拟南芥微丝结合蛋白Profilin3体内生理功能研究
摘要
微丝骨架是存在于所有真核细胞中高度动态变化的三维网状结构,不仅为细胞提供内在的机械支持,同时也参与了许多重要的细胞过程。微丝骨架这些功能的实现既依赖于微丝骨架自身的组织结构,又依赖于微丝结合蛋白(actin binding proteins, ABPs)和不同信号分子调节的微丝骨架动态变化。植物细胞中,许多ABPs已被发现并证明参与不同的生理功能。Profilin是真核生物中发现较早,高度保守且表达量较多的一类ABPs。拟南芥基因组中有5个高度保守的profilin异型体。目前对于该家族成员的生化功能有较多的报道,而对于其体内生理功能的研究却非常匮乏。Profilin3(PRF3)是一个组成型表达的profilin基因,与拟南芥其他profilin基因相比,该基因的N末端多了36个氨基酸残基,而其生理生化功能则完全未知。因此,本研究首次系统的研究了PRF3的组织定位、生理功能等内容,并得到了以下结论：
1.PRF3是一类组成型表达的profilin异型体。通过GUS活性分析和荧光定量PCR的方法证实PRF3主要在幼苗,莲座叶和茎生叶中表达,在14天的小苗和花中的表达量很少,并且该基因的内含子可能增强基因的表达。这一结果揭示了PRF3的组织表达特征。
2.体外生化功能表明PRF3增加actin聚合的临界浓度,并且PRF3-ΔN36的Cc值最大,对actin聚合的影响最明显,而PRF3的Cc值介于PRF3-ΔN36和actin之间。这说明了PRF3的N端36个氨基酸与其蛋白功能有密切的关系。
3.与野生型拟南芥相比,过表达全长PRF3的cDNA序列的转基因植株缩短小苗的根长,下胚轴和根毛的长度,延迟种子的萌发,但是PRF3-ΔN36转基因植株和pf3突变体没有明显的表型。这一结果说明PRF3是根,下胚轴和根毛生长的重要的负调节因子,并且只有当基因完整时才具有这种功能。
4.光学显微镜和扫描电镜的观察结果显示OE-PRF3植株的下胚轴细胞的长度明显较短,细胞伸长受到抑制,说明PRF3与下胚轴细胞的伸长有着密切的关系。
5.激光共聚焦扫描显微镜观察发现OE-PRF3过表达植株的下胚轴细胞中粗的微丝束消失,但是细的微丝依然存在没有明显的变化,说明过表达PRF3全长基因能够引发微丝发束解聚,造成F-actin的重组,而粗的微丝束结构可能与细胞伸长密切相关。
6.通过对温度胁迫引发的微丝骨架组织结构的重组进行了观察发现微丝骨架的组织结构参与响应植株的温度胁迫。低温处理下微丝骨架发生重组,并有部分解聚；而高温处理下微丝骨架发生重组。
7.不同温度处理条件下,促进微丝聚合的药物phalloidin能够增加拟南芥小苗的温度耐受性,而抑制微丝聚合的药物latrunculin B降低植株的温度耐受性。这一结果说明微丝骨架参与植物对温度胁迫的响应。
8.荧光定量PCR的结果证实了非生物胁迫诱导大部分组成型表达的profilin和ADF表达,并且某些特定基因响应特定的环境胁迫,如ADF6只有在热激胁迫下表达量才明显升高。这一结果揭示了非生物胁迫影响这些基因转录水平的变化,而这种变化证明同基因响应同一非生物胁迫的方式不同,而不同非生物胁迫下同一基因的响应方式也不同。
综上所述,本研究发现PRF3在拟南芥幼苗期的生长发育发挥重要的生理功能,对根,根毛,下胚轴的生长有负调节作用,过表达PRF3引起的矮化表型可能是由微丝骨架重组造成的,特别是纵向微丝束的减少甚至消失。体外实验也证明了PRF3的N端特异的36个氨基酸可能与PRF3的生理功能密切相关。此外,本研究证明了微丝骨架参与植物抵御温度胁迫,同时荧光定量PCR的结果为研究ABPs在非生物胁迫中的功能提供了详实的理论依据。本研究为进一步研究植物微丝骨架及其结合蛋白的体内生理功能奠定了理论基础,为揭示植物响应非生物胁迫的机制研究提供了科学依据。
The actin cytoskeleton has emerged as a very pivotal component of eukaryotic cells, which presents a highly dynamic three-dimensional mesh networks. The actin cytoskeleton is not only offers internal mechanical support for cells but also participates in a range of important cellular processes. The function of actin cytoskeleton relies on both actin cytoskeleton own organization and actin dynamics which is regulated by multiple actin binding proteins (ABPs) and many different signals. In plant cells, a large number of ABPs have been identified and involved in different physiological functions. Profilin (PRFs) is a small monomer actin sequestering protein family in all eukaryotes, which is found earlier, highly conserved and widely expressed. The Arabidopsis PRF family contains five highly conservate isoforms. The biochemical functions of this family have been studied for a long time, the physiological function are unclear. PRF3, one of the vegetative isoforms, has36more N-terminal amino acid residues than other PRFs. The physiological and biochemical functions of PRF3are completely unknown. In this study, for the first time, we systematically studied the tissue localization of PRF3, and its physiological function, etc. The major results are as follows:
1. PRF3s a vegetative profilin gene. GUS activities and real time PCR analysis were detected in the most parts of the plants demonstrated that PRF3was strongly expressed in young seedlings, rosette leaves, and cauline leaves but weakly expressed in14-d-old seedlings and flowers and that its expression was enhanced by the first intron of PRF3.
2. PRF3increased the critical concentration (Cc) of actin assembly in vitro. PRF3-AN36increased the Cc of actin assembly, which was consistent with PRF1and PRF2in Arabidopsis. The Cc of N36was closer to actin-only controls, and the Cc of PRF3was between them. This result may indicate that the N36of PRF3protein has a role for its functions.
3. Overexpression of the full-length PRF3cDNA decreased the lengths of roots and hypocotyls and delayed germination, but PRF3-ΔN36transgenic plants and pr/3-1mutants were not, compared with wild-type plants. These results showed that overexpression of the full-length PRF3cDNA affected the lengths of roots and hypocotyls, and PRF3was a key regulator of root and hypocotyl lengths.
4. Using light microscopy and scanning electron microscopy, we found that the cell lengths of OE-PRF3-3hypocotyls were much shorter than those in the other three plant lines and that the dwarf phenotype of the OE-PRF3seedlings can be largely accounted for by the reduction in cell length. The results implated that PRF3was important for the elongation of the hypocotyl cells.
5. Confocal observation showed that thick actin cables were absent and fine actin filaments appeared less affected in the hypocotyl cells of the OE-PRF3-3seedlings. These results showed that overexpression of PRF3induce F-actin arrangement in hypocotyl cells and that thick actin cables might be associated with cell elongation.
6. We observed that reorganization of the actin cytoskeleton was triggered by cold or heat stress. The actin cytoskeletonal organization responds to temperature stresses. The MFs were considerably rearranged, and some MFs were depolymerized under cold stress; the applied heat shock stress led to changes in the MF arrays in Arabidopsis hypocotyl cells.
7. Facilitation of microfilament (MF) assembly by phalloidin treatment increased the resistance of Arabidopsis seedlings to temperature stress, whereas blocking MF assembly with latrunculin B decreased their temperature tolerance.
8. Using real-time PCR, we found that the expression patterns of some of the vegetative profilins and actin depolymerizing factors (ADFs) were specifically altered under various stresses, such as ADF6. These results showed that the expression profiles of different genes were different under the same stress, and different expression patterns for the same gene were also observed.
Thus, our results show that PRF3is vital for the development of Arabidopsis seedlings., especially the growth of roots, root hairs, and hypocotyls. The dwarf phenotype of the PRF3-overexpression seedlings can be largely accounted for by the reduction in cell length and F-actin arrangement, especially the disappearance of thick, longitudinally oriented cables. The in vivo results may indicate that the N36of PRF3protein has a role for its functions. Our results show that the actin cytoskeleton is vital for the tolerance of Arabidopsis seedlings to heat/cold stresses. The results of real time PCR will be valuable for studying the functions of ABPs in abiotic stresses and uncovering new components of stress signal transduction. In this study, we provide a theoretical foundation for studying the physiological function of the plant actin cytoskeleton and its binding proteins, and exploring the mechanism of epigenetic regulation in plant abiotic stresses tolerance.
1.PRF3是一类组成型表达的profilin异型体。通过GUS活性分析和荧光定量PCR的方法证实PRF3主要在幼苗,莲座叶和茎生叶中表达,在14天的小苗和花中的表达量很少,并且该基因的内含子可能增强基因的表达。这一结果揭示了PRF3的组织表达特征。
2.体外生化功能表明PRF3增加actin聚合的临界浓度,并且PRF3-ΔN36的Cc值最大,对actin聚合的影响最明显,而PRF3的Cc值介于PRF3-ΔN36和actin之间。这说明了PRF3的N端36个氨基酸与其蛋白功能有密切的关系。
3.与野生型拟南芥相比,过表达全长PRF3的cDNA序列的转基因植株缩短小苗的根长,下胚轴和根毛的长度,延迟种子的萌发,但是PRF3-ΔN36转基因植株和pf3突变体没有明显的表型。这一结果说明PRF3是根,下胚轴和根毛生长的重要的负调节因子,并且只有当基因完整时才具有这种功能。
4.光学显微镜和扫描电镜的观察结果显示OE-PRF3植株的下胚轴细胞的长度明显较短,细胞伸长受到抑制,说明PRF3与下胚轴细胞的伸长有着密切的关系。
5.激光共聚焦扫描显微镜观察发现OE-PRF3过表达植株的下胚轴细胞中粗的微丝束消失,但是细的微丝依然存在没有明显的变化,说明过表达PRF3全长基因能够引发微丝发束解聚,造成F-actin的重组,而粗的微丝束结构可能与细胞伸长密切相关。
6.通过对温度胁迫引发的微丝骨架组织结构的重组进行了观察发现微丝骨架的组织结构参与响应植株的温度胁迫。低温处理下微丝骨架发生重组,并有部分解聚；而高温处理下微丝骨架发生重组。
7.不同温度处理条件下,促进微丝聚合的药物phalloidin能够增加拟南芥小苗的温度耐受性,而抑制微丝聚合的药物latrunculin B降低植株的温度耐受性。这一结果说明微丝骨架参与植物对温度胁迫的响应。
8.荧光定量PCR的结果证实了非生物胁迫诱导大部分组成型表达的profilin和ADF表达,并且某些特定基因响应特定的环境胁迫,如ADF6只有在热激胁迫下表达量才明显升高。这一结果揭示了非生物胁迫影响这些基因转录水平的变化,而这种变化证明同基因响应同一非生物胁迫的方式不同,而不同非生物胁迫下同一基因的响应方式也不同。
综上所述,本研究发现PRF3在拟南芥幼苗期的生长发育发挥重要的生理功能,对根,根毛,下胚轴的生长有负调节作用,过表达PRF3引起的矮化表型可能是由微丝骨架重组造成的,特别是纵向微丝束的减少甚至消失。体外实验也证明了PRF3的N端特异的36个氨基酸可能与PRF3的生理功能密切相关。此外,本研究证明了微丝骨架参与植物抵御温度胁迫,同时荧光定量PCR的结果为研究ABPs在非生物胁迫中的功能提供了详实的理论依据。本研究为进一步研究植物微丝骨架及其结合蛋白的体内生理功能奠定了理论基础,为揭示植物响应非生物胁迫的机制研究提供了科学依据。
The actin cytoskeleton has emerged as a very pivotal component of eukaryotic cells, which presents a highly dynamic three-dimensional mesh networks. The actin cytoskeleton is not only offers internal mechanical support for cells but also participates in a range of important cellular processes. The function of actin cytoskeleton relies on both actin cytoskeleton own organization and actin dynamics which is regulated by multiple actin binding proteins (ABPs) and many different signals. In plant cells, a large number of ABPs have been identified and involved in different physiological functions. Profilin (PRFs) is a small monomer actin sequestering protein family in all eukaryotes, which is found earlier, highly conserved and widely expressed. The Arabidopsis PRF family contains five highly conservate isoforms. The biochemical functions of this family have been studied for a long time, the physiological function are unclear. PRF3, one of the vegetative isoforms, has36more N-terminal amino acid residues than other PRFs. The physiological and biochemical functions of PRF3are completely unknown. In this study, for the first time, we systematically studied the tissue localization of PRF3, and its physiological function, etc. The major results are as follows:
1. PRF3s a vegetative profilin gene. GUS activities and real time PCR analysis were detected in the most parts of the plants demonstrated that PRF3was strongly expressed in young seedlings, rosette leaves, and cauline leaves but weakly expressed in14-d-old seedlings and flowers and that its expression was enhanced by the first intron of PRF3.
2. PRF3increased the critical concentration (Cc) of actin assembly in vitro. PRF3-AN36increased the Cc of actin assembly, which was consistent with PRF1and PRF2in Arabidopsis. The Cc of N36was closer to actin-only controls, and the Cc of PRF3was between them. This result may indicate that the N36of PRF3protein has a role for its functions.
3. Overexpression of the full-length PRF3cDNA decreased the lengths of roots and hypocotyls and delayed germination, but PRF3-ΔN36transgenic plants and pr/3-1mutants were not, compared with wild-type plants. These results showed that overexpression of the full-length PRF3cDNA affected the lengths of roots and hypocotyls, and PRF3was a key regulator of root and hypocotyl lengths.
4. Using light microscopy and scanning electron microscopy, we found that the cell lengths of OE-PRF3-3hypocotyls were much shorter than those in the other three plant lines and that the dwarf phenotype of the OE-PRF3seedlings can be largely accounted for by the reduction in cell length. The results implated that PRF3was important for the elongation of the hypocotyl cells.
5. Confocal observation showed that thick actin cables were absent and fine actin filaments appeared less affected in the hypocotyl cells of the OE-PRF3-3seedlings. These results showed that overexpression of PRF3induce F-actin arrangement in hypocotyl cells and that thick actin cables might be associated with cell elongation.
6. We observed that reorganization of the actin cytoskeleton was triggered by cold or heat stress. The actin cytoskeletonal organization responds to temperature stresses. The MFs were considerably rearranged, and some MFs were depolymerized under cold stress; the applied heat shock stress led to changes in the MF arrays in Arabidopsis hypocotyl cells.
7. Facilitation of microfilament (MF) assembly by phalloidin treatment increased the resistance of Arabidopsis seedlings to temperature stress, whereas blocking MF assembly with latrunculin B decreased their temperature tolerance.
8. Using real-time PCR, we found that the expression patterns of some of the vegetative profilins and actin depolymerizing factors (ADFs) were specifically altered under various stresses, such as ADF6. These results showed that the expression profiles of different genes were different under the same stress, and different expression patterns for the same gene were also observed.
Thus, our results show that PRF3is vital for the development of Arabidopsis seedlings., especially the growth of roots, root hairs, and hypocotyls. The dwarf phenotype of the PRF3-overexpression seedlings can be largely accounted for by the reduction in cell length and F-actin arrangement, especially the disappearance of thick, longitudinally oriented cables. The in vivo results may indicate that the N36of PRF3protein has a role for its functions. Our results show that the actin cytoskeleton is vital for the tolerance of Arabidopsis seedlings to heat/cold stresses. The results of real time PCR will be valuable for studying the functions of ABPs in abiotic stresses and uncovering new components of stress signal transduction. In this study, we provide a theoretical foundation for studying the physiological function of the plant actin cytoskeleton and its binding proteins, and exploring the mechanism of epigenetic regulation in plant abiotic stresses tolerance.
引文
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