转iaaM和ABP1基因对棉花和苎麻纤维及细胞发育的影响
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摘要
植物生长素(auxin)是调控植物生长发育最重要的植物激素,关于生长素及其信号途径的研究逾百年来历久弥新。近年来有关生长素结合蛋白ABP1作用于植物细胞伸长和分裂的研究深受植物分子生物学界的关注。本研究以棉花和苎麻两种具有长纤维细胞的植物为材料,通过对生长素合成基因iaaM及生长素结合蛋白基因ABP1的转基因研究,探索了生长素及其结合蛋白ABP1对棉纤维的伸长、苎麻韧皮纤维及细胞发育的影响。
棉花纤维是高度伸长的胚珠表皮细胞。生长素影响着棉纤维的发育和伸长。为了确证纤维细胞内的生长素对纤维伸长的作用,本实验将催化色氨酸转变为生长素的关键酶——色氨酸单加氧酶基因iaaM重组到棉花纤维特异表达启动子E6调控的表达载体中,构建了在纤维细胞中特异表达iaaM基因的载体并以HPTⅡ作为选择标记,采用农杆菌介导的子房注射法转化棉花“湘杂棉14”,经潮霉素抗性筛选,获得了8株抗性植株。其中5株植株基因组中检测到目的基因的整合。RT-PCR分析证明纤维细胞中iaaM基因在转录水平上能有效表达,纤维细胞中的生长素含量也相应提高。但在转基因株稳定后,连续两年的利植和分析,其棉纤维长度并未发生明显改变。
生长素结合蛋白ABP1是生长素信号途径新确定的成员,但由于在模式植物拟南芥中的突变致死性使其功能的认识严重不足。为了揭示ABP1在棉花发育,尤其是棉纤维发育中的功能,本研究以“湘杂棉14”为材料,采用RT-PCR法克隆了棉花ABP1基因,该基因全长cDNA序列为792bp,与GenBank中陆地棉“新陆中5号”ABP1基因序列存在7个碱基的差别,同源性为99%,但可以编码一个保守的ABP1蛋白。采用半定量PCR方法检测了ABP1基因在棉花叶、花和开花后15d的纤维细胞中都有表达。其中在叶片中的表达量最高,在花器官和纤维细胞中的表达量稍低。为了验证ABP1在棉花纤维发育中的功能,将克隆的ABP1cDNA构建成了35S启动子控制的过表达载体并以NPTⅡ作为选择标记,经农杆菌介导转化了棉花后,筛选获得了6株Kan抗性植株,在其中3株中检测到ABP1表达水平提高。对这些ABP1过表达植株的观察分析表明,植株整体生长发育正常,纤维产量和长度变化都不显著。与受体对照植株相比,仅观察到叶缘的表皮毛密度增加。
鉴于棉花纤维细胞的特殊性,我们还选取苎麻作为材料,开展ABP1与韧皮纤维及细胞发育相关的研究。利用本研究团队已克隆的苎麻ABP1cDNA (BnABP1)构建了BnABP1基因cDNA过表达载体和反义表达载体,以HPT Ⅱ作为选择标记,分别通过农杆菌介导的叶盘转化法转化苎麻“湘苎3号”,经潮霉素抗性筛选和PCR检测获得了4株ABPl过表达植株和4株ABP1反义表达植株。ABP1过表达植株整体生长发育也完全正常,未能观察到基因过表达引起的性状改变。但ABP1反义表达载体转化苎麻后,植株生长发育表现出显著抑制现象。反义转基因苗生长发育迟缓,成苗后叶片小而卷曲,叶表皮毛减少；叶脉发育不完全,叶脉小,并且在叶片的某些部位还缺乏叶脉；叶柄的发育也表现异常。对叶片进行组织切片观察到表皮细胞和栅栏细胞变小,组成主脉和侧脉的细胞小,并且主脉维管束不发达,韧皮部细胞和导管变小,还缺少一些典型细胞层,细胞间的耦合也发生了变化。叶柄切片也观察到细胞和组织的类似变化。茎的发育也表现出薄壁细胞、韧皮部细胞、导管和维管束细小,导致转基因植株茎明显小于对照；韧皮纤维细胞数量减少的同时细胞也更为短小。
上述实验结果表明,棉花纤维作为特化的胚珠表皮细胞,伸长可能受多种因素共同调控,单纯提高细胞内生长素浓度或整体ABP1的表达水平对纤维伸长都没有显著作用。但ABP1表达水平的提高对棉花叶表皮毛的发育有一定的影响。
对于苎麻韧皮纤维及其茎和叶细胞,它们的正常发育需要一定表达量的ABP1,过表达ABP1对其并没有剂量效应,但当ABP1表达受到抑制后其量低于一定的阈值时,细胞的分化和伸长会表现出显著异常,说明ABP1通过影响细胞的分化及伸长来影响苎麻纤维及其茎和叶的发育。
对ABP1反义表达苎麻组织切片观察发现,ABP1的抑制表达并不影响其组织的初级分化。尽管植株生长受到严重抑制,但植株各组织的细胞类型分化完整,只是各类细胞都更加细小,细胞数量减少。说明ABP1表达抑制后主要影响细胞的膨大、伸长和细胞分裂频次,导致各组织中分化的细胞数量减少,器官细小。
Auxin, the most important phytohormone, plays a key role in the regulation of plant growth and development. How the signaling between it and its downstream effectors occurs has attracted widespread attention of researchers. AUXIN BINDING PROTEIN1(ABP1) is the first characterized protein that binds auxin and has been implied as a receptor for a number of auxin responses. Several lines of evidence support a role for ABP1in control of cell expansion and cell division. In this study, we used the fiber plants-cotton and ramie as the model to dissect the roles of auxin and ABP1in cotton fiber elongation, the development of ramie phloem fiber and cell development.
Cotton fiber is a highly elongated cell derived from the ovule epidermis. Auxin plays an important role in cotton fiber development. It can promote fiber elongation. A Tryptophan monooxygenase encoded by iaaM gene is the key enzyme in converting tryptophane to indole-3-acetamide which is then hydrolyzed to the indole acetic acid (IAA). So the iaaM gene has been widely used to manipulate the IAA level in various plants. To identify the function of IAA in fiber elongation, we constructed a plant expression vector carrying auxin synthesis gene iaaM coding sequence under the control of the fiber-specific E6promoter from Gossypium hirsutum and it was introduced into the genome of a cotton cultivar,'Xiangzamian14', by an Agrobacterium-mediated transformation method. Transgenic seedlings were screened out by hygromycin resistance, and then subjected to molecular analysis. PCR analysis showed that iaaM gene was successfully integrated into the cotton genome in five transgenic lines. The result of RT-PCR also verified that iaaM gene was transcripted at mRNA level in transgenic line. The IAA level in fibers at15days post anthesis was increased. However, the length of fibers was not changed significantly. It is clearly shown that excess endogenous auxin can not promote fiber cell elongation.
Auxin binding protein1(ABP1) is recently recognized as an essential receptor in auxin signaling. ABP1is especially involved in cell elongation. However, it is lack of evidence on the role of ABP1during postembryonic development due to the embryo lethal in Arabidopsis. To identify the role of ABP1in cotton development, the ABP1gene of cotton was cloned by RT-PCR in Gossypium hirsutum cultivar 'Xiangzamian 14'and its transcription level was analyzed by semi-quantitative reverse transcription polymerase chain reaction. The cDNA of GhABP1was792bp in length and shared99%identity to the cDNA of FJ609678from Gossypium hirsutum cultivar 'Xinluzhong5".It is expressed in leaves, flowers and fibers at15days post anthesis. The transcription level in young leaves is higher than that in other tissues. An over expression vector carrying ABP1gene coding sequence was constructed, and was introduced into the genome of a cotton cultivar'Xiangzamian14'by an Agrobacteriwn-mediated transformation method. Transgenic lines over expressing ABP1were generated. RT-PCR analysis showed that the transcription level of ABP1gene was up-regulated in three transgenic lines which exhibited normal morphological phenotypes. The leaf size and shape were not altered obviously but with leaf trichome density on the leaf margin increased. Moreover, over expression of ABP1did not result in significant alteration in fiber length.
Ramie [Boehmeria nivea (L.) Gaud.] is a perennial fiber plant. Its fiber is the longest phloem fiber. The role of auxin signaling in ramie development remains unclear. We were cloned ABP1cDNA (BnABP1) from ramie several years ago. Here, we focused on analyzing the effect of ABP1on ramie development. We constructed an over expression and an antisense expression vectors carrying BnABP1coding sequence under the control of the CaMV35S promoter, and they were transformed into a ramie cultivar,'xiangzhu3', by an Agrobacterium-mediated transformation method. Transgenic plantlets were screened out by hygromycin resistance and four ABP1over expression plants were generated. The transgenic lines exhibited normal morphological phenotypes. PCR analysis showed that anti-ABP1gene was integrated into the ramie genome in four transgenic lines and the ABP1expression was repressed in ABP1antisense transgenic plantlets. The suppression of ABP1expression led to defects in plant growth including dwarf plant, and decreased apical dominance in the intact plant. In addition, ABP1antisense lines exhibited serious retard of leaf development involving small and twisted leaves, and sparse trichome. Though, the cell type in leaves of ABP1antisense plants was not changed, the cell in epidermis and palisade tissue growed smaller. Moreover, the decreased ABP1expression repressed the development of leaf venation. ABP1antisense lines developed an incomplete leaf venation, which became smaller and lacked in some regions of leaves. Though the cell arrangement was normal in midrib and lateral vein, cell size was decrease and some cell layers were missing in midrib. The alteration of petiole histology was similar to that of leaf histology. ABP1antisense plants had smaller stems, which appeared to be attributable to a decrease in cell size of stems. The decreased ABP1expression repressed the development of fiber. Fiber cells were decreased and smaller. The results showed that ABP1plays an important role in ramie fiber and cell development.
In conclusion, the elongation of cotton fiber, as specialized ovule epidermal cell, may be influenced by many factors. It has no significant effect on fiber elongation by increasing either the auxin concentration or ABP1expression level in fiber cell. But the increase of ABP1expression level can has an effect on the development of leaf trichome.
For ramie phloem fiber and cell, its development requires a minimum amount of ABP1expression. The over expression of the ABP1have no dose effect on its development. However, the fiber development is significant anomaly when the ABP1expression is below a certain threshold by antisense RNA. The differentiation and elongation of the fibers as well as other cells is affected apparently. It has shown ABP1should keep in an amount for the normal growth and development of ramie. ABP1is a major regulator for the phloem fiber cell differentiation and elongation.
All expected cell types in leaves and stems of ABP1antisense lines are present, suggesting that suppression of ABP1gene does not repress primary morphogenesis of the tissues. However, the size and number of cells in the tissues is decreased, which has showed that ABP1can act on cell division and elongation. The mutant phenotypes of ABP1antisense lines are attributed to abnormality of cell division and elongation.
棉花纤维是高度伸长的胚珠表皮细胞。生长素影响着棉纤维的发育和伸长。为了确证纤维细胞内的生长素对纤维伸长的作用,本实验将催化色氨酸转变为生长素的关键酶——色氨酸单加氧酶基因iaaM重组到棉花纤维特异表达启动子E6调控的表达载体中,构建了在纤维细胞中特异表达iaaM基因的载体并以HPTⅡ作为选择标记,采用农杆菌介导的子房注射法转化棉花“湘杂棉14”,经潮霉素抗性筛选,获得了8株抗性植株。其中5株植株基因组中检测到目的基因的整合。RT-PCR分析证明纤维细胞中iaaM基因在转录水平上能有效表达,纤维细胞中的生长素含量也相应提高。但在转基因株稳定后,连续两年的利植和分析,其棉纤维长度并未发生明显改变。
生长素结合蛋白ABP1是生长素信号途径新确定的成员,但由于在模式植物拟南芥中的突变致死性使其功能的认识严重不足。为了揭示ABP1在棉花发育,尤其是棉纤维发育中的功能,本研究以“湘杂棉14”为材料,采用RT-PCR法克隆了棉花ABP1基因,该基因全长cDNA序列为792bp,与GenBank中陆地棉“新陆中5号”ABP1基因序列存在7个碱基的差别,同源性为99%,但可以编码一个保守的ABP1蛋白。采用半定量PCR方法检测了ABP1基因在棉花叶、花和开花后15d的纤维细胞中都有表达。其中在叶片中的表达量最高,在花器官和纤维细胞中的表达量稍低。为了验证ABP1在棉花纤维发育中的功能,将克隆的ABP1cDNA构建成了35S启动子控制的过表达载体并以NPTⅡ作为选择标记,经农杆菌介导转化了棉花后,筛选获得了6株Kan抗性植株,在其中3株中检测到ABP1表达水平提高。对这些ABP1过表达植株的观察分析表明,植株整体生长发育正常,纤维产量和长度变化都不显著。与受体对照植株相比,仅观察到叶缘的表皮毛密度增加。
鉴于棉花纤维细胞的特殊性,我们还选取苎麻作为材料,开展ABP1与韧皮纤维及细胞发育相关的研究。利用本研究团队已克隆的苎麻ABP1cDNA (BnABP1)构建了BnABP1基因cDNA过表达载体和反义表达载体,以HPT Ⅱ作为选择标记,分别通过农杆菌介导的叶盘转化法转化苎麻“湘苎3号”,经潮霉素抗性筛选和PCR检测获得了4株ABPl过表达植株和4株ABP1反义表达植株。ABP1过表达植株整体生长发育也完全正常,未能观察到基因过表达引起的性状改变。但ABP1反义表达载体转化苎麻后,植株生长发育表现出显著抑制现象。反义转基因苗生长发育迟缓,成苗后叶片小而卷曲,叶表皮毛减少；叶脉发育不完全,叶脉小,并且在叶片的某些部位还缺乏叶脉；叶柄的发育也表现异常。对叶片进行组织切片观察到表皮细胞和栅栏细胞变小,组成主脉和侧脉的细胞小,并且主脉维管束不发达,韧皮部细胞和导管变小,还缺少一些典型细胞层,细胞间的耦合也发生了变化。叶柄切片也观察到细胞和组织的类似变化。茎的发育也表现出薄壁细胞、韧皮部细胞、导管和维管束细小,导致转基因植株茎明显小于对照；韧皮纤维细胞数量减少的同时细胞也更为短小。
上述实验结果表明,棉花纤维作为特化的胚珠表皮细胞,伸长可能受多种因素共同调控,单纯提高细胞内生长素浓度或整体ABP1的表达水平对纤维伸长都没有显著作用。但ABP1表达水平的提高对棉花叶表皮毛的发育有一定的影响。
对于苎麻韧皮纤维及其茎和叶细胞,它们的正常发育需要一定表达量的ABP1,过表达ABP1对其并没有剂量效应,但当ABP1表达受到抑制后其量低于一定的阈值时,细胞的分化和伸长会表现出显著异常,说明ABP1通过影响细胞的分化及伸长来影响苎麻纤维及其茎和叶的发育。
对ABP1反义表达苎麻组织切片观察发现,ABP1的抑制表达并不影响其组织的初级分化。尽管植株生长受到严重抑制,但植株各组织的细胞类型分化完整,只是各类细胞都更加细小,细胞数量减少。说明ABP1表达抑制后主要影响细胞的膨大、伸长和细胞分裂频次,导致各组织中分化的细胞数量减少,器官细小。
Auxin, the most important phytohormone, plays a key role in the regulation of plant growth and development. How the signaling between it and its downstream effectors occurs has attracted widespread attention of researchers. AUXIN BINDING PROTEIN1(ABP1) is the first characterized protein that binds auxin and has been implied as a receptor for a number of auxin responses. Several lines of evidence support a role for ABP1in control of cell expansion and cell division. In this study, we used the fiber plants-cotton and ramie as the model to dissect the roles of auxin and ABP1in cotton fiber elongation, the development of ramie phloem fiber and cell development.
Cotton fiber is a highly elongated cell derived from the ovule epidermis. Auxin plays an important role in cotton fiber development. It can promote fiber elongation. A Tryptophan monooxygenase encoded by iaaM gene is the key enzyme in converting tryptophane to indole-3-acetamide which is then hydrolyzed to the indole acetic acid (IAA). So the iaaM gene has been widely used to manipulate the IAA level in various plants. To identify the function of IAA in fiber elongation, we constructed a plant expression vector carrying auxin synthesis gene iaaM coding sequence under the control of the fiber-specific E6promoter from Gossypium hirsutum and it was introduced into the genome of a cotton cultivar,'Xiangzamian14', by an Agrobacterium-mediated transformation method. Transgenic seedlings were screened out by hygromycin resistance, and then subjected to molecular analysis. PCR analysis showed that iaaM gene was successfully integrated into the cotton genome in five transgenic lines. The result of RT-PCR also verified that iaaM gene was transcripted at mRNA level in transgenic line. The IAA level in fibers at15days post anthesis was increased. However, the length of fibers was not changed significantly. It is clearly shown that excess endogenous auxin can not promote fiber cell elongation.
Auxin binding protein1(ABP1) is recently recognized as an essential receptor in auxin signaling. ABP1is especially involved in cell elongation. However, it is lack of evidence on the role of ABP1during postembryonic development due to the embryo lethal in Arabidopsis. To identify the role of ABP1in cotton development, the ABP1gene of cotton was cloned by RT-PCR in Gossypium hirsutum cultivar 'Xiangzamian 14'and its transcription level was analyzed by semi-quantitative reverse transcription polymerase chain reaction. The cDNA of GhABP1was792bp in length and shared99%identity to the cDNA of FJ609678from Gossypium hirsutum cultivar 'Xinluzhong5".It is expressed in leaves, flowers and fibers at15days post anthesis. The transcription level in young leaves is higher than that in other tissues. An over expression vector carrying ABP1gene coding sequence was constructed, and was introduced into the genome of a cotton cultivar'Xiangzamian14'by an Agrobacteriwn-mediated transformation method. Transgenic lines over expressing ABP1were generated. RT-PCR analysis showed that the transcription level of ABP1gene was up-regulated in three transgenic lines which exhibited normal morphological phenotypes. The leaf size and shape were not altered obviously but with leaf trichome density on the leaf margin increased. Moreover, over expression of ABP1did not result in significant alteration in fiber length.
Ramie [Boehmeria nivea (L.) Gaud.] is a perennial fiber plant. Its fiber is the longest phloem fiber. The role of auxin signaling in ramie development remains unclear. We were cloned ABP1cDNA (BnABP1) from ramie several years ago. Here, we focused on analyzing the effect of ABP1on ramie development. We constructed an over expression and an antisense expression vectors carrying BnABP1coding sequence under the control of the CaMV35S promoter, and they were transformed into a ramie cultivar,'xiangzhu3', by an Agrobacterium-mediated transformation method. Transgenic plantlets were screened out by hygromycin resistance and four ABP1over expression plants were generated. The transgenic lines exhibited normal morphological phenotypes. PCR analysis showed that anti-ABP1gene was integrated into the ramie genome in four transgenic lines and the ABP1expression was repressed in ABP1antisense transgenic plantlets. The suppression of ABP1expression led to defects in plant growth including dwarf plant, and decreased apical dominance in the intact plant. In addition, ABP1antisense lines exhibited serious retard of leaf development involving small and twisted leaves, and sparse trichome. Though, the cell type in leaves of ABP1antisense plants was not changed, the cell in epidermis and palisade tissue growed smaller. Moreover, the decreased ABP1expression repressed the development of leaf venation. ABP1antisense lines developed an incomplete leaf venation, which became smaller and lacked in some regions of leaves. Though the cell arrangement was normal in midrib and lateral vein, cell size was decrease and some cell layers were missing in midrib. The alteration of petiole histology was similar to that of leaf histology. ABP1antisense plants had smaller stems, which appeared to be attributable to a decrease in cell size of stems. The decreased ABP1expression repressed the development of fiber. Fiber cells were decreased and smaller. The results showed that ABP1plays an important role in ramie fiber and cell development.
In conclusion, the elongation of cotton fiber, as specialized ovule epidermal cell, may be influenced by many factors. It has no significant effect on fiber elongation by increasing either the auxin concentration or ABP1expression level in fiber cell. But the increase of ABP1expression level can has an effect on the development of leaf trichome.
For ramie phloem fiber and cell, its development requires a minimum amount of ABP1expression. The over expression of the ABP1have no dose effect on its development. However, the fiber development is significant anomaly when the ABP1expression is below a certain threshold by antisense RNA. The differentiation and elongation of the fibers as well as other cells is affected apparently. It has shown ABP1should keep in an amount for the normal growth and development of ramie. ABP1is a major regulator for the phloem fiber cell differentiation and elongation.
All expected cell types in leaves and stems of ABP1antisense lines are present, suggesting that suppression of ABP1gene does not repress primary morphogenesis of the tissues. However, the size and number of cells in the tissues is decreased, which has showed that ABP1can act on cell division and elongation. The mutant phenotypes of ABP1antisense lines are attributed to abnormality of cell division and elongation.
引文
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