Dacey D M
Department of Biological Structure, University of Washington, Seattle 98195.
J Neurosci. 1993 Dec;13(12):5334-55. doi: 10.1523/JNEUROSCI.13-12-05334.1993.
To study their detailed morphology, ganglion cells of the human retina were stained by intracellular tracer injection, in an in vitro, whole-mount preparation. This report focuses on the dendritic morphology and mosaic organization of the major, presumed color-opponent, ganglion cell class, the midget cells. Midget cells in the central retina were recognized by their extremely small dendritic trees, approximately 5-10 microns in diameter. Between 2 and 6 mm eccentricity, midget cells showed a steep, 10-fold increase in dendritic field size, followed by a more shallow, three- to fourfold increase in the retinal periphery, attaining a maximum diameter of approximately 225 microns. Despite large local variation in dendritic field size, midget cells formed one morphologically distinctive class at all retinal eccentricities. Two midget cell types were distinguished by their dendritic stratification in either the inner or outer portion of the inner plexiform layer (IPL), and presumably correspond to ON- and OFF-center cells respectively. The mosaic organization of the midget cells was examined by intracellularly filling neighboring cells in small patches of retina. For both the inner and outer midget populations, adjacent dendritic trees apposed one another but did not overlap, establishing a coverage of no greater than 1. The two mosaics differed in spatial scale, however: the outer midget cells showed smaller dendritic fields and higher cell density than the inner midget cells. An outer:inner cell density ratio of 1.7:1 was found in the retinal periphery. An estimate of total midget cell density suggested that the proportion of midget cells increases from about 45% of total ganglion cell density in the retinal periphery to about 95% in the central retina. Nyquist frequencies calculated from midget cell spacing closely match a recent measure of human achromatic spatial acuity (Anderson et al., 1991), from approximately 6 degrees to 55 degrees eccentricity. Outside the central retina, midget cell dendrites arborized in clusters within the overall dendritic field. With increasing eccentricity, the dendritic clusters increased in number and remained small (approximately 10-20 microns diameter) relative to the size of the dendritic field. Because neighboring midget cell dendritic trees do not overlap, the mosaic as a whole showed a pattern of clusters and holes. We hypothesize that midget cell dendritic trees may contact individual axon terminals of some midget bipolar cells and avoid contacting others, providing a basis for the formation of cone-specific connections in the IPL.
为了研究人类视网膜神经节细胞的详细形态,我们采用细胞内示踪剂注射的方法,在体外全视网膜铺片标本中对其进行染色。本报告重点关注主要的、推测为颜色拮抗型的神经节细胞类型——侏儒细胞的树突形态和镶嵌组织。中央视网膜中的侏儒细胞通过其极其微小的树突野得以识别,直径约为5 - 10微米。在离中心凹2至6毫米的偏心度范围内,侏儒细胞的树突野大小急剧增加10倍,随后在视网膜周边区域增加较为平缓,增加三至四倍,最大直径达到约225微米。尽管树突野大小存在较大局部差异,但侏儒细胞在所有视网膜偏心度下都构成一个形态独特的类别。两种侏儒细胞类型通过其在内网状层(IPL)内部或外部的树突分层来区分,大概分别对应于ON中心细胞和OFF中心细胞。通过在视网膜小区域内对相邻细胞进行细胞内填充来检查侏儒细胞的镶嵌组织。对于内层和外层侏儒细胞群体,相邻的树突野相互邻接但不重叠,覆盖率不超过1。然而,这两种镶嵌组织在空间尺度上有所不同:外层侏儒细胞的树突野较小,细胞密度比内层侏儒细胞高。在视网膜周边区域发现外层与内层细胞密度比为1.7:1。对侏儒细胞总密度的估计表明,侏儒细胞的比例从视网膜周边区域占神经节细胞总密度的约45%增加到中央视网膜的约95%。根据侏儒细胞间距计算的奈奎斯特频率与最近对人类消色差空间敏锐度的测量结果(Anderson等人,1991)紧密匹配,范围从约6度偏心度到55度偏心度。在中央视网膜之外,侏儒细胞的树突在整个树突野内成簇分支。随着偏心度增加,树突簇数量增加,并且相对于树突野大小仍然较小(直径约为10 - 20微米)。由于相邻侏儒细胞的树突野不重叠,整个镶嵌组织呈现出簇状和空洞的模式。我们推测侏儒细胞的树突可能与某些侏儒双极细胞的单个轴突终末接触,而避免与其他轴突终末接触,这为在内网状层中形成视锥细胞特异性连接提供了基础。