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Development of the human retina: patterns of cell distribution and redistribution in the ganglion cell layer.

Neurogenesis in the ventricular layer and the development of cell topography in the ganglion cell layer have been studied in whole-mounts of human fetal retinae. At the end of the embryonic period mitotic figures were seen over the entire outer surface of the retina. By about 14 weeks gestation mitosis had ceased in central retina and differentiation of photoreceptor nuclei was evident within a well-defined area which constituted about 2% of total retina area. This area was approximately centered on the site of the putative fovea, identified by the exclusive development of cone nuclei at that location. The area of retina in which mitosis had ceased increased as gestation progressed. By mid-gestation mitosis in the ventricular layer occupied about 77% of the outer surface of the retina and by about 30 weeks gestation mitosis in the ventricular layer had ceased. Cell density distributions in the ganglion cell layer were nonuniform at all stages studied (14-40 weeks). Densities were highest at about 17 weeks gestation, and by mid-gestation the adult pattern of cell topography was present with maps showing elevated cell densities in posterior retina and along the horizontal meridian. Cell densities generally declined throughout the remainder of the gestation period, except in the posterior retina, where densities in the perifoveal ganglion cell layer remained high during the second half of gestation. There is a rapid decline in cell density in the foveal ganglion cell layer toward the end of gestation, and it is suggested that the persistence of high densities in the perifoveal region may be related to migration of cells away from the developing fovea. The total population of cells in the ganglion cell layer was highest (2.2-2.5 million cells) between about weeks 18 and 30 of gestation. After this the cell population declined rapidly to 1.5-1.7 million cells. It is suggested that naturally occurring neuronal death is largely responsible for this decline.

Cell Count↗

RNA content of normal and axotomized retinal ganglion cells of rat and goldfish.

The responses of rat and goldfish retinal ganglion cells to axotomy were examined by a quantitative cytochemical method for RNA and by morphometric measurement 1-60 (rat) and 3-90 (goldfish) days after interruption of one optic nerve or tract intracranially. Unoperated control animals were studied also. The RNA content of axotomized neurons of rat fell 7-60 days postoperatively. Additionally, atrophy of the axotomized somas occurred. Over time, neuronal atrophy approximately paralleled the loss of RNA, and mean cell area and RNA content were reduced by about 25% 60 days after axotomy. Incorporation of 3H-uridine by axotomized neurons declined also. Axotomized retinal ganglion cells of goldfish behaved differently from those of the rat and showed increases in RNA content, most conspicuously 14-60 days postoperatively. Enlargement of axotomized fish neurons occurred but was less proportionately than concomitant increases in RNA content. The nonaxotomized ganglion cells of goldfish displayed statistically significant increases in size and RNA content 14-49 days after unilateral optic nerve or tract lesions. In contrast, alterations in rat retinal ganglion cells contralateral to interruption of one optic nerve were of limited and questionable significance. The contrasting reactions to axotomy by the retinal ganglion cells of these two vertebrates, one of which regenerates optic axons and one of which does not, may support the proposition that the somal response to axon injury has an important bearing upon the success or failure of CNS regeneration.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Retinal ganglion cell death during optic nerve regeneration in the frog Hyla moorei.

In the frog Hyla moorei we have estimated there to be between approximately 450,000 and 750,000 cells in the retinal ganglion cell layer. Optic axon counts and retrograde transport of horseradish peroxidase (HRP) indicated that 72-76% of these were ganglion cells. Cells of this type were distributed as a temporally situated area centralis within a horizontal visual streak. Cell and optic axon counts showed that there was an approximately 40% loss of ganglion cells during optic nerve regeneration. Ganglion cells appeared chromatolysed by 6-8 days after an extracranial nerve crush but there was no indication of cell death until 15 days. By this stage anterograde transport of HRP indicated that axons had reached the chiasma. Death was first seen in the area centralis, extended along the streak, and finally was observed in the periphery by 65 days; cell counts demonstrated that at this time the wave of death was almost complete. We have previously shown by electrophysiological visual mapping (Humphrey and Beazley, '82) and confirmed in this study that visuotectal projections were retinotopically organized during regeneration. Multiunit receptive fields were initially large but progressively refined starting in nasal field (temporal retina) to restore a normal projection. The similar sequences whereby the visuotectal projection became refined and death took place in the retinal ganglion cell layer suggested that death may be related to a process of organization within the regenerating projection. In normal animals primary visual pathways revealed by anterograde transport of HRP were essentially similar to those of Rana pipiens and R. esculenta. Regenerating axons generally remained within optic pathways. Exceptions were a retinoretinal projection which was not completely withdrawn even after 1,028 days and a direct projection to the ipsilateral tectum via an inappropriate part of the optic tract.

Animals↗

Ganglion cell dendritic structure and retinal topography in the rat.

The dendritic field size, the distribution of the dendrites relative to the cell body, and the overall shape of the dendritic field of type I ganglion cells in the rat retina were analyzed. These features of neuronal structure were related to the topography of the rat retina. As in the cat, the cell bodies of type I ganglion cells are arranged in a nonrandom mosaic. Previous work has demonstrated that the density of type I cells in the rat retina does not covary with the density of all ganglion cells. Type I dendritic field size varies over the retina; the increase in dendritic field size is accounted for better by the decrease in type I density than by the decrease in overall ganglion cell density. The center of the dendritic field of most type I cells is displaced in the plane of the retina from the cell body. Unlike in carnivore retina (Schall and Leventhal: J. Comp. Neurol. 257:149-159, '87), the dendritic fields in the rat are not displaced down the ganglion cell density gradient. Rather, there is a tendency for the dendritic trees, especially in temporal retina, to be displaced toward dorsal retina. Most of the dendritic fields are elongated, but the degree of elongation is less than that observed in carnivore or primate retina. Unlike in carnivore and primate retina (Leventhal and Schall: J. Comp. Neurol. 220:465-475, '83; Schall et al.: Brain Res. 368:18-23, '86), there is no relationship between dendritic tree orientation and position relative to any point on the retina in the rat.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Functional morphology of frog retinal ganglion cells and their central projections: the dimming detectors.

Intracellular recordings were made from frog retinal ganglion cell axons in the optic nerve. Following electrophysiological characterisation of receptive field properties, HRP was injected into the axon, and the brain and retina were subsequently stained. The morphologies of retinal ganglion cells, their dendritic domains, and their central projections were determined with light microscopy, and the optic nerve portion of the ganglion cell axon was examined with electron microscopy. This paper describes the structural and functional features of one ganglion cell class, the off units (class IV or dimming detectors) whose characteristic response is a preferential sensitivity to decreasing light intensity within the receptive field. Typical receptive field diameter of these units was about 16 degrees with a range of 3 degrees to more than 30 degrees. Examination of the spatial characteristics of their receptive field centers and surrounds showed that the class IV cells could be divided into two broad categories. Linear class IV cells did not respond to phase-reversal of a fine grating pattern. These linear cells also tended to have clear surround suppression: illumination of the surround diminished their response to light off at the center. The second group responded briskly to each reversal of the fine grating pattern, whatever its position within the receptor field center. These nonlinear class IV cells did not show surround suppression, but rather they had surround antagonism and they responded to light on in the surround. Nonlinear units were much more frequently recorded in frogs maintained in summer conditions (12-hour days, constant 20 degrees C temperature). In spite of this functional heterogeneity, all cells had similar morphology consisting of a large ganglion cell with a large dendritic arbor (400-1,000 microns) confined to a single stratum in the outer third of the inner plexiform layer, a medium-sized axon (2.4-microns diameter), a smallish pretectal arbor, and a large tectal arbor (300-700 microns) at layer 8.

Animals↗

Effect of different optic nerve lesions on retinal ganglion cell death in the frog Rana pipiens.

Following optic nerve crush in various species of frog, a proportion of the retinal ganglion cells re-establishes functional contact with the optic tectum. However, as much as 50% of the retinal ganglion cells die during this process. The determinants of an individual ganglion cell's fate have not been established. In this study of Rana pipiens, cell survival after optic nerve crush was compared with that after nerve cut followed by stump separation, a procedure that considerably delayed entry of optic axons to the brain. It was also ascertained, in the case of delayed ingrowth, whether application of nerve growth factor immediately after lesion influenced the cell death process. This study confirmed that retinal ganglion cell death is a relatively late event in regeneration, because in several animals where anterograde HRP labeling demonstrated regenerating axons within the tectum, no cell death had occurred. There was no statistically significant difference in cell death at 75 days after lesion between animals receiving nerve crush and those receiving nerve cut with stump separation, even though most crush animals had regenerated a complete visual projection, whereas most nerve cut animals had not. The application of NGF did not influence the level of cell death at 75 days after lesion. These results suggest that contact of optic axons with the optic tract or tectum is not necessary for retinal ganglion cell death to occur. However, this does not necessarily mean that contact with the brain is not involved with cell death during regeneration following nerve crush because it is possible that the mechanisms of cell death are different when axons are prevented from regenerating. Further investigations are therefore required to establish the reasons for this cell death.

Animals↗

Retinal ganglion cell morphology in the frog, Rana pipiens.

The morphology of retinal ganglion cells in the frog, Rana pipiens, has been examined in retinal flatmounts following backfilling of axons with horseradish peroxidase (HRP). Size and shape of the cell body and of the dendritic arbor, the dendritic branching pattern, and the depth of dendritic arborization within the inner plexiform layer (IPL) were all used to classify these cells. All of the ganglion cells so visualized can be grouped into one of 7 distinct cell classes. Class 1 contains the largest ganglion cells, with a soma size of 323 +/- 5.3 microns2 and dendritic fields of 86,819 +/- 11,817 microns2; the dendrites branch within strata 1 and 2 of the IPL. The second largest cells are class 2, with somas of 245 +/- 19.7 microns2 and dendritic fields of 55,983 +/- 7,392 microns2; the dendrites also branch within strata 1 and 2 of the IPL. Class 3 cells are the next largest class with somas of 211 +/- 11.8 microns2 and dendritic fields of 18,186 +/- 1,394 microns2; there are three varieties of class 3 cells based on the depth of branching of the dendrites: some cells are bistratified, others are tristratified, while still other cells arborize diffusely within the IPL. Class 4 cells are intermediate in size, with somas of 113 +/- 7.4 microns2 and dendrites of 4800 +/- 759 microns2; the dendrites arborize within strata 4 and 5 of the IPL. Class 5 cells have not been quantitatively analyzed because they are heterogeneous in soma and dendritic size. However, class 5 cells all have cell bodies displaced in location into the inner nuclear layer and all have a unique dendritic specialization: they send from 1 to 3 processes into the outer plexiform layer. Class 6 cells are the second smallest cell class with somas of 68.1 +/- 5.13 microns2 and dendritic fields of 888 +/- 182 microns2; the dendrites arborize within strata 3, 4, and 5 of the IP. Class 7 contains the smallest ganglion cells with somas of 62.1 +/- 2.86 microns2 and dendritic fields of 831 +/- 74.2 microns2; the dendrites arborize within strata 3, 4, and 5 of the IPL. The frequency of each cell class is inversely proportional to the size of the dendritic field. Thus, class 7 cells are the most frequent; class 1 cells are the least frequent. Furthermore, each of these 7 classes of ganglion cells has representative cells located in the inner nuclear layer.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Retinal ganglion cell topography in teleosts: a comparison between Nissl-stained material and retrograde labelling from the optic nerve.

The retinal topography of cells within the ganglion cell layer of three teleost species is examined in Nissl-stained material in which all neuronal elements containing Nissl substance in the cytoplasm are counted. A topographic comparison is made with retrogradely labelled ganglion cells to differentiate the proportion of nonganglion cells not possessing an axon joining the optic nerve. In the three species studied 92%, 80%, and 66% were found to be the maximum proportion of true ganglion cells in the area centralis, horizontal streak, and periphery, respectively. The proportion of nonganglion cells in the total population of cells counted was 24%. The major contribution to this discrepancy is from peripheral nonspecialized regions of the retina. There is little difference in both topography and peak densities of retinal ganglion cells between the two techniques. The soma areas of both populations are analysed, with the homogeneous nonganglion cell population possessing cells between 5 and 15 micron2 and the heterogeneous ganglion cell soma between 5 and 68 micron2, increasing in size with eccentricity.

Animals↗

Morphologies of rabbit retinal ganglion cells with complex receptive fields.

Ganglion cells that had complex receptive field properties, namely, On-Off and On direction-selective cells, orientation-selective cells, local edge detectors, and uniformity detectors (suppressed by contrast cells) were recorded in an isolated superfused rabbit eyecup preparation. Cells were first classified by their characteristic extracellular responses to manually controlled stimuli similar to those which have been used in previous in vivo studies. Ganglion cells were then impaled, confirmed in identity by intracellular recording, and iontophoretically injected with horseradish peroxidase for staining. Twenty-two ganglion cells, which included members of all the major classes mentioned above, were recovered from the visual streak or near periphery. All recovered cells were drawn in camera lucida from flat-mounted retinas and entered into a computer as two-dimensional stick figures; nearly all were three-dimensionally reconstructed to determine the level and manner of dendritic ramification in the inner plexiform layer (IPL). The location of ganglion cell dendrites in sublaminar regions of the IPL was found to be consistent with the hypothesis of a division of the IPL into excitatory On (proximal) and Off (distal) sublaminae, with some qualifications for particular classes. Each of the complex receptive field ganglion cell classes exhibited a distinctive three-dimensional dendritic arborization pattern uniquely associated with that physiological class.

Action Potentials↗

Topography and morphology of retinal ganglion cells in the coral trout Plectropoma leopardus (Serranidae): a retrograde cobaltous-lysine study.

The retinal topography of the adult coral trout Plectropoma leopardus (Serranidae, Perciformes) is examined in Nissl-stained material and confirmed by means of retrograde labelling with cobalt-lysine from the optic nerve. Concentric isodensity contours surround a temporoventral area centralis of over 1.5 x 10(4) cells per mm2 which lies ventral of the elongated optic nerve head. Peripheral densities of ganglion cells fall to less than 0.25 x 10(4) cells per mm2. A total of 1.17 x 10(5) ganglion cells is found over the entire retina. Silver intensifications of cobalt-lysine-stained cells allow a detailed analysis of ganglion cell morphology. Soma and dendritic field size increase with eccentricity, and within the confines of the area centralis only the smallest cell size is found. Dendritic arbors of peripheral ganglion cells are all oriented parallel to the retinal margin with an axon lying close to the vitreous humor. Nine classes of cells are characterized on the criteria of soma size, dendritic field size, primary and secondary branching patterns, soma position, and dendritic termination areas. Possible morphological homologies with ganglion cells of other teleosts are also discussed.

Animals↗

Dendritic morphologies of retinal ganglion cells projecting to the nucleus of the optic tract in the rabbit.

Focal injections of Rhodamine-latex microspheres or Fast Blue were made in the nucleus of the optic tract (NOT) of four rabbits. After survival times of 8-10 days, both dyes were retrogradely transported to medium to large sized ganglion cell somas in the retinas contralateral, but not ipsilateral, to the injected NOT. Most labelled cells were located in or near the visual streak, but a significant percentage were also found in the midperiphery of both the inferior and superior retina. One hundred fifteen labelled cells in four living superfused retinas were impaled under visual control and successfully injected with Lucifer Yellow. The dendritic arborizations of 60 of these were drawn from photographic montages for morphological identification and analysis. Nearly all the injected ganglion cells had large, relatively dense dendritic trees that stratified narrowly in the proximal inner plexiform layer. The dendritic field size and dendritic density of these cells varied with eccentricity, but at all eccentricities their anatomical characteristics closely resembled those of intracellularly stained On directionally selective ganglion cells. In three of the four experiments, a small percentage of ganglion cells were also labelled in the visual streak that had bistratified morphologies resembling those of On-Off directionally selective ganglion cells.

Animals↗

Regular mosaics of large displaced and non-displaced ganglion cells in the retina of a cichlid fish.

Large retinal ganglion cells in the tilapid cichlid fish Oreochromis spilurus (standard length 15-54 mm) were filled with horseradish peroxidase and studied in flatmounts. Three types, with distinct patterns of dendritic stratification, formed spatially independent, nonrandom mosaics. One type (about 0.3% of all ganglion cells) resembled the outer (off) alpha cells of mammals. They were very large, with thick primary dendrites and large, sparsely branched planar trees in the outer part of the inner plexiform layer (IPL). About 300 were arrayed regularly across each retina, their exact number and spacing depending on its size. Their somata were often displaced into the IPL, even where neighbours in the mosaic were orthotopic. Another type (0.8%) resembled the inner (on) alpha cells of mammals. These had slightly smaller somata that were never displaced and smaller trees in the middle layers of the IPL. About 800 were arrayed uniformly and regularly across each retina. A rarer type (0.06-0.08%) had two planar trees: one forming a coarse mosaic in the outer part of the inner plexiform layer (co-planar with the trees of outer alpha-like cells) and another in the outer plexiform layer. These "biplexiform" cells were smaller and rounder than alpha-like cells and always displaced. The dendrites were finer and less tapered. Cells in which we could identify an outer plexiform tree failed to cover the retina completely, but were nonrandomly distributed. We draw three main conclusions: (1) some nonmammalian vertebrates have separate inner and outer mosaics of large ganglion cells like those of mammals, (2) the vertical displacement of ganglion cell somata can vary widely within a single mosaic and may thus be functionally irrelevant, and (3) biplexiform ganglion cells exist in fish but differ in morphology from the biplexiform types described in some other vertebrates.

Animals↗

Intraorbital transection of the rabbit optic nerve: consequences for ganglion cells and neuroglia in the retina.

Rabbit retinae were stained with antibodies to glial fibrillary acidic protein (GFAP) at various times up to 5 months after complete unilateral intraorbital optic nerve transection, which is known to induce degeneration of ganglion cell axons and perikarya in the retina. A transient immunoreactivity for GFAP was observed in Müller glial cells that normally lack this marker. Müller-cell GFAP immunoreactivity became detectable 4 days after the lesion, but Müller cells were no longer labeled 3 months later. GFAP-labeled astrocytes located in the nerve fiber layer showed no change in immunoreactivity at any stage after transection. Application of horseradish peroxidase to the left and right superior colliculus of a rabbit whose optic nerve had been transected unilaterally 2 years before confirmed the completeness of the transection. Yet electron microscopy showed the presence of some healthy-looking ganglion cell axons in the lesioned retina, although these cells were deprived of their target. Labeling retinal wholemounts with neurofilament antibodies confirmed the presence of some ganglion cell axons and perikarya in the retina more than 2 years after transection. The course of these axons suggested that they were remnants of axons. Using antibodies to galactocerebroside (GC) we found that, as in the normal rabbit, these persisting ganglion cell axons were myelinated in the medullary rays. Although many ganglion cell axons had disappeared after 2 years, the number of neuroglial cells (including astrocytes and oligodendrocytes) present in the medullary ray region was not altered. The cell bodies of some oligodendrocytes were covered with a myelin sheath, an aberrant feature not seen normally.

Animals↗

Distribution of protein gene product 9.5 (PGP 9.5) in the vertebrate retina: evidence that immunoreactivity is restricted to mammalian horizontal and ganglion cells.

Using light microscopic immunocytochemistry, we have studied the distribution of protein gene product 9.5 (PGP 9.5), a neuron-specific protein first extracted from human brain (Doran et al., '83:J. Neurochem. 40:1542-1547), in the vertebrate retina. Retinas were obtained from frog, chicken, rat, rabbit, cow, cat, dog, and human. No immunoreactivity was observed in frog and only a faint staining was present in chicken. In mammalian retinas, a strong positive reaction was restricted to horizontal and ganglion cells, with minor interspecies variations. Immunostaining was present throughout the cell body and the dendritic tree in horizontal cells. At the level of retinal ganglion cells, immunolabel was particularly abundant in cell bodies and axons forming the optic nerve. Only the main dendrites were stained, the remainder of the dendritic tree giving rise to a diffuse punctate reaction in the inner plexiform layer. In rats, displaced amacrine cells, which are known to contribute largely (40-50%) to the total neuronal population within the ganglion cell layer (Perry, '81: Neuroscience 6:931-944) were not immunoreactive, as demonstrated from (i) analysis of the morphology, cell size and cell density of immunoreactive neurons in wholemounts; (ii) colocalization of retrograde label and PGP 9.5 immunoreactivity in about 80% of ganglion cells after injection of peroxidase into the optic nerve; and (iii) reduction of immunoreactivity in the inner plexiform and ganglion cell layers following optic nerve transection. Western blot analysis of extracts from rabbit retinas indicated that the immunoreactive species is PGP 9.5 or a closely related molecule. Recent studies have demonstrated that PGP 9.5 is a ubiquitin carboxyl-terminal hydrolase (Wilkinson et al., '89:Science 246:670-673). The present results, therefore, suggest that differences in the ubiquitination process exist between retinal neurons.

Animals↗

Morphological types of ganglion cells in the dog and wolf retina.

The morphological types of ganglion cells in the dog and wolf retina were studied by intracellular staining with Lucifer Yellow. These retinae contain a range of ganglion cell types that closely correspond to those found in cat retina: alpha cells with large somata and large, relatively densely branched dendritic trees; beta cells with medium-sized somata and small, densely branched dendritic trees; and a variety of other types with smaller somata and varying dendritic branching patterns and dendritic field sizes. The correspondence of canine and cat ganglion cell types strengthens the view that there is a common set of ganglion cell types in carnivores. Alpha and beta cell dendritic trees of dog and wolf are monostratified in either the inner or the outer part of the inner plexiform layer, suggesting an on/off dichotomy in the response to light. Dendritic field sizes of dog alpha and beta cells increase from the central area to peripheral retina: alpha cell fields from 160-200 microns to about 1,100 microns diameter, and beta cell fields from 25 microns to about 360 microns diameter. These sizes are quantitatively very similar to those found in cat retina. The close qualitative and quantitative morphological correspondence of cat and dog ganglion cells suggests that they are also functionally very similar. It is likely that dog alpha cells have brisk-transient (Y), and dog beta cells brisk-sustained (X) concentric receptive fields. From the smallest beta cell sizes it is concluded that the visual acuity of the dog may be as good as that of the cat.

Animals↗

Location of retinal ganglion cells contributing to the early imprecision in the retinotopic order of the developing projection to the superior colliculus of the wallaby (Macropus eugenii).

The position of ganglion cells contributing to the early imprecision in retinotopic order in the developing retinocollicular projection in the wallaby (Macropus eugenii) has been determined. Deposits of horseradish peroxidase conjugated to wheat germ agglutinin (WGA-HRP) were made in the caudal pole of the superior colliculus (SC) at ages ranging from 22 days after birth, when sparse retinal axons have only just reached the caudal pole of the SC and are yet to cover its surface completely, to 96 days when the retinotopy of ganglion cell terminals in the SC is precise (Marotte, '90). From 30 days onwards, the deposit of WGA-HRP resulted in a dense patch of retrogradely labelled retinal ganglion cells that could be seen to be appropriately positioned in nasal retina. However, at all ages prior to 92 days, there were inappropriately positioned labelled cells between the densely labelled patch and the central retina and both dorsal and ventral to the patch. They were not found in far distant regions of retina and composed a relatively small proportion of labelled cells. They reached a peak at 45 days, had decreased to low levels by 63 days, were rare by 81 days, and by 92 days were absent. This latter age fits with the time when retinotopy was judged to be precise in a previous study (Marotte, '90). Inappropriately projecting cells never originate from the entire retina but only from regions adjacent to the appropriate region. Thus, during development there are no gross projection errors. Initially, ganglion cell axons are distributed on the colliculus in a coarse retinotopy. Refinement of the projection then follows, revealed by this technique as a loss of inappropriately projecting ganglion cells. This is complete by 92 days well before eye opening at around 140 days.

Aging↗

Central retinal area is not the site where ganglion cells are generated first.

The development of retinal ganglion cells (RGC) was studied in the chick from stage 18 to adulthood. Our main objectives were to identify the retinal site where the first RGCs differentiate, to locate this site relative to the optically defined central retinal area, and to map the spatial arrangement of the RGC field at different stages in development. The eyes of the experimental animals were fixed and serially sectioned. The borders of RGC fields were determined from the presence of either ganglion cell perikarya or ganglion cell axons. In seven cases between stages 21 and 26, the borders of the RGC fields were confirmed electron microscopically. The serial sections together with the RGC fields were then reconstructed in three dimensions. The reconstructed retinae were projected onto a plane by using the radially equidistant polar azimuthal projection. First, RGCs appear dorsal to the apex of the optic fissure. Ganglion cell development then initially spreads out symmetrically with respect to the optic fissure. However, from stage 29 on, the nasal half of the retina expands much more than the temporal half. This asymmetrical growth entails that the optic fissure is eventually located in the temporal half of the retina in the mature animal. The RGC fields of the embryonic stages were superimposed on the retina of a visually active animal according to their real size and position. It turned out that the central retinal area was at least 2 mm away from the site where the first RGCs were generated. It is not before stage 28 that the prospective central retinal area is included into the expanding ganglion cell field. The fact that RGCs at the central retinal area are generated 2.5 days later than first RGCs near the apex of the optic fissure has important implications for the formation of the retinotectal projection.

Animals↗

Ultrastructural and immunocytochemical analysis of the circuitry of two putative directionally selective ganglion cells in turtle retina.

Two well-stained, horseradish peroxidase-filled varieties of putative ON-OFF directionally selective ganglion cells, G14a and G15, that project to the dorsolateral optic tectum (Guiloff and Kolb [1992a] Vis. Neurosci. 8:295-313) were studied qualitatively and quantitatively. Both were bistratified ganglion cells with one tier of dendrites in the OFF sublamina and the other in the ON sublamina of the inner plexiform layer (IPL). The cells were serially sectioned and examined for synaptic inputs by electron microscopy. Portions of the dendritic trees were also analyzed after postembedding immunocytochemistry for neurotransmitter candidates gamma aminobutyric acid (GABA), glycine, choline acetyltransferase (ChAT), and glutamate in presynaptic neurons. Both G14a and G15 are dominated by amacrine cell inputs and have only minor bipolar cell involvement. Probably at least two different types of bipolar cell are presynaptic. Both ganglion cells receive some GABA-positive (GABA+) amacrine inputs and G14a receives ChAT+ amacrine inputs. Glycine+ and glutamate+ inputs could not be detected in either cell. The GABA+ inputs appeared to be regionally arranged in the dendritic trees. The general distribution of amacrine and bipolar inputs to the two tiers of dendrites in both cell types appeared to be asymmetrical, both along the radial extent of the dendritic trees and within the depth of the IPL. Our data support some aspects of the current models for directional selectivity. We suggest candidate bipolar and amacrine cells that could have input to these ganglion cells. Since many of the putative presynaptic amacrine cells coincide with directionally selective types recorded and stained by other authors, we propose that in turtle retina directional selectivity arises in neurons presynaptic to the ganglion cells.

Animals↗