Amacrine cells in the retina of a teleost fish, the roach ( Rutilus rutilus ) : a Golgi study on differentiation and layering

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Abstract
We studied 250 amacrine cells in the retina of the tetrachromatic cyprinid Rutilus rutilus (roach) after rapid Golgi impregnation. All cells were recorded in camera lucida drawings from 50 -80 μm sections. For classifications we used independent criteria of presumed functional relevance, most of which could be quantified. These included ‘gross morphological’ features such as size, symmetry and orientation of the dendritic field, pattern of branching and number of ramification points as well as fine structural details like process diameter and the occurrence of spines and varicosities. We also took into account the pattern of radial distribution of dendrites. To obtain information about the subdivision of the inner plexiform layer, we used the relative stratification levels of stratified amacrine cells to plot a frequency distribution diagram showing that the dendrites of these cells are clustered in seven discrete sublayers of unequal width; four sublayers occupy the distal half of the inner plexiform layer (sublamina a) and three sublayers are present in the proximal half (sublamina b). The subdivision was compared with densitometric data of the inner plexiform layer after various staining methods and with previous observations about the location of bipolar terminals and neurochemical bandings. Our findings suggest that this layer is composed of complementary structural components, each of which is subject to a specific layering pattern. On this basis we could distinguish 43 different types of amacrine cell. If individual types occurred in more than one sublayer, they were considered as subtypes; of these, we found 70 different ones. Among the 43 types, 6 were observed only once. In comparison with amacrine cells described in other species, six ‘new’ types were identified. For each individual type, an identity chart was prepared summarizing camera lucida drawings of tangential views at low and high magnification, a semischematical drawing of the radial location of the dendrites, and the most relevant quantitative data. Our observations are discussed in the context of available evidence about light-evoked responses of identified amacrine types in other species, and possible transmitter content. They substantiate a functional concept according to which amacrine cells provide (i) a multicellular aggregate for coupled membrane potential; (ii) unit activity by the action of entire individual cells; and (iii) local microcircuits caused by isolated activation of single dendrites or parts thereof. The great variety of morphological differentiation, and the numerous transmitters found, suggest that within this basic framework individual amacrine types serve highly complex and sophisticated roles in retinal information processing. Our attempt towards a detailed classification and description of amacrine cell types is intended to provide a reference for future intracellular and neurochemical work, to facilitate precise identification.