We have studied the distribution of astrocytes in the ganglion cell and nerve fibre layers of the retina in cat, rat, rabbit, and possum using anti-serum and a monoclonal antibody against glial fibrillary acidic protein (GFAP) and our own monoclonal antibody against glial filaments. The distribution of retinal astrocytes appears to be strongly determined by the vasculature of the retina; astrocytes are absent from almost all the retina of the possum and from the avascular regions of the rabbit retina. In the cat and rabbit, retinal astrocytes also show a strong affinity for the bundles of ganglion cell axons found at the inner surface of the retina. Retinal astrocytes do not invest the somas of ganglion cells, and even in areas of retina in which they are numerous, they are sharply confined to the layer of ganglion cell axons. It is suggested that retinal astrocytes are "immigrant" fibrous astrocytes that enter the retina with its vasculature.

A detailed comparison is made between astrocytes and Müller cells of the cat's retina, with emphasis on their structural specialisations. Evidence is presented that astrocytes and Müller cells both contribute to the formation of the inner glia limitans of the retina, the glia limitans of vessels, and the glial sheaths of neurones. In particular, it was noted that both astrocytes and Müller cells wrap bundles of ganglion cells axons, that both contribute processes to the glial convergence on the initial segments and node-like structures of axons, and that both wrap the somas of neurones in the ganglion cell layer. Further, it was noted that adherent junctions form between astrocytes, between Müller cells, and between astrocytes and Müller cells, but not between these cells and neurones, or among neurones. These similarities suggest that astrocytes and Müller cells function interchangeably in many respects, and we suggest that they be regarded as variants of macroglia. Quantitative differences between astrocytes and Müller cells were noted in their ensheathment of neurones. In particular, the glial sheaths around the somas of ganglion cells are formed predominantly by Müller cells, and the glial processes attached to node-like specialisations of their axons are formed mainly by astrocytes. One qualitative difference was noted between the two cell classes. The gap junctions which form between astrocytes do not form between Müller cells or between cells of the two classes. From these differences, and previously established features of their shape, orientation, distribution and origin, a hypothesis is developed of the specialisation of macroglia represented by Müller cells.

In this paper, for the first time a quantitative description of the morphology and distribution of Müller cells in the macaque monkey retina using immunohistochemistry and high resolution confocal laser scanning microscopy is given. By their morphological features Müller cells are ideally adapted to their neuronal environment in the various retinal layers, with a dense network of horizontal processes, especially in the inner plexiform layer, and close contacts to neuronal somata especially in the outer nuclear layer and ganglion cell layer. Morphology varies with retinal eccentricity. The thickness of the inner trunk increases significantly with increasing retinal eccentricity. According to the overall thickness of the retina, Müller cells in central retina are longer than in peripheral regions. In the parafoveal region, the outer trunks of Müller cells in the outer plexiform layer are immensely elongated. These Müller fibres can reach lengths of several hundred micrometers as they travel through the outer plexiform layer from the foveal centre towards the foveal border where they enter the inner nuclear layer. Müller cell density varies between 6000 cells/mm2 in far peripheral and peak densities of > 30,000 cells/mm2 in the parafoveal retina. There is a close spatial relationship between Müller cells and blood vessels in the monkey retina, suggesting a role of Müller cells in the formation of the blood-retinal barrier, in the uptake of nutrients and the disposal of metabolites.

We describe the morphology and distribution of Müller cells in wholemounts of rabbit retinae labelled with either monoclonal antibodies (anti-Vimentin, 3H3, 4D6, and 4H11), or intracellular horseradish peroxidase. Several new features of Müller cell organization are noted. First, Müller cells appear to compose a single morphological class and their morphology varies systematically with retinal thickness. Second, in contrast to other retinal glia, Müller cells have a neuronlike distribution, with a peak density of 10,700-15,000 cells per mm2 at the visual streak and a minimum density of 4,400-6,000 per mm2 at both the superior and inferior retinal edges. There are 4.2 +/- 0.5 x 10(6) Müller cells per retina. Third, unlike in other species, rabbit Müller cells do not contact blood vessels, suggesting that they do not participate in the transfer of metabolites or in the blood:retinal barrier. Fourth, each Müller cell has a vitread endfoot about 20-40 microns in diameter composed of numerous fimbriae. The fimbriae from a single Müller cell generally contact several axon fascicles in the nerve fibre layer, and at each point along its length each fascicle is enclosed by the overlapping fimbriae from several Müller cells. Fifth, in the inner and outer plexiform layers, numerous filamentous branchlets extend 20 microns or more from the radial trunk, interweaving with branchlets from nearby Müller cells to form dense and continuous strata. In the ganglion cell layer and outer nuclear layer, Müller cell processes completely wrap neuronal somata, whereas in the inner nuclear layer they partially wrap somata. We discuss the functional and developmental implications of these observations.

Eight monoclonal antibodies were used to label Müller cells in four mammals that have vascular retinae (cats, dogs, humans, and rats) and in three with avascular retinae (echidnas, guinea pigs, and rabbits). Müller cells were found to have a fairly uniform retinal distribution in seven species, with a mean density of 8,000-13,000 cells mm-2. Müller cells in avascular retinae differ from their vascular counterparts in four respects. First, they are shorter than those in vascular retinae. This difference is mainly due to a reduction in the thickness of the outer nuclear layer. Second, the trunks of Müller cells in avascular retinae tend to be thicker, although those in echidnas are an exception to this trend. Third, Müller cell rootlets in avascular retinae follow a more tortuous course than those in vascular retinae, reflecting the fact that photoreceptor nuclei in the two types of retina have different shapes and stacking patterns. Fourth, due to a reduction in the density of photoreceptors in avascular retinae, there are fewer neurones per Müller cell. Although these four features may enable Müller cells to assist the nutrition of neurones in the inner layers of avascular retinae, they are unlikely to be morphological specializations that have evolved for that purpose. Rather, these features appear to be a direct consequence of the fact that avascular retinae are thinner and have a differently organised outer nuclear layer. These features aside, Müller cells in avascular retinae closely resemble their counterparts in vascular retinae.