White J G
MRC Laboratory of Molecular Biology, Cambridge, England.
Ann N Y Acad Sci. 1990;582:50-9. doi: 10.1111/j.1749-6632.1990.tb21667.x.
Animal cells cleave by the progressive constriction of an equatorial region. The plane of the constricting equator is orthogonal to the mitotic spindle axis and is positioned so that the spindle is bisected, thus ensuring equipartitioning of the chromosomes. Constriction is achieved by the active contraction of a circumferential band of actomyosin-containing filaments. The mitotic apparatus is necessary for the formation of the contractile ring and specifies its position and orientation; however, it takes no active part in furrowing. One of the central questions that has to be answered to understand cytokinesis is: how does the mitotic apparatus act to organize the contractile ring? It has been suggested that the mitotic apparatus acts to modulate locally the force generated by the cortical contractile filaments. The resultant gradients of cortical tension cause filaments to be pulled into regions of higher tension. The geometry of the stimulus from the mitotic apparatus is such that the highest levels of tension occur in the equatorial regions. The geometric distortions that take place when contractile filaments are pulled into this region cause them to become partially oriented circumferentially. When furrowing commences, mechanical forces act to align the filaments further and concentrate them in a narrow band. The furrow therefore becomes self-sharpening. Similar lateral flows of contractile filaments may occur during cell locomotion and growth cone extension.
动物细胞通过赤道区域的逐渐收缩而分裂。收缩赤道面与有丝分裂纺锤体轴正交,其位置能使纺锤体一分为二,从而确保染色体的均等分配。收缩是由含肌动球蛋白丝的圆周带的主动收缩实现的。有丝分裂器对于收缩环的形成是必需的,并确定其位置和方向;然而,它在沟裂过程中并不发挥主动作用。要理解胞质分裂必须回答的核心问题之一是:有丝分裂器如何作用来组织收缩环?有人提出,有丝分裂器作用是局部调节皮质收缩丝产生的力。由此产生的皮质张力梯度会使丝被拉向张力更高的区域。来自有丝分裂器的刺激的几何形状使得赤道区域出现最高水平的张力。当收缩丝被拉到这个区域时发生的几何变形会使它们部分地沿圆周方向排列。当沟裂开始时,机械力会使丝进一步排列并将它们集中在一条窄带中。因此,沟裂会自我变窄。在细胞运动和生长锥延伸过程中可能会发生类似的收缩丝横向流动。