The anaxonic granule cell of the olfactory bulb is believed to inhibit mitral and tufted cells through reciprocal dendrodendritic synapses. However, little is known about the detailed input-output properties of the granule cell. This study explores the functional properties of granule cells by using detailed reconstructions of Golgi-impregnated granule cells as the basis for computational models. Three Golgi-impregnated granule cells from the olfactory bulbs of C57BL/6j mice were selected for detailed analysis. Measurements were made of the diameter and length of all spine heads, spine necks, and dendritic branches. These measurements formed the basis of a compartmental model of each cell in which simulations of the spread of synaptic potentials within the dendritic tree were performed with SABER (Analogy, Inc.), a circuit analysis program. The results show that the degree of spread of synaptic potentials can define functionally related subsets of spines within the dendritic tree. The size of these subsets varies with the anatomical location of the input spine, the magnitude of the input, the time course of the input, the size of the spine neck resistance, and the activity of other spines. The data indicate that the functional organization of granule cell dendritic arbors is more complex than previously thought: between the level of the individual spine and the entire dendritic tree are several levels of subsets of spines that can mediate discrete localized inhibition onto subsets of mitral or tufted cell secondary dendrites within the external plexiform layer of the olfactory bulb.
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