拟南芥微管去稳定蛋白MDP25调控花粉管生长机制的研究
摘要
花粉萌发和花粉管生长是植物有性生殖的重要环节。花粉粒在柱头上萌发并产生花粉管,花粉管沿着花柱快速生长到达珠孔,而后花粉管停止生长并破裂,释放精子进入胚珠完成双受精。因此,从正、负两个方面调控花粉管的生长有着重要的生理学意义。微丝骨架在花粉管生长过程中起着重要的作用,其组织排列及动态变化受到很多信号分子(如Ca2+,pH值等)的调控,但具体的机制仍需要进一步的研究。
本文以拟南芥花粉管为研究对象,对花粉管中微丝骨架的组织及动态进行研究,探讨了具有Ca2+结合活性的微管去稳定蛋白MICROTUBULE-DESTABILIZING PROTEIN25(MDP25)在抑制花粉管生长过程中的作用及调控花粉管内微丝骨架的分子机制。
Genevestigator (http://www.genevestigator.ethz.ch)数据库中基因芯片数据显示MDP25在花粉中有表达；GUS染色结果也表明MDP25在花粉粒和花粉管中均有表达。对花粉体外的萌发和生长进行分析及苯胺蓝染色实验均发现MDP25功能缺失突变体(mdp25-1)的花粉管的生长速率比野生型的高。
在本研究中,首先利用原核表达系统,从大肠杆菌中纯化出具有生物学活性的MDP25的融合蛋白。共沉淀分析结果表明MDP25可以在体外直接结合微丝。体外荧光观察和TIRFM实验结果表明MDP25可以切割微丝,而且MDP25切割微丝的功能可能不依赖于Ca2+,但能被Ca2+增强。
进一步对MDP25亚细胞定位进行分析,发现MDP25在花粉管内亚细胞定位受Ca2+调控,花粉管顶端和亚顶端区域中较高浓度的Ca2+使MDP25从细胞膜上脱离下来进入了细胞质中。对花粉管内微丝进行分析发现,与野生型花粉管内的微丝相比,mdp25-1花粉管亚顶端的微丝束更大,密度更高；实时动态观察进一步表明mdp25-1花粉管亚顶端微丝的切割频率明显降低。
此外,体外生化分析结果表明MDP25氨基酸序列中的微管解聚位点突变蛋白(MDP25K7A和MDP25K18A)对微丝的结合能力和切割活性没有明显的变化；而微丝切割位点突变蛋白(MDP25N5A)的微丝结合能力及切割活性显著降低,但其对微管的去稳定能力没有明显变化。遗传学分析表明PMDP25:MDP25,PMDP25:MDP25K7A和PMDP25:MDP25K18A能将mdp25-1花粉管的表型恢复到野生型水平,而PMDP25:MDP25N5A却不能互(?)mdp25-1花粉管的表型,说明MDP25对花粉管生长的影响是通过调控微丝骨架而不是通过调控微管骨架实现的。
综上所述,本论文的研究结果表明MDP25是一个新的受Ca2+调控的微丝结合蛋白,MDP25通过切割微丝调控花粉管亚顶端微丝的组织和动态,进而影响花粉管的生长。
The germination and growth of pollen tube play a crucial role in plant gamogenesis. Pollen grains germinate and produce pollen tubes on the stigma of the pistil, which grow through the style and are guided to the micropylar. Then the growth of pollen tubes is arrested and the tube tips rupture to release the sperm cells. Therefore, the positive and negative regulation of pollen tube growth have important physiological significances. Actin filaments play critical roles in regulation of pollen tube growth. The organization and dynamics of actin fimaments are regulated by many developmental signals and environmental cues. However, the regulation mechanisms of actin filament organization and dynamics are poorly understood.
In this study, we investigated the actin filament orgnization and dynamics in Arabidopsis pollen tubes and the role of a calcium-binding and microtubule-destabilizing protein (MDP25) in negative regulation of pollen tube growth by modulating the actin cytoskeleton in the subapical region.
The data from the Affymetrix AG and ATH1GeneChip arrays in the Genevestigator database (http://www.genevestigator.ethz.ch) indicate that MDP25is expressed in pollen. GUS staining aslo revealed that MDP25was expressed in the pollen and pollen tubes. Statistical analysis indicated that the growth rate of mdp25-1pollen tubes was significantly higher than that of wild-type. Furthermore, aniline blue staining showed that mdp25-1pollen tube grow faster than wild-type pollen tube.
Recombinant MDP25was expressed and purified from Escherichia coli. Cosedimentation assay showed that recombinant MDP25directly binds F-actin in vitro. Confocal observation and TIRFM assay revealed that MDP25is capable of severing actin filaments and the severing activity of MDP25is significantly enhanced in the presence of Ca2+in vitro.
Sub-cellular localization assay showed that MDP25-GFP was mainly localized in the plasma membrane at the shank region of the pollen tube, but was absent at the membrane of the subapical and apex regions. A fluorescence microscopy assay showed that the mutated MDP25protein (Ca2+-binding site) is predominantly localized to the plasma membrane along the whole length of the pollen tube. The F-actin-severing frequency was significantly decreased and high density of actin filaments was observed in the subapical region of pollen tubes from the mdp25-1mutant.
MDP25K7A and MDP25K18A exhibited similar actin filament binding and severing activity as wild-type MDP25. MDP25N5A maintained normal function on microtubules, however lost its actin filament binding and severing activity. Genetic evidence demonstrates that MDP25regulates pollen tube elongation by regulating actin filaments, but not by its microtubule-destabilizing activity.
Our study suggests that Arabidopsis MDP25is a novel actin binding protein (ABP) regulated by Ca2+and MDP25is involved in negative regulation of pollen tube growth by modulating the actin cytoskeleton in the subapical region via its F-actin-severing activity.
本文以拟南芥花粉管为研究对象,对花粉管中微丝骨架的组织及动态进行研究,探讨了具有Ca2+结合活性的微管去稳定蛋白MICROTUBULE-DESTABILIZING PROTEIN25(MDP25)在抑制花粉管生长过程中的作用及调控花粉管内微丝骨架的分子机制。
Genevestigator (http://www.genevestigator.ethz.ch)数据库中基因芯片数据显示MDP25在花粉中有表达；GUS染色结果也表明MDP25在花粉粒和花粉管中均有表达。对花粉体外的萌发和生长进行分析及苯胺蓝染色实验均发现MDP25功能缺失突变体(mdp25-1)的花粉管的生长速率比野生型的高。
在本研究中,首先利用原核表达系统,从大肠杆菌中纯化出具有生物学活性的MDP25的融合蛋白。共沉淀分析结果表明MDP25可以在体外直接结合微丝。体外荧光观察和TIRFM实验结果表明MDP25可以切割微丝,而且MDP25切割微丝的功能可能不依赖于Ca2+,但能被Ca2+增强。
进一步对MDP25亚细胞定位进行分析,发现MDP25在花粉管内亚细胞定位受Ca2+调控,花粉管顶端和亚顶端区域中较高浓度的Ca2+使MDP25从细胞膜上脱离下来进入了细胞质中。对花粉管内微丝进行分析发现,与野生型花粉管内的微丝相比,mdp25-1花粉管亚顶端的微丝束更大,密度更高；实时动态观察进一步表明mdp25-1花粉管亚顶端微丝的切割频率明显降低。
此外,体外生化分析结果表明MDP25氨基酸序列中的微管解聚位点突变蛋白(MDP25K7A和MDP25K18A)对微丝的结合能力和切割活性没有明显的变化；而微丝切割位点突变蛋白(MDP25N5A)的微丝结合能力及切割活性显著降低,但其对微管的去稳定能力没有明显变化。遗传学分析表明PMDP25:MDP25,PMDP25:MDP25K7A和PMDP25:MDP25K18A能将mdp25-1花粉管的表型恢复到野生型水平,而PMDP25:MDP25N5A却不能互(?)mdp25-1花粉管的表型,说明MDP25对花粉管生长的影响是通过调控微丝骨架而不是通过调控微管骨架实现的。
综上所述,本论文的研究结果表明MDP25是一个新的受Ca2+调控的微丝结合蛋白,MDP25通过切割微丝调控花粉管亚顶端微丝的组织和动态,进而影响花粉管的生长。
The germination and growth of pollen tube play a crucial role in plant gamogenesis. Pollen grains germinate and produce pollen tubes on the stigma of the pistil, which grow through the style and are guided to the micropylar. Then the growth of pollen tubes is arrested and the tube tips rupture to release the sperm cells. Therefore, the positive and negative regulation of pollen tube growth have important physiological significances. Actin filaments play critical roles in regulation of pollen tube growth. The organization and dynamics of actin fimaments are regulated by many developmental signals and environmental cues. However, the regulation mechanisms of actin filament organization and dynamics are poorly understood.
In this study, we investigated the actin filament orgnization and dynamics in Arabidopsis pollen tubes and the role of a calcium-binding and microtubule-destabilizing protein (MDP25) in negative regulation of pollen tube growth by modulating the actin cytoskeleton in the subapical region.
The data from the Affymetrix AG and ATH1GeneChip arrays in the Genevestigator database (http://www.genevestigator.ethz.ch) indicate that MDP25is expressed in pollen. GUS staining aslo revealed that MDP25was expressed in the pollen and pollen tubes. Statistical analysis indicated that the growth rate of mdp25-1pollen tubes was significantly higher than that of wild-type. Furthermore, aniline blue staining showed that mdp25-1pollen tube grow faster than wild-type pollen tube.
Recombinant MDP25was expressed and purified from Escherichia coli. Cosedimentation assay showed that recombinant MDP25directly binds F-actin in vitro. Confocal observation and TIRFM assay revealed that MDP25is capable of severing actin filaments and the severing activity of MDP25is significantly enhanced in the presence of Ca2+in vitro.
Sub-cellular localization assay showed that MDP25-GFP was mainly localized in the plasma membrane at the shank region of the pollen tube, but was absent at the membrane of the subapical and apex regions. A fluorescence microscopy assay showed that the mutated MDP25protein (Ca2+-binding site) is predominantly localized to the plasma membrane along the whole length of the pollen tube. The F-actin-severing frequency was significantly decreased and high density of actin filaments was observed in the subapical region of pollen tubes from the mdp25-1mutant.
MDP25K7A and MDP25K18A exhibited similar actin filament binding and severing activity as wild-type MDP25. MDP25N5A maintained normal function on microtubules, however lost its actin filament binding and severing activity. Genetic evidence demonstrates that MDP25regulates pollen tube elongation by regulating actin filaments, but not by its microtubule-destabilizing activity.
Our study suggests that Arabidopsis MDP25is a novel actin binding protein (ABP) regulated by Ca2+and MDP25is involved in negative regulation of pollen tube growth by modulating the actin cytoskeleton in the subapical region via its F-actin-severing activity.
引文
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