肌动蛋白网架体系在粘附培养哺乳动物细胞胞质分裂变形中的作用
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摘要
胞质分裂中的细胞变形是一个由多种因素共同调节,复杂而又井然有序的力学变形过程。细胞的形态改变需要在时间和空间上调节多种生物大分子。细胞肌动蛋白网架体系做为细胞的主要支架,是驱动细胞变形的主要因素,研究胞质分裂过程中肌动蛋白网架体系在细胞变形中的作用,不仅可以在细胞形态变化-骨架装配变化-细胞力学特性变化之间建立起力学-生物学联系,进一步了解胞质分裂的主动变形及调控方式,还能够为与胞质分裂相关的疾病提供理论支持,为组织、器官再生提供细胞力学依据。
本文以贴壁生长的正常大鼠肾上皮细胞(NRK)为研究对象,采用微管吸吮、局部生化试剂刺激、显微图像动态采集、分析和荧光染色等生物力学和生物学实验技术,研究了肌动蛋白网架体系在胞质分裂赤道板内陷发生前后的不同时期,细胞整体皮层和子细胞极区皮层的不同区域内对胞质分裂变形的作用。以细胞骨架对细胞力学特性的贡献为依据,对细胞骨架蛋白异常和相应细胞力学特性改变引起的胞质分裂变形进程缺陷、细胞形态改变、子细胞变形能力改变和分裂沟细胞间桥动态变形做出了相关研究和分析,探索哺乳动物细胞骨架在胞质分裂变形的作用机理。主要工作内容和结论如下：
1.对正常和细胞骨架抑制剂作用下的NRK细胞的力学特性进行了测量和比较。实验中采用抑制肌动蛋白聚合的细胞松弛素D、抑制微管蛋白聚合的秋水仙素和肌球蛋白Ⅱ的ATP酶活性抑制剂blebbistatin分别处理细胞,应用微管吸吮技术和相关力学模型测定了不同处理组细胞的弹性模量和表面张力,并且分析了NRK细胞在准静态吸入负压作用下的变形。发现肌动蛋白对细胞的力学特性贡献最为明显,肌球蛋白Ⅱ的马达活性对细胞力学特性的贡献具有独立性,并且肌球蛋白Ⅱ与肌动蛋白的作用强弱是细胞力学特性的重要调节因素；
2.通过生化抑制剂在胞质分裂变形的不同时期和区域作用,分析了胞质分裂过程中不同骨架蛋白对变形进程的影响。
在分裂沟内陷发生前后两个不同时期,向细胞整体或单侧极区皮层施加细胞骨架蛋白抑制剂,分析肌动蛋白和与之作用的微管蛋白、肌球蛋白Ⅱ在细胞变形中的作用。实验表明不同区域的细胞骨架蛋白在不同变形阶段对胞质分裂变形进程影响有显著差异,并且肌动蛋白和肌球蛋白Ⅱ的相互作用可以调节肌动蛋白抑制剂对胞质分裂变形进程的抑制作用。另一方面,发现子细胞的高变形能力是肌动蛋白纤维与肌球蛋白Ⅱ的马达活性共同调节的,并且低水平的肌球蛋白Ⅱ马达活性有利于细胞变形；
3.对单侧极区皮层骨架蛋白抑制剂作用下的细胞进行胞质分裂间桥变形和子细胞变形的分析。
结合细胞间桥变细形态学参数及细胞表面张力的变化,对比分析了细胞间桥变形曲线,结果显示极区皮层肌动蛋白网络决定着细胞间桥变形的趋势,单侧子细胞的表面张力控制着细胞间桥变细的速度。在高变形能力子细胞的形态分析中发现,细胞间桥变形中D0.5的点可以描述子细胞变形的趋势,并且依据实验结果对间桥变细动力学进行改良。改良后的结果显示了在D05位置,细胞完成了对整体表面张力的调节。
由此得出在细胞胞质分裂变形过程中,肌动蛋白网架体系中的肌动蛋白是细胞力学特性最主要决定因素,肌动蛋白和肌球蛋白Ⅱ的相互作用是细胞力学特性和变形的主要调节因素。在不同变形时期,肌动蛋白和肌球蛋白Ⅱ通过调节自身装配和相互之间的作用,控制着细胞不同区域的力学特性,协调整体皮层表面张力的改变,控制着胞质分裂中的变形步骤。
Cytokinesis of animal cells is a mechanical deformation process characterized by a complex coordinated sequence of cytoskeletal changes. The ability to change cellular shape requires that the distribution of different classes of molecules be regulated both spatially and temporally. The actin filament network contributes structure and support for the cells and is the major cytoskeletal system that drives cytokinesis. Analysis the roles of actin cytoskeletal network during cytokinesis not only establishes mechanical-biological relationship among cell shape, cytoskeletal organization and cellular mechanical properties which can help us understand the process and mechanism of active changing cellular shape, but also provide theoretical evidence for revealing the mystery of disease related cytokinesis failure and provide cell mechanical mechanism for replenishment of tissues and organs.
In our study, micropipette aspiration technique, local cytoskeletal inhibitors application technique, micro-image dynamic collecting and analysis system, and immunofluorescence method were used to explore the effects of actin network in different phases and different regions during cytokinesis using normal rat kidney epithelial cells (NRK). The investigation analyzed on cells shape changes, defects of cytokinesis, deformability of daughter cells and furrow-thinning dynamics based on the cellular mechanical properpties. The main work and conclusions were as follows:
1. Determined on the cellular mechanical properties of cells. Cells treated with cytochalasin D (inhibitor of actin polymerization), colchicine (inhibitor of microtubule polymerization) and blebbistatin (inhibitor of myosinⅡATPase activity). The difference of Elastic modulus and surface tension were tested with by micropipette aspiration technique and its relative theoretical model. The results indicated that disturbance of the process of assembling actin meshwork would cause most dramatic reduction of cellular mechanical. The activity of myosinⅡATPase performed an independent mechanism to influence the mechanical parameters without changing of cytoskeleton, and the interaction between actin and myosinⅡacted as an important regulator on cellular mechanical properties.
2. Discussed the effects of cytoskeletal disturbing to the process of cell shape changes by releasing cytoskeletal inhibitors in different phases and regions during cytokinesis.
Our study released cytoskeletal inhibitors to cells in two different phases during cytokinesis delimited by furrow ingression. In addition, cytoskeltal inhibitors were applied near the polar region of dividing cells. The results showed distinct roles of cytoskeleton in different phases during cytokinesis. Our experiment found that the interaction between actin and myosinⅡcould diluted the disrupt effects to cytokinesis arousing by abnormal assembling actin. On the other hand, actin-myosinⅡinteraction determined the higher level deformability of daughter cell, and lower activity of myosinⅡATPase facilitated this process.
3. Investigating the dynamic of intercellular bridge and deformation of daughter cells by polar cortical treating with cytoskeletal inhibitors.
Our results showed that locally released CD or colchicine at the polar region dramatically influenced the trajectories of intercellular bridge thinning. Disturbing single polar cortical actin induced transformation of the intercellular bridge thinning process, and polar cortical tension controlled deformation time of intercellular bridges.
On the other hand, our study analyzed the surface area changes of higher level deformability daughter cells and their intercellular bridge shape. The results showed that bridge morphology parameters D0.5 acted as a dividing point to describe shape changes of daughter cells. Moreover, we adjusted furrow-thinning dynamical parameters based on our experiment. The results indicated that cells had regulated the balance of cortical tension before D0.5.
Our study indicated that actin network controlled each step of cell shape changes duiring cytokinesis. Actin acted as the decisive factor in cell mechanical properties and cell shape changes. The interaction between actin and myosinⅡundertook the prominent regulator relying on dynamical organization. This interaction controlled mechanical properties of different cellular regions, regulated cortical tension, manipulated the process of cell shape changes.
本文以贴壁生长的正常大鼠肾上皮细胞(NRK)为研究对象,采用微管吸吮、局部生化试剂刺激、显微图像动态采集、分析和荧光染色等生物力学和生物学实验技术,研究了肌动蛋白网架体系在胞质分裂赤道板内陷发生前后的不同时期,细胞整体皮层和子细胞极区皮层的不同区域内对胞质分裂变形的作用。以细胞骨架对细胞力学特性的贡献为依据,对细胞骨架蛋白异常和相应细胞力学特性改变引起的胞质分裂变形进程缺陷、细胞形态改变、子细胞变形能力改变和分裂沟细胞间桥动态变形做出了相关研究和分析,探索哺乳动物细胞骨架在胞质分裂变形的作用机理。主要工作内容和结论如下：
1.对正常和细胞骨架抑制剂作用下的NRK细胞的力学特性进行了测量和比较。实验中采用抑制肌动蛋白聚合的细胞松弛素D、抑制微管蛋白聚合的秋水仙素和肌球蛋白Ⅱ的ATP酶活性抑制剂blebbistatin分别处理细胞,应用微管吸吮技术和相关力学模型测定了不同处理组细胞的弹性模量和表面张力,并且分析了NRK细胞在准静态吸入负压作用下的变形。发现肌动蛋白对细胞的力学特性贡献最为明显,肌球蛋白Ⅱ的马达活性对细胞力学特性的贡献具有独立性,并且肌球蛋白Ⅱ与肌动蛋白的作用强弱是细胞力学特性的重要调节因素；
2.通过生化抑制剂在胞质分裂变形的不同时期和区域作用,分析了胞质分裂过程中不同骨架蛋白对变形进程的影响。
在分裂沟内陷发生前后两个不同时期,向细胞整体或单侧极区皮层施加细胞骨架蛋白抑制剂,分析肌动蛋白和与之作用的微管蛋白、肌球蛋白Ⅱ在细胞变形中的作用。实验表明不同区域的细胞骨架蛋白在不同变形阶段对胞质分裂变形进程影响有显著差异,并且肌动蛋白和肌球蛋白Ⅱ的相互作用可以调节肌动蛋白抑制剂对胞质分裂变形进程的抑制作用。另一方面,发现子细胞的高变形能力是肌动蛋白纤维与肌球蛋白Ⅱ的马达活性共同调节的,并且低水平的肌球蛋白Ⅱ马达活性有利于细胞变形；
3.对单侧极区皮层骨架蛋白抑制剂作用下的细胞进行胞质分裂间桥变形和子细胞变形的分析。
结合细胞间桥变细形态学参数及细胞表面张力的变化,对比分析了细胞间桥变形曲线,结果显示极区皮层肌动蛋白网络决定着细胞间桥变形的趋势,单侧子细胞的表面张力控制着细胞间桥变细的速度。在高变形能力子细胞的形态分析中发现,细胞间桥变形中D0.5的点可以描述子细胞变形的趋势,并且依据实验结果对间桥变细动力学进行改良。改良后的结果显示了在D05位置,细胞完成了对整体表面张力的调节。
由此得出在细胞胞质分裂变形过程中,肌动蛋白网架体系中的肌动蛋白是细胞力学特性最主要决定因素,肌动蛋白和肌球蛋白Ⅱ的相互作用是细胞力学特性和变形的主要调节因素。在不同变形时期,肌动蛋白和肌球蛋白Ⅱ通过调节自身装配和相互之间的作用,控制着细胞不同区域的力学特性,协调整体皮层表面张力的改变,控制着胞质分裂中的变形步骤。
Cytokinesis of animal cells is a mechanical deformation process characterized by a complex coordinated sequence of cytoskeletal changes. The ability to change cellular shape requires that the distribution of different classes of molecules be regulated both spatially and temporally. The actin filament network contributes structure and support for the cells and is the major cytoskeletal system that drives cytokinesis. Analysis the roles of actin cytoskeletal network during cytokinesis not only establishes mechanical-biological relationship among cell shape, cytoskeletal organization and cellular mechanical properties which can help us understand the process and mechanism of active changing cellular shape, but also provide theoretical evidence for revealing the mystery of disease related cytokinesis failure and provide cell mechanical mechanism for replenishment of tissues and organs.
In our study, micropipette aspiration technique, local cytoskeletal inhibitors application technique, micro-image dynamic collecting and analysis system, and immunofluorescence method were used to explore the effects of actin network in different phases and different regions during cytokinesis using normal rat kidney epithelial cells (NRK). The investigation analyzed on cells shape changes, defects of cytokinesis, deformability of daughter cells and furrow-thinning dynamics based on the cellular mechanical properpties. The main work and conclusions were as follows:
1. Determined on the cellular mechanical properties of cells. Cells treated with cytochalasin D (inhibitor of actin polymerization), colchicine (inhibitor of microtubule polymerization) and blebbistatin (inhibitor of myosinⅡATPase activity). The difference of Elastic modulus and surface tension were tested with by micropipette aspiration technique and its relative theoretical model. The results indicated that disturbance of the process of assembling actin meshwork would cause most dramatic reduction of cellular mechanical. The activity of myosinⅡATPase performed an independent mechanism to influence the mechanical parameters without changing of cytoskeleton, and the interaction between actin and myosinⅡacted as an important regulator on cellular mechanical properties.
2. Discussed the effects of cytoskeletal disturbing to the process of cell shape changes by releasing cytoskeletal inhibitors in different phases and regions during cytokinesis.
Our study released cytoskeletal inhibitors to cells in two different phases during cytokinesis delimited by furrow ingression. In addition, cytoskeltal inhibitors were applied near the polar region of dividing cells. The results showed distinct roles of cytoskeleton in different phases during cytokinesis. Our experiment found that the interaction between actin and myosinⅡcould diluted the disrupt effects to cytokinesis arousing by abnormal assembling actin. On the other hand, actin-myosinⅡinteraction determined the higher level deformability of daughter cell, and lower activity of myosinⅡATPase facilitated this process.
3. Investigating the dynamic of intercellular bridge and deformation of daughter cells by polar cortical treating with cytoskeletal inhibitors.
Our results showed that locally released CD or colchicine at the polar region dramatically influenced the trajectories of intercellular bridge thinning. Disturbing single polar cortical actin induced transformation of the intercellular bridge thinning process, and polar cortical tension controlled deformation time of intercellular bridges.
On the other hand, our study analyzed the surface area changes of higher level deformability daughter cells and their intercellular bridge shape. The results showed that bridge morphology parameters D0.5 acted as a dividing point to describe shape changes of daughter cells. Moreover, we adjusted furrow-thinning dynamical parameters based on our experiment. The results indicated that cells had regulated the balance of cortical tension before D0.5.
Our study indicated that actin network controlled each step of cell shape changes duiring cytokinesis. Actin acted as the decisive factor in cell mechanical properties and cell shape changes. The interaction between actin and myosinⅡundertook the prominent regulator relying on dynamical organization. This interaction controlled mechanical properties of different cellular regions, regulated cortical tension, manipulated the process of cell shape changes.
引文
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