Several types of short axon cells of the mammalian olfactory bulb have been described after Golgi impregnation. Two of these types have been observed in our material after treatment with the NADPH-diaphorase procedure or after immunohistochemistry for neuropeptide-Y (NPY). The cells stained by the two procedures have similar morphologies and distributions. A less extensive series of observations confirms that similar cells also display somatostatin (SS)-like immunoreactivity. One of these cell types corresponds to the superficial short axon cell of Golgi and electron microscopic studies. The dendrites of this cell lie within the periglomerular region and in the superficial external plexiform layer (EPL), generally lying parallel to the glomerular layer. In some cases the axon has been traced across the EPL into the granule cell layer (GCL). This cell may provide another route of interaction between the periglomerular region and the granule cells in addition to the influences conducted by basal dendrites and axon collaterals of some mitral and tufted cells. A type of deep short axon cell is also visible with these two procedures. It lies deep in the granule cell layer, frequently near the ventricular layer and its dendrites lie parallel to that layer. This deep short axon cell is stained with much greater frequency by the NADPH-diaphorase and NPY procedures than is the superficial short axon cell. It corresponds most closely to the Blanes or Golgi cells of the Golgi impregnation literature, but it appears to differ from these cells in the position and orientation of its dendrites. No spines have been observed on either the superficial or deep cells in this series. Many glomeruli are also stained by the NADPH-diaphorase procedure, but are not NPY or SS immunoreactive. This may provide additional evidence for functional differences between glomeruli in local regions of the olfactory bulb.
The pattern of output of mitral and tufted cells of the rat olfactory bulb (OB) to layer Ia overlying the pars externa (pE) of the anterior olfactory nucleus (AON) has been studied in the rat by iontophoresis of horseradish peroxidase and Phaseolus vulgaris-leucoagglutinin. These agents labeled mitral and tufted cells and at least the proximal portion of their axons. In most cases we observed small branches from axons of the lateral olfactory tract that appear to terminate in the region of the pE AON, while the main axon could often be traced for considerable distances past these branches. These branches are assumed to terminate in the pE AON because they could not be traced to other terminal regions, because they ramify in layer Ia, and because they usually show small swellings characteristic of axons in terminal regions. Although each ramification could be extensive, we found that the positions of these small branches were related to the positions of the injections within the OB. Dorsal medial injections labeled dorsal branches. Ventral medial injections labeled ventral branches. Injections on the lateral face of the OB labeled intermediate branches. The centers of the regions within which branches were labeled were strongly correlated with the positions of the injection around the circumference. Comparison of the anterior-posterior axis of the OB produced no such strong correlation. Reconstructions of axons showed that terminal branches arise from both mitral and tufted cells, although at least some mitral cells are shown not to have such branches in the pE AON. Studies of the patterns of dendrites and terminals in the pE AON indicate that this region has the same pattern of layer Ia and Ib terminals seen in other olfactory cortical regions. The pE AON cell layer is intercalated just below the boundary between layers Ia and Ib. Since dendrites of the underlying pars lateralis of the AON (pL AON) penetrate into layer Ia over much of the pE AON, it is necessary to remember that at least part of the pL AON may also receive topographically organized inputs.
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