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Role of the Barhl2 homeobox gene in the specification of glycinergic amacrine cells.

The mammalian retina contains numerous morphological and physiological subtypes of amacrine cells necessary for integrating and modulating visual signals presented to the output neurons. Among subtypes of amacrine cells grouped by neurotransmitter phenotypes, the glycinergic and gamma-aminobutyric acid (GABA)ergic amacrine cells constitute two major subpopulations. To date, the molecular mechanisms governing the specification of subtype identity of amacrine cells remain elusive. We report here that during mouse development, the Barhl2 homeobox gene displays an expression pattern in the nervous system that is distinct from that of its homologue Barhl1. In the developing retina, Barhl2 expression is found in postmitotic amacrine, horizontal and ganglion cells, while Barhl1 expression is absent. Forced expression of Barhl2 in retinal progenitors promotes the differentiation of glycinergic amacrine cells, whereas a dominant-negative form of Barhl2 has the opposite effect. By contrast, they exert no effect on the formation of GABAergic neurons. Moreover, misexpressed Barhl2 inhibits the formation of bipolar and Müller glial cells, indicating that Barhl2 is able to function both as a positive and negative regulator, depending on different types of cells. Taken together, our data suggest that Barhl2 may function to specify the identity of glycinergic amacrine cells from competent progenitors during retinogenesis.

Amacrine Cells↗

Colocalization of retinal dystrophin and actin in postsynaptic dendrites of rod and cone photoreceptor synapses.

In this paper we demonstrate immunostaining specific for dystrophin in photoreceptor synapses of human, bovine and rat retinas. Cryosections of retinas incubated with dystrophin-specific monoclonal antibodies displayed a punctuate staining pattern in the outer plexiform layer. This pattern resulted from binding of the antibodies to synaptic complexes of both rods and cones, shown by double-labelling with antibodies to either synaptophysin or actin. Confocal laser fluorescence microscopy demonstrated that dystrophin staining colocalized predominantly with actin, which is concentrated in the postsynaptic portions of the synaptic complex. No significant dystrophin immunolabel was seen in the presynaptic terminals labelled with antibodies to synaptophysin, a marker of synaptic vesicles. Immunoblot analysis confirmed the presence of approximately 420 kDa and approximately 360 kDa dystrophin-like polypeptide bands associated with membranes of the bovine retina. We speculate that retinal dystrophin is involved in the linkage of actin filaments to the postsynaptic plasma membrane. Such a linkage may be important for the generation of synaptic microdomains and for certain phenomena of synaptic plasticity. The absence of dystrophin in patients suffering from Duchenne's muscular dystrophy is accompanied by visual problems and abnormalities of the electroretinogram. Therefore it is likely that retinal dystrophin plays a role in certain stages of synaptic transmission between photoreceptors and the postsynaptic dendritic complex formed by horizontal and bipolar cells.

Actins↗

The distribution and developmental regulation of NMDA receptor subunit proteins in the outer and inner retina of the rat.

In order to investigate whether N-methyl-D-aspartate (NMDA) receptors with distinct pharmacological properties are differentially distributed within the retinal layers, the spatial distribution and temporal regulation of all NMDA receptor subunits was analyzed in parallel on the protein level in the rat retina during development. Immunohistochemistry was performed on retinal sections at different developmental ages between embryonic (E) days 20/21 and the adult stage using specific antibodies against NMDA subunits (NR1, NR2A-D). All NMDA subunits were expressed in the rat retina postnatally but showed different spatial patterns. In particular, and in contrast to previous in situ hybridization studies, labeling of NR2 subunits was observed in horizontal cell bodies and in the outer plexiform layer, indicating that functional NMDA receptors are expressed in this retinal cell type in the rat. Expression of NR2D was restricted to the inner retina and seemed to be involved in neurotransmission within the rod pathway. In the inner plexiform layer (IPL), distinct patterns of labeling were observed for different NMDA subunits. NR1 was found in two bands which can be related to the off- and on-signal pathways, whereas NR2A and NR2B were located in two bands within the off-sublaminae of the IPL. The antibody against NR2C was distributed throughout the whole IPL, and NR2D was expressed exclusively in the innermost part of the IPL where rod bipolar cell terminals terminate. Distinct bands of immunoreactivity in the IPL were observed only from P14 on. In conclusion, there are clear differences in the spatial distribution and temporal expression of NMDA receptor subtypes in the rodent retina. This indicates that specific retinal cells selectively express glutamate receptors composed of different subunit combinations and thus display different pharmacological and kinetic properties.

Animals↗

GABA-induced increases in [Ca2+]i in retinal neurons of postnatal rabbits.

Previous studies have indicated that gamma-aminobutyric acid (GABA) plays an important trophic role in the synapse formation between horizontal cells and photoreceptors in postnatal rabbit retina. However, the mechanism of the GABA effect has not been identified. Using fluo-3 Ca2+ imaging and confocal laser scanning microscopy we examined the effect of GABA on [Ca2+]i during postnatal retinal development. GABA (100 microM) evoked a fast and transient increase of [Ca2+]i in selected populations of freshly dissociated retinal cells from postnatal rabbits. This increase was apparent on postnatal day 1 and reached a maximum on day 5. Little increase in [Ca2+]i was observed in retinal cells isolated from adult rabbits. GABA receptor antagonists, picrotoxin and bicuculline, significantly reduced the response. The GABAB agonist, baclofen, did not evoke any [Ca2+]i changes. The GABA-induced increase in [Ca2+]i was observed in all retinal layers in neonatal retinal whole-mount explants. In the outer retina, the increase was seen in cone photoreceptors which were specifically labeled with peanut agglutinin (PNA). The GABA-induced increase in [Ca2+]i may provide an important mechanism for regulating cone synaptogenesis in the outer plexiform layer of the postnatal retina.

Animals↗

Sources and sink of retinoic acid in the embryonic chick retina: distribution of aldehyde dehydrogenase activities, CRABP-I, and sites of retinoic acid inactivation.

Previous experiments in mice and zebrafish led to the hypothesis that an asymmetric distribution of the transcriptional activator retinoic acid (RA) causes ventral-dorsal polarity in the vertebrate eye anlage. A high concentration of RA in the ventral retinal neuroepithelium has been suggested to induce developmental events that finally establish topographic order in the retinotectal projection along the vertical eye axis. In the present study we have investigated potential sources and sinks of RA during embryonic development of the chick retina. At embryonic day (E)1 to E2, when the spatial determination of the eye primordia takes place, no RA synthesis by aldehyde dehydrogenases was detectable, and neither immunoreactivity for retinaldehyde dehydrogenase RALDH-2 nor for cellular retinoic acid binding protein CRABP-I was observed. These components of RA signal transduction appeared in the eye between E3 and E5. At later stages, RA-measurements with a reporter cell line showed highest synthesis in the retinal pigment epithelium (RPE) and at the ventral and dorsal poles of the retina. RA degradation occurred mostly in a horizontal region in the middle of the retina with only small differences along the nasal-temporal axis. CRABP-I immunoreactivity appeared first in differentiating retinal ganglion cells with no indication of a spatial gradient across the ventral-dorsal eye axis. RA-production depended on three NAD+-dependent enzyme activities, which could be competitively inhibited by citral. One enzyme, located in the dorsal retina (corresponding to mouse RALDH-1), and one enzyme in the RPE (RALDH-2) were aldehyde dehydrogenases of the same molecular weight (monomers about 55 kDa) but with different isoelectric points (6.5-6.9; 4.9-5.4). The third RA-synthesizing activity (pI 6.0-6.3) was limited to the ventral retina, and likely corresponded to mouse RALDH-3. The restricted localization of retinoid-metabolizing activities along the dorsal-ventral axis of the embryonic chick retina does support the idea that RA is involved in dorsal-ventral eye patterning. However, the late time of appearance of aldehyde dehydrogenase activities and CRABP-I points to functions in cellular differentiation that are distinct from the initiation of the dorsal-ventral polarity.

Aldehyde Dehydrogenase↗

Gain of rod to horizontal cell synaptic transfer: relation to glutamate release and a dihydropyridine-sensitive calcium current.

We related rod to horizontal cell synaptic transfer to glutamate release by rods. Simultaneous intracellular records were obtained from dark-adapted rod-horizontal cell pairs. Steady-state synaptic gain (defined as the ratio of horizontal cell voltage to rod voltage evoked by the same light stimulus) was 3.35 +/- 0.60 for dim flashes and 1.50 +/- 0.03 for bright flashes. Under conditions of maintained illumination, there was a measurable increment of horizontal cell hyperpolarization for each light-induced increment of rod hyperpolarization over the full range of rod voltages. In separate experiments we studied glutamate release from an intact, light-responsive photoreceptor layer, from which inner retinal layers were removed. Steady light reduced glutamate release as a monotonic function of intensity; spectral sensitivity measures indicated that we monitored glutamate release from rods. The dependence of glutamate release on rod voltage was well fit by the activation function for a high-voltage-activated, dihydropyridine-sensitive L-type calcium current, suggesting a linear dependence of glutamate release on [Ca]i in the synaptic terminal. A simple model incorporating this assumption accounts for the steady-state gain of the rod to horizontal cell synapse.

Animals↗

H1 horizontal cells of carp retina have different postsynaptic mechanisms to mediate short- versus long-wavelength visual signals.

Vertebrate photoreceptors release neurotransmitter substance(s) tonically in the dark and this release is curtailed by light. Recently, we have become increasingly aware of the possibility that short- and long-wavelength visual signals are mediated differently during the synaptic transmission to second-order retinal neurons. The experiment described here advances this notion further by demonstrating a postsynaptic difference. Treatment of the carp retina by dopamine reduced the gap-junctional coupling of horizontal cells, and we made use of this known effect to measure the input resistance (Rin) of H1-type horizontal cells. Flashes of light increased Rin. This increase, however, was found to be smaller with short wavelengths, even though the comparison was made when voltage responses were equal in amplitude. Often, Rin was even found to decrease at the blue end of spectrum. No single postsynaptic mechanism can account for any equal-voltage Rin difference such as this. The synaptic spectral segregation thus revealed is probably subserved by a dual scheme wherein the transmitter from blue-sensitive cone photoreceptors acts to decrease the membrane conductance of H1 cells whereas the synapses made by red- and green-sensitive cones are of a classical excitatory type.

Animals↗

Absence of circadian clock regulation of horizontal cell gap junctional coupling reveals two dopamine systems in the goldfish retina.

In fish and other vertebrate retinas, although dopamine release is regulated by both light and an endogenous circadian (24-hour) clock, light increases dopamine release to a greater extent than the clock. The clock increases dopamine release during the subjective day so that D2-like receptors are activated. It is not known, however, whether the retinal clock also activates D1 receptors, which display a much lower sensitivity to dopamine in intact tissue. Because activation of the D1 receptors on fish cone horizontal (H1) cells uncouples the gap junctions between the cells, we studied whether the clock regulates the extent of biocytin tracer coupling in the goldfish retina. Tracer coupling between H1 cells was extensive under dark-adapted conditions (low scotopic range) and similar in the subjective day, subjective night, day, and night. An average of approximately 180 cells were coupled in each dark-adapted condition. However, bright light stimulation or application of the D1 agonist SKF38393 (10 microM) dramatically reduced H1 cell coupling. The D2 agonist quinpirole (1 microM) or application of the D1 antagonist SCH23390 (10 microM) and/or the D2 antagonist spiperone (10 microM) had no effect on H1 cell coupling in dark-adapted retinas. These observations demonstrate that H1 cell gap junctional coupling and thus D1 receptor activity are not affected by endogenous dopamine under dark-adapted conditions. The results suggest that two different dopamine systems are present in the goldfish retina. One system is controlled by an endogenous clock that activates low threshold D2-like receptors in the day, whereas the second system is controlled by light and involves activation of higher threshold D1 receptors.

Adaptation, Ocular↗

Responses of bipolar cells in the retina of the turtle.

The responses of bipolar cells in the retina of the turtle have been studied by intracellular recording. Two types of bipolar cell have been identified: one gave graded depolarizing and the other graded hyperpolarizing responses to small circles of light (100 mum diameter). The responses of both types of cell were similar in the following respects.1. Both were extremely sensitive to dim light; the amplitude of response to a small circle of light increased with light intensity more steeply than the cone response.2. Enlarging the diameter of a spot added an antagonistic effect which decreased response amplitude. This decrease in response amplitude was more apparent at dim than at bright light. Stimulating only distant areas of retina with an annulus produced a response of polarity opposite to that normally produced by a central spot. However, the responses of bipolar cells did not appear to be due to a simple summation of opposite polarity signals contributed from central and peripheral parts of their receptive fields.3. When small spots or annuli of light were turned off there frequently occurred an overshooting OFF transient. The occurrence of OFF transients depended on the duration of the stimulus. Cones recorded under similar conditions produced an OFF depolarization. The size of cone OFF depolarizations increased with increasing duration of the preceding light; following approximately 3 sec of illumination their maximum amplitude was roughly 1/10 the amplitude of the preceding hyperpolarization. The size of OFF responses in both cone and bipolar cells was increased when horizontal cells were hyperpolarized by light. It is concluded that bipolar cells produce large responses for very small cone responses, and, as a consequence, a small depolarization in cones following illumination produces large OFF transients in bipolar cells. Furthermore, the responses of bipolar cells do not appear to represent a simple summation of opposite polarity input from receptor and horizontal cells.

Animals↗

Voltage-dependent sodium channel alpha subunit immunoreactivity is expressed by distinct cell types of the cat and monkey retina.

Polyclonal (7493 and 7317) and monoclonal (mAb3) antibodies, generated to the alpha subunit of the voltage-gated sodium channel (alpha NaCh), were employed to assess the cell types containing alpha NaCh-like immunoreactivity in the mature cat and monkey retina. Immunoblot analyses of retinal proteins in the cat revealed that the polyclonal and monoclonal antibodies we employed labeled a band in the 260-kDa region which corresponds to the molecular mass of the alpha subunit of the NaCh. In both the cat and monkey, these antibodies immunolabeled several distinct types of retinal cells. With the polyclonal antibodies immunoreactivity was observed in ganglion cells and their intraretinal axons, in horizontal cells, and unexpectedly, in cones. In addition, in both species, a limited number of heavily labeled profiles, presumed to be bipolar cells, were seen in the inner nuclear layer. In cat and monkey the monoclonal antibody labeled axons in the fiber layer, ganglion cell somata, and a continuous band of immunoreactive cell bodies (presumed bipolar cells) situated in the outer half of the inner nuclear layer. By immunolabeling isolated cells dissociated from the cat retina, it was possible to demonstrate unequivocally that a population of bipolar cells was labeled by the monoclonal and the polyclonal antibodies we employed. The differences in the labeling observed with the monoclonal antibody as compared to the polyclonal antibodies were interpreted as reflecting the presence of different alpha-subunit subtypes in the mammalian retina. Collectively, our findings suggest that alpha NaCh-like proteins are expressed by a more diverse population of retinal cells than expected on the basis of previous physiological and immunohistochemical studies.

Animals↗

Generalized progressive retinal atrophy in two Akita dogs.

Two distantly related, two- to three-year-old male Akita dogs developed partial or complete night blindness which progressed to day blindness. Clinical features were attenuated retinal blood vessels, diffuse exaggerated tapetal reflex, and hyperreflective horizontal lines in the tapetal fundus. The principal histologic change was regional photoreceptor cell degeneration. Areas of moderately atrophic sensory retina alternated, concentrically to the papilla, with areas of severely atrophic retina. Ultrastructurally, there was disorganization of photoreceptor outer segments, retinal gliosis, focal loss with reorganization of the outer limiting membrane, and retinal invasion by pigment epithelium-derived macrophages. Clinical observations on five additional Akitas with generalized retinal atrophy suggest that the condition is heritable.

Animals↗

Complexity and scaling properties of amacrine, ganglion, horizontal, and bipolar cells in the turtle retina.

In the present study we have evaluated the complexity and scaling properties of the morphology of retinal neurons using fractal dimension as a quantitative parameter. We examined a large number of cells from Pseudemys scripta and Mauremys caspica turtles that had been labeled using Golgi-impregnation techniques, intracellular injection of Lucifer Yellow followed by photooxidation, intracellular injection of rhodamine conjugated horseradish peroxidase, or intracellular injection of Lucifer Yellow or horseradish peroxidase alone. The fractal dimensions of two-dimensional projections of the cells were calculated using a box counting method. Discriminant analysis revealed fractal dimension to be a significant classification parameter among several other parameters typically used for placing turtle retinal neurons in different cell classes. The fractal dimension of amacrine cells was significantly correlated with dendritic field diameters, while the fractal dimensions of ganglion cells did not vary with dendritic field span. There were no significant differences between the same cell types in two different turtle species, or between the same types of neurons in the same species after labeling with different techniques. The application of fractal dimension, as a quantitative measure of complexity and scaling properties and as a classification criterion of neuronal types, appears to be useful and may have wide applicability to other parts of the central nervous system.

Animals↗

Golgi studies of the first optic ganglion of the ant, Cataglyphis bicolor.

The neurons of the first optic ganglion (the lamina) in the desert ant, Cataglyphis bicolor, have been studied with the light microscope after Golgi silver impregnation. The different types of retinal and laminal fibres and their configuration are compared with the results obtained in the bee. The first synaptic region in the visual system of the ant lies proximally to the fenestrated layer below the basement membrane and the layer containing the monopolar cell bodies. The synaptic region can be separated into three morphologically different zones: (1) The most distal layer where the short visual fibres end at two different levels. The short visual fibres and some laminal fibres (monopolar cell fibres) also show lateral elements in this region. (2) The second layer appears almost free of branches of retinal and laminal fibres. (3) The most proximal layer, which has a characteristically dense horizontal structure resulting from the lateral elements of long visual, centrifugal, monopolar and tangential fibres. Nine cell axons arising from each ommatidium leave the retina. Six of these are short visual fibres and end at two different levels in the lamina. Three different types of short visual fibres can be distinguished by their different terminal depths and lateral branching pattern. The remaining three fibres, the long visual fibres, terminate in the medulla. They can be distinguished from each other by their lateral elements in the lamina neuropile. The five morphologically different laminal fibre types (axons of the monopolar cells in the lamina) have different shapes and different arborizations at different levels. Tangential, centrifugal and incerta sedis-fibres, which originate either from cell bodies in the cell body layer at the periphery of the outer chiasma or more centrally, terminate in the synaptic region of the lamina. Consideration is given to the clearly demarkated arrangement and length of the branching pattern of retinal and laminal fibres at different levels of the synaptic region of the lamina. In addition, a hypothetical connectivity pattern is discussed.

Animals↗

Ultracytochemical distribution of ouabain-sensitive, K+-dependent, p-nitrophenylphosphatase in the synaptic layers of goldfish retina.

Ouabain-sensitive, K+-dependent p-nitrophenylphosphatase (K+-pNPPase) activity, which represents the second dephosphorylation step of Na+,K+-ATPase, was localized histochemically at the light and electron microscopical levels in the goldfish retina. K+-pNPPase staining was most intense in the outer and inner plexiform layers and less intense over the photoreceptor inner segments. K+-pNPPase staining was observed on the membranes of horizontal cell dendrites and presynaptic membrane of all cone pedicles but only rarely over rod spherules. Bipolar cell dendrites in the outer plexiform layer were not stained for K+-pNPPase. In the inner plexiform layer (IPL), K+-pNPPase staining was observed at 90% of the bipolar cell ribbon synapses but only at 40% of amacrine cell synapses. The proportion of K+-pNPPase staining at amacrine cell synapses increased from 26 to 49% as one progressed from the outer to inner layers of the IPL, while staining at bipolar cell synapses showed no such trend. Only 16% of the amacrine synapses onto mixed, rod-cone (mb) bipolar cell synaptic terminals were positive for K+-pNPPase. We suggest that the differential distribution of K+-pNPPase staining at retinal synapses can be explained, in part, by the ionic conductances gated at the postsynaptic sites. In addition, the presence of K+-pNPPase on lateral horizontal cell dendrites in cone pedicles is consistent with the hypothesis that the sodium pump is involved in the release of GABA at feedback synapses from horizontal cells to cone photoreceptors.

4-Nitrophenylphosphatase↗

Novel ryanodine-binding properties in mammalian retina.

The ryanodine receptor (RyR)/Ca2+ release channel mobilizes Ca2+ from internal calcium stores to support a variety of neuronal functions. To investigate the presence of such a protein in mammalian retina, we applied ryanodine binding, PCR and antibodies against known RyRs. Surprisingly, ryanodine-binding properties of retinal endoplasmic reticulum-enriched membrane fraction were vastly different from those of skeletal and cardiac muscles ryanodine-binding proteins. In common with the skeletal and cardiac muscle, ryanodine bound with high-affinity to two or more types of binding site (Kd1 = 20.6 and Kd2 = 114 nM); binding was strongly stimulated by high concentrations of NaCl; it was inhibited by tetracaine and the protein appeared to possess an ATP-binding site. Unlike cardiac and skeletal muscle, RyRs in retina binding was Ca2+-independent; inhibited by caffeine and dantrolene; less sensitive to ruthenium red; and unaffected by La3+. Also, in retina, ryanodine rapidly associated to and dissociated from its binding sites. Furthermore, although the protein bound the ATP analog BzATP, retinal ryanodine binding was not stimulated by nucleotides. Immunostaining of bovine retinal sections with anti-RyR2 showed a strong staining of amacrine, horizontal and ganglion cells. Finally, using RT-PCR, the three known RyR isoforms were identified in retina. However, consistent with the novel binding properties, the peptide maps yielded by trypsin treatment and Western blotting demonstrate different patterns. Together, the results suggest that retina expresses a novel ryanodine-binding protein, likely to be a ryanodine receptor. Its presence in retina suggests that this protein might play a role in controlling intracellular Ca2+ concentration.

Animals↗

Overlapping and specific patterns of GDNF, c-ret and GFR alpha mRNA expression in the developing chicken retina.

GDNF and the GDNF receptors, c-Ret, GFR alpha 1 and 2 mRNA is expressed in the developing chicken retina. GDNF labelling was mainly found in embryonic day 4-5 retina but weak labelling could also be found over scattered retinal cells at later stages. c-ret labelling was found over ganglion cells, amacrine and horizontal cells; the preferred GDNF receptor (GFR alpha 1) over amacrine and horizontal cells; and the less preferred GDNF receptor (GFR alpha 2) over ganglion cells, amacrine cells and photoreceptors.

Animals↗

Effects of betaxolol on light responses and membrane conductance in retinal ganglion cells.

PURPOSE: To examine the physiological effects of betaxolol, a beta1-adrenergic receptor blocker commonly used in the treatment of glaucoma, on retinal ganglion cells and to evaluate its potential to elicit responses consistent with a neuroprotective agent against ganglion cell degeneration. METHODS: Single-unit extracellular recording, electroretinogram (ERG), intracellular and whole-cell patch-clamp recording techniques were made from flatmounted, isolated retina, superfused eyecup, and living retinal slice preparations of the larval tiger salamander. RESULTS: Bath application of 20 microM betaxolol reduced the glutamate-induced increase of spontaneous spike rate in retinal ganglion cell by approximately 30%. The glutamate-induced postsynaptic current recorded under voltage-clamp conditions was reduced by 50 microM betaxolol, and the difference current-voltage (I-V) relation (I(Control)-I(betaxolol)) was N-shaped and AP5-sensitive, characteristic of N-methyl-D-aspartate receptor-mediated current. Application of 50 microM betaxolol reversibly reduced the voltage-gated sodium and calcium currents by approximately one third of their peak amplitudes. The times-to-action of betaxolol on ganglion cells are long (15-35 minutes for 20-50 microM betaxolol), indicative of modulation through slow biochemical cascades. Betaxolol, up to 100 microM, exerted no effects on horizontal cells or the ERG, suggesting that the primary actions of this beta1 blocker are restricted to retinal ganglion cells. CONCLUSIONS: These physiological experiments provide supporting evidence that betaxolol acts in a manner consistent with preventing retinal ganglion cell death induced by elevated extracellular glutamate or by increased spontaneous spike rates under pathologic conditions. The physiological actions of betaxolol lead to reducing neurotoxic effects in ganglion cells, which are the most susceptible retinal neurons to glutamate-induced damages under ischemic and glaucomatous conditions. Therefore, betaxolol has the potential to be a neuroprotective agent against retinal degeneration in patients with disorders mediated by such mechanisms.

Adrenergic beta-Antagonists↗

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↗