油菜质膜水孔蛋白BnPIP1基因的功能分析及其上游调控区的研究
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摘要
植物水孔蛋白在植物体内形成水选择性运输通道，介导胞内或胞间的水分跨膜运输，并参与植物体的许多重要生理过程及逆境应答反应。本研究的主要目的是通过在烟草中正义和反义表达油菜种子萌发特异性表达的质膜水孔蛋白BnPIP1基因来探讨水孔蛋白在植物体内的水分运输功能及其在植物耐干旱胁迫中的意义。并通过克隆BnPIP1基因的上游调控序列初步研究了BnPIP1基因的时空表达模式。本研究为了解植物干旱胁迫下水分运输的调控机制提供了新的思路，也为进一步研究BnPIP1基因的表达特性奠定了基础。主要研究结果如下：
1．利用RT-PCR方法从油菜幼苗叶片总RNA中克隆出油菜质膜水孔蛋白BnPIP1基因的全长cDNA片段。该片段与文献报道的序列具有99％的同源性。存在4处碱基突变，仅导致一处氨基酸突变，推测为是物种内不同品种间的遗传差异。
2．以CaMV35S为启动子，以Tnos为终止子，分别构建了BnPIP1基因的正义和反义植物表达载体，用以研究BnPIP1基因在植物体内的生理功能。通过农杆菌介导的方法转化烟草叶盘。研究结果表明：异源反义表达BnPIP1基因，可以抑伟烟草内源水孔蛋白的表达，使植物水分运输能力明显下降。表现为叶肉原生质体在低渗溶液中处理2小时后才有80％会破裂，整株抗旱性弱。转正义BnPIP1基因的植株则通过增加外源水孔蛋白的含量实现了对水分运输快速灵敏的调控，表现为在低渗溶液中，90％以上的叶肉原生质体在5分钟内快速破裂，整株抗旱性强及T_0代种子在高渗土壤中萌发早而整齐。
3．转基因T_0代种子在含抗生素培养基上的萌发试验表明，不同转基因株系中外源基因的整合方式不同。有的株系外源基因以单拷贝或多拷贝的形式整合在一条染色体上，T_0代种子中外源基因呈现出近3：1的分离比。有的转基因株系中，外源基因以多拷贝的形式整合在不同的染色体上，T_0代种子中外源基因的分离比例比较复杂。
4．利用连接介导PCR的染色体步行技术，我们克隆了长为1610bp的油菜质膜水孔蛋白BnPIP1基因的上游调控序列。该序列存在着两个可能的TATA-box序列、一个G-box序列、4个TGAC序列、若干个TGAC-like序列及较多的AT丰富序列，这些顺势元件分别是RNA聚合酶Ⅱ、bZIP蛋白及AT丰富序列结合蛋白的识别位点，可能会在基因的转录与表达中起到调控作用。Blast分析表明，该序列为一新的调控序列，GeneBank上的登录号为AF472487。
博士学位论文 中 文摘要 于秋菊
5．利用双元植物表达载体 pBll21，pCAMBIA1305．1及专门用于检测启动子活性的
pPR97载体，构建了 BnPNI上游 1．6kb调控序列与牙m的融合基因。在含适当激素
的MS培养基上，再生小芽的组织化学染色表明，其驱动》份基因的表达强度与
pBllZI中 CaMV 355相当。GUS染色主要分布在水分需求量大的细胞迅速增生的部
位及运输水分的微管组织中。并初步推断该启动子可能具有种子萌发特异性。
6．利用 PCR扩增技术，将 1．6kb的上游调控区从 5’端分别缺失 obp，580hp，708hp和
1246…，构建了四个双元表达载体p97－0，p97－580，p97－708和p97－1246。转化烟草
愈伤组织的 GUS染色结果初步将整个启动子分为 3个区段：区段 1，－16lffe－1030 hP，
是启动子的正调控区，能指导算m基因大量表达；区段 2，－103De－902 hp，可能含
有启动子的负调控元件，强烈J：llial4）制着算凹基因的表达；区段3，－902～－19 hp，该
区具有全长启动子片断同样的活性，亦可指导9吩基因的表达。
Aquaporins make water-selective channels in plants, facilitating the permeation of water through membranes, and are involved in many important physiological processes as well as the response to stresses. This study aimed at investigating the transport function in plants of seed-germination specific plasma membrane aquaporin BnPIPl from Brassica napus and its meanings to plant drought tolerance.
The upstream sequence of BnPIPl was cloned and used for the study of time-space expression pattern of BnPIPl gene. This work has a high academic value and practical significance, which provides a new way for understanding the mechanisms of water transport in plants under water stress and makes a foundation for further study of BnPIPl expression pattern. The followings are the main results:
1. A full length cDNA fragment encoding plasma membrane aquaporin BnPIPl was cloned from the leaf total RNA of Brassica napus by RT-PCR method. Sequencing analysis indicated that this fragment showed 99% identity to the previously reported one. There were four mutant sites in this sequence and resulted in only one amino acid change, which was deduced to be the genetic variation between different species.
2. Two plant expression vectors with sense or antisense BnPIPlgene, driven by CaMVSSS promoter and terminated by Tnos, were constructed and transformed into tobacco discs via the mediation of Agrobacteriwn tumefaciens to study of physiological functions of BnPIPl in plant. Form
the expression of sense or antisense BnPIPl gene in transgenic tobaccos, we could draw a conclusion that antisense expression of BnPIPl could suppress the expression of endoaquaporins in tobaccos, resulting in significant reduction in water transport, longer time needed for leaf protoplasts to burst in hypotonic solution and sensitivity to water stress. Whereas, the lines overexpressing sense BnPIPlgene could fast and finely control the transport of water, leading to quick burst of leaf protoplasts in hypotonic solution, tolerance to water stress and early and even germination of TO seeds in high osmotic pressure soil.
3. The study of T0 seeds germination on MS mediums with antibiotic indicated that transgene had different integration patterns in different transgenic lines. In some lines, the foreign gene integrated into one chromosome with one or mutiple copies at one or more loci and segregated in TO seeds with the ratio about 3:1. In some other lines, the foreign
gene integrated into different chromosomes with multiple copies at different sites, showing complex segregation ratio.
4. The upstream sequence of BnPIPl gene was cloned by genomic walking based on ligation- mediated PCR method. This sequence was 1610bp in length, containing two possible TATA-box, one G-box, four TGAC sequences and several TGAC-like sequences. Blast analysis showed that this was a novel sequence. The GenBank accession number was AF472487.
5. Binary vectors pBI121, pCAMBIA1305.1 and binary promoter indicator vector pPR97 were used to construct plant expression vectors, in which the foil length upstream sequence of BnPIPl was fused with gus gene. After the treatment on MS medium with proper hormones, the regenerated shoots from transgenic tobacco calli were detected for GUS activity. The result showed that the full length sequence had promoter activity, the intensity of GUS was as strong as that of GUS driven by CaMVSSS promoter. GUS staining was mainly localized in the parts in which cells grow and proliferate fast and need vast water supply, or in the vessel tissues which function in water transport. At the same time, we deduced that this promoter probably was a seed-germination specific promoter.
6. A series of deletion vectors named as p97-0, p97-581, p97-708 and p97-1246 were constructed, in which Obp,580bp, 708bp and 1246bp were deleted from 5' end of the upstream sequence of BnPIPl by PCR method, respectively. Calli from transgenic tobaccos were used for GUS staining and the promoter was divided into three regions accordingly. Region 1: -1
1．利用RT-PCR方法从油菜幼苗叶片总RNA中克隆出油菜质膜水孔蛋白BnPIP1基因的全长cDNA片段。该片段与文献报道的序列具有99％的同源性。存在4处碱基突变，仅导致一处氨基酸突变，推测为是物种内不同品种间的遗传差异。
2．以CaMV35S为启动子，以Tnos为终止子，分别构建了BnPIP1基因的正义和反义植物表达载体，用以研究BnPIP1基因在植物体内的生理功能。通过农杆菌介导的方法转化烟草叶盘。研究结果表明：异源反义表达BnPIP1基因，可以抑伟烟草内源水孔蛋白的表达，使植物水分运输能力明显下降。表现为叶肉原生质体在低渗溶液中处理2小时后才有80％会破裂，整株抗旱性弱。转正义BnPIP1基因的植株则通过增加外源水孔蛋白的含量实现了对水分运输快速灵敏的调控，表现为在低渗溶液中，90％以上的叶肉原生质体在5分钟内快速破裂，整株抗旱性强及T_0代种子在高渗土壤中萌发早而整齐。
3．转基因T_0代种子在含抗生素培养基上的萌发试验表明，不同转基因株系中外源基因的整合方式不同。有的株系外源基因以单拷贝或多拷贝的形式整合在一条染色体上，T_0代种子中外源基因呈现出近3：1的分离比。有的转基因株系中，外源基因以多拷贝的形式整合在不同的染色体上，T_0代种子中外源基因的分离比例比较复杂。
4．利用连接介导PCR的染色体步行技术，我们克隆了长为1610bp的油菜质膜水孔蛋白BnPIP1基因的上游调控序列。该序列存在着两个可能的TATA-box序列、一个G-box序列、4个TGAC序列、若干个TGAC-like序列及较多的AT丰富序列，这些顺势元件分别是RNA聚合酶Ⅱ、bZIP蛋白及AT丰富序列结合蛋白的识别位点，可能会在基因的转录与表达中起到调控作用。Blast分析表明，该序列为一新的调控序列，GeneBank上的登录号为AF472487。
博士学位论文 中 文摘要 于秋菊
5．利用双元植物表达载体 pBll21，pCAMBIA1305．1及专门用于检测启动子活性的
pPR97载体，构建了 BnPNI上游 1．6kb调控序列与牙m的融合基因。在含适当激素
的MS培养基上，再生小芽的组织化学染色表明，其驱动》份基因的表达强度与
pBllZI中 CaMV 355相当。GUS染色主要分布在水分需求量大的细胞迅速增生的部
位及运输水分的微管组织中。并初步推断该启动子可能具有种子萌发特异性。
6．利用 PCR扩增技术，将 1．6kb的上游调控区从 5’端分别缺失 obp，580hp，708hp和
1246…，构建了四个双元表达载体p97－0，p97－580，p97－708和p97－1246。转化烟草
愈伤组织的 GUS染色结果初步将整个启动子分为 3个区段：区段 1，－16lffe－1030 hP，
是启动子的正调控区，能指导算m基因大量表达；区段 2，－103De－902 hp，可能含
有启动子的负调控元件，强烈J：llial4）制着算凹基因的表达；区段3，－902～－19 hp，该
区具有全长启动子片断同样的活性，亦可指导9吩基因的表达。
Aquaporins make water-selective channels in plants, facilitating the permeation of water through membranes, and are involved in many important physiological processes as well as the response to stresses. This study aimed at investigating the transport function in plants of seed-germination specific plasma membrane aquaporin BnPIPl from Brassica napus and its meanings to plant drought tolerance.
The upstream sequence of BnPIPl was cloned and used for the study of time-space expression pattern of BnPIPl gene. This work has a high academic value and practical significance, which provides a new way for understanding the mechanisms of water transport in plants under water stress and makes a foundation for further study of BnPIPl expression pattern. The followings are the main results:
1. A full length cDNA fragment encoding plasma membrane aquaporin BnPIPl was cloned from the leaf total RNA of Brassica napus by RT-PCR method. Sequencing analysis indicated that this fragment showed 99% identity to the previously reported one. There were four mutant sites in this sequence and resulted in only one amino acid change, which was deduced to be the genetic variation between different species.
2. Two plant expression vectors with sense or antisense BnPIPlgene, driven by CaMVSSS promoter and terminated by Tnos, were constructed and transformed into tobacco discs via the mediation of Agrobacteriwn tumefaciens to study of physiological functions of BnPIPl in plant. Form
the expression of sense or antisense BnPIPl gene in transgenic tobaccos, we could draw a conclusion that antisense expression of BnPIPl could suppress the expression of endoaquaporins in tobaccos, resulting in significant reduction in water transport, longer time needed for leaf protoplasts to burst in hypotonic solution and sensitivity to water stress. Whereas, the lines overexpressing sense BnPIPlgene could fast and finely control the transport of water, leading to quick burst of leaf protoplasts in hypotonic solution, tolerance to water stress and early and even germination of TO seeds in high osmotic pressure soil.
3. The study of T0 seeds germination on MS mediums with antibiotic indicated that transgene had different integration patterns in different transgenic lines. In some lines, the foreign gene integrated into one chromosome with one or mutiple copies at one or more loci and segregated in TO seeds with the ratio about 3:1. In some other lines, the foreign
gene integrated into different chromosomes with multiple copies at different sites, showing complex segregation ratio.
4. The upstream sequence of BnPIPl gene was cloned by genomic walking based on ligation- mediated PCR method. This sequence was 1610bp in length, containing two possible TATA-box, one G-box, four TGAC sequences and several TGAC-like sequences. Blast analysis showed that this was a novel sequence. The GenBank accession number was AF472487.
5. Binary vectors pBI121, pCAMBIA1305.1 and binary promoter indicator vector pPR97 were used to construct plant expression vectors, in which the foil length upstream sequence of BnPIPl was fused with gus gene. After the treatment on MS medium with proper hormones, the regenerated shoots from transgenic tobacco calli were detected for GUS activity. The result showed that the full length sequence had promoter activity, the intensity of GUS was as strong as that of GUS driven by CaMVSSS promoter. GUS staining was mainly localized in the parts in which cells grow and proliferate fast and need vast water supply, or in the vessel tissues which function in water transport. At the same time, we deduced that this promoter probably was a seed-germination specific promoter.
6. A series of deletion vectors named as p97-0, p97-581, p97-708 and p97-1246 were constructed, in which Obp,580bp, 708bp and 1246bp were deleted from 5' end of the upstream sequence of BnPIPl by PCR method, respectively. Calli from transgenic tobaccos were used for GUS staining and the promoter was divided into three regions accordingly. Region 1: -1
引文
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