大豆MYB转录因子基因的克隆及其表达研究
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摘要
通过转录因子在转录水平上调控目的基因的表达，是植物对其生长发育及生理代谢调控的一种重要方式。MYB转录因子基因是最大的植物转录因子基因家族之一，参与植物次生代谢调控、激素和环境因子的应答，并对细胞分化、细胞周期以及植物叶片等器官的形态建成具有重要的调节作用。大豆是重要的油料作物，同时也是蛋白和异黄酮的重要来源，而大多数MYB转录因子对植物类黄酮的代谢均有调控作用。因此，本文以大豆为研究材料，从中分离克隆新的MYB转录因子基因，以期为MYB转录因子在类黄酮的生物合成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．根据植物中MYB基因DNA结合域保守区设计简并引物，以大豆品种中豆27、吉林3号、淮豆1号及张家口黑豆的叶片为材料，RT-PCR扩增出14个MYB基因同源片段；据此设计基因特异引物，通过RACE-PCR分离克隆出4个MYB转录因子基因，GmMYBZ1(DQ902862)、GmMYBZ2(DQ902861)、GmMYBJ6(DQ902863)和GmMYBJ7(DQ902864)；氨基酸序列比对表明，这4个基因分别具有两个MYB结构域，为典型的R2R3MYB转录因子基因。
2．半定量RT-PCR对这4个基因的组织特异性表达情况进行了检测，结果只在叶片中检测到了GmMYBJ6的表达，在茎和叶中分别检测到了GmMYBZ1和GmMYBJ7的表达，而GmMYBZ2在植物的根、茎、叶及未成熟种子中均有表达。对胁迫应答的检测结果显示，在紫外辐射、高盐及干旱(PEG)条件下，随着处理时间的增长，GmMYBJ6的表达量增加，而GmMYBZ2和GmMYBJ7的表达量降低。
3．以基因组DNA为模板对GmMYBZ1、GmMYBZ2、GmMYBJ6和GmMYBJ7进行了PCR扩增，扩增产物经序列分析表明，GmMYBZ1不含内含子，GmMYBZ2含有1个内含子，而GmMYBJ6和GmMYBJ7分别含有2个内含子。
4．构建四个酵母效应质粒pBridge-GmMYBZ1、pBridge-GmMYBZ2、pBriage-GmMYBJ6和pBridge-GmMYBJ7，转入酵母菌株AH109中，并分别在缺少色氨酸的SD培养基和缺少色氨酸、组氨酸的SD双缺陷培养基上筛选阳性转化子。酵母表达表明，GmMYBZ2、GmMYBJ6和GmMYBJ7均具有明显的转录激活功能，β-半乳糖苷酶活性分别为10.35U、28.48U和24.31U，Whatman滤纸显色反应均能呈现出明显的蓝色；而GmMYBZ1酵母转化株的β-半乳糖苷酶活性仅为3.59U，Whatman滤纸显色反应未能呈现出明显的蓝色。
5．构建绿色荧光蛋白融合表达载体，基因枪法转化洋葱表皮细胞进行瞬时表达，结果表明，GmMYBZ2、GmMYBJ6和GmMYBJ7蛋白均定位于细胞核中，与亚细胞定位预测一致。
6．构建植物表达载体pCAMBIA2301-GmMYBZ2、pCAMBIA2301-GmMYBJ6和pCAMBIA2301-GmMYBJ7，利用农杆菌介导法转化烟草NC89；通过Kan抗性、Gus染色及RT-PCR筛选鉴定阳性苗；RT-PCR检测结果显示，GmMYBJ6可提高类黄酮代谢途径中的苯丙氨酸氨基裂解酶(PAL)、肉桂酸-4-羟化酶(C4H)、4-香豆酰辅酶A连接酶(4CL)、查尔酮合成酶(CHS)、黄酮醇合成酶(FLS)等关键酶的表达，结果在GmMYBJ6表达的烟草中，总黄酮含量增加；而由于GmMYBZ2和GmMYBJ7抑制了多数类黄酮代谢途径中基因的表达，从而导致转化烟草中总黄酮含量减少。耐胁迫检测结果表明，GmMYBJ6在烟草中的表达，增强了转基因烟草对UV-B、高盐及干旱等非生物胁迫的耐受性。
Transcription factors play an important role in regulation of plant growth andphysiological metabolism by regulation of gene expression at the transcription level.MYB transcription factor gene is one of the largest families in plants, which involved inregulation of secondary metabolism, responding to hormones and environmental factors,also regulation of cell differentiation, cell cycles and the formation of leaves and otherorgans. Soybean is not only an important oil crop and a source of proteins, but also themost important source of isoflavonoids. Many MYB transcription factors were reported tohave ability to regulate the metabolism of flavonoids. In this paper, some novel genesencoding MYB transcription factors were isolated and characterized from soybean cultivars.The main results of this study are as follows:
1. A pair of degenerate primers was designed according to the conserved regions whichencoding the MYB DNA binding domains in plant MYB genes. Fourteen fragments wereamplified from leaves of soybean cultivars ZhongDou-27, JiLin-3, HuaiDou-1 andZhangJiaKou Black Soybean using RT-PCR. Four novel genes encoding MYBtranscription factors, GmMYBZ1, GmMYBZ2, GmMYBJ6 and GmMYBJ7 were isolated byRACE-PCR. Comparison of deduced amino acid sequences showed that they are typicalR2R3MYB transcription factors because of including two MYB domains respectively.
2. The expression pattern of the four members in different organs was studied usingsemi-quantitative RT-PCR. The results showed that GmMYBZ2 was expressed in roots,stems, leaves and immature seeds of soybean, the expression of GmMYBZ1 and GmMYBJ7was detected in the stems and leaves, whereas the expression of GmMYBJ6 was detectedonly in the leaves. Expression of GmMYBJ6 could be increased under UV-B radiation,drought and high-salt treatment, while the expression of GmMYBJ6 and GmMYBJ6 wouldbe decreased under the same treatments.
3. The full-length DNA sequences of GmMYBZ1、GmMYBZ2、GmMYBJ6 andGmMYBJ7 were also amplified using genomic DNA as templates. The results indicatedthat GmMYBZ2 contains one intron, GmMYBJ6 and GmMYBJ7 contain two intronsrespectively, whereas the gene GmMYBZ1 lacks intron.
4. The recombinant vectors pBridge-GmMYBZ1, pBridge-GmMYBZ2, pBridge-GmMYBJ6 and pBridge-GmMYBJ7 were constructed and transformed into the yeast stainsAH109, then the positive yeast transformants were selected using the SD/-Trp selectivemedium and the SD/-Trp/-His selective medium. The transcriptional activation ofGmMYBZ1, GmMYBZ2, GmMYBJ6 and GmMYBJ7 proteins was confirmed by theyeast system, and itsβ-galactosidase activity was detected as 3.59U, 10.39U, 28.48U and24.31U, respectively. The results of the colony-lift filters assay indicated that the yeasttransformants of GmMYBZ2, GmMYBJ6 and GmMYBJ7 showed a clear blue, whereas theyeast transformants of GmMYBZ1 didn't.
5. The green fluorescent protein expression vectors p163-GFP-GmMYBZ2、p163-GFP-GmMYBJ6 and p163-GFP-GmMYBJ7 were constructed and transformed into the epidermalcells of onion via particle bombardrnental method. The results of instantaneous expressionshowed that GmMYBZ2、GmMYBJ6 and GmMYBJ7 proteins all were localized in cellnucleus, which was consistent with the prediction of the subcellular localization.
6. GrnMYBZ2、GmMYBJ6 and GmMYBJ7 were transformed into tobacco NC89 withAgrobacterium LBA4404 containing the plant expression vectors pCAMBIA2301-GmMYBZ2, pCAMBIA2301-GmMYBJ6 and pCAMBIA2301-GmMYBJ7, respectively.The positive tobacco transformants were selected using MS medium containing kanamycin,Gus activity assay and RT-PCR. Semi-quantitative RT-PCR analysis indicated thatGmMYBJ6 could improve the expression of some flavonoid biosynthetic genes, such asPAL (Phenylalanine ammonia lyase), C4H (cinnamate-4-hydroxylase), 4CL (4-coumaroyl-CoA ligase), CHS (Chalcone Synthase), CHI (chalcone isomerase), F3H (flavanone3-hydroxylase), and FLS (flavonol synthase), resulting the increase of the total flavonoidlevels. Whereas, over exprssion of GmMYBZ2 and GmMYBJ7 could decrease the totalflavonoid level in the transgenic tobacco plants due to the repression of the genes inflavonoid biosynthesis. Results of anti-stress experiments showed that the expression ofGmMYBJ6 could improve resistance to UV-B radiation and drought of transgenic tobaccoobviously.
1．根据植物中MYB基因DNA结合域保守区设计简并引物，以大豆品种中豆27、吉林3号、淮豆1号及张家口黑豆的叶片为材料，RT-PCR扩增出14个MYB基因同源片段；据此设计基因特异引物，通过RACE-PCR分离克隆出4个MYB转录因子基因，GmMYBZ1(DQ902862)、GmMYBZ2(DQ902861)、GmMYBJ6(DQ902863)和GmMYBJ7(DQ902864)；氨基酸序列比对表明，这4个基因分别具有两个MYB结构域，为典型的R2R3MYB转录因子基因。
2．半定量RT-PCR对这4个基因的组织特异性表达情况进行了检测，结果只在叶片中检测到了GmMYBJ6的表达，在茎和叶中分别检测到了GmMYBZ1和GmMYBJ7的表达，而GmMYBZ2在植物的根、茎、叶及未成熟种子中均有表达。对胁迫应答的检测结果显示，在紫外辐射、高盐及干旱(PEG)条件下，随着处理时间的增长，GmMYBJ6的表达量增加，而GmMYBZ2和GmMYBJ7的表达量降低。
3．以基因组DNA为模板对GmMYBZ1、GmMYBZ2、GmMYBJ6和GmMYBJ7进行了PCR扩增，扩增产物经序列分析表明，GmMYBZ1不含内含子，GmMYBZ2含有1个内含子，而GmMYBJ6和GmMYBJ7分别含有2个内含子。
4．构建四个酵母效应质粒pBridge-GmMYBZ1、pBridge-GmMYBZ2、pBriage-GmMYBJ6和pBridge-GmMYBJ7，转入酵母菌株AH109中，并分别在缺少色氨酸的SD培养基和缺少色氨酸、组氨酸的SD双缺陷培养基上筛选阳性转化子。酵母表达表明，GmMYBZ2、GmMYBJ6和GmMYBJ7均具有明显的转录激活功能，β-半乳糖苷酶活性分别为10.35U、28.48U和24.31U，Whatman滤纸显色反应均能呈现出明显的蓝色；而GmMYBZ1酵母转化株的β-半乳糖苷酶活性仅为3.59U，Whatman滤纸显色反应未能呈现出明显的蓝色。
5．构建绿色荧光蛋白融合表达载体，基因枪法转化洋葱表皮细胞进行瞬时表达，结果表明，GmMYBZ2、GmMYBJ6和GmMYBJ7蛋白均定位于细胞核中，与亚细胞定位预测一致。
6．构建植物表达载体pCAMBIA2301-GmMYBZ2、pCAMBIA2301-GmMYBJ6和pCAMBIA2301-GmMYBJ7，利用农杆菌介导法转化烟草NC89；通过Kan抗性、Gus染色及RT-PCR筛选鉴定阳性苗；RT-PCR检测结果显示，GmMYBJ6可提高类黄酮代谢途径中的苯丙氨酸氨基裂解酶(PAL)、肉桂酸-4-羟化酶(C4H)、4-香豆酰辅酶A连接酶(4CL)、查尔酮合成酶(CHS)、黄酮醇合成酶(FLS)等关键酶的表达，结果在GmMYBJ6表达的烟草中，总黄酮含量增加；而由于GmMYBZ2和GmMYBJ7抑制了多数类黄酮代谢途径中基因的表达，从而导致转化烟草中总黄酮含量减少。耐胁迫检测结果表明，GmMYBJ6在烟草中的表达，增强了转基因烟草对UV-B、高盐及干旱等非生物胁迫的耐受性。
Transcription factors play an important role in regulation of plant growth andphysiological metabolism by regulation of gene expression at the transcription level.MYB transcription factor gene is one of the largest families in plants, which involved inregulation of secondary metabolism, responding to hormones and environmental factors,also regulation of cell differentiation, cell cycles and the formation of leaves and otherorgans. Soybean is not only an important oil crop and a source of proteins, but also themost important source of isoflavonoids. Many MYB transcription factors were reported tohave ability to regulate the metabolism of flavonoids. In this paper, some novel genesencoding MYB transcription factors were isolated and characterized from soybean cultivars.The main results of this study are as follows:
1. A pair of degenerate primers was designed according to the conserved regions whichencoding the MYB DNA binding domains in plant MYB genes. Fourteen fragments wereamplified from leaves of soybean cultivars ZhongDou-27, JiLin-3, HuaiDou-1 andZhangJiaKou Black Soybean using RT-PCR. Four novel genes encoding MYBtranscription factors, GmMYBZ1, GmMYBZ2, GmMYBJ6 and GmMYBJ7 were isolated byRACE-PCR. Comparison of deduced amino acid sequences showed that they are typicalR2R3MYB transcription factors because of including two MYB domains respectively.
2. The expression pattern of the four members in different organs was studied usingsemi-quantitative RT-PCR. The results showed that GmMYBZ2 was expressed in roots,stems, leaves and immature seeds of soybean, the expression of GmMYBZ1 and GmMYBJ7was detected in the stems and leaves, whereas the expression of GmMYBJ6 was detectedonly in the leaves. Expression of GmMYBJ6 could be increased under UV-B radiation,drought and high-salt treatment, while the expression of GmMYBJ6 and GmMYBJ6 wouldbe decreased under the same treatments.
3. The full-length DNA sequences of GmMYBZ1、GmMYBZ2、GmMYBJ6 andGmMYBJ7 were also amplified using genomic DNA as templates. The results indicatedthat GmMYBZ2 contains one intron, GmMYBJ6 and GmMYBJ7 contain two intronsrespectively, whereas the gene GmMYBZ1 lacks intron.
4. The recombinant vectors pBridge-GmMYBZ1, pBridge-GmMYBZ2, pBridge-GmMYBJ6 and pBridge-GmMYBJ7 were constructed and transformed into the yeast stainsAH109, then the positive yeast transformants were selected using the SD/-Trp selectivemedium and the SD/-Trp/-His selective medium. The transcriptional activation ofGmMYBZ1, GmMYBZ2, GmMYBJ6 and GmMYBJ7 proteins was confirmed by theyeast system, and itsβ-galactosidase activity was detected as 3.59U, 10.39U, 28.48U and24.31U, respectively. The results of the colony-lift filters assay indicated that the yeasttransformants of GmMYBZ2, GmMYBJ6 and GmMYBJ7 showed a clear blue, whereas theyeast transformants of GmMYBZ1 didn't.
5. The green fluorescent protein expression vectors p163-GFP-GmMYBZ2、p163-GFP-GmMYBJ6 and p163-GFP-GmMYBJ7 were constructed and transformed into the epidermalcells of onion via particle bombardrnental method. The results of instantaneous expressionshowed that GmMYBZ2、GmMYBJ6 and GmMYBJ7 proteins all were localized in cellnucleus, which was consistent with the prediction of the subcellular localization.
6. GrnMYBZ2、GmMYBJ6 and GmMYBJ7 were transformed into tobacco NC89 withAgrobacterium LBA4404 containing the plant expression vectors pCAMBIA2301-GmMYBZ2, pCAMBIA2301-GmMYBJ6 and pCAMBIA2301-GmMYBJ7, respectively.The positive tobacco transformants were selected using MS medium containing kanamycin,Gus activity assay and RT-PCR. Semi-quantitative RT-PCR analysis indicated thatGmMYBJ6 could improve the expression of some flavonoid biosynthetic genes, such asPAL (Phenylalanine ammonia lyase), C4H (cinnamate-4-hydroxylase), 4CL (4-coumaroyl-CoA ligase), CHS (Chalcone Synthase), CHI (chalcone isomerase), F3H (flavanone3-hydroxylase), and FLS (flavonol synthase), resulting the increase of the total flavonoidlevels. Whereas, over exprssion of GmMYBZ2 and GmMYBJ7 could decrease the totalflavonoid level in the transgenic tobacco plants due to the repression of the genes inflavonoid biosynthesis. Results of anti-stress experiments showed that the expression ofGmMYBJ6 could improve resistance to UV-B radiation and drought of transgenic tobaccoobviously.
引文
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