PubMed Health⌕ Search

Biomedical subjects

P Sterling

Publications and source records attributed to P Sterling.

At least 19 recordsLinked to original sources
PubMed1996

Absence of spectrally specific lateral inputs to midget ganglion cells in primate retina.

Visual information is conveyed to the brain by the retinal ganglion cells. Midget ganglion cells serve fine spatial vision by summing excitation from a receptive field 'centre', receiving input from a single cone in the central retina, with lateral inhibition from a receptive field 'surround', receiving input from many surrounding cones. Midget ganglion cells are also thought to serve colour opponent vision because the centre excitation is from a cone of one spectral type, while the surround inhibition is from cones of the other type. The two major cone types, middle(M)- and long-(L)wavelength sensitive, are equally numerous and randomly distributed in the primate central retina, so a spectrally homogeneous surround requires that the cells mediating lateral interactions (horizontal or amacrine cells) receive selective input from only one cone type. Horizontal cells cannot do this because they receive input indiscriminately from M and L cones. Here we report that the amacrine cells connected to midget ganglion cells are similarly indiscriminate. The absence of spectral specificity in the inhibitory wiring raises doubt about the involvement of midget ganglion cells in colour vision and suggest that colour opponency may instead be conveyed by a different type of ganglion cell.

Animals↗
PubMed1996

Foveal cones form basal as well as invaginating junctions with diffuse ON bipolar cells.

The response of a mammalian bipolar cell is generally thought to be determined by the location and morphology of synapses from the cone terminal: ON bipolar cells are believed to be depolarized strictly at invaginating contacts and OFF bipolar cells hyperpolarized at basal contacts. This hypothesis was re-investigated in the macaque fovea (1 deg nasal) using electron micrographs of serial sections. We determined the number of invaginating sites available and then identified the contacts to bipolar cells with axons in the ON level of the inner plexiform layer. A cone terminal forms about 20 active zones marked by ribbons. A few active zones house two invaginating dendrites, so there are 22 invaginating sites per cone. A midget ON bipolar cell collects 18 invaginating contacts from one cone, thus only about four invaginating sites remain for diffuse ON bipolar cells. Two diffuse ON cells were reconstructed; each collects about 25 contacts from an estimated 10 cones. Only three or four of these contacts are invaginating; the rest are basal, adjacent to the triad. This suggests that basal contacts can be depolarizing. The distance from the vesicle release site at active zones to an invaginating contact is 140 +/- 40 nm; to a basal contact adjacent to the triad is 500 +/- 160 nm, and to the next nearest basal contact is 950 +/- 370 nm.

Animals↗
PubMed1996

Evidence that vesicles on the synaptic ribbon of retinal bipolar neurons can be rapidly released.

We relate the ultrastructure of the giant bipolar synapse in goldfish retina to the jump in capacitance that accompanies depolarization-evoked exocytosis. Mean vesicle diameter is 29 +/- 4 nm, giving 26.4 aF/vesicle, so the maximum evoked capacitance (150 fF within 200 ms) represents fusion of about 5700 vesicles. Two terminals contained, respectively, 45 and 65 ribbon-type synaptic outputs, and a fully loaded ribbon tethers about 110 vesicles. Thus, the tethered pool, about 6000 vesicles, corresponds to the rapidly released pool. Further, the difference between small and large terminals in number of tethered vesicles matches their difference in capacitance jump. This suggests, within a "fire and reload" model of exocytosis, that the ribbon translocates synaptic vesicles very rapidly to membrane docking sites, supporting a maximum release rate of 500 vesicles/active zone/s, until the population of tethered vesicles is exhausted.

Animals↗
PubMed1995

Retinal neurons and vessels are not fractal but space-filling.

Many branched patterns in nature are hypothesized to be fractal, i.e., statistically self-similar across a range of scales. We tested this hypothesis on the two-dimensional arbors of retinal neurons and blood vessels. First, we measured fractalness on synthetic fractal and nonfractal patterns. The synthetic fractal patterns exhibited self-similarity over a decade of scale, but the nonfractal "controls" showed hardly any self-similarity. Neuronal and vascular patterns showed no greater self-similarity than the controls. Second, we manipulated a synthetic fractal pattern to remove its self-similarity and found this to be reflected in a loss of measured fractalness. The same manipulation of the nonfractal control and also of the neural and vascular patterns did not alter their measured fractalness. Third, we "grew" patterns of branched line segments according to a variety of nonfractal algorithms. These patterns were, if anything slightly more fractal than the neural and vascular patterns. We conclude that the biological patterns studied here are not fractal. Finally, we measured extended versions of these patterns: a contiguous array of homotypic neuron arbors and a vascular pattern with a high degree of total detail. These patterns showed a "fractal dimension" of 2, which implies that down to some cut-off scale they fill space completely. Thus, neural and vascular patterns might best be described as quasi-regular lattices.

Animals↗
PubMed1995

Mammalian rod terminal: architecture of a binary synapse.

The mammalian rod synapse transmits a binary signal (one photon or none) using tonic, rapid exocytosis. We constructed a quantitative, physical model of the synapse. Presynaptically, a single, linear active zone provides docking sites for approximately 130 vesicles, and a "ribbon" anchored to the active zone provides a depot for approximately 640 vesicles. Postsynaptically, 4 processes invaginate the terminal: 2 (known to have low affinity glutamate receptors) lie near the active zone (16 nm), and 2 (known to have high affinity glutamate receptors) lie at a distance (130-640 nm). The presynaptic structure seems designed to minimize fluctuations in tonic rate owing to empty docking sites, whereas the postsynaptic geometry may permit 1 vesicle to evoke an all-or-none response at all 4 postsynaptic processes.

Animals↗
PubMed1995

How retinal microcircuits scale for ganglion cells of different size.

Ganglion cell receptive field centers are small in central retina and larger toward periphery. Accompanying this expansion, the distribution of sensitivity across the centers remain Gaussian, but peak sensitivities decline. To identify circuitry that might explain this physiology, we measured the density of bipolar cell synapses on the dendritic membrane of beta (X) and alpha (Y) ganglion cells and the distribution of dendritic membrane across their dendritic fields. Both central and peripheral beta cells receive bipolar cell synapses at a density of approximately 28/100 microns2 of dendritic membrane; central and peripheral alpha cells receive approximately 13/100 microns2. The distribution of dendritic membrane across the dendritic field is dome-like; therefore, the distribution of bipolar cell synapses is also dome-like. As the dendritic field enlarges, total postsynaptic membrane increases with field radius, but only linearly. Consequently, density of postsynaptic membrane in the dendritic field declines, and so does density of synapses within the field. The results suggest a simple model in which the receptive field center's Gaussian profile and peak sensitivity are both set by the density of bipolar cell synapses across the dendritic field.

Animals↗
PubMed1994

M and L cones in macaque fovea connect to midget ganglion cells by different numbers of excitatory synapses.

Visual acuity depends on the fine-grained neural image set by the foveal cone mosaic. To preserve this spatial detail, cones transmit through non-divergent pathways: cone-->midget bipolar cell-->midget ganglion cell. Adequate gain must be established along each pathway; crosstalk and sources of variation between pathways must be minimized. These requirements raise fundamental questions regarding the synaptic connections: (1) how many synapses from bipolar to ganglion cell transmit a cone signal and with what degree of crosstalk between adjacent pathways; (2) how accurately these connections are reproduced across the mosaic; and (3) whether the midget circuits for middle (M) and long (L) wavelength sensitive cones are the same. We report here that the midget ganglion cell collects without crosstalk either 28 +/- 4 or 47 +/- 3 midget bipolar synapses. Two cone types are defined by this difference; being about equal in number and distributing randomly in small clusters of like type, they are probably M and L.

Animals↗
PubMed1994

Subcellular localization of GABAA receptor on bipolar cells in macaque and human retina.

The subcellular distribution of GABAA receptor in the macaque and human retina was studied by immunocytochemistry with monoclonal antibodies for the alpha and beta subunits with a particular focus on bipolar cells. Immunoreactivity to GABAA receptor was present on dendritic tips of all bipolar cells. The stain was strongest on bipolar membranes in apposition to horizontal cell processes. Stain was concentrated on the tips of flat and invaginating cone bipolar cells at the base of the cone pedicle and on the invaginating tips of rod bipolar cells. Stain on the cone pedicle membrane was restricted to sites of apposition to stained bipolar dendrites; pedicle membrane in apposition to horizontal cell processes was unstained. Stain was also present on bipolar axon terminals in both on and off strata of the inner plexiform layer. All bipolar cell somas stained faintly; horizontal and Müller cell somas were unstained. The alpha and beta subunits distributed similarly in monkey and human retina. Presence of GABAA receptor on the bipolar dendritic tips suggests that horizontal cells directly affect bipolar cells. Thus, GABAA receptor might mediate the receptive field surround of both off and on bipolar cells. Presence of GABAA receptor on bipolar axon terminals suggests that much of the inhibition feeding back from GABAergic amacrine to bipolar cells is GABAA-mediated.

Animals↗
PubMed1994

Rate of quantal transmitter release at the mammalian rod synapse.

Under scotopic conditions, the mammalian rod encodes either one photon or none within its integration time. Consequently the signal presented to its synaptic terminal is binary. The synapse has a single active zone that releases neurotransmitter quanta tonically in darkness and pauses briefly in response to a rhodopsin isomerization by a photon. We asked: what minimum tonic rate would allow the postsynaptic bipolar cell to distinguish this pause from an extra-long interval between quanta due to the stochastic timing of release? The answer required a model of the circuit that included the rod convergence onto the bipolar cell and the bipolar cell's signal-to-noise ratio. Calculations from the model suggest that tonic release must be at least 40 quanta/s. This tonic rate is much higher than at conventional synapses where reliability is achieved by employing multiple active zones. The rod's synaptic mechanism makes efficient use of space, which in the retina is at a premium.

Animals↗
PubMed1992

Immunoreactivity to GABAA receptor in the outer plexiform layer of the cat retina.

The distribution of GABAA receptor in the outer plexiform layer of cat retina was studied by immunocytochemistry with monoclonal antibodies. Staining was observed at the base of the cone pedicle, extracellularly, in association with the "triad" synaptic complex. Some bipolar dendrites and the basal processes that interconnect the cone pedicles were also stained. Rod spherules and horizontal cells were negative. The findings support the idea that the cone horizontal cells are GABAergic.

Animals↗
PubMed1992

Computational model of the on-alpha ganglion cell receptive field based on bipolar cell circuitry.

The on-alpha ganglion cell in the area centralis of the cat retina receives approximately 450 synapses from type b1 cone bipolar cells. This bipolar type forms a closely spaced array (9 microns), which contributes from 1 to 7 synapses per b1 cell throughout the on-alpha dendritic field. Here we use a compartmental model of an on-alpha cell, based on a reconstruction from electron micrographs of serial sections, to compute the contribution of the b1 array to the on-alpha receptive field. The computation shows that, for a physiologic range of specific membrane resistance (9500-68,000 omega.cm2) and a linear synapse, inputs are equally effective at all points on the on-alpha dendritic tree. This implies that the electrotonic properties of the dendritic tree contribute very little to the domed shapes of the receptive field center and surround. Rather, these shapes arise from the domed distribution of synapses across the on-alpha dendritic field. Various sources of "jitter" in the anatomical circuit, such as variation in bipolar cell spacing and fluctuations in the number of synapses per bipolar cell, are smoothed by the overall circuit design. However, the computed center retains some minor asymmetries and lumps, due to anatomical jitter, as found in actual alpha-cell receptive fields.

Animals↗
PubMed1992

Gap junctions between the pedicles of macaque foveal cones.

Cone terminals ("pedicles") in the fovea of macaque retina were studied in electron micrographs of serial sections. Pedicles were sheathed in glia except for small (0.2 microns 2) fenestrations, 4.8 +/- 1.7 per pedicle. At each fenestration the membranes of adjacent pedicles were contiguous and marked by an adherent junction, which in turn was invariably associated with gap junctions. There were 3.2 +/- 1.4 gap junctions per adherent junction and thus, about fifteen gap junctions per pedicle. The gap junctions were small, 1.6 x 10(-3) +/- 1.8 x 10(-3) microns 2 (mean +/- SD) and were formed indiscriminately with all neighboring pedicles. An upper bound was estimated of 170 connexons per pedicle and thus a coupling conductance of 1.7 x 10(4) pS. Available psychophysical data suggest that the junctions are uncoupled at high luminance. They may couple at lower luminance where spatial averaging would improve contrast sensitivity without cost to spatial acuity.

Animals↗
PubMed1991

Microcircuitry related to the receptive field center of the on-beta ganglion cell.

1. We have investigated the anatomic basis for the Gaussian-like receptive field center of the on-beta ("X") ganglion cell in the area centralis of cat retina. Three adjacent on-beta cells were reconstructed from electron micrographs of 279 serial sections cut vertically through a patch of retina at approximately 1 degree eccentricity. 2. All the bipolar synapses on these cells were identified, and about one-half of these were traced to type b1 bipolar cells, which formed a regular array in the plane of the retina. 3. On average, seven b1 cells contributed to a beta cell: bipolar axons near the middle of the beta dendritic field tended to give many contacts (12-33 contacts); axons near the edge of the field tended to give few contacts (3-4 contacts). 4. Each b1 cell collected from four to seven cones, and the mean number of cones converging through the b1 array to a beta cell was 30. 5. Assuming equal effectiveness for all b1----beta cell synapses, a spatial weighting function was derived from these results. The mean radius of this function at 1/e amplitude for three beta cells was 18.0 +/- 1.1 (SD) microns. This is considerably narrower than the corresponding measurements of the beta cell receptive field center (28 +/- 3 microns) at this eccentricity. 6. It is concluded, in agreement with previous work, that all cones encompassed by the beta cell's dendritic field and those slightly beyond it connect directly to the beta cell via the b1 bipolar cell array. However, the center of the beta cell receptive field is still broader by approximately 60%. This suggests that pooling of cone signals may occur at the level of the outer plexiform layer.

Animals↗
PubMed1990

Demonstration of cell types among cone bipolar neurons of cat retina.

We identified all the cone bipolar cells (80) in a small patch of one retina and then studied in detail the complete subset (42) that sends axons to sublamina b of the inner plexiform layer. The point was to learn whether the 'types' suggested previously, based on a few examples from a large population, could be substantiated or whether there would be intermediate forms. Tissue from the area centralis (1 degree eccentricity), was prepared as a series of 279 ultrathin sections and photographed in the electron microscope. Thirteen cells were reconstructed completely and parcelled into five categories (b1-b5) based on external morphology. For nine of these cells (two from categories b1-b4 and one from b5) most of the synaptic inputs and outputs were identified. When these nine cells were parcelled according to their synaptic patterns, they sorted into the same five categories. The remaining 29 cells in the population, though not reconstructed, were studied in detail by tracing their processes through the series. Ten of these cells, those near the margin of the series, were incomplete. The other 19 cells had essentially the same distribution of morphologies and synaptic patterns as the subset studied by total reconstruction: when plotted in multiparametric space, they formed distinct clusters corresponding to the five morphological categories. There was no hint of intermediate forms. That all the neurons in the population sort into some cluster (no intermediate forms), and that each neuron sorts into the same cluster by different criteria, argues that the clusters represent natural types. Each type forms a regular array in the region studied with an axonal 'coverage factor' that is close to one.

Animals↗
PubMed1990

Convergence and divergence of cones onto bipolar cells in the central area of cat retina.

In the central area of cat retina the cone bipolar cells that innervate sublamina b of the inner plexiform layer comprise five types, four with narrow dendritic fields and one with a wide dendritic field. This was shown in the preceding paper (Cohen & Sterling 1990 a) by reconstruction from electron micrographs of serial sections. Here we show by further analysis of the same material that the coverage factor (dendritic spread x cell density) is about one for each of the narrow-field types (b1, b2, and b4). The same is probably true for the other narrow-field type (b3), but this could not be proved because its dendrites were too fine to trace. The dendrites of types b1, b2, and b4 collect from all the cone pedicles within their reach and do not bypass local pedicles in favour of more distant ones. The dendrites of type b5, the wide-field cell, bypass many pedicles. On average 5.1 +/- 1.0 pedicles coverage on a b1 bipolar cell; 6.0 +/- 1.2 converge on a b2 cell and 5.7 +/- 1.5 converge on a b4 cell. Divergence within a type is minimal: one pedicle contacts only 1.2 b1 cells, 1.0 b2 cells, and 1.0 b4 cells. Divergence across types is broad: each pedicle apparently contacts all four types of the narrow-field bipolar cells that innervate sublamina b. Each pedicle probably also contacts an additional 4-5 types of narrow-field bipolar cell that innervate sublamina a. There are several possible advantages to encoding the cone signal into multiple, parallel, narrow-field pathways.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed1990

Cone receptive field in cat retina computed from microcircuitry.

The receptive-field profile of the cone in cat-retina was computed. The computation was based on (1) the known anatomical circuit connecting cones via narrow-field bipolar cells to the on-beta ganglion cell; (2) the known physiological receptive-field profile of the on-beta (X) cell at the corresponding eccentricity; and (3) a model in which the beta receptive field arises by linear superposition of cone receptive fields. The computed cone receptive field has a center/surround organization with a center almost as broad as that of the beta cell center. The cone surround is comparably broad to that of the beta cell but somewhat lower in peak amplitude. The problems to which the center/surround receptive field are the solution, namely, signal compression and noise reduction, apparently must be solved before the first synapse of the visual pathway.

Animals↗