Various types of non-pyramidal hippocampal neurons project to the septum and contralateral hippocampus

Abstract
The morphology was studied of hippocampal neurons which had their somata in the hilus of the area dentata, and in stratum radiatum or stratum oriens of Ammon's horn, and which sent projections to the septum and contralateral hippocampus, respectively. The fluorescent marker Fast Blue was injected into the septum or contralateral hippocampus. Somata were then identified by their fluorescent label in slices of perfused brains. After intracellular injection of these somata with Lucifer Yellow, it was found that contralaterally projecting neurons were pyramidal cells, inverted fusiform and multipolar cells in CA3c, and stellate, fusiform and multipolar cells in the hilus. After septal injections, we identified two groups of aspiny stellate cells in the hilus; pyramidal basket cells, polygonal basket cells, horizontal basket cells in stratum oriens; and stellate cells in stratum radiatum of CA1 and CA3, as well as pyramid-like aspiny cells in stratum radiatum of CA1. These cells also had short locally arborizing axons, thus probably contributing to local circuits. Such cells may constitute a third class of hippocampal neurons combining the properties of principal cells and interneurons. These results support the opinion that the simple concept of separating hippocampal cells into projection neurons and local-circuit neurons needs reconsideration.