CN112500461B - 一种水稻叶色相关蛋白las5在调控水稻叶色中的应用 - Google Patents
一种水稻叶色相关蛋白las5在调控水稻叶色中的应用 Download PDFInfo
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- CN112500461B CN112500461B CN202011326013.XA CN202011326013A CN112500461B CN 112500461 B CN112500461 B CN 112500461B CN 202011326013 A CN202011326013 A CN 202011326013A CN 112500461 B CN112500461 B CN 112500461B
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Abstract
本发明公开了一种水稻叶色相关蛋白LAS5在调控水稻叶色中的应用,属于分子生物学及基因工程技术领域。水稻叶色相关蛋白LAS5的氨基酸序列如SEQ ID No.2所示,在水稻中转入抑制LAS5蛋白表达的物质,例如转入具有靶标序列的sgRNA表达载体,可促使水稻叶绿体中类囊体发育异常而导致水稻叶片白化。通过将抑制水稻叶色相关蛋白LAS5的表达的物质导入杂交亲本中作为一种形态学标记,可以保持杂交种的纯度。水稻叶色相关蛋白LAS5的作用使得其发现和相关突变体的鉴定以及对水稻遗传改良具有重要的意义。
Description
技术领域
本发明属于分子生物学及基因工程技术领域,更具体地说,涉及一种水稻叶色相关蛋白LAS5在调控水稻叶色中的应用。
背景技术
水稻是最重要的粮食作物之一,保持与稳步提高水稻的产量对保障全世界的粮食安全具有重要的作用。叶色变异直接影响水稻的光合作用,最终导致产量降低。目前,水稻中已克隆了超过120个叶色相关基因,初步阐明了叶绿素合成和降解的分子机理。然而,水稻叶色形成的机理十分复杂,解析的叶色相关基因仍显较少,亟待克隆新的叶色变异控制基因。
叶绿体是绿色植物参与调节新陈代谢和调节功能的含有遗传物质的半自主细胞器。在拟南芥叶绿体中大约含有2500种蛋白,叶绿体内的蛋白酶识别介导叶绿体蛋白的降解。目前,叶绿体蛋白酶主要有Clp蛋白酶、Lon蛋白酶、FtsH金属蛋白酶、DegP蛋白酶、SppA和EGY。通过正向和反向遗传学的手段,拟南芥多数Clp蛋白酶基因的功能被解析。Clp突变体表现出胚/苗死或黄化叶的表型。在水稻中,仅有OsClpP5与VYL的相关报道,其他Clp蛋白酶的组成以及功能鲜有报道。借鉴拟南芥Clp蛋白酶参与调节植物光合作用和新陈代谢,对水稻Clp蛋白开展功能的研究,有助于解析植物光合作用的调控机制,从而为培育高光效的水稻奠定基础。
发明内容
针对现有技术存在的上述问题,本发明所要解决的技术问题在于提供一种水稻叶色蛋白LAS5在调控水稻叶色中的应用。
为了解决上述技术问题,本发明所采用的技术方案如下:
水稻叶色相关蛋白LAS5在调控水稻叶色中的应用,所述的水稻叶色相关蛋白LAS5的氨基酸序列如SEQ ID No.2所示。
进一步地,所述的水稻叶色相关蛋白LAS5在调控水稻叶色中的应用是在水稻中转入抑制所述的水稻叶色相关蛋白LAS5表达的物质,促使水稻叶绿体中类囊体发育异常而导致水稻叶片白化。
进一步地,所述抑制LAS5蛋白表达的物质为重组载体、含有重组载体的转基因细胞系或重组菌。
进一步地,所述重组载体为具有靶标序列的sgRNA表达载体,所述靶标序列为SEQID NO.1所示的自5′端第164位-193位核苷酸序列。
进一步地,所述的应用,包括以下步骤:
1)构建具有靶标序列的sgRNA表达载体;
2)将所构建的具有靶标序列的sgRNA表达载体转入根癌农杆菌;
3)将阳性根癌农杆菌转化到水稻细胞;
4)培育筛选,得到叶片白化的转基因水稻。
进一步地,所述根癌农杆菌为EHA105。
相比于现有技术,本发明的有益效果为:
在杂交育种过程种,通过将上述水稻叶色相关蛋白LAS5的表达的物质导入杂交亲本中作为一种形态学标记,可以保持杂交种的纯度。水稻叶色相关蛋白LAS5的作用使得其发现和相关突变体的鉴定以及对水稻遗传改良具有重要的意义。
附图说明
图1为实施例1中野生型Dongjin与突变体las5的叶色表型鉴定图;
图2为实施例1中野生型和突变体叶绿体透射电镜观察图;图中,A、B是野生型水稻叶绿体的类囊体和基粒图;C、D是las5突变的水稻叶绿体结构图,其类囊体和基粒缺乏;
图3为是实施例1中突变体las5的T-DNA插入位点鉴定图;
图4为是实施例1中CRISPR/Cas9敲除基因LAS5的表型鉴定图。
具体实施方式
下面结合具体实施例对本发明进一步进行描述。
实施例1:水稻叶色相关突变体las5的表型鉴定
从水稻品种Dongjin T-DNA插入突变体库中筛选获得叶色相关突变体las5(1ethal albinic seedling 5)。
与野生型相比,纯合突变体las5表现出叶片白化的表型,杂合突变体叶色正常(图1A-B)。采用95%无水乙醇提取色素,取约200mg叶片剪碎,置于装有相应体积95%无水乙醇中,室温黑暗放置48小时。利用紫外可见分光光度计(TU1901)测定663nm、645nm和470nm处的吸光值,计算出单位质量的叶绿素含量。结果表明,相比野生型,突变体las5叶绿素含量显著降低,导致突变植株不能进行光合作用,最终死亡(图1C)。
取2叶期水稻苗叶片切成小块,固定于含2.5%的戊二醛的固定液中,抽真空1小时,室温放置24小时,而后用1%锇酸固定4小时,经不同浓度梯度乙醇脱水后,进行包埋、切片,醋酸铀染色后,利用透射电子显微镜(JEM-1200EX)观察。透射电镜结果表明,野生型叶绿体结构完整,类囊体和基粒结构完整,而突变体没有成熟的叶绿体,类囊体和基粒缺乏,导致植物不能正常合成叶绿素(图2A-D)。
综上所述,突变体las5由于叶绿体发育异常,叶绿素不能正常合成,导致植物出现白化的表型。
实施例2:LAS5的克隆与基因编辑验证
1、LAS5的克隆
为了克隆突变体las5的突变基因,我们根据T-DNA插入突变体的侧翼序列设计三条检测引物P1(如SEQ ID No.3所示)、P2(如SEQ ID No.4所示)和P3(如SEQ ID No.5所示)(图3A)。PCR扩增结果表明,引物P1+P2在野生型中能扩增出目的片段,而在突变体中不能扩增出目的片段。同时,引物P2+P3在突变体中可以扩增出目的片段,而在野生型中则不能(图3B),证明该T-DNA真实插入,进一步对扩增片段测序,结合生物信息学分析,结果表明该T-DNA插在基因LOC_Os05g51450第一外显子上(图3A),LOC_Os05g51450转录后翻译表达的蛋白为LAS5,其氨基酸序列如SEQ ID No.2所示。根据已公布的水稻日本晴基因组序列,结合基因测序,获得了编码LAS5蛋白的基因LOC_Os05g51450全长cDNA序列,如SEQ ID No.1所示。
2、T-DNA插入位点基因的表达分析
T-DNA插入到基因LOC_Os05g51450第一外显子上,可能抑制其转录,因而我们对野生型和突变体中LOC_Os05g5145的表达水平进行分析。
取野生型及突变体2叶期时叶片液氮磨碎,使用康为世纪公司植物RNA小量提取纯化试剂盒提取总RNA。使用Nano Drop检测提取所获得的总RNA的浓度和质量,并使用凝胶电泳检测RNA的完整性。选用大连宝生物公司的反转录试剂盒(PrimeScript ReverseTranscriptase kit)对总RNA进行反转cDNA。以cDNA为模板,选用SYBR premix Ex TaqTM(TaKaRa,Japan)试剂盒进行Real-time PCR,在伯乐荧光定量PCR仪CFX96上扩增,采用2-ΔΔCT方法进行表达分析。结果表明,相比于野生型,在突变体中,LOC_Os05g5145的转录水平显著下调(图3C)。
3、LAS5基因编辑转基因验证
为了验证las5的突变表型是否是LOC_Os05g5145的抑制表达所引起的,在LAS5基因编码区设计一个CRISPR/Cas9基因敲除靶点,靶向第1外显子。合成基于CRISPR/Cas9系统的靶序列引物,序列如下:
对LAS5-F和LAS5-R进行退火程序,形成双链DNA,作为片段插入CRISPR/Cas9基因敲除载体pCAMBIA1305。用AarI限制性内切酶(NEB)在37℃酶切上述CRISPR/Cas9双元载体,得到载体片段。用T4连接酶(Takara)连接插入片段和载体,将重组表达载体转入大肠杆菌,提取重组表达质粒测序(安徽通用生物)验证正确后,转入根癌农杆菌菌株EHA105中,侵染日本晴愈伤组织进行水稻遗传转化。
上述重组载体遗传转化获得了19株T0代转基因水稻植株。利用聚合美DNA提取试剂盒(Mei5bio)提取T0代转基因水稻植株DNA,利用KOD DNA聚合酶(TOYOBO)和基因组引物对含靶标位点的DNA片段进行扩增测序鉴定。所述引物序列为:
测序结果表明,有10株转基因植株带有目标突变,其中纯合突变体3株、杂合突变体7株。3株纯合突变体(LAS5a、LAS5b、LAS5c),全部表现出白化的表型,结果如图4所示。
以上所述仅为本发明的较佳实施例而已,并不用以限制本发明,凡在本发明的精神和原则之内所作的任何修改、等同替换和改进等,均应包含在本发明的保护范围之内。
序列表
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Claims (5)
1.水稻叶色相关蛋白LAS5在调控水稻叶色中的应用,其特征在于:所述的水稻叶色相关蛋白LAS5的氨基酸序列如SEQ ID No.2所示,在水稻中转入抑制所述的水稻叶色相关蛋白LAS5表达的物质,促使水稻叶绿体中类囊体发育异常而导致水稻叶片白化。
2.根据权利要求1所述的水稻叶色相关蛋白LAS5在调控水稻叶色中的应用,其特征在于:所述抑制LAS5蛋白表达的物质为重组载体、含有重组载体的转基因细胞系或重组菌。
3.根据权利要求2所述的水稻叶色相关蛋白LAS5在调控水稻叶色中的应用,其特征在于:所述重组载体为具有靶标序列的sgRNA表达载体,所述靶标序列为SEQ ID NO .1所示的自5'端第164位-193位核苷酸序列。
4.根据权利要求3所述的水稻叶色相关蛋白LAS5在调控水稻叶色中的应用,其特征在于,包括以下步骤:
1)构建具有靶标序列的sgRNA表达载体;
2)将所构建的具有靶标序列的sgRNA表达载体转入根癌农杆菌;
3)用阳性根癌农杆菌转化水稻细胞;
4)培育筛选,得到叶片白化的转基因水稻。
5.根据权利要求4所述的水稻叶色相关蛋白LAS5在调控水稻叶色中的应用,其特征在于,所述根癌农杆菌为EHA105。
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