This study investigated the organization of cells in the ganglion cell layer (GCL) using Nissl staining, retrograde cell degeneration with axotomy of the optic nerve, and retrograde cell labeling by injections of horseradish peroxidase (HRP) into the optic nerve of chicks (posthatching day 1 and 8, P-1 and P-8). The total number of cells in the GCL was 6.1 × 106 (P-1) and 4.9 × 106 (P-8), and the cell density was 14,300 cells/mm2 (P-1) and 10,400 cells/ mm2 (P-8) on average. Two high-density areas, the central area (CA) and the dorsal area (DA), were observed in the central and dorsal retinas in both P-1 (22,000 cells/mm2 in CA, 19,000 cells/mm2 in DA) and P-8 chicks (19,000 cells/mm2 in CA, 12,800 cells/mm2 in DA). The cell densities in the temporal periphery (TP) and the nasal (NP) peripheral retinas were 7,800 cells/mm2 and 12,500 cells/mm2, respectively, in P-1 and 5,000 cells/ mm2 and 8,000 cells/mm2, respectively, in P-8 chicks. The cell density in the temporal periphery was 35% (P-8) lower than in the nasal periphery in both P-1 and P-8 chicks. Thirty percent (1.9 × 106 cells in P-1) of the total cells in the GCL were resistant to axotomy of the optic nerve. The distribution of the axotomy-resistant cells showed two high-density areas in the central and dorsal retinas, corresponding to the CA (5,800 cells/mm2) and the DA (3,200 cells/mm2). These cells also exhibited a center-peripheral increase (2,200 cells/mm2 in the TP) in P-1 chicks, but the high-density area was not found in the dorsal retina of P-8 chicks. From these data and the HRP study, the number of presumptive ganglion cells in P-8 chicks was estimated to be 4 × 106 (8,600 cells/mm2 on average), and the density in each area was 13,500 (CA), 10,200 (DA), and 4,300 (TP) cells/mm2. The peripheral/ center ratios of the density of ganglion cells were significantly different along the nasotemporal and dorsoventral axes. The density of ganglion cells decreased more rapidly toward the temporal periphery (TP/CA ratio: 0.47 in P-1 and 0.32 in P-8) than toward the nasal periphery (NP/CA ratio: 0.67 in P-1 and 0.52 in P-8). In contrast, there was no significant difference in the peripheral/center ratios between the dorsal retina (DP/CA ratio: 0.6 in P-1 and 0.56 in P-8) and ventral retina (VP/CA ratio: 0.58 in P-1 and 0.51 in P-8). A small peak in the density of the presumptive ganglion cells was detected in the dorsal retina of both P-1 chicks (10,800 cells/mm2) and P-8 chicks (10,200 cells/mm2). The HRP-labeled cells were small in the CA (M ± SD: 35.7 ± 9.1 μm2) and DA (40.0 ± 11.3 μm2), and their sizes increased toward the periphery (63.4 ± 29.7 μm2 in the TP) accompanied by a decrease in the cell density. However, the axotomy-resistant cells did not significantly increase in size toward the peripheral retina (12.2 ± 2.2 μm2 in the CA, 15.2 ± 3.2 μm2 in the DA, 15.1 ± 3.8 μm2 in the TP). The characteristic distribution of ganglion cells could be related to visual behavior based upon the specialization of avian visual fields.

1.
Binggeli, R.L., and W.J. Paule (1969) The pigeon retina: quantitative aspects of the optic nerve and ganglion cell layer. J. Comp. Neurol., 137: 1–18.
2.
Boycott, B.B., and H. Wässle (1974) The morphological types of ganglion cells of the domestic cat’s retina. J. Physiol. (Lond.), 240: 397–419.
3.
Budnik, V., J. Mpodozis, F.J. Varela, and H.R. Maturana (1984) Regional specialization of the quail retina: ganglion cell density and oil droplet distribution. Neurosci. Lett, 51: 145–150.
4.
Buhl, E.H., and L. Peichl (1986) Morphology of rabbit retinal ganglion cells projecting to the medial terminal nucleus of the accessory optic system. J. Comp. Neurol., 253: 163–174.
5.
Buhl, E.H., and J.F. Dann (1988) Morphological diversity of displaced retinal ganglion cells in the rat: a lucifer yellow study. J. Comp. Neurol., 269: 210–218.
6.
Bunt, A.H., and D.S. Minckler (1977) Displaced ganglion cells in the retina of the monkey. Invest. Ophthalmol. Vis. Sci., 16: 95–98.
7.
Cajal, S., and Y. Ramon (1955) Histologie du Système Nerveux. Vol. 2 (translated by L. Azoulay in 1972), Consejo Superior de Investigaciones Cientificas, Madrid.
8.
Doi, M., Y. Uji, and H. Yamamura (1995) Morphological classification of retinal ganglion cells in mice. J. Comp. Neurol., 356: 368–386.
9.
Ehrlich, D. (1981) Regional specialization of the chick retina as revealed by the size and density of neurons in the ganglion cell layer. J. Comp. Neurol., 195: 643–657.
10.
Ehrlich, D., and I.G. Morgan (1980) Kainic acid destroys displaced amacrine cells in posthatch chicken retina. Neurosci. Lett, 17: 43–48.
11.
Famiglietti, E.V. (1992) Dendritic co-stratification of ON an ON-OFF directionally selective ganglion cells with starburst amacrine cells in rabbit retina. J. Comp. Neurol., 324: 322–335.
12.
Fischer, Q.S., and A.M. Kirby (1991) Number and distribution of retinal ganglion cells in anubis baboons (Papio anubis). Brain Behav. Evol., 37: 189–203.
13.
Fite, K.V., and S. Rosenfield-Wessels (1975) A comparative study of deep avian foveas. Brain Behav. Evol., 12: 97–115.
14.
Galvez, J.M.G., L. Puelles, and C. Prada (1977) Inverted (displaced) retinal amacrine cells and their embryonic development in the chick. Exp. Neurol., 56: 151–157.
15.
Ghosh, K.K., A.K. Goodchild, A.E. Sefton, and P.R. Martin (1996) Morphology of retinal ganglion cells in a new world monkey, the marmoset Callithrix jacchus. J. Comp. Neurol., 366: 76–92.
16.
Hayes, B.P. (1984) Cell populations of the ganglion cell layer: displaced amacrine and matching amacrine cells in the pigeon retina. Exp. Brain Res., 56: 565–573.
17.
Hayes, B.P., and A.L. Holden (1980) Size classes of ganglion cells in the central yellow field of the pigeon retina. Exp. Brain Res., 39: 269–275.
18.
Hayes, B.P., and A.L. Holden (1983) The distribution of displaced ganglion cells in the retina of the pigeon. Exp. Brain Res., 49: 181–188.
19.
Hebel, R., and H. Holländer (1983) Size and distribution of ganglion cells in the human retina. Anat. Embryol., 168: 125–136.
20.
Holden, A.L. (1981) Classifying and comparing retinal ganglion cells. Brain Behav. Evol., 18: 188–193.
21.
Hughes, A. (1975) A quantitative analysis of the cat retinal ganglion cell topography. J. Comp. Neurol., 163: 107–128.
22.
Hughes, A. (1981) Population magnitudes and distribution of the major model classes of cat retinal ganglion cells as estimated from HRP filling and systematic survey of the soma diameter spectra for classical neurons. J. Comp. Neurol., 197: 303–339.
23.
Hughes, A., and D.I. Vaney (1980) Coronate cells: displaced amacrines of the rabbit retina? J. Comp. Neurol., 189: 169–189.
24.
Ikushima, M., M. Watanabe, and H. Ito (1986) Distribution and morphology of retinal ganglion cells in the Japanese quail. Brain Res., 376: 320–334.
25.
Kalloniatis, M., and G.A. Napper (1996) Glutamate metabolic pathways in displaced ganglion cells of the chicken retina. J. Comp. Neurol., 367: 518–536.
26.
Karten, H.J., K.V. Fite, and N. Brecha (1977) Specific projection of displaced retinal ganglion cells upon the accessory optic system in the pigeon (Columbia livia). Proc. Natl. Acad. Sci. USA, 74: 1753–1756.
27.
Kolb, H., R. Nelson, and A. Mariani (1981) Amacrine cells, bipolar cells and ganglion cells of the cat retina: a Golgi study. Vision Res., 21: 1081–1114.
28.
Koontz, M.A., L.E. Hendrickson, and S.T. Brace (1993) Immunocytochemical localization of GABA and glycine in amacrine and displaced amacrine cells of macaque monkey retina. Vision Res., 33: 2617–2628.
29.
Layer, P.G., and G. Vollmer (1982) Lucifer yellow stains all displaced amacrine cells of the chicken retina during embryonic development. Neurosci. Lett, 31: 99–104.
30.
Leventhal, A.G., R.W. Rodieck, and B. Dreher (1981) Retinal ganglion cell classes in the old world monkey: morphology and central projections. Science, 213: 1139–1142.
31.
Leventhal, A.G., K.G. Thompson, and D. Liu (1993) Retinal ganglion cells within the foveola of new world (Saimiri sciureus) and old world (Macaca fascicularis) monkeys. J. Comp. Neurol., 338: 242–254.
32.
Lima, S.M.A., L.C.L. Silveira, and V.H. Perry (1996) Distribution of M retinal ganglion cells in diurnal and nocturnal new world monkeys. J. Comp. Neurol., 368: 538–552.
33.
Linberg, K.A., S. Suemune, and S.K. Fisher (1996) Retinal neurons of the california ground squirrel, Spermophilus beecheyi: a Golgi study. J. Comp. Neurol., 365: 173–216.
34.
Mesulam, M.-M. (1976) The blue reaction product in horseradish peroxidase neurohistochemistry: incubation parameters and visibility. J. Histochem. Cytochem., 24: 1273–1280.
35.
Mesulam, M.-M. (1982) Principles of horseradish peroxidase neurohistochemistry and their applications for tracing neural pathways-axonal transport, enzyme histochemistry and light microscopic analysis. In Tracing Neural Connections with Horseradish Peroxidase (ed. by M.-M. Mesulam), John Wiley and Sons, Chichester, pp. 1–151.
36.
Muchnick, N., and E. Hibbard (1980) Avian retinal ganglion cells resistant to degeneration after optic nerve lesion. Exp. Neurol., 68: 205–216.
37.
Naito, J. (1989) Retinogeniculate projection fibers in the monkey optic nerve: a demonstration of the fiber pathways by retrograde axonal transport of WGA-HRP. J. Comp. Neurol., 284: 174–186.
38.
Naito, J. (1994) Retinogeniculate projection fibers in the monkey optic chiasm: a demonstration of the fiber arrangement conjugated to horseradish peroxidase. J. Comp. Neurol., 346: 559–571.
39.
Oyster C.W., F.R. Amthor, and E.S. Takahashi (1993) Dendritic architecture of ON-OFF direction-selective ganglion cells in the rabbit retina. Vision Res., 33: 579–608.
40.
Peichl, L. (1989) Alpha and delta ganglion cells in the rat retina. J. Comp. Neurol., 286: 120–139.
41.
Peichl, L., E.H. Buhl, and B.B. Boycott (1987a) Alpha ganglion cells in the rabbit retina. J. Comp. Neurol., 263: 25–41.
42.
Peichl, L., H. Ott, and B.B. Boycott (1987b) Alpha ganglion cells in mammalian retinae. Proc. Roy. Soc. (Lond.) B, 231: 169–197.
43.
Perry, V.H., R. Oehler, and A. Cowey (1984) Retinal ganglion cells that project to the dorsal lateral geniculate nucleus in the macaque monkey. Neurscience, 12: 1101–1123.
44.
Polyak, S.L. (1957) The Vertebrate Visual System. The University of Chicago Press, Chicago.
45.
Provis, J.M. (1979) The distribution and size of ganglion cells in the retina of the pigmented rabbit: a quantitative analysis. J. Comp. Neurol., 185: 121–138.
46.
Reiner, A., N. Brecha, and H.J. Karten (1979) A specific projection of retinal displaced ganglion cells to the nucleus of the basal optic root in the chicken. Neuroscience, 4: 1679–1688.
47.
Rodieck, R.W., K.F. Binmoeller, and J. Dineen (1985) Parasol and midget ganglion cells of the human retina. J. Comp. Neurol., 233: 115–132.
48.
Saito, H.-A. (1983) Morphology of physiologically identified X-, Y-, and W-type retinal ganglion cells of the cat. J. Comp. Neurol., 221: 279– 288.
49.
Stone, J. (1978) The number and distribution of ganglion cells in the cat’s retina. J. Comp. Neurol., 180: 753–772.
50.
Stone, J., B. Dreher, and A. Leventhal (1979) Hierarchical and parallel mechanisms in the organization of visual cortex. Brain Res. Rev., 1: 345–394.
51.
Tauchi, M., K. Morigiwa, and Y. Fukuda (1992) Morphological comparisons between outer and inner ramifying alpha cells of the albino rat retina. Exp. Brain Res., 88: 67–77.
52.
Vaney, D.I., L. Peichl, and B.B. Boycott (1981) Matching populations of amacrine cells in the inner nuclear and ganglion cell layers of the rabbit retina. J. Comp. Neurol., 199: 373–391.
53.
Vitek, D.J., J.D. Schall, and A.G. Leventhal (1985) Morphology, central projections, and dendritic field orientation of retinal ganglion cells in the ferret. J. Comp. Neurol., 241: 1–11.
54.
Wässle, H., L. Peichl, and B.B. Boycott (1981) Morphology and topography of on- and off-alpha cells in the cat retina. Proc. Roy. Soc. (Lond.) B, 212: 157–175.
55.
Wässle, H., and R.-B. Illing (1980) The retinal projection to the superior colliculus in the cat: a quantitative study with HRP. J. Comp. Neurol., 190: 333–356.
56.
Wässle, H., W.R. Levick, and B.G. Cleland (1975) The distribution of the alpha type of ganglion cells in the cat’s retina. J. Comp. Neurol., 159: 419–438.
57.
Wathey, J.C., and J.D. Pettigrew (1989) Quantitative analysis of the retinal ganglion cell layer and optic nerve of the barn owl Tyto alba. Brain Behav. Evol., 33: 279–292.
58.
Webb, S.V., and J.H. Kaas (1976) The sizes and distribution of ganglion cells in the retina of the owl monkey, Aotus trivirgatus. Vision Res., 16: 1247–1254.
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