CN107326033B - OsROC4基因在调控水稻表皮蜡质合成中的应用 - Google Patents
OsROC4基因在调控水稻表皮蜡质合成中的应用 Download PDFInfo
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Abstract
OsROC4基因在调控水稻表皮蜡质合成中的应用,本发明的目的是提供水稻HD‑ZIP Ⅳ家族转录因子OsROC4基因、其编码蛋白及其应用。该OsROC4基因的核苷酸序列如序列表中SEQ ID NO:1所示。编码蛋白的氨基酸序列如SEQ ID NO:2所示。本发明将OsROC4基因在水稻中过量表达或敲除,通过一系列实验证明OsROC4基因过量表达转基因水稻表皮蜡质致密,耐旱性增强,相反,roc4突变体表皮蜡质稀疏,耐旱性减弱。本发明OsROC4基因通过调控水稻表皮蜡质合成,进而调控植物对干旱胁迫的反应。
Description
技术领域
本发明涉及一种OsROC4基因、其编码蛋白及其应用。
背景技术
由于全球气候变化,淡水资源短缺,以及日益增长的人口,干旱已成为制约世界水稻产量的最主要因素之一。水稻自身形成多种抵御干旱胁迫的机制,其中表皮蜡质结构为防止水分通过非气孔途径蒸发提供了重要的屏障作用,增加蜡质含量可显著提高水稻的抗旱能力。近年来,研究人员克隆了许多蜡质合成途径的基因,这些基因能够调节蜡质数量或组成,改变表皮蜡质结构,从而调控植物对干旱胁迫的反应,然而对于调控蜡质合成的转录因子研究较少。
发明内容
本发明的目的是提供水稻HD-ZIPⅣ家族转录因子OsROC4基因、其编码蛋白及其应用。
本发明水稻HD-ZIPⅣ家族转录因子OsROC4基因的核苷酸序列如序列表中SEQ IDNO:1所示。
本发明水稻HD-ZIPⅣ家族转录因子OsROC4基因的编码蛋白的氨基酸序列如SEQID NO:2所示。
本发明OsROC4基因在调控植物对干旱胁迫的反应中的应用。其中所述植物为水稻。
本发明OsROC4基因在调控水稻表皮蜡质合成中的应用。
本发明的有益效果:
本发明在分子水平上首次成功地克隆出水稻HD-ZIPⅣ家族转录因子OsROC4基因。
本发明通过遗传转化手段,将OsROC4基因在水稻中过量表达或敲除,通过一系列实验证明OsROC4基因过量表达转基因水稻表皮蜡质致密,耐旱性增强,相反,roc4突变体表皮蜡质稀疏,耐旱性减弱。
通过转基因技术将水稻OsROC4基因在水稻中过量表达,并获得了稳定遗传的高世代转基因材料。表型实验分析表明,转基因材料与野生型相比,表皮蜡质致密,耐旱性增强;而利用CRISPR/CAS9介导的基因编辑技术,获得纯合的roc4突变体,表皮蜡质稀疏,耐旱能力减弱。水稻HD-ZIPⅣ家族转录因子OsROC4的发现,将充实水稻表皮蜡质合成网络在水稻干旱胁迫反应中作用的理论依据,对提高水稻抵御干旱等逆境胁迫能力具有重要的应用价值,并对水稻产量的提高提供较大的实践空间,具有广阔的应用前景。
附图说明
图1为OsROC4过表达植株基因表达水平分析;
图2为OsROC4过表达植株蛋白水平分析;
图3为roc4突变体打靶位点示意图;
图4为打靶效果检测结果;
图5为扫描电镜观察OsROC4-OE和roc4表皮蜡质结构图;
图6为OsROC4-OE和roc4叶绿素浸提速率测定结果;
图7为OsROC4-OE和roc4表皮蜡质组分含量测定结果;
图8为正常浇灌下OsROC4-OE和roc4的植株形态比较;
图9为失水3d的OsROC4-OE和roc4的植株形态比较;
图10为干旱处理7d,复水3d的OsROC4-OE和roc4的植株形态比较;
图11为OsROC4-OE和roc4干旱处理后复水成活率统计结果;
图12为OsROC4-OE和roc4叶片失水率测定结果;
图13为OsROC4蛋白稳定性分析;
图14为OsROC4蛋白自由降解速率;
图15为MG132处理对OsROC4蛋白稳定性的影响;
图16为MG132处理对OsROC4蛋白稳定性影响的数据统计;
图17为激光共聚焦显微镜观察MG132对OsROC4蛋白稳定性的影响;
图18为OsROC4蛋白的自身泛素化;
图19为酵母双杂交验证OsDHS和OsROC4的相互作用;
图20为体外Pull-down验证OsDHS和OsROC4的相互作用;
图21为双荧光素酶互补体内验证OsDHS和OsROC4的相互作用;
图22为OsDHS对OsROC4稳定性影响的体外验证;
图23为OsROC4在OsROC4-OE和roc4中的降解速率分析;
图24为OsDHS对OsROC4稳定性影响的体内验证;
图25为体内验证OsDHS对OsROC4稳定性影响的数据分析;
图26为dhs中OsROC4蛋白含量分析;
图27为dhs中OsROC4相对蛋白浓度分析;
图28为dhs中OsROC4基因相对表达水平;
图29为扫描电镜观察dhs、roc4和dhs roc4表皮蜡质结构图;
图30为dhs、roc4和dhs roc4叶绿素浸提速率分析;
图31为dhs、roc4和dhs roc4表皮蜡质组成含量分析;
图32为dhs、roc4和dhs roc4干旱处理表型分析;
图33为dhs、roc4和dhs roc4干旱处理后复水存活率分析;
图34为dhs、roc4和dhs roc4叶片失水速率分析。
具体实施方式
本发明技术方案不局限于以下所列举具体实施方式,还包括各具体实施方式间的任意组合。
具体实施方式一:本实施方式HD-ZIPⅣ家族转录因子OsROC4基因的核苷酸序列如序列表中SEQ ID NO:1所示。
本实施方式水稻HD-ZIPⅣ家族转录因子OsROC4基因能够与干旱超敏感的基因(Drought Hypersensitive,OsDHS)相互作用,共同调控水稻表皮蜡质合成。OsROC4过表达水稻表皮蜡质显著增加,从而增强水稻耐旱能力,反之,roc4突变体表皮蜡质减少,导致水稻耐旱能力降低。
具体实施方式二:本实施方式水稻HD-ZIPⅣ家族转录因子OsROC4基因的编码蛋白的氨基酸序列如SEQ ID NO:2所示。
具体实施方式三:本实施方式OsROC4基因在调控植物对干旱胁迫的反应中的应用。
具体实施方式四:本实施方式与具体实施方式三不同的是:所述植物为水稻。其它与具体实施方式三相同。
具体实施方式五:本实施方式OsROC4基因在调控水稻表皮蜡质合成中的应用。
以下实施例中如无特别说明均为常规方法。
实施例(1)水稻HD-ZIPⅣ家族转录因子OsROC4基因的克隆:
一、以水稻品种日本晴为材料,用购买自Invitrogen公司的TRIzol试剂盒的操作手册提取叶片总RNA;
二、采用DNaseⅠ处理步骤一提取的总RNA;
三、取1μg步骤二处理后的总RNA用于cDNA的合成,cDNA的合成操作按照购买自BDBiosciences Clontech公司的BD SMARTTM RACE cDNA Amplification Kit试剂盒的使用手册进行,获得cDNA;
四、以获得的cDNA为模板通过正向引物F1与反向引物R1扩增OsROC4基因,PCR反应条件如下:94℃预变性5min,94℃变性30s,58℃退火30s,72℃延伸35S,共38循环,再72℃延伸10min,将PCR产物在ABI3130测序仪(ABI公司)上进行测序;测序结果表明水稻HD-ZIPⅣ家族转录因子OsROC4基因由2442个碱基组成,其核苷酸序列如序列表中的SEQ ID NO:1所示。
正向引物F1:5'-CACCATGCAGTTCCCGTTCTCCGGC-3'
反向引物R1:5'-TCACACGTCGCAATGCAGCGC-3'
实施例(2)水稻HD-ZIPⅣ家族转录因子OsROC4过量表达水稻和roc4突变体水稻的获得
一、OsROC4过量表达水稻的获得
过量表达载体构建:将获得的OsROC4基因的cDNA,装入入门载体pENTR中,利用LR反应克隆到植物表达载体pH7WGF2中,形成一个CaMV 35S启动子驱动的融合GFP标签的OsROC4融合体质粒;
目的载体转化农杆菌EHA105,农杆菌介导法转入黑龙江水稻品种龙粳11中,针对OsDHS基因,鉴定3株以上单拷贝插入的纯合过量表达转基因水稻。
二、roc4突变体水稻的获得
CRISPR/Cas9编辑载体构建:设计一段OsROC4的打靶sgRNAs,连入CRISPR/Cas9双元载体pYLCRISPR/Cas9Pubi-H中,连接U3启动子的打靶引物F2和R2的核苷酸序列如下:
引物F2:5'-GGCAAGATGGTTACCGCGGCCCA-3'
引物R2:5'-AAACTGGGCCGCGGTAACCATCT-3'
目的载体转化农杆菌EHA105,农杆菌介导法转入黑龙江水稻品种龙粳11中,针对OsROC4基因,鉴定5株以上roc4纯合突变体。
实施例(3)OsROC4基因过表达转基因水稻分子鉴定:
一、以待鉴定OsROC4基因过表达转基因水稻和其对照为材料,用购买自Invitrogen公司的TRIzol试剂盒的操作手册提取叶片总RNA;
二、采用DNaseⅠ处理步骤一提取的总RNA;
三、取1μg步骤二处理后的总RNA用于cDNA的合成,cDNA的合成操作按照购买自BDBiosciences Clontech公司的BD SMARTTM RACE cDNA Amplification Kit试剂盒的使用手册进行,获得cDNA;
四、以获得的cDNA为模板通过正向引物F3与反向引物R3,水稻内参Actin正向引物F4与反向引物R4,以及SYBR Green PCR master mix(TransStart)进行Quantitativereal-time PCR。数据从Bio-Rad chromo 4real-time PCR detector上获得;用2-△△CT方法分析倍数变化。
正向引物F3:5'-GGTGTATGGGCTGTAGTGGA-3'
反向引物R3:5'-CCTTCGGCAGTTCATGTTT-3'
正向引物F4:5'-AGACCTTCAACACCCCTGCTATG-3'
反向引物R4:5'-TCACGCCCAGCAAGGTCG-3'
OsROC4基因过表达转基因水稻分子鉴定结果如图1(以4株具有代表性的能稳定遗传的OsROC4基因过表达转基因水稻为例)。OsROC4基因在对照Control中的表达水平设为1;OsDHS基因过量表达转基因水稻的4个株系中,OsROC4-OE1表达量是Control的9.3倍,OsROC4-OE2表达量是Control的16.6倍,OsROC4-OE3表达量是Control的12.7倍,OsROC4-OE4表达量是Control的20.8倍。
五、图2为OsROC4基因过表达转基因水稻的WESTERN检测结果。取野生型和上述OsROC4-OE4的水稻叶片,用SDS蛋白裂解液提取蛋白,用SDS-PAGE分离,用OsROC4特异抗体做WESTERN检测,结果显示四个OsROC4过表达株系中OsROC4蛋白含量显著高于野生型。
以上结果证明本实验所采用的实验材料确实是OsROC4基因过量表达的。
实施例(4)roc4突变体水稻分子鉴定:
提取转基因植物DNA,在打靶sgRNAs两端设计引物,做PCR扩展包含打靶区域的片段,测序分析打靶效果。打靶效果检测引物F5和R5序列如下:
正向引物F5:5'-CGAGTGATTCGGAAAGGC-3'
反向引物R5:5'-CATCTCGTTCTCCACCTCCAAC-3'
roc4突变体打靶位点示意图如图3,打靶效果检测结果如图4,3株roc4突变体为几个碱基缺失,选择三个不同缺失类型的roc4突变体用于下一步研究。
实施例(5)OsROC4过量表达植株和roc4突变体表皮蜡质结构观察。
扫描电镜高倍数下(1000倍)观察OsROC4过量表达植株和roc4突变体表皮蜡质结构。
图5为扫描电镜观察OsROC4过量表达植株和roc4突变体表皮蜡质结构,由图可见,OsROC4基因过量表达植株叶片表皮蜡质晶体结构比野生型对照更加致密;而roc4突变体叶片表皮蜡质晶体稀疏,局部表皮裸露在外。由此可见OsROC4基因正向调控水稻表皮蜡质发育。
实施例(6)OsROC4过量表达植株和roc4突变体叶片渗透性分析:
一、取OsROC4过量表达植株、roc4突变体和非转基因对照植株相同部位叶片,剪成2cm小段,于室温中用30ml 80%的乙醇浸泡,避光,每隔10min取0.5ml浸提液检测OD664和OD647波长下的吸光值,连续检测1h后,继续浸泡24h,取0.5ml浸提液检测上述吸光值,根据公式(7.93×A664+19.53×A647)计算各时间点叶绿素含量,除以总叶绿素含量,计算叶绿素渗出速率。
图6为叶绿素浸提速率分析(图6中曲线a表示WT,曲线b表示OsROC4基因过量表达植株,曲线c表示roc4突变体),可见OsROC4基因过量表达植株叶片的叶绿素浸提速率显著低于野生型,而roc4突变体叶片的叶绿素浸提速率显著高于野生型。以上结果可见OsROC4基因正向调控水稻表皮蜡质结构,从而影响表皮的渗透性。
实施例(7)OsROC4过量表达植株和roc4突变体蜡质组分含量测定。
一、取OsROC4过量表达植株、roc4突变体和野生型叶片,测量叶面积,然后浸入30ml正己烷中,67℃,30s,迅速取出叶片。
二、向提取液中加入50μg 24烷作为内标,液氮吹干。
三、加入100μl of bis-N,N-(trimethylsilyl)trifluoroacetamide(BSTFA,Sigma,USA)和100μl吡啶,70℃衍生化60min。
四、用Agilent 7000C气相色谱质谱联机(GC-MS/MS)检测分析表皮蜡质组分和含量。
OsROC4过量表达植株和roc4突变体表皮蜡质组分和含量分析结果如图7。OsROC4基因过量表达植株表皮蜡质几乎所有组分含量均高于野生型,而roc4突变体表皮蜡质各组分含量均低于野生型;表1是统计烷、伯醇和醛三种主要组分总含量,可以看出,OsROC4基因过量表达植株表皮蜡质中几个主要组分总含量均显著高于野生型,总蜡质含量显著增加,而roc4突变体几个主要组分总含量均显著低于野生型,总蜡质含量显著降低。由此进一步说明OsROC4基因正调控水稻表皮蜡质合成。
表1
实施例(8)OsROC4过量表达植株和roc4突变体对干旱胁迫的反应
一、每种材料挑选饱满一致的种子浸种2天,37℃催芽2天,露白后各选取生长一致的种子移栽到蛭石中,每天浇灌营养液,人工气候培养箱(型号:RXZ0450;light:14h,dark:10h)28℃进行培养。
二、培养4周后,停止浇灌,干旱处理3d后开始观察表型,干旱处理7d后,重新浇灌,复水3d后统计植株存活率。
图8-图10为OsROC4过量表达植株和roc4突变体干旱处理表型,其中图8为正常浇灌下植株形态,OsROC4过量表达植株和roc4突变体与对照相比在植株形态上没有显著差别;图9为失水3d的表型,可见野生型叶片明显卷曲,OsROC4过量表达植株叶片仍然舒展,roc4突变体叶片则严重卷曲下垂;图10为干旱处理7d,复水3d的表型,可见OsROC4过量表达植株大部分存活,存活率显著高于野生型,而roc4突变体存活率显著低于野生型;图11为复水3d植株存活率的统计结果,可以看出,野生型植株存活率为53%,OsROC4过量表达植株存活率为86%,而roc4突变体存活率为。由此可见OsROC4基因正向调控水稻对干旱胁迫的耐受能力。
实施例(9)OsROC4过量表达植株和roc4突变体的叶片失水率
一、每种材料挑选饱满一致的种子浸种2天,37℃催芽2天,露白后各选取生长一致的种子移栽到蛭石中,每天浇灌营养液,人工气候培养箱(型号:RXZ0450;light:14h,dark:10h)28℃进行培养。
二、培养3周后每个株系及其对照挑选均匀一致的叶片用电子天平(型号:FA2004)各称量0.5g,置于玻璃平皿上,然后将带有叶片的玻璃平皿置于室温、无通风环境下,每隔30min称量一次,最后计算失水率。
图12为OsROC4过量表达植株和roc4突变体叶片失水率测定结果(图12中曲线a表示WT,曲线b表示OsROC4基因过量表达植株,曲线c表示roc4突变体)。结果显示OsROC4过量表达植株叶片失水速率显著低于野生型,而roc4突变体叶片失水率显著高于野生型。综合以上结果说明OsROC4正向调控表皮蜡质合成,进而正向调控水稻的耐旱能力。
实施例(10)OsROC4蛋白稳定性分析
取OsROC4过表达水稻抗性愈伤组织,用Degradation Buffer(25mM Tris-HCl,pH7.5,10mM NaCl,10mM MgCl2,4mM PMSF,5mM DTT,and 10mM ATP)提取蛋白,离心取上请,置于室温中,间隔一定时间点取样,加入上样缓冲液,煮沸5min,用SDS-PAGE分离,用OsROC4蛋白特异的抗体做WESTERN杂交。
图13为OsROC4蛋白稳定性WESTERN检验结果,图14为OsROC4蛋白于室温中自由降解速率。由图可见,OsROC4蛋白在室温中迅速降解,置于室温2h,蛋白浓度降解为初浓度的20%,说明OsROC4是不稳定蛋白。
实施例(11)OsROC4蛋白是否经26S-蛋白酶体途径降解的验证
一、取等量的3份OsROC4-OE水稻的抗性愈伤组织,分别用蛋白合成抑制剂CHX和26S蛋白酶体途径阻断剂MG132处理4h,以同样浓度的DMSO作为对照,然后提取蛋白,用SDS-PAGE分离,用OsROC4蛋白特异的抗体做WESTERN杂交。所述OsROC4-OE水稻为为OsROC4基因过量表达转基因水稻。
二、取2-week龄的OsROC4-OE水稻幼苗,分别用CHX和MG132处理,用同样浓度的DMSO作为对照,处理4h后,取幼苗的根部,用激光共聚焦显微镜观察绿色荧光,检验OsROC4蛋白的表达情况。
图15为不同处理条件下OsROC4在植物体内浓度变化的WESTERN检测结果,图16为不同处理条件下OsROC4的相对蛋白浓度。由图可见,经MG132处理后,植物体内OsROC4蛋白浓度显著高于对照,而经CHX处理后,植物体内OsROC4蛋白浓度显著低于对照,由此可见MG132能够阻断OsROC4蛋白的降解,说明OsROC4是经26S蛋白酶体途径降解的。
图17为经CHX和MG132处理后OsROC4在植物体内的表达情况。由于OsROC4蛋白特异性的在细胞核内表达,在激光共聚焦显微镜下可以清楚地看到融合GFP标签的OsROC4在细胞核中的表达情况。从图中可以看出,经MG132处理后,带有绿色荧光的细胞核明显多于对照,而经CHX处理的材料,带有绿色荧光的细胞核显著少于对照,由此进一步说明,OsROC4的降解是经过26S-蛋白酶体途径。
实施例(12)OsROC4蛋白自身泛素化检测
取1g OsROC4-OE水稻愈伤组织,用20μM MG132处理4h,用Native extract buffer(100mM Sodium Phosphate,pH 7.8,100mM NaCl,0.1%NP-40,2mM PMSF,Completeprotease inhibitor cocktail,and 20μM MG132)提取蛋白,加入到融合OsROC4特异抗体的protein A magbeads(GenScript)中,4℃孵育3h,然后用wash buffer(100mM SodiumPhosphate,pH 7.8,100mM NaCl,0.5%NP-40,2mM PMSF,Complete protease inhibitorcocktail,and 20μM MG132)洗3次,加入上样缓冲液,煮沸5min,用SDS-PAGE分离,分别用OsROC4蛋白特异的抗体和泛素抗体做WESTERN杂交。
图18为OsROC4蛋白自身泛素化检验结果。无论用OsROC4蛋白特异的抗体,还是泛素抗体杂交结果,在OsROC4蛋白条带上方均可见弥散的条带,是OsROC4蛋白多泛素化的条带,说明OsROC4蛋白在植物体内自身具有多泛素化形式,进一步说明OsROC4是有26S蛋白酶体途径降解。
实施例(13)OsDHS与OsROC4的相互作用。
一、酵母双杂交:将OsDHS基因的CDS插入pGBKT7中,构建BD-DHS载体,将OsROC4的CDS插入到pGADT7中,构建AD-ROC4载体,用酵母菌Y2H gold做酵母双杂交,用SD/-Leu/-Trp筛选互作的阳性克隆。所述OsDHS基因的核苷酸序列如序列表中SEQ ID NO:13所示。
二、体外蛋白Pull-down:将OsDHS的CDS连接到融合MBP标配的原核表达载体pMal–c2x中,将OsROC4的CDS连接到融合GST的原核表达载体pDEST15中,分别在大肠杆菌BL21中表达,提取蛋白,纯化后,用Amylose Resin做Pull-down,取上清分别用MBP-抗体和GST-抗体杂交检验Input情况,沉淀用GST-抗体杂交,检验Pull-down结果。
三、双萤光素酶互补分析:将OsDHS的CDS连接到pCAMBIA-nLUC载体中,将OsROC4的CDS连接到pCAMBIA-cLUC载体中,分别转化农杆菌GV3101,共同注射烟草叶片,2d后观察烟草叶片的荧光素酶活性。
图19-21分别是酵母双杂交、Pull-down和双萤光素酶互补验证OsDHS与OsROC4的相互作用结果,经过酵母双杂交验证,OsDHS与OsROC4能够相互作用,通过体外Pull-down和双萤光素酶互补分析进一步验证,OsDHS与OsROC4能够相互作用。
实施例(14)OsDHS对OsROC4蛋白稳定性的影响
一、体外验证OsDHS对OsROC4蛋白稳定性的影响:分别取OsROC4-OE、OsDHS-OE、dhs突变体和野生型(WT)水稻的抗性愈伤组织,用Degradation Buffer(配方为:25mM Tris-HCl,pH7.5,10mM NaCl,10mM MgCl2,4mM PMSF,5mM DTT,30and10mM ATP)提取蛋白,将OsROC4-OE蛋白提取液平均分为3份,加入等量的OsDHS-OE、dhs和WT蛋白提取液,置于室温中,间隔一定时间点取样,加入上样缓冲液终止反应,煮沸5min,用SDS-PAGE分离,用OsROC4蛋白特异性抗体做WESTERN杂交。其中OsDHS-OE为OsDHS基因过量表达转基因水稻。
二、体内瞬时表达分析OsDHS对OsROC4蛋白稳定性的影响:将OsROC4,OsDHS andOsDHSC95S插入瞬时表达载体PRT107中,构建35Spro:OsROC4,35Spro:OsDHS和35Spro:OsDHSC95S载体;用野生型水稻提取原生质体,将35Spro:OsROC4分别与35Spro:OsDHS和35Spro:OsDHSC95S共同转化水稻原生质体,以35Spro:OsROC4和空PRT107共同转化的水稻原生质体作为对照,28℃孵育过夜。提取转化不同质粒的水稻原生质体蛋白,用OsROC4特异抗体做WESTERN杂交。
OsDHSC95S的制备方法为:将OsDHS全长编码区序列插入融合MBP标签的原核表达载体pMal-c2x中;同时将OsDHS蛋白RING结构域的保守氨基酸Cys95定点突变为Ser,表示为OsDHSC95S。
三、体内验证OsDHS对OsROC4蛋白稳定性的影响:分别提取野生型和dhs突变体蛋白,用OsROC4特异抗体做WESTERN杂交;提取同样材料的RNA,反转录为cDNA,用引物F3和R3,利用real-time QRT-PCR检验OsROC4基因表达水平(方法同前)。
图22和23为OsROC4蛋白在OsDHS-OE和dhs蛋白中的降解速率(图23中a表示WT,b表示OsDHS-OE,c表示dhs),可以看出OsROC4在OsDHS-OE中降解速度显著快于野生型,而在dhs中的降解速度显著慢于野生型,由此可见OsDHS促进OsROC4的降解。
图24和25为瞬时表达分析OsDHS对OsROC4蛋白稳定性的影响,可以看出OsROC4与OsDHS水稻原生质体中共表达,OsROC4相对蛋白含量显著低于对照,而当OsROC4与OsDHSC95S在水稻原生质体中共表达时,OsROC4相对蛋白含量与对照无显著差异,进一步证明OsDHS促进OsROC4的降解,并且OsROC4有可能是OsDHS的泛素化底物。
图26和27为OsROC4在dhs突变体中的蛋白含量,图28为dhs突变体中OsROC4基因表达水平,与野生型对照相比,OsROC4在dhs突变体中的相对蛋白含量显著提高,而dhs中OsROC4基因转录水平与野生型对照并无显著差异。
综合以上结果说明,OsDHS促进OsROC4的降解,并且OsROC4可能是OsDHS的泛素化底物。
实施例(15)OsDHS与OsROC4的遗传学关系分析
一、将dhs与roc4进行杂交,获得dhs roc4双突变体;
二、通过扫描电镜观察表皮蜡质结构,叶绿素浸提速率检测、表皮蜡质组成含量测定,分析dhs、roc4和dhs roc4的表皮蜡质结构和组成差异(方法同上)
三、对dhs、roc4和dhs roc4进行干旱处理,不同突变体对干旱胁迫的反应,检测3种突变体的叶片失水率(方法同上)。
图29为扫描电镜观察dhs、roc4和dhs roc4的表皮蜡质结构(图29中a表示WT,b表示dhs,c表示roc4,d表示dhs roc4),可以看出dhs roc4的表皮蜡质结构稀少与roc4类似;图30为dhs、roc4和dhs roc4的叶绿素浸提速率(图30中a表示WT,b表示dhs,c表示roc4,d表示dhs roc4),dhs的叶绿素浸提速率显著慢于野生型,而dhs roc4的叶绿素浸提速率显著快于dhs;图31和表2为dhs、roc4和dhs roc4的表皮蜡质组成含量测定和分析,与野生相比,dhs中蜡质含量显著增加,而dhs roc4中蜡质含量显著降低。
表2
图32为dhs、roc4和dhs roc4的干旱处理表型(图32中a表示干旱处理前,b表示干旱处理3d,c表示干旱处理7d后复水3d),dhs roc4对干旱胁迫的耐受能力与roc4类似,显著低于dhs的耐旱能力;图33为干旱处理7d复水3d后dhs、roc4和dhs roc4材料的存活率,干旱处理后,约86%的dhs存活,而roc4仅有约36%存活,dhs roc4仅有约48%存活;图34为dhs、roc4和dhs roc4的叶片失水速率,dhs的叶片失水速率限制慢于野生型,而dhs roc4的叶片失水速率显著快于dhs。
综合以上结果,说明dhs突变体中OsROC4蛋白积累较多,导致其蜡质含量增加,耐旱能力提高,说明OsROC4位于OsDHS的遗传学下游。
序 列 表
<110> 中国科学院东北地理与农业生态研究所
<120> 水稻HD-ZIP Ⅳ家族转录因子OsROC4基因、其编码蛋白及其应用
<160> 13
<210> 1
<211>2442
<212> DNA
<213> 粳亚种(Oryza sativa L. japonica. cv. Nipponbare)
<220>
<223> 水稻HD-ZIP Ⅳ家族转录因子OsROC4基因
<400> 1
atgcagttcc cgttctccgg cgctggcccg ggcgtcttca cgtcatcgcc ggcgctctcc 60
ctcgcgctgg cggacgcggt ggcaggccgg aacagcggcg gcggtgggaa gatggttacc 120
gcggcccatg gcggcgtcgg cggaggagga ggaggaggac gcgcgaaggc gagggacgcg 180
ttggaggtgg agaacgagat gagccggtcc gggagcgacc acctcgacgt cgtctcttgc 240
ggcgacgcgg gcggcggcgg cggcgacgac gacgatgacg aggacgccga gcacggcaac 300
ccgcccaagc gcaagaagcg gtaccaccgc cacacgccgc agcagatcca agagctggaa 360
gcgatgttca aggaatgccc ccacccagac gagaagcagc gcgccgagct gagcaagcgg 420
ctcggcctcg aaccccggca ggtcaagttc tggttccaga atcggcgaac gcagatgaag 480
atgcaactgg agcgacacga gaactcgctg ctgaagcagg agaacgataa gctgcggtcc 540
gagaacctgt caatccggga ggccacgagc aacgcggtgt gcgttggctg cggcggcccg 600
gcgatgctcg gggaggtgtc cctggaggag caccaccttc gcgtcgagaa cgcgaggctc 660
aaggacgagc tcagccgagt gtgcgcgctc gccgccaagt tccttggcaa gtccatctct 720
gtcatggcgc caccgcagat gcatcagcct catcctgtgc caggctcgtc gctggagctt 780
gcggttgggg gtatcggttc gatgccatca gccacgatgc ccatctcgac gatcactgat 840
tttgctggcg ccatgtccag ttcaatgggc acggtgatca cgcccatgaa gtctgaggct 900
gaaccatcgg caatggctgg cattgacaag tccttgttct tggagctagc aatgagtgca 960
atggatgagc tagtcaagat ggctcagatg ggggatccgc tatggattcc aggtgcctcc 1020
gtaccttcct cgccggcaaa ggagagtcta aacttcgagg agtacctaaa caccttccca 1080
ccttgcatcg gggtgaagcc tgaagggtat gtatcagagg catctagaga atctggcatt 1140
gtcatcattg acgatggcgc cgcgcttgtg gagaccctca tggatgagcg acggtggtcc 1200
gatatgttct catgcatgat tgccaaggca tcaaccactg aggagatttc tactggtgtt 1260
gctgggagta gaaatggtgc attgcttctt gtgagtgatg aacattctgt tatgcaggca 1320
gagctacagg tgctttctcc tcttgtgcct attagagagg tgaagtttct caggttctcc 1380
aaacagctgg ctgatggtgt atgggctgta gtggacgttt cggctgatga attgatgagg 1440
gatcagggca ttacttctgc atcctcgact gcaaacatga actgccgaag gctgccttct 1500
ggttgtgtgc tgcaggacac tccaaatggg tttgttaagg tcacatgggt tgaacataca 1560
gaatatgatg aggcatctgt gcacccgctc taccggcctc ttctccggtc tggtcttgcc 1620
cttggtgcag ggcgatggat cgcgacatta cagcggcagt gcgaatgctt ggcccttctc 1680
atgtcttcta ttgcattgcc agagaacgac tcatcagcta tccatcctga aggtaaacgg 1740
agcatgttga agttggcaag gaggatgacg gacaacttct gtgcaggggt gagcacatca 1800
tctacccgtg aatggagcaa actggttgga ttgacaggca acattgggga ggatgtgcat 1860
gtaatggcgc ggaagagtgt ggatgaacct ggaacgccgc caggtgtggt gcttagtgct 1920
gctacatctg tgtggatgcc tgtgatgcct gaacggctct tcaacttctt gcacaacaag 1980
gggctgcgtg ctgaatggga tatcctcagc aatggtggcc ctatgcagga ggtgacaagc 2040
attgccaagg ggcaacagaa tggcaatacc gtatgtctac tgaaggctag tcccaccaaa 2100
gacaagcaga acagcatgct gatcctacag gagacgtgtg cagacgcatc cggttcaatg 2160
gttgtgtatg ctcctgtaga catcccagca atgcaccttg tcatgagtgg tggggattcg 2220
tcatgcgtcg cccttcttcc atcaggtttt gccatcctgc ctgctgggcc tagcatcggc 2280
gcagatcaca agatgggcgg ttcattgctc accgttgcat tccagatact tgccaacagc 2340
cagcccagtg ctaagctcac ggtggagtca gtcgagaccg tgagcaacct tatctcctgc 2400
accatcaaga agatcaagac ggcgctgcat tgcgacgtgt ga 2442
<210> 2
<211>813
<212> PRT
<213>粳亚种(Oryza sativa L. japonica. cv. Nipponbare)
<220>
<223> 水稻HD-ZIP Ⅳ家族转录因子OsROC4基因编码蛋白
<400> 2
Met Gln Phe Pro Phe Ser Gly Ala Gly Pro Gly Val Phe Thr Ser
5 10 15
Ser Pro Ala Leu Ser Leu Ala Leu Ala Asp Ala Val Ala Gly Arg
20 25 30
Asn Ser Gly Gly Gly Gly Lys Met Val Thr Ala Ala His Gly Gly
35 40 45
Val Gly Gly Gly Gly Gly Gly Gly Arg Ala Lys Ala Arg Asp Ala
50 55 60
Leu Glu Val Glu Asn Glu Met Ser Arg Ser Gly Ser Asp His Leu
65 70 75
Asp Val Val Ser Cys Gly Asp Ala Gly Gly Gly Gly Gly Asp Asp
80 85 90
Asp Asp Asp Glu Asp Ala Glu His Gly Asn Pro Pro Lys Arg Lys
95 100 105
Lys Arg Tyr His Arg His Thr Pro Gln Gln Ile Gln Glu Leu Glu
110 115 120
Ala Met Phe Lys Glu Cys Pro His Pro Asp Glu Lys Gln Arg Ala
125 130 135
Glu Leu Ser Lys Arg Leu Gly Leu Glu Pro Arg Gln Val Lys Phe
140 145 150
Trp Phe Gln Asn Arg Arg Thr Gln Met Lys Met Gln Leu Glu Arg
155 160 165
His Glu Asn Ser Leu Leu Lys Gln Glu Asn Asp Lys Leu Arg Ser
170 175 180
Glu Asn Leu Ser Ile Arg Glu Ala Thr Ser Asn Ala Val Cys Val
185 190 195
Gly Cys Gly Gly Pro Ala Met Leu Gly Glu Val Ser Leu Glu Glu
200 205 210
His His Leu Arg Val Glu Asn Ala Arg Leu Lys Asp Glu Leu Ser
215 220 225
Arg Val Cys Ala Leu Ala Ala Lys Phe Leu Gly Lys Ser Ile Ser
230 235 240
Val Met Ala Pro Pro Gln Met His Gln Pro His Pro Val Pro Gly
245 250 255
Ser Ser Leu Glu Leu Ala Val Gly Gly Ile Gly Ser Met Pro Ser
260 265 270
Ala Thr Met Pro Ile Ser Thr Ile Thr Asp Phe Ala Gly Ala Met
275 280 285
Ser Ser Ser Met Gly Thr Val Ile Thr Pro Met Lys Ser Glu Ala
290 295 300
Glu Pro Ser Ala Met Ala Gly Ile Asp Lys Ser Leu Phe Leu Glu
305 310 315
Leu Ala Met Ser Ala Met Asp Glu Leu Val Lys Met Ala Gln Met
320 325 330
Gly Asp Pro Leu Trp Ile Pro Gly Ala Ser Val Pro Ser Ser Pro
335 340 345
Ala Lys Glu Ser Leu Asn Phe Glu Glu Tyr Leu Asn Thr Phe Pro
350 355 360
Pro Cys Ile Gly Val Lys Pro Glu Gly Tyr Val Ser Glu Ala Ser
365 370 375
Arg Glu Ser Gly Ile Val Ile Ile Asp Asp Gly Ala Ala Leu Val
380 385 390
Glu Thr Leu Met Asp Glu Arg Arg Trp Ser Asp Met Phe Ser Cys
395 400 405
Met Ile Ala Lys Ala Ser Thr Thr Glu Glu Ile Ser Thr Gly Val
410 415 420
Ala Gly Ser Arg Asn Gly Ala Leu Leu Leu Val Ser Asp Glu His
425 430 435
Ser Val Met Gln Ala Glu Leu Gln Val Leu Ser Pro Leu Val Pro
440 445 450
Ile Arg Glu Val Lys Phe Leu Arg Phe Ser Lys Gln Leu Ala Asp
455 460 465
Gly Val Trp Ala Val Val Asp Val Ser Ala Asp Glu Leu Met Arg
470 475 480
Asp Gln Gly Ile Thr Ser Ala Ser Ser Thr Ala Asn Met Asn Cys
485 490 495
Arg Arg Leu Pro Ser Gly Cys Val Leu Gln Asp Thr Pro Asn Gly
500 505 510
Phe Val Lys Val Thr Trp Val Glu His Thr Glu Tyr Asp Glu Ala
515 520 525
Ser Val His Pro Leu Tyr Arg Pro Leu Leu Arg Ser Gly Leu Ala
530 535 540
Leu Gly Ala GLy Arg Trp Ile Ala Thr Leu Gln Arg Gln Cys Glu
545 550 555
Cys Leu Ala Leu Leu Met Ser Ser Ile Ala Leu Pro Glu Asn Asp
560 565 570
Ser Ser Ala Ile His Pro Glu Gly Lys Arg Ser Met Leu Lys Leu
575 580 585
Ala Arg Arg Met Thr Asp Asn Phe Cys Ala Gly Val Ser Thr Ser
590 595 600
Ser Thr Arg Glu Trp Ser Lys Leu Val Gly Leu Thr Gly Asn Ile
605 610 615
Gly Glu Asp Val His Val Met Ala Arg Lys Ser Val Asp Glu Pro
620 625 630
Gly Thr Pro Pro Gly Val Vla Leu Ser Ala Ala Thr Ser Val Trp
635 640 645
Met Pro Val Met Pro Glu Arg Leu Phe Asn Phe Leu His Asn Lys
650 655 660
Gly Leu Arg Ala Glu Trp Asp Ile Leu Ser Asn Gly Gly Pro Met
665 670 675
Gln Glu Val Thr Ser Ile Ala Lys Gly Gln Gln Asn Gly Asn Thr
680 685 690
Val Cys Leu Leu Lys Ala Ser Pro Thr Lys Asp Lys Gln Asn Ser
695 700 705
Met Leu Ile Leu Gln GLu Thr Cys Ala Asp Ala Ser Gly Ser Met
710 715 720
Val Val Tyr Ala Pro Val Asp Ile Pro Ala Met His Leu Val Met
725 730 735
Ser Gly Gly Asp Ser Ser Cys Val Ala Leu Leu Pro Ser Gly Phe
740 745 750
Ala Ile Leu Pro Ala Gly Pro Ser Ile Gly Ala Asp His Lys Met
755 760 765
Gly Gly Ser Leu Leu Thr Val Ala Phe Gln Ile Leu Ala Asn Ser
770 775 780
Gln Pro Ser Ala Lys Leu Thr Val Glu Ser Val Glu Thr Val Ser
785 790 795
Asn Leu Ile Ser Cys Thr Ile Lys Lys Ile Lys Thr Ala Leu His
800 805 810
Cys Asp Val
816
<210> 3
<211> 25
<212> DNA
<213>人工序列
<220>
<223> 正向引物F1
<400> 3
caccatgcagttcccgttctccggc 25
<210>4
<211> 21
<212> DNA
<213>人工序列
<220>
<223> 反向引物R1
<400> 4
tcacacgtcgcaatgcagcgc 21
<210> 5
<211> 23
<212> DNA
<213>人工序列
<220>
<223> 正向引物F2
<400> 5
ggcaagatggttaccgcggccca 23
<210>6
<211> 23
<212> DNA
<213>人工序列
<220>
<223> 反向引物R2
<400> 6
aaactgggccgcggtaaccatct 23
<210> 7
<211> 20
<212> DNA
<213>人工序列
<220>
<223> 正向引物F3
<400> 7
ggtgtatgggctgtagtgga 20
<210>8
<211> 19
<212> DNA
<213>人工序列
<220>
<223> 反向引物R3
<400> 8
ccttcggcagttcatgttt 19
<210> 9
<211> 23
<212> DNA
<213>人工序列
<220>
<223> 正向引物F4
<400> 9
agaccttcaacacccctgctatg 23
<210>10
<211> 18
<212> DNA
<213>人工序列
<220>
<223> 反向引物R4
<400> 10
tcacgcccagcaaggtcg 18
<210> 11
<211> 18
<212> DNA
<213>人工序列
<220>
<223> 正向引物F5
<400> 11
cgagtgattcggaaaggc 18
<210>12
<211> 22
<212> DNA
<213>人工序列
<220>
<223> 反向引物R5
<400> 12
catctcgttctccacctccaac 22
<210> 13
<211> 495
<212> DNA
<213>粳亚种(Oryza sativa L. japonica. cv. Nipponbare)
<220>
<223> 水稻RING finge家族E3泛素连接酶OsDHS基因
<400> 13
atggggttcc ctctggtgtg ctactgcatg gcgatcccca agccgctcat cgccttggcc 60
aagctcctcg ccgccatcag ggaggccctc cagctgatgc tcttcgtcgt cgggatctgc 120
caccacccgg agcgatcggg ccgcccggct gccgtcgacg ccccgctgcc cgacgaggtg 180
aaggaccgcc tcccgcccct cgagttcgcc cagctgctcg cggcctcgga gcacggctgt 240
catggctgcg acgacgacga ggcggtggcg gggtgcatcg tgtgcctgga gaggctggag 300
gcggatgacg tggtgcggcg gctgggcaac tgcgcgcacg cgttccaccg cggctgcatc 360
gaccggtgga tcgacctcgg ccggttgacg tgcccgctgt gccgctccac cctgctgccg 420
cgcgcgcgcc ccgccgccgg cccgcgcggg cgactgggcc gcctcgccac ccgcctcacg 480
ggcgtcgttt ggtga 495
Claims (1)
1.OsROC4基因在调控水稻表皮蜡质合成中的应用,所述OsROC4基因的核苷酸序列如序列表中SEQ ID NO:1所示。
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