Different granule cell populations innervate superficial and deep regions of the external plexiform layer in rat...

Studies on the morphological organization of the main olfactory bulb have indicated that there are subpopulations of granule cells with different dendritic patterns in the external plexiform layer (EPL). Small, extracellular injections of horseradish peroxidase (HRP) were made iontophoretically into superficial and deep parts of the EPL and the granule cell layer (GCL) in adult rats. Superficial EPL injections principally labeled superficial granule cell somata, whereas deep EPL injections labeled both superficial and deep granule cell somata. Injections in the superficial GCL labeled granule cell dendritic processes extending across the entire EPL. However, deep GCL injections labeled few granule cell dendrites in the superficial EPL, but labeled many such processes in the deep EPL. These results were the same in material processed with the Hanker-Yates procedure, where the morphology of individual neurons could be studied, and in the more sensitive tetramethyl benzidine procedure. Serial reconstructions of individual granule cells were made from both HRP and Golgi-Kopsch material. The distal dendrites of deep granule cells reached only as far as the deep EPL, where they branched extensively and had many dendritic spines. The dendrites of superficial granule cells, however, reached the most superficial part of the EPL where they ramified most extensively. The superficial granule cells typically had a higher spine density in the superficial part of the EPL than in the deep part. On the basis of these results, we conclude that the superficial granule cells predominantly innervate the superficial EPL and that the deep granule cells exclusively influences on the bulbar output neurons, the mitral and tufted cells, through reciprocal dendrodendritic synapses. Since the secondary dendrites of the tufted cells ramify in the superficial EPL and the dentrites of most mitral cells ramify in deep EPL, the superficial and deep granule cells may preferentially modulate the responses of tufted and mitral cells, respectively.