Morphological and functional identifications of catfish retinal neurons. II. Morphological identification

Abstract
In this study the morphological origins of the responses from the catfish retinal neurons evoked by step inputs were determined by injecting intracellularly a dye, Procion yellow. A method was devised to view the dye-injected neurons in flat mount to study their dendritic expansion; later the same neurons could be sectioned radially to locate the levels of their somata or dendritic expansion. The results of this study show the inherent danger of identifying dye-injected neurons only in a radial or tangential view. Bipolar cells could be identified functionally without any ambiguity by changing widely the stimulus parameters, because the stimulation of their receptive-field center and surround gave rise to responses of opposing polarity. We found no exception to this rule. The neurons in the proximal layers produced a large variety of responses which could not be segregated into two such classes as the amacrine and ganglion cells. In this part II they were classified into three broad categories: neurons giving rise to sustained, transient, and spiking responses. The demarcation among the three types, morphologywise and functionwise, was vague and not well established. The sustained responses were recoreded from the starburst and spaghetti neurons (part I (9)) which correspond to Ramon y Cajal's (2) amacrine cells. The transient responses, whose patterns were largely invariant of the changes in the stimulus parameters, were recorded from a class of neurons with spindle-shaped somata in the INL. We do not know whether they had axons or not, but we will not be surprised if a future study defines them as a class of ganglion cells. Responses with or without spike discharges were recorded from a class of neurons which were identified as ganglion cells. Observations made on a large number of Procion-injected neurons in both flat-mount preparations and radial sections show that finer dendritic arborizations were not seen in the dye-injected neurons although the presence of such branches was proved in the Golgi preparations. Probably this was due to the weak contrast of the Procion-injected cell against the tissue background, rather than the failure of the dye to diffuse into finer branches. We recognize the severe difficulty involved in the traditional approach of identifying a class of neurons based on typical but subjectively selected functional and structural samples. Neurons have to be classified statistically according to their (quantitative) parameters. (cont'd)