The Pyramidal Cell and its Local-Circuit Interneurons: A Hypothetical Unit of the Mammalian Cerebral Cortex

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Abstract
A pyramidal cell with five of its local-circuit interneurons (Cajal–Retzius, Martinotti, Cajal double-bouquet, basket, and chandelier cells), constitutes a distinct structural/functional assemblage of the mammalian neocortex. This pyramidal/local-circuit neuronal assemblage is proposed herein as a basic neocortical unit. This unit is shared by all mammals, embodies both specific structural as well as functional elements, and constitutes an essential developmental building block of the neocortex. In the model, the pyramidal cell represents a distinct, stable, projective, excitatory neuron that has remained essentially unchanged in the course of mammalian phylogeny. On the other hand, its local-circuit interneurons are more likely to be inhibitory and less stable, designed perhaps to adapt, and modify in response to environmental needs. The proposed model infers that the number of pyramidal cells contacted by each local-circuit interneuron as well as the number of synaptic contacts established with each one are elements acquired post-natally in response to individual needs. Thereby, the overall three-dimensional distribution and extent of these pyramidal/local-circuit neuronal assemblages should be species-specific, variable among individual of the same species, and able to adapt in response to environmental needs. The model introduces a different approach, perhaps a new vantage point, for the study of the basic structural organization of the mammalian cerebral cortex. Relationships of the proposed model to cortical function in general and to learning behavior in particular are discussed.