CN113372420A - OsSG2在调控植物种子粒型中的应用 - Google Patents
OsSG2在调控植物种子粒型中的应用 Download PDFInfo
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Abstract
本发明涉及水稻基因工程领域,具体涉及OsSG2在调控植物种子粒型中的应用。本发明首次发现野生日本晴水稻OsSG2基因具有调控水稻粒型的用途,为一个控制水稻粒型新基因;并且进一步研究发现OsSG2在多个植物品种中具有极高的同源保守性,因此OsSG2可用于调控水稻、小麦、高梁、大麦等多种植物种子粒型,用于植物高产品种的培育。
Description
技术领域
本发明涉及水稻基因工程领域,具体涉及OsSG2在调控植物种子粒型中的应用。
背景技术
水稻(Oryza sativa L.)是我国最重要的粮食作物,水稻的栽培总面积约占主要粮食作物种植面积的三分之一,产量接近粮食总产量的三分之一(郭兆武,2008)。但随着社会的发展,人口增长的加剧和耕地面积的减少对水稻的生产造成了极大的压力。如何提高水稻产量一直是育种家和科学家们的关注重点。水稻产量是极为复杂的数量性状,影响因素众多,其构成因素主要包括千粒重、有效穗和穗粒数。粒型是决定千粒重的重要农艺性状,包括粒长、粒宽和粒厚三个指标(孙忠明,2012)。种子大小是由粒长、粒宽、粒厚和谷粒充实度共同决定的,是千粒重最主要的决定因素。种子大小也是构成水稻理想株型的主要农艺性状之一,长期以来一直是很多作物育种改良的重要目标,对产量和外观品质有很大的影响。近年来,伴随着分子生物学的快速发展,越来越多的农业科学家开始使用分子生物学的方法研究农作物的育种问题,利用基因编辑的方法,将原本5至10年的育种周期缩短到2至3年,大大加快了研究进度(Wen Z J,2013),但是,在很长一段时间内,培育高产品种仍是水稻育种的主要研究方向。对于我国而言,近几年,全国大米进口量已超过300万吨,稻米的国际贸易持续保持入口状态。因此,如何提高水稻的产量和促进水稻生产,是目前亟待解决的科学问题(周宏,2003)。水稻作为单子叶植物中的模式植物,水稻基因组的研究也可以为其他植物的研究提供参考和借鉴。粒型是决定水稻产量和品质的一个关键性因素,寻找到新的粒型基因并引入实际应用对水稻遗传育种具有重要的意义。近年来,随着分子生物学的快速发展,越来越多的粒型QTL和粒型基因被发现和克隆,但仍有许多粒型相关基因有待发掘及验证。
水稻的粒型基因主要影响三个方面的性状:粒长、粒宽和粒厚,这三个性状与千粒重呈正相关,并且受多个基因调控(Yong Z X,2010)。目前已经发现的粒型QTL已经超过400个,在水稻的12条染色体中都有分布(表1),主要集中在1、2、3、5号染色体上,9、10和12号染色体上相对较少。在这些粒型基因中,调控粒长的有D11、PGL1、PGL2、SRS3等基因;调控粒宽的有GW2、GS2、GS5、GW5等基因;调节粒厚的有BG2、OsBZR1、WTG1、GL6等基因。粒长、粒宽和千粒重相关基因的研究较多,而粒厚相关基因的研究较少(杨树明,2012)
表1.已克隆的控制水稻粒型的基因
目前已知的粒型调控基因主要通过以下几种途径调控籽粒大小:丝裂原活化蛋白激酶途径(MAPK)、G蛋白信号途径、E3泛素蛋白酶体途径、转录调节因子途径和植物激素途径等(Na L,2019)。通过对比研究表明,通过MAPK途径、G蛋白偶联受体途径和E3泛素蛋白酶体途径的基因是影响细胞增殖进而调控粒型的;而通过植物激素途径的基因是通过同时影响细胞的增殖和扩张来调控粒型的。有越来越多的研究证实,不同的粒型调控基因之间存在着互相调控的关系(Huang R Y,2013)。例如,G蛋白α亚基D1,其突变体对BR不敏感,表明D1参与G蛋白信号与BR信号转导途径共同调控水稻粒型(Wang L,2006)。受甲基化调控的RAV6,其突变体表现BR相关突变体的表型,表明表观修饰与BR途径存在联系(Zhang X,2015)。
对于水稻种子粒型的调控基因有待进一步挖掘,为高产水稻品种的培育奠定基础。
发明内容
本发明主要目的在于提供OsSG2在调控植物种子粒型中的应用。本发明首次发现OsSG2可调控种子粒型,为高产植物品种的培育又提供了一种新的途径。
为实现上述目的,本发明采用以下技术方案:
本发明提供OsSG2在调控植物种子粒型中的应用,所述OsSG2的氨基酸序列如SEQID No.1所示。
进一步地,编码OsSG2的基因为a)或b):
a)所述基因的核苷酸序列如SEQ ID No.2所示;
b)将SEQ ID No.2所示的基因序列经过一个或多个碱基的取代和/或缺失和/或添加得到的基因序列,其编码的蛋白质具有SEQ ID No.2所示基因编码的OsSG2蛋白活性。
进一步地,OsSG2通过以下一种或几种调控途径调控植物种子的粒型:
1)参与APG/PGL2的调控途径调控种子的粒型;
2)调控颖壳细胞的形态改变粒型;
3)与其他蛋白互作形成异源二聚体或多聚体进行调控。
进一步地,所述植物包括单子叶植物和双子叶植物;
所述单子叶植物包括水稻、小麦、大麦、高粱和玉米;
所述双子叶植物包括拟南芥、番茄、烟草、大豆和土豆。
进一步地,通过调控种子的粒长、粒宽来控制种子的粒型。
本发明还提供过表达OsSG2基因的产品在调控植物种子粒型中的应用,所述OsSG2的氨基酸序列如SEQ ID No.1所示。
进一步地,所述产品包括过表达载体、引物和试剂。
本发明还提供抑制OsSG2基因表达的产品在调控植物种子粒型中的应用,所述OsSG2的氨基酸序列如SEQ ID No.1所示。
进一步地,所述产品包括OsSG2敲除载体,引物和试剂。
本发明还一种调控植物种子粒型的方法,以OsSG2基因为目的基因,其核苷酸序列如SEQ ID No.2所示,采用CRISPR/Cas9基因编辑系统,构建所述目的基因的敲除载体,转化入植物中,对植物进行培育;
以SEQ ID No.7所示碱基序列为靶标序列,采用SEQ ID No.3或SEQ ID No.4所示gRNA序列进行敲除。
与现有技术相比,本发明具有以下有益效果:
本发明首次发现野生日本晴水稻OsSG2基因具有调控水稻粒型的用途,为一个控制水稻粒型新基因;并且进一步研究发现OsSG2在多个植物品种中具有极高的同源保守性,因此OsSG2可用于调控水稻、小麦、高梁、大麦等多种植物种子粒型,用于植物高产品种的培育。
附图说明
构成本发明的一部分的说明书附图用来提供对本发明的进一步理解,本发明的示意性实施例及其说明用于解释本发明,并不构成对本发明的不当限定。
图1为以OsSG2氨基酸序列构建系统进化发育树图。
图2为OsSG2的基因结构图及基因敲除靶位点设计图。
图3为野生日本晴和两个敲除突变体sc2-1、sc2-2的植株对比照片;其中WT为野生型日本晴,sc2-1、sc2-2为基因敲除转基因株系单株。
图4为野生日本晴和两个敲除突变体OsSG2-1、OsSG2-2的籽粒长度对比照片;其中WT为野生型日本晴,sc2-1、sc2-2为基因敲除转基因株系单株的种子。
图5为野生日本晴和两个敲除突变体sc2-1、sc2-2的籽粒宽度对比照片;其中WT为野生型日本晴,sc2-1、sc2-2为基因敲除转基因株系单株的种子。
图6为野生日本晴和两个敲除突变体sc2-1、sc2-2其他农艺性状统计图,其中包括株高、分蘖数、千粒重、粒长、粒宽、长宽比;其中WT为野生型日本晴,sc2-1、sc2-2为基因敲除转基因株系单株。
图7为野生日本晴和敲除突变体OsSG2-1花器官数目对比图;其中WT为野生型日本晴,sc2-1、sc2-2为基因敲除转基因株系单株。
图8为野生型、sc2-1、sc2-2颖壳细胞学对比;其中A-C分别是:野生型、sc2-1、sc2-2的颖壳细胞。D为外稃细胞纵向长度柱形统计图。E为外稃细胞横向长度柱形统计图。F为外稃细胞面积柱形统计图;其中WT为野生型日本晴,sc2-1、sc2-2为基因敲除转基因株系单株。
图9为野生型、sc2-1、sc2-2成熟籽粒横截面的淀粉颗粒对比图,其中A为野生型,B为sc2-1,C为sc2-2,D为野生型籽粒横截面淀粉颗粒放大图,E为Sc2-1籽粒横截面淀粉颗粒放大图,F为sc2-2籽粒横截面淀粉颗粒放大图;其中WT为野生型日本晴,sc2-1、sc2-2为基因敲除转基因株系单株。
图10为粒型基因在WT与sc2-1中的表达水平对比图,其中WT为野生型日本晴,sc2-1为基因敲除转基因株系单株。
图11为淀粉合成基因在WT与sc2-1中的表达水平对比图,其中WT为野生型日本晴,sc2-1为基因敲除转基因株系单株的种子。
图12为以OsSG2为诱饵蛋白进行酵母文库筛选的结果图。AD与BD均为酵母文库所用载体,CRD与GLLAH均为与OsSG2互作的蛋白;其中SC2为OsSG2蛋白。
具体实施方式
应该指出,以下详细说明都是示例性的,旨在对本发明提供进一步的说明。除非另有指明,本文使用的所有技术和科学术语具有与本发明所属技术领域的普通技术人员通常理解的相同含义。
需要注意的是,这里所使用的术语仅是为了描述具体实施方式,而非意图限制根据本发明的示例性实施方式。如在这里所使用的,除非上下文另外明确指出,否则单数形式也意图包括复数形式,此外,还应当理解的是,当在本说明书中使用术语“包含”和/或“包括”时,其指明存在特征、步骤、操作和/或它们的组合。
为了使得本领域技术人员能够更加清楚地了解本发明的技术方案,以下将结合具体的实施例详细说明本发明的技术方案。
实施例1
(一)试验材料
供试的水稻品种为水稻粳稻品种日本晴,即为水稻(Oryza sativa Japonica)。日本晴品种水稻于四川农业大学水稻研究所遗传研究室进行保存。每季新收的种子种植前先用质量百分比浓度为2%过氧化氢消毒30min,在供氧的纯净水(内含1%H2O2)中于28℃环境下浸种1d,然后将种子放入30℃培养箱下进行发芽培养,每天换水两到三次,看到露白后转入28℃下培养。待水稻芽长到5-7mm长的时候播种到纱窗布上,并在水稻营养液中培养到3叶期后,将水稻幼苗移入水稻栽培田进行管理,观察,性状统计。
(二)试验方法
2.1所用试剂和载体
质粒CRISPR/Cas9P35S-H;农杆菌(Agrobacterium)感受态EHA105购于昂羽生物技术有限公司;大肠杆菌(EScherichia coli)DH5α感受态购于北京全试金生物公司;质粒提取试剂盒购于诺唯赞(南京)公司Plasmid Mini Kit DC201,FastPureEndoFree Plasmid Maxi Kit DC202;NAprep Pure植物总RNA提取试剂盒购于北京全试金生物公司;反转录试剂盒购于诺唯赞(南京)公司HiScript II Q RT SuperMix for qPCR(+gDNA wiper)R223-01;qPCR试剂盒购于诺唯赞(南京)公司;AceQ qPCR SYBR Green MasterMix(without ROX);重组酶One Step Cloning Kit购于南京诺唯赞生物科技有限公司;
2.2 OsSG2生物信息学分析
利用GO富集数据库及KEGG富集数据库对粒型基因的挖掘过程中发现了OsSG2基因表现出一定的变化规律及高度的保守性,我们猜测其可能属于管家基因,且截至目前尚无该基因的自然突变体及EMS诱变突变体。在水稻国家数据库中下载OsSG2基因序列与蛋白序列,以该基因的蛋白质序列作在NCBI在线数据库中通过BLASTp比对方法进行比对,得到了95条来自不同植物中的同源蛋白序列,经过筛选同源性大于80%的序列,最终选取了56条蛋白序列,下载序列后,利用MEGA5.0软件进行多序列插入与比对,通过Neighbour-joining(NJ)方法进行系统进化树分析。如图1所示,在数据库中选择不同物种的OsSG2蛋白构建系统进化发育树,可以看出OsSG2蛋白的功能是保守的。水稻中粳稻亚种的OsSG2蛋白与野生稻中预测的OsSG2蛋白亲缘关系最近,但都未见相关报道。在这些序列中,OsSG2蛋白没有与苔藓、蕨类、藻类植物的同源蛋白,仅在种子植物的被子植物中有同源蛋白。OsSG2蛋白在单子叶和双子叶植物中均存在多个保守的分支,在单子叶植物中旁系关系更近,在双子叶植物中存在的数量更多。
2.2 OsSG2敲除载体的构建
依据现有公开的在线设计网站,结合实验需求,设计、选取合适靶位点并设计相关引物;
利用双重PCR构建靶标sgRNA表达盒。本实验采用双靶点敲除载体,构建了单靶点sgRNA表达盒(U6a-目的基因编码区序列):以SEQ ID No.7所示碱基序列为靶标序列,采用SEQ ID No.3或SEQ ID No.4所示gRNA序列进行敲除。
靶标sgRNA表达盒克隆到表达载体pYLCRISPR/Cas9P35S-H:BasI酶切载体pYLCRISPR/Cas9P35S-H,使用T4连接酶将构建完成的sgRNA表达盒连接到表达载体上,再经过大肠杆菌转化、单克隆菌落PCR鉴定和提取质粒后,测序验证重组敲除载体是否正确。
在OsSG2上游预测启动子关键区域设计第一个靶位点,在OsSG2编码区设计第二个靶位点构建了OsSG2的敲除载体,农杆菌转化日本晴。经过验证后筛选分类,获得2个敲除转基因株系。室内培养这2个敲除转基因株系并选取叶片提取DNA,以靶位点及附近序列为模板,设计PCR扩增引物,PCR扩增,电泳检测大小合适片段,测序验证,基因序列比对。根据2个敲除转基因株系靶位点突变方式的不同,分别将其命名为OsSG2-1和OsSG2-2(图2、图3)其核苷酸序列分别如SEQ ID No.5、SEQ ID No.6所示。
2.3农杆菌介导水稻遗传转化
消毒:挑选成熟饱满且胚完整的受体材料种子,脱壳,在75%酒精中浸泡1min,转到0.15%HgCl2中继续浸泡20min,再用无菌水漂洗多次,直至将消毒液清洗干净,放入铺好灭菌滤纸的培养皿中,置于超净工作台上吹10~15min至风干;
愈伤组织诱导:在每个配制好的诱导培养皿中接入15粒种子,27℃无光培养10~15d至长出愈伤组织颗粒后,继续培养15d后继代,选择第三代胚性愈伤组织用于后续实验;
预培养:选取分散、鲜黄的2~3mm的颗粒状愈伤组织,转到预培养培养基中,置于27℃无光培养3d;
侵染:将预培养的愈伤组织置于三角瓶中,加入包含目的基因的农杆菌菌液浸泡30min,干燥愈伤组织,并将其接种在共培养基上,19℃无光培养3d;
筛选:将愈伤组织转到筛选培养基,30℃持续光照培养15d,再转到新的筛选培养基上,30℃培养至新的愈伤组织长出;
分化:将筛选后的愈伤组织转到分化培养基上,30℃光照培养至愈伤组织转绿并分化;
生根:将愈伤组织转移至生根培养基,30℃光照培养至生根;
移栽:已生根的植株用水洗净培养基后,移栽至土壤中。
2.4敲除转基因植株的验证
提取CRISPR/Cas9敲除转基因植株DNA,PCR扩增对应片段,电泳检测,测序验证,利用DNA MAN软件进行序列比对,归纳各转基因株系的突变方式。
2.5 CTAB提取DNA
取0.1g叶片置入1.5mL离心管中,倒入液氮,用研磨棒磨碎、磨细;
加入700μL预热到65℃的2%CTAB缓冲液,摇匀;
65℃水浴30min,期间多次摇匀;
加入400μL氯仿-异戊醇(体积比24:1)混合液,轻轻摇匀,室温放置10min;
室温,10000r/min,离心10min;
加入700μL-20℃预冷的无水乙醇至新离心管中,将上清液转至新离心管中,摇匀,-20℃静置10min(可长期保存);
室温,10000r/min,离心10min,弃上清加入700μL 75%乙醇;
室温,10000r/min,离心10min,弃上清,室温阴干2~3h;
加入50~200μL ddH2O,室温溶解30min;
核酸检测仪检测DNA浓度与质量,4℃保存。
2.6基因的qPCR分析
2.6.1 RNA提取
在光照培养箱培养日本晴植株,在其三叶期取整株于液氮保存;在大田培养日本晴及其转基因敲除株系,在其分蘖期分别取根、茎、叶于液氮保存,在其孕穗期取幼穗于液氮保存。采用TRIZOL试剂提取法提取RNA,具体步骤如下:
在液氮中研磨水稻组织,磨细后倒入1.5mL离心管中,加入1mL TRIZOL液,室温放置5min;
加入200μL三氯甲烷,涡旋剧烈震荡30s,室温放置3min;
4℃,12000r/min离心15min,吸取上层水相(约400μL)至新离心管,加入500μL异丙醇,上下颠倒混匀,室温放置20min;
4℃,12000r/min离心10min,弃上清,加入1mL的75%乙醇(DEPC-H2O配置);
4℃,12000r/min离心5min,弃上清,加入700μL无水乙醇,15℃,12000r/min离心10min,弃上清;
室温放置5~10min,待无水乙醇挥发,加入20~40μL的DEPC-H2O,用枪头轻轻吹打,室温溶解10~20min;
核酸浓度检测仪检测RNA浓度与质量,-80℃保存。
RNA反转录
按照诺唯赞(南京)公司提供的HiOsSGript II Q RT SuperMix for qPCR(+gDNAwiper)试剂盒说明方法操作:在PCR管中加入总量1000ng的RNA,以RNase free ddH2O定量到16μL,加4μL的4×Gdna wiper Mix,移液器轻轻吹打混匀,42℃孵育2min去除DNA;加入4μL的HisOsSGript II qRT SuperMix II,移液器轻轻吹打混匀,50℃15min后85℃加热5s终止反应。核酸浓度检验仪上检验cDNA浓度与质量,-20℃保存。
2.6.3荧光定量PCR
利用primer premier 5软件根据每一个目的基因设计适合的qPCR引物,选用OsActin(Os03g0718100)为内参基因,以反转录的cDNA为模板,按照诺唯赞公司提供的SYBR Green Master Mix(without ROX)试剂盒说明方法操作。qPCR反应体系为20μL:2×SYBR Green Master Mix加10μL,ddH2O加7.7μL,cDNA模板加1.5μL,引物各加0.4μL。反应程序为95℃预变性5min;95℃变性15s;60℃退火延伸30s;40个循环,每个样品均设有3个重复。反应结束后,采用仪器自带的qPCRsoft 3.2软件定义样品并导出数据到EXCEL表格软件,利用EXCEL软件进行DDCT算法处理数据,计算3次生物学重复的标准误差,计算显著性差异,进行数据分析并绘制柱状图。
同时检测了一些调控籽粒粒长、粒宽及长宽比的基因APG、PGL1、PLG2、FUWA、SMG1、GL3-1、GLW7、GW6a、GW2、GS5和GL7在WT与OsSG2-1的3cm YP中表达水平,结果如图10所示:相对于野生型,APG与PGL2在sc2-1中的表达水平最高,分别上调62倍与17倍,SMG1、GL3-1、GLW7及GW6a中则无显著变化。PGL1、FUWA、GW2和GS5在sc2-1中无表达或低表达(CT>40)。由此可知,OsSG2参与APG/PGL2的调控途径。
如图9所示,sc2-1和sc2-2籽粒的淀粉颗粒的形态发生变化,为验证SG2与淀粉合成的关系,在sc2-1中对淀粉分支酶基因OsBE I和OsBE IIb、淀粉合酶基因SS I和SS II的表达量进行检测。如图11所示,相对于野生型,淀粉合酶基因OsBE I、OsBE IIb、SS I和SSII在OsSG2-1中的表达量无显著变化,说明OsSG2不影响淀粉分支酶和淀粉合酶途径,而是影响其他途径调控淀粉颗粒的形态。
2.7农艺性状调查
大田培养日本晴野生型及转基因敲除株系,在开花期随机选择21株发育正常、处于相同花期的日本晴野生型及转基因敲除植株,对其开花穗部做好标记后,分别在第3、5、7、10、15、20、25d取小穗调查灌浆情况,拍照记录,并使用电子天平称量鲜重与干重。后期用Adobe Photoshop CC 2019图像处理软件排版处理,EXCEL软件整理数据,计算平均值并绘制折线图。
大田培养日本晴野生型及转基因敲除株系,在成熟期随机选择10株发育正常的野生型及转基因植株,分单株调查统计株高、分蘖数、粒长、粒宽及千粒重等农艺性状,三次生物学重复,拍照记录,利用考种仪测量并记录各项数据,利用EXCEL软件进行处理数据,计算3次生物学重复的标准误差,计算显著性差异,进行数据分析并绘制柱状图。
为了分析OsSG2敲除株系与野生型的表型差异,测量了野生型、sc2-1和sc2-2的株高、籽粒长度、长宽比、籽粒宽度、分蘖数和千粒重.如图3-6所示,与野生型相比:两个突变体的分蘖数和株高无显著变化,整体株型无显著变化;sc2-1和sc2-2粒长增加5%和6%,粒宽降低6%,长宽比增加11%和14%,千粒重无显著变化。以上结果说明,相对于野生型,在sc2-1和OsSG2-2中的粒型由原本的短圆变为了细长,且对千粒重没有显著影响。我们推测,可能是由于OsSG2表达水平较高的部位集中在幼穗,当OsSG2失去部分结构域以致其功能性丧失,无法完成相关信号转导,导致粒型发生变化;其余部位由于OsSG2的表达水平较低,当其功能丧失时反而不受影响。
2.8雌雄蕊数量观察
在大田培养日本晴野生型及转基因敲除株系,在开花前分别取其颖花,用镊子剥下内外稃及护颖,在体视显微镜下观察并拍照记录雌雄蕊数量,后期用Adobe PhotoshopCC 2019图像处理软件排版处理。
OsSG2在水稻中的同源基因FON1控制水稻花器官的生长发育,其突变体fon1表现出雌雄蕊数量上的变异。为了验证OsSG2-1是否有同样的表型,我们选取了开花前的sc2-1颖花部位,剥去内外稃和护颖,在体视显微镜下观察。如图7所示,在sc2-1的颖花中有6个雄蕊、2个雌蕊,与野生型相一致。结果表明,OsSG2敲除没有导致花器官数目异常,证明OsSG2可能不参与控制水稻花器官的生长发育。
2.9籽粒横截面形态和颖壳细胞数量及形态观察
在大田培养日本晴野生型及转基因敲除株系,取开花前的颖花和成熟的籽粒,用3%戊二醛固定液将其过夜固定,依次使用50%、70%、85%、95%、100%乙醇溶液脱水15min,再经过临界点干燥、喷金后,使用扫描电镜观察并拍照记录其细胞数量及形态。
水稻颖壳的形态是影响粒型的一个因素,细胞的生长、繁殖会在一定程度上影响颖壳的形态。为了研究OsSG2敲除是否影响颖壳细胞的大小及数量,分别选取了野生型和敲除突变体的成熟籽粒,剥下外稃后通过电镜扫描观察。如图8所示,相对于野生型,突变体的颖壳细胞长度显著降低,宽度显著增加,细胞面积无显著变化,细胞密度无显著变化。但是同一视野内颖壳细胞列数减少,行数增加,导致籽粒长宽比增加,粒型最终表现为细长型。这些结果表明,OsSG2敲除后导致了颖壳细胞长度降低及宽度增加。由此可知,OsSG2通过调控细胞的形态改变粒型。
2.10酵母文库筛选以及酵母双杂交
粳稻穗部组织酵母文库构建:取宜香1B的5~6cm长幼穗组织,送往上海欧易生物医学科技有限公司进行酵母文库构建;
诱饵蛋白载体构建:扩增诱饵蛋白基因OsSG2,采用双酶切和同源重组的方式,将诱饵蛋白连在pGBKT7载体上,使用T7通用引物对其测序以检测载体是否正确;酵母文库筛库:使用Clontech公司基于GAL4的Matchmaker Gold Yeast Two Hybrid System进行酵母双杂交实验。
以OsSG2基因序列为诱饵蛋白,在酵母文库中筛选到2个互作蛋白(如图12所示),查询其基因序列,均为未报道基因,注释为:假定表达的未表征富含Cys的结构域蛋白CRD(Cys-Rich Domain)和假定表达的GDSL样脂肪酶/酰基水解酶GLLAH(GDSL-like lipase/acyl hydrolase)。由此可知,OsSG2不能单独行使功能,需要与其他蛋白互作形成异源二聚体或多聚体行使功能。
本发明通过试验发现OsSG2具有控制种子大小的作用,并对OsSG2基因的上下游基因及其参与的相关调控信号通路进行分析。通过分析该基因在稻种子大小调控机理研究及粒型改良种培育品种中的作用,以为丰富禾谷类作物种子大小的分子机理奠定理论基础。
上述实施例为本发明较佳的实施方式,但本发明的实施方式并不受上述实施例的限制,其他的任何未背离本发明的精神实质与原理下所作的改变、修饰、替代、组合、简化,均应为等效的置换方式,都包含在本发明的保护范围之内。
SEQUENCE LISTING
<110> 四川农业大学
<120> OsSG2在调控植物种子粒型中的应用
<130>
<160> 7
<170> PatentIn version 3.5
<210> 1
<211> 619
<212> PRT
<213> 人工序列
<400> 1
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Ala Leu Ala Leu Ala Pro Arg Pro Ala Ala Pro Ala Ala Thr Asp Arg
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Ala Ala Leu Leu Ala Phe Arg Ala Ser Leu Pro Pro Pro Ser Arg Ala
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Ala Leu Ala Ser Trp Arg Gly Pro Leu Ser Glu Ser Trp Arg Gly Val
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Ser Leu His Pro Pro Ala Ala Ala Gly Ala Pro Ala Pro Ala Pro Pro
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Pro Ser Val Ser Gly Leu Ala Leu Arg Gly Leu Asn Leu Ser Gly Gln
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Leu Pro Ala Ala Pro Leu Ala Leu Leu Arg Arg Val Arg Ala Leu Asp
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Leu Ser Ala Asn Ala Leu Ser Gly Glu Leu Pro Cys Ser Leu Pro Arg
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Ser Leu Leu Asp Leu Asp Leu Ser Arg Asn Ala Leu Ser Gly Ala Val
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Pro Thr Cys Phe Pro Ala Ser Leu Pro Ala Leu Arg Ala Leu Asn Leu
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Ser Ala Asn Ala Leu Arg Phe Pro Leu Ser Pro Arg Leu Ser Phe Pro
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Ala Ser Leu Ala Ala Leu Asp Leu Ser Arg Asn Ala Leu Thr Gly Ala
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Val Pro Pro Arg Val Val Ala Asp Pro Asp Ala Ser Gly Leu Leu Leu
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Leu Asp Leu Ser His Asn Arg Phe Ser Gly Glu Ile Pro Val Gly Ile
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Thr Ala Ile Arg Ser Leu Gln Gly Leu Phe Leu Ala Asp Asn Gln Leu
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Ser Gly Glu Ile Pro Thr Gly Ile Gly Asn Leu Thr Tyr Leu Gln Ala
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Leu Asp Leu Ser Arg Asn Arg Leu Ser Gly Val Val Pro Ala Gly Leu
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Ala Gly Cys Phe Gln Leu Leu Tyr Leu Arg Leu Gly Gly Asn His Leu
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Ser Gly Ala Leu Arg Pro Glu Leu Asp Ala Leu Asp Ser Leu Lys Val
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Leu Asp Leu Ser Asn Asn Arg Ile Ser Gly Glu Ile Pro Leu Pro Leu
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agtggagctg tggccaaatg gcagagcttg aggttcttat cactggctgg taaccagctc 1080
tctggtcagc taccggattg gatgttctcg ttcccaacac ttcagtggat tgatttgtcc 1140
ggcaataggt ttgtgggttt catcccggat ggtgggttca atgtcagtgc cgtccttaat 1200
ggtggaggca gtggtcaggg gagtccatca gaggctgtgc ttccacctca gctgtttgtg 1260
tcagtgtcca cggatatggc aggccggcaa ttggagctgg gctatgatct tcaagcagct 1320
accgggatag atctgtctag gaatgagctc cgtggggaga taccagacgg gttggttgca 1380
atgaagggac tggagtattt gaatctctct tgtaattatt tggatgggca gattccttca 1440
ggtattgggg ggatggggaa gctacggacc cttgacttct cacataatga gttgtcaggg 1500
gtggtacctc ctgagatagc tgccatgaca gagcttgagg tgcttaatct ctcctacaat 1560
agcctatctg ggcctttgcc gacaactgat gggttgcaga agtttccagg agcattggct 1620
ggaaaccctg ggatatgcag cggggaagga tgcagtgcgc attctaggat gccagaaggg 1680
aaaatggcag gaagcaatcg ccatggttgg cttggtggct ggcatggaga gaacggatgg 1740
gtatccctag gtgcattctg tatcagcaca atgactagct tctatgtatc attggcaact 1800
ttgctgtgct cccgcaaggc aaggaacttt gtgtttcggc ctggaaggat ggaatattaa 1860
<210> 3
<211> 30
<212> DNA
<213> 人工序列
<400> 3
gttttagagc tagaaatagc aagttaaaat 30
<210> 4
<211> 30
<212> DNA
<213> 人工序列
<400> 4
caaaatctcg atctttatcg ttcaatttta 30
<210> 5
<211> 1860
<212> DNA
<213> 人工序列
<400> 5
atgccgaacg cctctcccct ccaccacctc gcctccctcc tcctccttgc cctcgccctc 60
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tccctcccgc cgccctcccg cgcggcgctc gcgtcgtggc gcggcccgct ctcggagtca 180
tggcgcggcg tttcgctcca cccgccggcc gccgccggcg cccccgcccc ggccccgccg 240
ccctccgtct ccgggctcgc gctgcggggg ctcaacctgt cggggcagct cccggcggcg 300
ccgctcgcgc tgctccgccg cgtccgcgcg ctcgacctct cggccaacgc gctctcgggc 360
gaactgccgt gctccctgcc gcgctcgctc ctcgacctcg acctctcccg caacgcgctc 420
tccggggccg tcccgacctg cttcccggcc tcgctcccgg ctctccgcgc cctcaacctc 480
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gcgctcgacc tctcgcggaa cgcgctcacg ggcgccgtcc cgccgcgggt cgtcgccgac 600
cccgacgctt cgggcctcct cctcctcgac ctctcccaca atcgcttctc cggcgagatc 660
cccgtcggga tcaccgccat ccggagcctt cagggattgt ttcttgcgga taaccagctc 720
tccggggaga ttccgactgg gattgggaat ttaacctact tgcaggcgct tgatttgtcg 780
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ggcaataggt ttgtgggttt catcccggat ggtgggttca atgtcagtgc cgtccttaat 1200
ggtggaggca gtggtcaggg gagtccatca gaggctgtgc ttccacctca gctgtttgtg 1260
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accgggatag atctgtctag gaatgagctc cgtggggaga taccagacgg gttggttgca 1380
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<210> 6
<211> 1860
<212> DNA
<213> 人工序列
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gcgctcgacc tctcgcggaa cgcgctcacg ggcgccgtcc cgccgcgggt cgtcgccgac 600
cccgacgctt cgggcctcct cctcctcgac ctctcccaca atcgcttctc cggcgagatc 660
cccgtcggga tcaccgccat ccggagcctt cagggattgt ttcttgcgga taaccagctc 720
tccggggaga ttccgactgg gattgggaat ttaacctact tgcaggcgct tgatttgtcg 780
cgtaataggc tctctggtgt agtgcctgct ggccttgccg gctgcttcca gcttctgtac 840
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gccgggtgcc gatctctgga ggtggtgaac ttgtcaggaa acaagatcac aggggagctc 1020
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ggcaataggt ttgtgggttt catcccggat ggtgggttca atgtcagtgc cgtccttaat 1200
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ggaaaccctg ggatatgcag cggggaagga tgcagtgcgc attctaggat gccagaaggg 1680
aaaatggcag gaagcaatcg ccatggttgg cttggtggct ggcatggaga gaacggatgg 1740
gtatccctag gtgcattctg tatcagcaca atgactagct tctatgtatc attggcaact 1800
ttgctgtgct cccgcaaggc aaggaacttt gtgtttcggc ctggaaggat ggaatattaa 1860
<210> 7
<211> 17
<212> DNA
<213> 人工序列
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gacctgcttc ccgggcc 17
Claims (10)
1.OsSG2在调控植物种子粒型中的应用,其特征在于,所述OsSG2的氨基酸序列如SEQID No.1所示。
2.根据权利要求1所述的应用,其特征在于,编码OsSG2的基因为a)或b):
a)所述基因的核苷酸序列如SEQ ID No.2所示;
b)将SEQ ID No.2所示的基因序列经过一个或多个碱基的取代和/或缺失和/或添加得到的基因序列,其编码的蛋白质具有SEQ ID No.2所示基因编码的OsSG2蛋白活性。
3.根据权利要求1所述的应用,其特征在于,OsSG2通过以下一种或几种调控途径调控植物种子的粒型:
1)参与APG/PGL2的调控途径调控种子的粒型;
2)调控颖壳细胞的形态改变粒型;
3)与其他蛋白互作形成异源二聚体或多聚体进行调控。
4.根据权利要求1-3任一项所述的应用,其特征在于,所述植物包括单子叶植物和双子叶植物;
所述单子叶植物包括水稻、小麦、大麦、高粱和玉米;
所述双子叶植物包括拟南芥、番茄、烟草、大豆和土豆。
5.根据权利要求1-3任一项所述的应用,其特征在于,通过调控种子的粒长、粒宽来控制种子的粒型。
6.过表达OsSG2基因的产品在调控植物种子粒型中的应用,其特征在于,所述OsSG2的氨基酸序列如SEQ ID No.1所示。
7.根据权利要求6所述的应用,其特征在于,所述产品包括过表达载体、引物和试剂。
8.抑制OsSG2基因表达的产品在调控植物种子粒型中的应用,其特征在于,所述OsSG2的氨基酸序列如SEQ ID No.1所示。
9.根据权利要求8所述的应用,其特征在于,所述产品包括OsSG2敲除载体,引物和试剂。
10.一种调控植物种子粒型的方法,其特征在于,以编码OsSG2的基因为目的基因,其核苷酸序列如SEQ ID No.2所示,采用CRISPR/Cas9基因编辑系统,构建所述目的基因的敲除载体,转化入植物中,对植物进行培育;
以SEQ ID No.7所示碱基序列为靶标序列,采用SEQ ID No.3或SEQ ID No.4所示gRNA序列进行敲除。
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CN116121442A (zh) * | 2023-02-07 | 2023-05-16 | 宝清北方水稻研究中心 | 一种水稻粒型QTL的InDel分子标记SG2-InDel、试剂、试剂盒及其应用 |
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