Clark P, Britland S, Connolly P
Department of Anatomy and Cell Biology, St. Mary's Hospital Medical School, Imperial College of Science, Technology and Medicine, London, UK.
J Cell Sci. 1993 May;105 ( Pt 1):203-12. doi: 10.1242/jcs.105.1.203.
Neurite growth cones detect and respond to guidance cues in their local environment that determine stereotyped pathways during development and regeneration. Micropatterns of laminin (which was found to adsorb preferentially to photolithographically defined hydrophobic areas of micropatterns) were here used to model adhesive pathways that might influence neurite extension. The responses of growth cones were determined by the degree of guidance of neurite extension and also by examining growth cone morphology. These parameters were found to be strongly dependent on the geometry of the patterned laminin, and on neuron type. Decreasing the spacing of multiple parallel tracks of laminin alternating with non-adhesive tracks, resulted in decreased guidance of chick embryo brain neurons. Single isolated 2 microns tracks strongly guided neurite extension whereas 2 microns tracks forming a 4 microns period multiple parallel pattern did not. Growth cones appear to be capable of bridging the narrow non-adhesive tracks, rendering them insensitive to the smaller period multiple parallel adhesive patterns. These observations suggest that growth cones would be unresponsive to the multiple adhesive cues such as would be presented by oriented extracellular matrix or certain axon fascicle structures, but could be guided by isolated adhesive tracks. Growth cone morphology became progressively simpler on progressively narrower single tracks. On narrow period multiple parallel tracks (which did not guide neurite extension) growth cones spanned a number of adhesive/non-adhesive tracks, and their morphology suggests that lamellipodial advance may be independent of the substratum by using filopodia as a scaffold. In addition to acting as guidance cues, laminin micropatterns also appeared to influence the production of primary neurites and their subsequent branching. On planar substrata, dorsal root ganglion neurons were multipolar, with highly branched neurite outgrowth whereas, on 25 microns tracks, neurite branching was reduced or absent, and neuron morphology was typically bipolar. These observations indicate the precision with which growth cone advance may be controlled by substrata and suggest a role for patterned adhesiveness in neuronal morphological differentiation, but also highlight some of the limitations of growth cone sensitivity to substratum cues.
神经突生长锥能够检测并响应其局部环境中的导向线索,这些线索在发育和再生过程中决定了固定的路径。层粘连蛋白的微图案(发现其优先吸附到通过光刻定义的微图案疏水区域)在此用于模拟可能影响神经突延伸的粘附路径。生长锥的反应通过神经突延伸的导向程度以及检查生长锥形态来确定。发现这些参数强烈依赖于图案化层粘连蛋白的几何形状以及神经元类型。减小层粘连蛋白的多个平行条带与非粘附条带交替排列的间距,会导致鸡胚脑神经元的导向作用减弱。单个孤立的2微米条带强烈引导神经突延伸,而形成4微米周期多个平行图案的2微米条带则不然。生长锥似乎能够跨越狭窄的非粘附条带,使其对较小周期的多个平行粘附图案不敏感。这些观察结果表明,生长锥对诸如定向细胞外基质或某些轴突束结构所呈现的多个粘附线索无反应,但可由孤立的粘附条带引导。在逐渐变窄的单条带上,生长锥形态逐渐变得更简单。在狭窄周期的多个平行条带上(这些条带不引导神经突延伸),生长锥跨越多个粘附/非粘附条带,其形态表明片状伪足的推进可能通过丝状伪足以支架的形式独立于基质。除了作为导向线索外,层粘连蛋白微图案似乎还影响初级神经突的产生及其后续分支。在平面基质上,背根神经节神经元是多极的,神经突生长高度分支,而在25微米条带上,神经突分支减少或不存在,神经元形态通常为双极。这些观察结果表明基质对生长锥前进的控制精度,并暗示图案化粘附性在神经元形态分化中的作用,但也突出了生长锥对基质线索敏感性的一些局限性。