棉花蔗糖合成酶基因（Sucrose Synthase 3）启动子结构、功能及其调控分析
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摘要
棉花是一种种植非常广泛的重要经济作物,它主要提供天然的纺织纤维。棉纤维是一个单细胞体,因此可作为研究细胞伸长、细胞壁和纤维素合成的一个模式系统。其发育过程可分为4个相互交迭的阶段:起始、伸长、次生壁形成、成熟,由此得到成熟的棉纤维。棉花在开花时,大约有15-25%的胚珠表皮层细胞可分化发育成长绒纤维细胞。至今,人们对棉花纤维细胞起始的分子机制还了解的很少。棉花蔗糖合成酶(SuSy)是近十年的研究所发现的一系列在棉纤维起始阶段起调控作用的功能基因之一。SuSy催化蔗糖分解为果糖和UDP-葡萄糖,为纤维素的合成提供底物。研究者发现SuSy促进棉纤维细胞的分化与伸长,同时SuSy在棉纤维细胞的起始和伸长等一系列活动中起着至关重要的作用,以RNA干扰的方式抑制该基因的表达后会使转基因棉花出现无毛的表型。
研究与棉纤维发育相关基因的启动子元件及调节这些基因表达的转录因子有助于了解棉花纤维起始和发育相关的分子机制。拟南芥从结构和遗传学来看,它与棉花纤维有一定的相似性。拟南芥的转基因技术成熟,效率相对较高,且生长周期短,一般2-3个月即可完成一代。因此,本研究利用拟南芥来进行棉花SuSy启动子及相关调控的研究。结果如下:
1.根据Genbank中SuSy的cDNA序列(U73588)获得了该基因的基因组DNA全长序列,与拟南芥中的蔗糖合成酶家族进行比较分析后发现,棉花SuSy与拟南芥中的AtSus1和AtSus4在基因的结构及外显子的长度上都有非常高的相似性;
2.经过三次基因组步移后,成功获得了长为1554bp的SuSy基因5′上游序列,同时分离出了SuSy编码区的第一个内含子,构建了启动子不用长度的5′端缺失载体及内含子相关的载体,通过农杆菌介导转化野生型拟南芥,经过筛选后获得可以稳定遗传的各载体转基因拟南芥株系;
3.棉花SuSy启动子可以驱动GUS基因在转基因拟南芥的维管组织、毛刺、子叶、根、根毛以及花器官中表达。-554bp到-1554bp之间的片段对启动子的活性有非常明显的增强作用,可以达到20倍以上;
4.赤霉素只对1152bp的启动子片段有调控作用,可以提高2.5倍GUS的活性,但对全长及810bp以内的启动子片段没有调控作用;
5.启动子-296到-376之间的80bp片段对SuSy启动子在拟南芥毛刺中的活性起着决定作用,同时也对其在花粉中的活性有一定的影响;
6.启动子上-96bp处有一个W-box序列,将该序列突变后并不影响启动子的组织特异性,但可以降低启动子的活性为原来的1/3:
7.棉花SuSy基因编码区的第一个内含子在与其启动子协同下可以特异抑制GUS基因在花粉中表达,并且在其他组织中降低GUS的活性,这种抑制作用可以达到一个数量级;
8.以赤霉素对内含子相关的转基因拟南芥进行处理,可以使GUS的活性提高4倍,这种增强作用可能与内含子上的P-box有关。
Cotton fibers play an important economic role in the world, it supplys the crude fiber for textile. Each cotton fiber is a highly elongated single cell of maternal origin that initiates from the ovule epidermis outer layer at or just before anthesis. It is divided into four distinct but overlapping phases: initiation, elongation, secondary wall synthesis and maturation. About 15-25%of the epidermal layer cells differentiates and develops lint fibers. In recent decade many studies were focused on molecular mechanism of the cotton fiber development. The cotton sucrose synthase 3 (SuSy) is one of the functional genes which were found involving in cotton fiber cell initiation and elongation, plays a role in carbohydrate partitioning and ovule development. It catalyzes a reversible reaction from sucrose to fructose and UDP-glucose, and the latter is the prefect substrate for cellulose biosynthesis in cotton fiber. Several studies have indicated that SuSy performed a very important role during the fiber initiation and elongation. Furthermore, down-regulation of SuSy mRNA levels in the cotton ovular epidermis by RNAi leads a fiber-less phenotype.
Efforts to understand and manipulate fiber development would be enhanced by an improved knowledge of the promoter elements and transcription factors that regulate fiber relative gene expression. The structure and genetic control of trichome morphogenesis of Arabidopsis have been extensively characterized, showing a number of structural and genetic similarities to cotton fibers. Thus, it has been possible to monitor the activity of cotton fiber relative promoters in this hetreologous model specie. The results are described as following:
1. According to the sequence of SuSy cDNA (U73588), genomic DNA of this gene was isolated by PCR. There are 13 exons in the gene coding region and its genomic structure has a high similarity with Sus1 and Sus4 of Arabidopsis thaliana in the length of exons;
2. A1554bp promoter fragment was isolated by LA PCR~(TM) in vitro cloning kit after three times PCR- based genome walking. The promoter then fused to GUS gene with a part of the first exon of SuSy and three binary vectors related to intron were constructed. All resulting vectors were transformed into Arabidopsis thaliana mediated by agrobacterium;
3. The transgenic plants demonstrated highly specific GUS expression in the vascular tissue, trichome, cotyledon, root, root hair and flower. The region between -554 and -1554 enhanced the GUS activity to more than 20 fold more than that of 554bp promoter fragment;
4. GA_3 could make the GUS activity of 1152bp promoter fragment (Susp8::1301) increasing 2.5 times, but has no effect on that of the whole promoter (Susp9::1301)and the other fragments less than 810bp (Susp7::1301, Susp6::1301, Susp5::1301, Susp4::1301);
5. The fragment from the -296 to -396 is essential for the promoter activity in transgenic Arabidopsis trichome, and also play a role in driving GUS expression in pollen;
6. Mutation of a W-box in the position of -96bp reduced the GUS activity in transgenic plants, but no changes were found of the promoter tissue specificity;
7. The intron specific inhibited the GUS expression in mature pollen and reduced 90% GUS activity in other tissues;
8. GA_3 enhanced the GUS activity in Susp9fi::1301 transgenic plant with 4 fold.
研究与棉纤维发育相关基因的启动子元件及调节这些基因表达的转录因子有助于了解棉花纤维起始和发育相关的分子机制。拟南芥从结构和遗传学来看,它与棉花纤维有一定的相似性。拟南芥的转基因技术成熟,效率相对较高,且生长周期短,一般2-3个月即可完成一代。因此,本研究利用拟南芥来进行棉花SuSy启动子及相关调控的研究。结果如下:
1.根据Genbank中SuSy的cDNA序列(U73588)获得了该基因的基因组DNA全长序列,与拟南芥中的蔗糖合成酶家族进行比较分析后发现,棉花SuSy与拟南芥中的AtSus1和AtSus4在基因的结构及外显子的长度上都有非常高的相似性;
2.经过三次基因组步移后,成功获得了长为1554bp的SuSy基因5′上游序列,同时分离出了SuSy编码区的第一个内含子,构建了启动子不用长度的5′端缺失载体及内含子相关的载体,通过农杆菌介导转化野生型拟南芥,经过筛选后获得可以稳定遗传的各载体转基因拟南芥株系;
3.棉花SuSy启动子可以驱动GUS基因在转基因拟南芥的维管组织、毛刺、子叶、根、根毛以及花器官中表达。-554bp到-1554bp之间的片段对启动子的活性有非常明显的增强作用,可以达到20倍以上;
4.赤霉素只对1152bp的启动子片段有调控作用,可以提高2.5倍GUS的活性,但对全长及810bp以内的启动子片段没有调控作用;
5.启动子-296到-376之间的80bp片段对SuSy启动子在拟南芥毛刺中的活性起着决定作用,同时也对其在花粉中的活性有一定的影响;
6.启动子上-96bp处有一个W-box序列,将该序列突变后并不影响启动子的组织特异性,但可以降低启动子的活性为原来的1/3:
7.棉花SuSy基因编码区的第一个内含子在与其启动子协同下可以特异抑制GUS基因在花粉中表达,并且在其他组织中降低GUS的活性,这种抑制作用可以达到一个数量级;
8.以赤霉素对内含子相关的转基因拟南芥进行处理,可以使GUS的活性提高4倍,这种增强作用可能与内含子上的P-box有关。
Cotton fibers play an important economic role in the world, it supplys the crude fiber for textile. Each cotton fiber is a highly elongated single cell of maternal origin that initiates from the ovule epidermis outer layer at or just before anthesis. It is divided into four distinct but overlapping phases: initiation, elongation, secondary wall synthesis and maturation. About 15-25%of the epidermal layer cells differentiates and develops lint fibers. In recent decade many studies were focused on molecular mechanism of the cotton fiber development. The cotton sucrose synthase 3 (SuSy) is one of the functional genes which were found involving in cotton fiber cell initiation and elongation, plays a role in carbohydrate partitioning and ovule development. It catalyzes a reversible reaction from sucrose to fructose and UDP-glucose, and the latter is the prefect substrate for cellulose biosynthesis in cotton fiber. Several studies have indicated that SuSy performed a very important role during the fiber initiation and elongation. Furthermore, down-regulation of SuSy mRNA levels in the cotton ovular epidermis by RNAi leads a fiber-less phenotype.
Efforts to understand and manipulate fiber development would be enhanced by an improved knowledge of the promoter elements and transcription factors that regulate fiber relative gene expression. The structure and genetic control of trichome morphogenesis of Arabidopsis have been extensively characterized, showing a number of structural and genetic similarities to cotton fibers. Thus, it has been possible to monitor the activity of cotton fiber relative promoters in this hetreologous model specie. The results are described as following:
1. According to the sequence of SuSy cDNA (U73588), genomic DNA of this gene was isolated by PCR. There are 13 exons in the gene coding region and its genomic structure has a high similarity with Sus1 and Sus4 of Arabidopsis thaliana in the length of exons;
2. A1554bp promoter fragment was isolated by LA PCR~(TM) in vitro cloning kit after three times PCR- based genome walking. The promoter then fused to GUS gene with a part of the first exon of SuSy and three binary vectors related to intron were constructed. All resulting vectors were transformed into Arabidopsis thaliana mediated by agrobacterium;
3. The transgenic plants demonstrated highly specific GUS expression in the vascular tissue, trichome, cotyledon, root, root hair and flower. The region between -554 and -1554 enhanced the GUS activity to more than 20 fold more than that of 554bp promoter fragment;
4. GA_3 could make the GUS activity of 1152bp promoter fragment (Susp8::1301) increasing 2.5 times, but has no effect on that of the whole promoter (Susp9::1301)and the other fragments less than 810bp (Susp7::1301, Susp6::1301, Susp5::1301, Susp4::1301);
5. The fragment from the -296 to -396 is essential for the promoter activity in transgenic Arabidopsis trichome, and also play a role in driving GUS expression in pollen;
6. Mutation of a W-box in the position of -96bp reduced the GUS activity in transgenic plants, but no changes were found of the promoter tissue specificity;
7. The intron specific inhibited the GUS expression in mature pollen and reduced 90% GUS activity in other tissues;
8. GA_3 enhanced the GUS activity in Susp9fi::1301 transgenic plant with 4 fold.
引文
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