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PubMed2005

A comparison of release kinetics and glutamate receptor properties in shaping rod-cone differences in EPSC kinetics in the salamander retina.

Synaptic transmission from cones is faster than transmission from rods. Using paired simultaneous recordings from photoreceptors and second-order neurones in the salamander retina, we studied the contributions of rod-cone differences in glutamate receptor properties and synaptic release rates to shaping postsynaptic responses. Depolarizing steps evoked sustained calcium currents in rods and cones that in turn produced transient excitatory postsynaptic currents (EPSCs) in horizontal and OFF bipolar cells. Cone-driven EPSCs rose and decayed faster than rod-driven EPSCs, even when comparing inputs from a rod and cone onto the same postsynaptic neurone. Thus, rod-cone differences in EPSCs reflect properties of individual rod and cone synapses. Experiments with selective AMPA and KA agonists and antagonists showed that rods and cones both contact pharmacologically similar AMPA receptors. Spontaneous miniature EPSCs (mEPSCs) exhibited unimodal distributions of amplitude and half-amplitude time width and there were no rod-cone differences in mEPSC properties. To examine how release kinetics shape the EPSC, we convolved mEPSC waveforms with empirically determined release rate functions for rods and cones. The predicted EPSC waveform closely matched the actual EPSC evoked by cones, supporting a quantal release model at the photoreceptor synapse. Convolution with the rod release function also produced a good match in rod-driven cells, although the actual EPSC was often somewhat slower than the predicted EPSC, a discrepancy partly explained by rod-rod coupling. Rod-cone differences in the rates of exocytosis are thus a major factor in producing faster cone-driven responses in second-order retinal neurones.

Animals↗
PubMed2005

Targeted effects of retinoic acid signaling upon photoreceptor development in zebrafish.

Retinoic acid (RA) is a signaling molecule important for photoreceptor development in vertebrates. The purpose of this study was to examine the mechanisms of the effects of RA upon developing rod and cone photoreceptors in the embryonic zebrafish. Exposure to exogenous RA increased the number of photoreceptors expressing rod opsin and red cone opsin, and decreased the number of photoreceptors expressing the blue and UV cone opsins, suggesting targeted effects of RA on photoreceptor development. RA exposure also increased opsin expression in individual rods and red cones, but decreased opsin expression in individual blue and UV cones, as indicated by differences in the strength of opsin hybridization in identified photoreceptors. RA exposure did not, however, significantly alter quantitative measures of photoreceptor pattern in a manner expected for changes in photoreceptor fate. These observations collectively indicate that RA treatment does not affect photoreceptor fate, but rather differentially influences opsin transcription in determined photoreceptors. An enzyme involved in RA synthesis, RALDH2, was immunocytochemically localized to retinal progenitor cells and the retinal pigmented epithelium (RPE), suggesting the presence of RA in the vicinity of developing photoreceptors. However, expression of an RA response element-driven transgene was restricted to the RPE, retinal progenitors, and a small population of neurons in ventral retina, suggesting that the endogenous RA signaling system is spatially limited within the eye.

Aldehyde Oxidoreductases↗
PubMed1993

Presence and foveal enrichment of rod opsin in the "all cone" retina of the American chameleon.

The retinal photoreceptors of the eye of the American chameleon, Anolis carolinensis, have been considered to be exclusively cones. Its retina is unusual for possessing two foveas (areas associated with heightened visual acuity), with the major, central fovea deeply incised and very densely packed with photoreceptors. Immunoblotting and light- and electron microscopic-immunocytochemistry, using several opsin monoclonal antibodies previously found specific for rods, demonstrated the presence and localization of this protein in the Anolis retina. This visual pigment appears sparsely in a subpopulation of photoreceptors in the periphery but overwhelmingly in the central fovea. Complementary results with cone-specific antibody and lectin binding corroborated this spatial organization. These results, as well as those with geckos, suggest that photoreceptor morphology is not an accurate guide among the lacertilians to visual pigment content, and that this phylogenetic grouping may constitute a crossroads in vertebrate photoreceptor evolution.

Animals↗
PubMed2000

Acquired color vision loss and a possible mechanism of ganglion cell death in glaucoma.

PURPOSE: First, to study the cellular mechanisms of acquired color vision loss in retinal detachment and diabetic retinopathy. Second, to learn why, in glaucoma, the type of color vision deficit that is observed is more characteristic of a retinal injury than it is of an optic neuropathy. Third, to test a hypothesis of photoreceptor-induced, ganglion cell death in glaucoma. METHODS: Various histologic techniques were employed to distinguish the L/M-cones (long/medium wavelength-sensitive cones, or red/green sensitive cones) from the S-cones (short wavelength-sensitive cones, or blue sensitive cones) in humans and monkeys with retinal detachment, humans with diabetic retinopathy, and both humans and monkeys with glaucoma. To test if the photoreceptors were contributing to ganglion cell death, laser photocoagulation was used in a experimental model of glaucoma to focally eliminate the photoreceptors. As a control, optic nerve transection was done following retinal laser photocoagulation in one animal. RESULTS: Selective and widespread loss of the S-cones was found in retinal detachment as well as diabetic retinopathy. By contrast, in human as well as experimental glaucoma, marked swelling of the L/M-cones was the predominant histopathologic feature. Retinal laser photocoagulation followed by experimental glaucoma resulted in selective protection of ganglion cells overlying the laser spots. This was not seen with retinal laser photocoagulation by optic nerve transection. CONCLUSIONS: In retinal detachment and diabetic retinopathy, acquired tritan-like color vision loss could be caused, or contributed to, by selective loss of the S-cones. Both L- and M-cones are affected in glaucoma, which is also consistent with a tritan-like deficit. Although not a therapeutic option, protection of ganglion cells by retinal laser in experimental glaucoma is consistent with an hypothesis of anterograde, photoreceptor-induced, ganglion cell death.

Adult↗
PubMed1997

Localization of guanylate cyclase-activating protein 2 in mammalian retinas.

Guanylate cyclase-activating proteins (GCAP1 and GCAP2) are thought to mediate the intracellular stimulation of guanylate cyclase (GC) by Ca2+, a key event in recovery of the dark state of rod photoreceptors after exposure to light. GCAP1 has been localized to rod and cone outer segments, the sites of phototransduction, and to photoreceptor synaptic terminals and some cone somata. We used in situ hybridization and immunocytochemistry to localize GCAP2 in human, monkey, and bovine retinas. In human and monkey retinas, the most intense immunolabeling with anti-GCAP2 antibodies was in the cone inner segments, somata, and synaptic terminals and, to a lesser degree, in rod inner segments and inner retinal neurons. In bovine retina, the most intense immunolabeling was in the rod inner segments, with weaker labeling of cone myoids, somata, and synapses. By using a GCAP2-specific antibody in enzymatic assays, we confirmed that GCAP1 but not GCAP2 is the major component that stimulates GC in bovine rod outer segment homogenates. These results suggest that although GCAP1 is involved in the Ca2+-sensitive regulation of GC in rod and cone outer segments, GCAP2 may have non-phototransduction functions in photoreceptors and inner retinal neurons.

Animals↗
PubMed1961

The spectral sensitivity of crayfish and lobster vision.

(1) The spectral sensitivity function for the compound eye of the crayfish has been determined by recording the retinal action potentials elicited by monochromatic stimuli. Its peak lies at approximately 570 mmicro. (2) Similar measurements made on lobster eyes yield functions with maxima in the region of 520 to 525 mmicro, which agree well with the absorption spectrum of lobster rhodopsin if minor allowances are made for distortion by known screening pigments. (3) The crayfish sensitivity function, since it is unaffected by selective monochromatic light adaptation, must be determined by a single photosensitive pigment. The absorption maximum of this pigment may be inferred with reasonable accuracy from the sensitivity data. (4) The visual pigment of the crayfish thus has its maximum absorption displaced by 50 to 60 mmicro towards the red end of the spectrum from that of the lobster and other marine crustacea. This shift parallels that found in both rod and cone pigments between fresh water and marine vertebrates. In the crayfish, however, an altered protein is responsible for the shift and not a new carotenoid chromophore as in the vertebrates. (5) The existence of this situation in a new group of animals (with photoreceptors which have been evolved independently from those of vertebrates) strengthens the view that there may be strong selection for long wavelength visual sensitivity in fresh water.

Adaptation, Ocular↗
PubMedDate not supplied

Visual pigments, cone oil droplets and ocular media in four species of estrildid finch.

A microspectrophotometric study was conducted on the retinal photoreceptors of four species of bird: cut-throat finches (Amadina fasciata), gouldian finches (Erythrura gouldiae), white-headed munias (Lonchura maja) and plum-headed finches (Neochmia modesta). Spectral characteristics of the photoreceptors in all four species were very similar. Rods contained a medium-wavelength-sensitive visual pigment with a wavelength of maximum absorbance at 502-504 nm. Four spectrally distinct types of single cone contained a visual pigment with wavelength of maximum absorbance at either 370-373 nm (ultraviolet-sensitive), 440-447 nm (short-wavelength-sensitive); 500 nm (medium-wavelength-sensitive) or 562-565 nm (long-wavelength-sensitive). Oil droplets in the ultraviolet-sensitive single cones showed no detectable absorption between 330 nm and 800 nm. Oil droplets in the short-, medium-, and long-wavelength-sensitive single cones had cut-off wavelengths at 415-423 nm, 510-520 nm and 567-575 nm, respectively. Double cones contained the visual pigment with wavelength of maximum absorbance at 562-565 nm observed in long-wavelength-sensitive single cones. Only the principal member of the double cone pair contained an oil droplet (P-type, cut-off wavelength at 414-489 nm depending on species and retinal location). Spectral transmittance of the intact ocular media of each species was measured along the optic axis. Wavelengths of 0.5 transmittance for all species were very similar (316-318 nm).

Animals↗
PubMed1998

Regeneration of the newt retina: order of appearance of photoreceptors and ganglion cells.

The adult newt regenerates a functional retina following removal or destruction of the original retina. We studied the order of appearance of cell types in the regenerating retina by using immunohistochemical techniques. An antibody that recognizes the alpha subunit (260 kDa) of voltage-dependent Na+ channels was found to label a 255-kDa band in Western blots of crude membrane fractions from the normal retina. Cryosections of normal retina revealed intense Na+ channel immunoreactivity in somata and axons of ganglion cells, weaker immunoreactivity in somata of amacrine cells, and no immunoreactivity in the inner plexiform layer. In the same sections, immunoreactivity to a monoclonal antibody (RB-1) specific to newt cones was intense in the photoreceptor layer. In regenerating retinas, double staining with the Na+ channel antibody as a possible marker of ganglion cells and RB-1 antibody first revealed immunoreactive cells at the intermediate stage (three to five cells thick), which does not exhibit segregated synaptic layers. Na+ channel-immunoreactive ganglion cells appeared before the RB-1-immunoreactive photoreceptors. Because ganglion cells also appear before photoreceptor cells in normal development, common mechanisms may control both the generation and the regeneration of the newt retina.

Animals↗
PubMed2002

Calcium regulation in photoreceptors.

In this review we describe some of the remarkable and intricate mechanisms through which the calcium ion (Ca2+) contributes to detection, transduction and synaptic transfer of light stimuli in rod and cone photoreceptors. The function of Ca2+ is highly compartmentalized. In the outer segment, Ca2+ controls photoreceptor light adaptation by independently adjusting the gain of phototransduction at several stages in the transduction chain. In the inner segment and synaptic terminal, Ca2+ regulates cells' metabolism, glutamate release, cytoskeletal dynamics, gene expression and cell death. We discuss the mechanisms of Ca2+ entry, buffering, sequestration, release from internal stores and Ca2+ extrusion from both outer and inner segments, showing that these two compartments have little in common with respect to Ca2+ homeostasis. We also investigate the various roles played by Ca2+ as an integrator of intracellular signaling pathways, and emphasize the central role played by Ca2+ as a second messenger in neuromodulation of photoreceptor signaling by extracellular ligands such as dopamine, adenosine and somatostatin. Finally, we review the intimate link between dysfunction in photoreceptor Ca2+ homeostasis and pathologies leading to retinal dysfunction and blindness.

Adaptation, Physiological↗
PubMed2001

Activation, deactivation, and adaptation in vertebrate photoreceptor cells.

Visual transduction captures widespread interest because its G-protein signaling motif recurs throughout nature yet is uniquely accessible for study in the photoreceptor cells. The light-activated currents generated at the photoreceptor outer segment provide an easily observed real-time measure of the output of the signaling cascade, and the ease of obtaining pure samples of outer segments in reasonable quantity facilitates biochemical experiments. A quiet revolution in the study of the mechanism has occurred during the past decade with the advent of gene-targeting techniques. These have made it possible to observe how transduction is perturbed by the deletion, overexpression, or mutation of specific components of the transduction apparatus.

Animals↗
PubMed2005

Different functional types of bipolar cells use different gap-junctional proteins.

Rod signals are transmitted to ON retinal ganglion cells by means of gap junctions between AII amacrine cells and ON bipolars. The AII amacrine cells are known to express connexin36 (Cx36), but previous studies of Cx36 in ON cone bipolars have been ambiguous. Here, we studied bipolar cells in a transgenic mouse line that expresses high levels of green fluorescent protein (GFP) in one type of ON cone bipolar cell. We found strong Cx36 immunostaining in the axon terminals of the GFP-labeled type 357 bipolar cells in both vertical sections and whole mounts of the retina. This finding was confirmed by single-cell immunostaining and single-cell reverse transcription-PCR (RT-PCR). As reported previously (Maxeiner et al., 2005), Cx45 was found in some ON bipolar cells, but RT-PCR showed Cx36 and not Cx45 to be expressed by the type 357 bipolar cells. Some of the remaining GFP-negative bipolar cells expressed Cx45 but not Cx36. It appears that different types of ON cone bipolar cells express different connexins at their gap junctions with AII amacrine cells.

Amacrine Cells↗
PubMed1998

Comparative morphology of distal neurons in larval and adult zebrafish retinas.

Distal retinal cells from larval (7-10 days postfertilization) and adult zebrafish retinas were cultured in 70% L-15 medium for 4-5 d and comparable cell types identified. Four photoreceptor types were observed in adult retinal cultures, whereas only single cones were isolated from larval retinas. Horizontal cells in both larval and adult cultures were distinguished by their large size and stellate morphology and two subtypes, A and B, were recognized. Bipolar cells were readily identified in adult cultures, but rare in larval cultures. Two bipolar cell types, large and small, were distinguished. Measurements of the various cell types are provided.

Animals↗
PubMed2005

Characterization of a transgenic mouse line lacking photoreceptor development within the ventral retina.

A unique transgenic mouse line was generated by incorporating a minigene that contained a cone-specific human cone transducin alpha-subunit (GNAT2) promoter, an attenuated diphtheria toxin A (DTA) gene, and an enhancer element from human interphotoreceptor retinoid-binding protein (IRBP) gene. This transgenic mouse line is designated h-GNAT2pro-DTA. During postnatal retinal development, both transgenic and non-transgenic retinas showed similar morphology and thickness at P1. Between ages P8 and P30, all retinal layers became recognizable in non-transgenic and also in transgenic dorsal retinas. However, in the ventral retina of the transgenic mice the photoreceptor layers did not develop. This aberration occurred as a result of abnormal cellular development, rather than as a consequence of retinal degeneration. In adult transgenic animals, approximately 44% of the retina located dorsally appeared morphologically normal, whereas 32% of the retina located ventrally was completely lacking photoreceptor development. The 24% mid-retinal region exhibited transitional morphology containing malformed photoreceptors. At P360 or older, the thickness of retina layers was reduced in both dorsal and ventral regions. The abnormality observed in transgenic retinas involved mainly the photoreceptors; the other retinal cell types were all present in both dorsal and ventral retinas. Since the DTA gene was only expressed in cone cells, the absence of cone photoreceptors in the transgenic retina was to be expected. However, what was unexpected was the concomitant absence of rod photoreceptors in the ventral retina, suggesting that the presence of cones may be important for the development of rods. This new transgenic line can lead to better understanding of photoreceptor development, and may serve as a new animal model for studying photoreceptor-related retinal diseases.

Animals↗
PubMedDate not supplied

Patch-clamp recording of human retinal photoreceptors and bipolar cells.

Photoreceptors and retinal bipolar cells are considered as nonspiking neurons; however, we recently showed that human rod photoreceptors can generate sodium action potentials in response to membrane depolarization from membrane potentials of -60 or -70 mV (Kawai et al., Neuron 30 [2001] 451). We performed patch-clamp recording of human cone photoreceptors and retinal bipolar cells to examine whether functional voltage-gated sodium channels are expressed in these cells as well as rod photoreceptors. Under current-clamp conditions, the injection of depolarizing current steps into a cone photoreceptor-induced marked action potentials. These action potentials were blocked by 1 microM tetrodotoxin, a voltage-gated sodium channel blocker. Under voltage-clamp conditions, depolarizing voltage steps-induced a fast transient inward current in several bipolar cells (n = 4/78). This current was activated from -70 to + 20 mV (maximal at -10 mV) and inactivated within 5 ms. The 10-90% rise time of this current was shorter than another inward current (less than one-hundredth). These results indicate that human cones and bipolar cells express voltage-gated sodium channels as rod photoreceptors. Sodium channels may serve to amplify the release of a neurotransmitter and to accelerate the light-dark change in photosignals.

Adult↗
PubMed2006

Visual cycle: Dependence of retinol production and removal on photoproduct decay and cell morphology.

The visual cycle is a chain of biochemical reactions that regenerate visual pigment following exposure to light. Initial steps, the liberation of all-trans retinal and its reduction to all-trans retinol by retinol dehydrogenase (RDH), take place in photoreceptors. We performed comparative microspectrophotometric and microfluorometric measurements on a variety of rod and cone photoreceptors isolated from salamander retinae to correlate the rates of photoproduct decay and retinol production. Metapigment decay rate was spatially uniform within outer segments and 50-70 times faster in the cells that contained cone-type pigment (SWS2 and M/LWS) compared to cells with rod-type pigment (RH1). Retinol production rate was strongly position dependent, fastest at the base of outer segments. Retinol production rate was 10-40 times faster in cones with cone pigments (SWS2 and M/LWS) than in the basal OS of rods containing rod pigment (RH1). Production rate was approximately five times faster in rods containing cone pigment (SWS2) than the rate in basal OS of rods containing the rod pigment (RH1). We show that retinol production is defined either by metapigment decay rate or RDH reaction rate, depending on cell type or outer segment region, whereas retinol removal is defined by the surface-to-volume ratio of the outer segment and the availability of retinoid binding protein (IRBP). The more rapid rates of retinol production in cones compared to rods are consistent with the more rapid operation of the visual cycle in these cells.

Ambystoma↗