The synaptology of the granule cells of the olfactory bulb

ABSTRACT The synapses related to the granule cells of the olfactory bulb of rat brain have been studied in aldehyde-fixed material. The synapses can be divided into three classes: (1) those which have asymmetrical synaptic membrane thickenings and spheroidal synaptic vesicles; (2) those with symmetrical synaptic thickenings and flattened vesicles; and (3) the reciprocal synapses, one half of which (from mitral to granule cell) has an asymmetrical synaptic thickening associated with spheroidal vesicles, while the other half (from granule to mitral cell) has a symmetrical synaptic thickening and flattened vesicles. Qualitative observations, supported by preliminary quantitative measurements, suggest that it may be possible to divide both the spheroidal and flattened-vesicle types into two further varieties, on the basis of size, The smaller variety of spheroidal vesicles is found in most axon terminals, while the larger spheroidal vesicles are present in mitral cell dendrites and in some of the axon terminals. The flattened vesicles associated with symmetrical synapses which are oriented on to the granule cells are smaller than the spheroidal vesicles, but the flattened vesicles in the spines and gemmules of the granule cells are the same size or larger than the spheroidal vesicles. The division of flattened vesicles into two sizes is supported by statistical analysis of measurements of these vesicles, but because of difficulty in identifying the axon terminals with asymmetrical synapses there is no quantitative evidence for such a division of spheroidal vesicles. The asymmetrical synapses are found predominantly on spines, gemmules, and dendritic varicosities, although they are occasionally present on shafts of dendrites and on the cell somata. The symmetrical synapses are almost completely restricted to the shafts of the peripheral processes and the deep dendrites, and to the cell somata; only very rarely are synapses of this type found on spines, and then always in conjunction with an asymmetrical synapse.