Schmidt J, Coen T
Department of Biological Sciences, University at Albany, State University of New York 12222, USA.
J Neurobiol. 1995 Dec;28(4):409-18. doi: 10.1002/neu.480280402.
In adult goldfish, electrophysiological studies have shown that the retinotectal projection reorganizes, following removal of half of the tectum, to form a complete but compressed projection over the remaining half tectum. As a result, each fiber terminates more rostrally than normal. Electron microscopic studies suggest a competition between retinal fibers for a fixed number of synaptic sites. The current study examines whether retinal arbors in the compressed projection are smaller than normal in extent or branching and whether the fiber paths in the tectum show the rostral movements and the search strategy that the retinal fibers use. The caudal half tectum was removed without cutting retinal fibers except those at the cut edge. At 3 to 19 months afterward, retinal fibers were labeled with horseradish peroxidase. In whole-mounted tecta, fibers and terminals were drawn under camera lucida and compared with normal arbors. The axonal paths were also traced across the tectum to their termination sites. At 3 to 6 months (early stages of compression), the arbors were rather normal in appearance, although they were actually significantly larger (23%) than normal in linear extent, arborized somewhat deeper and had fewer branches (18%). The fibers normally terminating in the rostral tectum followed normal stereotyped paths, whereas those cut at the edge had grown back and forth loops (apparent searching behavior) with little branching. By 10 months when compression is complete, arbors were significantly smaller than normal (19%), were arborizing significantly deeper, and had significantly fewer branches (19%). The differences were more pronounced in arbors of coarse and medium caliber than in fine caliber axons. The axons still ran in stereotyped fascicles, but included an extrafascicular portion that, unlike any axons in normals, turned back in a rostral direction before branching. This striking effect, present even in far rostral tectum, indicated that arbors had been forced to move rostrally to accomodate those from the ablated half. The small effect on arbor extent suggests that this is influenced by factors other than the magnification factor of the map, perhaps postsynaptic dendritic extent. The increased depth of termination is consistent with the increased thickness of the retinal terminal layer. The decreased number of branches is consistent with the conclusion that the remaining fixed number of synaptic sites shared among the full complement of retinal fibers should result in fewer synapses per retinal fiber.
在成年金鱼中,电生理研究表明,在切除一半视顶盖后,视网膜-视顶盖投射会重新组织,在剩余的一半视顶盖上形成一个完整但压缩的投射。结果,每条纤维的终止位置比正常情况更靠前。电子显微镜研究表明,视网膜纤维之间存在对固定数量突触位点的竞争。本研究考察了压缩投射中的视网膜分支在范围或分支数量上是否小于正常情况,以及视顶盖中的纤维路径是否显示出视网膜纤维所采用的向前移动和搜索策略。切除了视顶盖的后半部分,除了在切口边缘处的视网膜纤维外,未切断其他视网膜纤维。在术后3至19个月,用辣根过氧化物酶标记视网膜纤维。在整个视顶盖上,在显微镜下绘制纤维和终末,并与正常分支进行比较。还追踪了轴突在视顶盖上的路径直至其终止位点。在3至6个月(压缩早期阶段),分支的外观相当正常,尽管它们的线性范围实际上比正常情况显著更大(23%),分支更深一些且分支数量更少(18%)。通常终止于视顶盖前部的纤维遵循正常的固定路径,而在边缘处被切断的纤维则来回生长出环状结构(明显的搜索行为)且分支很少。到10个月压缩完成时,分支明显小于正常情况(19%),分支深度显著增加,且分支数量显著减少(19%)。粗口径和中等口径轴突的分支差异比细口径轴突更明显。轴突仍沿固定的束状结构走行,但包括一个束外部分,与正常情况下的任何轴突不同,该部分在分支前向前折返。这种显著的效应,即使在视顶盖前部很远的地方也存在,表明分支被迫向前移动以容纳来自被切除一半的纤维。对分支范围的微小影响表明,这受到除图谱放大因子之外的其他因素影响,可能是突触后树突的范围。终止深度的增加与视网膜终末层厚度的增加一致。分支数量的减少与以下结论一致,即在完整的视网膜纤维中共享的固定数量的突触位点应导致每条视网膜纤维的突触减少。