A distinct subpopulation of bipolar cells in macaque monkey retina was labeled with antisera that recognize glycine-extended cholecystokinin precursors. The labeled bipolar cells were found throughout the retina and had dendrites contacting a subpopulation of cone pedicles and axons ramifying in the fifth stratum of the inner plexiform layer. Several lines of evidence indicate that the labeled bipolar cells are a single type despite some variations in their morphology. First, the density of perikarya and their diameters varied continuously as a function of eccentricity. Second, the positions of perikarya within the inner nuclear layer and the level at which the axons branched in the inner plexiform layer were constant at all eccentricities. Bipolar cells with similar morphology have been described previously as "blue cone bipolar cells" (Mariani, 1984b), but there was no direct evidence that this was the case. In this study, we show by light microscopy that labeled bipolar cells have dendrites ending exclusively upon presumptive blue cones labeled by Procion black dye. All blue cones were contacted by labeled bipolar cells, and virtually all bipolar cells contacted blue cones, the only exceptions being in regions where blue cones had been lost. Approximately 20% more labeled bipolar cells than blue cones were found at every eccentricity; thus, connections between blue cones and labeled bipolar cells were not strictly one to one. The mean number of cones presynaptic to each bipolar cell was 1.2, and the mean number of bipolar cells postsynaptic to each cone was 1.8. By an electron microscopic study of labeled bipolar cell dendrites, we determined that they became central elements of ribbon synapses in blue cones. Some of their ribbon synapses were unusual: in one type, a single, large labeled dendrite was postsynaptic to two or more ribbons, while in the other type, ribbons had two or more central elements. The presence of these invaginating contacts and the axonal terminals in the proximal inner plexiform layer suggest that the labeled bipolar cells depolarize to short-wavelength stimuli and function to relay information from blue cones to the inner plexiform layer. There were also other, unlabeled bipolar cell dendrites that received inputs from blue cones at basal junctions and triad-associated flat contacts, which suggests that there are additional types of bipolar cells conveying information from short-wavelength cones in the primate retina.
In primates, the retinal ganglion cells that project to the magnocellular layers of the lateral geniculate nucleus have distinctive responses to light, and one of these has been identified morphologically as the parasol ganglion cell. To investigate their synaptic connections, we injected parasol cells with Neurobiotin in lightly fixed baboon retinas. The five ON-center cells we analyzed by electron microscopy received approximately 20% of their input from bipolar cells. The major synaptic input to parasol cells was from amacrine cells via conventional synapses and, in this respect, they resembled alpha ganglion cells of the cat retina. We also found the gap junctions between amacrine cells and parasol ganglion cells that had been predicted from tracer-coupling experiments. To identify the presynaptic amacrine cells, ON-center parasol cells were injected with Neurobiotin and Lucifer yellow in living macaque retinas, which were then fixed and labeled by immunofluorescence. Two kinds of amacrine cells were filled with Neurobiotin via gap junctions: a large, polyaxonal cell containing cholecystokinin and a smaller one without cholecystokinin. There were also appositions between cholecystokinin-containing amacrine cell processes and parasol cell dendrites. Cholinergic amacrine cell processes often followed parasol cell dendrites and made extensive contacts. In other mammals, the light responses of polyaxonal amacrine cells like these and cholinergic amacrine cells have been recorded, and the effects of acetylcholine and cholecystokinin on ganglion cells are known. Using this information, we developed a model of parasol cells that accounts for some properties of their light responses.
The short wavelength-sensitive (blue) cone bipolar cells was found to have a nonrandom distribution by analyzing the nearest neighbors and by calculating the density recovery profile (DRP). Blue cones had been shown previously to have a nonrandom distribution (Curcio et al., 1991). The relationship between the two arrays was then analyzed by calculating the cross-correlational density recovery profile (cDRP), which indicates the local density of blue cones around each blue cone bipolar cell. Although both cell types appeared to be distributed uniformly at the macroscopic level, the cDRP was 1.7 times higher within 15 μm of each bipolar cell perikaryon than in the surrounding area. The area of higher density was approximately the same as that in which the blue cone bipolar cells made synaptic contacts with blue cones. The finding that the blue cones and the blue cone bipolar cells were closer together than expected suggests that the positions of the perikarya of these neurons were influenced by their synaptic connections or other developmental interactions.
The synaptic contacts between photoreceptors and horizontal cells in the retina of the turtle (Geoclemys) were studied. Horizontal cells were classified into three types according to their intracellularly recorded spectral responses: luminosity, biphasic chromaticity, and triphasic chromaticity horizontal cells (LHC, BHC, and THC). These cells were then iontophoretically filled with horseradish peroxidase (HRP). The various types of photoreceptors located within the dendritic field of the HRP-filled horizontal cells were identified either as rods or as one of the three chromatic types of cones; the latter were identified by the presence and colors of their oil droplets in the inner segments. The synaptic contacts between photoreceptors and labeled horizontal cells were then investigated by light and electron microscopy of serial sections on three LHCs, two BHCs, and one THC. LHCs made synaptic contacts with about 100 photoreceptors, including rods and three chromatic types of cones; two-thirds of these photoreceptors contacted the cell body and the remaining its axon terminal. BHCs contacted about 30 cones; red-, green-, and blue-sensitive cones in the ratio of 3:4:1. THC contacted 20 cones; red- and blue-sensitive cones in the ratio of 2:1. The dendritic processes of HRP-filled horizontal cells were found as the lateral element of the ribbon synaptic complexes. The present finding suggests that the responses of BHC and THC to red flashes are fed directly from red-sensitive cones in addition to the feedback pathway of cone-horizontal cell connections in the previous studies.
Targeting by using the Régis-Valliccioni frame was very accurate compared with targeting with coordinates based on brain maps used hitherto. Although targeting improved the accuracy, further effort will still be necessary to reduce errors along dorsoventral axis. The apomorphine test indicated a reduced dopaminergic function of the irradiated area including striatum, which accompanied histological changes after a high dose of irradiation (150 Gy).
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