Regional variability and postsynaptic targets of chandelier cells in the hippocampal formation of the rat.

Chandelier cells are specialized cortical GABAergic neurons that establish synaptic contacts exclusively with the axon initial segments of principal neurons. They are found in all regions of the hippocampal formation. Here we describe their morphological features in the hilus and in regions CA1 and CA3 by using Golgi/electron microscopy. Attempts were also made to identify the target neurons of chandelier cells in the hilus and entorhinal cortex. Golgi-impregnated chandelier cells display a complex axonal arbor in CA1, with many collaterals forming strings of boutons. The axon plexuses of such cells are less developed in CA3, whereas those in the hilus cover the entire region, although single collaterals are rather simple, with only a few boutons. The dendrites of chandelier cells in CA1 and CA3 have an orientation similar to that of pyramidal cell dendrites and are thus likely to be activated by the same afferent fiber systems. The hilar chandelier cells do not give rise to dendrites invading the molecular layer. Thus, these cells may not receive a dense input from the entorhinal cortex but may be driven by the abundant mossy fiber collaterals in the hilar region. In the CA1 and CA3 regions, the axons of chandelier cells contact the axon initial segments of pyramidal cells. In the hilar region, gold-toned boutons were found to impinge on the initial segments of neurons displaying characteristics of mossy cells. This notion was substantiated by electron microscopic analysis of mossy cells identified by intracellular injection of Lucifer yellow. Those cells regularly showed numerous symmetric synapses on their axon initial segments. Entorhinohippocampal projection cells, identified by injection of horseradish peroxidase into the hippocampus, were found to be preferential targets of chandelier cells in the entorhinal cortex. Our data point to regional variations in chandelier cell morphology and connectivity and indicate that chandelier cells are a principal component of inhibitory mechanisms in all stations of the main excitatory pathway of the hippocampal formation.