Guild Gregory M, Connelly Patricia S, Ruggiero Linda, Vranich Kelly A, Tilney Lewis G
Department of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018, USA.
Mol Biol Cell. 2005 Aug;16(8):3620-31. doi: 10.1091/mbc.e05-03-0185. Epub 2005 May 25.
Actin filament bundles can shape cellular extensions into dramatically different forms. We examined cytoskeleton formation during wing hair morphogenesis using both confocal and electron microscopy. Hairs elongate with linear kinetics (approximately 1 microm/h) over the course of approximately 18 h. The resulting structure is vividly asymmetric and shaped like a rose thorn--elongated in the distal direction, curved in two dimensions with an oval base and a round tip. High-resolution analysis shows that the cytoskeleton forms from microvilli-like pimples that project actin filaments into the cytoplasm. These filaments become cross-linked into bundles by the sequential use of three cross-bridges: villin, forked and fascin. Genetic loss of each cross-bridge affects cell shape. Filament bundles associate together, with no lateral membrane attachments, into a cone of overlapping bundles that matures into an oval base by the asymmetric addition of bundles on the distal side. In contrast, the long bristle cell extension is supported by equally long (up to 400 microm) filament bundles assembled together by end-to-end grafting of shorter modules. Thus, bristle and hair cells use microvilli and cross-bridges to generate the common raw material of actin filament bundles but employ different strategies to assemble these into vastly different shapes.
肌动蛋白丝束可将细胞突起塑造成截然不同的形态。我们使用共聚焦显微镜和电子显微镜研究了翅毛形态发生过程中的细胞骨架形成。翅毛在大约18小时的过程中以线性动力学(约1微米/小时)伸长。最终形成的结构明显不对称,形状像玫瑰刺——向远端伸长,在两个维度上弯曲,基部呈椭圆形,尖端呈圆形。高分辨率分析表明,细胞骨架由类似微绒毛的小突起形成,这些小突起将肌动蛋白丝投射到细胞质中。这些丝通过依次使用三种交联桥蛋白(绒毛蛋白、叉状蛋白和肌动蛋白结合蛋白)交联成束。每种交联桥蛋白的基因缺失都会影响细胞形状。丝束相互结合,没有侧向膜附着,形成一个重叠束的锥体,通过在远端不对称地添加束而成熟为椭圆形基部。相比之下,长刚毛细胞的延伸由同样长(长达400微米)的丝束支撑,这些丝束通过较短模块的端对端嫁接组装在一起。因此,刚毛细胞和毛细胞利用微绒毛和交联桥蛋白产生肌动蛋白丝束这一共同的原材料,但采用不同的策略将它们组装成截然不同的形状。