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PubMed1997

Quantitative and topographical study of retinal ganglion cells in the chameleon (Chameleo chameleon).

Chameleons display a number of well-described physiological peculiarities of their visual system, but there is no information on the topography of the retinal ganglion cell layer. In the present study, ganglion cell density of the chameleon retina was constructed from whole mounts of the retina stained with cresyl violet. For the identification of ganglion cells, these latter cells were labelled retrogradely with horseradish peroxidase applied to the optic nerve. Using this criterion, the proportion of ganglion cells was estimated to represent 80% of retinal cells, while glial cells and amacrine cells represented 14 and 6%, respectively, of the total cell population of the retina. As for the main features of the retinal map, first, ganglion cells were distributed inhomogeneously within the ganglion cell layer, and revealed the existence of a putative area centralis. Second, a horizontal visual streak, which showed two peak density areas, was identified. These features point out the degree of specialisation of the chameleon retina and the complexity of its visual system.

Animals↗

PubMedDate not supplied

Neurotransmitter-specific identification and characterization of neurons in the all-cone retina of Anolis carolinensis. II: Glutamate and aspartate.

Immunocytochemical and autoradiographic methods were used to identify neurons in the pure cone retina of the lizard (Anolis carolinensis) that are likely to employ glutamate (GLU) or aspartate (ASP) as a neurotransmitter. GLU immunocytochemistry demonstrated high levels of endogenous GLU in all cone types and numerous bipolar cells. Moderate GLU levels were found in horizontal and ganglion cells. Müller cells and most amacrine cells had very low GLU levels. GLU immunoreactivity (GLU-IR) in the cones was present from the inner segment to the synaptic pedicle. A large spherical cell type with moderate GLU-IR was identified in the proximal inner plexiform layer (IPL). These cells also contain ASP and have been tentatively identified as amacrine cells. Uptake of [3H]-L-GLU labeled all retinal layers. All cone types and Müller cells sequestered [3H]-D-ASP, a substrate specific for the GLU transporter. Anti-ASP labeling was observed in cones, horizontal cells, amacrine cells, and cells in the ganglion cell layer. ASP immunoreactivity (ASP-IR) in the cones was confined to the inner segment. One ASP-containing pyriform amacrine cell subtype ramifying in IPL sublamina b was identified. Analysis of GLU-IR, ASP-IR, and GABA-IR on serial sections indicated that there were two distinct populations of horizontal cells in the Anolis retina: one containing GABA-IR, GLU-IR, and ASP-IR; and another type containing only GLU-IR and ASP-IR. Light GLU-IR was frequently found in GABA-containing amacrine cells but ASP-IR was not. The distinct distributions of GLU and ASP may indicate distinctly different roles for these amino acids. GLU, not ASP, is probably the major neurotransmitter in the cone-bipolar-ganglion cell pathway of the Anolis retina. Both GLU and ASP are present in horizontal cells and specific subpopulations of amacrine cells, but it is unclear if GLU or ASP have a neurotransmitter role in these cells.

Animals↗

PubMed2001

Quantitative analysis of the retinal ganglion cell layer in the ostrich, Struthio camelus.

The total number, distribution and peak density of ganglion cells were evaluated in the Nissl-stained retina of the ostrich (Struthio camelus). The mean (n = 4) total number of retinal ganglion cells (RGC) was estimated at 2,274,128 (s.d. = 273, 152). The ostrich retina exhibited a prominent horizontal visual streak along which a central area located nasal to the pecten had a peak density of 9,500 cells/mm2. A high concentration of cells with a peak density of 2,646 cells/mm2 was also observed in the temporal retina, slightly dorsal to the visual streak. The results further showed that the ostrich eye has a 15-mm pupil entrance diameter, its mean axial length is 39.81 mm, the estimated retinal magnification factor is 0.4075 mm/deg and the maximum visual acuity along the well-defined visual streak was estimated to be 19.32 cycles/deg. The latter component of the retina might subserve vision along the horizon while the temporal region mediates binocular processing. The data also showed that the degree of retinal illumination in this bird could be comparable to that noted in some nocturnal species. The findings in this study suggest that the ostrich might not be restricted to diurnal activity.

Algorithms↗

PubMed1984

Influence of amacrine cells on receptive field organization of ganglion cells of the generalized vertebrate cone retina: electronic simulation.

Two classes of amacrine cells are simulated, small-field and large-field. Small-field amacrine cells are formed by input from a single bipolar cell, while large-field amacrine cell is formed by inputs from same 7 bipolar cells that form the ganglion cell. Only tonic amacrine cells are studied with both chromatic and luminosity types as well as double- and single-opponent receptive fields. Amacrine cells are used in both feedforward to ganglion cells and feedback to bipolar and horizontal cells. Feedback to bipolar cells or feedforward to ganglion cells affected steady state levels in a predictable fashion. Negative feedback to bipolar cells and positive feedforward to ganglion cells does not introduce transients to ganglion cells while negative feedback to horizontal cells and negative feedforward does. Feedback to horizontal cells produces complex effects on bipolar, amacrine and ganglion cells dependent on such factors as center-surround field balance and negative feedback from luminosity type of horizontal cell to cones.

Animals↗

PubMed2001

Characterization of a transformed rat retinal ganglion cell line.

The purpose of the present study was to establish a rat retinal ganglion cell line by transformation of rat retinal cells. For this investigation, retinal cells were isolated from postnatal day 1 (PN1) rats and transformed with the psi2 E1A virus. In order to isolate retinal ganglion cells (RGC), single cell clones were chosen at random from the transformed cells. Expression of Thy-1 (a marker for RGC), glial fibrillary acidic protein (GFAP, a positive marker for Muller cells), HPC-1/syntaxin (a marker for amacrine cells), 8A1 (a marker for horizontal and ganglion cells) and neurotrophins was studied using reverse transcriptase-polymerase chain reaction (RT-PCR), immunoblotting and immunocytochemistry. One of the retinal cell clones, designated RGC-5, was positive for Thy-1, Brn-3C, Neuritin, NMDA receptor, GABA-B receptor, and synaptophysin expression and negative for GFAP, HPC-1, and 8A1, suggesting that it represented a putative RGC clone. The results of RT-PCR analysis were confirmed by immunocytochemistry for Thy-1 and GFAP. Upon further characterization by immunoblotting, the RGC-5 clone was positive for Thy-1, negative for GFAP, 8A1 and syntaxin. RGC 5 cells were also positive for the expression of neurotrophins and their cognate receptors. To establish the physiological relevance of RGC-5, the effects of serum/trophic factor deprivation and glutamate toxicity were analyzed to determine if these cells would undergo apoptosis. The protective effects of neurotrophins on RGC-5 after serum deprivation was also investigated. Apoptosis was studied by terminal deoxynucleotidyl transferase-mediated fluoresceinated dUTP nick end labeling (TUNEL). Serum deprivation resulted in apoptosis and supplementation with both BDNF and NT-4 in the growth media, protected the RGC-5 cells from undergoing apoptosis. On differentiation with succinyl concanavalin A (sConA), RGC-5 cells became sensitive to glutamate toxicity, which could be reversed by inclusion of ciplizone (MK801). In conclusion, a transformed rat retinal cell line, RGC-5, has certain characteristics of retinal ganglion cells based on Thy-1 and Brn-3C expression and its sensitivity to glutamate excitotoxicity and neurotrophin withdrawal. These cells may be valuable in understanding of retinal ganglion cell biology and physiology including in vitro manipulations in experimental models of glaucoma.

Animals↗

PubMed2005

Infection of retinal neurons during murine cytomegalovirus retinitis.

PURPOSE: Previous results suggest that retinal neurons are infected early during murine cytomegalovirus (MCMV) infection of the inner retina. The purposes of this study were to identify which retinal neurons are infected and to determine the routes by which MCMV spreads in the retina. METHODS: Immunosuppressed (IS) BALB/c mice were inoculated with 5 x 10(3) PFU of MCMV (k181) through the supraciliary route. Injected eyes were collected at several times after inoculation, sectioned, and examined by electron microscopy and by staining for retinal cell antigens and for MCMV early (EA) or late (LA) antigen. RESULTS: MCMV-infected cells were observed in the choroid and RPE by day 3 after infection (PI) and in the inner retina beginning at day 5 PI. At this time, many horizontal and bipolar cells were MCMV-antigen-positive but only rare MCMV-infected amacrine cells (glycine positive or gamma-aminobutyric acid [GABA] positive) or MCMV-infected ganglion cells (NF positive) were observed in the inner retina. At day 10 PI, most virus-infected cells were glial fibrillary acidic protein (GFAP)- and GABA-positive glia. Virions were observed by electron microscopy in the choroid, RPE, and inner nuclear layer of the retina. Although virions were observed in the endothelium of the retinal vessels and the nearby retinal cells, the endothelial cell lining of the retinal vessels remained intact. Both apoptotic cells and necrotic cells were seen in the inner retina. CONCLUSIONS: In the inner retina, horizontal and bipolar cells were the early (< or = day 7 PI) targets of MCMV infection. Virus spread from the RPE and the photoreceptor layer to the inner retina through infected Muller cells and within the inner retina horizontally through infected horizontal cells.

Animals↗

PubMed1998

The embryogenesis of rod photoreceptors in the teleost fish retina, Haplochromis burtoni.

Development of the retina, like that of other tissues, occurs via an orderly sequence of cell division and differentiation, producing the functional retina. In teleost fish, however, cell division and differentiation in the retina continue throughout the life of the animal in two distinct ways. Stem cells in a circumferential germinal zone at the periphery of the retina give rise to all retinal cell types and progenitor cells located throughout the retina in the outer nuclear layer (ONL) produce new rod photoreceptors. These processes in adult retina recapitulate in space the embryonic events responsible for forming the retina. Analysis of these events in an African cichlid fish, Haplochromis burtoni, confirmed that cone photoreceptors differentiate first, followed by rod photoreceptors. Correspondingly, at the margin of the eye, cone photoreceptors differentiate nearer to the margin than do rods. Control of photoreceptor production is not understood. Here we present the time of appearance and distribution pattern of GABA and vimentin which are candidates for the control of retinal cell division and differentiation. Antibody staining reveals that both GABA and vimentin exhibit unique patterns of expression during embryonic retinal development. Vimentin immunoreactivity is evident throughout the retina in a spoke-like pattern between developmental Days 4 and 7, as both cone and rod photoreceptors are being formed. GABA is expressed in horizontal cells between Days 5 and 7, corresponding to the onset of rod differentiation in time and in position within the retina. Moreover, the wave of GABAergic staining in the horizontal cells parallels the wave of rod differentiation across the embryonic retina of H. burtoni. Thus, GABA may play a role in the development of rod photoreceptors.

Animals↗

PubMed1992

Translaminar deficits in the retinae of albinos.

The central retina in albino mammals is poorly developed. There is a general reduction in ganglion cell density compared with the normal animal, and there are irregularities in the center to periphery gradient in ganglion cell density. It is not known whether, and if so to what extent, this abnormality is associated with deficits in other retinal layers. In this study the distribution of cells in the ganglion cell layer has been determined in horizontally sectioned eyes from pigmented and albino ferrets. This was undertaken to define the location of the region of highest cell density and the relative cell gradients around it. Detailed counts and measurements were then undertaken to determine the cell density within, and thickness of, the inner and the outer nuclear layers in these animals. All the albino animals had an abnormal distribution of cells in the ganglion cell layer in the central retina when compared with pigmented animals. The extent of this abnormality was variable. No differences between pigmented and albino animals could be found in the packing density of cells within the inner or the outer nuclear layer. However, in the pigmented animals there was a clear increase in the thickness of these layers associated with the region of highest density in the ganglion cell layer. This feature was absent in the albino animals, where the gradient in layer thickness was less marked and frequently contained irregularities. These abnormalities were most obvious in the outer nuclear layer.

Albinism↗

PubMed1985

Effect of excitatory amino acids on gamma-aminobutyric acid release from frog horizontal cells.

The effects of excitatory amino acids, analogues and K on [3H]gamma-aminobutyric acid [3H]GABA) release from horizontal cells of the isolated superfused frog retina were studied. Exposure of the retina to medium containing high concentrations (25-100 mM) of KCl increased the release of [3H]GABA to a maximum which was 40 times the spontaneous resting release. The K-evoked release of [3H]GABA was almost abolished in high-Mg/low-Ca medium. Glutamate, aspartate, kainate and quisqualate also stimulated the release of [3H]GABA from horizontal cells, the maximum evoked release being similar to that produced by KCl. The release of [3H]GABA evoked by glutamate, aspartate, kainate and quisqualate was abolished in high-Mg/low-Ca medium and by Na-free medium. The evoked releases of [3H]GABA were not reduced by tetrodotoxin. N-Methyl-D-aspartate (NMDA) at concentrations up to 10 mM had virtually no effect on [3H]GABA release from horizontal cells. In Mg-free medium, NMDA stimulated [3H]GABA release, but the maximum release was only 10% of that produced by other agonists. Mg-free medium did not significantly affect the evoked release of [3H]GABA by other agonists. NMDA apparently possessed affinity for the kainate receptor, because in normal medium it antagonized the effects of kainate but not glutamate, aspartate or quisqualate. The non-selective antagonist of excitatory amino acids, (+/-)-cis-2,3-piperidine dicarboxylic acid (PDA) antagonized the action of glutamate, aspartate, kainate and quisqualate on horizontal cell [3H]GABA release. D(-)-2-Amino-4-phosphonobutyrate (APB) and D-gamma-glutamylglycine (D-gamma-GG) antagonized the actions of kainate on horizontal cell [3H]GABA release at concentrations which had little affect on quisqualate-evoked responses. Approximate estimates of pA2 values (Schild, 1947) showed that the specificity and potency of the antagonists was low. Nevertheless, the retinal 'non-NMDA' receptors can probably be subdivided into kainate and quisqualate types. Glutamate diethylester (GDEE) did not affect the action of any agonist. We conclude that glutamate (and aspartate) probably stimulate the release of [3H]GABA from frog horizontal cells by activating receptors of the non-NMDA type. This activation may trigger the opening of tetrodotoxin-insensitive Na channels, resulting in the depolarization of the cell membrane and an increase in the conductance of voltage-sensitive Ca-channels. An influx of Ca ions would then trigger the release of [3H]GABA. Our results are not consistent with previous suggestions that GABA release from horizontal cells involves an outwardly directed transport process.

Amino Acids↗

PubMed2003

Cell diversity in the retina: more than meets the eye.

Over 10 years ago, Pax-6 was shown to play an evolutionarily conserved role in controlling eye formation from Drosophila to humans.1 Since then, the identification of an entire cascade of conserved eye determination genes has brought a new understanding to the developmental relationship between the insect compound eye and the vertebrate camera eye.2 Additional studies are now beginning to suggest that even late aspects of eye development, including cell type specification, also share common molecular machinery. In this commentary, I will discuss some of these findings, with a particular focus on the recent study by Dyer et al.3 describing a novel role for the Prox1 transcription factor in specifying horizontal cells in the mouse retina. As Prospero, the Drosophila homolog of Prox1, also participates in retinal cell specification, these data provide a forum for asking new questions concerning pathways that may regulate retinogenesis across evolution.

Animals↗

PubMed1988

Synaptic transmission from rods to horizontal cells in dark-adapted tiger salamander retina.

Synaptic transmission between rods and horizontal cells (HCs) was studied by simultaneously recording from the pre- and postsynaptic cells in the living retinal slices of the larval tiger salamander. Voltage tails after the termination of a flash were observed in dark-adapted rods and HC somata, but not in cones. These simultaneously recorded voltage tails were used to isolate the rod input from the cone input to the HCs. The rod voltage versus the HC voltage in the response tails yielded the input-output relation of the rod-HC synapse. Two factors, a luminance-dependent signal delay between the rod and HC responses and a voltage-dependent depolarizing current in the HCs, complicate the analysis of the rod-HC synapse. A single input-output relation of the rod-HC synapse was obtained after these two factors had been corrected. The rod-HC synapse was approximately linear near the rod dark potential (-40 mV) with a synaptic gain around 12, but it became progressively nonlinear at more hyperpolarized potentials. The synaptic gain at Vrod = -50 mV was about 0.65.

Ambystoma↗

PubMed1998

GABA as a developmental neurotransmitter in the outer plexiform layer of the vertebrate retina.

GABA is present in certain retinal neurons before synapses are formed, and it has a variety of effects on the developing retina. Its exact role in retinal maturation is not clear; however, there is growing evidence from rabbit retina that neonatal horizontal cells produce GABA within the outer retina, which in turn is necessary for normal synapse formation of photoreceptors. In addition to its classic role as an inhibitory transmitter, GABA in the neonate increases intracellular calcium levels in selected cells. This latter property is lost during postnatal development, along with a general decrease in the level of expression of both pre- and postsynaptic GABAergic markers and a decrease in the number of GABAergic neurons. The adult GABAergic circuitry in adult retina may represent a restricted, perhaps simplified version of the more complex and diversified interactions of the GABA system during development.

Animals↗

PubMed1984

Electrical coupling between horizontal cell bodies in the tiger salamander retina.

We studied lateral interactions between horizontal cells in the tiger salamander retina in order to determine the underlying mode of synaptic transmission. Pairs of cells, visually identified by inspection of the in vitro retinal slices, were impaled with two separate electrodes and the transmission between them was examined by injecting current into one cell while recording the resulting voltage response from the other soma. It was found that coupling between horizontal cells remained intact and even increased somewhat during the light response and in the presence of cobalt or acetylcholine. It was also observed that a fluorescent dye injected into one cell would often migrate and stain neighbouring horizontal cells. These results suggest that synaptic transmission between horizontal cell bodies is mediated mainly by electrotonic conduction. The increase in apparent coupling observed under certain experimental conditions can be explained by the increase in input resistance of each horizontal cell.

Acetylcholine↗

PubMed2005

The final fates of neurogenin2-expressing cells include all major neuron types in the mouse retina.

The molecular mechanism underlying vertebrate retinal development is not well understood. To examine whether neurogenin2 (ngn2) expression determines cell fate in the retina, we mapped the final fates of cells that once expressed ngn2, using the conditional, binary CreER -LacZ system. We found LacZ+ cells in all 3 nuclear layers of the mouse retina and including all major types of neurons: photoreceptors, horizontal, bipolar, amacrine, and ganglion cells. The distribution of LacZ+ cells among the 3 nuclear layers closely resembled a theoretical distribution of total retinal cells. The temporal window in which each cell type was marked appeared nonrandom, and was similar to its birthdate and birth sequence. These data indicate that cells expressing ngn2 at some point in their life histories may later differentiate into a variety of cell types.

Amacrine Cells↗

PubMedDate not supplied

Retinal neurotransmission.

The mammalian retina is classically divided into ten layers which contain the neuronal elements identified as photoreceptors, horizontal cells, bipolar cells, amacrine cells and ganglion cells. Using various neuroscientific techniques possible neurotransmitter substances have been assigned to each of these cell types. Thus the localization of transmitter synthesizing enzymes and storage vesicles, the demonstration of release of transmitter in response to specific stimuli, the observation of post-synaptic events mimicked or blocked by the iontophoretic application of exogenous transmitter/agonist or antagonist drug respectively, and the identification of efficient transmitter inactivation mechanisms synaptically add evidence for the association of certain proposed transmitter substances with specific neuronal elements. The evidence for the proposal that the excitatory amino acids glutamate and aspartate are transmitters of photoreceptors, that gamma-aminobutyric acid (GABA) is the inhibitory transmitter of horizontal and amacrine cells, that acetylcholine is associated with the functioning of bipolar cells, and that taurine, glycine and dopamine may all also play neurotransmitter or neuromodulatory roles at amacrine cell synapses is discussed.

Acetylcholine↗

PubMed2000

The SCG10-related gene family in the developing rat retina: persistent expression of SCLIP and stathmin in mature ganglion cell layer.

Neuronal growth-associated proteins (GAPs), such as GAP-43 and SCG10, are thought to play crucial roles in both axonal and dendritic outgrowth during neural development and regeneration, although the underlying mechanisms remain largely unknown. The recent finding that SCG10 is a microtubule regulator and also the identification of RB3 and SCLIP as two new SCG10-related members prompted us to investigate the roles of SCG10-related family in neural development, using the retina as a model system. We determined the temporal expression and the spatial distribution of SCG10-related mRNAs in the developing rat retina. Semiquantitative analysis by RT-PCR revealed that in prenatal retina, levels of SCG10 and stathmin mRNAs were higher than those of RB3 and SCLIP. In the postnatal retina, the level of SCLIP increased, whereas the level of RB3 remained low. In situ hybridization revealed that GAP-43 and all of the SCG10-related family mRNAs were present in the retinal ganglion cells (RGCs) at all stages of retinal development, and that stathmin mRNA was present in mitotic neuroblastic cells. Differential expression of SCG10 and other members of the family became more evident as retinal development proceeded; SCG10 and RB3 expression were relatively specific in the RGCs and amacrine cells, whereas SCLIP was also evident in bipolar and horizontal cells. Stathmin mRNA was highly expressed both in the RGCs and other interneurons. These results indicate that multiple SCG10-related proteins are expressed in single neurons including RGCs, and suggest that these nGAPs play similar but distinct roles in differentiation and functional maintenance of retinal neurons.

Animals↗

PubMed1984

Cytochemical localization of 5'-nucleotidase activity in retinal photoreceptor cells.

Further cytochemical studies on the ultrastructural localization of 5'-nucleotidase in the rat retina have revealed activity to be associated with the complex synapses formed by the rod spherules of the receptors and the bipolar and horizontal cell processes. Activity was also seen on the axolemma of receptor fibers. In the rod inner segment strong reaction product is located intracellularly. In the rod outer segment the enzyme appears to be located only on the cytoplasmic side of the disc membrane and not intradiscally . Retinal pigment cells are rich in 5'-nucleotidase. Their microvilli accompany the tips of the receptor cells and show enzyme activity in an ecto position. A role for 5'-nucleotidase is possible in the metabolism of guanylate and adenylate nucleotides both of which are important for visual transduction processes.

5'-Nucleotidase↗

PubMed2006

Circadian organization of the mammalian retina.

The mammalian retina contains an endogenous circadian pacemaker that broadly regulates retinal physiology and function, yet the cellular origin and organization of the mammalian retinal circadian clock remains unclear. Circadian clock neurons generate daily rhythms via cell-autonomous autoregulatory clock gene networks, and, thus, to localize circadian clock neurons within the mammalian retina, we have studied the cell type-specific expression of six core circadian clock genes in individual, identified mouse retinal neurons, as well as characterized the clock gene expression rhythms in photoreceptor degenerate rd mouse retinas. Individual photoreceptors, horizontal, bipolar, dopaminergic (DA) amacrines, catecholaminergic (CA) amacrines, and ganglion neurons were identified either by morphology or by a tyrosine hydroxylase (TH) promoter-driven red fluorescent protein (RFP) fluorescent reporter. Cells were collected, and their transcriptomes were subjected to multiplex single-cell RT-PCR for the core clock genes Period (Per) 1 and 2, Cryptochrome (Cry) 1 and 2, Clock, and Bmal1. Individual horizontal, bipolar, DA, CA, and ganglion neurons, but not photoreceptors, were found to coordinately express all six core clock genes, with the lowest proportion of putative clock cells in photoreceptors (0%) and the highest proportion in DA neurons (30%). In addition, clock gene rhythms were found to persist for >25 days in isolated, cultured rd mouse retinas in which photoreceptors had degenerated. Our results indicate that multiple types of retinal neurons are potential circadian clock neurons that express key elements of the circadian autoregulatory gene network and that the inner nuclear and ganglion cell layers of the mammalian retina contain functionally autonomous circadian clocks.

Animals↗