Chemoanatomical organization of the noradrenergic input from locus coeruleus to the olfactory bulb of the adult rat

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Abstract
The locus coeruleus contains noradrenergic neurons which project widely throughout the CNS. A major target of locus coeruleus projections in the rat is the olfactory bulb (Shipley et al.: Brain Res. 329:294–299, '85) but the organization of the projections within the bulb has not been systematically examined. In this study, the laminar distribution and densities of locus coeruleus-noradrenergic fibers in the main and accessory olfactory bulbs were determined with anterograde tracing and immunocytochemical techniques. Following iontophoretic injections of 1% wheat germ agglutinin-horseradish peroxidase into the locus coeruleus, the densest anterograde label in the accessory olfactory bulb was observed in the external plexiform layer, granule cell layer, and especially in the internal part of the mitral cell layer. Virtually no label was observed in the glomerular layer. In the main olfactory bulb, labelled axons were observed in the granule cell layer, in the internal and external plexiform layers, occasionally in the mitral cell layer, and least often in the glomerular layer. Noradrenergic fibers in the olfactory bulb were identified by using immunocytochemistry with an antibody to dopamine-β-hydroxylase. Laminar patterns and densities of noradrenergic innervation were determined with quantitative image analysis. In the accessory olfactory bulb, the densest innervation was in the innermost portion of the mitral cell layer followed by the granule cell layer, the superficial part of the mitral cell layer, and the external plexiform layer. The density of fibers in the glomerular layer was least. The laminar pattern of noradrenergic fiber distribution in the main olfactory bulb was similar to that in accessory olfactory bulb. The present studies demonstrate that locus coeruleus-noradrenergic fibers terminate preferentially in the internal plexiform, granule cell, and external plexiform layers. This suggests that the major influence of the locus coeruleus input to both the main and accessory the olfactory bulbs is on the predominant neuronal element in those layers, the granule cells. Additional studies are needed to resolve how this input influences specific olfactory bulb circuits.