CN107868123A - 一种同时提高植物产量和抗性的基因及其应用 - Google Patents
一种同时提高植物产量和抗性的基因及其应用 Download PDFInfo
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Abstract
本发明公开了一种同时提高植物产量和抗性的基因及其应用。本发明提供的蛋白质,命名为AGO2蛋白,是如下(a)或(b):(a)由序列表中序列2所示的氨基酸序列组成的蛋白质;(b)将序列2氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的由序列2衍生的蛋白质。编码所述AGO2蛋白的基因(AGO2基因)也属于本发明的保护范围。本发明通过对单个基因的转基因即可达到同时增产高抗的目的,并对病害与盐胁迫均具有抗性,效果显著,为高产高抗作物育种提供了新的遗传位点。
Description
技术领域
本发明涉及一种同时提高植物产量和抗性的基因及其应用。
背景技术
在正常生长条件下,水稻的产量通常由粒重,有效分蘖数和每穗粒数直接决定。粒重或籽粒大小受到显著遗传控制,水稻作为单子叶模式植物,籽粒大小由多基因调控,目前已克隆了较多影响水稻籽粒大小的数量性状位点基因(QTL)。同时,水稻产量也受到环境的影响,恶劣的环境如盐碱、干旱、病害等会导致严重减产。在逆境条件下,植物能够在分子、细胞和整体水平上做出相应的调整,以最大程度上减少环境所造成的伤害并得以生存。许多基因受胁迫诱导表达,这些基因的产物不仅能够直接参与植物的胁迫应答,而且能够调节其它相关基因的表达或参与信号传导途径,从而使植物避免或减少伤害,增强对胁迫环境的抗性。植物对病害的调控机制更为复杂,与非生物逆境既有关联,又有极大的不同,不同病原菌的致病机理也并不相同,因此,抗病基因克隆显得更具有挑战性和重要意义。
提高作物产量和抗性是育种家长期追求的目标。目前,大多数产量和抗性相对平衡的品种都是通过传统育种选育而来。利用产量高的一个亲本品种与另外一个抗性强的亲本品种通过杂交,在后代选育同时具有高产和高抗的植株,进一步进行必要的回交选育,最终将一个亲本中的抗性或产量遗传位点导入到另外一个目的品种中,达到改良的目的。随着高产高抗位点的克隆鉴定,在此过程中越来越多的通过分子标记辅助进行选育。高产高抗通常由多个遗传位点控制,通过设计各个位点的分子连锁标记,可相对快速的将目标高产或高抗位点进行分子聚合。除此之外,在了解某一个基因的功能前提下,利用转基因技术操纵某个基因或对某个基因进行编辑也可快速对目标品种进行改造。
传统育种往往是经验育种,育种家需要有足够的经验积累,因此存在一定的盲目性,并且周期长,工作量大。特别是对产量和抗性有多方面的要求时,更需要扩大育种规模,包括目标亲本和筛选群体,在此基础上进行多点多年测试以获得期望的高产多抗品种。虽然分子标记辅助选育能一定程度上提高效率,但一方面目前克隆到的基因仍然相对较少,而在已经克隆的基因中,很多位点往往已经在传统育种过程中被利用,人们只是找到了这些被利用的位点,而真正可供利用的有效的分子标记并不多,另一方面由于多基因的互作效应往往非常复杂,遗传关系并不清楚,即便经过耗时费力的聚合后,最终也很难达到预期的目标。最后,通过单一位点的基因操纵可以定向改造某一性状,但很难同时调节抗性和产量以满足多方面的改造需求。
发明内容
本发明的目的是提供一种同时提高植物产量和抗性的基因及其应用。
本发明提供的蛋白质,获自水稻,命名为AGO2蛋白,是如下(a)或(b):
(a)由序列表中序列2所示的氨基酸序列组成的蛋白质;
(b)将序列2的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的由序列2衍生的蛋白质。
为了使(a)中的AGO2蛋白便于纯化和检测,可在由序列表中序列2所示的氨基酸序列组成的蛋白质的氨基末端或羧基末端连接上如表1所示的标签。
表1标签的序列
上述(b)中的AGO2蛋白可人工合成,也可先合成其编码基因,再进行生物表达得到。上述(b)中的AGO2蛋白的编码基因可通过将序列表中序列1所示的DNA序列中缺失一个或几个氨基酸残基的密码子,和/或进行一个或几个碱基对的错义突变,和/或在其5′端和/或3′端连上表1所示的标签的编码序列得到。
编码所述AGO2蛋白的基因(AGO2基因)也属于本发明的保护范围。
所述AGO2基因为如下(1)-(4)中任一所述的DNA分子:
(1)编码区如序列表中序列1所示的DNA分子;
(2)序列表中序列1所示的DNA分子;
(3)在严格条件下与(1)或(2)限定的DNA序列杂交且编码所述蛋白质的DNA分子;
(4)与(1)或(2)或(3)限定的DNA序列具有90%以上同源性且编码所述蛋白质的DNA分子。
上述严格条件可为用0.1×SSPE(或0.1×SSC),0.1%SDS的溶液,在DNA或者RNA杂交实验中65℃下杂交并洗膜。
含有所述AGO2基因的重组表达载体、表达盒、转基因细胞系或重组菌均属于本发明的保护范围。
可用现有的植物表达载体构建含有AGO2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微弹轰击的载体等。使用AGO2基因构建重组表达载体时,可在其转录起始核苷酸前加上任何一种增强型、组成型、组织特异型或诱导型启动子,它们可单独使用或与其它的植物启动子结合使用;此外,使用AGO2基因构建重组表达载体时,还可使用增强子,包括翻译增强子或转录增强子,这些增强子区域可以是ATG起始密码子或邻接区域起始密码子等,但必需与编码序列的阅读框相同,以保证整个序列的正确翻译。所述翻译控制信号和起始密码子的来源是广泛的,可以是天然的,也可以是合成的。翻译起始区域可以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,如加入在植物中表达可产生颜色变化的酶或发光化合物的基因、具有抗性的抗生素标记物或是抗化学试剂标记基因等。
所述重组表达载体可为将pCAMBIA2300-35S-eGFP载体的多克隆位点中插入了序列表的序列1的自5′端第1-3102位核苷酸所示的DNA分子得到的重组质粒。所述重组表达载体具体可为将pCAMBIA2300-35S-eGFP载体的XmaI和XbaI酶切位点之间的小片段取代为了序列表的序列1的自5′端第1-3102位核苷酸所示的DNA分子得到的重组质粒。
本发明还保护AGO2蛋白或AGO2基因的应用,为如下(c1)和/或(c2):
(c1)调控植物产量;
(c2)调控植物耐逆性。
本发明还保护一种培育转基因植物的方法,是将AGO2基因导入目的植物中,得到转基因植物;所述转基因植物具有如下(d1)和/或(d2)所述表型:
(d1)产量高于所述目的植物;
(d2)耐逆性高于所述目的植物。
所述方法中,所述AGO2基因可以通过以上任一所述重组表达载体导入目的植物。所述重组表达载体可通过Ti质粒、Ri质粒、植物病毒载体、直接DNA转化、显微注射、电导、农杆菌介导等常规生物学方法转化到植物细胞或组织中。
本发明还保护一种培育转基因植物的方法,是提高目的植物中AGO2蛋白的表达量和/或活性,得到转基因植物;所述转基因植物具有如下(d1)和/或(d2)所述表型:
(d1)产量高于所述目的植物;
(d2)耐逆性高于所述目的植物。
以上任一所述目的植物为单子叶植物或双子叶植物。所述单子叶植物可为禾本目植物。所述禾本目植物可为禾本科植物。所述禾本科植物可为稻属植物。所述稻属植物具体可为水稻,例如中花11水稻。
本发明还保护AGO2蛋白或AGO2基因或以上任一所述方法在植物育种中的应用。
所述育种的目的为选育产量高和/或耐逆性高的植物。
以上任一所述耐逆性具体可为耐盐性和/或抗病性。所述抗病性具体可为白叶枯病抗性和/或黑条矮缩病抗性。
以上任一所述所述植物为单子叶植物或双子叶植物。所述单子叶植物可为禾本目植物。所述禾本目植物可为禾本科植物。所述禾本科植物可为稻属植物。所述稻属植物具体可为水稻,例如中花11水稻。
本发明的发明人通过实验证明,AGO2基因在水稻(中花11背景)中过表达后可促进水稻籽粒大小与粒重,相对于野生型,转基因植株粒长增加,百粒重提高可达20%-30%;同时植物对高盐胁迫及白叶枯病菌及矮缩病的抗性都显著增强,在200mM氯化钠盐溶液中培养,在野生型接近100%死亡后,转基因植株存活率可达50%-80%;接种水稻黑条矮缩病毒后,转基因植株生长优于野生型,并且利用抗体检测病毒蛋白,转基因植株病毒积累显著下降。接种白叶枯病菌后,转基因植株对多达8个白叶枯生理小种都表现出一定抗性。本发明通过对单个基因的转基因即可达到同时增产高抗的目的,并对病害与盐胁迫均具有抗性,效果显著,为高产高抗作物育种提供了新的遗传位点。
附图说明
图1为野生型和转基因植株的AGO2基因相对表达量、籽粒大小和粒重统计结果。
图2为野生型和转基因植株的盐胁迫抗性检测结果。
图3为野生型和转基因植株的白叶枯病抗性检测结果。
图4为野生型和转基因植株的黑条矮缩病抗性检测结果。
具体实施方式
以下的实施例便于更好地理解本发明,但并不限定本发明。下述实施例中的实验方法,如无特殊说明,均为常规方法。下述实施例中所用的试验材料,如无特殊说明,均为自常规生化试剂商店购买得到的。以下实施例中的定量试验,均设置三次重复实验,结果取平均值。
中花11:参考文献:“Xiao Y,Liu D,Zhang G,Tong H and Chu C(2017)Brassinosteroids Regulate OFP1,a DLT Interacting Protein,to Modulate PlantArchitecture and Grain Morphology in Rice.Front.Plant Sci.8:1698.doi:10.3389/fpls.2017.01698”;公众可以从中国科学院遗传与发育生物学研究所获得。
pCAMBIA2300-35S-eGFP载体:参考文献:“Xiao Y,Liu D,Zhang G,Tong H andChu C(2017)BrassinosteroidsRegulate OFP1,a DLT InteractingProtein,to ModulatePlantArchitecture and Grain Morphologyin Rice.Front.Plant Sci.8:1698.doi:10.3389/fpls.2017.01698”;公众可以从中国科学院遗传与发育生物学研究所获得。
农杆菌AGL1:北京博迈德基因技术有限公司。公众可以从中国科学院遗传与发育生物学研究所获得。
实施例1、AGO2蛋白及其编码基因的获得
对水稻全长基因进行序列分析、区段截取和功能验证,得到候选克隆,测序得到目标克隆的全长序列,如序列表中序列1所示,编码序列表的序列2所示的蛋白质。
将序列表的序列2所示的蛋白质命名为AGO2蛋白,由1034个氨基酸残基组成。将AGO2蛋白的编码基因命名为AGO2基因。AGO2基因编码区如序列表中序列1所示。
实施例2、AGO2基因过表达转基因植株的获得
1、提取野生型水稻中花11幼苗的总RNA,并反转录为cDNA。
2、以步骤1得到的cDNA为模板,采用引物AGO2FL-F和引物AGO2FL-R组成的引物对进行PCR扩增,得到PCR扩增产物。
AGO2FL-F:5’-CCC GGG ATG GAG CAC GAG CGC GGT G-3’;
AGO2FL-R:5’-TCT AGA GAT GAA GAA CAT GTT GTC CAC CAG A G-3’。
引物AGO2FL-F和引物AGO2FL-R中,下划线分别为XmaI和XbaI酶切位点。
3、采用限制性内切酶XmaI和XbaI双酶切步骤2得到的PCR扩增产物,回收酶切产物。
4、采用限制性内切酶XmaI和XbaI双酶切pCAMBIA2300-35S-eGFP载体,回收约10kb的载体骨架。
5、将步骤3的酶切产物和步骤4的载体骨架连接,得到重组载体pCAMBIA2300-35S-eGFP-AGO2。根据测序结果,对重组载体pCAMBIA2300-35S-eGFP-AGO2进行结构描述如下:将pCAMBIA2300-35S-eGFP载体的XmaI和XbaI酶切位点之间的小片段取代为了序列表的序列1的自5′端第1-3102位核苷酸所示的DNA分子。
6、采用步骤5得到的重组载体pCAMBIA2300-35S-eGFP-AGO2转化农杆菌AGL1,得到重组菌。参照文献“Hiei Y,Ohta S,Komari T,Kumashiro T.Efficient transformationof rice(Oryza sativa L.)mediated by Agrobacterium and sequence analysis ofthe boundaries of the T-DNA.Plant J 1994;6:271–82”中的方法将重组菌转化中花11愈伤组织,具体如步骤7-14所示。
7、将步骤6得到的重组菌接种于含有50mg/ml卡那霉素和50mg/ml利福平的YEB液体培养基中,200rpm,暗培养3天,得到重组菌悬液,4,000rpm离心3min收集沉淀。
8、采用含有0.1mM乙酰丁香酮的AAM液体培养基重悬步骤7得到的沉淀,28℃,150rpm摇床避光培养至OD600nm=0.4,得到侵染液。
9、挑选生长状态良好、颗粒状中花11愈伤组织,浸入步骤8得到的侵染液中,28℃、150-200rpm培养20min,然后将愈伤组织取出,用无菌滤纸吸干多余菌液,将愈伤组织平铺在含多层滤纸的无菌平皿中,超净台上吹干(愈伤分散不结块),然后将愈伤组织转移到NB固体培养基上,26℃暗培养2-3天。
10、完成步骤9后,将愈伤组织接种于含150mg/L G418和400mg/L头孢霉素的NB固体培养基上,26℃暗培养3.5周。
11、完成步骤10后,将成活的愈伤组织转入含有200mg/L G418和200mg/L头孢霉素的NB固体培养基上,26℃暗培养3周。
12、完成步骤11后,将成活的愈伤组织转入含有200mg/L G418的分化培养基(NB基本培养基、2mg/L 6-BA、1mg/L NAA)上,26℃、弱光(光强约150umol/m2.s)培养得到再生植株。
13、完成步骤13后,将再生植株在含200mg/L G418的壮苗培养基(1/2MS、0.5mg/LNAA、0.25mg/L MET)上26℃、弱光(光强约150umol/m2.s)培养至生根后转移至温室中培养,得到T0代转基因植株。
14、提取步骤13得到的T0代转基因植株叶片总DNA为模板,采用引物NPT-F和引物NPT-R组成的引物对进行PCR扩增,筛选得到T0阳性转基因植株(阳性植株PCR扩增产物大小为582bp)。
NPT-F:5’-TCC GGT GCC CTG AAT GAA CT-3’;
NPT-R:5’-GGC GAT ACC GTA AAG CAC GA-3’。
T0代植株自交,得到T1代植株。T1代植株自交,得到T2代植株。
将T1代植株和T2代植株也采用引物NPT-F和引物NPT-R进行鉴定,如果对于某一T0代植株,其抽样检测的T1代植株和T2代植株的PCR鉴定结果均为阳性,该T0代植株及其自交后代为一个纯合的过表达转基因株系。
15、提取水稻中花11(ZH11)和步骤14得到的若干过表达转基因株系的T2代植株叶片总RNA,并反转录为cDNA;以cDNA为模板,采用qRT-PCR的方法检测AGO2基因的表达情况(以Actin基因为内参基因),采用引物ACT-F和引物ACT-R组成的引物对检测Actin基因的表达,采用引物AGO2-F和引物AGO2-R组成的引物对检测AGO2基因的表达。
AGO2-F:5’-AGC CAA GGT CAA ATT GTT GG-3’;
AGO2-R:5’-CTC CTT GTC TGA AGC CTT GG-3’;
ACT-F:5’-TGC TAT GTA CGT CGC CAT CCA G-3’;
ACT-R:5’-AAT GAG TAA CCA CGC TCC GTC A-3’。
结果如图1B所示。图1B中,纵坐标为AGO2基因相对表达量。结果表明,与野生型相比,AGO2基因在5个转基因株系(A2OX-4、A2OX-7、A2OX-8、A2OX-14、A2OX-18)中表达量均升高。
16、采用pCAMBIA2300-35S-eGFP载体替代重组载体pCAMBIA2300-35S-eGFP-AGO2,按照步骤6-步骤14进行操作过程中,得到空载体植株。
实施例2、AGO2基因过表达转基因植株的表型分析
一、籽粒大小与粒重统计
待测植株:野生型中花11(ZH11)、转基因株系(A2OX-4、A2OX-7、A2OX-8、A2OX-14、A2OX-18)的T2代植株、转空载体植株。
对待测植株成熟期收获的籽粒大小和粒重进行统计,结果如图1A、图1C和图1D所示。图1A位表型观察结果。图1C为粒长统计结果,纵坐标为粒长(mm)。图1D位百粒重统计结果,纵坐标为百粒重(g)。
结果表明,相对于野生型,转基因株系粒长粒重均显著增加,特别是A2OX-4和A2OX-8两个株系粒重增加最为明显,百粒重分别增加30.8%和22.7%。转空载体植株表型与野生型无显著差异。
二、抗盐胁迫检测
将野生型中花11(ZH11)和转基因株系(A2OX-4、A2OX-7、A2OX-8、A2OX-14、A2OX-18)的T2代植株的种子在37℃黑暗浸种萌发后,每个转基因株系与野生型各对半铺于去掉底部的96孔塑料板中,每个株系设4个重复,在光照培养箱中使用1/2MS液体培养基培养10天(温度30℃,光照12h/黑暗12h,每2天换新鲜1/2MS液体培养基),第10天换成含有200mM氯化钠的1/2MS液体培养基,继续培养10天,观察表型。
结果如图2所示。结果表明,盐胁迫下野生型接近完全死亡,而转基因株系大部分仍然存活,存活率达到50%-80%,采用转空载体植株替代转基因株系植株进行试验,存活率与野生型无显著差异,说明AGO2基因可以提高植物的盐胁迫抗性。
三、白叶枯病抗性检测
待测植株:野生型中花11(ZH11)、转基因株系(A2OX-4、A2OX-8)的T2代植株、转空载体植株。
检测待测植株白叶枯病抗性,检测方法和所用生物材料在“Chen H,Wang S,ZhangQ.2002.New gene for bacterial blight resistance in rice located on chromosome12identified from minghui 63,an elite restorer line.Phytopathology 92:750–754.doi:10.1094/PHYTO.2002.92.7.”中记载。在水稻生长正季,将田间生长2个月的待测植株分别接种不同白叶枯生理小种共4种,用剪刀分别沾取培养好的病原菌液,减去同一发育阶段的叶片的尖部同样位置,两周后,测量叶片受侵染后的病斑长度,除以叶片长度,计算病斑相对长度。
结果如图3所示。结果表明,转基因株系感病相对长度明显低于野生型,转空载体植株与野生型无显著差异,转基因植株对各个白叶枯生理小种均具有明显抗性。
四、黑条矮缩病抗性检测
检测植株的黑条矮缩病抗性,检测方法和所用生物材料在“Yuqing He,HehongZhang,Zongtao Sun,Junmin Li,Gaojie Hong,Qisong Zhu,XuebiaoZhou,StuartMacFarlane,Fei Yan and Jianping Chen.2016.Jasmonic acid-mediated defensesuppresses brassinosteroidmediatedsusceptibility to Rice black streaked dwarfvirusinfection in rice.New Phytologist.doi:10.1111/nph.14376”中记载。
1、成年无毒雌性灰飞虱接种于野生型中花11上产卵,生长10天成蛹虫后转移到已感染黑条矮缩病病毒的水稻苗上喂食4天,使其带毒,再转移到健康中花11水稻幼苗上生长12天到成年期,使用酶联免疫吸附测定法确定带毒。
2、将野生型中花11(ZH11)、转基因株系(A2OX-4、A2OX-8)的T2代植株种子萌发后置于同一玻璃烧杯中在光照培养箱中水培一周,用塑料网封闭后按每棵幼苗3只昆虫放入步骤1得到的带毒灰飞虱,培养3天,之后彻底除去昆虫,将幼苗移栽到田间培养一个月,观察表型,植物表现为显著矮化,叶片发黑,即为发病症状。从侵染当天开始计算,于30天和60天分别取叶片,利用免疫杂交检测病毒蛋白含量,比较野生型与转基因株系病毒存活水平。
结果如图4所示。结果表明,转基因株系带毒量明显减少,特别是在A2OX-8株系中,采用转空载体植株替代转基因株系植株进行试验,表型和病毒蛋白含量与野生型无显著差异,说明转基因株系对病毒有抗性。
<110> 中国农业科学院作物科学研究所
中国科学院遗传与发育生物学研究所
<120> 一种同时提高植物产量和抗性的基因及其应用
<160> 2
<210> 1
<211> 3105
<212> DNA
<213> 水稻(Oryza sativa)
<400> 1
atggagcacg agcgcggtgg cggtggccgc ggccgcggga ggggtcgcgg tggcgggcgt 60
ggcggcggtg gcggcgatgg tcgcggaggc ggttatggtg gtgctggtgg tggtggtgtc 120
ggcgggcgcg gtgggcgtgg gcctcctggt ggtggtggtg gacgcgggta cgagcccggc 180
ggcgggcgtg ggtacggtgg cggcggcggc ggtggtggac gtgggtatgg cggcggaggc 240
ggcggtggtg ggtacgagtc cggcggtggg cgtgggtatg gcggcggtgg acgtgggtat 300
gaatccggcg gtgggcgtgg acctggcggc ggcggccgtg ggcacgagtc cggcggtggc 360
ggtggccgcg gcgggaacgt gtgggcgcag ccggggagag ggcgcggagg agcccccgcc 420
ccggcgccgg cgccagcacc agcagcgagg aggatccagg acgagggggc cgcgaggtcg 480
tcgggtaccg ttgagcgcat tgcttctact gaggttgtaa gagtacaacc acctgcaccc 540
ccagttgctg tgtctcgtag tggcacgcgt gtgccaatgc gaagacctga tggtggaggc 600
tcagtatcga aagccaaggt caaattgttg gtgaaccatt ttatagttaa gtaccgacag 660
gcatcaactg tttttcacta tgacatagac atcaagcttg atataagttc ccccaaggct 720
tcagacaagg agctatccaa gggagatttt cttactgtca aggacgagct cttcaaggat 780
gagagctttc ggcggctttc atcagctgtt gcttatgatg gaaaaagaaa tttatttact 840
tgtgctgagc taccagatgg tttgtttcgt gtcaaagtcc gttcacggac ttacattgta 900
tctgtggagt tcaagaagaa gcttcctttg agccaactct cggaactgcc tgtgcccaga 960
gaggtcttgc aggggcttga tgtcattgtg cgtgaggcct ctagctggcg caagattatc 1020
attggtcagg gattttactc gcagggccgc agtgtgccca ttgggccgga tgttgtagct 1080
ctcaaaggaa cccagcagac cctgaaatgc actcagaaag gactgatcct ttgtgtggac 1140
tattcggtta tgccgtttcg caaagctgga cctgtgttgg atcttgttca gaagtctgtg 1200
agataccttg actacaggac aacactaaac aaacaccaat tggacacttt gaagaatgaa 1260
ctcaaaggcc agcgtgtcac tgtaaatcat aggaggacaa agcagaagta cattgttaaa 1320
ggtttgactg ataaacctgc aagtcagata acttttgtag attctgaatc aggacagacc 1380
aagaagcttc ttgattacta ttcgcagcag tatggcaagg ttattgagta tcaaatgctt 1440
ccatgcttgg atttgagcaa gagcaaggac aagcaaaact atgtgccgat tgaattgtgt 1500
gatcttcttg aagggcagag atacccaaaa gcaagcttaa ataggaattc tgataaaaca 1560
ctgaaagaaa tggctttgat ccctgcctca agtaggaagg aggagattct ggagttggtg 1620
aatgctgacg atgggccttg caggggtgaa attgctcagc agttcgggat ttctttggat 1680
gtacaaatga tggaagtcac tggtaggacc cttcctcctc ccagcctaaa acttggcacc 1740
tccagtggcc aaccccccaa attcaatatt gatcagccta actgccagtg gaaccttacg 1800
aggaaaagac tagcagaggg cggggtgcta cagtgctggg gcgttgtgga cttcagtgca 1860
gattctgggc agtacgccct gaatgggaac atgtttattg acaagattgt caggaagtgc 1920
tgcgaccttg gcgtacagat gaaccgtaac ccatgcattg tgcaactgtt agatatggag 1980
gtgctatccg atccacatca gctcttcgag gagcttaaca aagctaagca ggcggcagcc 2040
agtaagaaac agaagctgca gctcctcttc tgcccaatgt ctgatcagca tcctgggtac 2100
aagacgctga agcttatctg cgagacgcag ctggggatcc agacccagtg cttcttgagc 2160
ttcctcgcga acaaacaaca gggacaggac cagtacatgt ccaaccttgc tctgaagatc 2220
aacggcaaga ttggaggaag caacatccaa ctgtttggtg aatcgctccc gcggatctcc 2280
ggcgcgccat acatgttcat cggcgccgac gtgaatcacc catcgccggg gaacgtcgag 2340
agcccgtcga ttgcagcagt ggtggcctcg gtggatcaag gcgccagcaa gtacgtgcca 2400
agaatccgcg ctcagcctca ccgctgcgag gtgatccagc acctcggcga catgtgcaag 2460
gagctcatcg gcgtgttcga gaagcggaac cgcgtgaagc cccagaggat catctacttc 2520
cgcgacggcg tcagcgacgg tcagttcgac atggtgctga acgaggagct ggcggacatg 2580
gagaaggcga tcaagaccaa ggactactcc ccgacgatca ccgtgatcgt ggccaagaag 2640
cggcaccaca ccaggctgtt ccccaaggac ctgaaccagc agcagaccaa gaacggcaac 2700
gtgctccccg gcacggtggt ggacaccggc gtggtcgacc cggcggcgta cgacttctac 2760
ctgtgcagcc acaacgggct gatcgggacg agccggccga cgcactacta cagccttctg 2820
gacgagcacg gcttcgcctc cgacgacctg cagaagctgg tgtacaacct ctgcttcgtc 2880
ttcgcccgct gcaccaagcc ggtgtcgctg gccacgcccg tctactacgc cgacctcgcc 2940
gcctaccgcg gcaggctcta ctacgagggc atgatgatgt cgcagccgcc accgtcttcc 3000
gcggcgtcgg cgtcgtcggc atcctcctcc ggcgccggcg cttccgactt caggagcttc 3060
ccggcgctgc acgaggatct ggtggacaac atgttcttca tctga 3105
<210> 2
<211> 1034
<212> PRT
<213> 水稻(Oryza sativa)
<400> 2
Met Glu His Glu Arg Gly Gly Gly Gly Arg Gly Arg Gly Arg Gly Arg
1 5 10 15
Gly Gly Gly Arg Gly Gly Gly Gly Gly Asp Gly Arg Gly Gly Gly Tyr
20 25 30
Gly Gly Ala Gly Gly Gly Gly Val Gly Gly Arg Gly Gly Arg Gly Pro
35 40 45
Pro Gly Gly Gly Gly Gly Arg Gly Tyr Glu Pro Gly Gly Gly Arg Gly
50 55 60
Tyr Gly Gly Gly Gly Gly Gly Gly Gly Arg Gly Tyr Gly Gly Gly Gly
65 70 75 80
Gly Gly Gly Gly Tyr Glu Ser Gly Gly Gly Arg Gly Tyr Gly Gly Gly
85 90 95
Gly Arg Gly Tyr Glu Ser Gly Gly Gly Arg Gly Pro Gly Gly Gly Gly
100 105 110
Arg Gly His Glu Ser Gly Gly Gly Gly Gly Arg Gly Gly Asn Val Trp
115 120 125
Ala Gln Pro Gly Arg Gly Arg Gly Gly Ala Pro Ala Pro Ala Pro Ala
130 135 140
Pro Ala Pro Ala Ala Arg Arg Ile Gln Asp Glu Gly Ala Ala Arg Ser
145 150 155 160
Ser Gly Thr Val Glu Arg Ile Ala Ser Thr Glu Val Val Arg Val Gln
165 170 175
Pro Pro Ala Pro Pro Val Ala Val Ser Arg Ser Gly Thr Arg Val Pro
180 185 190
Met Arg Arg Pro Asp Gly Gly Gly Ser Val Ser Lys Ala Lys Val Lys
195 200 205
Leu Leu Val Asn His Phe Ile Val Lys Tyr Arg Gln Ala Ser Thr Val
210 215 220
Phe His Tyr Asp Ile Asp Ile Lys Leu Asp Ile Ser Ser Pro Lys Ala
225 230 235 240
Ser Asp Lys Glu Leu Ser Lys Gly Asp Phe Leu Thr Val Lys Asp Glu
245 250 255
Leu Phe Lys Asp Glu Ser Phe Arg Arg Leu Ser Ser Ala Val Ala Tyr
260 265 270
Asp Gly Lys Arg Asn Leu Phe Thr Cys Ala Glu Leu Pro Asp Gly Leu
275 280 285
Phe Arg Val Lys Val Arg Ser Arg Thr Tyr Ile Val Ser Val Glu Phe
290 295 300
Lys Lys Lys Leu Pro Leu Ser Gln Leu Ser Glu Leu Pro Val Pro Arg
305 310 315 320
Glu Val Leu Gln Gly Leu Asp Val Ile Val Arg Glu Ala Ser Ser Trp
325 330 335
Arg Lys Ile Ile Ile Gly Gln Gly Phe Tyr Ser Gln Gly Arg Ser Val
340 345 350
Pro Ile Gly Pro Asp Val Val Ala Leu Lys Gly Thr Gln Gln Thr Leu
355 360 365
Lys Cys Thr Gln Lys Gly Leu Ile Leu Cys Val Asp Tyr Ser Val Met
370 375 380
Pro Phe Arg Lys Ala Gly Pro Val Leu Asp Leu Val Gln Lys Ser Val
385 390 395 400
Arg Tyr Leu Asp Tyr Arg Thr Thr Leu Asn Lys His Gln Leu Asp Thr
405 410 415
Leu Lys Asn Glu Leu Lys Gly Gln Arg Val Thr Val Asn His Arg Arg
420 425 430
Thr Lys Gln Lys Tyr Ile Val Lys Gly Leu Thr Asp Lys Pro Ala Ser
435 440 445
Gln Ile Thr Phe Val Asp Ser Glu Ser Gly Gln Thr Lys Lys Leu Leu
450 455 460
Asp Tyr Tyr Ser Gln Gln Tyr Gly Lys Val Ile Glu Tyr Gln Met Leu
465 470 475 480
Pro Cys Leu Asp Leu Ser Lys Ser Lys Asp Lys Gln Asn Tyr Val Pro
485 490 495
Ile Glu Leu Cys Asp Leu Leu Glu Gly Gln Arg Tyr Pro Lys Ala Ser
500 505 510
Leu Asn Arg Asn Ser Asp Lys Thr Leu Lys Glu Met Ala Leu Ile Pro
515 520 525
Ala Ser Ser Arg Lys Glu Glu Ile Leu Glu Leu Val Asn Ala Asp Asp
530 535 540
Gly Pro Cys Arg Gly Glu Ile Ala Gln Gln Phe Gly Ile Ser Leu Asp
545 550 555 560
Val Gln Met Met Glu Val Thr Gly Arg Thr Leu Pro Pro Pro Ser Leu
565 570 575
Lys Leu Gly Thr Ser Ser Gly Gln Pro Pro Lys Phe Asn Ile Asp Gln
580 585 590
Pro Asn Cys Gln Trp Asn Leu Thr Arg Lys Arg Leu Ala Glu Gly Gly
595 600 605
Val Leu Gln Cys Trp Gly Val Val Asp Phe Ser Ala Asp Ser Gly Gln
610 615 620
Tyr Ala Leu Asn Gly Asn Met Phe Ile Asp Lys Ile Val Arg Lys Cys
625 630 635 640
Cys Asp Leu Gly Val Gln Met Asn Arg Asn Pro Cys Ile Val Gln Leu
645 650 655
Leu Asp Met Glu Val Leu Ser Asp Pro His Gln Leu Phe Glu Glu Leu
660 665 670
Asn Lys Ala Lys Gln Ala Ala Ala Ser Lys Lys Gln Lys Leu Gln Leu
675 680 685
Leu Phe Cys Pro Met Ser Asp Gln His Pro Gly Tyr Lys Thr Leu Lys
690 695 700
Leu Ile Cys Glu Thr Gln Leu Gly Ile Gln Thr Gln Cys Phe Leu Ser
705 710 715 720
Phe Leu Ala Asn Lys Gln Gln Gly Gln Asp Gln Tyr Met Ser Asn Leu
725 730 735
Ala Leu Lys Ile Asn Gly Lys Ile Gly Gly Ser Asn Ile Gln Leu Phe
740 745 750
Gly Glu Ser Leu Pro Arg Ile Ser Gly Ala Pro Tyr Met Phe Ile Gly
755 760 765
Ala Asp Val Asn His Pro Ser Pro Gly Asn Val Glu Ser Pro Ser Ile
770 775 780
Ala Ala Val Val Ala Ser Val Asp Gln Gly Ala Ser Lys Tyr Val Pro
785 790 795 800
Arg Ile Arg Ala Gln Pro His Arg Cys Glu Val Ile Gln His Leu Gly
805 810 815
Asp Met Cys Lys Glu Leu Ile Gly Val Phe Glu Lys Arg Asn Arg Val
820 825 830
Lys Pro Gln Arg Ile Ile Tyr Phe Arg Asp Gly Val Ser Asp Gly Gln
835 840 845
Phe Asp Met Val Leu Asn Glu Glu Leu Ala Asp Met Glu Lys Ala Ile
850 855 860
Lys Thr Lys Asp Tyr Ser Pro Thr Ile Thr Val Ile Val Ala Lys Lys
865 870 875 880
Arg His His Thr Arg Leu Phe Pro Lys Asp Leu Asn Gln Gln Gln Thr
885 890 895
Lys Asn Gly Asn Val Leu Pro Gly Thr Val Val Asp Thr Gly Val Val
900 905 910
Asp Pro Ala Ala Tyr Asp Phe Tyr Leu Cys Ser His Asn Gly Leu Ile
915 920 925
Gly Thr Ser Arg Pro Thr His Tyr Tyr Ser Leu Leu Asp Glu His Gly
930 935 940
Phe Ala Ser Asp Asp Leu Gln Lys Leu Val Tyr Asn Leu Cys Phe Val
945 950 955 960
Phe Ala Arg Cys Thr Lys Pro Val Ser Leu Ala Thr Pro Val Tyr Tyr
965 970 975
Ala Asp Leu Ala Ala Tyr Arg Gly Arg Leu Tyr Tyr Glu Gly Met Met
980 985 990
Met Ser Gln Pro Pro Pro Ser Ser Ala Ala Ser Ala Ser Ser Ala Ser
995 1000 1005
Ser Ser Gly Ala Gly Ala Ser Asp Phe Arg Ser Phe Pro Ala Leu
1010 1015 1020
His Glu Asp Leu Val Asp Asn Met Phe Phe Ile
1025 1030
Claims (10)
1.一种蛋白质,是如下(a)或(b):
(a)由序列表中序列2所示的氨基酸序列组成的蛋白质;
(b)将序列2的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的由序列2衍生的蛋白质。
2.编码权利要求1所述蛋白质的基因。
3.如权利要求2所述的基因,其特征在于:所述基因为如下(1)-(4)中任一所述的DNA分子:
(1)编码区如序列表中序列1所示的DNA分子;
(2)序列表中序列1所示的DNA分子;
(3)在严格条件下与(1)或(2)限定的DNA序列杂交且编码权利要求1所述蛋白质的DNA分子;
(4)与(1)或(2)或(3)限定的DNA序列具有90%以上同源性且编码权利要求1所述蛋白质的DNA分子。
4.含有权利要求2或3所述基因的重组表达载体、表达盒、转基因细胞系或重组菌。
5.权利要求1所述蛋白质,或,权利要求2或3所述基因的应用,为如下(c1)和/或(c2):
(c1)调控植物产量;
(c2)调控植物耐逆性。
6.如权利要求5所述的应用,其特征在于:所述植物为双子叶植物或单子叶植物。
7.一种培育转基因植物的方法,是将权利要求2或3所述基因导入目的植物中,得到转基因植物;所述转基因植物具有如下(d1)和/或(d2)所述表型:
(d1)产量高于所述目的植物;
(d2)耐逆性高于所述目的植物。
8.一种培育转基因植物的方法,是提高目的植物中权利要求1所述蛋白的表达量和/或活性,得到转基因植物;所述转基因植物具有如下(d1)和/或(d2)所述表型:
(d1)产量高于所述目的植物;
(d2)耐逆性高于所述目的植物。
9.权利要求1所述蛋白质,或,权利要求2或3所述基因,或,权利要求7或8所述方法,在植物育种中的应用。
10.如权利要求9所述的应用,其特征在于:所述育种的目的为选育产量高和/或耐逆性高和/或白叶枯病抗性和/或黑条矮缩病抗性提高的植物。
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CN109777792A (zh) * | 2018-09-29 | 2019-05-21 | 中国科学院动物研究所 | 一种rna解旋酶3及其编码基因和应用 |
CN110713527A (zh) * | 2018-07-11 | 2020-01-21 | 中国农业大学 | Bin2及其编码基因在调控植物耐盐中的应用 |
CN112812162A (zh) * | 2021-02-08 | 2021-05-18 | 江苏省农业科学院 | 一种水稻抗性相关基因及其应用 |
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CN110713527A (zh) * | 2018-07-11 | 2020-01-21 | 中国农业大学 | Bin2及其编码基因在调控植物耐盐中的应用 |
CN109777792A (zh) * | 2018-09-29 | 2019-05-21 | 中国科学院动物研究所 | 一种rna解旋酶3及其编码基因和应用 |
CN109777792B (zh) * | 2018-09-29 | 2020-12-11 | 中国科学院动物研究所 | 一种rna解旋酶3及其编码基因和应用 |
CN112812162A (zh) * | 2021-02-08 | 2021-05-18 | 江苏省农业科学院 | 一种水稻抗性相关基因及其应用 |
CN112812162B (zh) * | 2021-02-08 | 2022-07-19 | 江苏省农业科学院 | 一种水稻抗性相关基因及其应用 |
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