The neurons of the glomerular layer of the rat olfactory bulb have been studied using Nissl staining and Golgi-Kopsch impregnation in light microscopy to define the size, shape and morphological features of individual cell somata, dendrites and axons; these have been correlated with electron-microscopic material in which fine-structural characteristics were also noted for each cell type, particularly synaptic specializations. Three neuron types are described: the external tufted and periglomerular cells of classical microscopy, and additional, superficial short-axon cells; a description of the glomerular arborizations of the mitral and deep tufted cells is also included. The tufted and mitral cells show large, non-spiny glomerular dendritic arborizations, having terminal varicosities, the external tufted cells being more limited in their branching than the deeper cells. External tufted cells have large somata and abundant cytoplasm containing stacks of Nissl material; their main dendrites are characterized by pale cytoplasm and a regular array of neurotubules. Reciprocal dendro-dendritic and somato-dendritic synapses are commonly found, the tufted/mitral cells containing spherical vesicles and contacting by means of asymmetrical membrane thickenings; the other profile involved is a gemmule containing large flattened vesicles and associated with a symmetrical thickening. The periglomerular cells are smaller, with a spiny glomerular arborization, as well as some other dendrites; all the dendrites of these cells tend to be of irregular outline. They have a dark nucleus and very little somatic cytoplasm; somatic and dendritic appendages are common and often contain large flattened vesicles. Synapses oriented from the dendritic shaft or gemmule also show such vesicles, invariably associated with symmetrical thickenings. The superficial short-axon cells are characterized by the entirely periglomerular distribution of their dendrites, which are varicose and rarely branch. Of intermediate soma dimensions, but containing dispersed Nissl material, these cells and their stem dendrites show no regions that can be designated as presynaptic.
Features of axon initial segments, axo-somatic and axo-dendritic synapses are also described for each cell, as well as some unusual glial relationships. Reasons are adduced for relating the superficial short-axon cell to the axon terminal type containing small flattened vesicles, as well as for considering that the external tufted and periglomerular cells show the same synaptic specializations at their axon terminals as at their dendritic and somatic synapses. The cells of the glomerular layer are compared with those of the deeper layers of the bulb and atypical synaptic specializations discussed; some physiological implications of these findings are considered.
Differences in Glomerular-Layer-Mediated Feedforward Inhibition onto Mitral and Tufted Cells Lead to Distinct Modes of Intensity Coding
