Gaze R M, Grant P
Department of Zoology, University of Edinburgh, Scotland.
J Comp Neurol. 1992 Jan 15;315(3):264-74. doi: 10.1002/cne.903150303.
During development of the retina in mammals and birds, most retinal ganglion cells (RGC) that are produced are eliminated later in development by cell death. In lower vertebrates, however, such massive cell death has not been observed; total ganglion cell number increases linearly during most of development. Using 3H-thymidine or 5-bromodeoxyuridine labeling of retinal cell nuclei, we have been able to identify postmitotic RGC populations in Xenopus central retina at different developmental stages and follow their fate during development to postmetamorphic stages. RGC populations that become postmitotic between embryonic stages 32 and 49, during the initial stages of retinal growth, lose 40-77% of their cells during metamorphosis (approximately 4,000-5,000 cells). Twenty percent of the RGC present at stage 54, which later disappear, represent the same population of dying RGC that were present at stage 49. This suggests that the ganglion cells that became postmitotic between stage 49 and 53/54 show no apparent decline in numbers during metamorphosis. Since thyroxine is known to stimulate an increase in RGC number as well as the extent of fiber projection on the tectum, we suggest that this reduction in RGC numbers is not due to thyroxine-induced neuronal cell death. After stage 54, however, binocular vision develops in Xenopus (Keating, '74) and ipsilateral fibers begin to grow into thalamic visual neuropils (Hoskins and Grobstein, '85). We suggest, therefore, that as in mammals, in which RGC elimination correlates with binocular segregation of contralateral and ipsilateral retinal axons in visual centers, a similar process may occur in the frog among those RGC projecting to thalamic visual neuropils.
在哺乳动物和鸟类视网膜发育过程中,大多数产生的视网膜神经节细胞(RGC)在发育后期会通过细胞死亡被清除。然而,在低等脊椎动物中,尚未观察到如此大规模的细胞死亡;在大多数发育阶段,神经节细胞总数呈线性增加。通过用³H-胸腺嘧啶核苷或5-溴脱氧尿苷标记视网膜细胞核,我们能够鉴定非洲爪蟾中枢视网膜在不同发育阶段有丝分裂后的RGC群体,并追踪它们在发育至变态后阶段的命运。在视网膜生长初期,即在胚胎第32至49阶段之间进入有丝分裂后期的RGC群体,在变态过程中会损失40%-77%的细胞(约4000-5000个细胞)。第54阶段存在的RGC中有20%后来消失了,这些消失的RGC与第49阶段存在的即将死亡的RGC是同一群体。这表明在第49至53/54阶段进入有丝分裂后期的神经节细胞在变态过程中数量没有明显减少。由于已知甲状腺素会刺激RGC数量增加以及顶盖纤维投射范围扩大,我们认为RGC数量的减少并非由甲状腺素诱导的神经元细胞死亡所致。然而,在第54阶段之后,非洲爪蟾开始形成双眼视觉(基廷,1974年),同侧纤维开始向丘脑视觉神经毡生长(霍斯金斯和格罗布斯坦,1985年)。因此,我们认为,与哺乳动物一样,在哺乳动物中RGC的清除与视觉中枢中对侧和同侧视网膜轴突的双眼分离相关,在青蛙中,投射到丘脑视觉神经毡的那些RGC之间可能也会发生类似的过程。