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.     

The connectivity of cones and cone horizontal cells in a mosaic-type teleost retina

Summary A total of 20 Golgi-impregnated cone horizontal cells of Nannacara anomala (Cichlidae) were studied in alternating semi- and ultrathin sections in order to examine their connections with the overlying square mosaic of equal double and central single cones. Cone horizontal cells exhibit three types of processes: (a) the long horizontal axon, (b) short horizontal dendrites with a terminal swelling, and (c) cone contacting processes ascending towards the outer plexiform layer. As seen in tangential sections, the latter processes are arranged in the form of two concentric circles including a central spot. The processes of the inner circle contact the eight double cone pedicles of one square unit: processes of the outer circle contact eight more double cone pedicles which are directly adjacent to the square unit. The central spot represents a process which contacts the central single cone. Processes of the inner circle most often terminate in a dichotomous branching which represents the lateral elements to one ribbon synapse, whereas in the outer circle only a single terminal swelling is observed. Because of the mosaic of the cones and the constancy of this pattern of connectivity a model can be constructed where the dendritic fields of the cone horizontal cells overlap to a considerable extent. From this model, it follows that each double cone pedicle is contacted by four different horizontal cells. The functional significance of these findings for color vision is discussed in the light of recent work with the microspectrophotometer characterizing the cone system of this species as bichromatic. The mosaiclike arrangement of the horizontal cell dendrites supports the conclusion that the parallels between the patterns of receptor and horizontal cells are no coincidence but play an important role in lateral inhibition and neural adaptation of the retina.A preliminary report of this study was given at the international symposium Neural principles in vision held at the University of Munich in September 1975Supported by grant Wa 348/1 of the Deutsche Forschungsgemeinschaft     

Electronic simulation of cones,horizontal cells and bipolar cells of generalized vertebrate cone retina

Electronic analogue of my theoretical model of generalized vertebrate cone retina [Siminoff: J. Theor. Biol. 86, 763 (1980)] is presented. Cone mosaic is simulated by 25x21 grid of phototransistors that have colored filters mounted in front of then to produce red-, green-, and blue-sensitive cones arranged in a trichromatic retina. Each retinal element is simulated by Summator-Integrator and unit gain voltage invertes are used to give correct polarities to output voltages. Dynamic properties of retinal elements are developed solely by temporal interplay of antagonistic input voltages with differing time courses, and spatial organization of receptive fields is developed by unit hexagons that precisely define cone input voltages to subsequent elements. Electronic model contains both color- and non-colorcoded channels. Negative feedback from L-horizontal cells to cones, electrical coupling of like-cones, and electrical coupling of like-horizontal cells are simulated by feedfoward circuits. Stray light is present due to light scattering properties of colored filters used to simulate color selectivety of cones. Stationary and moving spots of white and colored lights of varied sizes and intensities are used to study characteristics of electronic analogue. Results demonstrate practicality of electronic simulation to function analogous to real cone retinas to process visual stimuli and give information to higher centers as to size, shape, color and motion of objects in visual world.     

Some horizontal cells of the bovine retina receive input synapses in the inner plexiform layer

Bovine retinae were stained immunocytochemically with antibodies against the calcium-binding protein, calbindin. Horizontal cells in the outer plexiform layer were heavily labelled. The processes of most horizontal cells were confined to the level of the outer plexiform layer, and the tips of their dendrites were positioned as the lateral elements of the cone triads, viz. the usual mammalian arrangement. However, some of the horizontal cells had additional thick processes descending to branch within the inner plexiform layer, where they were postsynaptic at bipolar cell dyads and where they also received input from amacrine cells. No output synapses of horizontal cells were observed in the inner plexiform layer.     

Changes in direct current input resistance in horizontal cells of fish retina during excitation

The input resistance of pike retinal horizontal cells was measured by means of coaxial electrodes under various conditions of illumination. With moderate intensities of illumination, the resistance (determined from a potential drop caused by the current passed through the microelectrode) increases, whereas at high saturating intensities it decreases, as compared with its value in darkness. Such changes in resistance of the horizontal cells explain the effects of input signals interaction in these cells, such as enhancement and complete saturation, observed earlier. Some properties of the horizontal cell response permit us to assume that the "active" cell response to polarization makes a substantial contribution to the measured resistance of these cells. Possible mechanisms of such changes in input resistance of horizontal cells are discussed.Institute of Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 3, No. 2, pp. 210–216, March–April, 1971.     

Displaced horizontal cells in the chick retina

The retina of the chick contains retinal cells of a morphology very similar to that of the horizontal cells, but the perikarya, axons, and axon terminals lie in the inner plexiform layer. The discovery of this neuronal ectopia appears to support the idea that some horizontal and amacrine cells derive from a common, freely migrating cell.     

Effect of acetylcholine on membrane potential of the horizontal cells of the goldfish retina and the electroretinogram of the frog

The action of acetylcholine on the horizontal cells of the goldfish retina and the electro-retinogram of the frog was studied. Acetylcholine in concentrations of 1·10⁻⁹–1·10⁻³ M depolarized these cells. The maximal level of depolarization never reached zero level of the membrane potential and was about equal to the membrane potential in darkness. In a concentration of 1·10⁻²–5·10⁻² M acetylcholine suppressed the b- and d-waves of the frog electro-retinogram, and as a result the stable P_III component was isolated from the ERG. A mediator role is ascribed to acetylcholine in the synapses of the outer plexiform layer.     

The cone synapses of cone bipolar cells of primate retina

One each of bipolar cell types DB2 and DB4, together with a flat and an invaginating midget bipolar cell, were taken from a Golgi-stained rhesus macaque retina; then serially sectioned for EM examination of their synapses with cone pedicles. The cone input to the dendrites of the DB2 cell was exclusively at basal junctions; it had a characteristic distribution. Fifty per cent of the basal synapses were with cone pedicle membrane immediately adjacent to the dendrite of a bipolar cell invaginating to end opposite the ribbon of a cone triad (this, therefore, is called triad-associated). The remainder were one or more synapses distant from the triad-associated position (and, therefore, non-triad associated). The DB4 cell had both basal (predominantly in the triad-associated position) and ribbon-related synaptic input. But the basal to invaginating ratio differed from that of our previously published cell; 56% basal, 43% invaginating, as compared with 31% basal and 69% invaginating. Like foveal IMB cells the synapses of the mid-peripheral invaginating midget bipolar cell were exclusively invaginating; but were about 25% more numerous. The flat midget bipolar cell made exclusively basal synapses. These were 2.5 times more numerous than those of foveal flat midget bipolar cells, and 3.5 times the number of invaginating midget bipolar synapses at equivalent eccentricity. The synapses between cones and diffuse and midget bipolar cells are characteristic for each particular bipolar cell type, but the details depend on a cell’s distance from the fovea (eccentricity). A rather constant number of cone pedicle synaptic ribbons 38.6±2.5 (n±:60) was found across mid-peripheral macaque and vervet monkey retinae. The smaller mean number for vervet monkey, 27.4±3.5 (n ±:23), suggests there can also be generic differences in synaptic detail at cone bipolar cell synapses.     

Enkephalin in the goldfish retina

Enkephalin-like immunoreactive amacrine cells were visualized using the highly sensitive avidin-biotin method. The somas of these cells were situated in the inner nuclear and ganglion cell layers. Enkephalin-stained processes were observed in layers 1, 3, and 5 of the inner plexiform layer. The biosynthesis of sulfur-containing compounds in the goldfish retina was studied by means of a pulse-chase incubation with 35S-methionine. A 35S-labeled compound, which comigrated with authentic Met5-enkephalin on high-performance liquid chromatography (HPLC), was synthesized and was bound competitively by antibodies to enkephalin and by opiate receptors. This compound was tentatively identified as "Met5-enkephalin." The newly synthesized 35S-Met5-enkephalin was released upon depolarization of the retina with a high K+ concentration. This K+-stimulated release was greatly suppressed by 5 mM Co2+, suggesting that the release was Ca2+ dependent. Using a double-label technique, enkephalin immunoreactivity and gamma-aminobutyric acid (GABA) uptake were colocalized to some amacrine cells, whereas others labeled only for enkephalin or GABA. The possible significance of enkephalin-GABA interactions is also discussed.     

Ion exchange in the horizontal cells of the retina

Experimental data indicate that the membrane potential of L-type horizontal cells of the retina to bright light (i.e., when synaptic inputs are completely closed) is close to the potassium equilibrium potential. From this observation the intracellular concentration of K⁺ and Na⁺ was estimated. The latter was found to be relatively high (tens of millimoles/liter), i.e., comparable with the intracellular K⁺ concentration. This result, coupled with data on closeness of the equilibrium potential of the photic response to zero, is evidence that besides sodium conductance, the potassium conductance of the subsynaptic membrane also participates in generation of the photic response by these cells. The steady-state sodium and potassium synaptic currents was shown to be relatively small and to vary only a little over the whole working range of potentials (from –72 to –16 mV), due to the nonlinear properties of the nonsynaptic cell membrane.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 14, No. 1, pp. 3–10, January–February, 1982.     

Germinal cells in the goldfish retina that produce rod photoreceptors

Dividing cells and their progeny in retinae of young goldfish were labeled with [3H]thymidine, and selected cells were reconstructed from serial sections processed for electron microscopic autoradiography. Our goals were to characterize the cells that were identified as rod precursors in previous light microscopic autoradiographical studies and to determine their origin and fate. (In fish the population of rods increases several-fold postembryonically by proliferation of rod precursor cells scattered across the retina). Over 200 labeled cells taken from 11 retinas were examined, and 20 of these were reconstructed in their entirety. Some retinas were examined at short intervals (1 to 48 hr) after [3H]thymidine injection in order to study mitotically active cells, and others were examined after longer intervals (9 or 14 days) to discover the nature of the progeny of labeled dividing cells. Previous evidence from thymidine studies in larval goldfish suggested that proliferating cells destined to produce rods appear first in the inner nuclear layer and later in the outer nuclear layer, where they continue to divide and generate new rods (P.R. Johns, (1982) J. Neurosci. 2, 179). The present results provide morphological evidence in support of the suggestion that rod precursors migrate from inner to outer nuclear layer and, furthermore, show that the precursors are closely associated with, and perhaps guided by, the radial processes of Müller glial cells. Examination of EM autoradiographs of labeled cells at 9 and 14 days after a pulse label with thymidine confirms that the differentiated progeny of dividing precursor cells are exclusively rods. To our knowledge, rod precursors are the first example of a neuronal germinal cell in the vertebrate central nervous system that under normal conditions produces only one type of neuron.     

Interaction between horizontal cells of the turtle retina

Interaction between horizontal cells of the turtle retina was studied by two microelectrodes (polarizing and recording), inserted into different cells at different distances apart. The presence of a direct electrical connection was demonstrated between the L cells of the same type (I, with large, and II, with small receptive fields). Its magnitude depends on the conditions of illumination and the level of the membrane potential, possibly because of the properties of the subsynaptic and nonsynaptic membranes of the horizontal cells. No direct electrical connection exists between L cells of different types. However, hyperpolarization of the type I cells through the microelectrode or by stimulation with a circle of light evoked depolarization in the type II cells. This indirect connection between the horizontal cells, also dependent on the conditions of illumination, can probably be explained by feedback to these cells from the photoreceptors. Polarization of L cells of both types had no effect on horizontal cells of color type.     

鲫鱼视网膜亮度型水平细胞上不同视锥信号的相互作用:实验及模型

本文应用胞内记录和动态模型分析方法,研究了离体鲫鱼视网膜视锥驱动的高度型水平细胞（ＬＨＣ）上不同视锥信号的相互作用。实验表明,绿背景光的作用可以提高ＬＨＣ的红光反应,这种增强作用与绿敏锥的活动程度密切相关,模型分析表明,背景光 的作用谷氨酸介导的前馈性通路和ＧＡＢＡ介导的反馈性通路活动同时得以增强,水平细胞对光反应的增强效应不能为外泊性ＧＡＢＡ所消除。则其程度为前馈性通路和反馈性通路活动增加的相对     

Dynamics of chromaticity horizontal cells in the freshwater turtle retina

A model of the cone-horizontal cell circuit is presented based on morphological evidence recently found in the Reeves' turtle: a luminosity horizontal cell (LHC) that receives inputs from red-, green-, and blue-sensitive cones in the ratio of 15:3:1, a triphasic horizontal cell (THC) that receives inputs from one class of red-sensitive and from blue-sensitive cones in the ratio of 2:1; and a biphasic chromaticity horizontal cell (BHC) that receives inputs from green-sensitive cones as well as from a special class of red-sensitive (i.e. the broad spectrum) and from blue-sensitive cones in the ratio of 3:2:1. A study of the simulated impulse responses strongly suggests that the basic response patterns of the BHC and THC can be readily explained by a simple wiring diagram consisting of direct hyper-polarizing inputs from the appropriate cones and a depolarizing input from the LHC which acts as a voltage inverter. A negative feedback circuit from the LHC to the cone pedicles is included and its negative feedback gain increases as the mean illuminance level (Io) increases. The negative feedback circuit, which promotes adaptation in the cones to changing Io's, is not necessary for opponent polarization in the BHC or THC, but does explain variabilities of impulse responses.     

Membrane potential clamping in horizontal cells of the fish retina

The membrane potential of horizontal cells of the retina was clamped by uniform polarization of the layer of these cells by a current passed through extracellular electrodes. The volt-ampere characteristic curve of the synaptic membrane of the horizontal cells in some cases had segments with negative slope. With a sharp change in the level of voltage clamping the time taken for the resistance of the membrane to change was under 20 msec. Comparison of responses to photic stimulation recorded with and without voltage clamping showed that participation of the nonsynaptic membrane in the generation of responses to photic stimulation can affect their shape substantially.Institute of Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 9, No. 4, pp. 402–407, July–August, 1977.