We quantified the spatial distribution of presumed ganglion cells and displaced amacrine cells in unstained whole mounts of six young normal human retinas whose photoreceptor distributions had previously been characterized. Cells with large somata compared to their nuclei were considered ganglion cells; cells with small somata relative to their nuclei were considered displaced amacrine cells. Within the central area, ganglion cell densities reach 32,000-38,000 cells/mm2 in a horizontally oriented elliptical ring 0.4-2.0 mm from the foveal center. In peripheral retina, densities in nasal retina exceed those at corresponding eccentricities in temporal retina by more than 300%; superior exceeds inferior by 60%. Displaced amacrine cells represented 3% of the total cells in central retina and nearly 80% in the far periphery. A twofold range in the total number of ganglion cells (0.7 to 1.5 million) was largely explained by a similar range in ganglion cell density in different eyes. Cone and ganglion cell number were not correlated, and the overall cone:ganglion cell ratio ranged from 2.9 to 7.5 in different eyes. Peripheral cones and ganglion cells have different topographies, thus suggesting meridianal differences in convergence onto individual ganglion cells. Low convergence of foveal cones onto individual ganglion cells is an important mechanism for preserving high resolution at later stages of neural processing. Our improved estimates for the density of central ganglion cells allowed us to ask whether there are enough ganglion cells for each cone at the foveal center to have a direct line to the brain. Our calculations indicate that 1) there are so many ganglion cells relative to cones that a ratio of only one ganglion cell per foveal cone would require fibers of Henle radiating toward rather than away from the foveal center; and 2) like the macaque, the human retina may have enough ganglion cells to transmit the information afforded by closely spaced foveal cones to both ON- and OFF-channels. Comparison of ganglion cell topography with the visual field representation in V1 reveals similarities consistent with the idea that cortical magnification is proportional to ganglion cell density throughout the visual field.

Topography of ganglion cells in human retina.

Golgi-impregnated ganglion cells were studied in two flat-mounted human retinas A number of different morphologic forms were observed, and those showing a thickly branching dendritic field with terminals that stratified within a narrow zone of the inner plexiform layer were selected for further investigation When the dendritic field diameter of these cells was plotted against distance from the fovea, the scatter diagram showed two distinct clusters At any given eccentricity, there was no overlap between the cell group with large dendritic fields and the group with small dendritic fields Those with the larger dendritic fields also tended to have larger somas and thicker axons than the group with the smaller dendritic fields The dendritic fields of both groups tended to be elongated, and the orientation and degree of this elongation were quantified by determining the best-fitting ellipse for each dendritic field The degree of elongation was independent of eccentricity The orientation of the dendritic field (major axis of the ellipse) of a cell did not appear to be independent of its position on the retina To test whether the major axes tended to be directed toward any particular point on the retina, the positions of the cells on the retinal flat mount were transformed to relative positions on the retinal hemisphere, and the orientations of the dendritic fields were expressed in a spherical coordinate system for this hemisphere A search was made for the position on the hemisphere which minimized the mean square deviation of the orientations from this point The group with the large dendritic fields showed a significant tendency to be radially oriented toward a specific location on the retinal hemisphere, and that location lay within a few degrees of the fovea Leventhal and Schall ('83) have reported a similar finding for ganglion cells of the cat retina For the group with small dendritic fields, the retinal location that minimized the mean square deviation was also near the fovea; however, the set of orientations showed no greater tendency for mutual alignment than did a randomized set The cell group with the large dendritic fields appears to correspond to Dogiel's (1891) type II cells, to Polyak's ('41) parasol cells, to the A cells of the monkey retina described by Leventhal et al ('81), observed following HRP injection to the magnocellular layer of the LGN, and to the P alpha cells of the monkey retina, observed by Perry and Cowey ('81), following HRP uptake by cut axons of the optic nerve(ABSTRACT TRUNCATED AT 400 WORDS)

Parasol and midget ganglion cells of the human retina.

The autonomic nervous system influences numerous ocular functions. It does this by way of parasympathetic innervation from postganglionic fibers that originate from neurons in the ciliary and pterygopalatine ganglia, and by way of sympathetic innervation from postganglionic fibers that originate from neurons in the superior cervical ganglion. Ciliary ganglion neurons project to the ciliary body and the sphincter pupillae muscle of the iris to control ocular accommodation and pupil constriction, respectively. Superior cervical ganglion neurons project to the dilator pupillae muscle of the iris to control pupil dilation. Ocular blood flow is controlled both via direct autonomic influences on the vasculature of the optic nerve, choroid, ciliary body, and iris, as well as via indirect influences on retinal blood flow. In mammals, this vasculature is innervated by vasodilatory fibers from the pterygopalatine ganglion, and by vasoconstrictive fibers from the superior cervical ganglion. Intraocular pressure is regulated primarily through the balance of aqueous humor formation and outflow. Autonomic regulation of ciliary body blood vessels and the ciliary epithelium is an important determinant of aqueous humor formation; autonomic regulation of the trabecular meshwork and episcleral blood vessels is an important determinant of aqueous humor outflow. These tissues are all innervated by fibers from the pterygopalatine and superior cervical ganglia. In addition to these classical autonomic pathways, trigeminal sensory fibers exert local, intrinsic influences on many of these regions of the eye, as well as on some neurons within the ciliary and pterygopalatine ganglia.

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Autonomic control of the eye

Publisher Summary This chapter discusses the structure and regulation of the corpus allatum. The corpora allata (CA) are endocrine glands in the posterior regions of the head, or in rare instances in the thorax, which are closely associated with the stomatogastric nervous system. This chapter focuses on the regulation of the CA. It also describes the embryology, innervations, and the relationships of the structure, particularly the ultrastructure, to its synthetic activity. The characteristic shape of the CA is ovoid to round but they may be elongate as in large larvae and adults of Libellula depressa. The size of the glands is frequently about the diameter of the aorta or smaller; however, there is much variation among species and even within a species, size differs with age, sex, polymorphism, and the activity cycle of the glands. Although only one type of glandular cell occurs in the CA, there are a variety of types of CA with respect to the number of cells per gland and the relative size of the cells. The CA are surrounded by a continuous noncellular basal lamina, roughly 0.1-1 μ m thick. This material occasionally projects between two glandular cells into the interior of the gland, forming trabeculae that may accompany nerves and trachea.

Structure and Regulation of the Corpus Allatum

There are few neurochemical markers that reliably identify retinal ganglion cells (RGCs), which are a heterogeneous population of cells that integrate and transmit the visual signal from the retina to the central visual nuclei. We have developed and characterized a new set of affinity-purified guinea pig and rabbit antibodies against RNA-binding protein with multiple splicing (RBPMS). On western blots these antibodies recognize a single band at 〜24 kDa, corresponding to RBPMS, and they strongly label RGC and displaced RGC (dRGC) somata in mouse, rat, guinea pig, rabbit, and monkey retina. RBPMS-immunoreactive cells and RGCs identified by other techniques have a similar range of somal diameters and areas. The density of RBPMS cells in mouse and rat retina is comparable to earlier semiquantitative estimates of RGCs. RBPMS is mainly expressed in medium and large DAPI-, DRAQ5-, NeuroTrace- and NeuN-stained cells in the ganglion cell layer (GCL), and RBPMS is not expressed in syntaxin (HPC-1)-immunoreactive cells in the inner nuclear layer (INL) and GCL, consistent with their identity as RGCs, and not displaced amacrine cells. In mouse and rat retina, most RBPMS cells are lost following optic nerve crush or transection at 3 weeks, and all Brn3a-, SMI-32-, and melanopsin-immunoreactive RGCs also express RBPMS immunoreactivity. RBPMS immunoreactivity is localized to cyan fluorescent protein (CFP)-fluorescent RGCs in the B6.Cg-Tg(Thy1-CFP)23Jrs/J mouse line. These findings show that antibodies against RBPMS are robust reagents that exclusively identify RGCs and dRGCs in multiple mammalian species, and they will be especially useful for quantification of RGCs.

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The RNA binding protein RBPMS is a selective marker of ganglion cells in the mammalian retina.

Retinal ganglion cells were labeled retrogradely by localized injections of HRP into different regions of the pretectum, tectum, and optic tract in 26 cats. Retinal projection zones in the pretectum were labeled anterogradely in the same cats by intravitreal injections of 3H-proline. This allowed the HRP injection sites to be located with respect to the retinal termination zones. The form of the projection zones from retina to pretectum was determined from serial reconstructions of either coronal or horizontal sections. The zones are best distinguished in horizontal sections, where they are seen as four roughly parallel strips on either side of the brain. They are more-or-less parallel to the anterior border of the tectum, and appear to traverse the entire width of the retinal projection to the tectum. Each zone is similar in form for the ipsilateral and contralateral projections, although the contralateral projection is thicker and denser. Binocular injections of 3H-proline showed that the projections from the two eyes were in register and did not interdigitate. Cells labeled by HRP injections in the anteromedial end of the pretectum were concentrated in the lower nasal quadrant of the contralateral retina, and the lower temporal quadrant of the ipsilateral retina. Posterolateral injections labeled cells in the upper quadrants. There is thus a rough retinotopic mapping along the elongated axis of the pretectum. When the distributions of ganglion cells labeled by HRP injections to different parts of the pretectum are combined, they show a concentration in both the visual streak and area centralis, and thereby reflect, at least qualitatively, the relative spatial distribution of the entire ganglion-cell population. About 85% of the retinal projection to the pretectum is contralateral. For all of the HRP injections, the spatial density of labeled cells was always low, accounting for no more than 3% of the total spatial density of ganglion cells in any retinal region. Several types of ganglion cells were labeled following injections to most regions of the pretectum; these included alpha, beta, and epsilon cells, as well as small-bodied cells showing a variety of morphologic forms. Alpha cells were labeled mainly from the anterolateral end of the pretectum, but other cell types were labeled from all injected regions. In the peripheral retina, 2% of the labeled cells were alpha cells, 32% were beta cells, 19% were epsilon cells, and the remaining 47% were small cells whose dendrites only occasionally filled to any significant extent.(ABSTRACT TRUNCATED AT 400 WORDS)

The retinal projection to the cat pretectum

The distribution patterns of GABA immunoreactive (+) and immunonegative (-) amacrine cell synapses and profiles in the inner plexiform layer (IPL) were analyzed in three macaque monkey retinas using postembedding electron-microscopic (EM) immunogold cytochemistry. Synapses and profiles were counted at 5% intervals throughout the IPL depth in three EM montages (total area = 6509 microns 2), with 0% depth at the inner nuclear layer/IPL border. Nearly 70% of all amacrine synapses were GABA+, and they contacted all major classes of neurons that arborize in the IPL: bipolars (45%), ganglion cells (25%), and GABA+ (20%) and GABA- (10%) amacrines. A major relationship was seen between GABA+ amacrine processes and bipolar terminals: 76% of all amacrine-to-bipolar synapses were GABA+, and 82% of bipolar output dyads contained at least one GABA+ amacrine. GABA+ amacrine profiles (N = 2455) were concentrated in three wide bands at IPL depths of 0-25%, 40-60%, and 75-100%, corresponding to the dense bands seen with light-microscopic immunocytochemistry. In contrast, GABA+ amacrine synapses (N = 1081) were distributed evenly throughout the IPL depth, rather than being confined to the three dense bands. GABA- amacrine synapses (N = 516) were concentrated at 40% and 60% depths. Each category of amacrine output synapses had a characteristic pattern of stratification in the IPL. GABA+ amacrine-to-bipolar synapses occurred throughout the IPL but were most frequent at 20% and 80% IPL depths, where the dendrites of midget cone bipolars arborize (Polyak, 1941). In contrast, GABA+ amacrine-to-ganglion cell synapses were concentrated at 30% and 70% IPL depths, near the dendritic arborizations of parasol ganglion cells (Watanabe & Rodieck, 1989). GABA+ synapses onto bipolars and amacrines were also concentrated at the level of rod bipolar terminals. GABA+ amacrines must play significant but different roles in ON and OFF midget and parasol pathways as well as the rod pathway.

Distribution of GABA-immunoreactive amacrine cell synapses in the inner plexiform layer of macaque monkey retina.

Immunocytochemical localization of GABA and glycine in amacrine and displaced amacrine cells of macaque monkey retina

Distributions of bipolar (B) and amacrine (A) synapses and postsynaptic ganglion cell (G) dendritic profiles in the inner plexiform layer (IPL) were analyzed in EM montages of monkey central and human foveal and peripheral retinae. Synapses and profiles were counted and plotted for each 5% interval of IPL, with 0% at the inner edge of the inner nuclear layer and 100% at the outer edge of the ganglion cell layer. In monkey and human retinae, both A and B synapses occur throughout the IPL, but the ratio of A to B synapses varies from 2:1 to more than 6:1. In the monkey central retina, four bands of A conventional synapses are concentrated at 15, 35, 60, and 80% depth. In the human foveal slope, there are two main A bands at 45 and 85%, whereas in the human periphery, there are five bands at 15, 35, 60, 75, and 90%. In both species, A processes containing large dense-core vesicles are concentrated in three bands at 10-20, 50, and 80-90% depth, corresponding to previously described levels of peptides, dopamine, and GABA. B ribbon synapses are distributed fairly evenly throughout the IPL, with a suggestion of four broadly overlapping bands. Most B ribbons are presynaptic to one A and one G (B----A/G). In the human, there are significantly more B dyads with postsynaptic G's (B----A/G, B----G/G) in the fovea (91%) than in the periphery (66%), implying greater A cell processing peripherally. Also in the human, B terminals containing glycogenlike granules are concentrated in the outer half of the IPL, with agranular terminals in the inner half. Our results demonstrate multiple strata containing different types of synaptic contacts in primate IPL.

Stratified distribution of synapses in the inner plexiform layer of primate retina

Synaptic contacts onto fibers and somata in the nerve fiber layer (NFL) and ganglion cell layer (GCL) of macaque and human retina were demonstrated at the electron microscopical (EM) level. Many presynaptic processes in monkey NFL are gamma aminobutyric acid (GABA) immunoreactive, using anti-GABA antiserum with an EM immunogold procedure. Immunocytochemistry at the light microscopic level revealed that many GABA-reactive cells in the GCL send branching processes into the NFL, forming a sparse synaptic plexus. The presence of long, unbranched GABA-reactive fibers running horizontally in the NFL and entering the optic nerve suggests that some ganglion cells may be GABAergic. GABA-reactive cells contributing to the plexus appear to be a new class of displaced amacrines that arborize in the NFL.

Margaret A. Koontz

Papers

The retinal projection to the cat pretectum

Distribution of GABA-immunoreactive amacrine cell synapses in the inner plexiform layer of macaque monkey retina.

Immunocytochemical localization of GABA and glycine in amacrine and displaced amacrine cells of macaque monkey retina

Stratified distribution of synapses in the inner plexiform layer of primate retina

GABA-immunoreactive synaptic plexus in the nerve fiber layer of primate retina.