A Golgi study of the sixth layer of the cerebral cortex. I. The lissencephalic brain of Rodentia, Lagomorpha, Insectivora and Chiroptera

Abstract

A study of the morphological characteristics of the neurons in layer VI of the cerebral cortex was carried out using the rapid Golgi method in several lissencephalic species including Rodentia (rat, mouse, vole (Microtus agrestis) and hamster), Lagomorpha (rabbit), Insectivora (hedgehog) and in the Chiroptera the dwarf bat (Pipistrellus pipistrellus). There was a basic uniformity in the structure of the sixth layer. Main neuronal types in lamina VIa were large pyramidal neurons, triangular or atypical pyramidal cells, multiapical pyramidal neurons, inverted pyramids, fusiform neurons, Martinotti cells and bi-tufted cells. Main neuronal types in lamina VIb were medium sized, flattened pyramids, large and small horizontal neurons, horizontal pyramidal cells, fan shaped neurons and multipolar spinous neurons with long descending axons. Sparsely spinous and spine-free multipolar neurons with short axons were present in the two laminae of layer VI, but sparsely spinous neurons with axons similar to those found in basket cells of other layers of the cortex were observed mainly in lamina VIa. Neuronal subsystems were tentatively classified on the basis of the course of the axons. Pyramidal neurons, fusiform neurons, multiapical pyramidal cells, inverted pyramidal cells, fan shaped neurons and multipolar neurons with large descending axons were interpreted as being the main source of long projection and association connections. Large horizontal neurons were interpreted as possible ipsilateral association neurons because the horizontal course of the axons over long distances followed the boundary of the deeper region of the sixth layer. Three intracortical (association) subsystems were included. Axons of Martinotti cells and collateral ascending axons of pyramidal neurons (including multiapical pyramidal neurons) formed the ascending interlaminar fibrillary subsystem. Axons of small horizontal cells and horizontal collaterals of pyramidal neurons formed the horizontal intracortical subsystem. Sparsely spinous and spine-free multipolar neurons and bi-tufted cells were the main source of the local, non-horizontal fibrillary subsystem.