The Neuropil of the Glomeruli of the Olfactory Bulb

Abstract

The neuropil of the glomeruli of the rat olfactory bulb has been studied with the electron microscope with a view to elucidating the type of processes involved - dendrites, appendages and axons - their cellular identity, and the synaptic relationships they establish. The problems encountered in defining these are considered and criteria based on the previous study of neuron types and on examination of serial sections are put forward. The glomeruli are large structures containing many thousands of processes and are the sole site of termination of the olfactory receptor axons. The terminals of the latter are characteristically electron-dense, allowing identification in normal material; they run through the glomeruli making many synapses by means of spherical vesicles and asymmetrical thickenings on to all types of dendritic profile. The glomerular dendritic arborizations of mitral and tufted cells, which are indistinguishable from each other, start as large, fairly regular, pale profiles but become increasingly varicose as they branch and diminish in size. They regularly show groups of spherical vesicles, often in association with asymmetrical synaptic thickenings directed from the dendrite; these are typically associated with return, reciprocal, synapses of the symmetrical type from profiles containing large flattened vesicles. These latter profiles are those of the dendrites and gemmules of periglomerular cells; the dendrites are of irregular outline and give rise to many appendages, mostly gemmules making synaptic contact with mitral or tufted cell dendrites. A small number of pale axon terminals containing either small or large flattened vesicles, derived from short-axon and periglomerular cells respectively, synapse with symmetrical thickenings on to the periglomerular cell dendritic processes. Close associations of particular types of axo-dendritic and dendro-dendritic synapses on interconnecting processes, termed synaptic patterns, are described and their significance considered. The nature of the glomerular interactions is discussed and then placed in the context of other, smaller glomeruli in the central nervous system; certain common principles of glomeruli are suggested.