马铃薯StCOL和StTFL1的克隆及其在块茎形成中的功能分析
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摘要
马铃薯(Solanum tuberosum L.)是仅次于水稻、小麦、玉米之后的第四大粮食作物。块茎是马铃薯的繁殖器官,也是产品器官。研究马铃薯块茎形成的调控机制,不仅具有理论价值,而且对马铃薯的产量提高具有指导意义。马铃薯块茎形成是一个复杂的生物学过程,并受内在因素及外界因素的调控。最近,有报道将拟南芥的AtCO基因导入马铃薯,在短日照条件下,AtCO过表达可以导致块茎形成减少,说明AtCO对块茎的形成有负调节作用。但是其调控块茎形成机制以及内源CO在块茎形成中作用仍不清楚。为了揭示CO在马铃薯块茎形成中的作用,本文克隆了马铃薯StCOL、StTFL1和StLFY基因,并通过构建StCOL反义抑制和StTFL1过表达转基因植株,初步分析了StCOL和StTFL1对块茎形成的作用。主要研究结果如下：
1.克隆了一个马铃薯StCOL cDNA序列(DQ882684)、基因组DNA序列及其启动子。该基因由一个内含子和两个外显子构成,StCOL cDNA序列含有一个1083 bp的开放阅读框,编码一个360个氨基酸残基构成的蛋白质分子,StCOL分子量为39.21 kD,等电点为5.09。采用DNAMAN对StCOL和其他同源蛋白的氨基酸序列比对,结果表明StCOL氨基酸序列与拟南芥的AtCO. AtCOL、AtCOL2和AtCOL3的氨基酸序列具有较高的一致性,在N-和C-末端分别具有保守的B-box和CCT结构域,在蛋白C-末端有一保守的7肽序列(YGVVPSF)；系统进化分析结果表明StCOL与豌豆PsCOLb及拟南芥AtCOL3分子的亲缘关系最近；StCOL基因启动子序列上游有8个TATA-box、4个CAAT-box、1个I-box、4个GATA-box等多个顺式作用元件。采用半定量RT-PCR方法对StCOL mRNA水平分析结果显示,StCOL在马铃薯的根、茎、叶、顶芽、花芽和花中都有表达,其中在叶片中的表达水平最高；对块茎形成过程中该基因的表达分析结果显示,在块茎形成不同时期都有StCOL表达,在块茎形成后期表达水平较弱,在成熟块茎中检测不到该基因的表达。另外,体外微型薯的暗诱导过程中该基因持续表达,但该基因的表达水平不受蔗糖和赤霉素的影响。
2.采用RT-PCR和RACE的方法克隆了马铃薯StTFL1 cDNA序列(DQ307621)。StTFL1 cDNA序列含有一个528 bp的开放阅读框,编码一个由175个氨基酸残基组成的蛋白,该蛋白分子量为19.86 kD,等电点为8.73,66～88位氨基酸间具有PEBP蛋白典型的特征结构,采用DNAMAN软件对StTFL1和其他同源蛋白的氨基酸序列比对,结果表明StTFL1和拟南芥TFL1、水稻CEN及烟草CEN在氨基酸水平有较高的一致性,而且拟南芥的TFL1和马铃薯的TFL1的二级结构基本一样。采用半定量RT-PCR方法对不同组织中该基因的表达水平检测结果显示,StTFL1基因在马铃薯的根、茎、叶、顶芽和花芽中均表达,其中在根中的表达水平最高；另外,发现在块茎形成的不同时期StTFL1都表达,尤其在匍匐茎形成时有高水平的表达。
3.采用RT-PCR和RACE的方法克隆了马铃薯StLFY cDNA序列(EF062357)及基因组DNA序列(EU371047)。该基因由三个外显子和两个内含子组成,StLFY cDNA含有一个1257 bp的开放阅读框,编码一个由418个氨基酸残基组成的蛋白,该蛋白分子量为46.37 kD,等电点为6.18。采用DNAMAN软件对StLFY和其他同源蛋白的氨基酸序列比对,结果表明StLFY和AtLFY、NtFL2及SILFY在氨基酸水平有较高的一致性,而且在氨基末端都有一个保守的富含脯氨酸的区域。半定量RT-PCR分析结果显示,StLFY仅在马铃薯顶芽和花芽有较弱的表达,而且在匍匐茎初始形成时也有较弱表达。
4.为了进一步了解StCOL和StTFL1与马铃薯块茎形成的关系,首先优化了一简便的农杆菌介导的马铃薯茎段遗传转化方法：茎段外植体暗中预培养2d,农杆菌侵染5～8 min,暗中共培养时间为2d,转入愈伤诱导和芽分化的培养基(MS+0.5 mg/LTDZ+0.3 mg/L GA3+50mg/L Kan+200 mg/L Car+200 mg/L Cef),光照培养4～5周即有不定芽的生成。采用PCR方法验证转基因植株,转化率约为5%。在此基础上,进行了StCO L和StTFL1基因的遗传转化,初步对StCO L和StTFL1在块茎形成中的作用以及它们之间的关系进行了分析。短日照条件下,与未转化对照组相比,表达StCOL反义片段的转基因株系叶片中StCOL mRNA水平下降,植株形成块茎个体总数增加,此结果表明StCO L对块茎的形成有负调节作用；过表达StTFL1转基因株系叶片中StTFL1 mRNA水平和形成的块茎个数均增加,表明StTFL1对块茎的形成有促进作用；分别对StCOL反义抑制和StTFL1过表达转基因植株中的StTFL1和StCOL转录表达分析,结果显示,StTFL1是StCOL的下游基因。
Potato (Solarium tuberosum L.) is ranked 4th in crop production following rice, wheat and maize. Potato tubers are propagation organ and product organ. Researches on the regulation mechanism of tuberization is not only of theoretical value but also of instructive significance for improving production. Potato tuberization is a complex biological process. Tuberization is regulated by many factors including the internal and the environmental factors. Recently, AtCO was introduced into potato, the over-expression of AtCO could reduce tuberization under short-day, this result showed that AtCO could negatively regulate tuber formation. However, roles of AtCO and inner CO in controlling tuber formation remain unclear. In order to reveal the role of CO in the tuber formation, this paper focused on cloning StCOL, StTFL1 and StLFY gene from potato, and preliminarily analyzing the roles of StCOL and StTFLl in tuberization by constructing over-expreession StTFLl and antisene repression StCOL transgenic plants. The main results are as follows:
1. A gene named StCOL(Solanum tuberosum CONSTANS-LIKE, DQ882684) and its promoter was isolated from potato, the full length of the cDNA contains an open reading frame of 1083 bp coding a protein of 360 amino acids, corresponding to a 39.21 kD polypeptide with an isoelectric point of 5.09. Alignment of the amino acid sequence of StCOL and that of other homologues showed that StCOL had a high identity with AtCO, AtCOLl, AtCOL2 and AtCOL3 from Arabidopsis at amino acid level, especially in their N-and C-terminal regions containing two highly conserved B-boxes and CCT domain, respectively. In addition, the conserved 7-amino acid stretch (YGVVPSF) was also observed in C-terminal end of this new protein, it suggested that the new protein belonged to the CO or COL protein family. The deduced amino acid sequence and clustering analysis revealed that StCOL was colse to PsCOLb in pea, and AtCOL3 in A.thiliana. Promoter of StCOL contained some cis-acting elements including eight TATA-boxes, four CAAT-boxes, one I-box, four GATA-boxes and so on. Analysis of StCOL mRNA level by semi-quantity RT-PCR showed that it was expresed in roots, stems, leaves, apical buds, floral buds, flowers, especially highly expressed in leaves, and it was expressed lightly in the whole process of tuber formation, the level of StCOL mRNA became weaker at the late stage of tuberization, and disapeared in mature tubers. In addition, it was expressed continuously in stems during darkness-induce microtuber formaiton in vitro, and was independent on the level of gibberellin A3 and sucrose.
2. A homologue of TFL/CEN, designated StTFL1 (Solanum tuberosum terminal flower 1, DQ307621), was isolated from potato by RT-PCR and RACE. The cDNA sequence of StTFLl was 746 bp long and contained an ORF of 528 bp. The predicted amino acid sequence of the gene was consisted of 175 amino acids, corresponding to a 19.86 kD polypeptide with an isoelectric point of 8.73, and had a conserved region at 66～88 amino residue when being aligned with sequences of other members of PEBP family. Alignment of the amino acid sequence of StTFLl and that of other homologues showed that StTFLl had a high identity with TFL/CEN homologous members from A. thiliana, O. sativa and N. tabacum. The secondary structure models of AtTFLl and StTFL1 were nearly same. Clustering analysis revealed that StTFL1 was close to CaSP. Analysis of StTFLl mRNA by semi-quantity RT-PCR showed that it was expressed in roots, stems, leaves, apical buds and floral buds but not expressed in flowers, especially highly expressed in roots, and that it was expressed continuously during tuber formation, especially the level of its mRNA was more higher in initial stolons than in other tissues.
3. A homologue of FLORICAULAILEAFY, designated StLFY(Solanum tuberosum LEAFY, EF062357), was isolated from potato by RT-PCR and RACE. The cDNA sequence of StLFY was 1417 bp long and contained an ORF of 1257 bp. StLFY gene contained three exons and two introns. The predicted amino acid sequence of the gene was consisted of 418 amino acids, corresponding to a 46.37 kD polypeptide with an isoelectric point of 6.18. Alignment of the amino acid sequence of StLFY and that of other homologues showed that StLFY had a high identity with LFY homologous members from A. thiliana, S. lycopersicum and N. tabacum, and had a conserved region in the N-termiusl when being aligned with sequences of other LFY homologues. Analysis of StLFY mRNA by semi-quantity RT-PCR showed that it was slightly expressed in apical buds, floral buds and initial stolons.
4. In order to further understand the regulation role of StTFL1 and StCOL in potato tuberizatin, first of all, a simple genetic transformation method of Agrobacterium-mediated potato stem cuttings was optimized:stem explants were precultured for two days in darkness, then infected by Agrobacterium for 5～8 min, co-cultured for two days, finally, transferred to the MS medium supplemented with 0.5 mg/L TDZ+0.3 mg/L GA3+50 mg/L Kan+200 mg/L Car+200 mg/L Cef. Adventitious buds could emerge after induction in light for 4～5 weeks. The transformation was tested by PCR, the transformation efficiency reached to 5%. Based on this, the genetic transformation of StCOL and StTFLl was carried. The role and relationship between StCOL and StTFL1 in tuber formation were preliminarily analyzed by being compared with the wild type, transgenic lines of antisense-stCOL had a lower level of StCOL mRNA in leaves but more tubers in average. It suggested that StCOL could negatively regulate tuberization; the transgenic lines of over-expression StTFL1 had a higher level of StTFL1 mRNA in leaves but also more tubers in average. It suggested that StTFL1 could promote tuberization. Analysis of the transcription level of StTFL1 mRNA and StCOL mRNA in transgenic lines of antisense-repression StCOL and over-expression StTFL1 showed that StTFL1 was a downstream gene of StCOL.
1.克隆了一个马铃薯StCOL cDNA序列(DQ882684)、基因组DNA序列及其启动子。该基因由一个内含子和两个外显子构成,StCOL cDNA序列含有一个1083 bp的开放阅读框,编码一个360个氨基酸残基构成的蛋白质分子,StCOL分子量为39.21 kD,等电点为5.09。采用DNAMAN对StCOL和其他同源蛋白的氨基酸序列比对,结果表明StCOL氨基酸序列与拟南芥的AtCO. AtCOL、AtCOL2和AtCOL3的氨基酸序列具有较高的一致性,在N-和C-末端分别具有保守的B-box和CCT结构域,在蛋白C-末端有一保守的7肽序列(YGVVPSF)；系统进化分析结果表明StCOL与豌豆PsCOLb及拟南芥AtCOL3分子的亲缘关系最近；StCOL基因启动子序列上游有8个TATA-box、4个CAAT-box、1个I-box、4个GATA-box等多个顺式作用元件。采用半定量RT-PCR方法对StCOL mRNA水平分析结果显示,StCOL在马铃薯的根、茎、叶、顶芽、花芽和花中都有表达,其中在叶片中的表达水平最高；对块茎形成过程中该基因的表达分析结果显示,在块茎形成不同时期都有StCOL表达,在块茎形成后期表达水平较弱,在成熟块茎中检测不到该基因的表达。另外,体外微型薯的暗诱导过程中该基因持续表达,但该基因的表达水平不受蔗糖和赤霉素的影响。
2.采用RT-PCR和RACE的方法克隆了马铃薯StTFL1 cDNA序列(DQ307621)。StTFL1 cDNA序列含有一个528 bp的开放阅读框,编码一个由175个氨基酸残基组成的蛋白,该蛋白分子量为19.86 kD,等电点为8.73,66～88位氨基酸间具有PEBP蛋白典型的特征结构,采用DNAMAN软件对StTFL1和其他同源蛋白的氨基酸序列比对,结果表明StTFL1和拟南芥TFL1、水稻CEN及烟草CEN在氨基酸水平有较高的一致性,而且拟南芥的TFL1和马铃薯的TFL1的二级结构基本一样。采用半定量RT-PCR方法对不同组织中该基因的表达水平检测结果显示,StTFL1基因在马铃薯的根、茎、叶、顶芽和花芽中均表达,其中在根中的表达水平最高；另外,发现在块茎形成的不同时期StTFL1都表达,尤其在匍匐茎形成时有高水平的表达。
3.采用RT-PCR和RACE的方法克隆了马铃薯StLFY cDNA序列(EF062357)及基因组DNA序列(EU371047)。该基因由三个外显子和两个内含子组成,StLFY cDNA含有一个1257 bp的开放阅读框,编码一个由418个氨基酸残基组成的蛋白,该蛋白分子量为46.37 kD,等电点为6.18。采用DNAMAN软件对StLFY和其他同源蛋白的氨基酸序列比对,结果表明StLFY和AtLFY、NtFL2及SILFY在氨基酸水平有较高的一致性,而且在氨基末端都有一个保守的富含脯氨酸的区域。半定量RT-PCR分析结果显示,StLFY仅在马铃薯顶芽和花芽有较弱的表达,而且在匍匐茎初始形成时也有较弱表达。
4.为了进一步了解StCOL和StTFL1与马铃薯块茎形成的关系,首先优化了一简便的农杆菌介导的马铃薯茎段遗传转化方法：茎段外植体暗中预培养2d,农杆菌侵染5～8 min,暗中共培养时间为2d,转入愈伤诱导和芽分化的培养基(MS+0.5 mg/LTDZ+0.3 mg/L GA3+50mg/L Kan+200 mg/L Car+200 mg/L Cef),光照培养4～5周即有不定芽的生成。采用PCR方法验证转基因植株,转化率约为5%。在此基础上,进行了StCO L和StTFL1基因的遗传转化,初步对StCO L和StTFL1在块茎形成中的作用以及它们之间的关系进行了分析。短日照条件下,与未转化对照组相比,表达StCOL反义片段的转基因株系叶片中StCOL mRNA水平下降,植株形成块茎个体总数增加,此结果表明StCO L对块茎的形成有负调节作用；过表达StTFL1转基因株系叶片中StTFL1 mRNA水平和形成的块茎个数均增加,表明StTFL1对块茎的形成有促进作用；分别对StCOL反义抑制和StTFL1过表达转基因植株中的StTFL1和StCOL转录表达分析,结果显示,StTFL1是StCOL的下游基因。
Potato (Solarium tuberosum L.) is ranked 4th in crop production following rice, wheat and maize. Potato tubers are propagation organ and product organ. Researches on the regulation mechanism of tuberization is not only of theoretical value but also of instructive significance for improving production. Potato tuberization is a complex biological process. Tuberization is regulated by many factors including the internal and the environmental factors. Recently, AtCO was introduced into potato, the over-expression of AtCO could reduce tuberization under short-day, this result showed that AtCO could negatively regulate tuber formation. However, roles of AtCO and inner CO in controlling tuber formation remain unclear. In order to reveal the role of CO in the tuber formation, this paper focused on cloning StCOL, StTFL1 and StLFY gene from potato, and preliminarily analyzing the roles of StCOL and StTFLl in tuberization by constructing over-expreession StTFLl and antisene repression StCOL transgenic plants. The main results are as follows:
1. A gene named StCOL(Solanum tuberosum CONSTANS-LIKE, DQ882684) and its promoter was isolated from potato, the full length of the cDNA contains an open reading frame of 1083 bp coding a protein of 360 amino acids, corresponding to a 39.21 kD polypeptide with an isoelectric point of 5.09. Alignment of the amino acid sequence of StCOL and that of other homologues showed that StCOL had a high identity with AtCO, AtCOLl, AtCOL2 and AtCOL3 from Arabidopsis at amino acid level, especially in their N-and C-terminal regions containing two highly conserved B-boxes and CCT domain, respectively. In addition, the conserved 7-amino acid stretch (YGVVPSF) was also observed in C-terminal end of this new protein, it suggested that the new protein belonged to the CO or COL protein family. The deduced amino acid sequence and clustering analysis revealed that StCOL was colse to PsCOLb in pea, and AtCOL3 in A.thiliana. Promoter of StCOL contained some cis-acting elements including eight TATA-boxes, four CAAT-boxes, one I-box, four GATA-boxes and so on. Analysis of StCOL mRNA level by semi-quantity RT-PCR showed that it was expresed in roots, stems, leaves, apical buds, floral buds, flowers, especially highly expressed in leaves, and it was expressed lightly in the whole process of tuber formation, the level of StCOL mRNA became weaker at the late stage of tuberization, and disapeared in mature tubers. In addition, it was expressed continuously in stems during darkness-induce microtuber formaiton in vitro, and was independent on the level of gibberellin A3 and sucrose.
2. A homologue of TFL/CEN, designated StTFL1 (Solanum tuberosum terminal flower 1, DQ307621), was isolated from potato by RT-PCR and RACE. The cDNA sequence of StTFLl was 746 bp long and contained an ORF of 528 bp. The predicted amino acid sequence of the gene was consisted of 175 amino acids, corresponding to a 19.86 kD polypeptide with an isoelectric point of 8.73, and had a conserved region at 66～88 amino residue when being aligned with sequences of other members of PEBP family. Alignment of the amino acid sequence of StTFLl and that of other homologues showed that StTFLl had a high identity with TFL/CEN homologous members from A. thiliana, O. sativa and N. tabacum. The secondary structure models of AtTFLl and StTFL1 were nearly same. Clustering analysis revealed that StTFL1 was close to CaSP. Analysis of StTFLl mRNA by semi-quantity RT-PCR showed that it was expressed in roots, stems, leaves, apical buds and floral buds but not expressed in flowers, especially highly expressed in roots, and that it was expressed continuously during tuber formation, especially the level of its mRNA was more higher in initial stolons than in other tissues.
3. A homologue of FLORICAULAILEAFY, designated StLFY(Solanum tuberosum LEAFY, EF062357), was isolated from potato by RT-PCR and RACE. The cDNA sequence of StLFY was 1417 bp long and contained an ORF of 1257 bp. StLFY gene contained three exons and two introns. The predicted amino acid sequence of the gene was consisted of 418 amino acids, corresponding to a 46.37 kD polypeptide with an isoelectric point of 6.18. Alignment of the amino acid sequence of StLFY and that of other homologues showed that StLFY had a high identity with LFY homologous members from A. thiliana, S. lycopersicum and N. tabacum, and had a conserved region in the N-termiusl when being aligned with sequences of other LFY homologues. Analysis of StLFY mRNA by semi-quantity RT-PCR showed that it was slightly expressed in apical buds, floral buds and initial stolons.
4. In order to further understand the regulation role of StTFL1 and StCOL in potato tuberizatin, first of all, a simple genetic transformation method of Agrobacterium-mediated potato stem cuttings was optimized:stem explants were precultured for two days in darkness, then infected by Agrobacterium for 5～8 min, co-cultured for two days, finally, transferred to the MS medium supplemented with 0.5 mg/L TDZ+0.3 mg/L GA3+50 mg/L Kan+200 mg/L Car+200 mg/L Cef. Adventitious buds could emerge after induction in light for 4～5 weeks. The transformation was tested by PCR, the transformation efficiency reached to 5%. Based on this, the genetic transformation of StCOL and StTFLl was carried. The role and relationship between StCOL and StTFL1 in tuber formation were preliminarily analyzed by being compared with the wild type, transgenic lines of antisense-stCOL had a lower level of StCOL mRNA in leaves but more tubers in average. It suggested that StCOL could negatively regulate tuberization; the transgenic lines of over-expression StTFL1 had a higher level of StTFL1 mRNA in leaves but also more tubers in average. It suggested that StTFL1 could promote tuberization. Analysis of the transcription level of StTFL1 mRNA and StCOL mRNA in transgenic lines of antisense-repression StCOL and over-expression StTFL1 showed that StTFL1 was a downstream gene of StCOL.
引文
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