Astrocyte subtypes in the rat olfactory bulb: Morphological heterogeneity and differential laminar distribution

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Abstract
Despite increased recognition of the importance and heterogeneity of astrocyte functions throughout the central nervous system (CNS) relatively little attention has been paid to morphological diversity among astrocytes. Recent studies have indicated that subsets of astrocytes are involved in glial‐axonal interactions critical to both development and reinnervation of the rat olfactory bulb. Here, we have characterized the morphologies and distribution of astrocytes within anatomically and functionally distinct layers of the adult main olfactory bulb (MOB). Using a known immunohistochemical marker for astrocytes, glial fibrillary acidic protein (GFAP), and the classic gold sublimate method, we identified six astrocyte subtypes based on their morphology and distribution: (1) unipolar, (2) irregular, (3) wedge‐shape, (4) circular, (5) semicircular, and (6) elongate. Unipolar, irregular and wedge‐shape astrocytes have not been previously described in the CNS. The unipolar and irregular types are located exclusively in the olfactory nerve layer. Wedge‐shape astrocytes are unique to, and are the major subtype in, the glomerular layer. These three morphologically unique astrocyte subtypes may correspond to olfactory nerve layer (ONL) and glomerular layer (GL) astrocytes, which express molecules that regulate axonal growth or synaptogenesis during development and/or regeneration of the olfactory nerve. In the glomerular layer, astrocytes are highly organized with respect to the glomeruli. Individual astrocytes are loyal to a single glomerulus. In the external plexiform layer, astrocytes are spaced relatively uniformly. In the granule cell layer, astrocytes appear to compartmentalize granule cell aggregates, recently shown to be coupled by tight junctions. The distribution and patterns of astrocyte processes and the density of GFAP immunoreactivity are distinctive for each of the layers of the olfactory bulb. The spacing of astrocytes and the organization of their processes may be important to compartmentalization of neuronal functions. High levels of GFAP immunoreactivity correlated with layers of high neuronal plasticity. The morphological diversity and differential distribution of astrocytes in the olfactory bulb reported here support growing evidence for functional diversity of astrocytes and important interactions among specific astrocyte and neuron subtypes. It is reasonable to hypothesize, therefore, that as for neurons, morphologically distinctive astrocyte subtypes may correspond to functionally specific classes.

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