CN110499326B - Rgga在调控作物农艺性状中的应用 - Google Patents
Rgga在调控作物农艺性状中的应用 Download PDFInfo
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- CN110499326B CN110499326B CN201910645392.XA CN201910645392A CN110499326B CN 110499326 B CN110499326 B CN 110499326B CN 201910645392 A CN201910645392 A CN 201910645392A CN 110499326 B CN110499326 B CN 110499326B
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Abstract
本发明提供RGGA在调控作物农艺性状中的应用,通过比较拟南芥RGGA突变体与野生型的生长状况发现,苗期在添加2%蔗糖的培养基上短期培养后,突变体的营养生长指标(莲座叶大小、叶绿素含量、地上部及地下部干鲜重)及种子产量指标(单株果荚数、单果荚种子数、百粒重)均明显优于野生型,且生长早期短暂供糖就能保持其整个生育期的生长优势。RGGA基因在单子叶、双子叶植物中普遍存在,在水稻、玉米等作物中也表现出明显优于野生型的生长表型。因此,通过敲除RGGA同源基因,结合早期供糖培养有望显著提高作物产量,对于RGGA功能和作用机制的分析为获得高产作物新品种提供了理论和方法基础,因此具有良好的实际应用之价值。
Description
技术领域
本发明属于分子生物学与生物技术领域,具体涉及RGGA在调控作物农艺性状中的应用。
背景技术
公开该背景技术部分的信息仅仅旨在增加对本发明的总体背景的理解,而不必然被视为承认或以任何形式暗示该信息构成已经成为本领域一般技术人员所公知的现有技术。
无论真核还是原核细胞都存在糖抑制现象,即糖的积累对大量基因的表达有抑制作用。蔗糖作为植物光合作用的终产物,在细胞质中合成后经韧皮部向其他组织、器官供应碳源和能量,是植物光合作用同化产物运输的主要形式,可以通过反馈抑制参与“源”和“库”之间的平衡调节,满足植物的生长及抗逆性的需要。早期研究证明,添加蔗糖降低藻类和植物组织培养中的光合速率和叶绿素积累,相反,糖的消耗提高了植物光合作用、分解代谢和碳水化合物运输过程中各种基因的表达。另一方面蔗糖作为信号分子,在植物受到机械损伤、病菌侵染等时,激活或阻遏多种基因的表达。
蔗糖在植物生长发育过程中有着举足轻重的作用,在外加蔗糖的培养条件下,植物选择性地激活或者抑制某些基因的表达。蔗糖调控基因广泛参与植物生命周期的各个阶段,如种子萌发、营养生长、生殖生长、植物衰老和胁迫应答等。因此,研究植物蔗糖代谢的机理及植物蔗糖代谢相关基因的表达调控,对于提高作物产量、培育高效利用蔗糖的作物新品种具有重要的理论和应用价值。
在真核生物中,RNA结合蛋白(RBPs)是一类重要的转录后调控因子,通过与RNA结合形成核糖核蛋白复合物来调节真核生物细胞的RNA代谢过程,包括RNA的转移、修饰、翻译及降解。RNA结合蛋白广泛存在于动物、植物以及微生物中,约占真核生物基因编码蛋白的2%-8%。RNA结合蛋白不仅在植物中大量存在,而且作为重要的调控因子在RNA代谢、生长发育以及应激反应过程中发挥重要作用,已引起人们的广泛关注。相对于动物RNA结合蛋白的大量研究,植物RNA结合蛋白的功能研究还相对较少。近几年,关于植物RNA结合蛋白的功能、作用机制以及与其它蛋白相互关系的研究逐渐增多。已有研究表明,拟南芥、水稻和小麦等模式植物或经济作物的RBPs在RNA代谢、生长发育以及逆境胁迫响应过程中发挥重要作用,从而为基础研究和农业生产实践提供了重要的理论依据。
发明内容
针对上述现有技术,本发明提供RGGA在调控作物农艺性状中的应用。发明人意外发现通过敲除或抑制作物RGGA同源基因,结合早期供糖培养能够显著提升作物农艺性状,促进作物生长发育,提高作物产量,从而能够为作物增产育种提供新策略,因此本发明具有良好的实际应用之价值。
本发明是通过如下技术方案实现的:
本发明的第一个方面,提供调控RGGA基因表达在提升作物农艺性状中的应用。
所述作物包括但不限于拟南芥、烟草、水稻和玉米;
进一步的,所述调控为负调控;即通过负调控RGGA基因的表达,达到提高农作物农艺性状的目的。
本发明的第二个方面,提供一种提升作物农艺性状的方法,所述方法为:通过抑制作物RGGA基因表达和/或活性降低和/或失活,实现提升作物农艺性状。
进一步的,所述提升作物农艺性状的方法还包括在作物生长发育早期(如苗期)对作物进行糖培养处理;
进一步的,所述糖培养处理为向作物施加蔗糖。
本发明的第三个方面,提供抑制作物RGGA基因表达和/或活性降低和/或失活的物质在提升作物农艺性状的应用。
本发明的第四个方面,提供一种作物生长促进剂,其活性成分为抑制作物RGGA基因表达和/或活性降低和/或失活的物质。
本发明有益效果:本发明通过比较拟南芥RGGA突变体(rgga)与野生型(Col-0)的生长状况发现,苗期在添加2%蔗糖的培养基上短期培养后,突变体的营养生长指标(莲座叶大小、叶绿素含量、地上部及地下部干鲜重)及种子产量指标(单株果荚数、单果荚种子数、百粒重)均明显优于野生型,且生长早期短暂供糖就能保持其整个生育期的生长优势。RGGA基因在单子叶、双子叶植物中普遍存在,分别利用RNAi技术在本生烟、水稻中下调RGGA和CRISPR/Cas9基因编辑技术在玉米中敲除RGGA,获得的RGGA下调和敲除植株,生长早期供糖培养,也表现出明显优于野生型的生长表型。因此,通过敲除RGGA同源基因,结合早期供糖培养有望显著提高作物产量,对于RGGA功能和作用机制的分析为获得高产作物新品种提供了理论和方法基础,因此具有良好的实际应用之价值。
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为本发明实施例1中AtRGGA缺失对拟南芥生长的影响图;(A)在不添加蔗糖和添加1%、2%、3%、4%、5%、6%蔗糖的MS培养基上播种并生长2w后的Col-0和rgga表型;(B)在2%蔗糖MS培养基生长2w后,移入蛭石中生长到3、4、5、6w苗龄的Col-0和rgga的表型;(C-G)依次为蔗糖添加培养Col-0和rgga的叶绿素含量、地上部分鲜重和干重、地下部分鲜重和干重的差异分析图;(H-J)依次为蔗糖添加培养2w的Col-0和rgga的单株果荚数、单果荚种子数和百粒重的差异分析图。w:周;d:天。
图2为本发明实施例1中蔗糖处理不同天数的4周苗龄Col-0和rgga表型差异图;具体的,2%蔗糖MS培养基上播种并生长2d、4d、5d和6d后移入蛭石中生长至4周苗龄的Col-0和rgga。d:天。
图3为本发明实施例2中蔗糖对本生烟NbRGGA RNAi株系生长的影响图;(A)2%蔗糖MS培养基培养14d的表型图;(B)qRT-PCR检测转基因株系NbRGGA表达水平图。d:天。
图4为本发明实施例3中蔗糖对水稻OsRGGA RNAi株系生长的影响图;(A)水稻wt及OsRGGARNAi株系2%蔗糖MS培养基上生长6d的表型图;(B)转基因水稻OsRGGA表达水平检测图。wt:野生型;d:天。
图5为本发明实施例4中蔗糖对玉米ZmRGGA敲除株系生长的影响图;具体为2%蔗糖MS培养基上生长6天的表型。
具体实施方式
应该指出,以下详细说明都是例示性的,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、器件、组件和/或它们的组合。应理解,本发明的保护范围不局限于下述特定的具体实施方案;还应当理解,本发明实施例中使用的术语是为了描述特定的具体实施方案,而不是为了限制本发明的保护范围。下列具体实施方式中如果未注明具体条件的实验方法,通常按照本领域技术内的分子生物学的常规方法和条件,这种技术和条件在文献中有完整解释。参见例如Sambrook等人,《分子克隆:实验手册》中所述的技术和条件,或按照制造厂商所建议的条件。
本发明的一个具体实施方式中,提供调控RGGA基因表达在提升作物农艺性状中的应用。
本发明的又一具体实施方式中,所述作物包括但不限于拟南芥、烟草、水稻和玉米;
其中,
拟南芥(Arabidopsis thaliana)AtRGGA基因序列如SEQ ID NO.1所示,AtRGGA基因编码氨基酸序列如SEQ ID NO.5所示。
本生烟(Nicotiana benthamiana)NbRGGA基因序列如SEQ ID NO.2所示,NbRGGA基因编码氨基酸序列如SEQ ID NO.6所示。
水稻(Oryza latifolia)OsRGGA基因序列如SEQ ID NO.3所示,OsRGGA基因编码氨基酸序列如SEQ ID NO.7所示。
玉米(Zea mays)ZmRGGA基因序列如SEQ ID NO.4所示,ZmRGGA基因编码氨基酸序列如SEQ ID NO.8所示。
本发明的又一具体实施方式中,所述调控为负调控;即通过负调控RGGA基因的表达,达到提高农作物农艺性状的目的。
本发明的又一具体实施方式中,所述负调控具体为通过RNAi或CRISPR/Cas9下调RGGA基因表达,从而提升作物农艺性状。
本发明的又一具体实施方式中,所述农艺性状包括但不限于营养生长指标性状和/或产量指标性状;
本发明的又一具体实施方式中,所述营养生长指标性状包括但不限于莲座叶大小、叶绿素含量、作物地上部及地下部干鲜重;
本发明的又一具体实施方式中,所述产量指标性状包括但不限于单株果荚数、单果荚种子数、百粒重。
本发明的又一具体实施方式中,提供一种提升作物农艺性状的方法,所述方法为:通过抑制作物RGGA基因表达和/或活性降低和/或失活,实现提升作物农艺性状。
本发明的又一具体实施方式中,所述抑制作物RGGA基因表达和/或活性降低和/或失活的方法可以为RNAi或CRISPR/Cas9。
本发明的又一具体实施方式中,所述提升作物农艺性状的方法还包括在作物生长发育早期(如苗期)对作物进行糖培养处理;
本发明的又一具体实施方式中,所述糖培养处理为向作物施加蔗糖。
本发明的又一具体实施方式中,提供抑制作物RGGA基因表达和/或活性降低和/或失活的物质在提升作物农艺性状的应用。
本发明的又一具体实施方式中,所述抑制作物RGGA基因表达和/或活性降低和/或失活的物质,包括但不限于RGGA蛋白特异的抗体、针对RGGAmRNA的RNA干扰分子或反义寡核苷酸、小分子抑制剂或siRNA等。
本发明的又一具体实施方式中,所述作物包括但不限于拟南芥、烟草、水稻和玉米;
本发明的又一具体实施方式中,所述农艺性状包括但不限于营养生长指标性状和/或产量指标性状;
本发明的又一具体实施方式中,所述营养生长指标性状包括但不限于莲座叶大小、叶绿素含量、作物地上部及地下部干鲜重;
本发明的又一具体实施方式中,所述产量指标性状包括但不限于单株果荚数、单果荚种子数、百粒重。
本发明的又一具体实施方式中,提供一种作物生长促进剂,其活性成分包含抑制作物RGGA基因表达和/或活性降低和/或失活的物质。
以下通过实施例对本发明做进一步解释说明,但不构成对本发明的限制。应理解这些实施例仅用于说明本发明而不用于限制本发明的范围。另外,实施例中未详细说明的分子生物学方法均为本领域常规的方法,具体操作可参看分子生物指南或产品说明书。
实施例1蔗糖处理拟南芥Col-0和rgga突变体表型观察
1、野生型和突变体拟南芥种子在含2%蔗糖的培养基中生长14天后,rgga地上部和地下部的干鲜重都明显高于野生型,叶绿素含量也高于野生型,另外种子产量的相关指标(包括百粒重、单果荚种子数、单株果荚数)均优于野生型(图1)。
2、野生型和突变体拟南芥种子在含2%蔗糖的培养基中,生长2、4、5、6天后转移到蛭石中生长至4周。生长状态如图2,经蔗糖处理5天、6天的突变体莲座叶明显比野生型大。
实施例2烟草NbRGGA RNAi株系的获得及表型观察
RNAi指由双链RNA(double-stranded RNA,dsRNA)诱发的、同源mRNA高效特异性降解的现象。选取NbRGGA cDNA靠近5'端的300bp序列设计引物,保证该序列在本生烟中是高度特异的。
引物序列
NbRGGA RNAi-F1:CTGCAG GCGTGGAGGAGGACGTGGGGG(SEQ ID NO.9);
NbRGGA RNAi-R1:GGATCC CCCACGAGGTCCGCCATATCCAC(SEQ ID NO.10);
NbRGGA RNAi-F2:GAGCTC GCGTGGAGGAGGACGTGGGGG(SEQ ID NO.11);
NbRGGA RNAi-R2:GGATCC CCCACGGCCAGCCACTTCGTCG(SEQ ID NO.12)。
通过农杆菌介导转化的方法获得NbRGGA RNAi本生烟株系(NbRGGAi),并通过qRT-PCR的方法检测下调效率,选取NbRGGA表达量下调70%以上的三个株系进行下一步试验。
获得的三个纯合NbRGGAi株系与野生型本生烟,在含有2%蔗糖培养基中生长14d后,观察表型,如图3,NbRGGA下调株系长势明显优于野生型。
实施例3水稻OsRGGA RNAi株系的获得及表型观察
1、OsRGGA RNAi载体构建
选取OsRGGA cDNA靠近5'端的500bp序列设计引物,保证该序列在水稻中是高度特异的。
引物序列
OsRGGA RNAi-F1:CTGCAGGAAGACAAGGAGATGAC(SEQ ID NO.13);
OsRGGA RNAi-R1:GGATCC GCGCTTGATCTTC(SEQ ID NO.14);
OsRGGA RNAi-F2:GAGCTCGAAGACAAGGAGATGAC(SEQ ID NO.15);
OsRGGA RNAi-R2:GGATCC CACCTGGGCTGTAG(SEQ ID NO.16);
2、OsRGGA RNAi水稻株系的获得
通过农杆菌介导转化的方法获得OsRGGA RNAi水稻株系(OsRGGAi),并通过qRT-PCR、ELISA和Western blot的方法检测下调效率,选取OsRGGA表达量下调70%以上的三个株系进行下一步试验。
3、OsRGGA RNAi株系表型及生物量检测
获得的三个纯合OsRGGAi株系与野生型水稻,在含有2%蔗糖培养基中生长10d后,观察表型,如图4,OsRGGA下调株系长势明显优于野生型。
实施例4玉米ZmRGGA敲除株系的获得与表型观察
1、利用CRISPR/Cas9技术敲除ZmRGGA
CRISPR/Cas9技术是指crRNA(CRISPR-derived RNA)通过碱基配对与tracrRNA(trans-activating RNA)结合形成tracrRNA/crRNA复合物,此复合物引导核酸酶Cas9蛋白再与crRNA配对的序列靶位点剪切双链DNA。而通过人工设计这两种RNA,可以改造形成具有引导作用的sgRNA(single-guide RNA),足以引导Cas9对DNA的定点切割。登录网站http://www.genome.arizona.edu/crispr/CRISPRsearch.html筛选靶点,并利用http://www.rgenome.net/cas-offinder/网站评估脱靶情况,筛选出合适的靶点后设计引物。
引物序列
MT1T2-BsF:AATAATGGTCTCAGGCGCCCGCCAGCCTTACCGTGG(SEQ ID NO.17);
MT1T2-F0:GCCCGCCAGCCTTACCGTGGGTTTTAGAGCTAGAAATAGC(SEQ ID NO.18);
MT1T2-R0:AACCCAAAGTCATCTCCTTATCGCTTCTTGGTGCC(SEQ ID NO.19);
MT1T2-BsR:ATTATTGGTCTCTAAACAACCCAAAGTCATCTCCTTAT(SEQ ID NO.20)。
PCR扩增:以pCBC-MT1T2为模板进行四引物PCR扩增。-BsF/-BsR为正常引物浓度;-F0/-R0稀释20倍。纯化回收PCR产物,建立如下酶切-连接体系:
取5μL连接产物转化大肠杆菌感受态,Kan板筛选。OsU3-FD3和TaU3-RD2菌落PCR鉴定,OsU3-FD3和TaU3-FD2测序确认。用于菌落PCR及测序鉴定的引物:
OsU3-FD3:GACAGGCGTCTTCTACTGGTGCTAC(测序引物,SEQ ID NO.21);
TaU3-RD2:CTCACAAATTATCAGCACGCTAGTC(SEQ ID NO.22);
TaU3-FD2:TTGACTAGCGTGCTGATAATTTGTG(SEQ ID NO.23);
OsU3-RD3:CACGGAGTTCAAATTCCCACAG(测序引物,SEQ ID NO.24)。
2、玉米ZmRGGA敲除株系的获得与鉴定
通过农杆菌介导转化的方法获得玉米ZmRGGA敲除株系(ZmRGGA),并通过ELISA(表1)和Western blot的方法鉴定是否敲除成功。
3、ZmRGGA敲除株系的表型
获得的ZmRGGA敲除株系与野生型玉米,在含有2%蔗糖培养基中生长6d后,观察表型,如图5,ZmRGGA敲除株系长势明显优于野生型。
表1玉米ZmRGGA敲除株系的ELISA检测
wt,wild type;nc,negative control
应注意的是,以上实例仅用于说明本发明的技术方案而非对其进行限制。尽管参照所给出的实例对本发明进行了详细说明,但是本领域的普通技术人员可根据需要对本发明的技术方案进行修改或者等同替换,而不脱离本发明技术方案的精神和范围。
SEQUENCE LISTING
<110> 山东农业大学
<120> RGGA在调控作物农艺性状中的应用
<130>
<160> 24
<170> PatentIn version 3.3
<210> 1
<211> 1068
<212> DNA
<213> 拟南芥(Arabidopsis thaliana)AtRGGA基因
<400> 1
atggcaactt tgaacccttt tgatttgttg gatgatgatg ctgaggatcc aagccagctc 60
gctgttgcca tcgagaagat tgataagtcc aagaaatctg gacaggtttc gagcttgcct 120
gctaagtcag ctcctaagct tccatcgaag ccacttcctc ctgctcaagc cgtgagagag 180
gcaaggagtg atgctccacg tggtggtgga ggccgtggag gatttaaccg tggtcgtggt 240
ggttacaacc gtgatgatgg taacaatgga tattcagggg gatacactaa accctcaggt 300
gaaggagatg tttcaaagtc ttcttacgag aggcgtggcg gtggtggtgc tcctcgtggt 360
tccttccgtg gtgaaggtgg tggacctggt ggtggtcgtc gtggtggatt cagcaatgag 420
ggtggtgatg gggaacgtcc tcgaagggcc tttgagcgtc gtagtggaac tggcagaggg 480
agtgacttca agcgtgacgg atctggtcgt gggaattggg gaactccagg ggaagagata 540
gctgctgaga ctgaagcagt agctggtgtt gagactgaga aggatgttgg agagaagcca 600
gctgttgatg atgtagctgc tgatgctaac aaggaggata ctgttgttga ggagaaagag 660
cctgaggata aggaaatgac tcttgatgag tatgagaaaa tactcgagga gaagaaaaag 720
gcacttcaat cattaactac ctctgagagg aaagttgata cgaaagtgtt tgaatcaatg 780
caacaactgt caaacaagaa gtctaatgat gaaatcttca tcaagctggg ttctgataag 840
gacaaacgca aagatgacaa agaagagaag gctaagaagg ctgtgagcat caatgagttt 900
ctgaagccag cagagggtgg gaactactac cgaggaggtc gtggtggccg tggacgtggt 960
ggtcgtggcc gtggaggtgt ttctagtggc gaatctggtg gttaccgtaa tgaagctgca 1020
ccggcaattg gagatgctgc tcagttccca tctcttgggg gcaagtaa 1068
<210> 2
<211> 1203
<212> DNA
<213> 本生烟(Nicotiana benthamiana)NbRGGA基因
<400> 2
atggctacac tgaatccatt tgacttattg gatgatgatg ctgaggatcc aagtttggat 60
gagttgaaac tggctcaagc tcaggcttca gctgctgctt ctgtacccaa gaaatcgggc 120
tcagttcagg cccaagccca gcccgttaag cctgctgcta agttgccttc caaacccgtt 180
ccaccttctc aggctgtaag ggatgcaagg agtgatagcc agcgtggagg aggacgtggg 240
ggtccacgtg ggggaggccg tggacgtgcg cctgggcgtg gacgtgggtt taaccaagaa 300
tctgctgatg atgagaatgc ttttggcagc aacaatggat tctctgggcg ctatagagtt 360
caagaagatg gagagtcagg aaagctctct gaaaggaaag gtggatatgg cggacctcgt 420
ggggggttcc gtggaggtcg tcgtggtggt ttcagtaatg gagatgctgt agaaggagaa 480
ggggaacggc cacggagagt gtttgatcga cgaagtggca ctggccgtgg gaatgagtat 540
attaaacggg agggttctgg tcgtggtaat tggggaactc ctgcagatga tattgcacaa 600
gagactgaag tggctgttaa tgatggcgag aagattgttg agactgagaa agaagctggg 660
caggaagatg ctgaagacac caataagaat tctactgctg ctgagccaga agagaaggaa 720
cctgaggaga aggagatgac ccttgaggag tatgagaagc taatggaaga gaagaggaag 780
gcgttgatgg ctctaaagcc tgaggaaaga aaggagatga cccttgagga gtatgagaag 840
ctaatggaag agaagaggaa ggcgttgatg gctctaaagc ctgaggaaag aaaggttaat 900
ttggacaaag aactccaatc catgcaactt ctctcaaaca agaaaaatga tgacgaaatc 960
ttcattaaat tgcgttccga gaaagaaaag agaaaagagg cagtggaaaa agccagaaag 1020
actcaaagca taaacgagtt cctgaagcct gcagagggag aaagtaacta tcgccgtggt 1080
ggtcgtggca ggggacaggg ccgtggaaga ggtggttttg gtggtggtgt tggtggaaac 1140
aaaaactatt cagccccatc tattgaagat gctggccagt tcccgtcctt ggttgccaaa 1200
taa 1203
<210> 3
<211> 1113
<212> DNA
<213> 水稻(Oryza latifolia)OsRGGA基因
<400> 3
atggtgacga acaaccagtt cgacctgctc gtcgatgtcg acaatgacga cccctcgcat 60
ctgatcgccg ccgccgagaa gatgaagaag gcggccgcgg ccgcgtcgcc cgcgccggtc 120
gtccaggcaa ggctgcccac caagccgccg ccgcccgcgc aggctgtaaa ggagtccagg 180
aactatggtg ctccagcccg cgatggggca ggccagaatg gaccagggca tgccaatggt 240
ggtttccgcg gaggcaggat gggccaaagg cgtgattttg gtgagggtga taccaatggt 300
gctagcggtt ttagggatgg aatagtgagg agggaggaag gtgagcgcag gccttcagaa 360
aggggccgtg gaccgcgcca gccttaccgt ggtggcggcc gacgtggtgg ctacactggt 420
ggggaggctg gggatgagtc tggccgtgcc ccttaccgtg cctatgaacg tcgcagcggc 480
acaggccgag ggtacgaaac gaaacgtgaa ggagcaggcc gtggtaactg gggaactgtc 540
accgatgaat ccctcgccca ggaatctggt gaggttgtta atattgaagt ggctgctgcc 600
gtgactgagg atgaaaccaa gcaggaagat gtgccacaat ctgaagttga gaaacgcaag 660
gagggtgaat caaatgaaga ggaagaaaag gaacctgaag ataaggagat gactttggag 720
gaatatgaga aagtactgga ggagaagagg aaagctttgc tttcactaaa ggctgaggag 780
agaaaggttg ttgtagacaa ggaattgcag tcaatgcaac agctgtcagt gaagaaggat 840
tctgatgagg tgttcatcaa gctgggttct gataaggaca aaaagaaaga aaatgttgaa 900
agagatgaac gcaccaggaa gtcccttagc atcaatgaat tcctgaagcc ggctgaaggt 960
gaaagatact acagcccagg tgggcgtggg cgtgggcgtg gtagaggccg tggtgaccgt 1020
ggaggattcc gggatggtta tagctctcga ggaccggttg ctgcaccggc aattgaagat 1080
caagcacagt ttcctagctt agctgggaaa tga 1113
<210> 4
<211> 1095
<212> DNA
<213> 玉米(Zea mays)ZmRGGA基因
<400> 4
atggcatcga gtaaccaatt cgatctactt ggagacgtcg agaacgacga ccctgcccac 60
ctcctcgcag ctgccgagaa gaaggcggcc gcggtgccta agcccgcgcc cgcgccggcc 120
aagttgccca ccaagcctcc gccgcccgcg caacctgtga gggaggaaag gaactatggt 180
gctccaccgc gcgatggtcc aggacgtgga agaggcggcc gcggtggcag gactggcccg 240
aggcgagatt ttggcgatgc tgatgcccac ggctttgagg gggggtatgg tggcggcggc 300
ggcggttttg gggacggtgg agttgctcgc ggtgaaaatg gtgagggtag gcaggcagag 360
aggggccgtg ggccccgcca gccttaccgt ggaggtgggc gacgtggtgg gtacagcgat 420
gggcaggctg ggaatgagtt cggacgtcct cgccgggcat atgagcgcca cagcggcaca 480
ggccgtggct acgggatgaa gcgtgaaggg gctgggcgtg gaaattgggg aactgcaact 540
gatgaaggcc ttgaacagga cactgtggat gctgttaatc ctgaggagac tactgccgtg 600
gtggaggatg aaaagaaacc tgaagatgca ccacagtctg aggttgagaa ggacaaggag 660
ggcgcggaga atgaagaaga aaaggaacct gaggataagg agatgacttt ggaggaatat 720
gaaaaggtac tggaggaaaa acgaaaagct ttactctccc ttaaggctga ggagagaaaa 780
gttgaaattg acaaggagct acaggccatg caacaactgt cagttaaaaa ggatggcgat 840
gaagtgttta tcaagctggg ctctgacaag gacttgaaaa agaaagaaag tgctgaaaga 900
gaagaacggg gcaagaagtc cctgagcatc aacgagttct tgaagccagc tgaaggcgag 960
aggtactata accctagtgg ccgtggccgt ggtaggggcc gtggggggtt ccgtggtggt 1020
tacaacggcg gttatcgggg accagctgct gctccagcga tcgaagacca ggctcagttc 1080
cctgccctgg cctaa 1095
<210> 5
<211> 355
<212> PRT
<213> 拟南芥(Arabidopsis thaliana)AtRGGA基因编码氨基酸
<400> 5
Met Ala Thr Leu Asn Pro Phe Asp Leu Leu Asp Asp Asp Ala Glu Asp
1 5 10 15
Pro Ser Gln Leu Ala Val Ala Ile Glu Lys Ile Asp Lys Ser Lys Lys
20 25 30
Ser Gly Gln Val Ser Ser Leu Pro Ala Lys Ser Ala Pro Lys Leu Pro
35 40 45
Ser Lys Pro Leu Pro Pro Ala Gln Ala Val Arg Glu Ala Arg Ser Asp
50 55 60
Ala Pro Arg Gly Gly Gly Gly Arg Gly Gly Phe Asn Arg Gly Arg Gly
65 70 75 80
Gly Tyr Asn Arg Asp Asp Gly Asn Asn Gly Tyr Ser Gly Gly Tyr Thr
85 90 95
Lys Pro Ser Gly Glu Gly Asp Val Ser Lys Ser Ser Tyr Glu Arg Arg
100 105 110
Gly Gly Gly Gly Ala Pro Arg Gly Ser Phe Arg Gly Glu Gly Gly Gly
115 120 125
Pro Gly Gly Gly Arg Arg Gly Gly Phe Ser Asn Glu Gly Gly Asp Gly
130 135 140
Glu Arg Pro Arg Arg Ala Phe Glu Arg Arg Ser Gly Thr Gly Arg Gly
145 150 155 160
Ser Asp Phe Lys Arg Asp Gly Ser Gly Arg Gly Asn Trp Gly Thr Pro
165 170 175
Gly Glu Glu Ile Ala Ala Glu Thr Glu Ala Val Ala Gly Val Glu Thr
180 185 190
Glu Lys Asp Val Gly Glu Lys Pro Ala Val Asp Asp Val Ala Ala Asp
195 200 205
Ala Asn Lys Glu Asp Thr Val Val Glu Glu Lys Glu Pro Glu Asp Lys
210 215 220
Glu Met Thr Leu Asp Glu Tyr Glu Lys Ile Leu Glu Glu Lys Lys Lys
225 230 235 240
Ala Leu Gln Ser Leu Thr Thr Ser Glu Arg Lys Val Asp Thr Lys Val
245 250 255
Phe Glu Ser Met Gln Gln Leu Ser Asn Lys Lys Ser Asn Asp Glu Ile
260 265 270
Phe Ile Lys Leu Gly Ser Asp Lys Asp Lys Arg Lys Asp Asp Lys Glu
275 280 285
Glu Lys Ala Lys Lys Ala Val Ser Ile Asn Glu Phe Leu Lys Pro Ala
290 295 300
Glu Gly Gly Asn Tyr Tyr Arg Gly Gly Arg Gly Gly Arg Gly Arg Gly
305 310 315 320
Gly Arg Gly Arg Gly Gly Val Ser Ser Gly Glu Ser Gly Gly Tyr Arg
325 330 335
Asn Glu Ala Ala Pro Ala Ile Gly Asp Ala Ala Gln Phe Pro Ser Leu
340 345 350
Gly Gly Lys
355
<210> 6
<211> 400
<212> PRT
<213> 本生烟(Nicotiana benthamiana)NbRGGA基因编码氨基酸
<400> 6
Met Ala Thr Leu Asn Pro Phe Asp Leu Leu Asp Asp Asp Ala Glu Asp
1 5 10 15
Pro Ser Leu Asp Glu Leu Lys Leu Ala Gln Ala Gln Ala Ser Ala Ala
20 25 30
Ala Ser Val Pro Lys Lys Ser Gly Ser Val Gln Ala Gln Ala Gln Pro
35 40 45
Val Lys Pro Ala Ala Lys Leu Pro Ser Lys Pro Val Pro Pro Ser Gln
50 55 60
Ala Val Arg Asp Ala Arg Ser Asp Ser Gln Arg Gly Gly Gly Arg Gly
65 70 75 80
Gly Pro Arg Gly Gly Gly Arg Gly Arg Ala Pro Gly Arg Gly Arg Gly
85 90 95
Phe Asn Gln Glu Ser Ala Asp Asp Glu Asn Ala Phe Gly Ser Asn Asn
100 105 110
Gly Phe Ser Gly Arg Tyr Arg Val Gln Glu Asp Gly Glu Ser Gly Lys
115 120 125
Leu Ser Glu Arg Lys Gly Gly Tyr Gly Gly Pro Arg Gly Gly Phe Arg
130 135 140
Gly Gly Arg Arg Gly Gly Phe Ser Asn Gly Asp Ala Val Glu Gly Glu
145 150 155 160
Gly Glu Arg Pro Arg Arg Val Phe Asp Arg Arg Ser Gly Thr Gly Arg
165 170 175
Gly Asn Glu Tyr Ile Lys Arg Glu Gly Ser Gly Arg Gly Asn Trp Gly
180 185 190
Thr Pro Ala Asp Asp Ile Ala Gln Glu Thr Glu Val Ala Val Asn Asp
195 200 205
Gly Glu Lys Ile Val Glu Thr Glu Lys Glu Ala Gly Gln Glu Asp Ala
210 215 220
Glu Asp Thr Asn Lys Asn Ser Thr Ala Ala Glu Pro Glu Glu Lys Glu
225 230 235 240
Pro Glu Glu Lys Glu Met Thr Leu Glu Glu Tyr Glu Lys Leu Met Glu
245 250 255
Glu Lys Arg Lys Ala Leu Met Ala Leu Lys Pro Glu Glu Arg Lys Glu
260 265 270
Met Thr Leu Glu Glu Tyr Glu Lys Leu Met Glu Glu Lys Arg Lys Ala
275 280 285
Leu Met Ala Leu Lys Pro Glu Glu Arg Lys Val Asn Leu Asp Lys Glu
290 295 300
Leu Gln Ser Met Gln Leu Leu Ser Asn Lys Lys Asn Asp Asp Glu Ile
305 310 315 320
Phe Ile Lys Leu Arg Ser Glu Lys Glu Lys Arg Lys Glu Ala Val Glu
325 330 335
Lys Ala Arg Lys Thr Gln Ser Ile Asn Glu Phe Leu Lys Pro Ala Glu
340 345 350
Gly Glu Ser Asn Tyr Arg Arg Gly Gly Arg Gly Arg Gly Gln Gly Arg
355 360 365
Gly Arg Gly Gly Phe Gly Gly Gly Val Gly Gly Asn Lys Asn Tyr Ser
370 375 380
Ala Pro Ser Ile Glu Asp Ala Gly Gln Phe Pro Ser Leu Val Ala Lys
385 390 395 400
<210> 7
<211> 370
<212> PRT
<213> 水稻(Oryza latifolia)OsRGGA基因编码的氨基酸
<400> 7
Met Val Thr Asn Asn Gln Phe Asp Leu Leu Val Asp Val Asp Asn Asp
1 5 10 15
Asp Pro Ser His Leu Ile Ala Ala Ala Glu Lys Met Lys Lys Ala Ala
20 25 30
Ala Ala Ala Ser Pro Ala Pro Val Val Gln Ala Arg Leu Pro Thr Lys
35 40 45
Pro Pro Pro Pro Ala Gln Ala Val Lys Glu Ser Arg Asn Tyr Gly Ala
50 55 60
Pro Ala Arg Asp Gly Ala Gly Gln Asn Gly Pro Gly His Ala Asn Gly
65 70 75 80
Gly Phe Arg Gly Gly Arg Met Gly Gln Arg Arg Asp Phe Gly Glu Gly
85 90 95
Asp Thr Asn Gly Ala Ser Gly Phe Arg Asp Gly Ile Val Arg Arg Glu
100 105 110
Glu Gly Glu Arg Arg Pro Ser Glu Arg Gly Arg Gly Pro Arg Gln Pro
115 120 125
Tyr Arg Gly Gly Gly Arg Arg Gly Gly Tyr Thr Gly Gly Glu Ala Gly
130 135 140
Asp Glu Ser Gly Arg Ala Pro Tyr Arg Ala Tyr Glu Arg Arg Ser Gly
145 150 155 160
Thr Gly Arg Gly Tyr Glu Thr Lys Arg Glu Gly Ala Gly Arg Gly Asn
165 170 175
Trp Gly Thr Val Thr Asp Glu Ser Leu Ala Gln Glu Ser Gly Glu Val
180 185 190
Val Asn Ile Glu Val Ala Ala Ala Val Thr Glu Asp Glu Thr Lys Gln
195 200 205
Glu Asp Val Pro Gln Ser Glu Val Glu Lys Arg Lys Glu Gly Glu Ser
210 215 220
Asn Glu Glu Glu Glu Lys Glu Pro Glu Asp Lys Glu Met Thr Leu Glu
225 230 235 240
Glu Tyr Glu Lys Val Leu Glu Glu Lys Arg Lys Ala Leu Leu Ser Leu
245 250 255
Lys Ala Glu Glu Arg Lys Val Val Val Asp Lys Glu Leu Gln Ser Met
260 265 270
Gln Gln Leu Ser Val Lys Lys Asp Ser Asp Glu Val Phe Ile Lys Leu
275 280 285
Gly Ser Asp Lys Asp Lys Lys Lys Glu Asn Val Glu Arg Asp Glu Arg
290 295 300
Thr Arg Lys Ser Leu Ser Ile Asn Glu Phe Leu Lys Pro Ala Glu Gly
305 310 315 320
Glu Arg Tyr Tyr Ser Pro Gly Gly Arg Gly Arg Gly Arg Gly Arg Gly
325 330 335
Arg Gly Asp Arg Gly Gly Phe Arg Asp Gly Tyr Ser Ser Arg Gly Pro
340 345 350
Val Ala Ala Pro Ala Ile Glu Asp Gln Ala Gln Phe Pro Ser Leu Ala
355 360 365
Gly Lys
370
<210> 8
<211> 364
<212> PRT
<213> 玉米(Zea mays)ZmRGGA基因编码的氨基酸
<400> 8
Met Ala Ser Ser Asn Gln Phe Asp Leu Leu Gly Asp Val Glu Asn Asp
1 5 10 15
Asp Pro Ala His Leu Leu Ala Ala Ala Glu Lys Lys Ala Ala Ala Val
20 25 30
Pro Lys Pro Ala Pro Ala Pro Ala Lys Leu Pro Thr Lys Pro Pro Pro
35 40 45
Pro Ala Gln Pro Val Arg Glu Glu Arg Asn Tyr Gly Ala Pro Pro Arg
50 55 60
Asp Gly Pro Gly Arg Gly Arg Gly Gly Arg Gly Gly Arg Thr Gly Pro
65 70 75 80
Arg Arg Asp Phe Gly Asp Ala Asp Ala His Gly Phe Glu Gly Gly Tyr
85 90 95
Gly Gly Gly Gly Gly Gly Phe Gly Asp Gly Gly Val Ala Arg Gly Glu
100 105 110
Asn Gly Glu Gly Arg Gln Ala Glu Arg Gly Arg Gly Pro Arg Gln Pro
115 120 125
Tyr Arg Gly Gly Gly Arg Arg Gly Gly Tyr Ser Asp Gly Gln Ala Gly
130 135 140
Asn Glu Phe Gly Arg Pro Arg Arg Ala Tyr Glu Arg His Ser Gly Thr
145 150 155 160
Gly Arg Gly Tyr Gly Met Lys Arg Glu Gly Ala Gly Arg Gly Asn Trp
165 170 175
Gly Thr Ala Thr Asp Glu Gly Leu Glu Gln Asp Thr Val Asp Ala Val
180 185 190
Asn Pro Glu Glu Thr Thr Ala Val Val Glu Asp Glu Lys Lys Pro Glu
195 200 205
Asp Ala Pro Gln Ser Glu Val Glu Lys Asp Lys Glu Gly Ala Glu Asn
210 215 220
Glu Glu Glu Lys Glu Pro Glu Asp Lys Glu Met Thr Leu Glu Glu Tyr
225 230 235 240
Glu Lys Val Leu Glu Glu Lys Arg Lys Ala Leu Leu Ser Leu Lys Ala
245 250 255
Glu Glu Arg Lys Val Glu Ile Asp Lys Glu Leu Gln Ala Met Gln Gln
260 265 270
Leu Ser Val Lys Lys Asp Gly Asp Glu Val Phe Ile Lys Leu Gly Ser
275 280 285
Asp Lys Asp Leu Lys Lys Lys Glu Ser Ala Glu Arg Glu Glu Arg Gly
290 295 300
Lys Lys Ser Leu Ser Ile Asn Glu Phe Leu Lys Pro Ala Glu Gly Glu
305 310 315 320
Arg Tyr Tyr Asn Pro Ser Gly Arg Gly Arg Gly Arg Gly Arg Gly Gly
325 330 335
Phe Arg Gly Gly Tyr Asn Gly Gly Tyr Arg Gly Pro Ala Ala Ala Pro
340 345 350
Ala Ile Glu Asp Gln Ala Gln Phe Pro Ala Leu Ala
355 360
<210> 9
<211> 27
<212> DNA
<213> 人工合成
<400> 9
ctgcaggcgt ggaggaggac gtggggg 27
<210> 10
<211> 29
<212> DNA
<213> 人工合成
<400> 10
ggatccccca cgaggtccgc catatccac 29
<210> 11
<211> 27
<212> DNA
<213> 人工合成
<400> 11
gagctcgcgt ggaggaggac gtggggg 27
<210> 12
<211> 28
<212> DNA
<213> 人工合成
<400> 12
ggatccccca cggccagcca cttcgtcg 28
<210> 13
<211> 23
<212> DNA
<213> 人工合成
<400> 13
ctgcaggaag acaaggagat gac 23
<210> 14
<211> 19
<212> DNA
<213> 人工合成
<400> 14
ggatccgcgc ttgatcttc 19
<210> 15
<211> 23
<212> DNA
<213> 人工合成
<400> 15
gagctcgaag acaaggagat gac 23
<210> 16
<211> 20
<212> DNA
<213> 人工合成
<400> 16
ggatcccacc tgggctgtag 20
<210> 17
<211> 36
<212> DNA
<213> 人工合成
<400> 17
aataatggtc tcaggcgccc gccagcctta ccgtgg 36
<210> 18
<211> 40
<212> DNA
<213> 人工合成
<400> 18
gcccgccagc cttaccgtgg gttttagagc tagaaatagc 40
<210> 19
<211> 35
<212> DNA
<213> 人工合成
<400> 19
aacccaaagt catctcctta tcgcttcttg gtgcc 35
<210> 20
<211> 38
<212> DNA
<213> 人工合成
<400> 20
attattggtc tctaaacaac ccaaagtcat ctccttat 38
<210> 21
<211> 25
<212> DNA
<213> 人工合成
<400> 21
gacaggcgtc ttctactggt gctac 25
<210> 22
<211> 25
<212> DNA
<213> 人工合成
<400> 22
ctcacaaatt atcagcacgc tagtc 25
<210> 23
<211> 25
<212> DNA
<213> 人工合成
<400> 23
ttgactagcg tgctgataat ttgtg 25
<210> 24
<211> 22
<212> DNA
<213> 人工合成
<400> 24
cacggagttc aaattcccac ag 22
Claims (4)
1.调控RGGA基因表达在提升作物农艺性状中的应用,其特征在于,所述调控为负调控;所述负调控为通过RNAi或CRISPR/Cas9下调RGGA基因表达;所述应用包括在作物生长发育早期对作物进行糖培养处理的步骤,所述糖培养处理为向作物施加蔗糖;所述作物为拟南芥、烟草、水稻或玉米;
其中,所述应用在拟南芥中,基因序列如SEQ ID NO.1所示,提升的农艺性状为营养生长指标和/或产量指标性状,所述营养生长指标性状为莲座叶大小、叶绿素含量、作物地上部及地下部干鲜重,所述产量指标性状为单株果荚数、单果荚种子数、百粒重;
所述应用在烟草、水稻或玉米中用于提升作物的生长,基因序列分别如SEQ ID NO.2-4所示。
2.一种提升作物农艺性状的方法,其特征在于,所述方法为:通过抑制作物RGGA基因表达、使得RGGA基因活性降低、使得RGGA基因失活的一种或多种,实现提升作物农艺性状;所述应用包括在作物生长发育早期对作物进行糖培养处理的步骤,所述糖培养处理为向作物施加蔗糖;所述作物为拟南芥、烟草、水稻或玉米;
其中,所述方法用于拟南芥中,基因序列如SEQ ID NO.1所示,提升的农艺性状为营养生长指标和/或产量指标性状,所述营养生长指标性状为莲座叶大小、叶绿素含量、作物地上部及地下部干鲜重,所述产量指标性状为单株果荚数、单果荚种子数、百粒重;
所述方法在烟草、水稻或玉米中用于提升作物的生长,基因序列分别如SEQ ID NO.2-4所示。
3.通过抑制作物RGGA基因表达、使得RGGA基因活性降低、使得RGGA基因失活的一种或多种方法在提升作物农艺性状的应用,其特征在于,所述抑制作物RGGA基因表达、使得RGGA基因活性降低或/和使得RGGA基因失活的物质,为RGGA蛋白特异的抗体、针对RGGA mRNA的RNA干扰分子或反义寡核苷酸、小分子抑制剂或siRNA;所述应用包括在作物生长发育早期对作物进行糖培养处理的步骤,所述糖培养处理为向作物施加蔗糖;所述作物为拟南芥、烟草、水稻或玉米;
其中,所述应用在拟南芥中,基因序列如SEQ ID NO.1所示,提升的农艺性状为营养生长指标和/或产量指标性状,所述营养生长指标性状为莲座叶大小、叶绿素含量、作物地上部及地下部干鲜重,所述产量指标性状为单株果荚数、单果荚种子数、百粒重;
所述应用在烟草、水稻或玉米中用于提升作物的生长,基因序列分别如SEQ ID NO.2-4所示。
4.一种作物生长促进剂,其特征在于,其活性成分包含抑制作物RGGA基因表达、使得RGGA基因活性降低或/和使得RGGA基因失活的物质,所述基因序列为如SEQ ID NO.1-4所示。
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Citations (6)
Publication number | Priority date | Publication date | Assignee | Title |
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