Johnson P T, Williams R R, Cusato K, Reese B E
Neuroscience Research Institute, University of California at Santa Barbara, Santa Barbara, California 93106-5060, USA.
J Comp Neurol. 1999 Nov 8;414(1):1-12.
Mature rod and cone photoreceptor cells extend terminals to the outer plexiform layer (OPL), where they form characteristic spherules or pedicles, synapsing with the second-order neurons of the inner nuclear layer (INL). The present study demonstrates that, prior to the formation of this connectivity, immature rods and cones in the ferret extend processes beyond the level of the horizontal cells and future OPL, reaching the inner plexiform layer (IPL). The number of processes extending to the IPL increases steadily as the population of photoreceptor cells expands postnatally, reaching a maximum 2 weeks after birth. These processes are immunopositive for synaptophysin, and they terminate in two strata occupied by the dendrites of amacrine cells and ganglion cells. The frequency of these processes declines rapidly during the third postnatal week, and they are no longer detectable by the fourth postnatal week. Their loss is neither a consequence of photoreceptor cell death nor is it due to selective protein trafficking mechanisms that render them immunonegative. Rather, these processes retract to the level of the OPL during this period, coincident with the maturation of bipolar and horizontal cell processes. These results demonstrate that, despite the clear presence of environmental signals presaging the formation of the OPL, photoreceptor terminals initially ignore them to grow beyond this level of the retina. Rather, they detect and respond to signals within the IPL during this period, terminating in proximity to the processes of other cells in the inner retina, where they may contribute to transient retinal circuitry during early development.
成熟的视杆和视锥光感受器细胞将其终末延伸至外网状层(OPL),在那里它们形成特征性的小球或足突,并与内核层(INL)的二级神经元形成突触。本研究表明,在这种连接形成之前,雪貂的未成熟视杆和视锥细胞会将突起延伸至水平细胞和未来OPL的水平之上,到达内网状层(IPL)。随着出生后光感受器细胞数量的增加,延伸至IPL的突起数量稳步增加,在出生后2周达到最大值。这些突起对突触素呈免疫阳性,并且它们终止于由无长突细胞和神经节细胞的树突所占据的两个层。这些突起的频率在出生后第三周迅速下降,到出生后第四周就不再能检测到。它们的消失既不是光感受器细胞死亡的结果,也不是由于使它们呈免疫阴性的选择性蛋白质运输机制所致。相反,在这段时间内,这些突起回缩至OPL的水平,这与双极细胞和水平细胞突起的成熟相一致。这些结果表明,尽管明显存在预示OPL形成的环境信号,但光感受器终末最初忽略了它们,生长至视网膜的该水平之上。相反,在此期间它们检测并响应IPL内的信号,终止于与视网膜内层其他细胞的突起相邻处,在早期发育过程中它们可能有助于形成短暂的视网膜回路。