甘蓝春化相关基因BoVIN3的克隆及分析
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摘要
结球甘蓝(Brassica oleracea var. captata L.)属于绿体春化型植物,幼苗需长到一定大小,方能感应低温通过春化。长时间低温处理促进植物开花的过程称为春化作用,在植物成花转变过程中起着重要的作用,该领域的研究近年来已取得了较大的进展,特别是对春化的分子机理探讨方面也逐渐凸显出其本质,虽然对春化作用机理的理解呈现出一些轮廓,但春化作用分子调控的过程却还很不清楚。
根据拟南芥及其芸薹属相关领域的研究报道,本试验重点针对开花网络核心基因FLC及其春化作用途径上游的关键基因VIN3进行研究,首次从十字花科芸薹属作物结球甘蓝中克隆了春化相关基因VIN3,通过生物软件对VIN3基因进行了序列分析,并通过RT-PCR方法检测在结球甘蓝中的表达情况及其与春化作用的关系,以为研究结球甘蓝绿体春化的分子机理提供理论依据。本试验主要研究结果如下：
本试验中对自然条件下生长的甘蓝幼苗可以感受低温春化的形态标准研究表明：甘蓝中熟品种“光荣一号”可以感受低温春化的低限苗龄为13片叶,茎粗达到12.83±0.14mm,低温处理48d春化作用结束,开始进入花芽分化临界期。甘蓝植株只有接受低温诱导达到春化饱和程度才能启动相应基因表达,所以本试验需采用可以感受低温春化的幼苗进行春化处理,以备下一步克隆春化相关基因VIN3。
根据拟南芥VIN3基因的mRNA序列(NM-125121.3)的ORF编码区,设计了VIN3基因的特异引物,以甘蓝RNA及DNA为模板克隆BoVIN3基因的cDNA和DNA序列。克隆得到两条甘蓝春化相关基因BoVIN3的cDNA ORF编码区序列,各1680bp,GenBank登录号分别为JQ394927和JQ394928,可编码由560个氨基酸组成的蛋白质；与拟南芥VIN3基因在核酸水平上同源性分别达到85.55%和79.95%；与拟南芥氨基酸序列同源性分别为80.72%和72.96%。同时获得BoVIN3基因的部分基因组片段,1405bp,GenBank登录号为JQ394929。
利用DNAMAN等软件以及在线分析程序对BoVIN3基因及其编码的蛋白进行分析和预测。结果表明,甘蓝BoVIN3-1、BoVIN3-2蛋白长度均为560AA,分子量约分别为62.116KD和62.434KD,等电点分别为6.56和5.63；通过磷酸化位点预测其蛋白存在多个磷酸化位点；通过NCBI保守结构域预测程序得知该蛋白具有PHD结构域。
半定量RT-PCR表达分析表明BoVIN3受春化作用的诱导,只在感应低温的茎尖生长点表达,在其它部位不表达,并且其转录产物随春化时间延长逐渐积累,在春化42d时达到峰值;而在不同浓度GA3和KT诱导下不表达;FLC受春化作用的抑制,并且BoVIN3受诱导表达后才能使FLC的表达受到抑制,暗示在结球甘蓝中BoVIN3基因可能参与春化调控开花的生物学过程,属于春化特异基因,只响应低温诱导而转录。
以pBI121植物表达载体和BoVIN3-1基因片段为基础,构建反义植物表达载体。将BoVIN3-1基因片段反向插入355启动子与GUS基因之间的限制性酶切位点XbaI和SmaI中,构建含反义BoVIN3-1基因的工程质粒pB135S-BoVIN3-1。利用转反义基因技术,即转化BoVIN3-1的反义片段进行了BoVIN3-1的功能分析,通过花蕾微量注射法转化甘蓝,观察到经春化处理的转反义基因植株与对照相比,春化一定程度被推迟,这一结果进一步表明,甘蓝BoVIN3-1基因可能直接参与了春化过程。
Cabbage (Brassica oleracea var. captata L.)belongs to green vernalizatio plants,young plants need to grow to certain sizes then can accept low temperature and go through vernalization.The pr-ocess of long colding time promoting plant flowering known as vernalization, playing an important role in transition to flowering, in recent years had made considerable progress in vernalization res-earch,especially the molecular mechanism discussion highlights its natural gradually.Although the understanding of the vernalization mechanism show some outline,but the molecular control process is not clear. According to related reports in the arabidopsis and brassica,this work focusing on core gene FLC in flowering regulation way and its upper key gene VIN3in vernalization to research,first isolate vernalization related gene VIN3from cabbage by Homology cloning, employ the biological software to analyze BoVIN3gene and deduced amino acid sequence,the expression patterns of the BoVIN3genes were investigated by semi-quantity RT-PCR, in an attempt to explore the relation be-tween its expression and vernalization, provide base theory for cabbage vernalization.The main conclusions of this study are as follows:
In this experiment, the research of morphological characters of the cabbage plants can accept low temperature growth under natural condition showed that medium maturing varieties of "GUANG RONG YI HAO", with13leaves and12.83±0.14mm in stem diameter, begin to be sensitive to vernalization,low temperature treatment48d vernalization complete, transition to flower bud differentiation. Cabbage plants only accept cold inducement reached saturation level to activate the corresponding gene expression,so this work must employ cabbage plants can feel low temperature to vernalize, prepared to following cloning the VIN3gene.
According to AtVIN3gene mRNA sequence (NM-125121.3) ORF coding region,design gene specific primer,take cabbage RNA and DNA as template to amplified BoVIN3gene.Two vernalization-related genes were isolated by RT-PCR. Sequencing analysis results showed that their ORF were both1680bp in length, which encoded560amino acids, and GenBank accession number are JQ394927and JQ394928respectively.Their nucleotide sequences shared85.55%and79.95%identity with AtVIN3respectively, and the deduced amino acid sequence showed homology to AtVIN3with80.72%and72.96%. At the same time we isolated BoVIN3part DNA segment,was1405bp, GenBank accession number is JQ394929.
Take advantage of DNAMAN software and on line analyze Bo VIN3gene and their coding protein.Result indicated BoVIN3protein560AA,molecular weigh is62.116KD and62.434KD respectively,PI is6.56and5.63respectively; Phosphorylation site analysis of BoVIN3,both have many Phosphorylation site; analysis of protein conserved domains of BoVIN3showed that they have Zn-finger domains.
The expression patterns of the BoVIN3genes were investigated by semi-quantity RT-PCR, results showed that BoVIN3gene could express induced by low temperature treatment, and expressed in the stem apex specifically, transcript levels increased with increasing treatment time.The transcription reach the peak at42d of vernalization. At the same time the expression of FLC was suppressed after BoVIN3was induced by low temperature treatment,results suggest that both genes maybe involved in the biological process of flowering regulation in vernalization. However both BoVIN3could not be induced by GA3and KT treatment.It indicated that cabbage BoVIN3was specific vernalization genes, whose transcripts could specially response to vernalization.
Employ pB1121plasmid and BoVIN3-l gene segment to construct anti-sense expression vector. Insert the BoVIN3-1anti-sense gene segment at Xbal and Smal restriction enzymes site between CaMV355promoter and GUS gene,construct plasmid pBI35S-BoVIN3-1.Transform it to cabbage flower bud by means of micro-injection,observed that after vernalization treatment the transgenic plant vernalization response be delayed at certain extent, the result further indicated that BoVIN3-1gene maybe part in vernalization directly.
根据拟南芥及其芸薹属相关领域的研究报道,本试验重点针对开花网络核心基因FLC及其春化作用途径上游的关键基因VIN3进行研究,首次从十字花科芸薹属作物结球甘蓝中克隆了春化相关基因VIN3,通过生物软件对VIN3基因进行了序列分析,并通过RT-PCR方法检测在结球甘蓝中的表达情况及其与春化作用的关系,以为研究结球甘蓝绿体春化的分子机理提供理论依据。本试验主要研究结果如下：
本试验中对自然条件下生长的甘蓝幼苗可以感受低温春化的形态标准研究表明：甘蓝中熟品种“光荣一号”可以感受低温春化的低限苗龄为13片叶,茎粗达到12.83±0.14mm,低温处理48d春化作用结束,开始进入花芽分化临界期。甘蓝植株只有接受低温诱导达到春化饱和程度才能启动相应基因表达,所以本试验需采用可以感受低温春化的幼苗进行春化处理,以备下一步克隆春化相关基因VIN3。
根据拟南芥VIN3基因的mRNA序列(NM-125121.3)的ORF编码区,设计了VIN3基因的特异引物,以甘蓝RNA及DNA为模板克隆BoVIN3基因的cDNA和DNA序列。克隆得到两条甘蓝春化相关基因BoVIN3的cDNA ORF编码区序列,各1680bp,GenBank登录号分别为JQ394927和JQ394928,可编码由560个氨基酸组成的蛋白质；与拟南芥VIN3基因在核酸水平上同源性分别达到85.55%和79.95%；与拟南芥氨基酸序列同源性分别为80.72%和72.96%。同时获得BoVIN3基因的部分基因组片段,1405bp,GenBank登录号为JQ394929。
利用DNAMAN等软件以及在线分析程序对BoVIN3基因及其编码的蛋白进行分析和预测。结果表明,甘蓝BoVIN3-1、BoVIN3-2蛋白长度均为560AA,分子量约分别为62.116KD和62.434KD,等电点分别为6.56和5.63；通过磷酸化位点预测其蛋白存在多个磷酸化位点；通过NCBI保守结构域预测程序得知该蛋白具有PHD结构域。
半定量RT-PCR表达分析表明BoVIN3受春化作用的诱导,只在感应低温的茎尖生长点表达,在其它部位不表达,并且其转录产物随春化时间延长逐渐积累,在春化42d时达到峰值;而在不同浓度GA3和KT诱导下不表达;FLC受春化作用的抑制,并且BoVIN3受诱导表达后才能使FLC的表达受到抑制,暗示在结球甘蓝中BoVIN3基因可能参与春化调控开花的生物学过程,属于春化特异基因,只响应低温诱导而转录。
以pBI121植物表达载体和BoVIN3-1基因片段为基础,构建反义植物表达载体。将BoVIN3-1基因片段反向插入355启动子与GUS基因之间的限制性酶切位点XbaI和SmaI中,构建含反义BoVIN3-1基因的工程质粒pB135S-BoVIN3-1。利用转反义基因技术,即转化BoVIN3-1的反义片段进行了BoVIN3-1的功能分析,通过花蕾微量注射法转化甘蓝,观察到经春化处理的转反义基因植株与对照相比,春化一定程度被推迟,这一结果进一步表明,甘蓝BoVIN3-1基因可能直接参与了春化过程。
Cabbage (Brassica oleracea var. captata L.)belongs to green vernalizatio plants,young plants need to grow to certain sizes then can accept low temperature and go through vernalization.The pr-ocess of long colding time promoting plant flowering known as vernalization, playing an important role in transition to flowering, in recent years had made considerable progress in vernalization res-earch,especially the molecular mechanism discussion highlights its natural gradually.Although the understanding of the vernalization mechanism show some outline,but the molecular control process is not clear. According to related reports in the arabidopsis and brassica,this work focusing on core gene FLC in flowering regulation way and its upper key gene VIN3in vernalization to research,first isolate vernalization related gene VIN3from cabbage by Homology cloning, employ the biological software to analyze BoVIN3gene and deduced amino acid sequence,the expression patterns of the BoVIN3genes were investigated by semi-quantity RT-PCR, in an attempt to explore the relation be-tween its expression and vernalization, provide base theory for cabbage vernalization.The main conclusions of this study are as follows:
In this experiment, the research of morphological characters of the cabbage plants can accept low temperature growth under natural condition showed that medium maturing varieties of "GUANG RONG YI HAO", with13leaves and12.83±0.14mm in stem diameter, begin to be sensitive to vernalization,low temperature treatment48d vernalization complete, transition to flower bud differentiation. Cabbage plants only accept cold inducement reached saturation level to activate the corresponding gene expression,so this work must employ cabbage plants can feel low temperature to vernalize, prepared to following cloning the VIN3gene.
According to AtVIN3gene mRNA sequence (NM-125121.3) ORF coding region,design gene specific primer,take cabbage RNA and DNA as template to amplified BoVIN3gene.Two vernalization-related genes were isolated by RT-PCR. Sequencing analysis results showed that their ORF were both1680bp in length, which encoded560amino acids, and GenBank accession number are JQ394927and JQ394928respectively.Their nucleotide sequences shared85.55%and79.95%identity with AtVIN3respectively, and the deduced amino acid sequence showed homology to AtVIN3with80.72%and72.96%. At the same time we isolated BoVIN3part DNA segment,was1405bp, GenBank accession number is JQ394929.
Take advantage of DNAMAN software and on line analyze Bo VIN3gene and their coding protein.Result indicated BoVIN3protein560AA,molecular weigh is62.116KD and62.434KD respectively,PI is6.56and5.63respectively; Phosphorylation site analysis of BoVIN3,both have many Phosphorylation site; analysis of protein conserved domains of BoVIN3showed that they have Zn-finger domains.
The expression patterns of the BoVIN3genes were investigated by semi-quantity RT-PCR, results showed that BoVIN3gene could express induced by low temperature treatment, and expressed in the stem apex specifically, transcript levels increased with increasing treatment time.The transcription reach the peak at42d of vernalization. At the same time the expression of FLC was suppressed after BoVIN3was induced by low temperature treatment,results suggest that both genes maybe involved in the biological process of flowering regulation in vernalization. However both BoVIN3could not be induced by GA3and KT treatment.It indicated that cabbage BoVIN3was specific vernalization genes, whose transcripts could specially response to vernalization.
Employ pB1121plasmid and BoVIN3-l gene segment to construct anti-sense expression vector. Insert the BoVIN3-1anti-sense gene segment at Xbal and Smal restriction enzymes site between CaMV355promoter and GUS gene,construct plasmid pBI35S-BoVIN3-1.Transform it to cabbage flower bud by means of micro-injection,observed that after vernalization treatment the transgenic plant vernalization response be delayed at certain extent, the result further indicated that BoVIN3-1gene maybe part in vernalization directly.
引文
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