Distribution Of Retinal Ganglion Cells Research Articles

Overlapping Ipsilateral and Contralateral Retinal Projections to the Lateral Geniculate Nucleus and Superior Colliculus in the Cat: A Retrograde Triple Labelling Study

To analyze the relative proportion and distribution of retinal ganglion cells projecting ipsilaterally and contralaterally in the cat, large injections of the fluorescent tracers Fluoro Gold, Fast Blue, and Diamidino Yellow were made in the main layers of the lateral geniculate nucleus (LGN) and superior colliculus (SC). One tracer was injected in both the LGN and SC on one side, and the other two tracers were injected contralaterally, in the LGN and SC, respectively; labelled ganglion cells were charted on retinal whole mounts. Ganglion cells labelled from the LGN and SC were highly intermingled in both the ipsilateral and contralateral retinae. The adopted combinations of tracers allowed the detection of cells double labelled from the SC and LGN, supporting the occurrence of branched retino-thalamic axons to the SC. About one-fourth of the ganglion cells labelled from the LGN and SC was located in the eye ipsilateral to the injection. Retrograde labelling from the ipsilateral side was almost entirely confined to the temporal hemiretina. In the contralateral eye, labelled cells were mainly concentrated in the nasal hemiretina, but more than 10% were also detected in the temporal half of the retina. In the latter area, cells displaying the entire range of sizes of the retinal ganglion cells, labelled from the contralateral LGN and SC, were found throughout the entire hemiretina. However, more than 50% of such “wrong” projecting cells were grouped in a strip of 2 mm closest to the naso-temporal division. Control experiments, in which the tracers injections were restricted to the rostral and dorsal portions of the LGN to avoid optic tract contamination, consistently confirmed the occurrence and distribution of the “wrong” projecting cells in the temporal hemiretina. Thus, these latter cells are not grouped in a central strip, where ganglion cells would have the same chance of projecting to the same or to the opposite side, and sparsely distributed in the temporal periphery, as previously believed. Instead, the present findings indicate that the retinal ganglion cells of origin of contralateral projections are distributed more in a continuum, with a naso-temporal gradient of density, across the temporal hemiretina.

Increased serotonin in the developing superior colliculus does not alter the number or distribution of retinotectal ganglion cells.

Administration of a single subcutaneous dose of 5,7-dihydroxytryptamine (5,7-DHT) to newborn hamsters results in a significant increase in the density of serotoninergic (5-HT) fibers in the superficial layers of the superior colliculus (SC) and marked abnormalities in both the crossed and uncrossed retinotectal projections when these animals reach adulthood (R. Rhoades, C. Bennett-Clarke, R. Lane, M. Leslie, and R. Mooney, 1993, J. Comp. Neurol. 334:397-409). The present study was undertaken to determine whether changes in the retinotectal projection of 5,7-DHT-treated animals were associated with alterations in the number or distribution of retinal ganglion cells in these animals. Nissl staining of retinae from normal adult and 5,7-DHT-treated hamsters revealed no differences between them in the number or average diameter of cells in the retinal ganglion cell layer. Retrograde labeling with horseradish peroxidase (HRP) demonstrated no effect of 5,7-DHT treatment on the number or distribution of ipsilaterally or contralaterally projecting ganglion cells. Neonatal 5,7-DHT administration also had no effect on the distribution of soma diameters for HRP-labeled retinal ganglion cells. Electron microscopic analysis demonstrated no significant difference between the number of optic nerve fibers in the normal and 5,7-DHT-treated hamsters. The results are consistent with the conclusion that the effect of 5,7-DHT on the retinotectal projection may primarily be a function of this toxin, or the increase in 5-HT it induces, on the terminal arbors of retinotectal axons rather than on their parent cells.

The visual system of the Florida garfish, Lepisosteus platyrhincus (Ginglymodi). IV. Bilateral projections and the binocular visual field.

Infusion of cobaltous-lysine into the optic nerve of juvenile Florida garfish reveals that the preoptic area, pretectum, thalamus and the mediorostral and ventrolateral poles of the optic tectum each receive bilateral, monosynaptic input. The bilateral input to the mediorostral pole of the tectum projects via the dorsal optic tract and terminates in the rostral half of the tectum. The bilateral input to the ventrolateral tectum projects via the ventral optic tract and extends the whole length of the tectum. The incidence of direct ipsilateral input to the tectum suggests binocular vision may play a functional role in the survival of this species. In order to test this hypothesis, we examined the retinal distribution of ipsilateral retinal ganglion cells that project to the mediorostral tectum and the size and location of the binocular visual field. Cell distribution was examined by retrograde labelling with rhodamine-conjugated dextran amine delivered by microinjections into the mediorostral pole of the right optic tectum. In the ipsilateral retina, ganglion cells are distributed in a narrow temporoventral area with a mean of 0.44 +/- 0.14 x 10(3) cells per mm2 apposing the retinal margin. In the contralateral retina, ganglion cells are also distributed within this temporoventral region with a mean peak density of 2.33 +/- 0.47 x 10(3) cells per mm2. Three broad classes of ipsilaterally projecting retinal ganglion cells (orthotopic cells within the ganglion cell layer, displaced cells within the inner nuclear layer, and giant cells within the ganglion and inner nuclear layers) are identified, intermingled in both the ipsilateral and contralateral retinae after retrograde labelling from the mediorostral pole of the tectum. Ophthalmoscopic mapping of the monocular and binocular visual fields reveals two small frontal wedges of binocular overlap. A dorsal wedge (12 degrees wide) extends from approximately 7 degrees above the horizontal axis to 10 degrees beyond the vertical axis, and a ventral wedge (20 degrees wide) extends approximately 10 degrees below the horizontal axis to 15 degrees beyond the vertical axis. The dorsal binocular visual field is subtended by the temporoventral region of the retina possessing both ipsilaterally and contralaterally projecting ganglion cells which terminate in the mediorostral pole of the optic tectum. Therefore, the partial decussation of retinal ganglion cell axons at the optic chiasm brings information from corresponding regions of the binocular visual field into register in the mediorostral pole of the optic tectum.(ABSTRACT TRUNCATED AT 250 WORDS)

The Visual System of the Florida Garfish, <i>Lepisosteus platyrhincus</i>(Ginglymodi) (Part 2 of 2)

Infusion of cobaltous-lysine into the optic nerve of juvenile Florida garfish reveals that the preoptic area, pretectum, thalamus and the mediorostral and ventrolateral poles of the optic tectum each receive bilateral, monosynaptic input. The bilateral input to the mediorostral pole of the tectum projects via the dorsal optic tract and terminates in the rostral half of the tectum. The bilateral input to the ventrolateral tectum projects via the ventral optic tract and extends the whole length of the tectum. The incidence of direct ipsilateral input to the tectum suggests binocular vision may play a functional role in the survival of this species. In order to test this hypothesis, we examined the retinal distribution of ipsilateral retinal ganglion cells that project to the mediorostral tectum and the size and location of the binocular visual field. Cell distribution was examined by retrograde labelling with rhodamine-conjugated dextran amine delivered by microinjections into the mediorostral pole of the right optic tectum. In the ipsilateral retina, ganglion cells are distributed in a narrow temporoventral area with a mean of 0.44±0.14 x 10³ cells per mm² apposing the retinal margin. In the contralateral retina, ganglion cells are also distributed within this temporoventral region with a mean peak density of 2.33 ± 0.47 x 10³ cells per mm². Three broad classes of ipsilaterally projecting retinal ganglion cells (orthotopic cells within the ganglion cell layer, displaced cells within the inner nuclear layer, and giant cells within the ganglion and inner nuclear layers) are identified, intermingled in both the ipsilateral and contralateral retinae after retrograde labelling from the mediorostral pole of the tectum. Ophthalmoscopic mapping of the monocular and binocular visual fields reveals two small frontal wedges of binocular overlap. A dorsal wedge (12° wide) extends from approximately 7° above the horizontal axis to 10° beyond the vertical axis, and a ventral wedge (20° wide) extends approximately 10° below the horizontal axis to 15° beyond the vertical axis. The dorsal binocular visual field is subtended by the temporoventral region of the retina possessing both ipsilaterally and contralaterally projecting ganglion cells which terminate in the mediorostral pole of the optic tectum. Therefore, the partial decussation of retinal ganglion cell axons at the optic chiasm brings information from corresponding regions of the binocular visual field into register in the mediorostral pole of the optic tectum. In addition, the size of the binocular visual field matches the relative proportion of the retina containing ipsilaterally projecting ganglion cells. The ontogeny and phylogeny of ipsilateral retinofugal projections in actinopterygians is discussed in relation to the putative secondary pathways mediating binocular vision.

Retinal ganglion cells labelled from the pulvinar nucleus in macaque monkeys

In order to study the distribution and morphological classes of retinal ganglion cells that can be retrogradely labelled from the pulvinar nucleus, we made two iontophoretic injections of horseradish peroxidase into the pulvinar in each hemisphere of five macaque monkeys. The retrogradely labelled ganglion cells projecting to or through the pulvinar nucleus were examined in retinal whole-mounts. They comprise all three major ganglion cell classes. Primate gamma cells formed the great majority of classifiable cells and, like the primate alpha cells that were found in much smaller numbers, they were already known to send axons to the superior colliculus and to the pretectal complex. In contrast, the primate beta cells were hitherto thought to project solely to the dorsal lateral geniculate nucleus. This primate beta cell projection to an extrageniculate target could account in part for the substantial number of primate beta cells that escape transneuronal retrograde retinal degeneration following striate cortical ablation, and might contribute to the residual visual sensitivity that survives destruction of striate cortex and the degeneration of the lateral geniculate nucleus.

Retinal ganglion cells within the foveola of New World (Saimiri sciureus) and Old World (Macaca fascicularis) monkeys.

The morphology and distribution of retinal ganglion cells within the foveola of New World (Saimiri sciureus) and Old World (Macaca fascicularis) monkeys labeled as a result of horseradish peroxidase injections into the dorsal lateral geniculate nucleus were studied. The results indicate that monkey's foveola normally contains significant numbers of retinal ganglion cells. Most of these project ipsilaterally. Cells within the foveola are larger than other cells in monkey central retina; their dendritic fields are up to 50 times larger in area than those of cells in the foveal slope. The dendritic fields of the ganglion cells within the foveola cover at least 70-100% of its area. Among ganglion cells within the foveola (as in most ganglion cells), there was a strong tendency for the axon and dendritic tree to arise from opposite poles of the soma. The axon-dendrite axes of ganglion cells within the foveola did not show a consistent pattern. In contrast, the axes of ganglion cells in the transition zone between the foveola and the foveal slope were directed tangentially to the circumference of the fovea. The dendritic coverage of the foveola by retinal ganglion cells suggests functional significance and provides a possible neural basis for 2-3 degrees of bilateral representation of the fovea within the central visual pathways. Alternatively, or in addition, these cells may be "remnants of foveation" and provide insight into the developmental processes that mediate the development of the fovea.

Number, distribution, and morphology of retinal ganglion cells with axons regenerated into peripheral nerve graft in adult cats

The optic nerve of adult cats was transected and an autologous peripheral nerve was transplanted to the stump. Two months later retinal ganglion cells (RGCs) that had regenerated their axons were retrogradely labeled with fluorescent dye injected into the graft. The more than 4000 labeled cells, which were 3.0-4.3% of total ganglion cell number in the intact retina, were distributed in a high-density area elongating horizontally, in a pattern similar to that observed in the intact retina. The dendritic morphology of RGCs with regenerated axons was examined after intracellular injections of HRP or Lucifer yellow in vitro. The morphology of alpha- and gamma-cells appeared comparable to normal, whereas the dendrites of some beta-cells seemed to degenerate. Some "unclassified" cells could not be identified as alpha-, beta-, or gamma-cells. The main dendrites of most unclassified cells were thin and less ramified than alpha- and beta-cells. Some unclassified cells had a vacant area in their dendritic fields, suggesting degenerative change in dendrites. The distal dendrites of some unclassified and gamma-cells had many fine processes that probably regrew during axonal regeneration. The proportion of alpha-, beta-, and other cells including gamma- and unclassified cells in the regenerating cell population was evaluated by examining all of the Lucifer yellow-injected cells in local patches. The average proportion of alpha-cells was 23.9%, about six times higher than in intact retinas. The average proportion of beta-cells was 50.3%, slightly higher than in intact retinas. The remaining cells (25.7%) could not be categorized as alpha or beta; this proportion is about half that found in intact retinas. These findings suggest that alpha-cells have the highest capacity to regenerate their axons into nerve grafts, rather than all types equally regenerating their axons.

Topography of ganglion cells in the dog and wolf retina.

The topographical distribution of retinal ganglion cells in seven breeds of dog (Canis lupus f. familiaris) and in the wolf (Canis lupus) was studied in retinal wholemounts stained with cresyl violet or with a reduced silver method. A prominent feature of all wolf retinae was a pronounced "visual streak" of high ganglion cell density, extending from the central area far into both temporal and nasal retina. By contrast, either a pronounced or a moderate visual streak was found in dog retinae. It is hypothesized that a pronounced streak is an archetypal feature of Canis lupus, and that the moderate streak in some dogs is a corollary of breeding during domestication. Irrespective of the differences in streak form and retinal area, the estimated total number of ganglion cells was about 200,000 cells in the wolf and 115,000 in the dog. Ganglion cell density maxima in the central area of the wolf were about 12,000-14,000/mm2, and in the dog they ranged from 6,400/mm2 to 14,400/mm2. This implies individual differences in visual acuity. Alpha ganglion cells constituted 3-14% of all ganglion cells in the dog and 1-18% in the wolf, depending on retinal location. A distinct feature of all dogs and wolves was the absence of alpha cells in a substantial region of temporal peripheral retina. This has not been found in any other mammalian species and suggests corresponding functional deficits.

Displaced retinal ganglion cells in normal frogs and those with regenerated optic nerves.

We have analysed the number and spatial distribution of displaced retinal ganglion cells in the frog Litoria (Hyla) moorei. A series of normal animals was compared with one in which the optic nerve was crushed and allowed to regenerate. Ganglion cells were labelled with horseradish peroxidase (HRP) applied to the optic nerve, and retinae were examined as sections or whole mounts. We analysed separately ganglion cells with somata displaced to the inner nuclear (Dogiel cells, DGCs) and to the inner plexiform layer (IPLGCs). These findings were related to data for the orthotopic ganglion cells (OGCs). The mean number of DGCs in the normal series was 2,550 (+/- 281) and fell to 1,630 (+/- 321) after regeneration, representing a mean loss of 36%. This reduction was not significantly different from the mean loss of 43% from the OGC population in which mean values fell from 474,700 (+/- 47,136) to 268,700 (+/- 54,395). In both the normal and the regenerate series, DGCs were estimated to represent means of only 0.6% of the OGC population. Densities of DGCs were highest in the naso-ventral and temporo-dorsal peripheries; densities of both DGCs and OGCs were lower after optic nerve regeneration. We conclude that the factors which affect ganglion cell death during optic nerve regeneration, do so to similar extents amongst the DGC and the OGC populations. The IPLGCs were very rare in normal animals with a mean of 420 (+/- 95). However, their numbers increased after regeneration to a mean of 3,350 (+/- 690), estimated to be 1.2% of the OGC population. These cells normally favoured peripheral retina but became pan-retinal after regeneration.(ABSTRACT TRUNCATED AT 250 WORDS)

Topography and morphology of retinal ganglion cells in Falconiforms: A study on predatory and carrion‐eating birds

The topographic distribution of retinal ganglion cells and their cell body size have been studied in five Falconiform species, including predatory (chilean eagle Buteo fuscenses australis, and sparrow hawk Falco sparverius) and carrion-eating (chimango caracara Milvago chimango; condor Vultur gryphus, and black vulture Coragyps atratus) birds. All these species had a well defined nasal fovea and a horizontal streak. Instead of a temporal fovea as in eagles and hawks, an afoveate temporal area is present in chimango, condor, and vulture. The highest ganglion cell density was found in the nasal fovea of Falco and Buteo with 65,000 and 62,000 cells/mm2, respectively. A negative correlation between ganglion cell density and cell body size was found in all the species studied. The specializations of the temporal retina showed a rather homogenous population of medium sized neurons, while the nasal foveas showed a homogeneous population of smaller ganglion cells. Finally, the peripheral retina showed a heterogeneous population of large, medium, and small ganglion cells. Predatory behavior appears to be closely related to foveal specializations, and is best exemplified in the eagle and hawk and to a lesser extent in the chimango.

Contralateral visual field representation in area 17 of the cerebral cortex of the agouti: A comparison between the cortical magnification factor and retinal ganglion cell distribution

The cortical representation of the contralateral visual field in area 17 of the agouti's brain was studied by multiunit recording. The borders of area 17 were determined by electrophysiological, cytoarchitectonic and myeloarchitectonic criteria. The results were plotted in flat, bidimensional representations of the cerebral cortex to minimize perspective distortions. The VI map, a first order topological transformation of the visual field, shows a magnified representation of the horizontal meridian that corresponds to a retinal specialization, the visual streak. The visual field representation has asymmetries that are not directly related to the topography of the retinal ganglion cell density. Whereas the ganglion cell density shows a plateau along the visual streak, the areal cortical magnification factor is higher in the region that corresponds to the intersection of the horizontal and vertical meridians. This suggests functional specializations that are not obvious when one considers the distribution of the whole ganglion cell population but which might be related to the distribution of specific ganglion cell classes.

Development of retinotopy in projections from the eye to the dorsal lateral geniculate nucleus and superior colliculus of the wallaby (Macropus eugenii)

The development of retinotopy in projections from the eye to the dorsal lateral geniculate (dLGN) and superior colliculus (SC) has been studied in the marsupial wallaby. Discrete retinal lesions were made and the remaining retinal projections were traced with horseradish peroxidase in animals at stages ranging from just after optic innervation of the dLGN and SC to the time when the projections are mature. Topographically organised projections could be recognized a few weeks after axons first reached the dLGN and SC with a topographically discrete projection from nasoventral retinal recognized later than from dorsal, dorso-temporal, temporal, and temporoventral retina. Over time there was an increase in precision of the retinotopy as judged by an increase in sharpness of the borders of filling defects in the projection labelled with horseradish peroxidase. Refinement of the projection from temporal retina preceded that from nasal retina in both the dLGN and SC and in the former occurred concomitantly with the segregation of eye-specific terminal bands. Refinement was complete 16 weeks after birth, prior to eye opening at around 20 weeks after birth. Inequalities in retinal representations in both nuclei were present from the time retinotopy could first be detected. This was before the inequalities in retinal ganglion cell distribution, which underly these representations in the adult, were obvious. Retinotopy and inequalities in retinal representation characteristic of the adult are present from a very early stage in the protracted development of visual projections in the wallaby. Refinement may involve death of inappropriately projecting cells, pruning of inappropriately projecting axon arborizations or could be achieved by growth of the retinorecipient neuropil. Temporonasal differences in the time course of refinement may reflect gradients of maturation in the retina.

Developmental changes in the distribution of retinal catecholaminergic neurones in hamsters and gerbils

Although Syrian hamsters and Mongolian gerbils are closely related, they have quite different patterns of retinal ganglion cell distribution and different patterns of retinal growth that produce their distributions. We have examined the morphology and distribution of catecholaminergic (CA) neurones in adult and developing retinae of these species in order to gain a more general understanding of the mechanisms producing cellular topographies in the retina. CA neurones were identified with an antibody to tyrosine hydroxylase (TH), the rate limiting enzyme in the production of catecholamines. In adult retinae of both hamsters and gerbils, most CA somata were located in the inner part of the inner nuclear layer (INL) and CA dendrites spread in a outer stratum of the inner plexiform layer (IPL). Their somata varied with retinal position, being largest in temporal and smallest in central retina. In hamsters, but not gerbils, a small number of CA interplexiform cells was also observed. In development, CA somata of hamster retinae were observed first in the middle and/or scleral regions of the cytoblast layer (CBL) at P (postnatal day) 8. By P12, CA somata were commonly located in the inner part of the INL and their dendrites spread into the outer region of the IPL. In developing gerbil retinae, CA somata were first observed at P6 in the middle of the CBL. Over subsequent days, they migrated into the inner part of the INL and spread their dendrites into the outer strata of the IPL. In both hamsters and gerbils, CA cells were initially concentrated in the superior temporal margin of the retina. In hamsters, this supero-temporal concentration persisted until adulthood, whereas in adult gerbils, the greatest density of CA cells was found just superior to the visual streak. These distributions were distinct from those of the ganglion cells in adult and developing retinae of each species. We discuss the role of maturational expression of TH, cell death, and retinal growth in the generation of the distinct distribution of the CA cells.

Retinal ganglion cell distribution and behaviour in procellariiform seabirds

Retinal ganglion cell distribution in nine species of procellariiform seabirds was studied by Nissl staining of retinal whole-mounts and the construction of density contour maps. Most species showed a horizontal linear area of high cell density, but concentric distributions with dorsal and central cell concentrations were found in two species. These results are discussed in relation to the birds' foraging behaviour.

Nonuniform retinal expansion during the formation of the rabbit's visual streak: Implications for the ontogeny of mammalian retinal topography

We have studied the distribution of retinal ganglion cells (RGCs) which have been retrogradely labeled from massive bilateral injections of the enzyme horseradish peroxidase into the retino-recipient nuclei of foetal and postnatal albino rabbits aged from the 24th postconceptional day (24PCD) to adulthood. The number of labeled RGCs increases from about 447,000 on the 24PCD to a peak of about 525,000 on the 27PCD. From the 29PCD to birth (31/32PCD), the number of RGCs rapidly declines to about 375,000. During the next 20 d, the number of RGCs stabilizes at about 335,000. After the 51PCD, the number of RGCs gradually declines to the adult value of about 280,000. Retinal area steadily increases from about 40 mm2 on the 24PCD to about 500 mm2 in the adult, while RGC density decreases. However, the reduction in RGC density is nonuniform: RGC density in the visual streak drops from 18,600 RGCs mm2 on the 24PCD to 4700 RGCs/mm2 in the adult, whereas RGC densities at the superior and inferior edges of the retina decreases proportionally much more (from 9300 to 105 RGCs/mm2 and from 12,000 to 170 RGCs/mm2, respectively). As a result of this differential reduction in RGC density, the streak/inferior edge ratio changes from 1.6:1 to about 28:1. In the periods from the 24PCD to the 29PCD and from the 32PCD to adulthood, the proportional increases in the streak/superior edge and streak/inferior edge RGC density ratios are linearly related to the proportional increases in retinal area. However, between the 29PCD and 32PCD, the RGC density ratios increase at a greater rate than retinal area. We conclude that (1) the centro-peripheral difference in RGC density that is already present on the 24PCD might be attributable to differential RGC generation; (2) the redistribution of RGCs between the 24PCD and adulthood is mainly due to nonuniform expansion of the retina, with minimal expansion of the visual streak and maximal expansion at the superior and inferior retinal edges; and (3) a small component of the increase in the centro-peripheral RGC density ratio, which becomes apparent between the 29PCD and 32PCD, is probably due to differential RGC loss. We discuss the pattern of retinal expansion in the rabbit and the factors which might contribute to it.

Retinal ganglion cell distribution in the cebus monkey: A comparison with the cortical magnification factors

The distribution of ganglion cells was determined in whole-mounted Cebus monkey retinae. Ganglion cell density along the horizontal meridian was asymmetric, being 1.2–4.3 higher in the nasal retinal region when compared to temporal retina at the same eccentricities. The total number of ganglion cells varied from 1.34 to 1.4 million. Ganglion cell density peaked at 49,000/mm 2 about 0.5 mm nasal to the fovea. Comparison between ganglion cell density and areal cortical magnification factors for V1 and V2 reveals that the relative representation of the fovea increases in the visual cortex. This effect seems to be a general feature of the visual system of primates.

Distribution of neurons in the pigmented rabbit's visual cortex

Past observations revealed a relationship between mammalian retinal ganglion cell distribution and cortical and subcortical projection of the rabbit's visual field (3, 4, 8). High ganglion cell density sectors of the retina claimed larger areas in their cortical and subcortical projections (with a high magnification factor) than retinal areas with low ganglion cell density. In this study, it was observed that distribution of nonpyramidal neurons in laminae IV and VI of the pigmented rabbit's visual cortex was not uniform. The nonuniformity in layer IV roughly followed the variation in the magnification factor in the cortical projection of the animal's visual field. The highest density of nonpyramidal neurons in layer IV was observed in the cortical area with the highest magnification factor.

Development of ganglion cell topography in ferret retina

The adult ferret has approximately 90,000 retinal ganglion cells, arranged in a prominent area centralis and visual streak. The role of differential cell generation, cell death, and retinal growth in the control of adult retinal ganglion cell number and distribution was evaluated by examining basic aspects of retinogenesis, including growth in retinal area, developmental changes in the number, size, and distribution of retinal ganglion cells (identification aided by retrograde transport of HRP), and the incidence of degenerating cells in the ganglion cell layer. Retinal development in the ferret was also compared to retinal development in the cat (which has an even more differentiated area centralis) to determine what alterations of developmental parameters are most closely associated with this species difference in adult morphology. The area of the retina increases linearly from birth (12 mm2) to postnatal day 24 (54 mm2), reaching an eventual adult value of 64 mm2. Ganglion cell numbers peak at 155,000 (approximately twice the adult number) on postnatal day 3, and fall to adult numbers by postnatal day 6. The remaining cells of the ganglion cell layer, principally displaced amacrine cells, reach their peak number on postnatal day 10 (approximately 280,000), falling to 200,000 by adulthood. Degenerating cells are abundant in the ganglion cell layer in the immediate postnatal period. A difference in the incidence of degenerating cells in the presumptive area centralis versus that in the retinal periphery was not observed postnatally, though there were other striking spatial nonuniformities, suggesting that differential cell loss might contribute to other features of retinal topographic organization. Ganglion cell density is virtually uniform across the retina at birth. Cell density is first reduced in the dorsal retina, resulting in a dorsal-to-ventral gradient in cell density that persists until day 10, when ganglion cell number has stabilized. By postnatal day 24, an area centralis and visual streak has emerged, but not of adult magnitude. Because ganglion cell number has stabilized long before the area centralis and visual streak emerge, we conclude that differential retinal growth is the principal mechanism producing this feature of retinal topography. Comparison with the cat suggests that the proportionately greater nonuniform growth of the cat's eye accounts for the greater differentiation of its area centralis.

Spatial organization of the retinal projection to the avian lentiform nucleus of the mesencephalon.

Utilizing the horseradish peroxidase retrograde tracing technique and the 2-deoxy-D-glucose metabolic mapping technique, we have demonstrated in chickens the distribution of retinal ganglion cells that project to the lentiform nucleus of the mesencephalon (LM) and the retinotopic organization of the projection in the LM. Retinal ganglion cells labeled after a nearly complete injection into the LM were found in the four quadrants, distributed in a wide horizontal belt lying along both sides of the retinal equator and stretching from the temporal to the nasal retina. The HRP-labeled cells, which appeared round or oval, ranged from 25 to 840 micron 2 in size with most in the smaller size range. Results of partial HRP injections into the LM and metabolic mapping patterns in the LM produced by stimulation of half the retina with horizontal visual motion suggest that there is an orderly mapping of the retina onto the LM. The inferior temporal quadrant projects to the rostrodorsal LM; the inferior nasal quadrant projects to the caudodorsal LM. The superior temporal quadrant projects to the middle and ventral LM, extending from the rostral to the caudal pole, whereas the superior nasal quadrant projects to a small zone in the caudal LM. The mapping of the retinal quadrants in the LM is remarkably similar to that reported in the optic tectum of birds. We suggest that a common embryological anlage with the optic tectum and the arrangement of retinal axons in the optic tract are important factors in establishing the retinotopic organization of the LM.

Asymmetric distribution of retinal ganglion cells in goldfish.

The distribution of retinal ganglion cells (RGCs) in goldfish was determined by removing an eye and applying cobaltous-lysine to the optic nerve for 24 hr. This procedure allowed the cobalt label to be in continuous contact with the cut ends of the optic axons and thereby backfilled many RGCs. RGC density was determined across three different sizes of retinae by using fish with different eye sizes. Confirming earlier work, we found that RGC density diminished as retinal area increased. However, irrespective of the retinal size, the density of RGCs was elevated along the temporal boundary between the dorsal and the ventral retina. A conservative estimate indicated that the RGC density in the temporal retina was at least 1.8-2.5 times higher than the mean RGC density of the entire retina. Thus, the goldfish retina does not appear to have a homogeneous distribution of RGCs as was previously considered. Small and large retinae differed with respect to the percentage of cells in the RGC layer that was RGCs. In small retinae, even when the noncobalt-filled cells (glia and displaced amacrine cells) were added to the cobalt-filled RGCs, the density of all cell types was elevated in the temporal retina relative to the remainder of the retina. Furthermore, in small retinae, the percentage of cells in the RGC layer that was RGCs (75%) was constant across the radial and circumferential aspects of the retina. In marked contrast, in medium-large retinae, a homogeneous distribution of cells across the entire retina resulted when the noncobalt-filled cells were added to the cobalt-filled cells. However, the percentage of cells that was cobalt-filled RGCs was significantly greater in the temporal retina (50%) than in the remainder of the retina (35%). In large retinae, as in small retinae, the percentage of cells that was RGCs did not vary as a function of distance from the optic disc. These data suggest that, in the course of retinal maturation, cell density in the temporal retina is elevated initially and then declines subsequently to the level of the surrounding retina. Over time, more displaced to the level of the surrounding retina. Over time, more displaced amacrine cells may be added to the tissue surrounding the temporal retina. Alternatively, more RGCs outside the temporal retina may become displaced amacrine cells. Such events could account for the growth-associated, disproportionate decrease in the percentage of cells that is RGCs in the tissue surrounding the temporal retina.(ABSTRACT TRUNCATED AT 400 WORDS)