Morphology and Synaptic Connections of Small Myelinated Primary Trigeminal Axons Arborizing among Neurons in the Border Zone of Rat Trigeminal Nucleus Oralis

Abstract

The anterograde transport of horseradish peroxidase (HRP) was used to examine the morphology and synaptic connections of a morphologically distinct group of small-diameter primary trigeminal axons that arborize throughout the border zone (BZ) of rat trigeminal nucleus oralis. Thinly myelinated parent branches (0.75-1.5 μm in diameter) descending in the spinal V tract (SVT) were seen to issue medially directed collaterals that entered BZ, where they branched and eventually terminated by giving rise to thin terminal strands characterized by several relatively widely spaced axonal endings. Based on the size and morphology of the parent branches in SVT, in the root entry zone, and in the sensory root of the trigeminal nerve, these primary axonal (P) endings are considered to be derived from small-diameter myelinated primary trigeminal axons (SDMA). The P endings measured 1-2 μm in diameter and contained numerous agranular spherical (40-60 nm) synaptic vesicles. In the BZ neuropil, most P endings lay in glomeruli, where each formed at least one asymmetrical axodendritic synapse on a dendritic shaft. It is at these synapses that this group of primary axons is thought to transfer its input directly to the dendritic arbors of BZ neurons. A small (0.5-1.5 μm) axonal (F) ending filled with flattened synaptic vesicles (29 × 60 nm) was observed to form at least one symmetrical to intermediate axoaxonic synapse on the P ending, as well as at least one axodendritic synapse on the same dendritic shaft receiving the primary input. Some F endings only contacted dendritic shafts. In view of their symmetrical to intermediate synaptic contacts, F endings are thought to belong to axons derived from at least one source that can inhibit or diminish the firing rate of BZ neurons in response to SDMA input. This would be accomplished either postsynaptically through the axodendritic synapses on the dendritic shafts, and/or presynaptically through the axoaxonic synapses on the P endings.