菊花CmMYB1和CmMYB2转录因子基因的克隆及功能鉴定
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摘要
植物通过转录因子在转录水平上调控目的基因的表达来调控其生长发育及生理代谢。MYB转录因子是最大的植物转录因子家族之一,参与植物次生代谢调控、激素和环境胁迫的应答,并对细胞分化和植物叶片等器官的形态建成起重要的调控作用。菊花原产我国,是我国十大传统名花和世界四大切花之一,具有很高的观赏和应用价值,在花卉生产中占有十分重要的地位。但菊花新基因的发掘和分子育种方始起步,本研究以菊花为研究材料,从中分离克隆出新的MYB转录因子,以期为MYB转录因子在植物次生代谢调控及其胁迫应答等方面的研究提供一定的理论基础。主要研究结果如下：
1.根据菊花EST文库筛选出2个MYB基因同源片段,据此设计基因特异引物,以自主选育的盆栽多头菊品种‘钟山紫桂’叶片为材料,通过3’、5’RACE-PCR分别得到全长为1,237bp和1,331bp的菊花MYB基因cDNA序列CmMYB1(JF795917)和CmMYB2(JF795918);其中CmMYB1具有一个846bp的ORF,编码281个氨基酸及95bp的5’非编码区和296bp包含了多聚A尾的3’非编码区；CmMYB2具有一个948bp的ORF,编码315个氨基酸及86bp的5’非编码区和297bp包含了多聚A尾的3’非编码区。氨基酸序列比对表明,这2个基因均具有两个MYB结构域,为典型的R2R3-MYB转录因子基因,其中CmMYB1基因属于第4亚群成员,而CmMYB2基因属于第22亚群成员。以基因组DNA为模板对CmMYB1和CmMYB2进行PCR扩增,扩增产物经序列分析表明,CmMYB1含有1个307bp的内含子,而CmMYB2不含内含子。
2.克隆了CmMYB1、CmMYB2基因的启动子序列并发现了一些与ABA及胁迫相关的作用元件。
3.构建2个酵母效应质粒pGBKT7-CmMYBl和pGBKT7-CmMYB2,转入酵母菌株Y2H中,并分别在缺陷培养基SD/-Trp上筛选后转入SD/-His-Ade上培养,结果表明,CmMYB1和CmMYB2均无明显的转录激活功能。
4.荧光定量PCR结果表明,CmMYB1和CmMYB2基因在菊花的根、茎、叶和花的5个发育阶段都有表达,且在茎中的表达量最高,CmMYBl在叶片和花的完全着色与初开期时表达量较高,在根中表达较低；CmMYB2基因在叶中表达量也较高,但在根和开花的各个时期表达量均较低。胁迫应答的检测结果显示,在高盐、干旱(PEG)、低温和ABA诱导条件下,CmMYB2的表达量在处理后1h均有所增加,而CmMYB1的表达量除了在低温胁迫下略有上升外,其他胁迫条件下均较低。
5.构建绿色荧光蛋白融合表达载体,农杆菌介导法转化洋葱表皮细胞进行瞬时表达,结果表明,CmMYB1、CmMYB2蛋白位于细胞核中。
6.构建植物表达载体pCAMBIA1301-CmMYB1和pCAMBIA1301-CmMYB2,利用农杆菌介导法转化拟南芥(Co1.0),通过潮霉素抗性、GUS染色及RT-PCR筛选鉴定阳性苗。转CmMYB1植株和野生型植株在表型上没有区别；qRT-PCR检测结果显示,CmMYB1降低木质素合成途径以及由总苯丙氨酸途径通往木质素合成途径的关键酶基因AtCAD (cinnamyl alcohol dehydrogenase,肉桂醇脱氢酶)、AtCOMT (caffeic acid o-methyl-transferase,咖啡酸-0-甲基转移酶)、AtC4H (cinnamate4-hydroxylase,肉桂酸-4-羟基酶)和At4CLl (4-coumarate-CoA ligase,4-香豆酸：辅酶A连接酶)的表达,结果在CmMYBl表达的拟南芥中,木质素含量降低。转CmMYB2植株较野生型开花延迟,叶片数目增多,耐胁迫检测结果表明,CmMYB2在拟南芥中的过表达,增强了转基因拟南芥对干旱和盐胁迫的耐受性；qRT-PCR检测结果显示,过表达CmMYB2拟南芥中RD22、RD29A、RAB18、COR47、ABA1和ABA2等胁迫相关基因的表达量增强；此外,CO(CONSTANS)、FT (FLOWERING LOCUS T)、SOC1(SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1)、LFY(LEAFY)、API (APETALA1)等开花相关基因的表达量降低,与转CmMYB2植株晚花表型相吻合。
The growth and metabolism of plants are regulated by controlling the expression of targeted gene on the transcript level. MYB transcription factor gene is one of the largest families in plants, which involved in regulation of secondary metabolism, responding to hormones and environmental stresses, also regulation of cell differentiation and organs morphogenesis. Chrysanthemum morifolium is one of ten most famous flowers in China and one of the top four cut flowers in the world with high ornamental and practical value. It occupies an important position in flower production. However, the discovery of new genes and the methods for molecular breeding of Chrysanthemum just started. In this study, two genes of MYB transcription factors of Chrysanthemum were cloned and characterized. These are expected to provide some basic theories for further research of MYB transcription factors on regulation of plant secondary metabolism and responding to abiotic stresses. The main results are as follows:
1. Two homologous MYB genes were screened from the EST libraries of Chrysanthemum and amplified by3'and5'RACE to obtain the full-length cDNA of CmMYB1(JF795917) and CmMYB2(JF795918). CmMYB1contains an846bp ORF, encoding a281residue peptide, with a95bp5'untranslated region and a296bp3' untranslated region with a poly(A) tail. CmMYB2contains an948bp ORF, encoding a315residue peptide, with a86bp5'untranslated region and a297bp3'untranslated region with a poly(A) tail. Comparison of deduced amino acid sequences showed that the two genes are typical R2R3-MYB transcription factors and belong to the subgroup4and22, respectively. Then the full-length DNA sequences of CmMYB1and CmMYB2were also amplified using genomic DNA as templates. The results indicated that CmMYBl contains a307bp intron, whereas the gene CmMYB2lacks intron.
2. By TAIL-PCR and5'RACE, full-length promoter sequences of CmMYB1and CmMYB2were obtained and submitted into promoter database for element forecast analysis. The results showed that there were some elements for response to ABA, drought and low temperature.
3. Recombinant vectors pGBKT7-CmMYB1and pGBKT7-CmMYB2were constructed and transformed into the yeast strains Y2H. The positive yeast transformants were selected using troptophan-deficient medium SD/-Trp, then transformed on SD/-His-Ade medium. The results showed that CmMYB1and CmMYB2have no significantly transcriptional activity.
4. Quantitative real-time PCR (qRT-PCR) showed CmMYB1and CmMYB2were expressed in roots, stems, leaves and flowers of plants with the highest expression in stems. CmMYB1has higher expression level at the early opening and full color of the flowers, but little level in roots. The transcript product of CmMYB2was higher in leaves, but lower in roots and at any time of the flowering. The results from the responses to abiotic stress showed that the expression of CmMYB2increased rapidly in response to drought, salinity or cold stress within1h, while the expression of CmMYB1was induced only under the low temperature condition.
5. The green fluorescent protein expression vectors pBI121-GFP-CmMYB1and pBI121-GFP-CmMYB2were constructed and transformed into the epidermal cells of onion by Agrobacterium-mediated transformation system. The results of instantaneous expression showed that CmMYB1and CmMYB2proteins were both localized in cell nucleus.
6.CmMYB1and CmMYB2were transformed into Arabidopsis thaliana (Col-0) with Agrobacterium EHA105containing the expression vectors pCAMBIA1301-CmMTB1and pCAMBIA1301-CmMYB2, respectively. The positive Arabidopsis transformants were selected using MS medium containing hygromycin resistance, GUS activity assay and RT-PCR. There was no phenotypic difference between the CmMYB1transgenic Arabidopsis and the wild-type plants. The qRT-PCR analysis indicated that CmMYB1could down-regulate some important genes involved in phenylpropanoid pathway and lignin biosynthetic pathway, such as CAD (cinnamyl alcohol dehydrogenase)、COMT (caffeic acid o-methyl-transferase)、C4H (cinnamate4-hydroxylase) and4CL1(4-coumarate-CoA ligase), resulting the decrease of lignin content. The transgenic Arabidopsis overexpressing CmMYB2exhibited delayed flowering and the increasing of leaves number compared with the wild-type plants. Results of abiotic-stress experiments showed that heterologous expression of CmMYB2could enhance drought and salt tolerance of Arabidopsis. In line with the above findings, the expression levels of several stress-responsive genes RD22^RD29A、RAB18、COR47、ABA1and ABA2were strongly increased, while the expression of genes related to flowering, such as CO (CONSTANS)、FT (FLOWERING LOCUS T)、 SOC1(SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1)、LFY(LEAFY) and AP1(APETALA1) reduced.
1.根据菊花EST文库筛选出2个MYB基因同源片段,据此设计基因特异引物,以自主选育的盆栽多头菊品种‘钟山紫桂’叶片为材料,通过3’、5’RACE-PCR分别得到全长为1,237bp和1,331bp的菊花MYB基因cDNA序列CmMYB1(JF795917)和CmMYB2(JF795918);其中CmMYB1具有一个846bp的ORF,编码281个氨基酸及95bp的5’非编码区和296bp包含了多聚A尾的3’非编码区；CmMYB2具有一个948bp的ORF,编码315个氨基酸及86bp的5’非编码区和297bp包含了多聚A尾的3’非编码区。氨基酸序列比对表明,这2个基因均具有两个MYB结构域,为典型的R2R3-MYB转录因子基因,其中CmMYB1基因属于第4亚群成员,而CmMYB2基因属于第22亚群成员。以基因组DNA为模板对CmMYB1和CmMYB2进行PCR扩增,扩增产物经序列分析表明,CmMYB1含有1个307bp的内含子,而CmMYB2不含内含子。
2.克隆了CmMYB1、CmMYB2基因的启动子序列并发现了一些与ABA及胁迫相关的作用元件。
3.构建2个酵母效应质粒pGBKT7-CmMYBl和pGBKT7-CmMYB2,转入酵母菌株Y2H中,并分别在缺陷培养基SD/-Trp上筛选后转入SD/-His-Ade上培养,结果表明,CmMYB1和CmMYB2均无明显的转录激活功能。
4.荧光定量PCR结果表明,CmMYB1和CmMYB2基因在菊花的根、茎、叶和花的5个发育阶段都有表达,且在茎中的表达量最高,CmMYBl在叶片和花的完全着色与初开期时表达量较高,在根中表达较低；CmMYB2基因在叶中表达量也较高,但在根和开花的各个时期表达量均较低。胁迫应答的检测结果显示,在高盐、干旱(PEG)、低温和ABA诱导条件下,CmMYB2的表达量在处理后1h均有所增加,而CmMYB1的表达量除了在低温胁迫下略有上升外,其他胁迫条件下均较低。
5.构建绿色荧光蛋白融合表达载体,农杆菌介导法转化洋葱表皮细胞进行瞬时表达,结果表明,CmMYB1、CmMYB2蛋白位于细胞核中。
6.构建植物表达载体pCAMBIA1301-CmMYB1和pCAMBIA1301-CmMYB2,利用农杆菌介导法转化拟南芥(Co1.0),通过潮霉素抗性、GUS染色及RT-PCR筛选鉴定阳性苗。转CmMYB1植株和野生型植株在表型上没有区别；qRT-PCR检测结果显示,CmMYB1降低木质素合成途径以及由总苯丙氨酸途径通往木质素合成途径的关键酶基因AtCAD (cinnamyl alcohol dehydrogenase,肉桂醇脱氢酶)、AtCOMT (caffeic acid o-methyl-transferase,咖啡酸-0-甲基转移酶)、AtC4H (cinnamate4-hydroxylase,肉桂酸-4-羟基酶)和At4CLl (4-coumarate-CoA ligase,4-香豆酸：辅酶A连接酶)的表达,结果在CmMYBl表达的拟南芥中,木质素含量降低。转CmMYB2植株较野生型开花延迟,叶片数目增多,耐胁迫检测结果表明,CmMYB2在拟南芥中的过表达,增强了转基因拟南芥对干旱和盐胁迫的耐受性；qRT-PCR检测结果显示,过表达CmMYB2拟南芥中RD22、RD29A、RAB18、COR47、ABA1和ABA2等胁迫相关基因的表达量增强；此外,CO(CONSTANS)、FT (FLOWERING LOCUS T)、SOC1(SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1)、LFY(LEAFY)、API (APETALA1)等开花相关基因的表达量降低,与转CmMYB2植株晚花表型相吻合。
The growth and metabolism of plants are regulated by controlling the expression of targeted gene on the transcript level. MYB transcription factor gene is one of the largest families in plants, which involved in regulation of secondary metabolism, responding to hormones and environmental stresses, also regulation of cell differentiation and organs morphogenesis. Chrysanthemum morifolium is one of ten most famous flowers in China and one of the top four cut flowers in the world with high ornamental and practical value. It occupies an important position in flower production. However, the discovery of new genes and the methods for molecular breeding of Chrysanthemum just started. In this study, two genes of MYB transcription factors of Chrysanthemum were cloned and characterized. These are expected to provide some basic theories for further research of MYB transcription factors on regulation of plant secondary metabolism and responding to abiotic stresses. The main results are as follows:
1. Two homologous MYB genes were screened from the EST libraries of Chrysanthemum and amplified by3'and5'RACE to obtain the full-length cDNA of CmMYB1(JF795917) and CmMYB2(JF795918). CmMYB1contains an846bp ORF, encoding a281residue peptide, with a95bp5'untranslated region and a296bp3' untranslated region with a poly(A) tail. CmMYB2contains an948bp ORF, encoding a315residue peptide, with a86bp5'untranslated region and a297bp3'untranslated region with a poly(A) tail. Comparison of deduced amino acid sequences showed that the two genes are typical R2R3-MYB transcription factors and belong to the subgroup4and22, respectively. Then the full-length DNA sequences of CmMYB1and CmMYB2were also amplified using genomic DNA as templates. The results indicated that CmMYBl contains a307bp intron, whereas the gene CmMYB2lacks intron.
2. By TAIL-PCR and5'RACE, full-length promoter sequences of CmMYB1and CmMYB2were obtained and submitted into promoter database for element forecast analysis. The results showed that there were some elements for response to ABA, drought and low temperature.
3. Recombinant vectors pGBKT7-CmMYB1and pGBKT7-CmMYB2were constructed and transformed into the yeast strains Y2H. The positive yeast transformants were selected using troptophan-deficient medium SD/-Trp, then transformed on SD/-His-Ade medium. The results showed that CmMYB1and CmMYB2have no significantly transcriptional activity.
4. Quantitative real-time PCR (qRT-PCR) showed CmMYB1and CmMYB2were expressed in roots, stems, leaves and flowers of plants with the highest expression in stems. CmMYB1has higher expression level at the early opening and full color of the flowers, but little level in roots. The transcript product of CmMYB2was higher in leaves, but lower in roots and at any time of the flowering. The results from the responses to abiotic stress showed that the expression of CmMYB2increased rapidly in response to drought, salinity or cold stress within1h, while the expression of CmMYB1was induced only under the low temperature condition.
5. The green fluorescent protein expression vectors pBI121-GFP-CmMYB1and pBI121-GFP-CmMYB2were constructed and transformed into the epidermal cells of onion by Agrobacterium-mediated transformation system. The results of instantaneous expression showed that CmMYB1and CmMYB2proteins were both localized in cell nucleus.
6.CmMYB1and CmMYB2were transformed into Arabidopsis thaliana (Col-0) with Agrobacterium EHA105containing the expression vectors pCAMBIA1301-CmMTB1and pCAMBIA1301-CmMYB2, respectively. The positive Arabidopsis transformants were selected using MS medium containing hygromycin resistance, GUS activity assay and RT-PCR. There was no phenotypic difference between the CmMYB1transgenic Arabidopsis and the wild-type plants. The qRT-PCR analysis indicated that CmMYB1could down-regulate some important genes involved in phenylpropanoid pathway and lignin biosynthetic pathway, such as CAD (cinnamyl alcohol dehydrogenase)、COMT (caffeic acid o-methyl-transferase)、C4H (cinnamate4-hydroxylase) and4CL1(4-coumarate-CoA ligase), resulting the decrease of lignin content. The transgenic Arabidopsis overexpressing CmMYB2exhibited delayed flowering and the increasing of leaves number compared with the wild-type plants. Results of abiotic-stress experiments showed that heterologous expression of CmMYB2could enhance drought and salt tolerance of Arabidopsis. In line with the above findings, the expression levels of several stress-responsive genes RD22^RD29A、RAB18、COR47、ABA1and ABA2were strongly increased, while the expression of genes related to flowering, such as CO (CONSTANS)、FT (FLOWERING LOCUS T)、 SOC1(SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1)、LFY(LEAFY) and AP1(APETALA1) reduced.
引文
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