Theta ganglion cell type of cat retina

We define a new bistratified ganglion cell type of cat retina using intracellular staining in vitro. The theta cell has a small soma, slender axon, and delicate, highly branched dendritic arbor. Dendritic fields are intermediate in size among cat ganglion cells, with diameters typically two to three times those of beta cells. Fields increase in size with distance from the area centralis, ranging in diameter from 70 to 150 microns centrally to a maximum of 700 microns in the periphery. Theta cells have markedly smaller dendritic fields within the nasal visual streak than above or below it and smaller fields nasally than temporally. Dendritic arbors are narrowly bistratified. The outer arbor lies in the lower part of sublamina a (OFF sublayer) of the inner plexiform layer where it costratifies with the dendrites of OFF alpha cells. The inner arbor occupies the upper part of sublamina b (ON sublayer), where it costratifies with ON alpha dendrites. The outer and inner arbors are composed of many relatively short segments and are densely interconnected by branches that traverse the a/b sublaminar border. Experiments combining retrograde labeling with intracellular staining indicate that theta cells project to the superior colliculus and to two components of the dorsal lateral geniculate nucleus (the C laminae and medial interlaminar nucleus). Theta cells project contralaterally from the nasal retina and ipsilaterally from the temporal retina. They apparently correspond to a sluggish transient or phasic W-cell with an ON-OFF receptive field center.     

The Eta ganglion cell type of cat retina.

We define a morphologic type of ganglion cell in cat retina by using intracellular staining in vitro. The eta cell has a small soma, slender axon, and delicate, highly branched dendritic arbor. Dendritic fields are intermediate in size among cat ganglion cells, with diameters typically two to three times those of beta cells. Fields increase in size as a function of distance from the area centralis, ranging in diameter from 90 microm to 200 microm centrally to a maximum of 600 microm in the periphery. This increase is unusually radially symmetric. By contrast with other cat ganglion cell types, eta cells do not have markedly smaller dendritic fields within the visual streak than above or below it nor much smaller fields nasally than temporally. Dendrites ramify broadly throughout sublamina a (OFF sublayer) of the inner plexiform layer. They arborize most densely in S2, where they costratify with dendrites of OFF alpha cells. There is apparently no matching ON variety of eta cell. Experiments combining retrograde labeling with intracellular staining indicate that eta cells project to the superior colliculus and to two components of the dorsal lateral geniculate nucleus (the C laminae and medial interlaminar nucleus). Eta cells apparently project contralaterally from the nasal retina and ipsilaterally from the temporal retina. The morphology and projection patterns of the eta cell suggest that its physiologic counterpart is a type of sluggish or W-cell with an OFF center, an ON surround, and possibly a transient light response.     

Extrinsic determinants of retinal ganglion cell structure in the cat

The degree to which a retinal ganglion cell's environment can affect its morphological development was studied by manipulating the distribution of ganglion cells in the developing cat retina. In the newborn kitten there is an exuberant ganglion cell projection from temporal retina to the contralateral lateral geniculate nucleus (LGNd) (Leventhal et al., 1988) and from nasal retina to the ipsilateral LGNd. Neonatal, unilateral optic tract section results in the survival of many of these ganglion cells (Leventhal et al., 1988). The morphology of ganglion cells which survive in regions of massively reduced ganglion cell density was studied. As reported previously (Linden and Perry, 1982; Perry and Linden, 1982; Ault et al., 1985; Eysel et al., 1985), we found that the dendritic fields of all types of ganglion cells on the border of an area depleted of ganglion cells extended into the depleted area. The cell bodies and dendritic fields of alpha and beta cells within depopulated areas, as well as on the borders of the depopulated areas, were larger than normal. The dendritic fields of these cells also exhibited abnormal branching patterns. For alpha and beta cell types the relative increase in size tended to be greatest where the relative change in density was the greatest. In fact, isolated beta cells within the cell-poor area centralis region resembled normal central alpha cells in the cell-rich region of the area centralis in the same retina. Interestingly, in the same regions of reduced density where alpha and beta cells were dramatically larger than normal, the cell body and dendritic field sizes of other cell types (epsilon, g1 and g2 were unchanged. These results indicate that neuronal interactions during development contribute to the morphological differentiation of retinal ganglion cells and that different mechanisms mediate the morphological development of different classes of cells in cat retina.     

Morphology of ganglion cell dendrites in the albino rat retina: an analysis with fluorescent carbocyanine dyes

The ganglion cell dendrites of the rat retina were investigated by means of the strongly fluorescent, non-polar carbocyanine dye 1,1-dioctadecyl-3,3,3',3'-tetramethyl-indocarbodyanine perchlorate (diI or diI-C18-3 or D282) which was taken up by retinofugal axons and transported in the retrograde direction. The dye completely outlined the somata, the axons and the dendritic trees of several retinal ganglion cells and allowed qualitative and quantitative investigations. By means of this labeling technique, the diameters were determined in 272 dendrites and somata of various ganglion cell sizes. A comparison of the measurements with those reported in the literature revealed that the diI could be taken up by all classes of retinal ganglion cells. The most frequently labeled cells were those of class II, which have small to middle-sized perikarya (16.7 +/- 2.5 microns in diameter) and small to middle-sized dendritic trees (187 +/- 70 microns in diameter) with a high branching frequency (88 +/- 19 branching points). Retinal ganglion cells of class I were less frequent and have large perikarya (21.9 +/- 3.4 microns in diameter) with large dendritic trees (318 +/- 55 microns in diameter) and medium branching frequency (60 +/- 19 branching points). Class III cells which were described incompletely in the literature, appeared to be small to middle-sized in their perikaryal diameter (15.9 +/- 2.5 microns) but have large dendritic trees (299 +/- 63 microns in diameter) and a low branching frequency (40 +/- 10 branching points). In about 10% of the retinal ganglion cells with completely filled dendritic fields, the somata were situated outside the dendritic extensions, as viewed on the whole mounted retina. These "asymmetric" retinal ganglion cells appeared to belong to class II cells and were evenly distributed throughout the entire retina and were not related to neighboring blood vessels. The orientation of the asymmetric dendrites was random in relation to the optic disc. The axons of asymmetric retinal ganglion cells were almost always oriented opposite to the direction of the dendritic trees. If the dendrites extended towards the optic disc, the proximal parts of the corresponding axons were oriented towards the periphery of the retina, turning then at 180 degrees to the optic disc. Less than 1.5% of the retrogradely filled cells were displaced ganglion cells and extended dendritic trees within the deep inner plexiform larger.(ABSTRACT TRUNCATED AT 400 WORDS)     

Alpha ganglion cells in the rabbit retina

In the rabbit retina a distinctive morphological class of large ganglion cells was demonstrated by a combination of intracellular staining with Lucifer Yellow and the quantification of reduced silver-stained preparations. The class is called alpha because of the qualitative and quantitative resemblance to the alpha cells of the cat's retina. Rabbit alpha cells change their size with location on the retina. In the high ganglion cell density region of the visual streak, their somata are about 15 micron in diameter, and their dendritic fields have diameters as small as 180-220 micron. The largest alpha cells in the inferior periphery have soma diameters of 30 micron and dendritic field diameters of 960 micron. There is a considerable scatter of sizes at any retinal location. Alpha cell density changes from about 55/mm2 in the streak to about 3/mm2 in far periphery, and the cells make up 1-1.4% of the ganglion cell population. Dendritic trees stratify in either an inner or an outer sublamina of the inner plexiform layer, suggesting an on/off dichotomy in the response to light. Each of the inner and outer branching subtypes is distributed in a regular mosaic, and the dendritic trees cover the retina completely and economically. The possibility is discussed that the alpha cells are the brisk transient/Y cells of physiology.     

Alpha and delta ganglion cells in the rat retina

In the rat retina a distinctive class of large ganglion cell was demonstrated by intracellular staining with Lucifer Yellow and with reduced silver staining. They are referred to as alpha cells because they resemble the alpha cells of other mammalian retinae. A second class, called delta cells, is also described. Both classes belong to the type I group defined by Perry (Proc. R. Soc. Lond. [Biol.] 204:363-375, '79). The dendritic trees of both classes stratify in either an inner or outer lamina of the inner plexiform layer which presumably corresponds to an on/off dichotomy in the response to light. Rat alpha cells constitute 2-4% of all ganglion cells, and their density, size, and detailed morphological appearance change with retinal location. Inner and outer stratifying alpha cells of the rat show significant differences compared to those of other mammals. In central retina (at the large cell density maximum) the densities and dendritic field sizes of inner and outer alpha cells are approximately equal. However, in peripheral retina outer alpha cells are up to three times more numerous and have dendritic field areas only one-third the size of those of the inner alpha cells. The maximal density is about 110 alpha cells/mm2; peripheral densities are about 30/mm2. The smallest central dendritic field diameters are 220 microns. Peripheral dendritic field diameters are 350-550 microns for outer and 570-790 microns for inner alpha cells. Each subpopulation is distributed in a regular mosaic, and the territorial arrangement of the dendritic fields provides a homogeneous coverage of the retina. The dendritic coverage is three- to 3.6-fold for each subpopulation, irrespective of their other quantitative differences. Eccentricity-dependent receptive field sizes of the alpha cells are predicted from the morphological data.     

Postnatal dendritic maturation of alpha and beta ganglion cells in cat retina

Of the 3 anatomically defined classes of ganglion cell in adult cat retina, the alpha and beta cells are the most well documented, thus providing a basis of comparison for developing ganglion cells. Alpha and beta ganglion cells in cat retinae at various ages from birth (P0) to adult were intracellularly injected with Lucifer yellow. At all ages, both cell types strongly resembled their adult counterparts. However, transient developmental characteristics established their immaturity. These features included spiny protuberances and "rings" along the dendritic surface that were no longer detectable after 3 weeks of age. In a small proportion of both inner and outer stratifying alpha ganglion cells, there was aberrant dendritic arborization. However, by P5 there was no remaining evidence of this deviant stratification pattern and all alpha and beta cells displayed the adult pattern of unistratification (present among the majority of these cells from birth). For both alpha and beta cells, the area of greatest development was the retinal periphery. In this region alpha cell dendritic trees continued to grow until 3 weeks postnatally, when they approached the adult dendritic field size; around this time, the major period of beta cell dendritic expansion began. From birth to adulthood, the distance between alpha cell dendritic branching points increased, while the number of nodes and tips decreased with age. The temporal disparity between alpha and beta cell dendritic expansion suggests that postnatal dendritic development involves an active process of growth, rather than merely passive stretching.     

Dendritic field development of retinal ganglion cells in the cat following neonatal damage to visual cortex: Evidence for cell class specific interactions

A well-known feature of the mammalian retina is the inverse relation that exists in central and peripheral retina between the density of retinal ganglion cells and their dendritic field sizes. Functionally, this inverse relation is thought to represent a means by which retinal coverage is maintained, despite significant changes in ganglion cell density. While it is generally agreed that the dendritic fields of mature retinal ganglion cells reflect, in part, competitive interactions that occur during development, the issue of whether these interactions are cell class specific remains unclear. In order to examine this question, we used intracellular staining techniques and an in vitro, living retina preparation to compare the soma and dendritic field sizes of alpha and beta ganglion cells from normal retinae with those of cells located in matched areas of retinae in which the density of beta ganglion cells had been reduced selectively by neonatal removal of visual cortex areas 17, 18, and 19. Our intracellular data show that while an early, selective, reduction in beta cell density has little or no effect on the cell body and dendritic field sizes of mature alpha cells, it results in a 13% increase in the mean soma area and an 83% increase in the mean dendritic field area of surviving beta cells. This differential effect suggests that the soma and dendritic field sizes of alpha and beta ganglion cells in the mature cat retina result primarily from competitive interactions during development that are cell class specific. J. Comp. Neurol. 390:470–480, 1998. © 1998 Wiley-Liss, Inc.     

Axon-bearing amacrine cells of the macaque monkey retina

A new and remarkable type of amacrine cell has been identified in the primate retina. Application of the vital dye acridine orange to macaque retinas maintained in vitro produced a stable fluorescence in the somata of apparently all retinal neurons in both the inner nuclear and ganglion cell layers. Large somata (approximately 15-20 microns diam) were also consistently observed in the approximate center of the inner plexiform layer (IPL). Intracellular injections of horseradish peroxidase (HRP) made under direct microscopic control showed that the cells in the middle of the IPL constitute a single, morphologically distinct amacrine cell subpopulation. An unusual and characteristic feature of this cell type is the presence of multiple axons that arise from the dendritic tree and project beyond it to form a second, morphologically distinct arborization within the IPL; these cells have thus been referred to as axon-bearing amacrine cells. The dendritic tree of the axon-bearing amacrine cell is highly branched (approximately 40-50 terminal dendrites) and broadly stratified, spanning the central 50% of the IPL so that the soma is situated between the outermost and innermost branches. Dendritic field size increases from approximately 200 microns in diameter within 2 mm of the fovea to approximately 500 microns in the retinal periphery. HRP injections of groups of neighboring cells revealed a regular intercell spacing (approximately 200-300 microns in the retinal periphery), suggesting that dendritic territories uniformly cover the retina. One to four axons originate from the proximal dendrites as thin (less than 0.5 microns), smooth processes. The axons increase in diameter (approximately 1-2 microns) as they course beyond the dendritic field and bifurcate once or twice into secondary branches. These branches give rise to a number of thin, bouton-bearing collaterals that extend radially from the dendritic tree for 1-3 mm without much further branching. The result is a sparsely branched and widely spreading axonal tree that concentrically surrounds the smaller, more highly branched dendritic tree. The axonal tree is narrowly stratified over the central 10-20% of the IPL; it is approximately ten times the diameter of the dendritic tree, resulting in a 100 times greater coverage factor. The clear division of an amacrine cell's processes into distinct dendritic and axonal components has recently been observed in other, morphologically distinct amacrine cell types of the cat and monkey retina and therefore represents a property common to a number of functionally distinct cell types. It is hypothesized that the axon-bearing amacrine cells, like classical neurons,     

Regular mosaics of large displaced and non-displaced ganglion cells in the retina of a cichlid fish

Large retinal ganglion cells in the tilapid cichlid fish Oreochromis spilurus (standard length 15-54 mm) were filled with horseradish peroxidase and studied in flatmounts. Three types, with distinct patterns of dendritic stratification, formed spatially independent, nonrandom mosaics. One type (about 0.3% of all ganglion cells) resembled the outer (off) alpha cells of mammals. They were very large, with thick primary dendrites and large, sparsely branched planar trees in the outer part of the inner plexiform layer (IPL). About 300 were arrayed regularly across each retina, their exact number and spacing depending on its size. Their somata were often displaced into the IPL, even where neighbours in the mosaic were orthotopic. Another type (0.8%) resembled the inner (on) alpha cells of mammals. These had slightly smaller somata that were never displaced and smaller trees in the middle layers of the IPL. About 800 were arrayed uniformly and regularly across each retina. A rarer type (0.06-0.08%) had two planar trees: one forming a coarse mosaic in the outer part of the inner plexiform layer (co-planar with the trees of outer alpha-like cells) and another in the outer plexiform layer. These "biplexiform" cells were smaller and rounder than alpha-like cells and always displaced. The dendrites were finer and less tapered. Cells in which we could identify an outer plexiform tree failed to cover the retina completely, but were nonrandomly distributed. We draw three main conclusions: (1) some nonmammalian vertebrates have separate inner and outer mosaics of large ganglion cells like those of mammals, (2) the vertical displacement of ganglion cell somata can vary widely within a single mosaic and may thus be functionally irrelevant, and (3) biplexiform ganglion cells exist in fish but differ in morphology from the biplexiform types described in some other vertebrates.     

Morphology of retinal ganglion cells in the ferret (Mustela putorius furo)

The ferret is the premiere mammalian model of retinal and visual system development, but the spectrum and properties of its retinal ganglion cells are less well understood than in another member of the Carnivora, the domestic cat. Here, we have extensively surveyed the dendritic architecture of ferret ganglion cells and report that the classification scheme previously developed for cat ganglion cells can be applied with few modifications to the ferret retina. We confirm the presence of alpha and beta cells in ferret retina, which are very similar to those in cat retina. Both cell types exhibited an increase in dendritic field size with distance from the area centralis (eccentricity) and with distance from the visual streak. Both alpha and beta cell populations existed as two subtypes whose dendrites stratified mainly in sublamina a or b of the inner plexiform layer. Six additional morphological types of ganglion cells were identified: four monostratified cell types (delta, epsilon, zeta, and eta) and two bistratified types (theta and iota). These types closely resembled their counterparts in the cat in terms of form, relative field size, and stratification. Our data indicate that, among carnivore species, the retinal ganglion cells resemble one another closely and that the ferret is a useful model for studies of the ontogenetic differentiation of ganglion cell types. J. Comp. Neurol. 517:459–480, 2009. © 2009 Wiley‐Liss, Inc.     

Independent mosaics of large inner- and outer-stratified ganglion cells in the goldfish retina.

Goldfish retinal ganglion cells were filled with horseradish peroxidase and studied in flatmounts. Two regular mosaics of large neurons with many of the properties of mammalian alpha ganglion cells were found, differing from each other in spacing, size, and dendritic stratification. The existence of biplexiform ganglion cells with additional dendrites in the outer plexiform layer was also confirmed. One of the two alpha-like mosaics consisted of giant ganglion cells with thick primary dendrites and large, sparsely branched dendritic trees in the outer sublamina of the inner plexiform layer (IPL). In fish 55-65 mm long, about 300 formed a tessellated array across each retina. Their somata (mean area 277 +/- 6 microns 2) were displaced to varying degrees into the IPL, neighbours in the mosaic often occupying different levels. Their dendrites ramified in one stratum near the inner nuclear layer, at a mean depth of 70.8 +/- 0.5% of the IPL. The other alpha-like mosaic comprised about 900 large ganglion cells, with slightly smaller somata (mean area 193 +/- 4 microns 2) in the ganglion cell layer. Most of their dendrites lay in a narrow stratum at 41.9 +/- 0.5% of the depth of the IPL. However, deviations (usually into more vitread strata) were common, which was not true for similar cells in the distantly related cichlid fish Oreochromis. Measurements of nearest neighbour distance (NND) for 4 outer and 4 inner mosaics showed that they were at least as regular as the alpha cell mosaics of mammals: the ratio of the mean NND to the standard deviation ranged from 4.03 for the least regular outer mosaic to 6.47 for the most regular inner mosaic. The wide phylogenetic distribution of these paired, regular mosaics points to a fundamental role in vision. The presence of some variability in dendritic stratification even within the exceptionally regular inner-stratified mosaic suggests that classifications based entirely on the detailed morphology of individual neurons may not always correlate well with their primary functional roles. Where possible, neuronal morphology and spatial distribution should be studied together.     

Morphology of retinal ganglion cells in the flying fox (Pteropus scapulatus): a lucifer yellow investigation

The morphology of retinal ganglion cells was determined in megachiroptera, commonly known as flying foxes. Retinal ganglion cells were intracellularly injected with the fluorescent dye Lucifer yellow in fixed retinae from adult little red flying foxes (Pteropus scapulatus) captured in their natural habitat. Ganglion cells closely resembled the three main classes of cat retinal ganglion cells, and therefore were classified into alpha-, beta-, and gamma-type cells. The size of the alpha- and beta-type somas and dendritic fields increased with increasing distance from the area centralis. However, this eccentricity dependence was not as pronounced as in the cat. The gamma-type cells were sub-divided into mono-, bi-, and diffusely stratified, in accordance with the ramification of their dendrites within the inner plexiform layer. The alpha- and beta-type cells were uni-stratified in either the sublamina of the inner plexiform layer closest to the ganglion cell layer or in that closest to the inner nuclear layer. These laminae correspond to those in the cat retina which contain the dendritic ramifications of ganglion cells whose central receptive fields respond best to onset of light (the "on-centre" cells), or to ganglion cells whose centres respond optimally to light being extinguished (the "off-centre" cells). Thus the flying fox retina contains a morphological correlate of the "on"/"off" dichotomy of alpha and beta cells in the cat retina. In general the flying fox retinal ganglion cells exhibit a degree of morphological complexity reminiscent of cat retinal cells and this may reflect similar functional properties.     

Morphology of rabbit retinal ganglion cells projecting to the medial terminal nucleus of the accessory optic system

Rabbit retinal ganglion cells were retrogradely labeled following injection of rhodamine-labeled microspheres into the medial terminal nucleus. The small fraction of rhodamine-labeled neurons reached their peak concentration within the visual streak and then decreased with increasing eccentricity until none were encountered in the far periphery. The same rabbits also received injections of the fluorescent tracer Fast Blue into the superior colliculus. No double-labeled neurons were observed, i.e., ganglion cells projecting to the medial terminal nucleus (MTN) had no axon collaterals to the superior colliculus. In fixed retinae rhodamine-labeled ganglion cells were intracellularly injected with the fluorescent dye Lucifer Yellow to reveal their full dendritic arborization. MTN-projecting cells had medium-sized to large somata with thin and frequently branched dendrites. The large dendritic trees had a distinct morphology and were predominantly unistratified in a narrow band that presumably corresponded to the electrophysiologically determined on-sublamina of the inner plexiform layer. Dendritic field sizes were inversely related to ganglion cell density, thus providing an eccentricity-independent, constant dendritic coverage factor. Approximately five to six dendritic fields from neighboring cells cover every point of the retina. Published reports claim that the physiological class of on-direction-selective ganglion cells provides the sole retinal input to the MTN in the rabbit. In this context morphological features of MTN-projecting cells and their presumed functional correlation with on-direction-selective ganglion cells are discussed.     

Subgroup of alpha ganglion cells in the adult cat retina is immunoreactive for somatostatin.

We have previously shown that two types of cells in the ganglion cell layer of the adult cat retina are immunoreactive for somatostatin (White et al., '90). One of the types was identified by morphological criteria as a wide-field amacrine cell. The other cell type had a large, angular soma that resembled the alpha ganglion cell, but evidence was not available to identify it definitively as a ganglion cell. Both cell types were distributed preferentially in the inferior retina. In this report, we demonstrate that the two types of cell are, indeed, displaced amacrine cells and alpha ganglion cells. First, when retrograde tracers were injected into central visual targets, the immunoreactive large cells but not the displaced amacrine cells were found to be labeled. Second, after unilateral section of the optic nerve, the immunoreactive large cells disappeared from the retina on the lesioned side, but the displaced amacrine cells occurred in the same numbers in both retinae. In the periphery, the large cells ranged in diameter from 33 to 47 microns, comparable only to alpha ganglion cells (Boycott and W?ssle, '74). An antiserum to parvalbumin was used to visualize the dendrites (R?hrenbeck and W?ssle, '88) of somatostatin-immunoreactive large cells. Based on dendritic stratification within the inner plexiform layer (Famiglietti and Kolb, '76), the somatostatin-immunoreactive large cells were found to include both on-center cells and off-center cells, but were predominantly of the off-center type. Within a local region, they were found to be arrayed with greater regularity than the overall population of alpha ganglion cells. These results indicate that alpha ganglion cells of the cat retina can be subdivided on the basis of their immunoreactive staining for somatostatin and suggest that the diversity of ganglion cells in the cat retina may be greater than has been recognized on the basis of morphological criteria alone.     

The zeta cell: A new ganglion cell type in cat retina

We define a new morphological type of ganglion cell in cat retina by using intracellular staining in vitro. The zeta cell has a small soma, slender axon, and compact, tufted, unistratified dendritic arbor. Dendritic fields were intermediate in size among cat ganglion cells, typically twice the diameter of beta cell fields. They were smallest in the nasal visual streak (<280 μm diameter), especially near the area centralis (60–150 μm diameter), and largest in the nonstreak periphery (maximum diameter 570 μm). Fields sizes were symmetric about the nasotemporal raphe except near the visual streak, where nasal fields were smaller than temporal ones. Zeta-cell dendrites ramified near the boundary between sublaminae a and b(OFF and ON sublayers) of the inner plexiform layer, occupying the narrow gap separating the dendrites of ON and OFF alpha cells. There was no evidence for separate ON and OFF types of zeta cell. Retrograde labeling studies revealed that both nasally and temporally located zeta cells project to the contralateral superior colliculus, whereas few project to the ipsilateral colliculus or to any subdivision of the dorsal lateral geniculate nucleus. The zeta cell's morphology and projection patterns suggest that it corresponds to the ON-OFF phasic W-cell (also known as the local edge detector) of physiological studies. Zeta cells have particularly small dendritic fields in the visual streak, presumably because they are disproportionately represented in the streak in comparison with other ganglion cell types. These conditions are consistent with optimal spatial resolution along the retinal projection of the visual horizon rather than principally at the center of gaze. Strong commonalities with similar ganglion cell types in ferret, rabbit, and monkey suggest that “zeta-like” cells may be a universal feature of the mammalian retina. J. Comp. Neurol. 399:269–288, 1998. © 1998 Wiley-Liss, Inc.     

Morphology and tracer coupling pattern of alpha ganglion cells in the mouse retina

Alpha cells are a type of ganglion cell whose morphology appears to be conserved across a number of mammalian retinas. In particular, alpha cells display the largest somata and dendritic arbors at a given eccentricity and tile the retina as independent on- (ON) and off-center (OFF) subtypes. Mammalian alpha cells also express a variable tracer coupling pattern, which often includes homologous (same cell type) coupling to a few neighboring alpha cells and extensive heterologous (different cell type) coupling to two to three amacrine cell types. Here, we use the gap junction-permeant tracer Neurobiotin to determine the architecture and coupling pattern of alpha cells in the mouse retina. We find that alpha cells show the same somatic and dendritic architecture described previously in the mammal. However, alpha cells show varied tracer coupling patterns related to their ON and OFF physiologies. ON alpha cells show no evidence of homologous tracer coupling but are coupled heterologously to at least two types of amacrine cell whose somata lie within the ganglion cell layer. In contrast, OFF alpha cells are coupled to one another in circumscribed arrays as well as to two to three types of amacrine cell with somata occupying the inner nuclear layer. We find that homologous coupling between OFF alpha cells is unaltered in the connexin36 (Cx36) knockout (KO) mouse retina, indicating that it is not dependent on Cx36. However, a subset of the heterologous coupling of ON alpha cells and all the heterologous coupling of OFF alpha cells are eliminated in the KO retina, suggesting that Cx36 comprises most of the junctions made with amacrine cells.     

Distribution and coverage of beta cells in the cat retina

We have reexamined the retinal distribution and dendritic field dimensions of beta cells in the cat retina. Beta cells were labeled by retrograde transport from the A-layers of the lateral geniculate nucleus and distinguished from alpha cells on the basis of soma size. Dendritic fields of beta cells were visualized by intracellular staining in vitro. The fraction of cat ganglion cells that were beta cells varied with retinal location. Except near the area centralis, beta cells represented about half of all ganglion cells in the nasal hemiretina. They contributed as heavily as the other major ganglion cell classes to the nasal visual streak. In and near the area centralis and in the temporal retina, beta cells represented about two-thirds of all ganglion cells. The areas of beta cell dendritic fields were reciprocally related to beta cell density. For example, they were 3-fold smaller within the visual streak than at matched eccentricities outside it. For many cells, we could estimate both local beta cell density and dendritic field area. Coverage factor (dendritic field area × local density) remained constant at about 4 despite 100-fold variations in beta cell density, and was independent of eccentricity, nasotemporal location, or position relative to the visual streak. Analysis in terms of sampling theory suggests that the beta cell array is matched to X-cell spatial resolution so as to optimize acuity. The beta cell distribution and its systematic reflection in dendritic architecture predict acuity levels that apparently correlate well with actual visual performance across the cat's visual field. © 1996 Wiley-Liss, Inc.     

Somatostatin-immunoreactive cells in the adult cat retina

Peptides have been found in the retinas of all mammalian species studied to date, but little is known about their localization and function in the cat. Using two mouse monoclonal antibodies directed to somatostatin 14, we have observed two sparse groups of somatostatin-immunoreactive neurons in the cat, both distributed preferentially in the inferior retina. The more numerous cell type is characterized by a small- to medium-sized soma (mean diameter = 16.3 +/- 9.0 microns; n = 186) with sparsely branching, far-reaching varicose processes that ramify mainly in the inner plexiform layer. The majority of these cells are located in the ganglion cell layer, with the remainder in the proximal inner nuclear layer and the inner plexiform layer. They are in especially high density at the retinal margin. In morphology and soma size, these cells resemble wide-field amacrine cells. The second cell type has a large, granular-staining soma (mean diameter = 29.7 +/- 14.8 microns; n = 145) with poorly stained primary processes and is found only in the ganglion cell layer. Cells of this type are most similar in their size and morphology to alpha ganglion cells. In contrast to the location of somatostatin-immunoreactive somata, a dense meshwork of immunoreactive processes was observed at all eccentricities within the inner plexiform layer, adjacent to the inner nuclear layer and to the ganglion cell layer. Labeled processes arising from the inner plexiform layer were also occasionally detected in the outer plexiform layer and the nerve fiber layer. Additional processes of unknown origin were observed in the nerve fiber layer and the optic nerve head. The extensive distribution of immunoreactive processes suggests that somatostatin-immunoreactive somata located preferentially in the inferior half of the retina have a widespread influence on neural activity.     

Dendritic remodelling of retinal ganglion cells during development of the rat

Investigation of the morphology of ganglion cells in the cat retina has shown that a remarkable reduction in the number of dendritic spines and branches occurs during development of the alpha and beta cell classes. To learn whether dendritic remodelling represents a generalized mechanism of mammalian retinal ganglion cell development, we have examined the morphology of ganglion cells in the retina of the developing rat. The present study has concentrated on type II cells, which retain a great number of dendritic spines and branches in the adult and comprise a large proportion of the population of rat retinal ganglion cells. To reveal fine dendritic and axonal processes, Lucifer yellow was injected intracellularly in living retinae maintained in vitro. Size and complexity of the dendritic trees were found to increase rapidly during an intial stage of development lasting from late fetal life until approximately postnatal day 12 (P12). Dendrites and axons of immature ganglion cells expressed several transient morphological features comprising an excessive number of dendritic branches and spine-like processes, and short, delicate axonal sidebranches. The following developmental stage was characterized by a remarkable decrease in the morphological complexity of retinal ganglion cells and a slowed growth of their dendritic fields. The number of dendritic branches and spines of types I and II retinal ganglion cells declined after P12 to reach a mature level by the end of the first postnatal month. Thus, even cells that retain a highly complex dendritic tree into the adult state undergo extensive remodelling. These results suggest that regressive modifications at the level of the dendritic field constitute a generalized mechanism of maturation in mammalian retinal ganglion cells. © 1993 Wiley-Liss, Inc.