Hitchcock P F, Easter S S
School of Medicine, Department of Ophthalmology, W.K. Kellogg Eye Center, University of Michigan, Ann Arbor 48105.
J Comp Neurol. 1987 Dec 22;266(4):556-64. doi: 10.1002/cne.902660408.
In larval frogs the retina and tectum grow in topologically dissimilar patterns: new cells are added as peripheral annuli in the retina and as caudal crescents in the tectum. Retinotopy is maintained by the continual caudalward shifting of the terminals of the optic axons. After metamorphosis the pattern of growth changes. The retina continues to add new ganglion cells peripherally, but there is no neurogenesis in the tectum. To maintain retinotopy in postmetamorphic frogs, the terminals of the optic axons must continually shift toward the central tectum. We tested the proposal of centripetally shifting axons by making punctate injections of horseradish peroxidase (HRP) in the tectum of adult Rana pipiens and observing the patterns of filled cells in the contralateral retina, as was done in the goldfish (Easter and Stuermer, '84). Punctate applications of HRP in the tectum should be taken up: 1) by fascicles, and label a partial anulus of cells, 2) by terminals, and label a cluster of cells in the corresponding retinotopic site, and 3) by the extrafascicular axonal segments, and label a band of cells connecting the partial annulus to the cluster. If the terminals have shifted centripetally, the band of cells labeled through their extrafascicular segments should have a spoke-like orientation, with the center of the retina as the hub. As the tectal site moves from rostral to caudal, this band of cells should move, pendulum-like, from temporal to nasal retina. In general, the patterns of HRP-filled retinal cells we observed were consistent with our predictions. In addition, HRP taken up by the oldest (rostral) tectal axons produced more complex patterns of filled cells that indicated that these axons had shifted both caudally before metamorphosis and centripetally after.
在幼蛙中,视网膜和视顶盖以拓扑学上不同的模式生长:新细胞在视网膜中作为周边环带添加,在视顶盖中作为尾侧新月形添加。视网膜拓扑结构通过视神经轴突终末不断向尾侧移动得以维持。变态后,生长模式发生变化。视网膜继续在周边添加新的神经节细胞,但视顶盖中不再有神经发生。为了在变态后的青蛙中维持视网膜拓扑结构,视神经轴突终末必须不断向视顶盖中央移动。我们通过在成年豹蛙的视顶盖中进行辣根过氧化物酶(HRP)的点状注射,并观察对侧视网膜中填充细胞的模式,来测试轴突向心移位的提议,就像在金鱼中所做的那样(伊斯特和施图尔默,1984年)。在视顶盖中进行HRP的点状应用应该被摄取:1)通过束,标记部分细胞环带;2)通过终末,标记相应视网膜拓扑位置的一群细胞;3)通过束外轴突段,标记连接部分环带和这群细胞的一条细胞带。如果终末已经向心移位,通过其束外段标记的细胞带应该具有辐条状取向,以视网膜中心为中心。随着视顶盖部位从吻侧移动到尾侧,这条细胞带应该像钟摆一样从颞侧视网膜移动到鼻侧视网膜。一般来说,我们观察到的HRP填充视网膜细胞的模式与我们的预测一致。此外,最老(吻侧)的视顶盖轴突摄取的HRP产生了更复杂的填充细胞模式,表明这些轴突在变态前已经向尾侧移位,在变态后又向心移位。