Operating principles of the cerebral cortex as a six-layered network in primates: beyond the classic canonical circuit model.

The cerebral cortex performs its computations with many six-layered fundamental units, collectively spreading along the cortical sheet. What is the local network structure and the operating dynamics of such a fundamental unit? Previous investigations of primary sensory areas revealed a classic "canonical" circuit model, leading to an expectation of similar circuit organization and dynamics throughout the cortex. This review clarifies the different circuit dynamics at play in the higher association cortex of primates that implements computation for high-level cognition such as memory and attention. Instead of feedforward processing of response selectivity through Layers 4 to 2/3 that the classic canonical circuit stipulates, memory recall in primates occurs in Layer 5/6 with local backward projection to Layer 2/3, after which the retrieved information is sent back from Layer 6 to lower-level cortical areas for further retrieval of nested associations of target attributes. In this review, a novel "dynamic multimode module (D3M)" in the primate association cortex is proposed, as a new "canonical" circuit model performing this operation.