The ON-alpha ganglion cell of the cat retina and its presynaptic cell types

Anatomical circuits converging onto the ON-alpha (Y) ganglion cell were studied by computer-assisted reconstruction of substantial portions of 2 alpha cells from electron micrographs of serial sections. The alpha cells in the area centralis were labeled by a Golgi-like retrograde filling with horseradish peroxidase, and certain presynaptic amacrine processes were labeled by uptake of 3H-glycine. About 4400 synapses contacted the alpha cell. Eighty-six percent were from amacrine cells; the rest were from bipolar cells. About one-quarter of the amacrine synapses were specifically labeled by 3H-glycine and probably belong to the A4 amacrine. The bipolar inputs were provided by several types: cone bipolar CBb1 (85%), cone bipolar CBb5 (2%), the rod bipolar (5%), and some unidentified cone bipolars (11%). Contacts from each type occurred in specific strata, with the consequence that they tended to form spots or annulli over the alpha dendritic field. The CBb1 bipolars formed a moderately dense array (8000/mm2), with a nearest-neighbor distance of 8.6 +/- 1.3 microns. Most members of the array (84%) contacted the alpha cell, providing 1-7 synapses (average, 2.7 +/- 1.6). The placement of contacts from an individual CBb1 followed certain rules: they were restricted to a parent branch of the alpha arbor or to 2 daughter branches, but almost never crossed a branch of the alpha arbor. The synaptic territory of an individual CBb1 was not shared with other b1s (or cone bipolars of any sort), although it was shared with amacrine contacts. Rod bipolar cells also formed a very dense array (54,500/mm2) in the alpha dendritic field, but only a few of these (3%) contacted the alpha cell. The concentric receptive field of the CBb1, combined with the spatial organization of its array, is used to predict the CBb1 contribution to the alpha cell receptive field; this contribution resembles the spatial and temporal organization of the alpha receptive field itself.