Dopaminergic Interplexiform Cells Research Articles

Sensory Integration: Cross-Modal Communication Between the Olfactory and Visual Systems in Zebrafish.

Cross-modal sensory communication is an innate biological process that refers to the combination and/or interpretation of different types of sensory input in the brain. Often, this process conjugates with neural modulation, by which the neural signals that convey sensory information are adjusted, such as intensity, frequency, complexity, and/or novelty. Although the anatomic pathways involved in cross-modal sensory integration have been previously described, the course of development and the physiological roles of multisensory signaling integration in brain functions remain to be elucidated. In this article, I review some of the recent findings in sensory integration from research using the zebrafish models. In zebrafish, cross-modal sensory integration occurs between the olfactory and visual systems. It is mediated by the olfacto-retinal centrifugal (ORC) pathway, which originates from the terminalis nerve (TN) in the olfactory bulb and terminates in the neural retina. In the retina, the TNs synapse with the inner nuclear layer dopaminergic interplexiform cells (DA-IPCs). Through the ORC pathway, stimulation of the olfactory neurons alters the cellular activity of TNs and DA-IPCs, which in turn modulates retinal neural function and increases behavioral visual sensitivity.

Zebrafish Olfacto-Retinal Centrifugal Axon Projection and Distribution: Effects of Gonadotropin-Releasing Hormone and Dopaminergic Signaling

The terminalis neurons (TNs) have been described in teleost species. In zebrafish, the TNs are located in the olfactory bulb. The TNs synthesize and release gonadotropin-releasing hormone (GnRH) as one of the major neurotransmitters. The TNs project axons to many brain areas, which include the neural retina. In the retina, the TN axons synapse with dopaminergic interplexiform cells (DA-IPCs) and retinal ganglion cells (RGCs). In this research, we examine the role of GnRH and dopaminergic signaling in TN axon projection to the retina using the transgenic zebrafish Tg(GnRH-3::GFP). While the TNs developed at 34 h postfertilization (hpf), the first TN axons were not detected in the retina until 48-50 hpf, when the first DA-IPCs were differentiated. In developing embryos, inhibition of retinal GnRH signaling pathways severely interrupted the projection of TN axons to the retina. However, inhibition of retinal dopaminergic signaling produced little effect on TN axon projection. In adult retinas, inactivation of GnRH receptors disrupted the patterns of TN axon distribution, and depletion of DA-IPCs abolished the TN axons. When DA-IPCs regenerated, the TN axons reappeared. Together, the data suggest that in developing zebrafish retinas GnRH signaling is required for TN axon projection, whereas in adult retinas activation of GnRH and dopaminergic signaling transduction is required for normal distribution of the TN axons.

Highlights of Swedish ophthalmology in the 20th century

Highlights of Swedish ophthalmology in the 20th century

Novel processes invaginate the pre-synaptic terminal of retinal bipolar cells

Mixed-rod cone bipolar (Mb) cells of goldfish retina have large synaptic terminals (10 microm in diameter) that make 60-90 ribbon synapses mostly onto amacrine cells and rarely onto ganglion cells and, in return, receive 300-400 synapses from gamma-aminobutyric acid (GABA)-ergic amacrine cells. Tissue viewed by electron microscopy revealed the presence of double-membrane-bound processes deep within Mb terminals. No membrane specializations were apparent on these invaginating processes, although rare vesicular fusion was observed. These invaginating dendrites were termed "InDents". Mb bipolar cells were identified by their immunoreactivity for protein kinase C. Double-label immunofluorescence with other cell-type-specific labels eliminated Müller cells, efferent fibers, other Mb bipolar cells, dopaminergic interplexiform cells, and somatostatin amacrine cells as a source of the InDents. Confocal analysis of double-labeled tissue clearly showed dendrites of GABA amacrine cells, backfilled ganglion cells, and dendrites containing PanNa immunoreactivity extending into and passing through Mb terminals. Nearly all Mb terminals showed evidence for the presence of InDents, indicating their common presence in goldfish retina. No PanNa immunoreactivity was found on GABA or ganglion cell InDents, suggesting that a subtype of glycine amacrine cell contained voltage-gated Na channels. Thus, potassium and calcium voltage-gated channels might be present on the InDents and on the Mb terminal membrane opposed to the InDents. In addition to synaptic signaling at ribbon and conventional synapses, Mb bipolar cells may exchange information with InDents by an alternative signaling mechanism.

Local Retinal Circuits of Melanopsin-Containing Ganglion Cells Identified by Transsynaptic Viral Tracing

Intrinsically photosensitive melanopsin-containing retinal ganglion cells (ipRGCs) control important physiological processes, including the circadian rhythm, the pupillary reflex, and the suppression of locomotor behavior (reviewed in [1]). ipRGCs are also activated by classical photoreceptors, the rods and cones, through local retinal circuits [2, 3]. ipRGCs can be transsynaptically labeled through the pupillary-reflex circuit with the derivatives of the Bartha strain of the alphaherpesvirus pseudorabies virus(PRV) [4, 5] that express GFP [6-12]. Bartha-strain derivatives spread only in the retrograde direction [13]. There is evidence that infected cells function normally for a while during GFP expression [7]. Here we combine transsynaptic PRV labeling, two-photon laser microscopy, and electrophysiological techniques to trace the local circuit of different ipRGC subtypes in the mouse retina and record light-evoked activity from the transsynaptically labeled ganglion cells. First, we show that ipRGCs are connected by monostratified amacrine cells that provide strong inhibition from classical-photoreceptor-driven circuits. Second, we show evidence that dopaminergic interplexiform cells are synaptically connected to ipRGCs. The latter finding provides a circuitry link between light-dark adaptation and ipRGC function.

Expression of the dopamine transporter in rat and bullfrog retinas

We examined the expression of the dopamine transporter in rat and bullfrog retinas by immunohistochemistry. In both species, the dopamine transporter was strongly expressed in somata and processes of all dopaminergic amacrine cells. In contrast, no immunoreactivity for dopamine transporter was observed in cholinergic amacrine cells. In rat dopaminergic interplexiform cells, dopamine transporter immunoreactivity was also observed on the ascending processes terminating in the outer plexiform layer. Furthermore, the labeling for dopamine transporter diffusely appeared in both outer and inner plexiform layers. This expression profile of the dopamine transporter suggests that dopamine may be taken up not only in the synapses but also extrasynaptically by dopamine transporter, diffusely distributed in both plexiform layers.

Olfactoretinal centrifugal input modulates zebrafish retinal ganglion cell activity: a possible role for dopamine‐mediated Ca2+ signalling pathways

The vertebrate retina receives centrifugal input from the brain. In zebrafish, the major centrifugal input originates in the terminal nerve (TN). TN cell bodies are located in the olfactory bulb and ventral telencephalon. The TN projects axons to the retina where they branch in the inner plexiform layer (IPL) and synapse onto several inner retinal cell types, including dopaminergic interplexiform cells (DA-IPCs). This olfactoretinal centrifugal input plays a role in modulating retinal ganglion cell (RGC) activity, probably via dopamine-mediated Ca2+ signalling pathways. Normally, dopamine inhibits RGC firing by decreasing the inward Ca2+ current. Olfactory stimulation with amino acids decreases dopamine release in the retina, thereby reducing dopaminergic inhibition of RGCs. This model of olfacto-visual integration was directly tested by recording single-unit RGC activity in response to olfactory stimulation in the presence or absence of dopamine receptor blockers. Stimulation of the olfactory neurones increased RGC activity. However, this effect diminished when the dopamine D1 receptors were pharmacologically blocked. In isolated RGCs, the application of dopamine or a dopamine D1 receptor agonist decreased voltage-activated Ca2+ current and lowered Ca2+ influx. Together, the data suggest that olfactory input has a modulatory effect on RGC firing, and that this effect is mediated by dopamine D1 receptor-coupled Ca2+ signalling pathways.

Two Visual Processing Pathways Are Targeted by Gonadotropin-Releasing Hormone in the Retina

In fish the terminal nerve is comprised of a group of cells with somata adjacent to the olfactory bulb and processes that extend both anteriorly to the olfactory mucosa and posteriorly to the telencephalon. In teleost fish an additional group of axons extends along the optic tract and delivers putative neuromodulators to the retina. One peptide – gonadotropin-releasing hormone (GnRH) – has been implicated as a prime candidate neuromodulator based on electrophysiological evidence that exogenous application influences neural activity. Here we describe the expression patterns of two GnRH receptor subtypes in the retina of a teleost fish, Astatotilapia (Haplochromis) burtoni. The type 1 GnRH receptor (GnRH-R1) was expressed in cells of the amacrine cell layer – where lateral inputs affect the flow of visual information from photoreceptors to the brain – and in a distribution and location pattern similar to dopaminergic interplexiform cells. Immunohistochemical labeling of GnRH fibers revealed varicosities along terminal nerve axons near the amacrine cell layer and near cells immunoreactive for tyrosine hydroxylase, a dopaminergic cell marker. This finding supports an existing model that the terminal nerve forms synapses with dopaminergic interplexiform cells. Surprisingly, the type 2 GnRH receptor (GnRH-R2) was abundantly expressed in ganglion cells, which lie along the direct pathway of visual information to the brain. These data suggest that GnRH from the TN could broadly influence processing of retinal signals both in lateral processing circuits through GnRH-R1 and in the vertical throughput pathway through GnRH-R2.

Terminal nerve and vision

The vertebrate retina receives efferent input from different parts of the central nervous system. Efferent fibers are thought to influence retinal information processing but their functional role is not well understood. One of the best-described retinopetal fiber systems in teleost retinae belongs to the terminal nerve complex. Gonadotropin-releasing hormone (GnRH) and molluscan cardioexcitatory tetrapeptide (FMRFamide)-containing fibers from the ganglion of the terminal nerve form a dense fiber plexus in the retina at the border of the inner nuclear and inner plexiform layer. Peptide-containing fibers surround and contact perikarya of dopaminergic interplexiform cells in teleost retina. In vitro experiments demonstrated that exogenously supplied GnRH mediates dopaminergic effects on the membrane potential and on the morphology of dendritic tips (spinules) of cone horizontal cells. These effects can be specifically blocked by GnRH-antagonists, indicating that the release of dopamine and dopamine-dependent effects on light adaptation of retinal neurons are affected by the terminal nerve complex. Recent data have shown that olfactory information has an impact on retinal physiology, but its precise role is not clear. The efferent fiber of the terminal nerve complex is one of the first retinopetal fiber systems for which the sources of the fibers, their cellular targets, and several physiological, morphological, and behavioral effects are known. The terminal nerve complex is therefore a model system for the analysis of local information processing which is influenced by a distinct fiber projection.

Olfactory input increases visual sensitivity in zebrafish: a possible function for the terminal nerve and dopaminergic interplexiform cells.

Centrifugal innervation of the neural retina has been documented in many species. In zebrafish Danio rerio, the only so-far described centrifugal pathway originates from terminal nerve (TN) cell bodies that are located in the olfactory bulb. Most of the TN axons terminate in the forebrain and midbrain, but some project via the optic nerve to the neural retina, where they synapse onto dopaminergic interplexiform cells (DA-IPCs). While the anatomical pathway between the olfactory and visual organs has been described, it is unknown if and how olfactory signals influence visual system functions. We demonstrate here that olfactory input is involved in the modulation of visual sensitivity in zebrafish. As determined by a behavioral assay and by electroretinographic (ERG) recording, zebrafish visual sensitivity was increased upon presentation of amino acids as olfactory stimuli. This effect, however, was observed only in the early morning hours when zebrafish are least sensitive to light. The effect of olfactory input on vision was eliminated after lesion of the olfactory bulbs or after the destruction of DA-IPCs. Intraocular injections of a dopamine D(2) but not a D(1) receptor antagonist blocked the effect of olfactory input on visual sensitivity. Although we cannot exclude the involvement of other anatomical pathways, our data suggest that the TN and DA-IPCs are the prime candidates for olfactory modulation of visual sensitivity.

Synaptic plasticity and functionality at the cone terminal of the developing zebrafish retina.

Previous studies have analyzed photoreceptor development, some inner retina cell types, and specific neurotransmitters in the zebrafish retina. However, only minor attention has been paid to the morphology of the synaptic connection between photoreceptors and second order neurons even though it represents the transition from the light sensitive receptor to the neuronal network of the visual system. Here, we describe the appearance and differentiation of pre- and postsynaptic elements at cone synapses in the developing zebrafish retina together with the maturation of the directly connecting second order neurons and a dopaminergic third order feedback-neuron from the inner retina. Zebrafish larvae were examined at developmental stages from 2 to 7dpf (days postfertilization) and in the adult. Synaptic maturation at the photoreceptor terminals was examined with antibodies against synapse associated proteins. The appearance of synaptic plasticity at the so-called spinule-type synapses between cones and horizontal cells was assessed by electron microscopy, and the maturation of photoreceptor downstream connection was identified by immunocytochemistry for GluR4 (AMPA-type glutamate receptor subunit), protein kinase beta(1) (mixed rod-cone bipolar cells), and tyrosine hydroxylase (dopaminergic interplexiform cells). We found that developing zebrafish retinas possess first synaptic structures at the cone terminal as early as 3.5dpf. Morphological maturation of these synapses at 3.5-4dpf, together with the presence of synapse associated proteins at 2.5dpf and the maturation of second order neurons by 5dpf, indicate functional synaptic connectivity and plasticity between the cones and their second order neurons already at 5dpf. However, the mere number of spinules and ribbons at 7dpf still remains below the adult values, indicating that synaptic functionality of the zebrafish retina is not entirely completed at this stage of development.

Dopamine Has a Critical Role in Photoreceptor Degeneration in the rd Mouse

Photoreceptors receive paracrine input from dopaminergic interplexiform cells. Rod photoreceptors in the rd mouse degenerate rapidly due to a specific gene defect. We investigated the effects of dopamine on rd mouse photoreceptors in retinal organ culture. Retinas were harvested from rd or wild-type mice at postnatal day 2 and grown in organ culture for 27 days. When antagonists for either D1- or D2-family dopamine receptors were added to the media, photoreceptor degeneration was blocked. Furthermore, when dopamine was depleted by the addition of 6-hydroxydopamine and pargyline, photoreceptor survival appeared comparable to wild-type retinal cultures. The addition of a dopamine agonist induced photoreceptor degeneration in dopamine-depleted rd organ cultures. In all cases, photoreceptors maintained robust staining of opsin. These results demonstrate that dopamine antagonists or dopamine depletion blocks photoreceptor degeneration and that dopamine is necessary for photoreceptor degeneration in the rd mouse retinal organ culture model, indicating that dopamine antagonists may represent a therapeutic strategy in retinal degenerative disease.

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.

Influence of light and neural circuitry on tyrosine hydroxylase phosphorylation in the rat retina

Light has been shown to increase dopamine synthesis and release in vertebrate retinas, but the retinal circuits mediating the light signal are unknown. We utilized three antibodies which recognize phosphorylated forms of tyrosine hydroxylase (TH) at serines 19, 31, and 40 to study the effects of light and neuroactive drugs on TH phosphorylation in the rat retina. Phosphorylated TH and total TH immunoreactivities were co-localized exclusively in retinal neurons whose shape and location are characteristic of dopaminergic interplexiform cells. Phosphorylated TH was weak to absent in darkness, but light strongly stimulated phosphorylation in all the three serine residues. Light-induced phosphorylation of TH induction by light was uniformly blocked by a combination of NMDA and AMPA glutamate receptor antagonists. In darkness, the combination of NMDA+AMPA induced phosphosphorylation of TH at serines 19 and 40 but it was weak at serine 31. A GABA A antagonist had the same effect. An agonist of depolarizing (ON) bipolar cells, l-(+)-2-amino-4-phosphonobutyric acid, did not prevent light-induced phosphorylated TH formation. Carbachol, a non-specific cholinergic agonist, selectively induced phosphorylation of TH at serine 31 in darkness, an effect which was blocked by the nicotinic antagonist, d-tubocurarine. These results show that retinal circuits involving glutamatergic, GABAergic and cholinergic synapses influence phospho-TH formation at different serine residues in this enzyme. Gamma amino butyric acid (GABA) and glutamate influence TH phosphorylation at serines 19 and 40, whereas cholinergic inputs affect its phosphorylation at serine 31.

GABA enhances short wavelength-sensitive cone input and reduces red cone input to carp L-type horizontal cells

Light responses of cone-driven horizontal cells were recorded intracellularly in the isolated superfused carp retina and the effects of γ-aminobutyric acid (GABA) on signals from red-sensitive (R-) and short-wavelength-sensitive (S-) cones (green cones and/or blue cones) were studied. In the presence of a bright red (694 nm) background light, which substantially suppressed signal from R-cones, the responses of L-type horizontal cells (L-HCs) to 532-nm flashes, predominantly driven by the S-cone input, were potentiated by application of GABA. In contrast, the responses of these cells to 694-nm flashes driven by the R-cone input, were suppressed, when signal from S-cones was suppressed by a bright 532-nm background light. Both the effects could be reversed by co-application of bicuculline, suggesting the involvement of GABA A receptors. It was unlikely that the potentiation by GABA of the S-cone driven responses of the L-HCs was mediated by actions of GABA on the cone photoreceptors. The dual action of GABA persisted in the dopamine-depleted retina, indicating no involvement of the dopaminergic interplexiform cells. We speculate that this dual action may be partially due to differential modulation by GABA of different postsynaptic mechanisms respectively mediating signal transfer from R-cones and S-cones to L-HCs.

Effects of Dopamine Depletion on Visual Sensitivity of Zebrafish

The visual sensitivity of zebrafish in which the retinal dopaminergic interplexiform cells (DA-IPCs) were destroyed by 6-hydroxydopamine was measured behaviorally. During the first 6-8 min of dark adaptation, visual thresholds of DA-IPC-depleted animals were similar to those of control animals. Thereafter, their visual thresholds were elevated so that by 14-18 min of dark adaptation, they were 2-3 log units above those of control animals. In DA-IPC-depleted animals, the electroretinogram was normal in terms of light sensitivity and waveform, but the light threshold for eliciting a ganglion cell discharge was raised by 1.8 log units as compared with control animals. No obvious rod system function was detected in DA-IPC-depleted animals as measured behaviorally. Partial rescue of the behavioral visual sensitivity loss in DA-IPC-depleted animals occurred when dopamine or a long-acting dopamine agonist (2-amino-6, 7-dihydroxy-1, 2, 3, 4-tetrahydronaphthalene hydrobromide) were injected intraocularly. Our data suggest that the principal visual defect shown by DA-IPC-depleted animals is attributable to effects occurring in the inner retina, mainly on rod signals. We also show that dopamine is involved in mediating the effect of the circadian clock on visual sensitivity.

Disruption of the Olfactoretinal Centrifugal Pathway May Relate to the Visual System Defect innight blindness bMutant Zebrafish

We describe here a dominant mutation, night blindness b (nbb), which causes an age-related visual system defect in zebrafish. At 4-5 months of age, dark-adapted nbb(+/-) mutants show abnormal visual threshold fluctuations when measured behaviorally. Light sensitizes the animals; thus early dark adaptation of nbb(+/-) fish is normal. After 2 hr of dark adaptation, however, visual thresholds of nbb(+/-) mutants are raised on average 2-3 log units, and rod system function is not detectable. Electroretinograms recorded from nbb(+/-) mutants are normal, but ganglion cell thresholds are raised in prolonged darkness, suggesting an inner retinal defect. The visual defect of nbb(+/-) mutants may be likely caused by an abnormal olfactoretinal centrifugal innervation; in nbb(+/-) mutants, the olfactoretinal centrifugal projection to the retina is disrupted, and the number of retinal dopaminergic interplexiform cells is reduced. A similar visual defect as shown by nbb(+/-) mutants is observed in zebrafish in which the olfactory epithelium and olfactory bulb have been excised. Homozygous nbb fish display an early onset neural degeneration throughout the CNS and die by 7-8 d of age.

Differential reinnervation of retinal bipolar cell dendrites and axon terminals by dopamine interplexiform cells following dopamine depletion with 6-OHDA

Depletion of retinal dopamine in goldfish increases light sensitivity at photopic backgrounds. As horizontal cells appear not to be involved with this effect (Yazulla and Studholme [1995] Vis. Neurosci. 12:827-837), we investigated the innervation patterns of the ON rod/cone bipolar cells (ON-BC) by dopaminergic interplexiform cells (DA-IPCs) normally and during the period of neogeneration of new DA-IPCs at the marginal zone following DA-IPC destruction. DA-IPCs were destroyed via intraocular injection of 6-hydroxydopamine over 2 successive days. Controls and 1 year post-injection retinas were double labeled for protein kinase C and tyrosine hydroxylase (TH) immunocytochemistry to identify the ON-BCs and the DA-IPCs, respectively. Double-labeled 25 microns tissue sections were examined on a confocal laser scanning microscope by using dual channel immunofluorescence acquisition. Image stacks were analyzed for DA-IPC/ON-BC contacts in the distal inner nuclear layer (INL) and inner plexiform layer (IPL). Image stacks were rotated 180 degrees with respect to each other and reanalyzed to determine potential randomness of the contacts. For control retinas there were 1.8 contacts/axon terminal in the IPL (n = 165) and 9.4 contacts/ON-BC in the distal INL (n = 28). At 1 year after injection, reinnervation of TH-immunoreactive boutons in the retina recovered to 16% of control in the IPL but only 10% in the distal INL. Establishment of DA-IPC/ON-BC contacts recovered to 36% of control for ON-BC axon terminals (n = 103), whereas there was no recovery of contacts in the distal INL (n = 30). Reinnervation of ON-BC by DA-IPCs preferentially targets the axon terminals. The absence of reinnervation of bipolar cell dendrites by DA-IPCs may account for the persistence of the increased light sensitivity following retinal dopamine depletion. Thus, dopamine input to ON-BCs in the outer retina maybe involved in setting background sensitivity under photopic conditions.

A MODEL OF AN ADAPTIVE RECEPTOR BASED ON THE RETINA

A model of the retina based on biological data is developed to study the processing of images by the retina as a model for an adaptive receptor. The direct visual pathway (photoreceptors, bipolar and ganglion cells) and the horizontal units and their gap-junctions, as well as the dopaminergic interplexiform cells and dopamine release are modeled. A small network of 12 units is first simulated in order to study the behavior of the different components involved, and the role played by lateral inhibition is analyzed. A 64×64 photoreceptors model, containing 16,448 units, is then simulated. The following properties were found for stimuli that remain constant in time: (i) The difference between the time constants of the photoreceptors and the horizontal units lead to both the formation of a transient output image that contains pertinent features of the input, and the formation of a reverse after-image when the stimulus is removed. (ii) Global modifications of lateral inhibition strength allow either contrast rendering or contour enhancement. Local modifications of lateral inhibition strength bring about adaptation to local input characteristics.

Horizontal cell spinule dynamics in fish are affected by rearing in monochromatic light

Blue acaras ( Aequidens pulcher, Cichlidae) were reared for 1 yr in white or monochromatic “red”, “green” and ”bluerd lights to study the function and control mechanisms of horizontal cell (HC) spinules in the synaptic pedicles of cones. Ratios of spinules per synaptic ribbon (S/R) were determined in tangential sections in both single and double cones. We found that the S/R ratios in light adapted retinae decreased with decreasing wavelength of the rearing light in all cone types. Conversely, there was an increasing number of incompletely formed spinules with the highest frequency in the blue light group. Dark adaptation resulted in the complete degradation of mature spinules. However, significant numbers of incompletely degraded spinules were observed in the group reared in blue light. Fish reared in blue light which were transferred to white light formed mature spinules when light adapted and still had vestigial spinules when dark adapted. The mechanisms of spinule formation and degradation and the control of spinule dynamics appear to be fully developed in fish reared in monochromatic light. However, long-term chromatic deprivation seems to induce a compensatory modulation of spinule dynamics. A working hypothesis is formulated that interprets the observed effects as manifestations of differences in the activition of dopaminergic interplexiform cells (light adapted) and the sensitivity to glutamate of HCs (dark adapted). Our findings are consistent with the hypothesis that spinules are involved in signin-verting feedback transmission from HCs to cones.