Dendritic and axonal organization of mitral and tufted cells in the rat olfactory bulb

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Abstract
The output cells of the main olfactory bulb, the mitral and tufted cells, can be categorized into subclasses on the basis of their intrabulbar dendritic and axonal characteristics. Their form was studied in rats following labeling by iontophoretic injection of horseradish peroxidase into the external plexiform layer (EPL). The fact that these extracellular injections labeled small numbers of neurons permitted reconstruction of individual cells. The injection depth within the EPL determined the type of cells labeled. Secondary dendrites of each cell type lay in one of three partially overlapping zones in the EPL. The deepest zone contained the secondary dendrites of one group of mitral cells (Type I), which had the deepest and longest dendrites of the output cells. An intermediate zone of the EPL contained the secondary dendrites of middle tufted and a second class of mitral cells (Type II). The superficial zone, adjacent to the glomerular layer, contained the relatively short, asymmetric dendritic fields of external tufted cells. The few labeled internal tufted cells had secondary dendrites in either the intermediate or deep zones. Every cell type, except the Type I mitral cells, had axon collaterals in the internal plexiform and upper granule cell layers. No cell types had axons re-entering the EPL. These results for output cells combined with our previous observations on granule cells point to a functional sublaminar organization of the EPL that has not previously been proposed.