Shaping the Cortical Landscape: Functions and Mechanisms of Top-Down Cortical Feedback Pathways

Cortical feedback pathways are proposed to guide cognition and behavior according to context and goal-direction. At the cellular level, cortical feedback pathways target multiple excitatory and inhibitory populations. However, we currently lack frameworks that link how the cellular mechanisms of cortical feedback pathways underlie their cognitive/behavioral functions. To establish this link, we expand on the framework of signal routing, the ability of cortical feedback pathways to proactively modulate how feedforward signals are propagated throughout the cortex. We propose that cortical feedback modulates routing through multiple mechanisms: preparing intended motor representations, setting the trigger conditions for evoking cortical outputs, altering coupling strengths between cortical regions, and suppressing expected sensory representations. In developing this framework, we first define the anatomy of cortical feedback pathways and identify recent advances in studying their functions at high specificity and resolution. Second, we review the diverse functions of cortical feedback pathways throughout the cortical hierarchy and evaluate these functions from the framework of signal routing. Third, we review the conserved cellular targets and circuit impacts of cortical feedback. Fourth, we introduce the concept of the "cortical landscape," a graphical depiction of the routes through cortex that are favored at a specific moment in time. We propose that the cortical landscape, analogous to energy landscapes in physics and chemistry, can capture important features of signal routing including coupling strength, trigger conditions, and preparatory states. By resolving the cortical landscape, we may be able to quantify how the cellular processes of cortical feedback ultimately shape cognition and behavior.