Tilney Lewis G, Connelly Patricia S, Ruggiero Linda, Vranich Kelly A, Guild Gregory M, Derosier David
Department of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018, USA.
Mol Biol Cell. 2004 Dec;15(12):5481-91. doi: 10.1091/mbc.e04-06-0472. Epub 2004 Sep 15.
Drosophila bristles display a precise orientation and curvature. An asymmetric extension of the socket cell overlies the newly emerging bristle rudiment to provide direction for bristle elongation, a process thought to be orchestrated by the nerve dendrite lying between these cells. Scanning electron microscopic analysis of individual bristles showed that curvature is planar and far greater near the bristle base. Correlated with this, as development proceeds the pupa gradually recedes from the inner pupal case (an extracellular layer that encloses the pupa) leading to less bristle curvature along the shaft. We propose that the inner pupal case induces elongating bristles to bend when they contact this barrier. During elongation the actin cytoskeleton locks in this curvature by grafting together the overlapping modules that comprise the long filament bundles. Because the bristle is curved, the actin bundles on the superior side must be longer than those on the inferior side. This is accomplished during grafting by greater elongation of superior side modules. Poor actin cross-bridging in mutant bristles results in altered curvature. Thus, the pattern of bristle curvature is a product of both extrinsic factors-the socket cell and the inner pupal case--and intrinsic factors--actin cytoskeleton assembly.
果蝇刚毛呈现出精确的方向和曲率。套细胞的不对称延伸覆盖新出现的刚毛原基,为刚毛伸长提供方向,这一过程被认为是由位于这些细胞之间的神经树突协调的。对单个刚毛的扫描电子显微镜分析表明,曲率是平面的,且在刚毛基部附近更大。与此相关的是,随着发育进行,蛹逐渐从内蛹壳(包围蛹的细胞外层)后退,导致沿刚毛轴的曲率减小。我们提出,内蛹壳在伸长的刚毛接触到这个屏障时会诱导其弯曲。在伸长过程中,肌动蛋白细胞骨架通过将构成长丝束的重叠模块嫁接在一起,锁定这种曲率。由于刚毛是弯曲的,上侧的肌动蛋白束必须比下侧的长。这是在嫁接过程中通过上侧模块的更大伸长来实现的。突变刚毛中肌动蛋白交叉桥接不良会导致曲率改变。因此,刚毛曲率模式是外在因素(套细胞和内蛹壳)和内在因素(肌动蛋白细胞骨架组装)共同作用的产物。