tcgtaatccc cagttacagg ctttccttca gcagcagcag 720
cagagactga gacaacaaca aatgtttcag caaatgccac aattacaccg agcacacttg 780
cagcagcagc aacaacaaca acaaatgcaa ttgaggcagc agcagcagca gcagcaacaa 840
caacaacaac aacaacaaca acaacaacaa caacaacaac aacaacaaca acaacagcaa 900
cagcaagtga tgcagccctc ttctgtggtc aagcgtccat atgacagtag tgttagtggg 960
gtatgtgccc gtcgattgat gcagtatctc tatcatcaaa ggcaacgacc aaatgataat 1020
agtattgcct attggagaaa atttgtggct gaatattact ctcctcgtgc aaagaaacgg 1080
tggtgcttgt cattatatag taatgttggg catcacgcac ttggtgtttt tccccaagca 1140
tctatggatg catggcactg tgacatatgt ggttctaaat ctggaagggg atttgaggca 1200
acttatgaag tattacctag acttaatgaa atcaaatttg gcagtggagt aattgatgaa 1260
ctattgtttc tggacatgcc acgtgaaatg agattcgctt ctggtgcgat gatgttagaa 1320
tatggaaaag cagttcaaga gagtgtatat gagcagcttc gtgttgttcg tgaaggtcaa 1380
cttcgtatca tattcactca agacttgaag atattatctt gggagttctg tgcaaggtgc 1440
catgaggaac ttcttcctcg aaggttggtt gcaccacagg tcaaccagtt agttcaggta 1500
gctaaaaaat gtcaaagtac aattgctgaa agtggttctg atggggtttc tcaacaagac 1560
atccaaacaa acagcaacat gttgttgaca gctgggggtc agcttgcgaa gattttggag 1620
atgcaatcgc taaatgagtt gggcttttct aaaagatacg tgagatgttt gcaaatttcg 1680
gaggttgtca atagcatgaa agacctaata gatatctgtg cagatcacaa aattggtgcc 1740
attgagagtt tgaaaaattt tcctcgtcta gcaacagcct caaaggtcca gatgcagaag 1800
atgcaggaaa tggaacagct agcaaatgtt caaggtctgc caactgatcg aaacacactc 1860
aataagctaa tggcactgaa tcctggattg aacaaccata taaacaaccc tcataatatg 1920
gttaatcgtg gtgctttgag tgggtcagcc caagctgctt tagcactgaa caactaccaa 1980
aatcttctca tgaggcaaaa ttcaatgaac tctagccctg gctcacttca gcgcgaaggg 2040
tcctctttca ataattcaaa ccagagtcca tcttcagctt tgcaaggagc tggtcctgct 2100
ttaattccag gcccaatgca gaattcatct gttagtggtt tcccaagccc ccgtctaccc 2160
ccacagcagc agcaacacca cctacaacag ccctcattaa gtgcaaatgc tttactgcaa 2220
caaaatcatt cacagggttc ccaaggaaat caagctctgc agcagcagat gatccatcaa 2280
ctactgcagg agatgtcaaa taacaacggg ggagtgcaac cacagtctct tggtggaccc 2340
agtgcaaata tggcaaagaa tgcactgggg tttgggggcc attatccatc cttaagtgga 2400
ggttctgcca atgttacagg aaacaatgga cctatgtcaa ggaataatag cttcaaaaca 2460
actgcaaata gtgattcttc tgctgctggt ggcaacaatg gattaaacca gagaacatct 2520
gagatgccac aaaatctaca tttgcaagat gtggttcagg atattggcaa tgaattcacg 2580
gataatccct tccttaacag tgatcttgat gataacatgg gttttggctg gaaggca 2637
<210> 3
<211> 880
<212> PRT
<213> 大豆(Glycine max)
<400> 3
Met Thr Pro Leu Arg Val Ala Gly Gly Leu Thr Gln Ser Ser Ser Asn
1 5 10 15
Ser Gly Ile Phe Tyr Gln Gly Asp Gly Gln Ser Gln Asn Val Val Asn
20 25 30
Ser His Leu Ser Ser Ser Phe Val Asn Ser Ser Ser Thr Val Ser Gly
35 40 45
Ala Ser Arg Ser Asn Leu Gly Pro Val Ser Gly Asp Met Asn Asn Ala
50 55 60
Val Leu Asn Ser Val Ala Asn Ser Ala Pro Ser Val Gly Ala Ser Ser
65 70 75 80
Leu Val Thr Asp Ala Asn Ser Ala Leu Ser Gly Gly Pro His Leu Gln
85 90 95
Arg Ser Ala Ser Val Asn Thr Asp Ser Tyr Leu Arg Leu Pro Ala Ser
100 105 110
Pro Met Ser Phe Thr Ser Asn Asn Ile Ser Ile Ser Gly Ser Ser Val
115 120 125
Met Asp Val Ser Ser Val Val Gln Gln Ser Ser His Gln Asp Gln Asn
130 135 140
Val Gln Gln Leu Gln Gln Asn Gln Gln Gln Pro Gln Gly Ala Ser Ser
145 150 155 160
Ala Met Ser Leu Ser Ala Ser Gln Thr Gly Pro Ser Met Leu Gln Met
165 170 175
Gly Ala Gln Ile Pro Gly Ser Phe Ile Gln Asp Pro Asn Asn Met Ser
180 185 190
His Leu Ser Lys Lys Pro Arg Met Asp Ile Lys Gln Glu Asp Met Met
195 200 205
Gln Gln Gln Val Ile Gln Gln Ile Leu Gln Arg Gln Asp Ser Met Gln
210 215 220
Phe Gln Gly Arg Asn Pro Gln Leu Gln Ala Phe Leu Gln Gln Gln Gln
225 230 235 240
Gln Arg Leu Arg Gln Gln Gln Met Phe Gln Gln Met Pro Gln Leu His
245 250 255
Arg Ala His Leu Gln Gln Gln Gln Gln Gln Gln Gln Met Gln Leu Arg
260 265 270
Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln
275 280 285
Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Val
290 295 300
Met Gln Pro Ser Ser Val Val Lys Arg Pro Tyr Asp Ser Ser Val Ser
305 310 315 320
Gly Val Cys Ala Arg Arg Leu Met Gln Tyr Leu Tyr His Gln Arg Gln
325 330 335
Arg Pro Asn Asp Asn Ser Ile Ala Tyr Trp Arg Lys Phe Val Ala Glu
340 345 350
Tyr Tyr Ser Pro Arg Ala Lys Lys Arg Trp Cys Leu Ser Leu Tyr Ser
355 360 365
Asn Val Gly His His Ala Leu Gly Val Phe Pro Gln Ala Ser Met Asp
370 375 380
Ala Trp His Cys Asp Ile Cys Gly Ser Lys Ser Gly Arg Gly Phe Glu
385 390 395 400
Ala Thr Tyr Glu Val Leu Pro Arg Leu Asn Glu Ile Lys Phe Gly Ser
405 410 415
Gly Val Ile Asp Glu Leu Leu Phe Leu Asp Met Pro Arg Glu Met Arg
420 425 430
Phe Ala Ser Gly Ala Met Met Leu Glu Tyr Gly Lys Ala Val Gln Glu
435 440 445
Ser Val Tyr Glu Gln Leu Arg Val Val Arg Glu Gly Gln Leu Arg Ile
450 455 460
Ile Phe Thr Gln Asp Leu Lys Ile Leu Ser Trp Glu Phe Cys Ala Arg
465 470 475 480
Cys His Glu Glu Leu Leu Pro Arg Arg Leu Val Ala Pro Gln Val Asn
485 490 495
Gln Leu Val Gln Val Ala Lys Lys Cys Gln Ser Thr Ile Ala Glu Ser
500 505 510
Gly Ser Asp Gly Val Ser Gln Gln Asp Ile Gln Thr Asn Ser Asn Met
515 520 525
Leu Leu Thr Ala Gly Gly Gln Leu Ala Lys Ile Leu Glu Met Gln Ser
530 535 540
Leu Asn Glu Leu Gly Phe Ser Lys Arg Tyr Val Arg Cys Leu Gln Ile
545 550 555 560
Ser Glu Val Val Asn Ser Met Lys Asp Leu Ile Asp Ile Cys Ala Asp
565 570 575
His Lys Ile Gly Ala Ile Glu Ser Leu Lys Asn Phe Pro Arg Leu Ala
580 585 590
Thr Ala Ser Lys Val Gln Met Gln Lys Met Gln Glu Met Glu Gln Leu
595 600 605
Ala Asn Val Gln Gly Leu Pro Thr Asp Arg Asn Thr Leu Asn Lys Leu
610 615 620
Met Ala Leu Asn Pro Gly Leu Asn Asn His Ile Asn Asn Pro His Asn
625 630 635 640
Met Val Asn Arg Gly Ala Leu Ser Gly Ser Ala Gln Ala Ala Leu Ala
645 650 655
Leu Asn Asn Tyr Gln Asn Leu Leu Met Arg Gln Asn Ser Met Asn Ser
660 665 670
Ser Pro Gly Ser Leu Gln Arg Glu Gly Ser Ser Phe Asn Asn Ser Asn
675 680 685
Gln Ser Pro Ser Ser Ala Leu Gln Gly Ala Gly Pro Ala Leu Ile Pro
690 695 700
Gly Pro Met Gln Asn Ser Ser Val Ser Gly Phe Pro Ser Pro Arg Leu
705 710 715 720
Pro Pro Gln Gln Gln Gln His His Leu Gln Gln Pro Ser Leu Ser Ala
725 730 735
Asn Ala Leu Leu Gln Gln Asn His Ser Gln Gly Ser Gln Gly Asn Gln
740 745 750
Ala Leu Gln Gln Gln Met Ile His Gln Leu Leu Gln Glu Met Ser Asn
755 760 765
Asn Asn Gly Gly Val Gln Pro Gln Ser Leu Gly Gly Pro Ser Ala Asn
770 775 780
Met Ala Lys Asn Ala Leu Gly Phe Gly Gly His Tyr Pro Ser Leu Ser
785 790 795 800
Gly Gly Ser Ala Asn Val Thr Gly Asn Asn Gly Pro Met Ser Arg Asn
805 810 815
Asn Ser Phe Lys Thr Thr Ala Asn Ser Asp Ser Ser Ala Ala Gly Gly
820 825 830
Asn Asn Gly Leu Asn Gln Arg Thr Ser Glu Met Pro Gln Asn Leu His
835 840 845
Leu Gln Asp Val Val Gln Asp Ile Gly Asn Glu Phe Thr Asp Asn Pro
850 855 860
Phe Leu Asn Ser Asp Leu Asp Asp Asn Met Gly Phe Gly Trp Lys Ala
865 870 875 880
<210> 4
<211> 879
<212> PRT
<213> 大豆(Glycine max)
<400> 4
Met Thr Pro Leu Arg Val Ala Gly Gly Leu Thr Gln Ser Ser Ser Asn
1 5 10 15
Ser Gly Ile Phe Tyr Gln Gly Asp Gly Gln Ser Gln Asn Val Val Asn
20 25 30
Ser His Leu Ser Ser Ser Phe Val Asn Ser Ser Ser Thr Val Ser Gly
35 40 45
Ala Ser Arg Ser Asn Leu Gly Pro Val Ser Gly Asp Met Asn Asn Ala
50 55 60
Val Leu Asn Ser Val Ala Asn Ser Ala Pro Ser Val Gly Ala Ser Ser
65 70 75 80
Leu Val Thr Asp Ala Asn Ser Ala Leu Ser Gly Gly Pro His Leu Gln
85 90 95
Arg Ser Ala Ser Val Asn Thr Asp Ser Tyr Leu Arg Leu Pro Ala Ser
100 105 110
Pro Met Ser Phe Thr Ser Asn Asn Ile Ser Ile Ser Gly Ser Ser Val
115 120 125
Met Asp Val Ser Ser Val Val Gln Gln Ser Ser His Gln Asp Gln Asn
130 135 140
Val Gln Gln Leu Gln Gln Asn Gln Gln Gln Pro Gln Gly Ala Ser Ser
145 150 155 160
Ala Met Ser Leu Ser Ala Ser Gln Thr Gly Pro Ser Met Leu Gln Met
165 170 175
Gly Ala Gln Ile Pro Gly Ser Phe Ile Gln Asp Pro Asn Asn Met Ser
180 185 190
His Leu Ser Lys Lys Pro Arg Met Asp Ile Lys Gln Glu Asp Met Met
195 200 205
Gln Gln Gln Val Ile Gln Gln Ile Leu Gln Arg Gln Asp Ser Met Gln
210 215 220
Phe Gln Gly Arg Asn Pro Gln Leu Gln Ala Phe Leu Gln Gln Gln Gln
225 230 235 240
Gln Arg Leu Arg Gln Gln Gln Met Phe Gln Gln Met Pro Gln Leu His
245 250 255
Arg Ala His Leu Gln Gln Gln Gln Gln Gln Gln Gln Met Gln Leu Arg
260 265 270
Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln
275 280 285
Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Gln Val Met
290 295 300
Gln Pro Ser Ser Val Val Lys Arg Pro Tyr Asp Ser Ser Val Ser Gly
305 310 315 320
Val Cys Ala Arg Arg Leu Met Gln Tyr Leu Tyr His Gln Arg Gln Arg
325 330 335
Pro Asn Asp Asn Ser Ile Ala Tyr Trp Arg Lys Phe Val Ala Glu Tyr
340 345 350
Tyr Ser Pro Arg Ala Lys Lys Arg Trp Cys Leu Ser Leu Tyr Ser Asn
355 360 365
Val Gly His His Ala Leu Gly Val Phe Pro Gln Ala Ser Met Asp Ala
370 375 380
Trp His Cys Asp Ile Cys Gly Ser Lys Ser Gly Arg Gly Phe Glu Ala
385 390 395 400
Thr Tyr Glu Val Leu Pro Arg Leu Asn Glu Ile Lys Phe Gly Ser Gly
405 410 415
Val Ile Asp Glu Leu Leu Phe Leu Asp Met Pro Arg Glu Met Arg Phe
420 425 430
Ala Ser Gly Ala Met Met Leu Glu Tyr Gly Lys Ala Val Gln Glu Ser
435 440 445
Val Tyr Glu Gln Leu Arg Val Val Arg Glu Gly Gln Leu Arg Ile Ile
450 455 460
Phe Thr Gln Asp Leu Lys Ile Leu Ser Trp Glu Phe Cys Ala Arg Cys
465 470 475 480
His Glu Glu Leu Leu Pro Arg Arg Leu Val Ala Pro Gln Val Asn Gln
485 490 495
Leu Val Gln Val Ala Lys Lys Cys Gln Ser Thr Ile Ala Glu Ser Gly
500 505 510
Ser Asp Gly Val Ser Gln Gln Asp Ile Gln Thr Asn Ser Asn Met Leu
515 520 525
Leu Thr Ala Gly Gly Gln Leu Ala Lys Ile Leu Glu Met Gln Ser Leu
530 535 540
Asn Glu Leu Gly Phe Ser Lys Arg Tyr Val Arg Cys Leu Gln Ile Ser
545 550 555 560
Glu Val Val Asn Ser Met Lys Asp Leu Ile Asp Ile Cys Ala Asp His
565 570 575
Lys Ile Gly Ala Ile Glu Ser Leu Lys Asn Phe Pro Arg Leu Ala Thr
580 585 590
Ala Ser Lys Val Gln Met Gln Lys Met Gln Glu Met Glu Gln Leu Ala
595 600 605
Asn Val Gln Gly Leu Pro Thr Asp Arg Asn Thr Leu Asn Lys Leu Met
610 615 620
Ala Leu Asn Pro Gly Leu Asn Asn His Ile Asn Asn Pro His Asn Met
625 630 635 640
Val Asn Arg Gly Ala Leu Ser Gly Ser Ala Gln Ala Ala Leu Ala Leu
645 650 655
Asn Asn Tyr Gln Asn Leu Leu Met Arg Gln Asn Ser Met Asn Ser Ser
660 665 670
Pro Gly Ser Leu Gln Arg Glu Gly Ser Ser Phe Asn Asn Ser Asn Gln
675 680 685
Ser Pro Ser Ser Ala Leu Gln Gly Ala Gly Pro Ala Leu Ile Pro Gly
690 695 700
Pro Met Gln Asn Ser Ser Val Ser Gly Phe Pro Ser Pro Arg Leu Pro
705 710 715 720
Pro Gln Gln Gln Gln His His Leu Gln Gln Pro Ser Leu Ser Ala Asn
725 730 735
Ala Leu Leu Gln Gln Asn His Ser Gln Gly Ser Gln Gly Asn Gln Ala
740 745 750
Leu Gln Gln Gln Met Ile His Gln Leu Leu Gln Glu Met Ser Asn Asn
755 760 765
Asn Gly Gly Val Gln Pro Gln Ser Leu Gly Gly Pro Ser Ala Asn Met
770 775 780
Ala Lys Asn Ala Leu Gly Phe Gly Gly His Tyr Pro Ser Leu Ser Gly
785 790 795 800
Gly Ser Ala Asn Val Thr Gly Asn Asn Gly Pro Met Ser Arg Asn Asn
805 810 815
Ser Phe Lys Thr Thr Ala Asn Ser Asp Ser Ser Ala Ala Gly Gly Asn
820 825 830
Asn Gly Leu Asn Gln Arg Thr Ser Glu Met Pro Gln Asn Leu His Leu
835 840 845
Gln Asp Val Val Gln Asp Ile Gly Asn Glu Phe Thr Asp Asn Pro Phe
850 855 860
Leu Asn Ser Asp Leu Asp Asp Asn Met Gly Phe Gly Trp Lys Ala
865 870 875
<210> 5
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
atgacacctt tgcgagtggc 20
<210> 6
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 6
tcatgccttc cagccaaaac cc 22
<210> 7
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
acaattacac cgagcaca 18
<210> 8
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 8
cttgaccaca gaagaggg 18
Claims (3)
1.核苷酸序列如SEQ ID No.2所示的大豆百粒重增效基因的分子标记，该分子标记位于大豆染色体第16号染色体Chr16: 35923082-35923257bp的位置，该分子标记的引物如SEQ ID No.7和SEQ ID No.8所示。
2.鉴定大豆是否含有核苷酸序列如SEQ ID No.2所示的大豆百粒重增效基因的方法，其特征在于，以待鉴定大豆的基因组DNA为模板，以SEQ ID No.7和SEQ ID No.8所示的引物进行PCR扩增，若能得到175bp的扩增产物，则该大豆含有所述的大豆百粒重增效基因。
3.核苷酸序列如SEQ ID No.2所示的大豆百粒重增效基因或含有SEQ ID No.2所示核苷酸序列的重组载体或SEQ ID No.5和SEQ ID No.6所示的引物对或权利要求1所述的分子标记在大豆百粒重性状育种中的应用。
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| CN105543222B (zh) * | 2016-02-29 | 2019-05-07 | 南京农业大学 | 大豆百粒重主效QTL的分子标记InDeL_33及其应用 |
| CN106755368B (zh) * | 2016-12-09 | 2020-09-04 | 哈尔滨师范大学 | 一种辅助鉴定大豆百粒重性状的分子标记hnusoy05及其应用 |
| CN108411021A (zh) * | 2018-03-13 | 2018-08-17 | 山东省农业科学院作物研究所 | 一种大豆百粒重相关的分子标记方法及其标记组合 |
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