Mosaics and territories of cat retinal ganglion cells.
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Biomedical subjects
L Peichl
Publications and source records attributed to L Peichl.
The structural correlate of the receptive field centre of alpha ganglion cells in the cat retina.
The correlation between the receptive field centre and the dendritic tree of individual brisk transient, or alpha, ganglion cells in the cat retina was investigated by a combination of physiological and anatomical techniques. The sizes of receptive field centres of brisk transient (Y) cells were measured with a flickering spot of light. Contour maps and response (or sensitivity) profiles were measured at mesopic and scotopic backgrounds. Recording positions on the retina and nearby blood vessels were back-projected onto the receptive field plots on the tangent screen. After recording, whole amount preparations of the retinae were stained by a reduced silver method to stain all alpha cells together with their dendritic trees. By comparing the landmarks on the screen plot with those of the whole mount it was possible to identify the recorded cells in the preparation and to study their morphology. The dendritic tree of an alpha cell determines the position, size and shape of its receptive field centre. The mesopic receptive field centres were found to be a factor of 1.4 +/- 0.13 larger than their respective dendritic fields. It is suggested that the dendritic fields of presynaptic neurones (bipolar and amacrine cell processes) add to the ganglion cell dendritic tree to produce the larger centre summating area.
Action Potentials↗
Dendritic territories of cat retinal ganglion cells.
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Morphological identification of on- and off-centre brisk transient (Y) cells in the cat retina.
Brisk transient (Y) cells were recorded extracellularly in the cat retina. The position and shape of their receptive field centres were plotted on a tangent screen, together with retinal landmarks, such as blood vessels adjacent to the recording area. After recording the retina was processed as a whole mount and stained with a reduced-silver method (see appendix). This technique stains the entire alpha cell population including the dendritic trees. Alpha cells are the morphological correlate of the brisk transient cells (Boycott & Wässle 1974; Cleland et al. 1975). Maps of the screen plot and the histological preparation could be accurately superimposed by means of the retinal landmarks and each recorded brisk transient unit could unequivocally be attributed to a particular alpha cell. Alpha cell dendritic trees are unistratified in either of two laminae within the inner plexiform layer: (1) close to the inner nuclear layer border, 'outer alpha cells', or (2) about 10 micrometers further towards the ganglion cell layer, 'inner alpha cells'. This stratification difference can be observed in whole mounts for large populations of cells (Wässle et al. 1981). Of the recorded brisk transient cells, all on-centre units were inner alphas and all off-centre units outer alphas.
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Morphology and topography of on- and off-alpha cells in the cat retina.
Neurofibrillar staining methods were found to stain all alpha cells of the cat retina completely, that is the perikaryon, the axon and the dendritic branches. The dendrites of the alpha cells in vertical sections were found to be unistratified and to occupy two narrow strata in the outer half of the inner plexiform layer. This difference in branching level could also be observed in whole-mount preparations and it has been demonstrated in the preceding paper (Peichl & Wässle 1981) that it corresponds to the physiological on-off dichotomy. Thus the topographical distribution of on- and off-alpha cells could be studied. They are found to occur in about equal numbers. Both on- and off-alpha cell perikarya form a regular lattice and both lattices are superimposed independently. The dendritic branches of neighbouring alpha cells overlap and each retinal point is covered by the dendritic field of at least one on- and one off-alpha cell. The dendritic trees of on-alpha cells seem to have more small branches and are on the average smaller than those of off-alpha cells. The density of alpha cells was found to peak in the central area whence it continuously decreased towards the retinal periphery.
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Almost all ganglion cells in the rabbit retina project to the superior colliculus.
After a localized injection of horseradish peroxidase into the superior colliculus of the rabbit almost all of the ganglion cells in the monocular portion of the contralateral retina were retrogradely labelled with the enzyme. Of the few ganglion cells which were not labelled, most had large cell bodies.
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Quantification of horizontal cells in the domestic cat retina.
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Size, scatter and coverage of ganglion cell receptive field centres in the cat retina.
1. Receptive field centre sizes of brisk-sustained (X) and brisk-transient (Y) ganglion cells of the cat retina were assessed by three different methods: small spot mapping, area threshold method and spatial resolution. 2. Centre sizes of brisk-sustained (X) cells increased from 20' in the central area to about 70' at an eccentricity of 4.5 mm, centre sizes of brisk-transient (Y) cells from 50' in the central area to about 140' at 5 mm eccentricity. 3. The scatter of centre sizes at one retinal location was measured by recording as many ganglion cells as possible in one cat in a small field of retina. The centre sizes of the individual classes were homogeneous and exhibited only a small amount of scatter. 4. The coverage of the retina by the different ganglion cell classes was assessed from their density and their receptive field centre area. At every retinal location the receptive field centres of seven to twenty brisk-sustained (X) cells and of three to six brisk-transient (Y) cells were found to overlap. Sluggish concentric and non-concentric cells taken together have a coverage factor of about 60.
Action Potentials↗
Morphological types of horizontal cell in the retina of the domestic cat.
Two morphologically distinct types of horizontal cell are described from Golgi-stained whole mounts of the cat retina. They are referred to as A-type and B-type cells. The two types differ in their dendritic branching pattern, their overall size and the absence or presence of an axon. At every retinal position the dendrites of B-type cells branch more densely and overlap each other more frequently than do the dendrites of A-type cells. At equivalent retinal positions the dendritic field size of A-type cells is greater than that of B-type cells by a factor of about 1.5. Only B-type cells have an axon, which branches at the end into a large axon terminal system. The axons have no preferred direction of orientation. The stain-ability of horizontal cells by different Golgi methods is discussed.
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Receptor contacts of horizontal cells in the retina of the domestic cat.
The terminal aggregations of A- and B-type horizontal cells, stained by the Golgi-Colonnier method, have been analysed. The pattern of the aggregations is regular and is shown to be in register with the cone mosaic. Both tyes of horizontal cell are in contact with at least 80% of the cones above their dendritic fields. Therefore, the different horizontal cell classes cannot be selective for a special kind of cone but must have at least 60% of the cone input in common. Each A-type horizontal cell makes contacts with between 120 and 170 cones, and each B-type horizontal cell with 60-90 cones. An individual A-type horizontal cell occupies an average of 20% of the lateral elements of the triads in a cone pedicle, but an individual B-type cell fills only some 13%. Each and every cone is connected with several of both types of horizontal cell. An estimation of the number of rods converging onto a single axon terminal system showed that it could be as many as 3000.
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Topography of horizontal cells in the retina of the domestic cat.
Neurofibrillar methods stain a class of horizontal cells in the cat retina which are shown to be identical with the A-type horizontal cell of Golgi-staining. Thus all of the A-type cells of a single retina can be observed. On this basis the changes in density and dendritic field size of A-type horizontal cells with respect to retinal eccentricity were measured. The decrease in density from centre to periphery is balanced by a corresponding increase in size of the dendritic field. Consequently each retinal point--independent of retinal position--is covered by the dendritic fields of three of four A-type horizontal cells. The nuclei and nucleoli of B-type horizontal cells could also be recognized in neurofibrillar-stained material and thus their distribution was determined. The density ratio B-type: A-type is 2.8 +/- 0.4 and does not vary much from the centre to the periphery of the retina. Each retinal point is also covered by four B-type horizontal cells. Thus a single cone can contact a maximum of eight horizontal cells. The rate of density decrease from centre to periphery is closely similar in cones and horizontal cells but greater in ganglion cells.
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Are field potentials an appropriate method for demonstrating connections in the brain?
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Morphological types of horizontal cell in rodent retinae: a comparison of rat, mouse, gerbil, and guinea pig.
Retinal horizontal cells of four rodent species, rat, mouse, gerbil, and guinea pig were examined to determine whether they conform to the basic pattern of two horizontal cell types found in other mammalian orders. Intracellular injections of Lucifer-Yellow were made to reveal the morphologies of individual cells. Immunocytochemistry with antisera against the calcium-binding proteins calbindin D-28k and parvalbumin was used to assess population densities and mosaics. Lucifer-Yellow injections showed axonless A-type and axon-bearing B-type horizontal cells in guinea pig, but revealed only B-type cells in rat and gerbil retinae. Calbindin immunocytochemistry labeled the A- and B-type populations in guinea pig, but only a homogeneous regular mosaic of cells with B-type features in rat, mouse, and gerbil. All calbindin-immunoreactive horizontal cells in the latter species were also parvalbumin-immunoreactive; comparison with Nissl-stained retinae showed that both antisera label all of the horizontal cells. Taken together, the data from cell injections and the population studies provide strong evidence that rat, mouse, and gerbil retinae have only one type of horizontal cell, the axon-bearing B-type, whereas the guinea pig has both A- and B-type cells. Thus, at least three members of the family Muridae differ from other rodents and deviate from the proposed mammalian scheme of horizontal cell types. The absence of A-type cells is apparently not linked to any peculiarities in the photoreceptor populations, and there is no consistent match between the topographic distributions of the horizontal cells and those of the cone photoreceptors or ganglion cells across the four rodent species. However, the cone to horizontal cell ratio is rather similar in the species with and without A-type cells.
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Development of dendritic trees of rabbit retinal alpha ganglion cells: relation to differential retinal growth.
To provide a quantitative description of the postnatal development of dendritic trees in alpha ganglion cells of the rabbit retina, these cells were stained either by intracellular injection of Lucifer yellow or by application of the lipophilic dye DiI. This was done at three developmental stages: postnatal day (P) 8/9, P 16/17, and in adults. For different retinal locations we quantified the alpha cell dendritic field area, the number of dendritic branch points, and the average dendritic length between branch points. According to the alpha cell location, the data were collected in three groups representing the retinal center, midperiphery, and far periphery, respectively. The data were then correlated with the postnatal retinal expansion which is known to differ among the above topographic regions of the retinae (Reichenbach et al., 1993). Our results show that the growth of alpha ganglion cell dendrites is not proportional to, but significantly exceeds, that of the local retinal tissue. Between P 8/9 and adulthood, the area of central alpha cells increases almost six-fold from 26,000 to 144,000 microns 2 (retinal expansion: 2.2-fold), and that of peripheral cells more than 15-fold from 35,000 to 556,000 microns 2 (retinal expansion: four-fold). During this period, the coverage factor of alpha cell dendritic fields increases about three-fold, and reaches adult levels of about 3 (retinal center) and 2.2 (periphery), respectively. The number of dendritic branch points remains nearly constant, and the distance between them increases by a factor close to the square root of the factor by which the dendritic field area grows. Thus, it appears that, from the second postnatal week on, dendritic trees of rabbit alpha ganglion cells increase by intense "interstitial growth," rather than by outgrowth of (new) dendritic branches. This growth pattern is different from that of some other rabbit retinal ganglion cell types, and of alpha ganglion cells of the cat retina, whose dendritic trees expand at a rate equal to or less than that of the surrounding retinal tissue. The consequences for synaptic contacts with bipolar and amacrine cells are discussed; they suggest a high degree of synaptic plasticity during normal postnatal retinal growth.
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The horizontal cells of artiodactyl retinae: a comparison with Cajal's descriptions.
The morphology of horizontal cells in ox, sheep, and pig retinae as observed after Lucifer Yellow injections are described and compared with the descriptions of Golgi-stained cells by Ramón y Cajal (1893). Horizontal cells in the retinae of less domesticated species, wild pig, fallow and sika deer, mouflon, and aurochs were also examined. All these retinae have two types of horizontal cell; their morphologies are in common, although with some familial differences. Their basic appearance is as Cajal described; except in one important respect, a single axon-like process could not be identified on the external horizontal cells. It is concluded that external horizontal cells of artiodactyls correspond to the axonless (A-type) cells of other mammals. Cajal's internal horizontal cells have a single axon which contacts rods. This type corresponds to the B-type cells of other mammalian retinae. Artiodactyl A- and B-type horizontal cells differ from those of many other mammals in that the B-type dendritic tree is robust and the A-type dendritic tree is delicate. Historically, this morphological difference between orders of mammals has led to some confusion. The comparisons presented here suggest that the morphological types of primate horizontal cells can be integrated into a general mammalian classification.
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Alpha ganglion cells in mammalian retinae: common properties, species differences, and some comments on other ganglion cells.
A specific morphological class of ganglion cell, the alpha cell, was first defined in cat retina. Alpha cells have since been found in a wide range of mammalian retinae, including several orders of placental and marsupial mammals. Characteristically, they have the largest somata and a large dendritic field with a typical branching pattern. They occur as inner and outer stratifying subpopulations, presumably corresponding to ON-center and OFF-center receptive fields. In all species, alpha cells account for less than 10% of the ganglion cells, their somata are regularly spaced, and their dendritic fields evenly and economically cover the retina in a mosaic-like fashion. The morphology of alpha cells and many features, both of single cells and of the population, are conserved across species with different habitats and life-styles. This suggests that alpha cells are a consistent obligatory ganglion cell type in every mammalian retina and probably subserve some fundamental task(s) in visual performance. Some general rules about the construction principles of ganglion cell classes are inferred from the alpha cells, stressing the importance of population parameters for the definition of a class. The principle, that a functionally and morphologically homogeneous population should have a regular arrangement and a complete and even coverage of the retina to perform its part in image processing at each retinal location, is especially evident across species and across ganglion cell types.
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Photoreceptor types and distributions in the retinae of insectivores.
The retinae of insectivores have been rarely studied, and their photoreceptor arrangements and expression patterns of visual pigments are largely unknown. We have determined the presence and distribution of cones in three species of shrews (common shrew Sorex araneus, greater white-toothed shrew Crocidura russula, dark forest shrew Crocidura poensis; Soricidae) and in the lesser hedgehog tenrec Echinops telfairi (Tenrecidae). Special cone types were identified and quantified in flattened whole retinae by antisera/antibodies recognizing the middle-to-long-wavelength-sensitive (M/L-)cone opsin and the short-wavelength-sensitive (S-)cone opsin, respectively. A combination of immunocytochemistry with conventional histology was used to assess rod densities and cone/rod ratios. In all four species the rods dominate at densities of about 230,000-260,000/mm2. M/L- and S-cones are present, comprising between 2% of the photoreceptors in the nocturnal Echinops telfairi and 13% in Sorex araneus that has equal diurnal and nocturnal activity phases. This suggests dichromatic color vision like in many other mammals. A striking feature in all four species are dramatically higher S-cone proportions in ventral than in dorsal retina (0.5% vs. 2.5-12% in Sorex, 5-15% vs. 30-45% in Crocidura poensis, 3-12% vs. 20-50% in Crocidura russula, 10-30% vs. 40-70% in Echinops). The functional and comparative aspects of these structural findings are discussed.
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