Receptive Field Size Of Horizontal Cells Research Articles

The light-induced reduction of horizontal cell receptive field size in the goldfish retina involves nitric oxide

Horizontal cells of the vertebrate retina have large receptive fields as a result of extensive gap junction coupling. Increased ambient illumination reduces horizontal cell receptive field size. Using the isolated goldfish retina, we have assessed the contribution of nitric oxide to the light-dependent reduction of horizontal cell receptive field size. Horizontal cell receptive field size was assessed by comparing the responses to centered spot and annulus stimuli and from the responses to translated slit stimuli. A period of steady illumination decreased the receptive field size of horizontal cells, as did treatment with the nitric oxide donor (Z)-1-[N-(2-aminoethyl)-N-(2-ammonioethyl)amino]diazen-1-ium-1,2-diolate (100 μM). Blocking the endogenous production of nitric oxide with the nitric oxide synthase inhibitor, N(G)-nitro-L-arginine methyl ester (1 mM), decreased the light-induced reduction of horizontal cell receptive field size. These findings suggest that nitric oxide is involved in light-induced reduction of horizontal cell receptive field size.

Strongly pH-Buffered Ringer's Solution Expands the Receptive Field Size of Horizontal Cells in the Carp Retina

By intracellular recordings, we studied the effects of pH buffering on the size of the receptive field and the extent of dye coupling of horizontal cells (HCs) in the light-adapted carp retina. These parameters were compared between data obtained in fortified Ringer's solution and those obtained in control bicarbonate Ringer's of the same pH (7.60). In Ringer's fortified with 10 mM HEPES or 15 mM Tris, the dye-coupling ratio of HCs increased by 71% and 70%, respectively. These fortified Ringer's solutions also depolarized the dark membrane potential and increased the light-evoked response. The HC receptive field profile could be described by the exponential decline in peak response amplitude to a slit of light moved tangentially from the recording electrode. Thus, the receptive field size was determined as a space constant proportional to (gj/gm)(1/2), where gj and gm denote gap and non-gap-junctional conductances. The HEPES- or Tris-fortified Ringer's significantly increased the space constant by 43% and 41%, respectively. Since dye coupling was increased in the fortified Ringer's, it is likely that gj increased more than gm as a result of alkalinization of the cytosol. Since HEPES has an aminosulfonate moiety, it has been assumed to close the hemi-channels of connexin 26, but the pH-buffering effects were essentially the same as those of Tris that has no aminosulfonate moiety. Therefore, it is unlikely that the closure of connexin 26 hemichannels is responsible for the change in the receptive field size of HCs.

Horizontal cell receptive fields are reduced in connexin57‐deficient mice

Horizontal cells are coupled by gap junctions; the extensive coupling of the horizontal cells is reflected in their large receptive fields, which extend far beyond the dendritic arbor of the individual cell. In the mouse retina, horizontal cells express connexin57 (Cx57). Tracer coupling of horizontal cells is impaired in Cx57-deficient mice, which suggests that the receptive fields of Cx57-deficient horizontal cells might be similarly reduced. To test this hypothesis we measured the receptive fields of horizontal cells from wildtype and Cx57-deficient mice. First, we examined the synaptic connections between horizontal cells and photoreceptors: no major morphological alterations were found. Moreover, horizontal cell spacing and dendritic field size were unaffected by Cx57 deletion. We used intracellular recordings to characterize horizontal cell receptive fields. Length constants were computed for each cell using the cell's responses to concentric light spots of increasing diameter. The length constant was dependent on the intensity of the stimulus: increasing stimulus intensity reduced the length constant. Deletion of Cx57 significantly reduced horizontal cell receptive field size. Dark resting potentials were strongly depolarized and response amplitudes reduced in Cx57-deficient horizontal cells compared to the wildtype, suggesting an altered input resistance. This was confirmed by patch-clamp recordings from dissociated horizontal cells; mean input resistance of Cx57-deficient horizontal cells was 27% lower than that of wildtype cells. These data thus provide the first quantification of mouse horizontal cell receptive field size and confirm the unique role of Cx57 in horizontal cell coupling and physiology.

Modulation of horizontal cell receptive fields in the light adapted goldfish retina

In the isolated goldfish retina, 700 nm background illumination increases the horizontal cell receptive field size, as measured with 565 nm slits of light, but decreases the receptive field size, when measured with 660 nm slits. These background-induced changes in receptive field size are absent when the depolarizing responses in bi- and triphasic horizontal cells are blocked by lowering the [Ca 2+] in the Ringer's solution from 1.0 to 0.1 mM. These results cannot be explained by the linear properties of the horizontal cell layers, nor by slow adaptational processes, but are consistent with the concept that feedback from horizontal cells to cones modifies the horizontal cell receptive field properties.

Modulation of synaptic transfer between retinal cones and horizontal cells by spatial contrast.

We studied the influence of steady annular light on the kinetics and sensitivity of horizontal cell (HC) responses to modulation of the intensity of small concentric spots in the turtle retina. As shown by previous investigators, when the intensity of the annulus was equal to the mean spot intensity, spot response kinetics were the same as those for the modulation of spatially uniform light. Turning off the annulus attenuated dramatically high-frequency flicker sensitivity and enhanced somewhat low-frequency sensitivity. This phenomenon reflects a modulation of synaptic transfer between cones and second-order neurons that is mediated by cones, and it will be referred to as cone-mediated surround enhancement (CMSE). Our main results are as follows: (a) The change in test-spot response sensitivity and kinetics upon dimming a steady surrounding annulus is a consequence of the change in spatial contrast rather than change in overall light level. (b) Introduction of moderate contrast between the mean spot intensity and steady surrounding light intensity causes a marked change in spot response kinetics. (c) The dependence of spot response kinetics on surrounding light can be described by a phenomenological model in which the steady state gain and the time constant of one or two single-stage, low-pass filters increase with decreasing annular light intensity (d) The effect of surrounding light on spot responses of a given HC is not determined by change in the steady component of the membrane potential of that cell. (e) Light outside the receptive field of an HC can affect that cell's spot response kinetics. (f) In an expanding annulus experiment, the distance over which steady annular light affects spot response kinetics varies among HCs and can be quite different even between two cells with closely matched receptive field sizes. (g) The degree of CMSE is correlated with HC receptive field size. This correlation suggests that part of the enhancement mechanism is located in the HC. Taken together, our results suggest the involvement of the inner retina in CMSE.

Effects of norepinephrine on [ 3H]dopamine release and horizontal cell receptive-field size in the goldfish retina

Norepinephrine increased the release of pre-loaded [ 3H]dopamine from goldfish retinas. Pharmacological studies suggested that the n norepinephrine-induced [ 3H]dopamine release was due to an exchange mechanism between norepinephrine and pre-loaded [ 3H]dopamine. Norepinephrine also depolarized and reduced the receptive-field size of horizontal cells in goldfish retinas. The action of norepinephrine on horizontal cells was probably not due to the release of endogeneous dopamine because the effect of norepinephrine was not abolished in retinas in which all dopaminergic neurons had been destroyed by prior treatment with 6-hydroxydopamine. The pharmacology of the effect of norepinephrine on horizontal cells suggested that it was due to an agonist action of norepinephrine acting at horizontal cell dopamine receptors. It is still unclear whether endogenous norepinephrine is a regulator of dopamine release in the fish retina. Consequently, the function of the putative norepinephrine-containing amacrine cells of the fish retina remains to be elucidated.

Background illumination reduces horizontal cell receptive-field size in both normal and 6-hydroxydopamine-lesioned goldfish retinas

The effect of background illumination on horizontal cell receptive-field size and dye coupling was investigated in isolated superfused goldfish retinas. Background illumination reduced both horizontal cell receptive-field size and dye coupling. The effect of light on horizontal cell receptive-field size was mimicked by treating the retina with 20 microM dopamine. To test the hypothesis that the effects of light were due to endogenous dopamine release, the effect of light was studied in goldfish retinas in which dopaminergic interplexiform cells were lesioned using 6-hydroxydopamine treatment. In lesioned retinas, background illumination reduced both horizontal cell receptive-field size and dye coupling. Furthermore, the effect of background illumination on unlesioned animals could not be blocked by prior treatment with the D1 dopamine receptor antagonist SCH-23390. These results suggest that, in goldfish retina, dopamine release is not the only mechanism by which horizontal cell receptive-field size could be reduced by light.

Effects of light stimuli on the release of dopamine from interplexiform cells in the white perch retina

Interplexiform cells are centrifugal neurons in the retina carrying information from the inner to the outer plexiform layers. In teleost fish, interplexiform cells appear to release dopamine in the outer plexiform layer after prolonged darkness that modulates the receptive-field size and light responsiveness of horizontal cells (Mangel & Dowling, 1985; Yang et al., 1988a, b). It has been proposed that interplexiform cells may also release dopamine upon steady illumination because horizontal cells' receptive fields shrink in the light (Shigematsu & Yamada, 1988). Here, we report the shrinkage of the receptive fields of horizontal cells seen in the presence of background illumination is not blocked by dopamine antagonists, indicating that dopamine does not underlie the receptive-field size changes observed during steady illumination. Flickering light, however, does appear to stimulate the release of dopamine from the interplexiform cells, resulting in a marked reduction of horizontal cell receptive-field size. Taken together, experiments on horizontal cells indicate that dopamine is released from interplexiform cells in the teleost retina after prolonged darkness and during flickering light, but that dopamine release from interplexiform cells during steady retinal illumination is minimal.