Quantitative, three-dimensional analysis of granule cell dendrites in the rat dentate gyrus

The three-dimensional organization of dentate granule cell dendritic trees has been quantitatively analyzed with the aid of a computerized microscope system. The dendrites were visualized by iontophoretic injection of horseradish peroxidase into individual granule cells in the in vitro hippocampal slice preparation. Selection criteria insured that the analyzed cells were completely stained and that only neurons with two or fewer cut dendrites in the distal portion of the molecular layer were analyzed. Twenty-nine of the 48 sampled granule cells had no cut dendrites. The granule cells had between one and four primary dendrites. Granule cell dendritic branches were covered with spines and most extended to the hippocampal fissure or pial surface. The mean total dendritic length was 3,221 microns with a range from 2,324 microns to 4,582 microns. The dendrites formed an elliptical plexus with the transverse spread averaging 325 microns and the spread in the septotemporal axis averaging 176 microns. On individual neurons, the maximum branch order ranged from four to eight and the number of dendritic segments ranged from 22 to 40. Approximately 63% of the dendritic branch points occurred in a zone that included the granule cell layer and the inner one-third of the molecular layer. The dendritic tree was organized so that, on average, 30% of the length was in the granule cell layer and proximal third of the molecular layer, 30% was in the middle third, and 40% was in the distal third. Comparisons were made between the dendrites of granule cells in the suprapyramidal and infrapyramidal blades of the dentate gyrus. Suprapyramidal cells had a significantly greater total dendritic length than infrapyramidal cells, their transverse spread was higher, and they had a greater number of dendritic segments. When neurons in the suprapyramidal blade were further subdivided on the basis of somal position within the depth of the cell body layer, superficial neurons were found to have a greater number of primary dendrites, more elliptical trees, and larger transverse spreads of their dendrites. There were no significant differences in dendritic segment number or total dendritic length between superficial and deep cells.