Teleost Retina Research Articles

Dopamine and nitric oxide control both flickering and steady-light-induced cone contraction and horizontal cell spinule formation in the teleost (carp) retina: serial interaction of dopamine and nitric oxide.

Adaptation to ambient light, which is an important characteristic of the vertebrate visual system, involves cellular and subcellular (synaptic) plasticity of the retina. The present study investigated dopamine (DA) and nitric oxide (NO) as possible neurochemical modulators controlling cone photomechanical movements (PMMs) and horizontal cell (HC) spinules in relation to steady and flickering light adaptation in the carp retina. Haloperidol (HAL; a nonspecific DA receptor blocker) or cPTIO (a NO scavenger) largely inhibited the cone PMMs and HC spinule formation induced by either steady or flickering light. These results suggested that both DA and NO could be involved in the light-adaptation changes induced by either pattern of input and that DA and NO effects may not be completely independent. The possibility that NO and DA interact serially was evaluated pharmacologically by cross-antagonist application (i.e., DA + cPTIO or NO + HAL). When a NO donor was coapplied with HAL to dark-adapted eyecups, normal light-adaptive cone PMMs and HC spinules occurred. In contrast, when DA was applied in the presence of cPTIO, the dark-adapted state persisted. It was concluded 1) that DA and NO are both light-adaptive neurochemicals, released in the retina during either steady or flickering light; 2) that the effects of DA and NO on light-adaptive cone PMMs and HC spinules do not occur in parallel; and 3) that NO and DA act mainly in series, specifically as follows: Light --> DA --> NO --> Cone PMMs + HC spinules.

Interocular effect of intravitreal injection of 6-hydroxydopamine and dopamine on spinule formation in teleost retina.

Terminal dendrites of cone horizontal cells (HCs) in teleost retinas show numerous spine-like protrusions named spinules, which are invaginated into the cone pedicles during light-adaptation, but retracted during dark-adaptation. Somata of HC show nematosomes whose size decreases as the number of spinules increases. Mechanisms regulating these changes in nematosomes and spinules are only partially understood, being an area of controversy in retinal cell biology. It has been suggested that efferent fibres from the brain to the retina might be involved in the control of spinule formation. Moreover, we have reported that actin depolymerization has an interocular effect on spinule formation, which could be mediated by these fibres. In the present report, we show an interocular effect on spinule dynamics: the monocular intravitreal injection of dopamine (DA) and 6-hydroxydopamine (6-OHDA), two drugs that affect the spinule formation, produces the same effects in the contralateral, untreated eye as in the injected eye. Our results reinforce the idea of an interocular central control of this phenomenon of synaptic plasticity. Dopamine-dependent events in the retina appear to be necessary to forge the afferent signals eliciting this interocular effect.

Characterization of retinal guanylate cyclase-activating protein 3 (GCAP3) from zebrafish to man.

Calmodulin-like neuronal Ca2+-binding proteins (NCBPs) are expressed primarily in neurons and contain a combination of four functional and nonfunctional EF-hand Ca2+-binding motifs. The guanylate cyclase-activating proteins 1-3 (GCAP1-3), the best characterized subgroup of NCBPs, function in the regulation of transmembrane guanylate cyclases 1-2 (GC1-2). The pairing of GCAPs and GCs in vivo depends on cell expression. Therefore, we investigated the expression of these genes in retina using in situ hybridization and immunocytochemistry. Our results demonstrate that GCAP1, GCAP2, GC1 and GC2 are expressed in human rod and cone photoreceptors, while GCAP3 is expressed exclusively in cones. As a consequence of extensive modification, the GCAP3 gene is not expressed in mouse retina. However, this lack of evolutionary conservation appears to be restricted to only some species as we cloned all three GCAPs from teleost (zebrafish) retina and localized them to rod cells, short single cones (GCAP1-2), and all subtypes of cones (GCAP3). Furthermore, sequence comparisons and evolutionary trace analysis coupled with functional testing of the different GCAPs allowed us to identify the key conserved residues that are critical for GCAP structure and function, and to define class-specific residues for the NCBP subfamilies.

Nitric oxide donor stimulated increase of cyclic GMP in the goldfish retina

Nitric oxide (NO) activates soluble guanylyl cyclase (sGC) and the resulting increase in cyclic guanosine monophosphate (cGMP) is an important intracellular signalling pathway in the vertebrate retina. Immunocytochemical detection of cGMP following exposure to NO donors has proven an effective method of identifying cells that express sGC. While such an approach has proven useful for the study of several vertebrate retinas, it has not been applied to the well-characterized teleost retina. Therefore, in the present study, we have applied this approach to the retina of the goldfish (Carassius auratus). In the presence of the phosphodiesterase (PDE) inhibitor 3-isobutyl-1-methylxanthine (IBMX), incubation of goldfish eyecups in Ringer's solution containing (+/-)-S-nitroso-N-acetylpenicillamine (SNAP) increased cGMP-like immunoreactivity (cG-ir) in bipolar, horizontal, amacrine, and ganglion cells and in ganglion cell axons and optic nerve. Weak labeling was observed in horizontal cells but no change in cG-ir was noted within photoreceptors. The NO donor-stimulated increases of cG-ir in horizontal, bipolar, amacrine, and ganglion cells are consistent with known physiological effects of NO on these neurons. The physiological significance of NO action at the level of optic nerve is not known. The lack of an effect of SNAP on cG-ir in photoreceptors was unexpected, as there are known physiological actions of NO, mediated by cGMP, on these neurons. Although this may be due to insufficient sensitivity of immunolabeling, this result may indicate a difference between isoforms of sGC or cGMP PDE in these neurons, compared to neurons where exogenous NO increased cG-ir.

Identification of rod- and cone-specific phosducins in teleost retinas

Phosducin (PD) is a regulatory protein of vertebrate phototransduction cascades. In mammalian retina, it has been thought that only one kind of PD commonly exists in both rods and cones. However, we have found two kinds of PD (OlPD-R and OlPD-C) in the retina of a teleost, medaka ( Oryzias latipes). In situ hybridization and immunohistochemical analysis demonstrated that OlPD-R and -C are selectively expressed in rods and cones, respectively. The antiserum against medaka PDs recognized two kinds of proteins in bluegill ( Lepomis macrochirus) retina. These results suggest that rod- and cone-specific PDs exist in teleost retinas, probably creating differences in light adaptation between rods and cones.

Expression of pineal ultraviolet- and green-like opsins in the pineal organ and retina of teleosts.

In teleostean bony fishes, studies on the adults of various species have shown that pineal photoreceptors are maximally sensitive to short- and middle-wavelength light, possibly utilising both rod-like and pineal-specific opsins. Until recently, however, very little was known about the pineal opsins present in embryonic and larval teleosts and their relationships to opsins expressed by retinal photoreceptors. Our immunocytochemical studies have revealed that, in Atlantic halibut, herring and cod, pineal photoreceptors express principal phototransduction molecules during embryonic life before they appear in retinal photoreceptors. In cDNA from embryonic and adult halibut, we identified two partial opsin gene sequences, HPO1 and HPO4, with highest homology to teleost green and ultraviolet cone opsins (72-83% and 71-83% amino acid identity, respectively). In halibut, these opsins are expressed in the pineal organ of embryos and appear in the retina of larvae. Our recent in situ hybridisation studies with RNA probes for HPO1 and HPO4 demonstrate the presence of green-like opsin mRNAs in the pineal organ and the retina of herring, cod, turbot, haddock, Atlantic salmon, zebrafish and three species of cichlid, and of ultraviolet opsins in the retinas of zebrafish, Atlantic salmon, turbot and the three cichlid species. We conclude that the halibut pineal organ appears to have the potential for both ultraviolet and green photosensitivity from the embryonic stage and that the retina may acquire the same potential during the larval stages. In the other teleosts studied, although both pineal and retinal photoreceptors seem to utilise a green-like opsin from the larval stage, ultraviolet photoreception appears to be restricted to the retina.

Mouse horizontal cells do not express connexin26 or connexin36.

Gap junctions between neurons function as electrical synapses, and are present in all layers of mammalian and teleost retina. These synapses are largest and most prominent between horizontal cells where they function to increase the receptive field of a single neuron beyond the width of its dendrites. Receptive field size and the extent of gap junctional coupling between horizontal cells is regulated by ambient light levels and may mediate light/dark adaptation. Furthermore, teleost horizontal cell gap junction hemichannels may facilitate a mechanism of feedback inhibition between horizontal cells and cone photoreceptors. As a prelude to using mouse genetic models to study horizontal cell gap junctions and hemichannels, we sought to determine the connexin complement of mouse horizontal cells. Cx36, Cx37, Cx43, Cx45 and Cx57 mRNA could be detected in mouse retina by RT-PCR. Microscopy was used to further examine the distribution of Cx26 and Cx36. Cx26 immunofluorescence and a beta-gal reporter under regulatory control of the Cx36 promoter did not colocalize with a horizontal cell marker, indicating that these genes are not expressed by horizontal cells. The identity of the connexin(s) forming electrical synapses between mouse horizontal cells and the connexin that may form hemichannels in the horizontal cell telodendria remains unknown.

A molecular phenotype atlas of the zebrafish retina.

The rasborine cyprinid Danio rerio (the zebrafish) has become a popular model of retinal function and development. Its value depends, in part, on validation of homologies with retinal cell populations of cyprinine cyprinids. This atlas provides raw and interpreted molecular phenotype data derived from computationally classified sets of small molecule signals from different cell types in the zebrafish retina: L-alanine, L-aspartate, L-glutamine, L-glutamate, glutathione, glycine, taurine and gamma-aminobutyrate. This basis set yields an 8-dimensional signature for every retinal cell and formally establishes molecular signature homologies with retinal neurons, glia, epithelia and endothelia of other cyprinids. Zebrafish photoreceptor classes have been characterized previously: we now show their metabolic profiles to be identical to those of the corresponding photoreceptors in goldfish. The inner nuclear layer is partitioned into precise horizontal, bipolar and amacrine cell layers. The horizontal cell layer contains at least three and perhaps all four known classes of cyprinine horizontal cells. Homologues of cyprinid glutamatergic ON-center and OFF-center mixed rod-cone bipolar cells are present and it appears likely that all five classes are present in zebrafish. The cone bipolar cells defy simple analysis but comprise the largest fraction of bipolar cells, as in all cyprinids. Signature analysis reveals six molecular phenotypes in the bipolar cell cohort: most are superclasses. The amacrine cell layer is composed of approximately equal 64% GABA+ and 35% glycine+ amacrine cells, with the remainder being sparse dopaminergic interplexiform cells and other rare unidentified neurons. These different amacrine cell types are completely distinct in the dark adapted retina, but light adapted retinas display weak leakage of GABA signals into many glycinergic amacrine cells, suggesting widespread heterocellular coupling. The composition of the zebrafish ganglion cell layer is metabolically indistinguishable from that in other cyprinids, and the signatures of glial and non-neuronal cells display strong homologies with those in mammals. As in most vertebrates, zebrafish Müller cells possess a high glutamine, low glutamate signature and contain the dominant pool of glutathione in the neural retina. The retinal pigmented epithelium shows a general mammalian signature but also has exceptional glutathione content (5-10 mM), perhaps required by the unusually high oxygen tensions of teleost retinas. The optic nerve and the marginal zone of the retina reveal characteristic metabolic specializations. The marginal zone is strongly laminated and its nascent neurons display their characteristic signatures before taking their place in the retina proper.

Cellular proliferation and neurogenesis in the injured retina of adult zebrafish.

The retinas of adult teleost fish can regenerate neurons following a chemical or mechanical injury. Previous studies have demonstrated that mechanical excision of fish retina induces a hyperplasia within the retinal sheet, including the formation of a proliferative blastema from whence new retinal cells are produced to fill the excision site. The current study was designed to address two issues regarding injury-induced retinal hyperplasia: (1) Retinas of adult zebrafish can regenerate following a surgical excision, but compared to other fish they contain very few proliferative cells: Might retinal injury in adult zebrafish therefore induce minimal, or perhaps no, hyperplasia? (2) The fate of injury-induced, proliferative retinal cells outside surgical excision sites has yet to be determined. Do such cells produce retinal neurons? Evidence is presented that mechanical injury to the adult zebrafish retina induces a dramatic increase in the number of proliferative cells both within and external to the lesion site, and some of these cells apparently migrate within the radial dimension of the retina. Evidence is also presented that injury-induced proliferative cells outside a lesion site can produce retinal neurons--including cone photoreceptors, interplexiform cells, and amacrine cells--that are incorporated into the extant retina. The results suggest that the adult zebrafish retina contains a latent population of cells that is induced to proliferate following retinal injury, and that these cells might represent a novel avenue for pluripotent neurogenesis within the intact adult teleost retina.

Taurine blocks spontaneous cone contraction but not horizontal cell dark suppression in isolated goldfish retina.

The objective of this study was to investigate the effects of taurine on cone retinomotor movements and the responses of cone-driven horizontal cells in dark-adapted teleost retina. In isolated goldfish retina preparations maintained in the dark, cones spontaneously contracted, and the responses of horizontal cells were suppressed. Addition of 5 mM taurine to the physiological solution blocked the spontaneous contraction of cones in the dark but did not block the dark-suppression of horizontal cell responses. These results indicate that the mechanism that leads to horizontal cell dark suppression is not sensitive to taurine. Although both cone retinomotor position and horizontal cell responsiveness are known to be modulated by dopamine, the present results do not support the hypothesis that taurine inhibits dopamine release in the dark because only spontaneous cone contraction was affected by taurine. These results also indicate that spontaneous cone contraction in the dark is not the cause of horizontal cell dark suppression because, in the presence of taurine, cones were elongated yet horizontal cell responses were still suppressed. Consequently, these results make it clear that horizontal cell dark suppression is not an artifact produced by incubating isolated teleost retina preparations in taurine-free physiological solution.

Inherited retinal degeneration and apoptosis in mutant zebrafish

The mechanism of retinal cell death was studied in mutant zebrafish (Danio rerio) which undergo inherited degeneration of the retina and the brain. The shrunken head (shrm33) mutation was isolated as part of a large scale mutagenesis screen. The yellow head (yhd) mutation arose spontaneously among inbred wild type zebrafish. Although the mutants share many morphological features, including small eyes, a small brain and an enlarged pericardial sac, crossing shrm33 and yhd heterozygotes results in phenotypically normal fish. The retinae of both mutant lines of fish begin to develop normally and then undergo massive degenerative changes. Pyknotic cells first appear in the retina of the shrm33 fish by 3 days post-fertilization and in the yhd fish by 1.5 days post-fertilization. By 5 days post-fertilization the outer nuclear layer containing the photoreceptor cells has largely disappeared in both mutants. The inner nuclear layer and ganglion cell layer are also severely affected. By 6–7 days post-fertilization, the retina has been largely cleared of pyknotic cells by retinal pigment epithelial cells and by rare macrophage-like cells. Both mutations are lethal by 7–8 days post-fertilization. Two independent measures, TdT-mediated dUTP-biotin nick end label (TUNEL) and transmission electron microscopy, indicate that the pyknotic cells in the mutant retinae are apoptotic. Apoptosis is very rarely observed during normal development of the teleost retina and was not observed in age-matched wild type zebrafish retinae examined for comparison. Our results indicate that a genetic defect can induce massive apoptosis in cell populations that do not normally undergo apoptosis during development. Anat Rec 258:145–155, 2000. © 2000 Wiley-Liss, Inc.

Inherited retinal degeneration and apoptosis in mutant zebrafish.

The mechanism of retinal cell death was studied in mutant zebrafish (Danio rerio) which undergo inherited degeneration of the retina and the brain. The shrunken head (shr(m33)) mutation was isolated as part of a large scale mutagenesis screen. The yellow head (yhd) mutation arose spontaneously among inbred wild type zebrafish. Although the mutants share many morphological features, including small eyes, a small brain and an enlarged pericardial sac, crossing shr(m33) and yhd heterozygotes results in phenotypically normal fish. The retinae of both mutant lines of fish begin to develop normally and then undergo massive degenerative changes. Pyknotic cells first appear in the retina of the shr(m33) fish by 3 days post-fertilization and in the yhd fish by 1.5 days post-fertilization. By 5 days post-fertilization the outer nuclear layer containing the photoreceptor cells has largely disappeared in both mutants. The inner nuclear layer and ganglion cell layer are also severely affected. By 6-7 days post-fertilization, the retina has been largely cleared of pyknotic cells by retinal pigment epithelial cells and by rare macrophage-like cells. Both mutations are lethal by 7-8 days post-fertilization. Two independent measures, TdT-mediated dUTP-biotin nick end label (TUNEL) and transmission electron microscopy, indicate that the pyknotic cells in the mutant retinae are apoptotic. Apoptosis is very rarely observed during normal development of the teleost retina and was not observed in age-matched wild type zebrafish retinae examined for comparison. Our results indicate that a genetic defect can induce massive apoptosis in cell populations that do not normally undergo apoptosis during development.

Effect of melatonin agonists and antagonists on horizontal cell spinule formation and dopamine release in a fish retina.

The crucian carp retina was used to study the effects of the melatonin antagonist p697 (N-pentanoyl 2-benzyltryptamine) and the melatonin agonists [+]- and [-]-AMMTC (N-acetyl-4-aminomethyl-6-methoxy-9-methyl-1,2,3,4-tetrahydrocarbazol e) on horizontal cell spinule formation, an indicator of the state of retinal adaptation. DH97 was capable of both counteracting dark-adaptive spinule degradation and inducing light-adaptive spinule formation at the beginning of the dark phase. Addition of dopamine receptor blockers opposed the action of DH97 on spinules, with SCH 23930, a D1 dopamine receptor antagonist, being more effective than the D2 receptor antagonist sulpiride. DH97 induced a twofold increase in dopamine release. We conclude that melatonin acts as a dark signal within the teleost retina by inhibiting the dopaminergic system. In accordance with this, both enantiomers of AMMTC prevented light-induced spinule formation, and reduced dopamine release to below dark-adaptive baseline levels. We suggest that the suppression of spinule formation by AMMTC may be due to either a direct inhibitory interaction between the melatonin agonist and horizontal cell dopamine D1 receptors, or an inhibitory effect on the activity of the dopamine-releasing interplexiform cells.

Effects of genistein, a tyrosine kinase inhibitor, on light adaptive cellular and synaptic plasticity in the outer retina of a teleost fish (carp)

The effects of genistein, a protein tyrosine kinase inhibitor, on light adaptation-induced morphological (cellular and synaptic) plasticity were studied in the retina-eyecups of carp ( Cyprinus carpio ). In particular, the effects of genistein on the photomechanical movements of cones and the spinules of horizontal cells were quantitatively evaluated. The data suggested that genistein significantly blocked both light adaptive processes. It is concluded, therefore, that light adaptation of the teleost retina could involve activation of tyrosine kinases(s). This conclusion agrees with previous findings that multiple neuromodulators and protein kinases control retinal light adaptation. Keywords : Genistein, tyrosine kinase, light adaptation, plasticity, cone, photomechanical, movement, horizontal cell, spinules, retina, carp.

Antibody labeling of the blue-sensitive cones in the retinae of teleost fishes

Characterization of visual opsins can be achieved with the use of specific polyclonal or monoclonal antibodies. In this study, antiserum was raised against a conjugate protein including an N-terminal sequence of the short wavelength sensitive (SWS, commonly referred to as blue-sensitive) visual pigment opsin in goldfish (Carrasius auratus). The antiserum showed immunoreactivity to retinae of phylogenetically distant teleosts, including the goldfish, rainbow trout (Oncorhynchus mykiss), and bluehead wrasse (Thalassoma bisasciatum). Complementary DNA (cDNA) was synthesized from goldfish retinal messenger RNA. An N-terminal sequence of the SWS opsin was PCR-amplified from the cDNA and subsequently cloned into an expression vector. Expressed protein was purified by metal chelate affinity chromatography and used to immunize New Zealand white rabbits. Immune serum was collected and used in immunocytochemical assays, which revealed that the antiserum contains antibodies specific for an opsin that resides in single-cone photoreceptors of the goldfish, rainbow trout, and bluehead wrasse. The results confirm those from earlier studies of goldfish in which short-wavelength sensitivity was localized to single cones, and suggest that the association between short-wavelength sensitivity and single cones may be a feature common to many teleosts. The production of antiserum specific for SWS opsins provides a useful tool in the characterization of opsin expression within teleost retinae.

Cone specific arrestin and G protein-coupled receptor kinase isolated from teleost retina

Cone specific arrestin and G protein-coupled receptor kinase isolated from teleost retina

AMPA-preferring receptors with high Ca2+ permeability mediate dendritic plasticity of retinal horizontal cells.

The synaptic complex formed by the cone photoreceptor pedicles and the dendrites of horizontal cells in the teleost retina undergoes structural changes during light adaptation. Numerous spinules are formed by the terminal dendrites, and they are subsequently retracted during dark adaptation. In a retina kept under continuous illumination, the retraction process can be initiated by analogues of the neurotransmitter glutamate acting at AMPA/kainate receptors. On the other hand, the retraction process depends on calcium influx and the subsequent activation of CaMkII. We show here that the retraction of spinules induced by AMPA or kainate is not impaired in the presence of cobalt, making an involvement of voltage-gated calcium channels unlikely. Using calcium imaging techniques with isolated horizontal cells, we demonstrate that AMPA and kainate, but not NMDA, increase [Ca2+]i in the presence of nicardipine, caffeine and thapsigargin. The increase of [Ca2+]i under these conditions depends on [Ca2+]o and on the agonist in a dose-dependent manner, suggesting that the increase of [Ca2+]i is largely due to calcium influx through the agonist-gated channel. Pharmacological studies were performed to determine whether AMPA- and/or kainate-preferring receptors mediate the calcium influx. The AMPA-preferring receptor antagonist LY303070 blocked glutamate- and kainate-evoked increases of [Ca2+]i in a concentration-dependent manner, indicating that kainate-preferring receptors contributed little or nothing to the observed [Ca2+]i increase. This was supported by experiments where cyclothiazide (which blocks the desensitization of AMPA receptors) and concanavalin A (which potentiates responses mediated by kainate receptors) were applied. In all cases, LY303070 blocked the agonist-evoked increase of [Ca2+]i. The presence of AMPA-preferring receptors with high Ca2+ permeability on horizontal cells was also supported by measuring agonist-induced currents using whole-cell recording techniques. Furthermore, LY303070 was able to impair the retraction of spinules during dark adaption in the in vivo situation.

Activation of metabotropic glutamate receptors modulates the voltage-gated sustained calcium current in a teleost horizontal cell.

In the teleost retina, cone horizontal cells contain a voltage-activated sustained calcium current, which has been proposed to be involved in visual processing. Recently, several studies have demonstrated that modulation of voltage-gated channels can occur through activation of metabotropic glutamate receptors (mGluRs). Because glutamate is the excitatory neurotransmitter in the vertebrate retina, we have used whole cell electrophysiological techniques to examine the effect of mGluR activation on the sustained voltage-gated calcium current found in isolated cone horizontal cells in the catfish retina. In pharmacological conditions that blocked voltage-gated sodium and potassium channels, as well as N-methyl-D-aspartate (NMDA) and non-NMDA channels, application of L-glutamate or 1-aminocyclopentane-1,3-dicarboxylic acid (1S,3R-ACPD) to voltage-clamped cone horizontal cells acted to increase the amplitude of the calcium current, expand the activation range of the calcium current by 10 mV into the cell's physiological operating range, and shift the peak calcium current by -5 mV. To identify and characterize the mGluR subtypes found on catfish cone horizontal cells, agonists of group I, group II, or group III mGluRs were applied via perfusion. Group I and group III mGluR agonists mimicked the effect of L-glutamate or 1S,3R-ACPD, whereas group II mGluR agonists had no effect on L-type calcium current activity. Inhibition studies demonstrated that group I mGluR antagonists significantly blocked the modulatory effect of the group I mGluR agonist, (S)-3,5-dihydroxyphenylglycine. Similar results were obtained when the group III mGluR agonist, L-2-amino-4-phosphonobutyric acid, was applied in the presence of a group III mGluR antagonist. These results provide evidence for two groups of mGluR subtypes on catfish cone horizontal cells. Activation of these mGluRs is linked to modulation of the voltage-gated sustained calcium current.

Spatiotemporal development of the fish retina: Distribution of calbindin D-28K

Using antibodies to the calcium-binding protein calbindin (CB), we have studied the embryonic and postnatal development of retinal cells using the trout as an animal model. Our results reveal that the time course of expression of CB parallels the vitreal to scleral progression of differentiation of the teleost retina. The maturation of the retina also follows a spatial, centro-peripheral gradient of CB-expression with peripheral regions of the more mature retina undergoing maturation processes characteristic of the central retina of younger animals. Moreover, CB-expression within the optic tectum and the further lamination of this centre starts to take place only after the labeled ganglion cell axons arrived at the tectum.

Interocular effect of actin depolymerization on spinule formation in teleost retina

Teleost retinas adapted to light show numerous spinules invaginated in the cone pedicles whereas darkness induces a reduction in the number of spinules. Horizontal cells show nematosomes whose size decreases as the number of spinules increases. We have investigated the involvement of actin filaments in spinule formation, by using cytochalasin D through intraocular injection into an eye. The ultrastructural analysis reveals that cytochalasin D impairs spinule formation and nematosome-size reduction in both treated and contralateral untreated retinas.