CN107253980B - OsGRF7基因在水稻株型调控中的应用 - Google Patents
OsGRF7基因在水稻株型调控中的应用 Download PDFInfo
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- CN107253980B CN107253980B CN201710608507.9A CN201710608507A CN107253980B CN 107253980 B CN107253980 B CN 107253980B CN 201710608507 A CN201710608507 A CN 201710608507A CN 107253980 B CN107253980 B CN 107253980B
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Abstract
本发明公开了一种OsGRF7基因在水稻株型调控中的应用,属于作物基因工程技术领域。本发明通过将OsGRF7基因在水稻中过表达后,水稻分蘖减少、茎秆粗壮、粒子变宽、株高降低、千粒重增加;而将OsGRF7基因在水稻中敲除、失活或者降低活性后,分蘖增多、茎秆变细、粒子变窄、千粒重下降,说明该基因可控制水稻的株型。OsGRF7基因还可以作为分子标记用于水稻育种中。OsGRF7基因为利用分子标记辅助育种及利用基因工程的方法培育高产水稻新品种,提供了强有力的手段和工具,具有巨大的应用潜力。
Description
技术领域
本发明涉及植物基因工程领域,特别涉及OsGRF7基因(Oryza sativa Growth-regulating factor 7)在水稻株型调控中的应用。
背景技术
水稻的株型包括分蘖数目、分蘖角度、穗形和株高等性状。良好的株型是提高水稻产量的关键因素,同时也是水稻应对环境的关键因素之一。当前生产上应用的大部分栽培品种为含有半矮杆基因SD1的矮杆品种。与传统的高杆品种相比,矮杆品种具有较多的优势。但是,矮杆品种固有的缺点却限制了其产量的进一步提高,这包括分蘖数较少、穗子较小等缺点。为了克服当前大部分栽培品种增产潜力有限的缺点,进一步满足人们对粮食的需求,国际水稻所的育种学家提出了水稻新株型的概念,新株型的主要特点是分蘖少,没有无效分蘖;穗子大,穗粒数多;茎秆粗壮,抗倒伏。
水稻分蘖数目是水稻生产中的一个重要农艺性状。单位面积的有效分蘖数决定了有效穗数,而有效穗又是决定单位面积上水稻产量的四个关键因素之一。因此,合理的控制水稻分蘖的发生,尽量减少无效分蘖具有重要的生产意义。
抗倒伏能力一直是水稻育种学家十分重视的方面。虽然其不能直接提高水稻的产量,但它对于产量的稳定具有十分重要的作用,是产量进一步提高的限制性因素。与传统高杆品种相比,矮杆品种通过降低株高,提高了植株的抗倒伏能力,是水稻更高产的前提。在这种情况下,改变茎秆的性状,培育更加粗壮,抗倒伏能力更强的品种一直是育种学家努力的目标。
国际水稻所新株型的基本特点为少分蘖、粗杆和大穗。模拟研究表明,在热带地区、干季的条件下,具有新株型品种的产量可以比当前品种再提高25%。因此,阐明分蘖、茎秆和穗子发育的遗传基础和分子机理,对于获得更加高产的品种具有十分重要的意义。当前,尽管已经克隆了很多产量相关的基因,但是能够从整体上改变水稻株型,使之产生新株型特点的基因还未见报道。
发明内容
本发明的目的在于提供水稻生长调节因子基因7(OsGRF7)与水稻分蘖数目、株高、粒径大小,千粒重等株型性状的关系,进而提供OsGRF7在提高水稻产量上的应用。
本发明的目的通过以下技术方案实现:
第一方面,提供一种OsGRF7基因在改变水稻株型中的应用,所述的改变水稻株型包括分蘖数目降低、株高降低、茎秆直径及粒径变宽、千粒重增加;OsGRF7基因编码的蛋白质GRF7的氨基酸序列如下(1)或(2)所示:
(1)SEQ ID NO.1所示的氨基酸序列;
(2)将SEQ ID NO.1所示序列的氨基酸残基经过一个或几个氨基酸残基的取代和/或缺失/或添加,仍能发挥与SEQ ID NO.1所示蛋白质一样的改变水稻株型的作用。
优选地,本发明提供编码SEQ ID NO.1所示的氨基酸序列的cDNA核苷酸序列,如SEQ ID NO.2所示。
优选地,所述的水稻为粤泰B。
在本发明的一个具体实施例中,在水稻体内过表达OsGRF7基因发现:所述转基因水稻与所述目的水稻相比,分蘖减少、茎秆粗壮、粒子变宽、千粒重增加、株高降低。
在本发明的一个具体实施例中,在水稻体内通过携带OsGRF7基因干扰载体的表达,使OsGRF7基因失活或降低活性后,分蘖增多、茎秆变细、粒子变窄、千粒重下降。
第二方面,本发明提供扩增OsGRF7基因全长或其中任一片段的引物在改变水稻株型中的应用。
第三方面,本发明提供含有OsGRF7基因的表达盒、重组载体、转基因细胞系和重组菌在提高水稻产量上的应用。
第四方面,本发明提供一种提高水稻产量的方法,为提高水稻体内OsGRF7基因的表达量,具体为将OsGRF7基因通过表达载体导入到水稻体内使OsGRF7过表达。
具体为,可用现有的植物表达载体构建含有OsGRF7基因的重组表达载体,所述植物表达载体包括双元农杆菌载体和可用于植物瞬时表达的载体等,如pCAMBIA1301-UBI、pCAMBIA1391、pGWB或其他衍生植物表达载体。
使用OsGRF7基因构建重组表达载体时,可在其转录起始核苷酸前加任何一种增强型、组成型、组织特异型或诱导型启动子,如花椰菜花叶病毒(CAMV)35S启动子、泛素(Ubiquitin)基因启动子(pUBI)等,它们可单独使用或与其他的植物启动子结合使用;翻译起始区域可来自转录起始区域或结构基因。
为了便于对转基因植物细胞或植物进行鉴定及筛选,可对所用植物表达载体进行加工,如加入在植物总表达可产生颜色变化的酶或发光化合物的基因(GUS基因、GFP基因、FLAG基因等)、具有抗性的抗生素标记物(庆大霉素标记物、卡那霉素标记物、潮霉素标记物等)。
所述重组表达载体具体可为在植物表达载体pCAMBIA1301的多克隆位点间或者pH7WG2D重组位点插入上述与水稻株型调控基因得到的重组表达载体。
所述与水稻株型相关的基因OsGRF7是通过上述重组表达载体导入目的水稻中的。携带有本发明的与水稻株型调控基因OsGRF7的植物表达载体可通过Ti质粒、Ri质粒、直接DNA转化、农杆菌介导等常规生物学方法转化到水稻细胞或组织中。
第五方面,OsGRF7基因还可以作为分子标记用于水稻育种中。
与现有技术相比,本发明具有如下优势:
本发明通过将OsGRF7基因在水稻中过表达之后,水稻分蘖减少、茎秆粗壮、粒子变宽、千粒重增加,株高降低;而将本发明保护的基因在水稻中失活或降低活性后,分蘖增多、茎秆变细、粒子变窄、千粒重下降,说明该基因可控制水稻的株型。因此,OsGRF7基因为分子标记辅助育种及利用基因工程的方法培育新株型水稻品种,从而进一步提高水稻的产量提供了有力的手段,具有重要的理论意义和巨大的应用潜力。
附图说明
图1水稻OsGRF7基因过表达、干涉和常规籼稻粤泰B的表型。
图2 OsGRF7基因的cDNA和蛋白序列图及miRNA结合OsGRF7转录本的示意图
A、蓝色的核苷酸代表5’和3’非编码区,下滑线所标注的蛋白序列代表QLQ结构域和WRC结构域,红色星号代表miRNA396靶位点;
B、RACE的结果证明miRNA结合了OsGRF7转录本。
图3过表达及干涉的转基因水稻相关农艺性状的比较。
具体实施方式
通过以下详细说明结合附图可以进一步理解本发明的特点和优点。所提供的实施例仅是对本发明方法的说明,而不以任何方式限制本发明揭示的其余内容。
下述实施例中,如无特殊说明,均为常规方法。
下述实施例中水稻材料的田间栽培:水稻种子浸种2天后,30度催芽2天,然后播种,25天后移栽到大田。
【实施例1】转基因植物的获得及其阳性检测
一、水稻基因组DNA的提取:
采用CTAB方法从水稻叶片中提取基因组DNA。取1g水稻叶片,加0.1mlCTAB和一颗钢珠,在植物组织打样机中5min,之后65度水浴1h提取DNA,获得的DNA沉淀溶解于200μl的ddH2O中。
二、转基因植物的获得
1、重组表达载体的构建
1)基因的克隆
利用表2中的引物组合GRF7OE-F/GRF7OE-R从粤泰B的幼穗cDNA中扩增出了OsGRF7的CDS区段,通过回收连接到T载体上得到最终的CDS区段
表2、引物序列
2)构建表达载体
将步骤1)得到的包含全长OsGRF7基因的CDS区段重组到pH7WG2D中,得到重组表达载体GRF7OE,经验证载体构建正确。
2、转基因植物的获得
将质粒GRF7OE通过电击的方法转入农杆菌(Agrobacterium tumefaciens)EHA105中,筛选得到含有重组质粒GRF7OE的重组农杆菌菌株。
用含有重组质粒GRF7OE的重组农杆菌菌株侵染粤泰B愈伤组织,在黑暗处25度共培养3天后,在含有50mg/l潮霉素和400mg/l的头孢的筛选培养基上筛选抗性愈伤进一步得到转基因植株。将潮霉素抗性植株在组培室炼苗3天后移栽到水田中,获得的转基因植株为T0代。收获T0代植株的种子,按照上述田间栽培的方法进行栽培,并通过常规分子检测,获得含GRF7OE的T1代转基因植株。
三、转基因植株的株型检测
1、通过qRT–PCR检测OsGRF7基因的表达量
利用Trizol(购自Invitrogen公司)提取转基因植株和对照粤泰B植株的totalRNA,并利用反转录试剂盒(购自Invitrogen公司)进行反转录,得到cDNA。利用引物GRF7RT-F和GRF7RT-R进行定量检测OsGRF7的表达量。利用引物UBI-F和UBI-R扩增Ubiquitin基因作为内参,引物序列如表3。结果显示转基因植株中OsGRF7表达量增加。(图3)
表3、引物序列
引物名称 | 引物序列(5’-3’) |
GRF7RT-F | AAGTACTGCGAGCGGCACATA |
GRF7RT-R | AGATTGGCCTTCCACATGCT |
UBI-F | ACCACTTCGACCGCCACTACT |
UBI-R | ACGCCTAAGCCTGCTGGTT |
2、转基因植株的株型检测
对GRF7OE的T4代转基因植株、粤泰B对照植株进行株型的统计,每种材料统计15个单株。结果如图3所示,GRF7OE转基因T4代植株与粤泰B对照植株相比,其分蘖数目降低,粒子变宽,株高降低,千粒重增加。
【实施例2】OsGRF7基因干涉载体转基因植株的获得及其检测
一、转基因植株的获得
1、干涉片段的获得
利用OsGRF7全长cDNA作为模板,用表4中的引物对GRF7RNAi-F/GRF7RNAi-R进行PCR扩增,获得的产物进行测序。
表4、引物序列
SEQ ID NO.3是SEQ ID NO.2的第662bp到978bp的317bp片段,见图2A,经全基因组比对分析确认在水稻基因组中无其他同源序列。
2、干涉载体的构建
将引物对GRF7RNAi-F/GRF7RNAi-R扩增得到的产物重组到表达载体pANIC 8A载体中,得到重组表达载体GRF7RNAi,插入片段表达后形成发夹结构。
3、转基因植株的获得
将质粒GRF7RNAi通过电击的方法转入农杆菌(Agrobacterium tumefaciens)EHA105中,筛选得到含有重组质粒GRF7RNAi的重组农杆菌菌株。
用含有重组质粒GRF7RNAi的重组农杆菌菌株侵染粤泰B愈伤组织,在黑暗处25度共培养3天后,在含有50mg/l潮霉素和400mg/l的头孢的筛选培养基上筛选抗性愈伤进一步得到转基因植株。将潮霉素抗性植株在组培室炼苗3天后移栽到水田中,获得的转基因植株为T0代。收获T0代植株的种子,按照上述田间栽培的方法进行栽培,并通过常规分子检测,获得含GRF7RNAi的T1代转基因植株。RACE的结果证明miRNA结合OsGRF7转录本,见图2B。
二、转基因植株的检测
1、通过qRT–PCR检测OsGRF7基因的表达量
利用Trizol(购自Invitrogen公司)提取转基因植株和对照粤泰B植株的totalRNA,并利用反转录试剂盒(购自Invitrogen公司)进行反转录,得到cDNA。利用引物GRF7RT-F和GRF7RT-R进行定量检测OsGRF7的表达量。利用引物UBI-F和UBI-R扩增Ubiquitin基因作为内参,引物序列如表3。结果显示转基因植株中OsGRF7表达量下降。(图3)
2、转基因植株的株型检测
对GRF7RNAi的T4代转基因植株、粤泰B对照植株进行株型的统计,每种材料统计15个单株。结果如图3所示,GRF7RNAi转基因T4代植株与粤泰B对照植株相比,其分蘖数目增加,粒子变窄,株高降低,千粒重下降。
SEQUENCE LISTING
<110> 武汉大学
<120> OsGRF7基因在水稻株型调控中的应用
<160> 11
<170> PatentIn version 3.3
<210> 1
<211> 430
<212> PRT
<213> 水稻(Oryza Sativa)
<400> 1
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Arg Ser Gln Asp Tyr Thr Asn Gln Gln His Asn Ile Leu Gln Asn Asn
225 230 235 240
Thr Lys Gly Asp Asn Trp Ser Glu Glu Met Ser Ser Gln Ala Asp Tyr
245 250 255
Ala Val Ile Pro Ala Gly Ser Leu Met Asn Thr Pro Gln Ser Ala Asn
260 265 270
Leu Asn Pro Ile Pro Gln Gln Gln Arg Cys Lys Gln Ser Leu Phe Gly
275 280 285
Lys Gly Ile Gln His Asp Asp Ile Gln Leu Ser Ile Ser Ile Pro Val
290 295 300
Asp Asn Ser Asp Leu Pro Thr Asn Tyr Asn Lys Ala Gln Met Asp His
305 310 315 320
Val Val Gly Gly Ser Ser Asn Gly Gly Asn Asn Thr Arg Ala Ser Trp
325 330 335
Ile Pro Gly Ser Trp Glu Ala Ser Ile Gly Gly Pro Leu Gly Glu Phe
340 345 350
Phe Thr Asn Thr Ser Ser Ala Ser Asp Asp Lys Gly Lys Ser Arg His
355 360 365
Pro Pro Ser Leu Asn Leu Leu Ala Asp Gly His Thr Thr Ser Pro Gln
370 375 380
Leu Gln Ser Pro Thr Gly Val Leu Gln Met Thr Ser Phe Ser Ser Val
385 390 395 400
Pro Ser Ser Thr Val Ser Ser Pro Ala Gly Ser Leu Cys Asn Gly Leu
405 410 415
Leu Thr Ser Gly Leu Val Asn Ala Gln Thr Val Gln Thr Leu
420 425 430
<210> 2
<211> 1293
<212> DNA
<213> 水稻(Oryza Sativa)
<400> 2
atggcaatgg cgacccctac gaccaacggc agcttccttc ttggatcagg tggctatccc 60
ggtgcccaga ttctaagctt ctcctcctca ggtcacagcg gcaatgggtt ggattgtgga 120
agctcagatg tggcaagaat gcagggggtt ttagcaaggg ttagggggcc attcacacca 180
acacaatgga tggagctgga gcaccaggct ctgatctaca agcacattgt ggcgaatgcg 240
ccggtaccgg ccggcttgct cctccccatc aggagaagcc tccatccacc agtgttccca 300
cacttctcct ctggtggcat tcttggctcc agctccttgg gatgggggtc atttcagctg 360
ggctattctg ggagtgctga ctccgagccc gggagatgcc gtcgaaccga tggcaagaaa 420
tggcggtgct cgagagacgc agttgtcgac caaaagtact gcgagcggca cataaaccgg 480
ggtcgccacc gttcaagaaa gcatgtggaa ggccaatcta gccatgccgc aaaagcaacg 540
gttcccgcca tagcacaacc acccattggt gcatctaatg gcaaattgtc aggcagccat 600
ggtgtgtcaa atgagctcac gaaaaccttg gctactaaca ggatgatgtt ggataaagca 660
aatcttattg aacgctccca ggactacact aatcagcaac acaacatcct acagaacaac 720
acaaaaggtg ataattggtc tgaagagatg tcctcacaag cagactatgc agtaatccct 780
gctggctctc tcatgaacac accgcaatcg gcgaatttaa atccaattcc ccagcaacaa 840
cgctgtaagc agtcactctt tggcaaaggg atacagcatg atgacattca gctgtcgata 900
tccattcccg tggataactc cgacttaccc actaactaca acaaggctca aatggaccat 960
gtagtaggcg gttcatcgaa tggcggaaac aacacgcgag caagttggat accgggctcc 1020
tgggaagcgt ccataggtgg acctctgggt gagttcttca ccaacaccag cagcgcatca 1080
gacgacaaag gcaaaagccg ccacccgcca tctttgaacc tcttagctga tggacatact 1140
acaagtccac agctgcaatc gcccaccgga gtcctgcaga tgactagctt cagttcagtg 1200
cccagcagca ctgttagtag tcctgcaggc agcctctgca atggcttgct cacttcaggc 1260
ctggtgaatg cccagactgt ccaaacactg tga 1293
<210> 3
<211> 317
<212> DNA
<213> 水稻(Oryza Sativa)
<400> 3
atcttattga acgctcccag gactacacta atcagcaaca caacatccta cagaacaaca 60
caaaaggtga taattggtct gaagagatgt cctcacaagc agactatgca gtaatccctg 120
ctggctctct catgaacaca ccgcaatcgg cgaatttaaa tccaattccc cagcaacaac 180
gctgtaagca gtcactcttt ggcaaaggga tacagcatga tgacattcag ctgtcgatat 240
ccattcccgt ggataactcc gacttaccca ctaactacaa caaggctcaa atggaccatg 300
tagtaggcgg ttcatcg 317
<210> 4
<211> 59
<212> DNA
<213> 人工序列
<400> 4
ggggacaagt ttgtacaaaa aagcaggctt cgaaggagat aggatggcaa tggcgaccc 59
<210> 5
<211> 55
<212> DNA
<213> 人工序列
<400> 5
ggggaccact ttgtacaaga aagctgggtc tcacagtgtt tggacagtct gggca 55
<210> 6
<211> 21
<212> DNA
<213> 人工序列
<400> 6
aagtactgcg agcggcacat a 21
<210> 7
<211> 20
<212> DNA
<213> 人工序列
<400> 7
agattggcct tccacatgct 20
<210> 8
<211> 21
<212> DNA
<213> 人工序列
<400> 8
accacttcga ccgccactac t 21
<210> 9
<211> 19
<212> DNA
<213> 人工序列
<400> 9
acgcctaagc ctgctggtt 19
<210> 10
<211> 49
<212> DNA
<213> 人工序列
<400> 10
ggggacaagt ttgtacaaaa aagcaggctt catcttattg aacgctccc 49
<210> 11
<211> 46
<212> DNA
<213> 人工序列
<400> 11
ggggaccact ttgtacaaga aagctgggtc cgatgaaccg cctact 46
Claims (4)
1. 一种OsGRF7基因在改变水稻株型中的应用,其特征在于,所述的改变水稻株型包括分蘖数目降低、株高降低、茎秆直径变宽;OsGRF7基因编码的蛋白质GRF7的氨基酸序列如SEQ ID NO.1所示的氨基酸序列。
2.根据权利要求1所述的OsGRF7基因在改变水稻株型中的应用,其特征在于,编码蛋白质GRF7的氨基酸序列SEQ ID NO.1的cDNA核苷酸序列,如SEQ ID NO.2所示。
3.根据权利要求1所述的OsGRF7基因在改变水稻株型中的应用,其特征在于,所述的水稻为粤泰B。
4.含有OsGRF7 基因的表达盒、重组载体、转基因细胞系和重组菌在改变水稻株型中的应用,其特征在于,所述的改变水稻株型包括分蘖数目降低、株高降低、茎秆直径变宽。
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CN114940708B (zh) * | 2022-04-08 | 2023-09-19 | 扬州大学 | OsPIL16基因或其蛋白在调控水稻茎粗中的应用 |
Citations (2)
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CN101855355A (zh) * | 2007-09-14 | 2010-10-06 | 巴斯夫植物科学有限公司 | 具有提高的产量相关性状的植物和用于制备该植物的方法 |
CN105647940A (zh) * | 2014-11-11 | 2016-06-08 | 武汉大学 | OsGRF6基因提高水稻产量的方法及其应用 |
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CN101855355A (zh) * | 2007-09-14 | 2010-10-06 | 巴斯夫植物科学有限公司 | 具有提高的产量相关性状的植物和用于制备该植物的方法 |
CN105647940A (zh) * | 2014-11-11 | 2016-06-08 | 武汉大学 | OsGRF6基因提高水稻产量的方法及其应用 |
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Arabidopsis growth-regulating factor7 functions as a transcriptional repressor of abscisic acid- and osmotic stress-responsive genes, including DREB2A;Kim JS等;《Plant Cell》;20120831;第24卷(第8期);3393-3405 * |
Growth-Regulating Factors (GRFs): A Small Transcription Factor Family with Important Functions in Plant Biology;Omidbakhshfard MA等;《Mol Plant》;20150122;第8卷(第7期);998-1010 * |
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