New metrics for analysis of dendritic branching patterns demonstrating similarities and differences in ON and ON‐OFF directionally selective retinal ganglion cells
Authors: Famiglietti E. V. 
Published: May 2005

The morphology and dendritic branching patterns of retinal ganglion cells have been studied in Golgi‐impregnated, whole‐mount preparations of rabbit retina. Among a large number of morphological types identified, two have been found that correspond to the morphology of ON and ON‐OFF directionally selective (DS) ganglion cells identified in other studies. These two kinds of DS ganglion cell are compared with each other, as well as with examples of class I, class II, and class III cells, defined here with reference to our previous studies. Cell body, dendritic field size and branching pattern are analyzed in this paper and levels of dendritic stratification are examined in the following paper. ON DS ganglion cells are about 10% larger in soma size and about 5 times the dendritic field area of ON‐OFF DS ganglion cells, when compared at the same retinal location. These two morphological types of ganglion cell can be said to define the upper and lower bounds of an intermediate range of cell body and dendritic field sizes within the whole population of ganglion cells. Nevertheless, in previous physiological studies receptive field sizes of the two types were shown to be similar. This discrepancy between morphological and physiological evidence is considered in the Discussion in terms of a model of the excitatory receptive field of ON‐OFF DS ganglion cells incorporating starburst amacrine cells.

A new set of metrics is introduced here for the quantitative analysis and characterization of the branching pattern of neuronal arborizations. This method compares the lengths of terminal and preterminal dendritic branches (treated separately), as a function of the distances of their origins from the soma, viewed graphically in a two‐dimensional scatter plot. These values are derived from computer‐aided 3D logging of the dendritic trees, and distance from the soma is measured as the shortest distance tracked along the dendritic branches. From these metrics of the “branch length distributions,” scale‐independent branching statistics are derived. These make use of mean branch lengths and distances, slopes of lines fitted to the distributions, and elliptical indices of scatter in the distributions. By these measures, ON and ON‐OFF DS ganglion cells have similar branching patterns, which they share to varying degrees with functionally unrelated class III.1 ganglion cells. The scale of the branching patterns of ON and ON‐OFF DS cells and their degree of uniformity are different, however. ON‐OFF DS ganglion cells are the most uniform of all the cells examined, and epitomize the “tufted” branching pattern, while class Ia2 cells represent the other extreme of the “radiate” pattern. ON DS cells, like class III.1 cells, exhibit inhomogeneities or “patchiness” in the distribution of short dendritic branches within the dendritic tree. The functional significance of these inhomogeneities and the uniformity of branching in ON‐OFF DS cells is discussed, and the merits of the new methods of branching analysis are compared with methods previously used.