Morphology and physiology of the polyaxonal amacrine cells in the rabbit retina

We examined the morphology and physiological response properties of the axon-bearing, long-range amacrine cells in the rabbit retina. These so-called polyaxonal amacrine cells all displayed two distinct systems of processes: (1) a dendritic field composed of highly branched and relatively thick processes and (2) a more extended, often sparsely branched axonal arbor derived from multiple thin axons emitted from the soma or dendritic branches. However, we distinguished six morphological types of polyaxonal cells based on differences in the fine details of their soma/dendritic/axonal architecture, level of stratification within the inner plexiform layer (IPL), and tracer coupling patterns. These morphological types also showed clear differences in their light-evoked response activity. Three of the polyaxonal amacrine cell types showed on-off responses, whereas the remaining cells showed on-center responses; we did not encounter polyaxonal cells with off-center physiology. Polyaxonal cells respected the on/off sublamination scheme in that on-off cells maintained dendritic/axonal processes in both sublamina a and b of the IPL, whereas processes of on-center cells were restricted to sublamina b. All polyaxonal amacrine cell types displayed large somatic action potentials, but we found no evidence for low-amplitude dendritic spikes that have been reported for other classes of amacrine cell. The center-receptive fields of the polyaxonal cells were comparable to the diameter of their respective dendritic arbors and, thus, were significantly smaller than their extensive axonal fields. This correspondence between receptive and dendritic field size was seen even for cells showing extensive homotypic and/or heterotypic tracer coupling to neighboring neurons. These data suggest that all polyaxonal amacrine cells are polarized functionally into receptive dendritic and transmitting axonal zones.