Interaction between the horizontal cells of the turtle retina]

The interaction between horizontal cells in the turtle retina was tested by means of two microelectrodes, polarizing and recording ones impaling two cells at different distances between each other. The direct electrical coupling is shown to exist between L-cells of the same type (type I - with big receptive fields and type II - with small receptive fields). The value of this coupling changes with the conditions of illumination as well as with the level of the membrane potential. This can be accounted for by the known properties of subsynaptic and nonsynaptic membranes of the horizontal cells. There is no direct electrical coupling between L-cells of different types. However strong hyperpolarization of L-cells of type I by extrinsic current or by a light annulus evokes a depolarization in L-cells of type II. This indirect interaction between L-cells also dependent on the conditions of illumination may be explained by a mechanism of feedback between the horizontal cells and photoreceptors. Polarization of L-cells of both types has no effect on the cells of C-type.     

Dark- and light-induced changes in coupling between horizontal cells in mammalian retina

Retinal horizontal cells exhibit large receptive fields derived from their extensive electrical coupling by means of gap junctions. The conductance of these gap junctions seems to be regulated by dopamine acting through a cAMP-mediated cascade. There is now abundant evidence that extracellular dopamine levels vary with changes in ambient light intensity, suggesting that changes in the dark/light adaptational state of the retina can modulate coupling between horizontal cells. We studied this question in the mammalian retina by determining the effects of ambient light levels, in the form of changing background light intensity, on the coupling profiles of A- and B-type horizontal cells in the rabbit. Changes in coupling were assessed by measurements of the space constants of the syncytium formed by horizontal cells and the intercellular spread of the biotinylated tracer Neurobiotin. Our results indicate that dark-adapted horizontal cells show relatively weak coupling. However, presentation of background lights as dim as one-quarter log unit above rod threshold resulted in increases in both the averaged extent of tracer coupling and space constants of A- and B-type horizontal cells. Coupling expanded further as background light intensities were increased by 1-1.5 log units, after which additional light adaptation brought about an uncoupling of cells. Coupling reached its minimum at light intensities about 3 log units above rod threshold, after which, with further light adaptation, it stabilized at levels close to those seen in dark-adapted retinas. Our results indicate that electrical coupling between mammalian horizontal cells is modulated dramatically by changes in the adaptational state of the retina: coupling is maximized under dim ambient light conditions and diminishes as the retina is dark or light adapted from this level.     

Horizontal cell electrical coupling in the giant danio: synaptic modulation by dopamine and synaptic maintenance by calcium

Electrical synapses, and their structural manifestation, gap junctions, are critical elements of retinal circuitry. These synapses are subject to both rapid modulation and slower structural changes by physiological signals which mediate changes in the adaptational state of the retina. The electrical synapses of fish retinal horizontal cells are an excellent preparation for in vitro studies of electrical synapses. We have examined the rapid modulation of electrical coupling by dopamine and effects on the expression and maintenance of electrical synapses by cell calcium in pairs of horizontal cells isolated from retinas of the giant danio (Danio aquipinnatus). We report that rapid modulation by dopamine reduces junctional conductance by modifying gap junction channel gating, while maintaining cells in reduced calcium medium, and lowering; intracellular calcium concentration, results in the loss of electrical coupling. The effects of calcium on synaptic maintenance may be related to structural changes observed in horizontal cell electrical synapses during light adaptation.     

Junctions form between catfish horizontal cells in culture

Cone horizontal cells from the catfish retina extend out processes after a few days in culture that sometimes contact adjacent cone horizontal cells. Two types of specialized junctions were observed by electron microscopy along the newly formed contact areas. One junctional type consisted of prominent electron-dense material along and just under the plasma membrane of one or both of the contacting elements. Sometimes vesicle clusters were associated with these junctions. The other type of junction showed some electron-dense material along the membranes of both processes and patchy areas of close membrane apposition resembling gap junctions. In about half of the cases tested, electrical coupling was detected between cone horizontal cells that had made contact in culture. In no case was the coupling as tight as is typically found between horizontal cells that had formed gap junctions in vivo.     

Endogenous dopaminergic regulation of horizontal cell coupling in the mammalian retina

Horizontal cells in an isolated wholemount preparation of the mouse retina were injected with Lucifer yellow and neurobiotin to characterize both the pattern of gap junctional connectivity and its regulation by dopamine. The injected horizontal cells had a uniform morphology of a round cell body, a compact dendritic tree, and an axon, which could sometimes be traced to an expansive terminal system. The dendro-dendritic gap junctions between neighboring cells mediated both weak Lucifer yellow dye coupling and strong neurobiotin tracer coupling. The extent of the tracer coupling was decreased by either exogenous dopamine (100 microM) or cyclic adenosine monophosphate (cAMP) analogs and was significantly increased by the D1 antagonist SCH 23390 (10 microM). These results provide the first evidence in the mammalian retina that the gap junctions between horizontal cells are endogenously regulated by dopamine, which acts through D1 receptors to increase the intracellular cAMP. It has been proposed that the gap junctional coupling between horizontal cells is mediated by connexin 32 (Cx32), but the pattern and dopaminergic regulation of horizontal cell coupling were unaffected in Cx32-knockout mice, ruling out the possible involvement of Cx32. Every tracer-coupled horizontal cell showed calbindin immunoreactivity, and vice versa, providing strong evidence that the horizontal cells in the mouse retina comprise a single cell type. Like the axonless horizontal cells in other mammalian retinas, the axon-bearing horizontal cells in the mouse retina are coupled by gap junctions that are permeable to Lucifer yellow and dopamine sensitive, suggesting that the mouse horizontal cells have hybrid properties to compensate for the absence of axonless horizontal cells.     

Functional expression of connexin57 in horizontal cells of the mouse retina

Horizontal cells are interneurons of the vertebrate retina that exhibit strong electrical and tracer coupling but the identity of the channel-forming connexins has remained elusive. Here we show that horizontal cells of the mouse retina express connexin57 (Cx57). We have generated Cx57-deficient mice by replacing the Cx57 coding region with a lacZ reporter gene, expressed under control of the endogenous Cx57 promoter. These mice were fertile and showed no obvious anatomical or behavioural abnormalities. Cx57 mRNA was expressed in the retina of wild-type littermates but was absent from the retina of Cx57-deficient mice. Previously reported results that the Cx57 gene was very weakly expressed in several other mouse tissues turned out to be unspecific. Cx57 mRNA is abundantly expressed in the retina and weakly in the thymus of adult mice but absent in all other adult tissues tested, including brain. Furthermore, Cx57 is expressed in embryonic kidney at E16.5 to E18.5 days post-conception, as indicated by the pattern of lacZ expression. Within the retina, lacZ signals were assigned exclusively to horizontal cells based on co-localization with cell-type-specific marker proteins. Microinjection of Neurobiotin into horizontal cells of isolated retinae revealed less than 1% of tracer coupling in Cx57-deficient retinae compared with wild-type controls. Cx57 is the first connexin identified in mammalian horizontal cells and the first connexin whose expression is apparently restricted to only one type of neuron.     

Retinoic Acid Modulates Gap Junctional Permeability Between Horizontal Cells Of The Mammalian Retina

In the retina, all-trans retinoic acid (at-RA) could function as a light signal because its production increases with the level of illumination. Given the well-established effects of retinoic acid on cell coupling in other tissues, it is possible that the changing levels of at-RA modulate the gap junctional permeability between retinal neurons. This study examines the effects of retinoic acid on horizontal cell coupling, which is known to be modulated by the ambient light level. Single horizontal cells were injected under visual control with either Neurobiotin (mouse retina) or Lucifer yellow (rabbit retina) and the extent of tracer coupling or dye coupling was used to monitor the gap junctional permeability. In the mouse retina, the injection of Neurobiotin revealed a network of approximately 150-250 tracer-coupled horizontal cells. The tracer coupling was completely abolished by incubating the retina in 150 microM at-RA for 35 min. In the rabbit retina, the injection of Lucifer yellow into A-type horizontal cells revealed networks of approximately 15-30 dye-coupled horizontal cells. Incubation in 150 microM at-RA reduced the dye coupling within 12 min and complete uncoupling was achieved after 35 min. The uncoupling effects of at-RA in the mouse and rabbit retinas were concentration- and time-dependent and they were reversible after washout. The coupling was not affected by either the 9-cis form of retinoic acid or by at-RA that had been isomerized by intensive light. The uncoupling effect of at-RA persisted following treatment with a D1 receptor antagonist and thus was dopamine-independent. This study has established that at-RA is able to modulate the gap junctional permeability between horizontal cells in the mammalian retina, where its light-dependent release has already been demonstrated.     

Choline acetyltransferase is found in terminals of horizontal cells that label with GABA, nitric oxide synthase and calcium binding proteins in the turtle retina

In this study, we discriminated the various types of horizontal cell in the turtle retina on their content of neuroactive substances. Double label immunocytochemistry was performed on sectioned and wholemount retina using antisera to neural- and endothelial-nitric oxide synthase (nNOS, and eNOS), calretinin (CR), calbindin (CB), gamma-aminobutyric acid (GABA) and choline acetyltransferase (ChAT). H1 cells and their axon terminals label with CR, CB and GABA. Only H1 axon terminals label with eNOS. H2 cells contain CB, CR, nNOS and GABA maybe in their dendrites. H3 cells label only with nNOS. The localization of nNOS in the H2 and H3 cells is a novel finding. None of these antibodies labels H4 cells. The photoreceptor subtypes have been differentiated by different intensity of labeling with CB. The accessory member of the double cone is less intensely labeled with CB than the principal member and rods and blue cones do not appear to label at all. ChAT-IR is located in terminal boutons of H1 and H2 horizontal cells and H1 axon terminals and these boutons contact rods and all spectral types of cones. Clearly, GABA is present in H1 horizontal cells and may be used in neurotransmission between horizontal cells and possibly for feedback pathways to photoreceptors. The evidence of nNOS immunoreactivity in H2 and H3 horizontal cells, combined with available physiological evidence, suggests that NO may be involved in electrical coupling and/or modulation of synaptic input to these types of cells. Furthermore, our results raise the possibility that cholinergic synaptic transmission may occur from horizontal cell processes to photoreceptors in the outer plexiform layer of the turtle retina.     

Modulation of cone horizontal cell activity in the teleost fish retina. III. Effects of prolonged darkness and dopamine on electrical coupling between horizontal cells

The effects of prolonged darkness and dopamine on the coupling between horizontal cells in the isolated, superfused white perch retina were studied. Two assays of coupling were employed; area versus amplitude relationships (area-response curves) and the diffusion of the fluorescent dye Lucifer yellow from intracellularly injected cells to neighboring cells. In prolonged dark-adapted retinas, area-response curves were difficult to determine because of the small light responses; however, light-evoked responses did not increase in size when light spots were larger than 0.8 mm in diameter. Following the presentation of dim background illumination that partially sensitized the retina, responses to light spots of various sizes were enhanced and an area-response curve could be constructed. Subsequent presentation of moderate background illumination that more fully sensitized the retina resulted in reduced responses to small spots (less than 1.6 mm in diameter) and enhanced responses to large spot or full-field stimuli. In retinas exposed to moderate background illumination, Lucifer yellow injected intracellularly into cone horizontal cells diffused into many neighboring horizontal cells. The coupled cells were very similar in morphology, suggesting they were of the same type. In prolonged dark-adapted retinas, on the other hand, the dye was usually restricted to the injected cell and a few adjacent cells. These results indicate that coupling between cone horizontal cells is modulated by prolonged darkness and background illumination. Following dopamine (50 microM) application, in both 6-OHDA-treated and untreated retinas, changes in area-response curves of cone horizontal cells were observed just opposite to those that occurred when prolonged dark-adapted retinas were exposed to background illumination. That is, following 5 min application of dopamine to the retina, responses to small spots (less than 2 mm in diameter) increased in size while responses to larger spots decreased in amplitude compared with control responses. Following 20 min of superfusion with dopamine, the recorded responses were very small, and an accurate area-response curve could not be determined. Following dopamine application to light-sensitized retinas, Lucifer yellow was restricted to the injected cells or to the injected cell and a few neighboring cells. The results suggest that the modulation of coupling between cone horizontal cells by prolonged darkness and background illumination may be mediated by dopamine. Spatial properties of rod horizontal cells were also examined.(ABSTRACT TRUNCATED AT 400 WORDS)     

Dopaminergic mechanisms underlying the reduction of electrical coupling between horizontal cells of the turtle retina induced by d-amphetamine, bicuculline, and veratridine

Previous studies have shown that dopamine, bicuculline, or d-amphetamine reduce the electrical and dye-coupling between the axon terminals of the horizontal cells of the turtle retina (see Piccolino et al., 1984). In the present study we observed similar effects following the application of veratridine. The actions of all these drugs were prevented by dopamine antagonists acting on D1 receptors such as flupenthixol and SCH 23390. However, in contrast to dopamine, the actions of d-amphetamine, bicuculline, and veratridine were attenuated or abolished by pharmacological agents (such as 6-OH-dopamine, alpha-methyl-p-tyrosine, or reserpine) known to reduce the release of dopamine from dopaminergic neurons. Moreover, the actions of veratridine and bicuculline were prevented by tetrodotoxin, indicating that one or more neurons in the dopamine pathway are spike-generating. We conclude that d-amphetamine, bicuculline, and veratridine reduce electrical coupling between the axon terminals of the turtle horizontal cells by promoting the release of endogenous dopamine from the dopaminergic amacrine cells previously identified (Witkovsky et al., 1984). Electron-microscopic observations revealed that 6-OH-dopamine selectively attacked this population of amacrine cells. No degenerating terminals were found adjacent to the horizontal cell axon terminals. On this basis, we postulate that dopamine reaches the horizontal cell by diffusion through the extracellular space.     

Spatial density and immunoreactivity of bipolar cells in the macaque monkey retina.

The anatomical substrates of spatial and color vision in the primate retina are investigated by measuring the immunoreactivity and spatial density of bipolar, amacrine and horizontal cells in the inner nuclear layer of the macaque monkey retina. Bipolar cells can be distinguished from amacrine and horizontal cells by their differential immunoreactivity to antisera against glutamate, glycine, GABA, parvalbumin, calbindin (CaBP D-28K), and the L7 protein from mouse cerebellum. The spatial density of bipolar cells is compared to the densities of photoreceptors and ganglion cells at different retinal eccentricities. In the centralmost 2 mm, cone bipolar cells outnumber ganglion cells by about 1.4:1. The density of cone bipolar cells is thus high enough to allow for input to different (parasol and midget) ganglion cell classes by different (diffuse and midget) bipolar cell classes. The density gradient of cone bipolar cells follows closely that of ganglion cells in central retina but falls less steeply in peripheral retina. This suggests that the convergence of cone signals to the receptive fields of ganglion cells in the peripheral retina occurs in the inner plexiform layer. The density of cone bipolar cells is 2.5-4 times that of cones at all eccentricities studied, implying that cone connectivity to bipolar cells remains constant throughout the retina. Different subgroups of bipolar cells are distinguished by their relative immunoreactivity to the different antisera. All rod and cone bipolar cells show moderate to strong glutamate-like immunoreactivity. The bipolar cells that show weak to moderate GABA-like immunoreactivity are also labeled with the antiserum to the L7 protein and are thus identified as rod bipolar cells. Nearly half of all cone bipolar cells showed glycine-like immunoreactivity. The results suggest that the inhibitory neurotransmitter candidates GABA and glycine are segregated respectively in rod and cone bipolar cell pathways. A diffuse, cone bipolar cell type can be identified by the anti-parvalbumin and the anti-calbindin antisera. All horizontal cells show parvalbumin-like immunoreactivity. Nearly all amacrine cells show GABA-like or glycine-like immunoreactivity; a variety of subpopulations also show immunoreactivity to one or more of the other markers used.     

Horizontal Cells in the Monkey Retina: Cone connections and dendritic network

Horizontal cells of the macaque monkey retina were quantified and the number of cones converging onto an individual horizontal cell as well as the number of horizontal cells contacting a single cone were determined. This was done by combining data from individual horizontal cells stained by the Golgi method with the results of immunocytochemical staining described in the preceding paper (Röhrenbeck et al., 1989). The observation (Boycott et al., 1987) that all horizontal cells contact all cones in their dendritic field irrespective of cone type was confirmed. The particular cones contacted by the terminal aggregates of each horizontal cell were found. The dendritic fields of H1 and H2 cells increase with increasing eccentricity; close to the fovea H1 cells are smaller than H2 cells, at 6 mm eccentricity they are about the same size and in peripheral retina H1 cells are much larger than H2 cells. The density gradients of the two cell types balance their denritic field changes so that throughout the retina each and every cone synapses with 3–5 horizontal cells of each type. Horizontal cells of both cat (Wässle et al., 1978) and monkey retina follow the general rule that all cones in the dendritic fields are contacted, their perikarya form a regular mosaic and the boundaries of their dendritic fields are marked by the perikarya of their homologous neighbours.     

Gap junction particle density of horizontal cells in goldfish retinas lesioned with 6-OHDA

The I1 dopaminergic interplexiform cells of the fish retina are believed to modulate horizontal cell coupling by increasing gap junction resistance. Dopamine also modulates the morphology of horizontal cell gap junctions and mimics the effects of light adaptation. To determine whether the light-dependent changes in gap junction morphology are due to endogenous dopamine release, horizontal cell gap junctions were studied in goldfish retinas lacking dopaminergic neurons. Dopaminergic interplexiform cells were destroyed by intraocular injections of 6-hydroxydopamine in both eyes. After lesioning, fish were treated in one of four ways: (1) light-adapted, (2) dark-adapted (1 hour), (3) light-adapted and given an intraocular injection of dopamine, or (4) dark-adapted (1 hour) and injected with dopamine. The effectiveness of lesioning was evaluated by autoradiographic detection of [3H]-dopamine uptake in the retina of one eye. Retinas in which lesioning of the contralateral eye was deemed effective were processed for freeze-fracture electron microscopy and the particle density of horizontal cell gap junctions determined. Lesioned retinas, whether light- or dark-adapted, had elevated horizontal cell soma gap junction particle densities compared to lesioned retinas treated with dopamine. These results demonstrate that high soma gap junction particle densities can be correlated with the absence of dopamine and low densities associated with the presence of dopamine. The differences in gap junction particle density between lesioned and lesioned + dopamine-treatment were similar to differences between nonlesioned dark-adapted (1 hour) and light-adapted retinas, respectively. Therefore, the particle density of light- and dark-adapted soma gap junctions suggests a greater release of dopamine in light-adapted fish than in 1 hour dark-adapted fish.(ABSTRACT TRUNCATED AT 250 WORDS)     

Decrease of gap junction permeability induced by dopamine and cyclic adenosine 3':5'-monophosphate in horizontal cells of turtle retina

The axon terminals of the H1 horizontal cells of the turtle retina are electrically coupled by extensive gap junctions. Dopamine (10 nM to 10 microM) induces a narrowing of the receptive field profile of the H1 horizontal cell axon terminals, increases the coupling resistance between them, and decreases the diffusion of the dye Lucifer Yellow in the network formed by the coupled axon terminals. These actions of dopamine involve the activation of D1 receptors located on the membrane of the H1 horizontal cell axon terminals proper. Increases of the intracellular cyclic AMP concentration induced by either stimulating the adenylate cyclase activity with forskolin or inhibiting the phosphodiesterase activity with isobutylmethylxanthine, theophylline, aminophylline, or compound RO 20-1724 elicit effects similar to those of dopamine on the receptive field profile of the H1 horizontal cell axon terminals, on their coupling resistance, and on the diffusion of Lucifer Yellow in the axon terminal network. It is concluded that dopamine decreases the permeability of the gap junctions between the axon terminals of the H1 horizontal cells of the turtle retina and that this action probably involves cyclic AMP as a second messenger.     

Tracer coupling patterns of the ganglion cell subtypes in the mouse retina

It is now clear that electrical coupling via gap junctions is prevalent across the retina, expressed by each of the five main neuronal types. With the introduction of mutants in which selective gap junction connexins are deleted, the mouse has recently become an important model for studying the function of coupling between retinal neurons. In this study we examined the tracer‐coupling pattern of ganglion cells by injecting them with the gap junction‐permanent tracer Neurobiotin to provide, for the first time, a comprehensive survey of ganglion cell coupling in the wildtype mouse retina. Murine ganglion cells were differentiated into 22 morphologically distinct subtypes based on soma‐dendritic parameters. Most (16/22) ganglion cell subtypes were tracer‐coupled to neighboring ganglion and/or amacrine cells. The amacrine cells coupled to ganglion cells displayed either polyaxonal or wide‐field morphologies with extensive arbors. We found that different subtypes of ganglion cells were never coupled to one another, indicating that they subserved independent electrical networks. Finally, we found that the tracer‐coupling patterns of the 22 ganglion cell populations were largely stereotypic across the 71 retinas studied. Our results indicate that electrical coupling is extensive in the inner retina of the mouse, suggesting that gap junctions play essential roles in visual information processing. J. Comp. Neurol. 512:664–687, 2009. © 2008 Wiley‐Liss, Inc.     

Horizontal cells of the turtle retina. I. Light microscopy of golgi preparations

In Golgi preparations of turtle retina, four types of horizontal cells were observed and their morphological characteristics determined in vertical thick sections, whole mount preparations, and reconstructions from serial 1-μm sections. H1 consists of a nucleated, stellate cell body (H1CB) and an irregular, tuberous axon terminal (H1AT) connected by a slender axon. Both parts of these cells make contact with receptor cells. H1CB's appear to correspond to “L2-type cells” while H1AT's correspond to “L1-type cells” described in the physiological literature. H2 and H3 are axonless stellate cells which are similar to one another in vertical profile and may occasionally appear similar in horizontal view. In general, the dendritic tree is more densely branched and the density of receptor cell contacts is higher for H2 than for H3. H2-type cells may correspond to “R/G C-type cells.” H4 is also an axonless stellate cell type which is smaller than H2 or H3 at equivalent retinal locations. The dendritic fields of H1CB's vary widely, but systematically, in size and shape over the retina. Their size is inversely related to receptor cell density, and the shape of the dendritic tree varies from roughly circular in the central area to elliptical in the periphery of the retina.     

Cat cones have rod input: A comparison of the response properties of cones and horizontal cell bodies in the retina of the cat

The responses of horizontal cell bodies and cones in the retina of the cat have been studied by means of intracellular recording and Procion dye injection In an isolated, arterially perfused eyecup preparation. Comparison of the hyperpolarizing responses of these units to red and blue stimuli of different intensities indicated that all morphological varieties of horizontal cells and, additionally, cones themselves, had mixed rod and cone input. The rod input into horizontal cell bodies is thus explained on the basis of cone physiology. The half-saturating intensity of 441 nm stimuli for the rod input into cones and horizontal cells was about 400 quanta/μm₂/sec and about 160,000 quanta/μm₂/sec for the cone input. Little of this difference can be related to the different quantum catching abilities of rods and cones. The spatial properties of horizontal cell bodies and cones have been characterized using stimuli consisting of long slits in conjunction with a continuous cable model. Space constants for horizontal cells ranged from 210 μm to 410pm, whereas those for cones ranged from 50μm, or possibly less, to 180 μm. It is argued that horizontal cell bodies of the cat retina form electrical networks, and that the sizes of the receptive fields generated in these networks may be limited by the diameters of the primary and secondary dendrites of horizontal cells. The rod and cone fields of horizontal cell bodies were found to be nearly coextensive in space, arguing against the notion that substantial rod input came from distant, rod-dominated terminal arborizations.     

Opposite effects of ammonia and carbon dioxide on dye coupling between horizontal cells in the carp retina

Effects of ammonia (NH3) and carbon dioxide (CO2) on the membrane potential of horizontal cells and on dye coupling between the cells in isolated retinas of the carp (Cyprinus carpio) were investigated. Ammonia (less than 300 ppm NH3 in air) initially depolarized and subsequently hyperpolarized, while CO2 (10% in air) hyperpolarized the membrane potential of horizontal cells, accompanied by a diminution of both center and surround responses to spot and annular light stimuli. During the course of amplitude diminution, the center response consistently became smaller with NH3 and larger with CO2 than the surround response. In the presence of intravitreally applied DA (50 microM) or amphetamine (100 microM), a fluorescent dye Lucifer Yellow CH (LY) was found to be restricted to single injected horizontal cells. The presence of intravitreal haloperidol (100 microM) for 20-25 min or an exposure of the retina to NH3 for 5-10 min diffused the restricted LY from single injected cells to numerous neighboring cells. On the other hand, CO2 was found to restrict the injected dye to single cells, an effect similar to that of DA and opposite to that of NH3 and haloperidol. The results suggest that NH3 appears to act as a coupler while CO2 acts as an uncoupler on gap junctions between horizontal cells in the carp retina, presumably by changing the intracellular pH. In addition, a brief exposure of cells, marked with LY in the presence of DA, to the exciting light 426 nm was found to prevent the NH3-induced dye diffusion from single cells to their neighbors; the reason is unknown.     

Gap-junctional coupling of mammalian rod photoreceptors and its effect on visual detection

The presence of gap junctions between rods in mammalian retina suggests a role for rod-rod coupling in human vision. Rod coupling is known to reduce response variability, but because junctional conductances are not known, the downstream effects on visual performance are uncertain. Here we assessed rod coupling in guinea pig retina by measuring: (1) the variability in responses to dim flashes, (2) Neurobiotin tracer coupling, and (3) junctional conductances. Results were consolidated into an electrical network model and a model of human psychophysical detection. Guinea pig rods form tracer pools of 1 to ～20 rods, with junctional conductances averaging ～350 pS. We calculate that coupling will reduce human dark-adapted sensitivity ～10% by impairing the noise filtering of the synapse between rods and rod bipolar cells. However, coupling also mitigates synaptic saturation and is thus calculated to improve sensitivity when stimuli are spatially restricted or are superimposed over background illumination.